Orchids of Britain and Ireland

Orchids
of Britain and Ireland

A Field and Site Guide
000 prelims intro.indd 1
29/1/09 12:53:14
For Rita
29/1/09 12:53:14
Orchids
of Britain and Ireland
A Field and Site Guide
Second Edition
Anne and Simon Harrap
A & C BLACK
LOndOn
12/2/09 17:29:38
ACKNOWLEDGEMENTS
We would especially like to thank Dr Tony Leech for much wisdom as well as practical help and advice (he read and re-read
most of the book), and Lenny Thomson for her equally good-humoured helping hand. Julie Dando very, very patiently performed her design alchemy and has worked wonders, and at A&C Black Nigel Redman can claim the credit for initiating the
project and provided help and support throughout, while Jim Martin exercised his considerable editorial skills. For the provision of additional photographs we would like to thank Richard Bateman, Robin Chittenden www.harlequinpictures.co.uk,
Sean Cole, Bob Gibbons/Natural Image, Richard Gulliver, Michael Frost, Nigel Redman, Craig Robson, Paul Sterry/Nature
Photographers Ltd and Peter Wakely (English Nature). We also thank Richard Millington for his delightful illustration of Bee
Orchid varieties. We are very grateful to the Botanical Society of the British Isles and CEH Monks Wood for allowing us
reproduce the maps contained in the recent New Atlas of the British and Irish Flora (Preston et al. 2002), and for facilitating
their reproduction here; we would particularly like to thank Henry Arnold, the Database Manager, for his time and trouble.
Mark Van Beirs assisted with French translations, Keith and Karen Ashby with German, and Gareth Burnell with Latin. For
providing references we thank the Library of the Royal Botanic Gardens, Kew and also Jan Gebbie (Scottish Natural Heritage)
and Joan Chapman (Glasgow Natural History Society). The staff at Holt Public Library in Norfolk assisted cheerfully with
our many requests. Regarding matters botanical we especially thank Richard Bateman for answering many queries regarding
taxonomy and nomenclature and for providing advance copies of papers; similarly Pete Hollingsworth assisted with our
research on helleborines. Mavis and Richard Gulliver gave advice and unpublished material on Irish Ladys Tresses and Richard
Hedley provided much information on Sword-leaved Helleborine (and suggested a translation for Cephalanthera), as did
David Ball (Hampshire County Council). Plantlife International kindly provided copies of their Back from the Brink dossier
on Sword-leaved Helleborine and Reg Land advised on Fen Orchid. Sean Cole supplied much information on Ghost Orchid
as well as other species. Jill Sutcliffe (English Nature) kindly read through all the proofs and made many useful suggestions.
We would also like to thank Richard Abbott, Steve Alton, Debbie Allan, Norman Baldock, Ian Bonner, Janet
Canning, David Carrington, Clive Chatteris, Robin Chittenden, Peter Clarke, Sean Cole, Les Colley, Richard Collingridge,
David Cottridge, Phil Davey, Charles David, Ian Denholm, Tim Dixon, Stuart Dunlop, Bob Ellis, Tom Ennis, Rachael
Fickweiler, Michael Foley, Maggie Gamble, Paul Hackman, Frank Hunt, James R. Hunter, Kevan Joynes, Brian Laney, Mandy
Leivers, Ian Livermore, Alan Lewis, Alex Lockton, Tim McGrath, Steve Madge, Stephen Martin, Nigel Milbourne, Dr E.
Charles Nelson, Malcolm Ogilvie, John Oxenford, Gavin Peplow, Michael Parsons, E. G. Philip, Chris Pogson, Rachel
Remnant, A. J. Richards, Craig Robson, Dr Francis Rose, Martin Sanford, Brendan Sayers, M.-A. Selosse, Alan Showler, Terry
Smith, Ron Stanbridge, Jon and Lucinda Starling, Malcolm Storey, Giles Strother, Ian Taylor, Peter Thompson, Chris Sydes,
Darrell Watts, Dave White, Steve Whitehouse, Martin Woodcock and Julian Woodman.
Of course, we take responsibility for any errors in the text and would be delighted to receive comments and
corrections. We can be contacted via our website: www.norfolknature.co.uk, or via the publishers. For assistance in the preparation of the second edition we would like to thank Richard Bosanquet, Tom Ennis, John Knowler, Geoff Musker and Andy
Scobie, but especially Richard Bateman and Sean Cole for their continuing support and for answering many questions, and
Roland de la Moussaye for facilitating a visit to North Uist.
Published in 2005 by A&C Black Publishers Ltd,

36 Soho Square, London W1D 3QY
www.acblack.com
Second edition published in 2009; electronic edition 2010
Copyright 2005 and 2009 Anne and Simon Harrap
Photographs 2005 and 2009 Simon Harrap except where indicated otherwise
ISBN 978-1-408-10571-9
eISBN 978-1-4081-3323-1
A CIP catalogue record for this book is available from the British Library
All rights reserved. No part of this publication may be reproduced or used in any form or by any means photographic,
electronic or mechanical, including photocopying, recording, taping or information storage and retrieval systems
without permission of the publishers. This book is produced using paper that is made from wood grown in managed, sustainable forests. It is natural, renewable and recyclable. The logging and manufacturing processes conform to the environmental regulations of the country of origin.
Printed in China by Lion Productions Ltd.
Designed by Fluke Art, Cornwall
10 9 8 7 6 5 4 3 2 1
Contents
INTRODUCTION
ORCHID BIOLOGY
ORCHID HABITATS
ORCHID CONSERVATION
FIELD GUIDE
Notes on the species accounts
Species Accounts
Ladys-slipper
Red Helleborine
Sword-leaved Helleborine
White Helleborine
Lesser Twayblade
Common Twayblade
Birds-nest Orchid
Marsh Helleborine
Dark-red Helleborine
Broad-leaved Helleborine
Violet Helleborine
Narrow-lipped Helleborine
Dune Helleborine
Lindisfarne Helleborine
Green-flowered Helleborine
Ghost Orchid
Fen Orchid
Bog Orchid
Coralroot Orchid
Creeping Ladys-tresses
Irish Ladys-tresses
Autumn Ladys-tresses
Summer Ladys-tresses
Musk Orchid
Man Orchid
Monkey Orchid
Military Orchid
Lady Orchid
Early Purple Orchid
Small White Orchid
Lesser Butterfly Orchid
Greater Butterfly Orchid
Common Fragrant Orchid
Heath Fragrant Orchid
Marsh Fragrant Orchid
Early Marsh Orchid
7
8
12
18
20
Cypripedium calceolus
Cephalanthera rubra
Cephalanthera longifolia
Cephalanthera damasonium
Neottia cordata
Neottia ovata
Neottia nidus-avis
Epipactis palustris
Epipactis atrorubens
Epipactis helleborine
Epipactis purpurata
Epipactis leptochila
Epipactis dunensis
Epipactis sancta
Epipactis phyllanthes
Epipogium aphyllum
Liparis loeselii
Hammarbya paludosa
Corallorhiza trifida
Goodyera repens
Spiranthes romanzoffiana
Spiranthes spiralis
Spiranthes aestivalis
Herminium monorchis
Orchis anthropophora
Orchis simia
Orchis militaris
Orchis purpurea
Orchis mascula
Pseudorchis albida
Platanthera bifolia
Platanthera chlorantha
Gymnadenia conopsea
Gymnadenia borealis
Gymnadenia densiflora
Dactylorhiza incarnata
25
34
41
48
56
62
67
78
85
92
101
107
113
122
126
136
142
152
160
166
174
181
188
193
202
207
214
222
228
236
243
249
256
263
267
276
29/1/09 12:53:14
Frog Orchid
Common Spotted Orchid
Heath Spotted Orchid
Southern Marsh Orchid
Pugsleys Marsh Orchid
Hebridean Marsh Orchid
Northern Marsh Orchid
Irish Marsh Orchid
Dense-flowered Orchid
Burnt Orchid
Lizard Orchid
Loose-flowered Orchid
Pyramidal Orchid
Green-winged Orchid
Small-flowered Tongue Orchid
Greater Tongue Orchid
Fly Orchid
Bee Orchid
Late Spider Orchid
Early Spider Orchid
Other species
Orchid family tree
Map of vice-counties
Orchid flowering periods
Dactylorhiza viridis
Dactylorhiza fuchsii
Dactylorhiza maculata
Dactylorhiza praetermissa
Dactylorhiza traunsteinerioides
Dactylorhiza ebudensis
Dactylorhiza purpurella
Dactylorhiza occidentalis
Neotinea maculata
Neotinea ustulata
Himantoglossum hircinum
Anacamptis laxiflora
Anacamptis pyramidalis
Anacamptis morio
Serapias parviflora
Serapias lingua
Ophrys insectifera
Ophrys apifera
Ophrys fuciflora
Ophrys sphegodes
288
294
303
309
316
325
328
334
341
345
354
363
366
372
381
384
392
398
405
411
416
417
418
420
SITE GUIDE
Introduction to the site accounts
Useful addresses
Botanical organisations
Abbreviations
Site Accounts
Southeast England
Southwest England and Channel Islands
East Anglia
Central England
Northern England and Isle of Man
Wales
Scotland
Ireland
424
432
438
442
447
452
456
460
GLOSSARY
465
SOURCES OF INFORmATION AND BIBLIOGRApHY
469
INDEx OF SpECIES
479
421
422
423
423
29/1/09 12:53:15
InTROdUCTIOn
Introduction
why this book?
Orchid! The very word conjures up an image of the exotic, beautiful and, above all, rare. It is associated
with wealth, power and the eminently desirable. Few people realise that orchids are very much part of
the natural heritage of Britain and Ireland, and that there are no fewer than 56 species of native wild
orchid. They range from the tiny, green Bog Orchid to the flamboyant Marsh Helleborine and the
gorgeous Green-winged Orchid. They also include Ladys-slipper and Ghost Orchid, probably the
two rarest native wild flowers in the British Isles, and Common Spotted and Early Purple Orchids,
species found throughout the land.
The aim of this book is both to introduce wild orchids to a wider audience and to show those who
think they know about orchids that there is always more to learn. For orchids need friends. Despite
the many schemes designed to alleviate its effects, modern industrial agriculture, combined with
urbanisation, insensitive forestry practices and a rain of nutrients from car exhausts, has resulted in an
ever-increasing homogenisation of the countryside. The beautiful, delicately woven tapestry of fields,
pastures, woods and marshes created unwittingly by the hand of man over many generations is being
put into the equivalent of a food blender that is reducing everything to a monotonous and anonymous
wildlife desert. In the face of this assault, the majority of wild plants are in retreat, including all
orchids. Do not be fooled by cheeky television presenters and glossy wildlife magazines. Although
there is now more money for conservation in general and nature reserves are at last receiving some
desperately needed cash, orchids and much other wildlife besides have disappeared from most of the
landscape. If we can help more people to enjoy, appreciate and value orchids and the places where they
grow, our purpose will have been served.
upper sepal
what is an orchid?
Orchids (the family Orchidaceae) are among
the most diverse groups of plants, with over
1000 genera and at least 25,000 species. Indeed,
the Orchidaceae is the largest and most highlyevolved family of flowering plants.
The family derives its name from the
Greek orchis, meaning testicle, a reference to
the appearance of the underground tubers of
some species. The term orchis was first used by
Theophrastus (c. 370-285 BC) in his Natural
History of Plants; he was a student of Aristotle
and is considered to be the father of botany.
Most people would recognise an orchid, even
those without any particular interest in botany or
gardening at least the gaudy, hot-house hybrids
and some of the more colourful wild orchids.
Giving a precise definition of an orchid is more
difficult, especially a non-technical definition, but
European orchids all share the following features:
l
l
l
l
petal
petal
lateral
sepal
lateral
sepal
column
lip
pollen
masses
caudicle
viscidium
pollinia
rostellum
bursicle
stigma
mouth of
the spur
pollinium (detail)
column (detail)
M A typical orchid flower.
They are perennial herbs rather than trees or shrubs and lack any woody parts.
The leaves are simple and not divided into lobes or smaller leaflets.
The leaves have no stalk and are arranged alternately along the stem.
The flowers are carried in a single spike at the tip of the stem.
29/1/09 12:53:15
8
l
l
ORCHIdS OF BRITAIn And IRELAnd
The ovary is inferior, that is, placed below the sepals and petals.
Male and female parts of the flower, the stamens and the stigma, are not separate but are fused
together into a single structure, the column, that lies in the centre of the flower.
The flower is made up of three sepals and three petals but one of the petals differs from the others,
usually significantly so, and forms the lip (sometimes known as the labellum). This is often
brightly coloured and patterned, and intricately shaped. The lip is actually the uppermost petal
but usually lies at the bottom of the flower because either the ovary or its stalk is twisted (the
flower is therefore resupinate).
Orchid Biology
orchids And fungi
The relationship that orchids have with fungi impacts on all aspects of their biology and, more than
anything else, defines them.
Their seeds require fungi in order to germinate and grow. The seedling spends months or years
underground and during this period is completely dependent on the nutrients that it obtains from
fungi and is mycotrophic (a term deriving from the Greek mukes meaning fungus and trephein to
feed). Seed must, however, be produced in large quantities to ensure that some, at least, will find the
correct conditions for successful germination and growth, including the presence of the correct fungi.
In turn, the need to produce large quantities of seed has powered the evolution of elaborate flowers
and complex pollination mechanisms.
Even when the orchid has appeared above ground as an adult or near-adult plant and is able
to photosynthesise and manufacture its own carbohydrates, in many species it still maintains a
relationship with fungi. In a few orchids the adult plant continues to be entirely dependent on fungi
(the so-called saprophytic orchids). In others, it is probably largely independent of fungi and gains
its nutrients almost entirely from photosynthesis (these are phototrophic). Most orchids, however,
fall somewhere between these two extremes, with both sources of nutrition being utilised, perhaps
in varying proportions depending on the season. The ability to utilise two sources of nutrition allows
orchids to thrive in marginal habitats; some grow in heavy shade and many are found on poor soils.
In the tropics, orchids have extensively colonised the soil-less trunks and branches of trees and are
epiphytes. Finally, the ability to fall back on fungi as a source of nutrition explains why many orchids
are able to become dormant underground for a year or sometimes longer.
mycorrhizas
It is thought that around 90% of the worlds plants have a relationship with fungi. Such a relationship
is known as a mycorrhiza and the fungi that form these attachments are known as mycorrhizal.
Mycorrhizal fungi live in the soil and, unable to manufacture their own carbohydrates by photosynthesis
or obtain sufficient for their needs by the decomposition of organic matter, they invade the root
systems of green plants. But, rather than being parasitic, the fungus actually benefits the host plant
by functioning as an extended root system. The mycelium of the fungus extends far into the soil and
is able to provide the plant with minerals, especially phosphorus; it may also confer some degree of
drought, pest or disease resistance. The plant in turn provides the fungus with carbohydrate that it
has produced through the process of photosynthesis; most plants are able to divert up to 20% of their
carbohydrate to fungi without coming to harm. The relationship between the plant and the fungus
is therefore mutualistic (such mutually beneficial relationships were once termed symbiotic, but this
term is now used for a wider range of interactions).
29/1/09 12:53:15
ORCHId BIOLOGY
l
l
Fungi form several kinds of mycorrhiza:

Ectomycorrhizal fungi form a sheath or mantle over the plants roots (the Greek ectos means
outside). Ectomycorrhiza are the dominant type of mycorrhiza formed by forest trees and
are critical to their growth. Most ectomycorrhizal fungi are macrofungi and many produce
recognisable mushrooms.
Endomycorrhizal fungi (also known as VAM fungi) penetrate the cells of the plants roots (the
Greek endon means within). They are microfungi and do not produce distinctive fruiting bodies.
Poorly-known, they cannot be seen without a microscope and have proved impossible to cultivate
and study in vitro.
Ericaceous mycorrhiza are formed with the roots of various ericaceous plants.
Orchidaceous mycorrhiza are formed with orchids.
orchid fungi
Until very recently, orchid mycorrhizas were thought to differ fundamentally from other mycorrhizal
systems in that the orchid does not provide the fungus with carbohydrates: this must be true of orchid
seedlings which develop entirely underground and have no green leaves. Rather, it is the fungus that
provides the orchid with energy; there is good evidence that carbohydrate obtained by fungi from the
decomposition of organic matter (or from trees, see below) is transferred to the orchid. In short, it
seemed that orchids are parasitic on fungi (or, to use the terminology of recent scientific papers, they
cheat in their relationship with the fungus). There was no evidence for the transfer of nutrients from
the orchid to the fungus, even in mature plants which are able to photosynthesise, but very recently it
was shown that carbon does pass from Creeping Ladys-tresses to their associated fungi, and this has
re-opened the debate regarding the orchid-fungus relationship. Throughout the text we have placed
infection and partner in inverted commas to emphasise uncertain nature of the relationship between
orchids and fungi.
The physical relationship between orchids and fungi is very sophisticated. Fungal hyphae pass
through the outer layers of the orchids root, rhizome or other underground organs and penetrate
the cell walls to form loops and coils, called pelotons. At intervals, the orchid digests these pelotons
and receives water, mineral salts, carbohydrate and other organic compounds from the fungus. So
sophisticated is the orchids use of fungi that it is able to control its spread and confine it to specialist
cells; indeed, some orchids produce phytoalexins which act as a fungicide and prevent the fungi from
reaching tubers and other storage organs. The main fungal associates of orchids are Basidiomycetes
of the Rhizoctonia group (other members of the Rhizoctonia group are soil saprotrophs or
pathogens).
saprophytic orchids
Some species of orchid take the relationship with fungi to an extreme. Birds-nest, Coralroot and
Ghost Orchids have no green leaves (or have the green pigments very much reduced) and throughout
their lives depend entirely on their fungal partner for nutrition (they are fully mycotrophic).
Furthermore, these species do not form associations with the usual orchid fungi of the Rhizoctonia
group. It has been shown that both Birds-nest and Coralroot Orchids form relationships instead with
ectomycorrhizal fungi which are simultaneously in partnership with nearby trees. Via these fungi the
orchids acquire carbohydrates from the trees and therefore they are, in effect, parasitic on the trees.
It has been suggested that forming associations with such ectomycorrhizal fungi may provide the
orchid with a stabler and more reliable source of nutrients, which is particularly important when it
has no other source of nutrition.
Birds-nest, Coralroot and Ghost Orchids are frequently but incorrectly described as saprophytic.
Saprophytes (nowadays more properly known as saprotrophs) derive their nutrition from dead
organic matter; these orchids acquire nutrients from a living fungus.
29/1/09 12:53:15
10
woodland orchids
It has also been shown recently that some other orchids, such as Red, White and Broad-leaved
Helleborines, also form relationships with ectomycorrhizal fungi and are therefore able to utilise
nutrients provided unwittingly by nearby trees. This may explain their ability to thrive in low light
levels and to become dormant underground for long periods. Many are particularly associated with
Beech trees.
fungi and rare orchids

There is mounting evidence that some orchids are very specific about their fungal partners. Various
orchids in the genera Liparis, Goodyera and Spiranthes have been shown to associate with just one species
of fungus and to use the same species, both as germinating seeds and as adult plants. Other orchids
form a relationship with many different fungi as adult plants but are very specific as germinating seeds
(and may associate as seeds with a different species of fungus to any of those used by adult plants). Yet
other orchids seem to use a broad range of fungi for both germination and as adults.
Orchids that require a specific fungus in order to germinate and grow may be very limited in where
they can live, compared to the more generalist orchids, and this has clear conservation implications.
We would venture to guess that some of Britain and Irelands orchids are rare and localised because
their fungal partners are rare and localised. This may well explain why, for example, Burnt Orchid
seldom colonises new sites. Any conservation programme, especially if it involves a reintroduction,
may ultimately be unsuccessful if it fails to take this into account.
orchid seeds
Orchid seeds are rather small and, indeed, are often known as dust seeds. They typically weigh 2-8
micrograms and, in British species, are 0.35-1.4mm in length. They are made up of a relatively simple
embryo enclosed in a hardened carapace and surrounded by the much larger testa, a honeycomb of
dead cell walls that traps air.
Small size confers several advantages. Large
numbers of seeds can be produced at relatively
testa
little cost; in British orchids counts of between
376 and 25,000 seeds per capsule have been
embryo
recorded (Lesser Twayblade and Greater Butterfly Orchid being the extremes). With many
0.5mm
air spaces the seeds are ideally suited to wind
dispersal and can travel long distances. They can M A typical orchid seed.
also float on water, another effective means of
dispersal. However, small size also imposes limitations. The seeds are so tiny that they contain very
little in the way of food reserves (merely a few lipids and proteins) and they depend entirely upon
fungi to provide their nutrients when they germinate.
germinAtion And underground growth

The orchid seed germinates 2-10cm below the surface of the soil. The seedlings are very vulnerable to
desiccation and require the presence of fungi, so it is important for the seed to avoid germinating on
the soils surface. To ensure this, almost all orchid seeds will only germinate in darkness, once rain has
washed them down into the soil or they have been covered by fallen leaves. Unlike other plants, they
do not require light following germination because they do not produce green leaves.
It is thought that most European orchids germinate in the spring (although there is only limited
direct evidence for this) and therefore the seeds must have a mechanism to keep them dormant over
the winter. Germination depends upon the uptake of water by the seed and the hard carapace forms
a barrier against water, slowing or preventing germination for a while. In some orchids the carapace is
incomplete and these seeds germinate much more easily and rapidly.
29/1/09 12:53:16
ORCHId BIOLOGY
11
With its very limited reserves of nutrients, the orchid seed is dependent on fungi from the outset.
In some species, such as Birds-nest Orchid, seeds start to germinate before infection by a fungus,
although the breaking of the seeds dormancy almost certainly requires the fungus to be present in the
immediate vicinity of the seed, which perhaps responds to a chemical signal from the fungus. In other
species, such as Coralroot Orchid, the seeds will only start to germinate after it has been infected by
the specific fungal partner. From the outset the seedling is able to control the extent of infection by
the fungus, confining it to certain areas.
Some species, including Birds-nest and Coralroot Orchids, form relationships with a very
restricted range of fungi, perhaps just one species. Their seeds will only germinate in the presence
of the appropriate fungus. (The implication is that the seeds can remain dormant for relatively long
periods, waiting for the correct fungus to appear.) Other orchids form a relationship with a variety of
common soil fungi that probably occur in all suitable habitats. Examples include the marsh orchids;
their seeds will germinate and start to grow in the absence of the appropriate fungi, presumably
because there is a very high probability that a suitable fungus will be encountered very soon.
Upon germination the seed forms a protocorm. This is a small, often parsnip- or top-shaped
structure with a scatter of root-hairs on its surface; these are single-celled projections that facilitate
fungal infection. The protocorm usually goes on to develop roots and at this stage is often known as
a mycorhizome. The primary function of these first roots is to host fungal activity rather then the
supply of nutrients and water. In most orchids, as the seedling continues to develop, fungal activity is
increasingly confined to the roots, and the mycorhizome, now free of fungal infection, is known as a
rhizome. In some orchids the adult plant grows from a rhizome, but in many species the rhizome is
largely replaced by tubers. In adult orchids, whether they grow from a rhizome or from a tuber, fungal
activity is usually confined to the roots and, in some species, to the slender extremities of the tubers.
PollinAtion
Orchids are renowned for the beauty and complexity of their flowers. These flowers have not, however,
evolved to amaze and delight us, but to fulfil the primary function of the plant, which is to reproduce
itself.
cross-pollination versus self-pollination

Most flowering plants reproduce sexually and the production of the next generation depends upon
the successful unification of male and female gametes to produce seed; the male gamete is pollen
and the female gametes are contained within the ovary. In most plants, including orchids, each
flower produces both male and female gametes. It would seem a simple process therefore for pollen
to be transferred within the same flower to the ovary. Indeed, why bother with the resources and
energy needed to produce a flower at all when a small drab structure would be sufficient to bring
pollen and ovary together? The answer lies in the advantages that cross-pollination brings to the
species.
If pollen and ovary come from different individuals they will each carry a different set of genetic
material and the resulting offspring will not, therefore, be genetically identical to its parents. This
continual mixing of genetic material creates a large gene pool which gives the species flexibility and
adaptability. It also means that individual mutations are disseminated through the population rather
than being passed, unchanged, to the offspring; cross-pollinated species therefore tend to be relatively
uniform in appearance.
If a flower is pollinated with its own pollen (i.e. it is self-pollinated) its offspring will have limited
genetic variation. In turn, their offspring will be similarly limited. In self-pollinated species the overall
genetic variation is rather low and the gene pool is limited, giving the species very little flexibility.
Paradoxically, however, self-pollinated orchids can show more variation between colonies and are
more likely to produce distinct varieties as mutations; these might produce, for example, differently
coloured flowers and are passed down unchanged to the next generation. Bee Orchid is perhaps the
best example of this.
29/1/09 12:53:16
12
There are circumstances where self-pollination is an advantage. If, for example, a single windblown seed produced a flowering Lizard Orchid many miles from other plants of the same species, it
would have no chance of reproducing if it could only be cross-pollinated. If it is able to self-pollinate,
it can produce seed and reproduce itself. Not surprisingly, self-pollination tends to be commoner in
species that have scattered populations, or are at the edge of their range. Another factor may be a
lack of pollinators, and species adapted to deep shade may well benefit from self-pollination; White
Helleborine, a species which is routinely self-pollinated, is successful in Britain, whereas Red and
Sword-leaved Helleborines, which are always cross-pollinated, have declined as increasing shade has
made their woodland homes too dark for their pollinators. Self-pollination may therefore be a useful
strategy in the short-term and it may only be over an evolutionary time scale, involving thousands of
generations, that the advantages of cross-pollination come to the fore.
Most orchids are cross-pollinated but, rather cunningly, a large proportion have adaptations that
prevent self-pollination but then allow the flowers to self-pollinate if, after a few days, a suitable
pollinator has not come along. Conversely, the small number of routinely self-pollinated orchids
are, at least occasionally, cross-pollinated by insects. In the world of orchids, nothing is straightforward.
insects as pollinators
Orchid flowers have evolved to use insects to carry pollen from one plant to another and employ
a variety of mechanisms to attract suitable pollinators. Bright colours and scents advertise their
presence and the lip of the flower acts as a convenient landing platform. Many offer their insect
visitors a reward of nectar but a large proportion do not. Their bright colours and scents are instead
a deceit and they rely on the stupidity of insects, which are slow to learn that the flowers offer no
reward. Fly, Early Spider and Late Spider Orchids (genus Ophrys), have evolved even more elaborate
mechanisms that take advantage of the sexual behaviour of insects (see p.388).
Compared to most other flowering plants, orchids produce large quantities of seeds. Therefore
large quantities of pollen have to be moved between the flowers because each pollen grain can only
produce one seed. In most orchids the pollen grains are amalgamated together in large numbers to
form pollinia. Due to their size and weight, these pollinia must be carried to other flowers by an insect
and must be very securely attached if the process is to be completed successfully. The insect must be of
the right size and shape, both to pick up the pollinium in the first place and to be positioned correctly
in the next flower so that the pollinium makes contact with the stigma and effects fertilisation.
European orchids are not usually pollinated with a whole pollinium, rather just fragments, and thus
a single pollinium can, provided it remains on the insect, pollinate several flowers. But, if too small
a quantity of pollen is deposited the capsule will mature without all the ovules being fertilised, and
there will be many non-viable seeds.
Orchid Habitats
Britain and Ireland form a landscape that has been tamed, with virtually no truly wild places; even
the moors and mountains of Wales and Scotland are not natural, having been stripped of their trees
many generations ago. Orchids therefore have to find a niche in the habitats which mankind has
moulded; some of these habitats are very rich in orchids whereas others are devoid of their grace and
beauty.
It is possible to identify five factors which determine the suitability of any habitat.
Soil chemistry: Many orchids are fussy about the soil chemistry and, as any gardener knows, whether
the soil is acid or alkaline determines to a great extent which plants will thrive. A high proportion of
orchids favour soils that are rich in calcium; these are known as calcareous soils and they are alkaline
with a high pH. Calcium is a major component of chalk and limestone and therefore soils derived
29/1/09 12:53:16
ORCHId HABITATS
13
from these are often excellent for orchids. A similar chemistry is produced by magnesium and some
limestones are rich in this mineral; although they are not calcareous, the terms alkaline or base-rich
can also be used to cover these soils. In practice, however, calcareous, alkaline and base-rich are used
interchangeably. A small minority of orchids favour acid conditions, usually soils that are damp or
wet, and these are usually to be found where peat is formed from bog mosses (Sphagnum spp.), or in
pinewoods.
Soil fertility: Like most wild plants, orchids do best in a soil that is very low in nutrients. This is
largely because a soil that is high in nutrients, either naturally or through the application of fertilisers,
allows a limited range of larger and more aggressive species to thrive, and these will swamp any
orchids. There is also evidence that some nutrients are actually toxic to some orchids (for example,
high levels of phosphorus may be toxic to Green-winged Orchid).
Age: Orchids are great colonists and their dust-seeds enable them to spread long distances.
Nevertheless, there are some unknown factors which limit the ability of many species to move into
new habitats. Colonisation often takes place very slowly, probably over hundreds of years for some
species. It follows that the older a habitat is, the more likely it is to support orchids and the greater
the diversity of species.
Stability: Orchids are relatively long-lived plants and most have a period of immaturity lasting several
years which they spend underground. Both as underground seedlings and as adult plants they cannot
tolerate ground disturbance which would destroy their rhizomes or tubers. Therefore orchids can
only occupy a habitat if it has a degree of permanence; ground that is regularly ploughed or otherwise
disturbed cannot support orchids.
Disturbance: Despite the preference of many species for older, undisturbed habitats, our impression
is that orchids are often found on areas that have been disturbed in the past, albeit a long time ago
in some cases. Old quarries and sand and chalk pits are good examples of this. It is also increasingly
obvious that the continuous, low-level ground disturbance produced by grazing animals is critical to
the long-term survival of many species.
deciduous woodland
Deciduous woodland can be rich in orchids but is, all too often, a disappointment. Woodland in
Britain and Ireland falls into two broad categories; ancient woodland and secondary woodland.
Ancient woodland is found on sites which have probably had an uninterrupted cover of trees
for thousands of years. Throughout Europe, only fragments of this primeval wildwood remain and
Britain and Ireland are no exception, with just 1-2% of the original cover left. Man has continuously
been at work in these woodland fragments, which have been cut for timber, coppiced and grazed,
but the continuity of tree cover has allowed a rich flora and fauna to survive and an ancient wood
will undoubtedly have the greatest diversity of orchids. Such woods can usually be recognised on
the ground by their substantial boundary banks and diverse flora, and on a map will often have a
somewhat irregular outline; historical research, looking at old maps and documents, is often needed
to confirm their provenance, but this is not necessary for the orchid hunter.
Secondary woodland grows where the continuity of tree cover has been broken. If a field that
has been cultivated for decades or even centuries is planted with trees or allowed to tumble down
to woodland it may look like a wood but it will seldom, if ever, acquire the richness and diversity of
ancient woodland (hence the planting of new community forests and millennium woodlands will do
little for orchids). Many woodland plants and animals are very poor colonisers and are unable cross
stretches of inhospitable territory. Importantly, a secondary wood may well lack the complex soil
structure and diversity of soil fungi that is found in an ancient wood and thus lack suitable fungal
partners for some orchids. Overall, with the exception of a few species, secondary woodland will be
poor in orchids (and will often have none).
Management is another factor that can determine how orchid-rich a wood is, whether ancient
or secondary. Until the early 20th century most woodland was intensively managed. Many woods
29/1/09 12:53:17
14
were coppiced cut in small blocks on a rotation of around ten years to provide timber for handcrafts, charcoal and firewood. Coppice stools grow again after each round of cutting, providing an
endless supply of timber. In coppiced woodland periods of sunny, well-lit conditions are followed
by increasing shade as the stools grow again, ideal for some orchids. Abundant labour also allowed
rides to be cut and cleared for access and the limited technology of the day meant that felling and
replanting for large timber was carried out at a relatively slow pace and in a piecemeal manner. All
in all, the more extensive management of this period resulted in woods that were a patchwork of
habitats and much sunnier due to the larger number of open rides and glades. Modern management
techniques depend on the use of minimal labour supplemented by machinery. Coppicing has largely
been abandoned (although now increasingly reinstated by conservation bodies), ride management is
negligible and felling is undertaken in large blocks using heavy machinery. Under such regimes woods
are darker than before and often suffer heavy ground disturbance.
A final factor to affect the suitability of woods is biological. There are ever increasing numbers
of deer in Britain; indeed, it is said that there are more deer now than at any time since William the
Conqueror. Deer eat orchids and some scarce species, for example Narrow-lipped Helleborine, can
lose all their flowers to deer year after year. Another increasingly negative factor, at least in some parts
of the country, is pheasant rearing. In a commercial pheasant-shoot large numbers of young birds are
released into a wood and these can decimate the flora as they scratch and root en masse.
In spite of the many factors that can negatively effect a woods suitability, there are still many
wonderful woods in Britain and Ireland, and the best are often reserves. Characteristic species of
ancient woodland include Common Twayblade, Common Spotted and Fly Orchids, and, especially in
coppiced woodland, Early Purple and Greater Butterfly Orchids. In Kent, Lady Orchid is widespread
and some favoured woods, from southern England to northern Scotland, hold the exquisite, sunloving Sword-leaved Helleborine. Two of our rarest orchids, Red Helleborine and Ghost Orchid, are
confined to deciduous woodland. The overall increase in the level of shade in modern woodland is not
bad for all orchids: Birds-nest Orchid and White, Violet and Narrow-lipped Helleborines can thrive
in very shady woods.
Secondary woodland and plantations of deciduous trees are likely to be much poorer in species,
although two of the more mobile species, Common Spotted Orchid and, in beech plantations, White
Helleborine, can occur in large numbers. The strangely fickle Broad-leaved Helleborine and the
enigmatic Green-flowered Helleborine seem to occur at random.
coniferous woodland
The only coniferous trees that are native to Britain and Ireland are Juniper, Yew and Scots Pine and it
is the last of these that forms a special habitat for orchids. Native Scots Pine is confined to Scotland,
where the remnants of the so-called ancient Forest of Caledon are concentrated on Speyside and
Deeside in the east and around Beinn Eighe in Ross & Cromarty. These are home to Creeping Ladystresses and, more locally, Lesser Twayblade and Coralroot Orchid, with Heath Spotted Orchid in
some of the more open areas.
Conifer plantations have been established all over Britain and Ireland but are usually planted with
non-native species such as Sitka Spruce and Douglas Fir. They were often established on areas that had
been open ground, such as heaths, moorland and sand dunes, but in the post-war period the Forestry
Commission also had a policy of coniferisation and converted large areas of ancient woodland into
conifer plantations. Plants, including orchids, are great survivors and where an ancient woodland has
been coniferised there may sometimes be orchids in rides and along edges, with Common and Heath
Spotted Orchids and Broad-leaved Helleborine the likeliest species. Otherwise, conifer plantations
south of the Scottish border hold little interest with one notable exception: in parts of Norfolk and
northern England, mature pine plantations hold some impressive colonies of Creeping Ladys-tresses.
In Scotland, plantations of pines may acquire a great deal of interest as they mature, taking on some
of the characteristics of native pine woods.
29/1/09 12:53:17
ORCHId HABITATS
15
carr woodland
Carr woodland is dominated by willows and Alder and develops in the wet, waterlogged conditions
found on and around mires, bogs, rivers and lakes. These fast-growing trees are pioneer species and
in a natural system carr woodland usually eventually dries out and other species of tree become
dominant. Carr may hold orchids such as Common and Heath Spotted Orchids, inherited from
the open ground it colonised, but it also has two specialities. In Scotland, Coralroot Orchid is
most abundant in carr and in southern England carr may be the natural habitat of Green-flowered
Helleborine.
grassland
Grassland is a prime orchid habitat, with a large range of species possible, including Common and
Heath Spotted Orchids, Greater and Lesser Butterfly Orchids, Common Fragrant, Green-winged,
Early Purple, Pyramidal, Bee and Frog Orchids and Common Twayblade. Calcareous grassland,
especially the chalk downs of southern England, is the richest; the specialities are Musk, Burnt,
Man, Monkey, Early Spider and Late Spider Orchids and Autumn Ladys-tresses. Neutral grassland
can also be productive and even acid grassland can hold species such as Heath Spotted and Heath
Fragrant Orchids and, in northern Britain, Small White Orchid.
Grassland will always tend to develop into scrub and eventually woodland unless prevented by
either grazing or mowing. It is possible to distinguish between a pasture, grassland which is grazed
for some or all of the year but not cut, and a meadow, grassland from which stock are excluded so
that it can be cut for hay (and often then grazed later in the year). The timing and intensity of grazing
and mowing is very important in determining which orchids are able to thrive and even the animals
concerned can be important; cattle are heavy beasts and can damage the turf but they are selective
eaters and do not eat orchids (although they may stand on them); sheep, lighter on their feet, eat
everything. Some grazing regimes produce a very short turf beloved of Musk and Frog Orchids and
Autumn Ladys-tresses, while longer grass can support Pyramidal, Man and even Lizard Orchids.
It is always worth investigating the various micro-habitats in any area of grassland, which may
be favoured by different orchids. Steeper slopes are often particularly interesting, as are any ancient
earthworks, which may have remained undisturbed for centuries.
limestone pavements
This very specialised habitat is found widely in the north and west of Britain and in Ireland. The
limestone bedrock was first scoured flat by the ice sheets and then weathered by the action of water
and frost to produce blocks of rock (clints) with deep and sometimes treacherous crevices between
them (grykes). Over time, soil may accumulate in the grykes and most pavements would, in a natural
state, be wooded, but many were cleared of trees long ago and some are now rather bare. The range
of orchids present on a limestone pavement depends in part on how much naked rock is exposed;
many pavements have a substantial covering of turf and this holds all the species typical of calcareous
grassland. Limestone pavements also have two specialities: the very local Dark-red Helleborine and,
in the Burren in western Ireland, Dense-flowered Orchid, which grows on short turf.
marshes and fens

Marsh, swamp, fen and bog are all vague terms and are used interchangeably by the layman (and
most botanists too) to describe any sort of waterlogged or seasonally flooded ground. In terms of
their flora, however, there are many different varieties, only some of which are good for orchids. Wet
ground can be categorised in terms of the source of its water, the water chemistry and the management
that it receives. Broadly, the same factors which determine the orchids that can thrive in grassland
operate in marshes too.
All the water may come from rainfall, in which case a bog is formed, either a blanket bog of the
sort which carpets the landscape in northern and western Britain and Ireland, or a raised bog, a much
29/1/09 12:53:17
16
rarer habitat. Bogs are acidic and usually dominated by bog-mosses Sphagnum spp., the dead remains
of which lead to an accumulation of peat. In general, bogs are poor for orchids, although Heath
Spotted Orchid may be frequent on the higher and drier hummocks.
Alternatively, the bulk of the water may come from springs and seepages in the ground, in
which case its chemistry is strongly influenced by the rocks it has passed through on the way to the
surface. This flushing water may be acid or very alkaline or anything in between, but is usually low in
nutrients. Such spring-fed marshes are often called mires in the technical literature and can support
some scarce and local orchids. In southern England valley mires are commonest, in which water seeps
from the ground along the sides of a valley at the boundary of pervious and impervious rocks to form
a long, narrow mire with a central stream. Valley mires are often found in heathland but formerly
would have been widespread and almost every parish would have had its area of boggy ground; most
were reclaimed for agriculture long ago. In the north and west spring-fed mires can be found within
more extensive areas of acid bog and often form a focus for interesting plants.
Because there is a whole range of water chemistry each variety of mire grades into the next in a
complex and often poorly understood manner. Similarly the mix of orchids changes subtly as the
habitat becomes more or less favourable. And, even within a particular mire, there may be many
micro-habitats; in an otherwise very alkaline mire hummocks of Sphagnum may produce locally
acidic conditions.
Where mires are acidic they may hold the diminutive Bog Orchid, as well as the purple-flowered
form of Early Marsh Orchid (subspecies pulchella) and Heath Spotted Orchid. Slightly more neutral
conditions and drier ground favour Lesser Butterfly Orchid and the very local Heath Fragrant Orchid,
as well as the ubiquitous Common Spotted Orchid. Alkaline mires support Marsh Helleborine, and
Marsh Fragrant, Southern Marsh and Northern Marsh Orchids and, in some areas, the very local
Pugsleys Marsh Orchid. Of the real rarities, Fen Orchid and the creamy-flowered subspecies of Early
Marsh Orchid cling on in East Anglia.
As a third alternative, the water in a marsh may come from rivers and streams, either overflowing
their banks from time to time or percolating through the soil. In this case the water may be acid or
alkaline but is usually nutrient-rich, either naturally or frequently due to the run-off of fertilsers from
agricultural land or the discharge of treated sewage. The nutrient-rich water leads to the growth of
lush, tall vegetation, often dominated by Common Reed or by Nettles, Meadowsweet and Hemp
Agrimony. This vegetation may be controlled, however, by mowing or grazing, in which case such
waterside meadows can hold Southern, Northern and Irish Marsh Orchids, Early Marsh Orchid of
the pink-flowered subspecies incarnata and Common Spotted Orchid.
heaths and moors

Heathland is dominated by heather, one or more species of gorse and sometimes various grasses, and
is generally found on sandy or gravely soils in the lowlands of southern England, whilst moorland is
dominated by heather, grasses and a variety of dwarf shrubs and is generally found in the uplands of
northern and western Britain and Ireland. Both heaths and moors are acidic and are likely to hold
few orchids. Heathland itself is usually very dry but is frequently interspersed by valley mires, which
can be rich in orchids (see above). Moorland tends to be wetter and may hold Heath Spotted Orchids
where it grades into bog. In the north and west, damp moorland, especially on north-facing slopes,
can hold Lesser Twayblade.
Pits, quarries and railway lines

When extraction has come to an end, quarries have large areas of bare rock or sparse soil and where
the bedrock is chalk and limestone these can, over time, form superb orchid habitats. Any of the
grassland orchids can occur and, if there is scrub, woodland species such as Fly, Military and Lady
Orchids are also possible. The timescale involved can be lengthy, with medieval workings now forming
excellent sites. Railway lines go through cuttings which may also expose bare rock or produce thin,
29/1/09 12:53:17
ORCHId HABITATS
17
skeletal soils, while embankments may have been built up from calcareous spoil; disused railway lines
are always worth investigating. Sandpits are unlikely to be calcareous but the bare ground can be
favourable to some species, such as Bee Orchid, which can form substantial colonies so long as scrub
does not take over.
Sadly, even these man-made habitats are under threat as any hole in the ground may be seen as a
suitable site for landfill and many important orchid sites have been buried by rubbish.
road verges, green lanes, churchyards and lawns

In the last few decades most hay meadows and permanent pastures have been lost, especially in
England. These unimproved grasslands were ideal for orchids and species such as Green-winged
Orchid, which were common and widespread not so long ago, are now highly localised. Road verges,
green lanes and churchyards are now often the only areas where permanent, unimproved grassland
can be found, although in many cases they are drenched with nutrients from farming operations
and car exhausts, and too overgrown with rank vegetation to support orchids or, in the case of
some churchyards, over-tidied and mown far too often. Nevertheless, Early Purple, Green-winged,
Pyramidal and Bee Orchids and even rarities such as Man and Lizard Orchids can be found in these
habitats. Some Wildlife Trusts have conservation churchyard schemes that may help to identify
the best sites and most counties operate a Roadside Nature Reserve policy to protect and correctly
manage the best verges. Garden lawns are usually cut too frequently and managed with herbicides
and fertilisers and are therefore unlikely to support orchids. On suitable soils, however, unimproved
lawns, even relatively new ones, can be graced by Autumn Ladys-tresses. Increasingly, Common
Spotted and Bee Orchids seem to be turning up on lawns and, once found, are often cherished by the
householders.
dunes and dune slacks

In coastal sand dunes the depressions are often damp and may be flooded in the winter. If the dune
systems are made up from sand that is rich in shell fragments they will be neutral or calcium-rich, and
such dune slacks and the surrounding slopes can then support a very rich variety of orchids. Specialities
include Dune Helleborine, a British endemic found in the dune systems of Anglesey and northwest
England; Lindisfarne Helleborine, endemic to the dunes of Holy Island in Northumberland, and Fen
Orchid, found on the coast of South Wales, as well as the red-flowered subspecies of Early Marsh
Orchid (subspecies coccinea) and, in northern England and Scotland, Coralroot Orchid. Other species
may occur, sometimes in large numbers; Marsh Helleborine, and Common Spotted, Southern Marsh
and Northern Marsh Orchids can be abundant and other species to look for include Broad-leaved
and Green-flowered Helleborines, Common Twayblade, the pale pink subspecies of Early Marsh
Orchid (subspecies incarnata), Frog, Pyramidal and Bee Orchids and even, in one or two areas, Man
Orchid. In many places dune systems have been planted with conifers and although this is bad news
for the specialised dune flora, Dune, Green-flowered and Broad-leaved Helleborines all take readily
to the plantations.
machair
Machair is species-rich grassland found on the wet and windy west-facing coasts of Scotland and
Ireland but nowhere else in the world. Sand that contains many fragments of seashells, and hence rich
in calcium, is blown onshore by Atlantic gales and settles on the low-lying coastal areas. Grassland
develops on these low dunes and periods of extensive grazing have been interspersed with cultivation
for crops. The uncultivated areas often support large numbers of orchids, including two specialities,
the Hebridean Spotted Orchid (the hebridensis subspecies of Common Spotted Orchid) and the very
localised Hebridean Marsh Orchid, endemic to North Uist in the Outer Hebrides. Other orchids
which can be found include the red-flowered subspecies of Early Marsh Orchid, Northern Marsh,
Heath Fragrant, Lesser Butterfly and Frog Orchids, and Common Twayblade.
29/1/09 12:53:17
18
Orchid Conservation
Orchids face three major threats: habitat destruction, habitat change and human predation.
Habitat destruction has clearly taken the greatest toll. Farming, forestry and other developments
have destroyed innumerable orchid sites, especially in the period since World War Two. Most of the
destruction has been state-sponsored through the operations of the Common Agricultural Policy
(CAP) and the Forestry Commission. Between 1945 and 1980 the Forestry Commission attempted
to destroy and re-plant with conifers 200,000 hectares of ancient woodland, to say nothing of the
tens of thousands of hectares of heathland, moorland and sand dunes that were destroyed. The
CAP has been reformed in recent years and attitudes and policies at the Forestry Commission have
changed (although there is still a great reluctance to undo much of the damage done in the name
of near-worthless timber). But, despite much lip-service in recent years, few politicians have any
commitment whatsoever to conservation and when push comes to shove development almost always
takes precedence over wildlife. No wonder the government conservation agencies (English Nature,
the Countryside Council for Wales and Scottish Natural Heritage) have often been accused of being
lame ducks in the face of their political masters, despite the best efforts of their staff. In terms of
habitat destruction, the prospects for orchids remain bleak.
Habitat change has only recently been acknowledged as a major issue. It has come to be recognised
that orchids do not live in stable, climax communities of plants, at least in the British Isles, rather
in habitats that were created and maintained, albeit inadvertently, by people. Grassland, marshes,
heathland and woodland are all the product of traditional land-use. Once these traditions died out,
habitats started to change, slowly at first but then rapidly, and many have become unsuitable for
orchids. Ironically, this applied especially to reserves, where a fence and a keep out sign were often
the limit of any management. Now, conservationists try to replicate the traditional land-uses, often at
great expense, that created and maintained the habitats they manage. The reinstatement of grazing is
often the single most important measure that can be taken to help orchids.
Human predation has often been seen as a major threat to orchids, be it innocent ramblers picking
bunches of flowers or avaricious botanists determined to get another specimen for their collections or
gardens. More recently, photographers and even visitors keen to merely look at plants have joined the
list of threats. The answer has traditionally been secrecy, and details of the locations of the greatest
rarities were and still are jealously guarded; even the location of huge colonies of species such as Burnt
and Early Spider Orchids was veiled in secrecy.
Human predation certainly poses a threat to those species that occur in such small numbers that
a significant part of a population (or even the whole population) can be stolen. There are still cases
where plants are dug up illegally, from Bog Orchid to Lizard Orchid. Perhaps the most notorious in
recent years was the attack on the single Ladys-slipper growing in Silverdale in Lancashire. English
Nature had made the bold decision to allow limited publicity and a large number of people had been
able to admire this beautiful orchid. The fact that this particular plant was probably originally of
garden origin does not in any way lessen the damage done.
Despite the odd incident, however, we are convinced that for most orchids human predation is, in
the final analysis, irrelevant to their fortunes, especially in the face of habitat destruction and habitat
change. Unnecessary secrecy has indeed probably led to the destruction or degradation of many sites,
as those responsible for the land remain in ignorance of its importance. It has also deprived many
people of the enjoyment of seeing the orchids and many potential friends for orchid conservation
have surely been lost in this way.
There have been a few special conservation initiatives involving orchids. In 1983 the Sainsbury
Orchid Conservation Project was established at the Royal Botanic Gardens, Kew. This has involved
research into the propagation of orchids with a view to reintroducing some of the rarer species. A
range of orchids has been involved, including Military and Fen Orchids, with a substantial effort
going into the reintroduction of Ladys-slipper. Reintroductions are controversial, however, with
29/1/09 12:53:18
ORCHId COnSERVATIOn
19
some conservationists arguing that the time and effort could be better spent on conserving existing
populations. A second initiative at the Royal Botanic Gardens was the establishment in 1997 of the
Millennium Seed Bank. This is intended to store viable seeds for as many of the worlds plants as
possible, including, of course, British wild orchids. Techniques have been developed which should allow
orchid seeds to be stored for long periods, although some species cannot yet be cultivated successfully.
what you can do

The first step in orchid conservation is accurate and up-to-date information on their distribution
and abundance. Amateur botanists provide the vast majority of information on plant distribution
in Britain and Ireland via the system of county recorders organised by the Botanical Society of the
British Isles (BSBI); they are always pleased to receive records, with details of the species involved,
numbers, date and location (for contact details see p.422). There is also always a need for volunteers
to undertake practical habitat management on reserves and other sites; the local wildlife trusts are the
first contact if you are keen to get involved.
The greatest contribution individuals can make to orchid conservation is, in our opinion, to
become a local champion. Getting to know an area intimately, finding and recording orchids and other
wildlife, and then badgering local councils, wildlife trusts, government agencies or church-wardens to
sit up and do what is necessary to safeguard the good areas. This may not make you popular in some
quarters but may, in the end, get things done.
orchids and the law

All orchids and, indeed, almost all wild plants, are protected in Britain by the Wildlife and Countryside
Act, 1981 and cannot be uprooted unless you are the owner or occupier of the land or have their
permission to do so (although it is legal to pick them). In addition, some of the rarer orchids enjoy
much greater protection under Schedule 8 of the Act and it is illegal for anyone, even the owner or
occupier of the land, to uproot, destroy or pick these orchids. The term pick is defined to include
gathering or plucking any part of the plant, including collecting seeds. It is also illegal to posses any
live or dead wild plant in Schedule 8, or any part of or anything derived from such a plant, or to trade
in such items. Schedule 8 includes: Ladys-slipper, Red Helleborine and Ghost, Fen, Monkey, Military,
Lizard, Late Spider and Early Spider Orchids (it also currently includes Youngs Helleborine, now
shown to be a poorly defined variant of Broad-leaved Helleborine, and Lapland Marsh Orchid,
now a subspecies of Pugsleys Marsh Orchid). In Northern Ireland, Birds-nest, Bog, Small White,
Pugsleys Marsh, Green-winged and Bee Orchids, Marsh and Green-flowered Helleborines and Irish
Ladys-tresses are specially protected under Schedule 8 of the Wildlife (NI) Order, 1985. In the
Republic of Ireland, Sword-leaved Helleborine, Irish Ladys-tresses, Bog, Small White and Greenwinged Orchids are specially protected under the Flora Protection Order, 1999.
M Derbyshire, June. Grazing is critical to the survival of many orchid populations and is increasingly the linchpin of
conservation management. Paradoxically, however, in the north and west, overgrazing has decimated species such as
Small White Orchid.
29/1/09 12:53:18
20
Field Guide
notes on the sPecies Accounts
conservation designations
The latest UK conservation designations are indicated in red. Several species are also BAP species,
that is included in the UK List of Priority Species and Habitats.
introductions to the genera

Each genus has an introductory section giving some general information about the genus, including
notes on identification for the more difficult species, such as the Epipactis helleborines and the marsh
and spotted orchids. It also includes notes on various aspects of biology, such as growth, reproduction
and pollination, that are common to all the members of the genus. For some genera that contain just
one species worldwide, or where only one species is represented in Britain and Ireland, much briefer
notes on distribution and the generic name are given.
names
The scientific names used in the text and the order of the species accounts follow the list produced by
Richard Bateman (Bateman 2005). Scientific names reflect the relationships between species and in
recent years new evidence, especially from genetic studies, has greatly improved our understanding of
these. This is reflected in new scientific names for some species.
Under the heading Formerly we give scientific names as used in publications that pre-date the
Bateman list. We have tried to include most older scientific names if they have been in general use
since the publication in 1968 of the second edition of Summerhayes Wild Orchids of Britain. We
have not, however, provided an exhaustive list of synonyms. In addition, some older English names
are also mentioned. Under the heading Other names we give some alternative names in current use,
especially in North America.
habitat
The likely habitats for each species are outlined and more detail on many orchid habitats can be
found above. The information applies to Britain and Ireland; in some cases orchids occupy a rather
broader range of habitats elsewhere. Most maximum recorded heights above sea level come from the
New Atlas (Preston et al. 2002).
flowering period
A guide is given to the period in which the species is likely to be in flower (see also chart of flowering
periods on p.420). Flowering times do vary, however, both predictably and unpredictably:
l
Orchids growing further to the north and at higher altitudes tend to flower a little later (although
there is often surprisingly little difference between southern England and Scotland).
l
Orchids growing in wetter habitats will flower later than the same species growing in drier habitats.
l
Orchids growing on or very near to the coast will tend to come into flower a little earlier than
those inland.
l
Orchids growing on sheltered south-facing slopes will flower earlier than those with an exposed,
westerly aspect or those facing north.
There can also be marked and unpredictable variations between colonies, even those close to each
other, and, in recent years, perhaps as a result of global warming, many orchids have been coming into
flower earlier; it is worth bearing this in mind if you are hoping to see a species at its best.
Orchids are notorious for the wide variations from year to year in the number of plants in flower.
It used to be thought that this was related to fluctuations in the size of the population and that some
species, such as Bee Orchid, were monocarpic and therefore flowered just once before dying. It is now
29/1/09 12:53:19
SpECIES ACCOUnTS
21
known that most orchids are relatively long-lived and the total population, including non-flowering
plants and those dormant underground, is often fairly stable. Fluctuations in the numbers flowering
are related to growing conditions both in the current year and in the previous growing season (which
may be either the previous summer or the previous winter, depending on the species). Growing
conditions are, in turn, usually related to rainfall. Wet weather is conducive to growth but prolonged
dry spells can be very bad for orchids and in some cases can severely restrict flowering.
range
Details of the range is given for Britain and Ireland. A problem faced by all biologists (and many other
people as well) is how to sort and classify records. County boundaries have changed several times over
the years and it becomes very difficult to keep track; was a plant recorded in the Lincolnshire of the
19th century in the same region as one recorded in Humberside or North Lincolnshire today? This
problem is particularly acute in Wales and Scotland where there have been radical changes, and some
of the modern administrative units are very large.
Wisely, most botanical recording uses the system of Watsonian vice-counties. This was devised
in 1852 by Hewett Cottrell Watson, who divided Britain into 112 similarly-sized areas. He followed
traditional county boundaries where he could and divided larger counties such as Yorkshire into
smaller units. The value of the vice-county system is its stability, allowing past and present to
be compared. In the majority of cases we have given records by vice-county, although we have
amalgamated some (e.g. we often refer to Norfolk rather than West Norfolk and East Norfolk) and
we have amended some vice-county names to become more recognisable (for example, West and
East Ross becomes Ross & Cromarty; Westerness and Easterness becomes Inverness-shire). See
p.418 for full details.
We have used two main sources for distributional information: The New Atlas of the British and
Irish Flora (Preston et al. 2002) and the Vice-county Census Catalogue (Stace et al. 2003). We have
also consulted a variety of county floras. These sources do not always agree, especially regarding the
validity of older records and the origin of out-of-range species.
Orchid seeds have evolved for wind or water dispersal and can, in some circumstances, travel
long distances. Nevertheless, a great deal of scepticism attaches to records of orchids away from
their normal range. In some quarters, these are inevitably attributed to deliberate introductions, even
where there is no evidence whatsoever for this (for example, the two tongue orchids). On the other
hand, a non-natural origin has either been proven or is very strongly suggested for some controversial
records (see Other species p. 416). It seems best to keep an open mind but our inclination is to give
such records the benefit of the doubt.
Under the subheading World range, details of the orchids distribution outside Britain and
Ireland is given. We have used several sources for this, including Brown (2003), Davies et al. (1983),
Delforge (1995, 2001) and Hultn & Fries (1986). Again, these sources do not always agree and we
have tried to present the best and most likely compromise.
Accompanying the text is a range map, where the distribution is given by 10km squares in three
data classes: 1987-1999, 1970-1986 and pre 1970. These maps have been supplied by the Biological
Records Centre.
Key to range maps:
Native Distribution
1987-99
1970-86
pre 1970
description
A detailed description of the orchid is given, taken in many cases from the living plant. We have
depended on the literature for some details, especially measurements. It is worth remembering that
many orchids can be taller than the range quoted; even in our limited experience we have found plants
that are bigger than the books suggest.
29/1/09 12:53:19
22
subspecies
Some orchids show definite patterns of variation which may be either geographical or ecological;
plants in a particular region or a particular habitat may differ consistently in appearance from other
areas. In these cases we have recognised different subspecies. There is much disagreement about
which subspecies are worthy of recognition and we have not followed any one authority in this.
Variation and varieties

We give brief details of the normal range of variation which can be encountered. We also give details
of some of the varieties. Like all organisms, orchids are subject to random mutations of their DNA
which may produce a variety of aberrant and deformed plants, and, like stamp collectors (and the
butterfly collectors of old), orchidologists have traditionally been fascinated by these abnormalities.
We do not share that obsession and consider that the odd plant that produces flowers that are deformed
or unusually coloured in some way to be of little special interest. However, some deformities shed
light on more primitive stages in the orchids evolution. In addition, species such as Bee Orchid throw
up the same mutation again and again in widely separated localities, and in such cases these varieties
(var. for short) are often given names. Unlike subspecies, varieties do not show definite geographical
or ecological patterns and can pop up anywhere. In most cases we have followed Ettlinger (1997) in
the use of varietal names.
hybrids
Hybrids have, like varieties, long fascinated orchidologists and in some cases are either very attractive
or throw light on the relationships between orchids. Hybrids between species in the same genus tend
to occur much more frequently than hybrids between species in different genera and, for example,
the frequency with which Frog Orchid hybridised with the spotted and marsh orchids was a longstanding clue to its true relationships. On the other hand, many hybrids are undistinguished and
their true parentage, and even their status as a hybrid, may be the subject of guesswork, sometimes
highly ambitious guesswork. All too often, plants are diagnosed as hybrids when they are merely
aberrant individuals (or even within the range of normal variation). We have usually only included
hybrids that are listed in Stace (2004) and have given the names used in that work.
Name and classification

The origin of the scientific name is discussed under this heading. In some cases the derivation is
clear but in others there is disagreement (or the stated origin seems highly unlikely). We have used
Delforge (1995), Gledhill (2002) and Socit Franaise dOrchidophilie (1998) as our main sources.
Notes are also given on the classification of the species, including any recent changes, and on its
relationships (see also the orchid family tree on p.417).
history and conservation

The date and, where known, the location of the first record for Britain and Ireland is given (largely
from Clarke 1900). For the rarer species we have also given extensive details of their past and current
distribution.
The conservation status of each species is mentioned. Nationally Scarce denotes that the species
is included in Scarce Plants in Britain (Stewart et al. 1994) which indicated that it has been recorded
from 16 to 100 10km squares in Britain from 1970 onwards (the project did not include Ireland).
Rarer species were treated in the British Red Data Book (Wiggington 1999), which includes all plants
recorded from 15 or fewer 10km squares in the period from 1987 onwards. Each species is assigned
to a threat category (Endangered, Vulnerable etc.).
We have extracted data from the New Atlas (Preston et al. 2002) to draw up a table for each species
indicating the extent of its historic and current range and the amount by which it has declined. This
information is put in context by a discussion of the possible reasons for decline (all orchids have
declined), and current threats and conservation measures.
29/1/09 12:53:19
GenUs CYPRIPEDIUM
Ladys-sLippers
001 systematic.indd 23
29/1/09 11:59:25
24
GENUS CYPRIPEDIUM
Ladys-slippers are the most primitive orchids in Europe, differing markedly in their
floral structures from the other species, and have a cunning method of trapping insect
pollinators.
Distribution
The genus Cypripedium contains 45 species
distributed in Europe, temperate Asia and
North and Central America. Around 30 species
are found in China and 12 in North America
but just three extend to Europe and only one
occurs in Britain.
Classification
The ladys-slippers belong to the subfamily
Cypripedioideae and are the only representatives
of that subfamily in Britain and Europe.
Floral structures
Ladys-slippers are distinguished from all other
British and European species by having two
fertile stamens, one on either side of the column,
plus a large, modified, third, sterile stamen
known as the staminode. This is flattened,
tongue-like, often boldly marked and projects
into the large opening of the slipper. The pollen
is granular, sticky and glutinous. There are
three receptive stigmas and the lip is shaped
into a slipper or moccasin-like pouch. The two
lateral sepals are joined for almost their entire
length with just a notch at the tip to suggest
their origin (this structure is called a synsepal).
Members of the other orchid subfamilies have
just one fertile stamen, pollen that is aggregated
together into two pollinia and two stigmas
(although these are often joined together).
Pollination
There is no nectar, and pollinating insects are
tricked into entering the flower through the
large opening on the top of the lip. They can
only leave via the small rear openings, picking
up pollen on the way.
Growth pattern
All ladys-slippers grow from a rhizome that
has slender, fleshy roots. Each year the rhizome
produces two buds at its tip: the larger will

produce the next aerial shoot, the smaller may
form a new branch of the rhizome but usually
eventually dies. The aerial shoot will also die
off at the end of each season, but the rhizome
continues to grow from a bud that forms on
its base (sympodial growth, as opposed to
monopodial growth, in which the main stem
continues to grow indefinitely behind the
same growing point). In the ladys-slipper,
the rhizome grows 5-10mm each year in a
characteristic zig-zag pattern as the position of
the larger bud alternates from left to right. The
rhizome may die off slowly and rot at the rear
while growing at the front, but up to 20 living
segments may be present at any one time
(and by counting the number of segments
its age may be estimated). The rhizome also
branches, roughly every five years, and each
branch may eventually put up an aerial stem.
Thus as a plant ages, the number of flowering
shoots increases.
Fungal partners
Poorly known, and this lack of information is
an obstacle to the conservation of the species.
Vegetative reproduction
The rhizome branches as it grows and
eventually the basal portion dies, leaving the
terminal branches as independent plants, clones
of the mother plant. In at least some species of
Cypripedium, buds can also form on the tips of
the roots as a means of propagation.
Name
The generic name Cypripedium derives from the
Greek Kypris, a name for Aphrodite (goddess
of love), and Latin pes, meaning foot and thus
Aphrodites slipper.
O 31 May, Khabarovsk, eastern Sibera. Ladys-slipper

occurs right across Europe and Asia but is threatened
almost everywhere.
29/1/09 11:59:25
LADYS-SLIPPER
Ladys-sLipper
25
Critically Endangered, BAP
Cypripedium calceolus
This spectacular species is Britains rarest orchid with just one group of plants of native
origin surviving at a closely guarded site in Yorkshire. Originally fairly widespread in the
limestone districts of northern England, its large, showy flowers have been its downfall.
Plants were pillaged from the wild for hundreds of years, either to be dried, pressed and
stored away in dusty herbaria or to be transplanted to gardens.
Identification
Unmistakable.
Similar species
The leaves of non-flowering plants could be
overlooked and resemble the leaves of Lily-ofthe-valley.
Habitat
The surviving native plants are found in speciesrich grassland on a fairly steep, well-drained,
north-facing slope. They nestle in a sheltered

limestone valley that has some trees and lies on
the northern edge of a large block of woodland.
In Europe, Ladys-slipper favours a variety of
woodland, usually on calcareous soils, but in
the northern parts of the range is also found in
the open, in fens and marshy grassland. Overall,
it appears to require relatively well-lit areas
but likes to have its roots in cool, moist soil. In
England suitable areas would have been found
in open woodland on north or northwest facing
slopes or where subsurface flushing kept the
ground damp but not waterlogged; the former
English, sites were in ash, hazel and oak woods
on steep, rocky slopes, always on limestone, at
150-260m above sea level. The correct balance
of sun and shade seems to be critical. Too little
light and the plant fails to flower and may even
become dormant underground, too much
light and rank herbs and shrubs will swamp it,
although in a more natural environment grazing
animals may have suppressed much of the
competing vegetation.
Flowering period
Late May to early or mid-June. Plants are in
flower for two or three weeks and each flower
lasts 11-17 days, withering on the sixth day
after pollination.
Range
M 30 May, Lancashire (Sean Cole). This plant, probably

of garden origin, had been growing in this spot for perhaps
a century but was vandalised a few weeks after this
picture was taken in 2004.
Confined to a single site in Yorkshire,

although formerly more widespread.
Artificially propagated plants have recently
been planted out at several localities within
its former range. World range: Europe and
southern Siberia, extending eastwards to
29/1/09 11:59:26
26
GENUS CYPRIPEDIUM
1987-99
1970-86
pre 1970
northern Mongolia, northeast China, Korea,

the Russian Far East and Sakhalin. It is
essentially a northern or boreal species, and
in Europe the Ladys-slipper ranges north to
northernmost Scandinavia. It is absent from
the Mediterranean lowlands but is found in
the mountains as far south as northern Spain,
central Italy, northern Greece and Bulgaria.
There are also outposts in the Crimea and
Caucasus. It is also absent from the Atlantic
fringes of Europe, for example central and
western France, and the English populations
were therefore always out on a limb.
How to find it
Potential visitors are asked to keep away from
the native site in Yorkshire due to the fragility
of the habitat. Visitors were welcome at a site at
Silverdale in Lancashire where one can admire
a plant, albeit probably of European origin, in
a natural setting (it has hopefully survived the
attack by vandals of 2004). Otherwise, we must
hope that the reintroduction programme is a
success and that self-sustaining populations of
flowering plants will once more grace the dales
of northern England. At present, just one of
the reintroduction sites has public access
Ingleton Glen in North Yorkshire but
the young Ladys-slipper plants have not yet
flowered there.
DESCRIPTION
Height: 15-70cm but usually around 30cm and
rarely more than 60cm.
Stem: Glandular-hairy with three to four green
or brown sheaths at the base. The species forms
clumps of shoots, sometimes quite large, which
may belong to one or more plants.
Leaves: Green and oval, elongated into a
pointed tip; the three to four (sometimes five)
leaves are arranged alternately up the stem.
They are sparsely hairy (especially on the
underside), ciliate along the margins, wavy
edged and very prominently veined.
Spike: Each stem usually produces one or two
flowers, very rarely three.
Bract: Leaf-like, longer than the flower and
held erect behind it.
Ovary: Long, slender, six-ribbed, curved (but not
twisted), with glandular hairs and a short stalk.
Flower: Large and conspicuous. The sepals
are purplish-brown or claret with wavy edges,
downy on their inner surface and hairy at
the base. The upper sepal is lanceolate with
a pointed tip and is held erect, whereas the
two lateral sepals are fused and hang vertically
below the lip (their tips forming two small teeth
at the tip of the combined synsepal). The petals
are purplish-brown, mottled with olive-yellow
towards the base and with downy midribs and
long hairs at the base. They are strap-shaped
with pointed tips, variably twisted (through up
to 360) and hang at four and eight o-clock
on either side of the lip. The lip is yellow and
looks like a bag or clog (the slipper). There is
a large entrance on the upper side towards the
rear and two small openings on either side of
the column at the base. The edges of the large
upper opening are rolled down and under, and
the interior of the slipper is covered in sticky
hairs with lines of reddish dots along its floor.
The column projects forwards into the slipper
and is divided into two parts: the staminode,
which is yellowish-white variably marked with
red spots and very conspicuous, and the large
P Yorkshire (Peter Wakely, English Nature).
29/1/09 11:59:27
29/1/09 11:59:36
28
GENUS CYPRIPEDIUM
fleshy stigma, which lies on the lower part of

the column hidden inside the slipper. The two
remaining stamens lie on either side of the base
of the staminode, adjacent to the two small
rear openings into the slipper. The flowers are
delicately scented, and the scent is said to be
sweet, recalling oranges.
Subspecies
None in Britain.

None in Britain.
BIOLOGY
Pollination and reproduction
It is thought that Ladys-slipper is pollinated
by small bees, especially bees of the genus
Andrena. The flowers do not produce nectar,
however, and there is debate as to what it is
that attracts the insects. A strong contender is
the flowers scent, which comprises a complex
mixture of chemicals that may mimic the
bees pheromones, chemical signals that are
associated with feeding and mating behaviours.
Other possible attractions, which could
indeed combine with the scent lure, include
the red spots on the staminode and floor of
the slipper that may act as false nectar guides.
Alternatively, the bees may actually receive a
reward for their visit, perhaps the oil secreted
by the small hairs inside the slipper. Bees may
also find shelter in the slipper in cold weather
or overnight.
Whatever the attraction, bees land on the
edge of the large opening or try to land on
the staminode and fall into the slipper. After
a few minutes the bee tries to leave. However,
the sides of the slipper are very smooth and
slippery and the rim of the large opening curls
over and inwards, making escape via this route
impossible. The bee can only leave through the
small openings on either side of the column
where there are small stiff hairs to give it a
foothold. The openings are only just big enough
for the bee, which is forced to make contact
with one of the stamens as it makes its escape,
picking up a load of pollen in the process. The

bee goes on to visit another flower; when it
eventually leaves this its back rubs against the
stigma, which projects down into the slipper,
and pollen from the first flower is deposited
there; the surface of the stigma has minute, stiff,
pointed papillae that act as a brush to remove
pollen from the bees back. As it escapes, more
pollen is carried away, ready to be deposited on
the next flower and continue the process.
The mechanism is precise and in order to
effect pollination the bee has to be a specific
species that is the right size; bees that are
too large or too small can escape without
pollinating the flower. A wide variety of other
insects also enters the slipper but these too
are the wrong size and shape and either leave
unharmed or may be trapped and die. Selfpollination is unlikely; the bee would have to
reverse back into the flower just as it was on the
point of escape. In addition, it seems that the
flowers are, to a great extent, self-sterile.
The pollination strategy is not efficient
and seed set is rather poor with few fertile
capsules being produced. Bees are attracted
to large groups of flowers, especially those
in sunlight, but even in large populations in
Europe an average of just 10% of flowers set
seed. Nevertheless, each capsule contains 6,00017,000 seeds which may be dispersed by rain
as the seedpods seem to close up when dry and
open when wet.
Ladys-slipper also reproduces vegetatively
through division of the branching rhizome,
and in many populations in Europe this is
thought to be more important than seed in the
recruitment of new plants to the population.
Development and growth

The Ladys-slipper grows from a slender,
creeping, branched rhizome. Each branch of
the rhizome may eventually put up an aerial
stem, and so as a plant ages the number of
flowering shoots increases. Alternatively,
a clump of shoots may arise from several
separate clones, each produced by vegetative
reproduction, or by the development of
29/1/09 11:59:37
LADYS-SLIPPER
29
M 6 June, Lancashire.The large opening of the flower, with the tongue-like staminode to the rear, is obvious.
29/1/09 11:59:39
30
GENUS CYPRIPEDIUM
seedlings at the base of the mother plant.

The first green leaves are reported to appear
in the fourth year after germination by some
authors and in the first year by others. The
immature plants have a slender stem with one
or two small leaves and may remain in this state
for several years. In England a seedling has been
noted to flower nine years after it first appeared
above ground, and in Europe the young plant
takes six to ten years to produce flowers. Plants
are long-lived and many are over 30 years old,
with some over 100 years. Indeed, a lifespan
of 192 years has been determined from the
examination of the growth of a single rhizome
in Estonia.
Hybrids
None in Britain.

The specific name calceolus means little shoe
or little slipper and like the English name
refers to the slipper-like appearance of the
lip. An old name for the species was Calceolus
Mariae or Marys Shoe. Yellow Ladys-slipper
C. parviflorum of North America is very closely
related and is sometimes amalgamated with
this species.
HISTORY AND
CONSERVATION
A Red Data Book species that is classed as
Critically Endangered and fully protected
under Schedule 8 of the Wildlife and
Countryside Act 1981. The species is also rare
and threatened throughout much of its range
in Europe and Asia.
The first British record dates from 1629
when John Parkinson, a London apothecary,
recorded the species in his Paradisi in Sole;
Paradisus Terrestris (Park-in-suns Earthly
Paradise): In a wood called the Helkes in
Lancashire neere the border of Yorkeshire.
This was the same wood that supplied the first
British record of Sword-leaved Helleborine in
1666. Both species are now gone from there.
The Ladys-slipper was subsequently found
widely but locally in the limestone districts
Past and present occurrence of Ladys-slipper in Britain

and Ireland (based on presence or absence in 10km
squares of the National Grid; data from the New Atlas).
total historical range,

1500-1999
current range
% lost, 1500-1969
Britain
Ireland
22
1 (0.04%*)
95%
% lost, 1970-1986
0%
% lost, total
95%
* current range as a % of the total number of 10km squares
of northern England. There was a single site

in Derbyshire at the Heights of Abraham
overlooking Matlock, and it was recorded in
southern Cumbria around Whitbarrow and
Scout Scar (northwest of Levens). The bulk of
the records came, however, from three areas:
first, West Yorkshire around Ingleborough
(including Helkes Wood) and the Upper
Wharfedale region around Litton, Kettlewell
and Grassington; second, North Yorkshire
on the southern flanks of the Cleveland Hills,
especially the valley of the River Rye and its
tributaries north and west of Helmsley; third,
Castle Eden Dene in Co. Durham.
Such a large, conspicuous and attractive
flower as the Ladys-slipper was an obvious
subject for curiosity and avarice, and plants
were picked or dug up from at least the 16th
century onwards. In the late 18th and early
19th centuries, they were ruthlessly stripped
from the wild, both for horticulture and as
herbarium specimens. By the mid-19th century
the species was rare and in 1917 the Ladysslipper was declared extinct in Britain.
In 1930, Ladys-slipper was resurrected
from the dead when a single plant was found
in a remote Yorkshire dale (this is the plant
that has survived to the present day). However,
the last note of its rediscovery in print was
in 1937, and the species quietly slipped from
the botanical worlds attention. As well as the
site being a closely-guarded secret, the Ladysslipper itself was being very coy. From 14 stems
and just one flower in 1930 it dwindled to two
29/1/09 11:59:40
LADYS-SLIPPER
to five stems by the late 1940s and 1950s and

hardly ever bothered to flower; a single bloom
was produced in 1934 and 1943 but not again
until 1959.
Despite the secrecy, in the 1960s word
started to get out and the Ladys-slipper
again faced the old threat from collectors
and a new threat from visiting botanists with
big feet and heavy cameras. Indeed, the site
was raided and half the plant was removed.
Various individuals and groups attempted to
protect the orchid and the ensuing conflict
and confusion prompted the late Edgar MilneRedhead, President of the Botanical Society
of the British Isles, to set up the Cypripedium
Committee in 1970. The committee included
representatives of various conservation and
botanical interests with the aim of coordinating
the orchids conservation. The first priority
was to safeguard the sole remaining wild plant,
and this has been guarded every year since
then. Visitors are asked to keep away, and
even the committee has only rarely visited the
site, whilst management visits have also been
curtailed for fear of damaging seedlings.
With careful protection and habitat
management the Ladys-slipper has slowly
increased in vigour at the wild site, with a
steady increase in the number of shoots and
flowers. In 1996, the peak year to date, there
were 65 shoots and 23 flowers on the main
clump (which may be just one plant, several
clones or even include seedlings). Few or
no flowers were being pollinated naturally,
however, and hand-pollination began in 1970.
This has resulted in good seed set and the
production of many capsules, some of which
are left to mature on the plant while others are
sent to Kew Gardens.
As part of the Species Recovery Programme,
organised by English Nature, and with the goal
of establishing self-sustaining populations of
Ladys-slipper in the wild, ex-situ propagation
began in 1983, when a donation from Sir
Robert and Lady Sainsbury established the
Sainsbury Orchid Conservation Project at the
Royal Botanic Gardens, Kew. The fungus that
31
aids the Ladys-slipper seeds in germination and

growth could not be identified, however, and
this initially thwarted efforts to cultivate plants
from seed. But, after much trial and error, a
method of germinating seed in the absence of
fungi was developed. This involves supplying
the nutrients directly to the seedling in a sterile,
asymbiotic medium. Although only about 10%
of seeds germinate, large numbers of seedlings
can now be produced from the hand-pollinated
capsules of the wild plant.
The major problem has become the
development of techniques to introduce these
laboratory-grown seedlings into a natural
environment. The first six seedlings were
planted out at the native site in Yorkshire in
autumn 1989, and by 2003 approximately
2,000 seedlings had been introduced into
23 locations (although some of the sites are
very close to each other). Survival rates are
not high, and slugs and snails are a particular
problem. Up to 2004, just 105 plants survived
at ten sites. Two have flowered, the first in the
summer of 2000, 11 years after being planted
out. Early reintroductions used two-year old
seedlings (one year in flask and another in
compost), and survival rates were particularly
poor as the tiny seedlings fell foul of slugs or
bacterial and fungal infections. Fewer older
seedlings are now used (three or even more
years old). These are being planted in drier and
more open micro-habitats and appear to have
much better prospects for survival.
A single plant has been present at Silverdale
in Lancashire for many years, although it is
thought that it was planted there in the late
19th or early 20th century; its DNA suggests
that it is from either Austria or possibly the
Pyrenees. It did not flower for many years
but slowly increased in vigour and by 2004
produced nine flowers. Sadly, later in the 2004
season this plant was vandalised and probably
partially removed although fortunately it has
survived this.
29/1/09 11:59:40
GENUS
CEPHALANTHERA
HELLEBORINES (I)
12/2/09 16:51:29
GENUS CEPHALANTHERA
33
A genus of relatively primitive woodland orchids that are probably heavily dependent on
fungi throughout their lives. Their flowers look very simple but are surprisingly similar to
those of the apparently more complex and highly-evolved Epipactis helleborines.
Distribution
Around 15 species are found in Europe,
North Africa and temperate Asia with a single
representative in western North America. Eight
species are found in Europe, with three in Britain.
Classification
Considered a relatively primitive genus due
to the rather unspecialised reproductive
structures. Closely related to the helleborines of
the genus Epipactis, but the flowers are stemless
(sessile), the ovary is often twisted and the
column is simpler in structure.
Floral structures
upper
sepal
anther
column
petal
pseudopollen
ovary
ovary
lip
Pollination
White Helleborine is largely self-pollinated but
Red and Sword-leaved Helleborines are crosspollinated. Their flowers do not produce nectar,
however, and depend upon deceit, displaying
bright yellow pseudopollen on the lip to entice
potential pollinators to visit. The mechanism is
rather simple: visiting insects brush against the
stigma and pick up some of the sticky substance
that it exudes, then, as the insects back out of
the flower, the pollinia stick to this sticky mess
and are carried away.
Growth pattern
The aerial stems grow from rhizomes that put
out slender, fleshy roots.
Fungal partners
lateral sepal
upper sepal
pollinia. As in the Epipactis helleborines, the lip

is divided by a narrow waist into the hypochile
at the base and epichile at the tip.
lateral
sepal
petals
They are probably heavily dependent on fungi,

and recent studies have shown that adult plants
of both Red and White Helleborines do acquire
nutrients via a fungal route and also have an
association with ectomycorrhizal fungi; they
may gain nutrients from the roots of nearby
trees via these fungi. One member of the genus,
the Phantom Orchid C. austiniae of western
North America, produces no green leaves and is
fully dependent on fungi.
Development from seed

bract
lateral sepals
The column has a simple, sticky stigma and no

rostellum or viscidium. The pollen is aggregated
together into pollinia but as single grains
that form a powdery mass; in more advanced
orchids the pollen is grouped into tetrads that
are held together by elastic threads, and it is
these tetrads that are bound together in the
O 26 May, Cambridgeshire.White Helleborine
Poorly known, and seedlings are rarely found.
White Helleborine may produce new shoots
from buds on the roots, but Sword-leaved
Helleborine is thought to lack any method of
vegetative propagation.
Name
The name Cephalanthera means floweryheaded and derives from the Greek kephale
head and antheros flowery or blooming.
29/1/09 11:59:42
34
GENUS CEPHALANTHERA
red HeLLeborine
Critically Endangered, BAP
Cephalanthera rubra
With flowers that are closer to pink than red, this is one of the most striking and attractive
of all British and Irish orchids. Sadly, it is also one of the rarest, with just a handful of plants
known at three sites in southern England. Red Helleborine has always been rare in Britain
but even in its traditional stronghold in the Cotswolds it has now declined to just a single
site. On the edge of its range in Britain, it is capricious and seems to require exactly the right
habitat with exactly the right mix of light and shade, but in Europe it is much less picky.
Identification
This species is so rare that identification is likely
to be academic. If you are fortunate enough to
be looking at a Red Helleborine you will have
travelled specifically to see it. It is, however,
possible that new sites may still be found in
or around suitable woodland in southern
England. Flowering plants are unmistakable,
but non-flowering plants are very hard to find
and difficult to distinguish with certainty from
White Helleborine or even indeed from the
Epipactis helleborines.
Habitat
Found in beechwoods growing either on
chalk, as in the Chilterns and Hampshire, or
on limestone, as in the Cotswolds, usually on
free-draining slopes. At all three current sites it
grows at the boundary of the chalk or limestone
and the overlying acidic clay drift. All three sites
are probably ancient woodland.
Although a woodland orchid, too much
shade will prevent it from flowering regularly
or successfully. It has long been known that
the finest specimens were to be found in rather
open spots within the woods, sometimes
amongst tall grass, brambles and other
undergrowth alongside paths or in scrubby
places and on open banks. In such open
situations, however, the growth of scrub can
eventually overwhelm the helleborines. It may
well be adapted to flower in the gaps caused
by tree-falls, where a sudden increase in light
allows it to flower for a year or two before the
canopy closes again, light levels fall and the
plant retreats to a dormant state underground
M 24 June, Buckinghamshire. The plant appears spindly

and fragile, almost top-heavy, and seemingly struggles to
support such a large, elegant flower spike.
29/1/09 11:59:43
RED HELLEBORINE
35
until conditions become favourable again.

Alternatively, grazing animals in the primeval
forests may have kept the development of rank
vegetation and scrub at bay for long periods
of time. In France and Germany, where Red
Helleborine is locally common, it thrives best in
situations where it receives a few hours of direct
sunlight each day, in glades or along forest roads
and tracks.
Flowering period
Mid-June to mid-July, sometimes late July, but
usually at its best in the last week of June and
first week of July.
Range
Currently known from single sites in north
Hampshire, Buckinghamshire and east
Gloucestershire. World range: Essentially a
European species, ranging north to c. 60N in
southern Scandinavia, southernmost Finland
and the Baltic States, east to the Urals and
south to the Mediterranean, including Corsica,
Sardinia, the Balearic Islands, Crete and
Cyprus, and also to the Crimea and Caucasus.
Found in North Africa in Morocco, Algeria
and Tunisia and in Turkey, Israel, Syria and
northern Iran. It appears to prefer a continental
climate and is largely absent from the Low
Countries, the milder western seaboard of
France and, of course, Britain.
1987-99
1970-86
pre 1970
M 28 June, Buckinghamshire. Confined to beechwoods,

Red Helleborine acquires nutrients from its fungal
partner throughout its life.
How to find it
Red Helleborine is a shy flowerer, and even
when it flowers, there may be many more
non-flowering plants in the vicinity, and it can
remain in a vegetative state for many years. It
is nevertheless worth looking for, especially
in beechwoods on chalk or limestone, where
recent felling or tree-falls have let in more
light. But realistically, the only chance to see
this species is at one of the known sites. The
populations are very small and vulnerable and
although visitors are not encouraged, low-key
open days are organised at the Hampshire
site during the flowering season (contact the
Hampshire Wildlife Trust for details).
DESCRIPTION
Height: 15-60cm.
Stem: Slender, often wavy and with abundant
short glandular hairs on the upper portion. The
stem is dusky green, variably washed brownishpurple towards the tip and sometimes with
violet sheaths at the base.
29/1/09 11:59:44
29/1/09 11:59:48
RED HELLEBORINE
Leaves: Dusky grey-green, held in two ranks

alternately up the stem at about 45. Most of
the five to eight leaves are long, narrow and
lanceolate, but the lower ones are shorter,
blunter and duller green.
Spike: Rather open, with two to nine flowers
(occasionally as many as 17). The ovaries are
held more-or-less erect, and the flowers face
upwards and outwards. The unopened
buds are purplish-pink, becoming creamy at
the base.
Bract: Dusky green, with numerous short
glandular hairs, especially towards the base.
The bracts are very narrow and pointed, the
lower about one-and-a-half times the length
of the ovary, the upper roughly equal in length.
Ovary: Dusky green, becoming brownishpurple at the base and with the ribs variably
washed purplish. Slim, cylindrical, ribbed
and twisted, the ovary has numerous
glandular hairs.
Flower: A beautiful shade of pink, the flowers
open widely. The sepals are elongated ovals,
narrowing both towards the pointed tips and
towards the base. They are lilac-pink, becoming
whiter around the base, with numerous short
glandular hairs on their outer surface. The
petals are similar, although shorter, broader
and less tapering at the base. When fully
open the lateral sepals are held horizontally
like outstretched arms, while the upper sepal
M 28 June, Buckinghamshire.The lip has a series of parallel raised ridges, probably related to the attraction of
pollinating insects.
O 24 June, Buckinghamshire.
37
and petals form a rather loose hood. The lip

is divided into hypochile and epichile by a
constriction around the mid-point. The basal
portion (hypochile) is gutter-shaped and the
sides curl up on either side of the purplish
column and anther; it is white with fine yellow
veins and pale-pink sides. The epichile is flatter
(although still concave), arrow-shaped and
tapers to a pointed tip that is bent downwards.
It is whitish with variably pinker edges, a deep
lilac-pink tip and seven to nine longitudinal
yellow ridges. The column is violet-rose.
Subspecies
None.

None.
BIOLOGY
Red Helleborine is pollinated by small solitary
bees, with members of the genus Chelostoma
being important, at least in Europe. The flowers
do not produce nectar, but studies in Sweden
suggest that the bees are attracted to the
helleborines because to the bees eyes (which are
not sensitive to the red end of the spectrum)
they resemble certain blue bellflowers
Campanula sp. that do produce nectar. And,
not only do bees visit the bellflowers to collect
nectar, but also male bees search for females
around the flowers. The Red Helleborine may
therefore be able to exploit the bees sexual
urges as well as its foraging behaviour. The
pollination mechanism is, however, not efficient
in England, perhaps due to an absence of bees
of the right size and shape, and few flowers set
seed. Many flowers remain unfertilised and fall
off the plant complete with their ovaries.
Vegetative reproduction may be more
important than reproduction by seed. If the
central rhizome dies off, the short side roots,
densely infected with fungi, can remain alive
and produce a bud at the tip that will grow into
a new rhizome and eventually produce a new
leafy shoot.
29/1/09 11:59:50
38
GENUS CEPHALANTHERA
O 28 June, Buckinghamshire. The lateral sepals

are spread wide.

The aerial stems grow singly from a slender,
horizontal rhizome. There is a heavy presence
of fungi in the roots, especially the short side
roots. Recent isotope studies have confirmed
that mature plants acquire nitrogen and carbon
via their fungal partner, gaining around 60%
of their nitrogen and about 25% of their
carbon in this way. Unexpectedly, the Red
Helleborine has a particular association with
ectomycorrhizal fungi and thus, like Coralroot
and Birds-nest Orchids, may gain nutrients
via these fungi from the roots of nearby trees,
especially Beech. But unlike the tree, it cheats
the fungus by giving nothing in return.
Red Helleborine may become dormant
underground, and gaps of up to four years
between appearances have been recorded
from Hampshire (although it is possible that
shoots did emerge but were quickly grazed off ).
Indeed, it may be able to persist underground
for much longer periods. After some trees were
felled at the Hampshire site, a plant appeared
in 2003 that had not been recorded in any
of the previous 17 years. With 14 flowers, it
was one of the most robust plants ever seen
at this site, suggesting that it was not a newly
emerged seedling. In addition, Red Helleborine
may be relatively long-lived, and again at the
Hampshire site the plant first discovered in

bloom in 1986 survived for at least another 18
years and flowered in all but four seasons.
There is no information on germination or
early development. It has been stated that the
first leaves are not produced until about six
years after germination, and flowering does not
occur until the plant is ten years of age. But, as
with all such reports, the actual time-scale may
be much shorter.
Hybrids
None.

The specific name rubra means red.
HISTORY AND
CONSERVATION
A Red Data Book species that is classed as
Critically Endangered and fully protected under
Schedule 8 of the Wildlife and Countryside
Act 1981. The population is tiny, and in 2004
there were just over 30 shoots at three sites,
producing ten flower spikes.
The first British record was published in
1797 in English Botany: Gathered last June
on Hampton Common, Gloucestershire,
by Mrs. Smith, of Barnham House in that
neighbourhood. The Cotswolds were for a long
29/1/09 11:59:51
RED HELLEBORINE
time the stronghold of the species with records

from a great many of the Beech woods from
Nailsworth to Birdlip, but it has seldom been
seen in quantity and flowering is extremely
erratic (Lousley 1969). It was common enough
at times, however: I once saw some fifty or sixty
plants together, but only about ten bore spikes
of flowers, and somebody cut those before the
next morning (Riddelsdell et al. 1948). There
are also old records from south Somerset, west
Gloucestershire, West Sussex and Kent, but
no specimens have survived from the latter
two counties to confirm the identifications
and although this is a very distinctive plant the
reliability of these reports is uncertain.
Following its discovery, the number of
sites in the Cotswolds gradually dwindled
and there is now just one (although it is quite
possible that the plant is lurking unseen at
others). At this beechwood reserve there were
up to 40 shoots in the 1970s, but by the late
1990s, probably due to increased shade, it had
declined to only three shoots with just one of
these flowering. Happily, there has since been
a partial recovery to around ten shoots with at
least one flowering each year. This may have
been encouraged by some limited felling to let
in more light.
Red Helleborine was found in the Chilterns
in Buckinghamshire in 1955 when three
flowering spikes were discovered in a clearing
in a beechwood. There were ten the following
year, together with 64 non-flowering plants
within about 25m. After this spectacular
discovery, however, there was a sharp decline,
and no flowers were recorded from 1960.
Indeed, no plants were seen at all for several
years in the 1970s. But from 1980 onwards odd
non-flowering plants reappeared and flowering
resumed in 1983. There was a distinct increase
from 1989 (probably encouraged by the felling
of a handful of trees in 1987), and since then
about 10-14 plants have appeared each year, a
little over half of them producing flowers.
Red Helleborine was recorded in
Hampshire in 1926 and then refound in north
Hampshire in 1986 when a single flower spike
39
was located. It is thought that this plant may

have been stimulated into flowering by the
felling of a few trees some years before; the
wood where it grows had been heavily shaded
by Beech and Yew. Subsequently the whole
area was thinned and then clear-felled, mown
and hand-weeded. Wire cages have been used
to protect the plants, but despite this the
flower spikes have been damaged by careless
photographers on three occasions. And, despite
the careful management, there has been just
a modest increase with one to eight plants
appearing and producing one or two flower
spikes almost every year.
Red Helleborine is clearly sensitive to light
levels. Another factor that affects the plants is
grazing: deer, rabbits and slugs undoubtedly
damage the helleborines and prevent flowering.
Slug pellets are used in the Cotswolds for this
reason, although their effect on the fungal
partner is unknown. Conversely, grazing
also keeps down the surrounding vegetation
and reduces competition, and the scratching
and scraping of rabbits may provide suitable
conditions for the establishment of new plants.
Thus in the Cotswolds the management regime
now includes some limited raking and also
fencing the area against deer over the summer
and then removing the fences in an attempt to
get the best of both worlds.
British populations of Red Helleborine
produce few young plants and seed production
is low. An individual plant in Hampshire
produced 96 flowers over 14 seasons but just
nine capsules were recorded, despite some
flowers being hand-pollinated in nine of those
years. In the Chilterns natural pollination levels
are also low, and seed is only occasionally set;
hand-pollination was also undertaken for a
while but analyses of the seeds showed that
only around 20% were viable.
The capricious flowering and poor seedset of Red Helleborine in England recalls
Sword-leaved Helleborine, a species that is
also pollinated by small solitary bees. Both
helleborines will flower once a certain level
of light is achieved, but much brighter, sunlit
29/1/09 11:59:52
40
GENUS CEPHALANTHERA
conditions with a variety of genuinely nectarproducing flowers in the vicinity are probably
required if they are to be visited by the correct
pollinators and set seed. The spectacle of
large numbers of flowering plants may also
boost pollination rates and the tiny British
populations of Red Helleborine may be
especially handicapped if this is true.
With so few plants, management of the three
populations is necessarily tentative. The most
effective techniques may be to mimic natural
systems: limited felling to produce a mosaic of
woodland and glades; winter grazing to suppress
rank vegetation; and protection from deer and
rabbits when flowering and fruiting.
Past and present occurrence of Red Helleborine in Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
10
3 (0.1%*)
% lost, 1500-1969
50%
% lost, 1970-1986
20%
% lost, total
70%
O 28 June, Buckinghamshire. As in all helleborines,

the lip is divided into two
about half way along its
length.
29/1/09 11:59:54
SWORD-LEAVED HELLEBORINE
41
sword-Leaved HeLLeborine
Cephalanthera longifolia
Vulnerable, BAP
Other names: Narrow-leaved Helleborine

This is among the most striking and attractive orchids, but sadly also one of the most
threatened. Sword-leaved Helleborine occurs very widely in Britain and Ireland but has
suffered a dramatic decline over the last 200 years, and sites for the species are now few
and far between. It has only been recorded recently from around a quarter of its historical
range, and many populations are so small that they are in imminent danger of extinction.
Fortunately, our understanding of the biology of this species has improved, and with
appropriate management some sites have made a spectacular recovery.
Identification
A distinctive orchid, with graceful, gently
arching foliage, long leaves alternating all the
way up the stem and spires of pure white
flowers that open just enough to display a large
golden-yellow patch on the lip.
Similar species
White Helleborine is rather similar and
sometimes found growing together with
Sword-leaved Helleborine in southern England.
It has duller, creamier flowers, blunter sepals,
and its leaves, especially the lower ones, are
on average shorter and broader. The best
distinction is the length of the bracts, which are
longer than the ovary in all White Helleborine
flowers, even at the top of the spike. On Swordleaved Helleborine the bracts are shorter than
the ovary, at least in the upper part of the spike
(they may be very long and leaf-like on the
lowest two or three flowers), and therefore the
flower spike is well-demarcated from the leafy
part of the stem.
Habitat
Sword-leaved Helleborine is a woodland
orchid but its optimum habitat is the interface
of woodland and grassland. It does best in
glades, clearings, rides and on the margins of
roads and tracks, in areas where the ground
vegetation is not too dense. It depends on small
solitary bees for pollination and these are found
in species-rich grassland, visiting the sunnier
parts of adjacent woodland to forage. Suitable
M 27 May, Hampshire. The stately, leafy stems, clearly

distinct from the leafless flower spike, are characteristic.
29/1/09 11:59:55
42
GENUS CEPHALANTHERA
sunlit, open conditions are naturally transient

in woods as open areas are invaded by scrub
or the canopy closes overhead, increasing the
level of shade. Sword-leaved Helleborine will
persist and can flower well under the shade
of a high, closed canopy but in the absence
of successful pollination and reproduction,
will eventually disappear from such sites. It
can also persist in dense shade under scrub,
but flowering will be very much reduced, the
plants remaining in a vegetative state or even
becoming dormant underground. However, it
can flower spectacularly if the scrub is cleared
and the light intensity rises above a certain level.
Notably, in Hampshire, rather than being in
ancient woodland many of its sites are along
ancient trackways or in secondary woods that
were arable fields in the 19th century but were
allowed to tumble down to woodland.
In southern England, Sword-leaved
Helleborine favours beechwoods on chalk, but
elsewhere it grows under a variety of deciduous
trees. In the Wyre Forest in Worcestershire,
it once thrived in coppice cut on a relatively
long, 18-year cycle. It is sometimes found in
scrub and occasionally on chalk grassland, but

the plants are very small on grassland with few
flowers. On the coast of Co. Mayo in western
Ireland, it once grew on wind-blown shell-sand
overlying peat, both in stunted hazel and oak
scrub and on an exposed summit (where the
plants were even smaller, just 5-7.5cm high
with only one or two flowers). It has also been
found in pine plantations at Newborough
Warren on Anglesey.
Many sites for Sword-leaved Helleborine
are on calcareous soils overlying chalk and
limestone, but in the north and west it is found
on a variety of other soils, although probably
still with an alkaline influence. It is tolerant of
both wet and dry conditions, from damp woods
or wet scrub in the north and west to dry chalky
slopes in southern England. Sword-leaved
Helleborine is confined to the lowlands.
Flowering period
Mid-May to mid-June, being at its best in
the last two weeks of May, even in Scotland,
although sometimes still in flower in early July
at exposed sites in the north.
M 27 May, Hampshire. Although sadly much reduced, Sword-leaved Helleborine responds very favourably to appropriate
management, especially good levels of light combined with grazing.
29/1/09 11:59:56
43
Range
Sword-leaved Helleborine has always had
an unusually scattered distribution in the
British Isles but is now very much reduced.
The strongholds are in Hampshire and Argyll,
each with around a dozen sites. Other minor
concentrations are found in Worcestershire (the
Wyre Forest holds a few small populations),
Merionethshire and Cumbria. There are
one or two sites each in West Sussex,
Surrey, west Gloucestershire, Warwickshire,
Montgomeryshire, Caernarvonshire, Anglesey
and, in Scotland, Perthshire, Inverness-shire,
Ross & Cromarty and west Sutherland, and
also Arran, Islay, Jura and Skye in the Inner
Hebrides. In Ireland it is very scattered, with
a handful of sites in Co. Kerry, Co. Clare, Co.
Wexford, Co. Galway, Westmeath and Co.
Donegal. World range: Essentially a European
species but with scattered records to the east,
in northern Iran, the mountains of Central
Asia and through the Himalayas to southwest
China. In Europe it occurs north to 63N in
Scandinavia, southern Finland and the Baltic
States, east to western Russia (around Moscow)
and south to the Mediterranean, including the
Balearic Islands, Corsica, Sardinia and Sicily,
and also to the Crimea and Caucasus. It is
found in North Africa in Morocco, Algeria
and Tunisia, in a few places in Turkey and in
Lebanon and Israel.
M 27 May, Hampshire. The lower flowers can have very

long bracts, but they are always very short on the upper
flowers, a clear distinction from White Helleborine.
How to find it
1987-99
1970-86
pre 1970
The Hampshire Wildlife Trust reserve at

Chappetts Copse holds by far the largest British
population (2,185 flowering plants in 2003)
and is undoubtedly the best place in southern
England to look for this stunning orchid. In
Scotland it can be found at Knapdale and at
Ballachuan Hazelwood (both Argyll & Bute).
DESCRIPTION
Height: 15-65cm but can be as short as 5cm in
very exposed situations.
Stem: Green, with some short hairs on the
upper part and two to four whitish, often greentipped sheaths at the base. Stems grow singly.
Leaves: Clear grass green with fairly prominent
29/1/09 11:59:57
44
GENUS CEPHALANTHERA
29/1/09 12:00:00
veins, strap-shaped, keeled, long and rather

narrow, the upper ones with a pointed tip, the
lowermost shorter, broader and blunter. The
seven to 20 leaves are spaced alternately up
the stem and arranged in two variably defined
opposite rows that may twist around the stem in
a spiral pattern. They are held at 30-60 above
the horizontal and arch gracefully outwards.
Spike: Rather loose, with three to 15 flowers,
sometimes 25 or very exceptionally even 40, the
lower are held at 30-45 above the horizontal,
the unopened upper buds tending to be nearer
the vertical.
Bract: Green, narrow and pointed. On the lowest
few flowers the bract can be either very long and
leaf-like or much shorter, only half to two-thirds
the length of the ovary, but on all plants the bracts
are very short at the tip of the spike.
Ovary: Green, long and slender, prominently
twisted and boldly six-ribbed, with short hairs;
the very short stalk is minutely hairy.
Flower: Large and white with a golden-yellow
patch on the lip. The sepals and petals are a
delicate white, the sepals lanceolate and pointed
and the petals slightly shorter and broader. The
vast majority of flowers do not open widely,
the sepals and petals cupping the lip with the
tips of the sepals flared outwards, but in some
plants the lateral sepals are held spreading. The
lip is white, short and broad and divided into
inner (hypochile) and outer (epichile) halves
by a distinct narrowing or waist. The hypochile
and base of the epichile are held parallel to the
column with their sides curved upwards and
inwards to form a deep gutter. The epichile is
spade-shaped with a projecting central lobe or
tooth. The base of the hypochile is washed a
rich golden-yellow, and towards the tip of the
epichile there is an extensive patch of dense
papillae with a golden-yellow frosting, although
the fringe of the lip remains white. Towards the
base of this golden-yellow patch there are five
to six (rarely seven) parallel longitudinal ridges
that are also washed golden-yellow. The column
is whitish, long and projects forward (like a
boats figurehead) and the anther is tinged paleyellowish. There is no scent.
45
Subspecies
None.

None.
BIOLOGY
The flowers are pollinated by solitary bees.
Sword-leaved Helleborine produces no nectar
and relies on deceit to attract its pollinators;
the golden-yellow pseudopollen on the lip
may be especially important in luring insects.
However, for the deceit to function effectively
there must be a good supply of genuinely
nectar-producing flowers in the vicinity.
Furthermore, bees will only visit flowers in
bright sunlight and forage mostly between
10am and 3pm, and both the helleborines and
the surrounding nectar-producing plants have
to be in sunlight for some of that period; sites
that catch the morning sun may be especially
favoured. Pollination rates in Britain are very
variable, and a study in Hampshire found that
an average of 35% of flowering plants in sunny
glades produced at least one seed capsule;
under a high canopy this fell to 16% and in
scrub to just 7%. The maximum recorded in
any one season was 56% of flowering plants in a
M 27 May, Hampshire. The flowers sometimes open

more widely, showing the yellow pseudopollen on the lip.
O 27 May, Hampshire.
29/1/09 12:00:01
46
GENUS CEPHALANTHERA
There is no information from wild plants on

the length of the period between germination
and the first appearance above ground but it
may be four or more years. In the laboratory the
interval is two and a half years. Plants need to
have reached a certain height and minimum leaf
area before they can flower but do sometimes
bloom in their first year above ground.
Sword-leaved Helleborine is thought to lack
any method of vegetative propagation.
Hybrids
C. x schulzei, the hybrid with White
Helleborine, has been recorded very rarely from
Hampshire and West Sussex.

The specific name longifolia means simply
long-leaved.
HISTORY AND
CONSERVATION
M 27 May, Hampshire. Some plants are very robust.
sunny glade, although many of these may have

produced just one seed capsule. As well as sunlit
sites, higher rates of pollination are associated
with larger congregations of flowers, and it may
be that the spectacle of many helleborines in
flower is more attractive to bees. Conversely, at
many sites, especially where there are very small
populations, virtually no seed is produced.
Self-pollination has not been recorded and is
unlikely due to the structure of the flower.

Poorly known. The aerial stems grow from
a rhizome that produces long and sparsely
branched roots, with fungi concentrated in the
tips of the roots and in the few short side roots.
Plants are able to remain dormant underground
for about one year. Nothing specific is known
about its fungal associates but it seems
probable that like Red and White Helleborines
it acquires a substantial proportion of its
nutrients via a fungal route.
The first British record was published by

Christopher Merrett in 1666 in his Pinax
Rerum Naturalium Britannicarum (A picture
of British natural history): In Helk wood in
Yorkshire, not far from Ingleborough.
Sword-leaved Helleborine is classified as
Vulnerable in Britain and is specially protected
in Eire under the Flora (Protection) Order. It
is the subject of a Plantlife International Back
from the Brink project.
Past and present occurrence of Sword-leaved Helleborine in
Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
131
31
34 (1.2%*)
7 (0.7%*)
% lost, 1500-1969
60.5%
68%
% lost, 1970-1986
13.5%
9.5%
74%
77.5%
% lost, total
Sword-leaved Helleborine has been in decline

throughout the 19th and 20th centuries, initially
due to woodland losses and the replanting of
29/1/09 12:00:02
woodlands with conifers. At a few sites, small

populations were given the final death-knell
by collectors. The decline has continued in
the last 30 years, probably due to the lack
of woodland management. This has led to
the disappearance of glades and rides and
the development of a denser canopy, and the
management that does take place often involves
clear-felling large areas at once. The loss of
nearby hedgerows and permanent pastures may
have indirectly affected the species too, reducing
the populations of suitable pollinating bees.
Other threats include heavy browsing by deer,
now commoner in Britain than since the time
of William the Conqueror.
Sword-leaved Helleborine can now be found
at less than 50 sites, with around 20 each in
England and Scotland and five in Wales.
Research in the 1990s showed that 31% of sites
supported just one plant and 61% held less than
ten. Many are likely to be long-lived individuals
clinging on at a site no longer suitable for
reproduction, and the prospects of such tiny
populations surviving are bleak. By 2003
only nine sites in Britain held more than 50
plants, and in Ireland the species was formerly
recorded from 15 counties but is now extinct in
ten of them.
It is difficult to fashion effective
conservation measures for this orchid. Its
long-term survival depends on adequate levels
of flowering and fruiting and the subsequent
recruitment of enough new plants to the
population. It will flower well once the light
levels reach a critical threshold (which is rather
lower than the levels produced, for example,
by coppicing). Its pollinating bees require
higher levels of sunlight, however, and at these
higher light levels there is a tendency for rank
vegetation and scrub to take over and this will
overwhelm the helleborines. Nutrient creep
from surrounding agricultural land can also
promote the growth of rank vegetation that can
smother the orchids.
The most effective management regime
probably comprises limited felling to produce a
mosaic of woodland and glades, winter grazing
47
ExtINCt?
(the date of the last record is given where known)
Devon (before 1939)

Dorset (1876)
Wiltshire (1958)
East Sussex (1982)
Kent (1980s)
Oxfordshire (1970)
Norfolk (1883)
Herefordshire (1986)
Shropshire (last date uncertain)
Staffordshire (before 1901)
Derbyshire (before 1903)
Nottinghamshire (before 1839)
Lancashire (1898)
Yorkshire (1941)
Co. Durham (1882)
Northumberland (1841)
Monmouthshire (1987)
Cardiganshire (1985)
Montgomeryshire (1978)
Dumfries-shire (1854)
Ayrshire (1972)
Midlothian (1892)
Fife (1847)
Co. Cork
Co. Carlow
Co. Wicklow
Co. Dublin
Co. Tipperary
Co. Roscommon
Co. Mayo
Co Down
Co. Antrim (prior to c. 1840)
to suppress rank vegetation and protection

from deer and rabbits when flowering and
fruiting. Recent successes give hope for this
very special orchid: at the Little Shoulder of
Mutton in Hampshire, 31 plants were found in
1987 after Beech trees and scrub were cleared
to increase the size of an existing small patch of
chalk grassland. A programme of further scrub
control, mowing and protecting individual
plants with wire netting resulted in a negligible
increase, but dramatic results came when winter
grazing was introduced, with a jump to 240
plants by 2004. It now seems to be the perfect
site with a sunny, south-facing scrubby edge
adjoining chalk grassland (which can provide
nectar and nest sites for the pollinating bees).
29/1/09 12:00:02
48
GENUS CEPHALANTHERA
wHite HeLLeborine
Vulnerable, BAP
Cephalanthera damasonium
This subtly attractive orchid is relatively common in beechwoods in southeast England and
is sometimes almost the only flowering plant to be found beneath the dense canopy of the
trees. It has recently been found to have a special relationship with nearby trees, extracting
nutrients from them via a mutual fungal partner.
Identification
The loose spike of rather egg-shaped, upwardpointing, creamy-white flowers, most of which
do not open widely, is very distinctive. Most
flowers set seed, and the stout, elongated capsules, held upright, can identify the species late in
the season, well after the flowers have withered.
Similar species
Sword-leaved Helleborine is much rarer but is
sometimes found together with White
Helleborine. The flowers of Sword-leaved
Helleborine are always pure white and tend to
open more widely. In addition, the sepals have
more pointed tips, and the leaves, especially the
lower ones, are on average rather longer and
narrower. The best distinction is the length of the
bracts: on Sword-leaved Helleborine these are
shorter than the ovary, at least in the upper part
of the spike (on the lowest two or three flowers
they may be very long and leaf-like); on White
Helleborine the bracts are all longer than the ovary.
Habitat
M 26 May, Cambridgeshire. The flowers are usually

creamy and egg-shaped, with the sepals and petals hardly
opening at all.
Strictly confined to well-drained calcareous

soils on chalk and limestone and found in
woodland and shelter belts, sometimes in scrub
and occasionally on nearby grassland (especially
on north-facing slopes). It tolerates quite deep
shade, but the most robust plants are found
where the shade is not too intense. It is strongly
associated with Beech, even solitary trees, and
can happily grow in the dense shade cast by
this species. Its truly classic habitat is a beech
hanger on a steep slope with a sparse or
non-existent ground cover, the helleborines
growing through a carpet of dead leaves or on
bare stony or mossy ground.
29/1/09 12:00:03
WHITE HELLEBORINE
Flowering period
Mid-May to late June, exceptionally from late
April or to mid-July, with flowers in the open or
in very dry woods being the earliest.
49
1987-99
1970-86
pre 1970
Range
Southern England, especially the North and
South Downs, the chalk of Hampshire and
Wiltshire, and the Chilterns and Cotswolds.
Ranges west to Dorset, Somerset and
Gloucestershire, south to the Isle of Wight
and north to Herefordshire, Worcestershire,
Northamptonshire and Cambridgeshire. World
range: Almost confined to central and southern
Europe, extending north to Denmark, Gotland
in southern Sweden and the Baltic States, east
to Belarus and the western Ukraine and south
to the Mediterranean, including the Balearic
Islands, Corsica, Sardinia, Sicily and Crete, and
to the Crimea and Caucasus. Also found in
Turkey, Syria, Israel and northern Iran.
How to find it
Usually an easy species to find in suitable habitat,
although often growing well away from paths
and tracks in heavy shade in the depths of the
forest. Notable sites away from its heartland in
the southeast include Beechwood in Cambridgeshire and Browns Folly in Avon, and at Friston
Forest in East Sussex it is rapidly colonising a
large area of beech plantations.
DESCRIPTION
Height: 8-67.5cm but typically 15-40cm, and
many are just 13-18cm tall with only one or
two flowers.
M 26 May, Cambridgeshire. White Helleborine has a special association with Beech trees, extracting nutrients via a
mutual fungal link; it may occur in large numbers where little else can grow.
29/1/09 12:00:04
50
GENUS CEPHALANTHERA
29/1/09 12:00:08
WHITE HELLEBORINE
Stem: Green, the upper part is ridged and

either hairless or slightly hairy. There are one
to three brownish, membranous sheaths at the
base, the upper of which is sometimes tipped
green. Stems grow singly.
Leaves: Oval to broadly lanceolate, tapering to
a moderate point and becoming narrower and
more bract-like towards the spike; the lowest
leaf is rather short and cowl-like. The three to
five leaves are placed alternately up the stem,
more or less in two opposite rows, and curve
gracefully upwards to lie horizontally. They
are greyish-green, sometimes with a bluish
tinge, and have prominent veins (especially
on the underside).
Spike: Loose, with one to12 relatively large
flowers, sometimes as many as 16, most
pointing vertically upwards.
Bract: Greenish, narrowly lanceolate and
relatively long, often much longer than the
flower, but becoming shorter towards the tip of
the spike, although still longer than the ovary.
Ovary: Pale green, either not twisted or only
moderately so, slender, cylindrical and boldly
six-ribbed.
Flower: Relatively large and creamy-white.
The sepals are white to creamy-white with a
hint of green. They are tear-shaped to ovallanceolate with the broader end at the base and
a blunt tip. The petals are slightly shorter and
more oval in shape. The lip is short and broad
and divided into inner (hypochile) and outer
(epichile) halves by a distinct narrowing or
waist. The hypochile and base of the epichile
are held parallel to the column with their sides
curved upwards and inwards to form a gutter,
the base of which is washed golden-yellow. The
epichile is heart-shaped, rather broader than
long, with the tip curving gently downwards.
Towards the tip of the epichile there is an
extensive patch of golden-yellow frosting that
has three to five longitudinal ridges and furrows
towards its base; these ridges are also washed
golden-yellow. The column is long, slender and
whitish, with the anther tinged pale yellow. The
flowers do not normally open widely, the sepals,
petals and lip forming an egg-shape around the
51
M 31 May, Cambridgeshire. Although often forming a tall

spire, the shape of the spike is variable.
column, but in a few plants the lateral sepals

spread apart like outstretched arms and the
upper sepal and petals then form a loose hood.
There is no scent.
Subspecies
None.
29/1/09 12:00:09
52
GENUS CEPHALANTHERA
flower is fully fertilised, the lip folds up again to

shut the door. The mechanism is very efficient,
unlike Red and Sword-leaved Helleborines,
and almost all flowers produce seed. Vegetative
propagation also occurs. The roots produce
buds, and these develop into new aerial shoots.
M 31 May, Cambridgeshire.The spout-like tip of the lip is

revealed.

Var. chlorotica is deficient in chlorophyll, and
its stem and leaves are very pale green or even
yellowish-white. It has rarely been recorded.
BIOLOGY
The aerial stem grows from a deeply buried,

short, woody rhizome. This sends up to 95
roots vertically down into the soil; these are
thick, corky and 30-60cm long. The mature
plant was thought to be independent of fungi,
but recent isotope studies have shown that
it acquires the majority of its nitrogen and
roughly half its carbon from a fungal partner.
This undoubtedly explains the ability of
White Helleborine to thrive in densely shaded
sites where it is sometimes the only green
plant present. Albino, chlorophyll-less White
Helleborines are sometimes found and these
must be fully mycotrophic and completely
dependent on their fungal partner.
Although it appears to form
relationships with a wide range of fungi,

Probably largely self-pollinated in a process
that was described by Charles Darwin. The
anther opens while the flower is still in bud
and releases the pollinia, which sit on top
of the column (the flower is held pointing
upwards) with their front edge resting against
the stigmatic zone. The pollinia are very friable
and at least some of the pollen adheres to the
stigma and effects pollination. Once the flower
is mature, however, the outer part of the lip
folds down to form a landing platform for
insects, especially bees. These are attracted by
the golden-yellow mass of pseudopollen on
the tip of the lip (the ridges are said to taste of
vanilla and are often gnawed by some creature;
Darwin found minute, bitten-off fragments but
could not be sure of the culprit). It seems that
visiting insects may not only spread more pollen
onto the stigma within the flower but also
sometimes transfer pollen from flower to flower,
thus causing cross-pollination. But, once the
M 31 May, Cambridgeshire. Rather rarely, the flowers

open widely, showing the yellow pseudopollen and raised
ridges on the lip and the long column projecting into the
flower.
29/1/09 12:00:14
WHITE HELLEBORINE
White Helleborine preferentially forms

associations with ectomycorrhizal fungi,
especially Basidiomycetes; these in turn have
a relationship with nearby trees. Thus, like
Coralroot and Birds-nest Orchids, it gets
nutrients from the roots of these trees via the
fungi but unlike the trees, it cheats the fungus
by giving nothing in return. In view of this, the
well-known association of White Helleborine
with Beech takes on a more sinister significance.
After germination the seedling is reported
to spend eight years underground before
sending up aerial shoots, and flowers are not
produced for another two or three years. There
is, however, probably little direct evidence for
these timings.
53
Nottinghamshire, Derbyshire, Warwickshire,

Essex and Devon. This species is still, however,
relatively common within its core range and
can occur in large numbers. Furthermore, it
can colonise newly available habitats, such as
maturing beech plantations.
Past and present occurrence of White Helleborine in Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
% lost, 1500-1969
Britain
Ireland
233
136 (4.8%*)
33%
% lost, 1970-1986
8.5%
Hybrids
% lost, total
41.5%
C. x schulzei, the hybrid with Sword-leaved

Helleborine, has been recorded rarely from
Hampshire and West Sussex.

The origin of the specific name damasonium is
obscure. The usual derivation has no connection
whatsoever with orchids: damasonium was used
by Pliny (AD 23-79, author of the encyclopedic
Natural History) for Alisma, which in turn was
a name given by Dioscorides in around AD 64
to a plantain-leaved water plant. An alternative
derivation is from damaso meaning to subdue,
i.e. subdue evil, as the plant was considered an
antidote to the venom of toads.
HISTORY AND
CONSERVATION
1670 when John Ray noted this species in
his Catalogus Plantarum Angliae et Insularum
adjecentium (A catalogue of plants found
around Cambridge), In the woods near
Stokenchurch, Oxfordshire...
White Helleborine has been lost from
over 40% of its historical range, largely due to
woodland clearance and coniferisation, and
is classified as Vulnerable. Many of the losses
have been on the edges of the range, and it is
now extinct in southern Yorkshire, Shropshire,
M 31 May, Cambridgeshire. Plants may be tiny, with just

one flower (this was 11cm high).
29/1/09 12:00:14
GenUs NEOTTIA
birds-nest orCHids
and twaybLades
29/1/09 12:00:18
GENUS NEOTTIA
55
This genus contains two apparently very different groups, the birds-nest orchids and the
twayblades, but they are united by a very similar flower structure and almost identical
pollination mechanism.
Distribution
Pollination
Europe, temperate Asia and North America,

just creeping into North Africa. There are
c. 33 species in this genus, but only three occur
in Europe, all of which are found in Britain
and Ireland.
All three British species produce nectar and

have a hair-trigger mechanism. This fires a
drop of glue onto the head of a visiting insect,
and at the same time the pollinia are released.
Pollination is efficient and seed-set is high.
Classification
Growth pattern
The genus can be conveniently divided

into two groups. There are nine species of
birds-nest orchids from Europe and Asia,
all of which lack green leaves and are fully
dependent on fungi throughout their life,
and 25 species of twayblades, all of which
have a pair of green leaves held opposite each
other roughly midway up the stem. Until
very recently the twayblades were placed in
a separate genus, Listera, although they had
long been considered close to the birds-nest
orchids (the original Neottia), due to their
very similar flower structure and pollinating
mechanism. Recent genetic research has
shown that the two groups are so closely
related that they should all be in the same
genus, which takes the name Neottia, the older
and more senior of the two names available.
The aerial stem grows from a rhizome that

puts out numerous roots. The growth pattern
is sympodial, and each year the rhizome grows
upwards to form a new aerial stem that may
flower and always dies off. Underground the
rhizome continues to grow from a lateral bud at
the base of the aerial stem.
Floral structures
The column has a wide flat rostellum and
there are two stalkless pollinia. There is,
however, no detachable viscidium, rather the
rostellum expels its contents in a sticky drop
when triggered. There is no spur, and nectar is
produced in a central groove or slight hollow at
the base of the lip. The pollen is grouped into
tetrads and these are loosely connected by a few
weak threads. The pollen is rather powdery.
Fungal partners
In adult plants the roots have a heavy fungal
infection, and Birds-nest Orchid is dependent
on fungi throughout its life and is fully
mycotrophic.
The roots are heavily infected with fungi and
are nutritionally independent. They are also
able to form buds at their tips (a facility that is
unique to orchids) and these can develop into
new rhizomes. Thus, if the central rhizome dies
after flowering (as in Birds-nest Orchid) or is
otherwise fragmented, each piece can grow into
a new plant.
Name
The generic name Neottia means nest-offledglings, a reference to the appearance of the
roots of Birds-nest Orchid. (The genus Listera
was named after Martin Lister (1638-1711),
the English doctor and botanist.)
O 20 May, Norfolk. Birds-nest Orchid has a complex

relationship with fungi and forest trees, in particular
Beech. It can be hard to spot among the leaf-litter of the
woodland floor.
29/1/09 12:00:19
56
GENUS NEOTTIA
Lesser twaybLade
Neottia cordata
Formerly: Listera cordata: Other names: Heart-leaved Twayblade (North America)
This species occurs widely on wet moorland and woodland in Scotland, as well as in northern
England, Wales, Ireland and on Exmoor. It is very small and often hard to find, but it is well
worth the effort. On close examination one can see that the flower is a perfect miniature,
with every part reproduced exactly.
Identification
This little orchid is very distinctive. It has two
heart-shaped leaves set opposite each other
rather high on the stem and tiny, more-or-less
reddish flowers, each sitting on a large, globular
ovary. On close inspection with a hand-lens the
flowers resemble a tiny elfin figure. The deeply
forked lip forms the legs, the two hornlike
projections at its base the arms, while the sepals
and petals spread star-like around the column
to form a hat around the head.
Similar species
None, but there are usually a significant number
of non-flowering plants in any population, with
the paired leaves lying at the tip of the stem.
These are very like young Bilberry plants.
Habitat
Lesser Twayblade is found in two, apparently
distinct, habitats, but both offer the same
combination of cool, humid shade and acid
soils. The first and most frequent habitat is wet
moorland or peat bog, where it grows on the
cushions of moss, usually Sphagnum, found
under and between mature, leggy bushes of
Heather, Bell Heather and Bilberry. The best
conditions are usually found on north-facing
slopes. In the oceanic climate of Shetland it is
sometimes also found on short, heathy pastures.
The second habitat is damp woodland, where
the orchid can be found growing among a
variety of mosses, sometimes in open areas
and sometimes again among an understorey of
Heather, Bilberry and scattered Bracken fronds.
Willow, birch and alder woods are favoured
but Lesser Twayblade is also found in ancient
Caledonian pinewoods and mature pine
M 7 June, Merioneth. A tiny plant, sometimes with just

a few flowers in a reddish spike; no wonder they are
hard to spot.
29/1/09 12:00:20
LESSER TWAYBLADE
57
plantations. It occurs up to 1,065m above sea

level (Stob Coire Easain, Inverness-shire), and
most sites are now in the hills, as those in the
lowlands have largely been destroyed.
Flowering period
Mid-May to mid-July, exceptionally from
late April, but generally peaking from late
May. Once the flower has been pollinated the
column quickly withers and blackens but the
remainder of the flower sometimes persists
until September.
1987-99
1970-86
pre 1970
Range
Lesser Twayblade occurs throughout Scotland,
including the Outer Hebrides, Orkney and
Shetland, although it is absent from the central
lowlands. In England it is found from north
Lancashire and Yorkshire northwards, with an
isolated population in southwest England on
Exmoor (Somerset/Devon). In Wales it occurs
very locally from northern parts of Breconshire
and Cardiganshire northwards, including
Anglesey. In Ireland Lesser Twayblade is fairly
widespread from Co. Sligo, Co. Cavan and Co.
Down northwards but is very local in the south:
in Co. Dublin and Co. Wicklow in the east, Co.
Galway in the west, and from Co. Limerick and
Co. Tipperary southwards.
There are a few records of Lesser Twayblade
from elsewhere in southern England. In
M 7 June, Midlothian. Lesser Twayblade needs cool, moist

conditions and is found on north-facing slopes on damp
moorland or in woodland.
29/1/09 12:00:22
58
GENUS NEOTTIA
29/1/09 12:00:24
LESSER TWAYBLADE
Hampshire it was recorded in 1853 and 1895

near Bournemouth and then it turned up in
the New Forest near Brockenhurst in 1927-30
and was reported again there in the 1970s (a
record from Bratley in about 1980 is suspected
to have been a deliberate introduction). The
species has also been recorded from Baldwins
Wood in Hertfordshire in 1980 and from
Gravetye Woods in East Sussex in about 1975
and then again in 1989. Some of these records
are assumed to involve plants introduced
accidentally when pines or rhododendrons
were planted but wind-blown seed is a possible
source and in Hampshire there could have been
relict populations on the New Forest heaths.
World range: Found throughout the boreal
regions of the Northern Hemisphere in Europe,
Asia and North America, ranging furthest
south in the mountains. In Europe it extends
south to the Pyrenees, northern Italy, northern
Greece, the Crimea and Caucasus and adjacent
northern Turkey, and north to Greenland,
Iceland and northernmost Scandinavia. In
Asia it is found in Siberia eastwards to Lake
Baikal and in the Russian Far East, Sakhalin
and Japan. In North America it occurs south
to California and New Mexico in the west and
along the Appalachians to North Carolina in
the east.
How to find it
Locally common in Scotland, it is worth
looking for Lesser Twayblade wherever there
is rank Heather on damp, north-facing slopes,
as well as in suitable woodland, although the
number of spikes can fluctuate markedly from
year to year. It is often to be found under the
Heather, or at least just under its eaves, and
it may be necessary to move the vegetation
aside to see the orchids. Scattered plants can
be exceedingly difficult to find but, fortunately,
the species often occurs in loose colonies and at
least one or two plants may be more obvious.
The key to success is patience and perseverance.
Once a Lesser Twayblade is spotted, a careful
search of the area will usually produce more,
although a majority may be non-flowering and
59
even harder to spot. This species is very local in

Wales but one well-known site is Roman Steps
(Gwynedd).
DESCRIPTION
Height: 3-25cm but usually 5-10cm; tends to
be tallest in sheltered woodland.
Stem: Green or reddish-purple, ridged
towards the tip and with fine glandular hairs
for a short distance above the leaves. There are
one or two membranous, brownish sheaths at
the base of the stem. Usually grows singly but
occasionally two or three stems grow from the
same rhizome.
Leaves: Two, lying opposite each other,
one third to halfway up the stem and held
either horizontally or at up to 45 above the
horizontal. They are dark, shiny green and
roughly heart-shaped, with a prominent midrib
that terminates in a tiny projecting point
(mucro). The leaves are also faintly net-veined
(reticulate) and often have undulating margins.
Spike: Relatively open, with three to 20 flowers.
Bract: Tiny, triangular and greenish.
Ovary: Green, spherical, with six reddish ribs,
held on a reddish or greenish stalk that is a little
longer than the ovary, ribbed and twisted.
Flower: Very small and variable in overall
coloration; it usually has a pronounced reddish
tone but can be much plainer and greener. The
M 7 June, Midlothian.The flowers form tiny, elfin figures.

O 7 June, Merioneth.
29/1/09 12:00:25
60
GENUS NEOTTIA
sepals are greenish, variably washed reddish in

the centre and around the edges, and are oval
with blunt tips. The petals are narrower and
more strap-shaped and tend to be redder. Both
sepals and petals are widely spread and form
a star-like pattern around the column. The lip
is coppery or pale green, washed red, relatively
large and triangular and divided more than
halfway to the base into two sharply pointed
lobes. Tiny amounts of nectar are produced
in a disc-shaped nectary at the base of the

lip just below the column; there are two very
short horn-shaped lobes on either side of this
nectary and a longitudinal nectar-filled groove
running from it to the base of the fork. The lip
is held pointing downwards, more or less at
right angles to the column. The column is short,
stubby and whitish, with a large, thin, leaf-like
rostellum that extends forward over the base
of the lip, above which lie the yellow anther cap
and yellow pollinia. The pollinia are shed by
the anther when the flower is still in bud and lie
loose on top of the rostellum, held in position
by its incurved margins. The flowers have a
faint but unpleasant foetid odour, probably
originating from the nectar.
Subspecies
None.

Var. trifoliata has a third leaf above the
usual two. It is rare, but has been recorded in
Scotland.
BIOLOGY
M 7 June, Midlothian. The ovary is roughly spherical and

almost as big as the flower.
The flowers are pollinated by a variety of small

insects, including flies and gnats, attracted by
the nectar. Three pressure-sensitive hairs project
from the tip of the rostellum and these act as a
trigger. The slightest touch by an insect causes
a droplet of glue to be squirted explosively
onto the insects head, and the pollinia are
simultaneously released and fall onto this glue.
The glue dries in just a few seconds and the
pollinia are carried off by the startled insect.
When the flower first opens the flaplike rostellum physically blocks access to the
stigma and any insect visitor will trigger the
mechanism the moment it touches the hairs on
the rostellum. The rostellum remains in place
once the pollinia have been removed but is now
spread flat, having released the pollinia. This
prevents self-pollination should the pollencarrying insect return to the flower immediately
after it has left. About 24 hours later the
rostellum slowly moves upwards, allowing
29/1/09 12:00:26
LESSER TWAYBLADE
insects to deposit pollen from other flowers

onto the stigma, which has now become very
sticky. Insects are reported to work upwards
from the bottom of the spike towards the
top. They therefore start with the lowest and
most mature flowers (i.e. those likely to have
receptive stigmas), before moving on to younger
flowers that still have pollinia waiting to be
removed. If this is so, an insect cannot pollinate
flowers on the same plant. If by any chance
the pollinia are not removed by an insect, the
rostellum lifts upwards anyway after a few days,
allowing the flower to be self-fertilised.
The individual flowers of Lesser Twayblade
are self-compatible and artificial self-pollination
will produce viable seed, but studies in
California suggested that it is not usually selfpollinating. However, it is possible that, as in
Common Twayblade, small quantities of pollen
may occasionally fall from the pollinia on to the
stigma or be carried there by tiny insects.
Seed-set is very efficient and the capsules
mature and split open within five weeks. The
lowest, oldest capsules on a spike may be
ripe and shedding seed before the uppermost
flowers are even pollinated. Indeed, the capsules
swell so quickly and seed is produced so
efficiently that some authors suggest that selfpollination must occur routinely.
Lesser Twayblade also reproduces
vegetatively, from buds on the roots.

The aerial stem grows from a slender, creeping
rhizome that lies near the surface of the soil.
This puts out a few long, slender, hairy roots.
Usually only one stem is produced, but buds
can form at the tip of the roots and develop
into additional aerial shoots that flower in
their third year. Lesser Twayblade is apparently
short-lived and there is little information on the
interval between germination and flowering,
although the first green leaf is reported to
appear after two or three years of underground
development.
61

The specific name cordata means heart-shaped,
a reference to the shape of the leaves.
Together with Common Twayblade, this
species was formerly placed in the genus Listera,
but genetic studies have indicated that Listera
should be united with Birds-nest Orchid in the
genus Neottia, as it was in the past.
HISTORY AND
CONSERVATION
The first British record was published in 1666
by Christopher Merrett in his Pinax Rerum
Naturalium Britannicarum (A picture of British
natural history): Neer the Beacon on Pendle
Hill in Lancashire.
Past and present occurrence of Lesser Twayblade in Britain and Ireland (based on presence or absence in 10km
Britain
Ireland
822
96
454 (16%*)
48 (4.8%*)
% lost, 1500-1969
25%
41.5%
% lost, 1970-1986
19.5%
8.5%
% lost, total
44.5%
50%

1500-1999
current range
Since 1666 the species has disappeared

from many sites, especially in the lowlands
and on the periphery of the range in northern
England, where the drainage and reclamation
of bogs and heaths caused many losses in the
19th century. Lesser Twayblade has gone from
44.5% of its historical range in Britain and
50% in Ireland, with a rather large proportion
of the losses in Britain being comparatively
recent. Given that it has been better recorded
in recent years because more people are actively
looking for it, the actual decline has surely been
substantially greater. It is extinct in Shropshire,
Derbyshire, Cheshire, Flintshire, southern parts
of Yorkshire and the Isle of Man, and probably
also Montgomeryshire.
Hybrids
None.
29/1/09 12:00:26
62
GENUS NEOTTIA
Common twaybLade
Neottia ovata
Formerly: Listera ovata
This rather unassuming orchid is one of the commonest and most widely distributed species
in Britain and Ireland. Its small, green, inconspicuous flowers belie the fact that it is very
long-lived and has an intricate and efficient means of using insects as pollinators.
Identification
Straightforward. It is green or greenish-yellow
overall with two large, egg-shaped leaves held
opposite each other at the base of the stem
and a tall spike of small flowers, each of which
resembles a tiny green figure. Non-flowering
plants, with just two leaves opposite each other
at the tip of the stem, are fairly frequent.
Similar species
Lesser Twayblade is rather similar but tiny, with
heart-shaped leaves, and flowers that are usually
reddish and have sharply pointed tips to the
lobes of the lip.
Man, Frog, Fen and Bog Orchids have
greenish flowers but all of these differ markedly
in the structure of the lip. Man and Frog
Orchids also differ in having a basal rosette of
leaves and Bog Orchid has merely tiny clasping
leaves at the base of the stem.
Habitat
Possibly more varied than any other British
orchid. It is found on short chalk grassland,
machair, dune slacks, limestone pavements,
permanent pastures, road verges and fens, and
also in scrub, hedgerows and moist deciduous
woodland, sometimes in deep shade. It has a
preference for calcareous soils but will grow in
mildly acidic conditions, occasionally amongst
Bracken and Heather. It can sometimes be
found in relatively new habitats, such as disused
railway lines, quarries and sand-pits or in
plantations, even of pine. It occurs up to 670m
above sea level (Ben Lawers, Perthshire).
Flowering period
Late April to early August, latest in the north,
exceptionally even to September.
M 29 June, Hampshire. Twayblades have two leaves and

the English name originates from tway, an archaic and
obsolete word for two.
P 23 May, Co. Clare.
29/1/09 12:00:27
COMMON TWAYBLADE
63
29/1/09 12:00:29
64
GENUS NEOTTIA
1987-99
1970-86
pre 1970
Range
Occurs almost throughout Britain and Ireland,
including the Outer Hebrides and Orkney, but
absent from Shetland, much of the Scottish
Highlands and many coastal regions of southern
Ireland. World range: Essentially a European
species but the range extends into Asia. In
Europe it occurs north to Iceland and c. 70
N in northern Scandinavia and south to the
Mediterranean, including Corsica, Sardinia,
Sicily and Crete, and to the Crimea and
Caucasus. In Siberia it ranges to c. 95 E,
and in southern Asia it is found in Turkey,
northern Iran, the Altai Mountains and western
Himalayas. Introduced in southern Ontario in
Canada.
Stem: Green to mid-brown, thicker and whiter

at the base with two or three membranous,
scale-like basal sheaths and numerous short,
white, glandular hairs. The stems usually
grow singly, although not infrequently there
are clusters.
Leaves: The two large, egg-shaped or elliptical
sheathing leaves are held opposite each other
towards the base of the stem; sometimes they
lie flat and sometimes they are angled at up to
45 above the horizontal. The leaves are green
in the shade and more yellowish-green in sunny
places, with three to five prominent veins. There
are also one to three tiny, triangular, bract-like
leaves higher on the stem.
Spike: Loose to fairly compact, with 15-30
small green flowers, sometimes as many as 100.
Bract: Lanceolate and short; usually shorter than
the flower stalk and sometimes much shorter.
How to find it
One of the commonest and most widespread
orchids and usually fairly easy to find, although
in regions away from chalk and limestone
soils it is often rather local. The best places to
look will be in ancient woodlands, marshes
or in grasslands that have not been improved,
ploughed or cut too closely. Being green,
including the flowers, odd plants can be
inconspicuous but Common Twayblades are
often found in large numbers.
DESCRIPTION
Height: 10-75cm but usually 20-60cm.
M 15 June, North Uist. Twayblades are found in a wide

variety of habitats, from woodland to chalk downland or
(as here) coastal machair.
29/1/09 12:00:31
COMMON TWAYBLADE
65
Ovary: Green, short and rounded, with six

prominent reddish-brown ribs. The reddishbrown flower stalk is twisted and a little longer
than the ovary; both are variably hairy.
Flower: Small, green and vaguely man-like.
The sepals are bluntly oval and dull green,
sometimes tinged or fringed dull reddish
or brown. The petals are dull green and
narrower and more strap-shaped. Both sepals
and petals curve inwards to form a very
loose hood around the short, thick column.
The lip is yellowish-green, long and strapshaped, sharply folded down and backwards
below the flower and divided for around half
its length into two blunt-tipped lobes (the
legs); a shallow nectar-bearing groove runs
down the centre of the lip to the base of the
fork. The column is greenish. The flowers are
variously described as scented (light, musky or
repellent) or as odourless.
Subspecies
None.

Var. trifoliata has three leaves and is not
uncommon. The third leaf is smaller, more
pointed and placed above or, less frequently,
below the main leaves.
Var. platyglossa has a short lip that broadens
into blunt diverging lobes, sometimes with
a small tooth between them. It was formerly
found in dunes in South Wales.
BIOLOGY
Pollination was intensively studied by C. K.
Sprengel (published in 1793) and later by
Charles Darwin. The flowers are pollinated
by small insects, especially ichneumon wasps
but also by sawflies and beetles, attracted
by the flowers scent. Once it has landed the
insect follows the nectar-filled groove up the
lip and makes contact with the projecting
rostellum. The pollinia are then stuck to its
head by the sudden secretion or explosion
of a drop of sticky liquid that dries in just
two or three seconds. The mechanism is
M 5 July, Norfolk. The tiny green flowers look very much

like a human figure.
extremely sensitive and the slightest touch will

trigger it. The startled insect flies away with
the pollinia attached, often to another plant
altogether. Meanwhile, once the pollinia have
been removed, the rostellum bends forwards
and downwards, thereby hindering access to
the stigma and preventing self-pollination. It
then slowly shrinks away upwards again to
expose the stigma to visiting insects. If they
are carrying pollinia these can then pollinate
the flower as they try to get at the nectar at the
base of the lip. The mechanism is efficient and
many flowers set seed. Self-pollination may also
occur occasionally; if the pollinia dry out small
fragments of pollen may fall on to the stigma
and effect pollination. The seed capsules each
contain an average of 1,240 seeds.
29/1/09 12:00:32
66
GENUS NEOTTIA
study in Sweden 20 out of 29 plants were still

alive after 40 years. Mature plants have been
recorded spending one or two years dormant
underground and then reappearing.
Hybrids
None.

The specific name ovata meaning egg-shaped is
a reference to the shape of the leaves. Together
with Lesser Twayblade, this species was
formerly placed in the genus Listera, but recent
genetic studies have indicated that this should
be united with the genus Neottia.
HISTORY AND
CONSERVATION
M 27 June, Norfolk.Vegetative multiplication can produce

clumps of spikes.
Vegetative propagation also occurs, with

buds on the roots producing new rhizomes, and
group of clones can be formed, sometimes a
dense circular cluster of dozens of plants

The aerial stem grows from a short, thick
rhizome which has large numbers of long, fairly
thick roots. Seeds are thought to germinate
in spring, and early estimates of the interval
between germination and flowering were seven
to 20 years or more, with the seedling spending
the first three or four years underground. These
estimates may be rather inaccurate, however,
and in the laboratory plants can produce leaves
less than a year after germination.
Common Twayblade can be extremely longlived. The remains of 24 old flower spikes have
been counted on a single rhizome, and in a
Together with Autumn Ladys-tresses, this

was the first orchid to be recorded in Britain.
In 1548 William Turner noted in his Names of
Herbes: Martagon...in many places of Englande
in watery middowes and in woddes.
Common, widespread and with a very
catholic choice of habitats, this species would
seem to be well-placed to survive changes
to the countryside. Nevertheless, Common
Twayblade has vanished from almost 30%
of its historical range in Britain and Ireland,
with a relatively large proportion of the British
losses being recent.
Past and present occurrence of Common Twayblade in Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
1,869
512
1,354 (47.5%*) 362 (36%*)
% lost, 1500-1969
16%
22.5%
% lost, 1970-1986
11.5%
6.5%
% lost, total
27.5%
29%
29/1/09 12:00:33
BIRDS-NEST ORCHID
birds-nest orCHid
67
Near Threatened
Neottia nidus-avis
Spotting a Birds-nest Orchid in the woodland gloom is somehow very reassuring; it is a
sign that you are in a special place. The species is locally common in mature woodland in
parts of southern England, but in much of the remainder of the British Isles it is scarce and
very local. One of the three British orchids that have no green leaves and are completely
dependent throughout their life on nutrients provided by fungi, Birds-nest Orchid only
appears above ground in order to flower and set seed. Both the English and the scientific
name relate to this subterranean existence, as the roots of this orchid form an untidy mass
that vaguely resembles the nest of a Wood Pigeon or Rook.
Identification
The honey-brown spikes are unique among
British orchids. On careful examination the
flowers are typical of an orchid, with the petals
and sepals forming an open, fan-shaped hood,
and the lip vaguely resembling a human torso.
The dried stem and open seed capsules of the
previous seasons blooms may be found nearby
and these can remain intact for almost two years.
Similar species
Confusion is possible with Yellow Birds-nest,
a more-or-less similarly coloured but totally
unrelated plant that is often found in the
same habitats, although the spike of Yellow
Birds-nest is bent over and nodding until it is
very mature. The various broomrapes (family
Orobanchaceae) also superficially resemble
Birds-nest Orchid, especially Knapweed
Broomrape, which is closest in colour. These
chlorophyll-less parasitic herbs are found in
open, grassy habitats and could occur on the
edge of woods or in woodland rides. A quick
look at the structure of the flower will settle the
matter, however, as neither Yellow Birds-nest
nor the broomrapes have a flower with a hood
and a two-lobed lip.
Habitat
The classic habitat for Birds-nest Orchid is the
heavy shade of a mature beechwood, the orchids
emerging from the leaf-litter and deep humus of
a woodland floor otherwise devoid of vegetation.
It also grows in mixed deciduous woodland and
M 2 May, Oxfordshire.
29/1/09 12:00:34
68
GENUS NEOTTIA
bloom in the latter half of May. Birds-nest

Orchids have been recorded flowering and
setting seed underground if the route to the
surface has become blocked by a large stone
or other obstacle. As no systematic searches
have been made, it is not known whether this
is exceptional or a regular event (Australian
orchids of the genus Rhizanthella routinely
flower underground).
Range
M Yellow Birds-nest, 16 June, Norfolk. Another fully

mycotrophic (fungus-dependent) species of the woodland floor, the similarity to Birds-nest Orchid ends with
the honey colour.
overgrown hazel coppice or sometimes under

shady old hedges, shelter-belts or planted
conifers, especially if there are still deciduous
trees present (and in Europe it regularly grows
in conifer woods). Rarely, it has been recorded
from grassland just outside woods, but this is
exceptional, as exposure to direct sunlight tends
to dry out the ground too much. Conversely,
it is not found in areas where the soil becomes
waterlogged. Birds-nest Orchid is commonest
on chalk and limestone soils but also grows on
clays and sands that have a chalky or limestone
component, such as boulder clay. Generally it is
a lowland species, but it has been recorded up to
250m above sea level in Cumbria.
Found almost throughout Britain and Ireland

but absent from the Isle of Man, Shetland,
Orkney and the Inner and Outer Hebrides
(with the exception of Skye and Mull); there
are, however, only a few sites in northern
Scotland, much of Ireland and in a belt from
the northern parts of East Anglia through
Lincolnshire and the north Midlands to
north and west Wales. Its strongholds are
undoubtedly southern England (especially
central-southern England) and southeast
Wales. World range: Occurs in Europe
and western Siberia, reappearing in the Far
East. In Europe found north to c. 65N in
Scandinavia and Finland and south to the
Mediterranean, including Corsica, Sardinia,
Sicily and the Balearic Islands, and to the
Crimea and Caucasus; it is largely confined to
the mountains in the south of the range and
absent from the Mediterranean lowlands. In
1987-99
1970-86
pre 1970
Flowering period
Early May to late June, exceptionally from
late April or to early July, but most are in
29/1/09 12:00:35
BIRDS-NEST ORCHID
Siberia it extends to c. 85E and it is also found

in Algeria in North Africa and at scattered sites
in Turkey and northwest Iran. In eastern Asia it
occurs in the Russian Far East, Sakhalin, Japan
and Korea.
How to find it
In most of Britain and Ireland the species is
rather local and only found in small numbers.
It often emerges in the darker and shadier
parts of a wood and this, combined with
its pale-brown colour, can make it hard to
find. Birds-nest Orchid is totally dependent
on the activity of its fungal partners and
not surprisingly does best in the warm, wet
conditions beloved of fungi in general. Warm,
wet springs can encourage larger numbers
of plants to flower, and conversely very dry
periods can result in a reduction in the aboveground population.
69
DESCRIPTION
Stem: Yellowish-brown, slightly glandularhairy towards the tip. Grows singly, although
two spikes occasionally develop from the same
rhizome.
Leaves: No green leaves are present. The lower
part of the stem is enclosed by three to five long,
roughly oblong, scale-like, yellowish-brown,
loosely sheathing leaves (the upper ones longer
and blunter).
Spike: Cylindrical and crowded, with up to 100
flowers in large plants, but the lower flowers
in the spike are usually more widely spaced
and there are odd single flowers further down
the stem.
Bract: Papery, lanceolate and roughly as long as
the ovary and stalk together, but inconspicuous.
Ovary: Oval, subtly six-ribbed, glandular-hairy
M 3 June, Norfolk.Whether through seed or vegetative reproduction via division of the underground rhizome, Birds-nest
Orchid sometimes occurs in groups.
29/1/09 12:00:38
70
GENUS NEOTTIA
BIOLOGY
M 26 May, Norfolk. Comparison of the flower structure

with Common Twayblade shows how close they are.
and held on a twisted stalk that is about half

the length of the ovary.
Flower: Entirely yellowish-brown. The sepals
and petals are roughly oval-spatulate in shape
and form a loose fan-shaped hood over the
column. The lip is slightly darker brown and
has a nectar-producing bowl-shaped depression
at the base (representing a rudimentary spur).
The lip is divided towards the tip into two
broad, rounded lobes that spread widely,
especially on the lower flowers, to form a lyreshape; there may also be a subtle point or tooth
on either side of the lip half way towards the
base. The lip is held pointing outwards and
downwards at c. 90 to the column, which is
pale brown, long and slender. The pollinia are
yellow and project conspicuously from beneath
the anther cap. The flowers have a pleasant but
sickly, honey-like scent.
Subspecies
None.

Var. pallida has a yellowish-white stem and
flowers, and white pollinia. It is rare.
Pollination is by insects, including flies,

attracted by the nectar. The mechanism is
very similar to that of Common Twayblade.
The visiting insect makes contact with the
projecting rostellum where there are six minute,
rough, touch-sensitive points and the pollinia
are stuck to its head by the sudden secretion
or explosion of a drop of sticky liquid. After a
while the rostellum, which has hitherto blocked
access to the stigma, rises to allow visiting
insects, complete with pollinia attached to their
heads, to make contact with the stigma. If the
mechanism is not triggered, after a few days
the pollinia fragment and pollen can then fall
onto the stigma below, effecting self-pollination
(autogamy); pollen may also be carried to
the stigma by small insects such as thrips.
Ants have been noted carrying pollen from
one flower to another on the same spike and
this may also effect self-pollination (this time
geitonogamy, as it is pollen from a different
flower on the same plant). Occasionally, selfpollination may take place in the bud before the
flowers have even opened. Pollination is very
efficient and almost all flowers will set seed.
Birds-nest Orchid is thought to be
monocarpic, that is the plant dies after
flowering once. But, although the rhizome dies,
the numerous roots can remain alive and go on
to produce new plants from buds at their tip.

The aerial stem develops from a short rhizome
that lies horizontally in the soil and is almost
entirely surrounded by an untidy mass of short,
thick, fleshy roots that stick out more or less at
right-angles.
Throughout its life Birds-nest Orchid is
entirely dependent on fungi for nutrition;
in adult plants fungi are found exclusively in
the roots of the orchid (in the three cortical
cell layers, just below the epidermis). Recent
studies have shown that the orchid is very
specific about its fungal partner and only
forms an association with a species of Sebacina.
29/1/09 12:00:39
BIRDS-NEST ORCHID
This obtains its carbohydrates by forming a

symbiotic ectomycorrhizal association with
the roots of trees, particularly Beech: the tree
produces carbohydrates through the process of
photosynthesis and passes these to the fungus,
which in turn contributes mineral nutrients to
the tree. The orchid invades this relationship
and, via the fungus, gets nutrients from the tree.
Birds-nest Orchid cheats in its partnership
with the fungus because, unlike the tree, it
does not contribute anything to the fungus
(see also p.9).
It is often stated that Birds-nest Orchid
lacks chlorophyll, but a small amount of
chlorophyll is present, although no effective
photosynthesis can occur. It is also often said
that Birds-nest Orchid is saprophytic and
obtains its nutrients from decaying organic
material. This is not correct. The orchid gets its
nutrients by digesting the living fungus.
Seeds germinate in the spring and require
the presence of the same fungus that supports
the growth of the adult plant. The seedling
initially takes the form of a torpedo-shaped
protocorm a few millimetres long. It then
begins to develop short, fat rootlets that stick
out at about 90 and start to take on the
appearance of a birds nest. At an early stage
the bud that will produce the flower spike
appears in the axil of a scale leaf at the tip of
the rhizome. As the rhizome grows the number
of roots progressively increases and the period
from germination to flowering is probably three
to five years.
Seed can only germinate where the Sebacina
fungus is present and this may have a localised
and restricted distribution, even within a
single wood, at least partly because the fungus
is entirely dependent on its host tree species
and cannot grow without it. The distribution
of the orchid is therefore also tied to that of
suitable host trees. Adult Birds-nest Orchids
always harbour the correct fungus and form
a convenient source of infection for the seed,
and germination is far more prolific in the
immediate vicinity of adult plants. The clusters
of Birds-nest Orchids that often grow on the
71
M 13 May, Norfolk. When shoots emerge in the spring

the flower buds are already fully formed.
exact spot that previously held a flowering plant

may be the product of this, or alternatively, of
the break-up of a single rhizome to produce
several new plants.
Hybrids
None.

The specific name nidus-avis means birds nest
(as the generic named Neottia means nest-offledglings the scientific name is a tautology).
With a similar flower structure and
pollination mechanism, the Birds-nest
Orchid is clearly related to the twayblades.
DNA studies have recently shown that the
relationship is a close one, and they are all now
placed in the same genus, Neottia.
29/1/09 12:00:41
72
GENUS NEOTTIA
HISTORY AND
CONSERVATION
First recorded from Britain in 1597 by John
Gerard in his Herball: I found it growing in
the middle of a wood in Kent two miles from
Gravesend.
With its requirement for heavy shade and a
stable, generally moist environment, the species
has undergone a significant decline, especially
in the period after 1945. This is due to the
grubbing out of woodland and the conversion
of deciduous woodland to conifer plantations.
More subtle changes, such as the use of heavy
machinery in forestry operations, could also
have been detrimental. The decline has been
most marked in southeast England, especially
Kent and the eastern Chilterns, and Birds-nest
Orchid has gone from 54% of its total historical
range in Britain and 45% in Ireland. As well as
being more widespread, populations may have
been larger in the past. For example, thousands
were recorded on the south-facing slopes of Box
Hill in Surrey in 1947 at a site which nowadays
only supports about 200 spikes. Birds-nest
Orchid is classified as Near Threatened in
Britain and is specially protected in Northern
Ireland under Schedule 8 of the 1985 Wildlife
Order (NI).
Past and present occurrence of Birds-nest Orchid in
Britain and Ireland (based on presence or absence in
10km squares of the National Grid; data from the New
Atlas).
Britain
Ireland
742
99
340 (12%*)
54 (5.4%*)
% lost, 1500-1969
41%
29.5%
% lost, 1970-1986
13%
16%
% lost, total
54%
45.5%

1500-1999
current range
O 6 June, Norfolk.The dead spike may persist for at least

a year.
P 8 July, Norfolk. Marsh Helleborines.
29/1/09 12:00:42
GENUS EPIPACTIS
HELLEBORINES (II)
12/2/09 16:53:23
74
GENUS EPIPACTIS
A genus of essentially woodland orchids which has diversified to occupy a variety of

other situations. Recent research has shown that fungi contribute very significantly to
their nutritional budgets, even as adults, with several having connections via their fungal
partners to neighbouring trees. As a genus, Epipactis is distinctive, with flowers that are
rather different to those of typical orchids, but it is necessary to have a good grasp of its
flower structure in order to identify some of the species with confidence.
Distribution
Largely confined to Europe and Asia with single
representatives in Africa and North America.
The number of species is uncertain, with the
most conservative authors listing around 11
species in Europe, whereas Delforge (2005)
details 59. There are eight species in the British
Isles, including two endemics.
Classification
Epipactis is closely related to the genus
Cephalanthera, and, although the flowers look
quite different, both have the lip divided into
outer and inner portions by a narrow waist.
Epipactis differs in having more complex
reproductive structures and an ovary that is not
twisted; rather it has a distinct stalk which is
twisted to bring the lip around to the bottom
of the flower.
The genus is sometimes divided into two
sections: Arthrochilium, which contains Marsh
Helleborine, and Euepipactis, which contains the
remainder of our species. In Marsh Helleborine
the inner and outer halves of the lip are joined
by a flexible hinge, whereas in all the other
species the joint is rigid and the inner portion of
the lip does not form such a distinct bowl.
Identification
Epipactis helleborines are relatively easy to
recognise as such. They have upright stems 10120cm tall and oval leaves with obvious parallel
veins. In some species, several spikes can arise
from the same rootstock. When it emerges
from the soil the stem is bent double and as it
grows it continues to weep. Eventually, however,
the stem becomes fully upright and the flowers,
which remain in bud for a frustratingly long
time, begin to open. In many species the flowers
are relatively small and drab but they may be
M Violet Helleborine, 29 June, Buckinghamshire. The

flowers spike is fully formed when the stem breaks
throughthesoilsurfacebutitmaybeseveralweeksbefore
itelongates,becomesuprightandtheflowersopen.
very numerous. All the helleborines bloom

from mid- to late summer, later than most
other orchids.
Floral structures
All the Epipactis helleborines have a similar
flower structure. The ovary forms the
apparent stalk of the flower. It actually
narrows into the real stalk, the pedicel,
just before it reaches the main stem. Once
fertilisation has occurred, the ovary swells
conspicuously to form a capsule while the
flower shrivels to a few brown wisps. In all
species the ovary is usually obviously ribbed.
The three sepals and two petals are rather
similar in size, shape and coloration; in
29/1/09 12:11:01
GENUS EPIPACTIS
some species they form a five-pointed star

around the column and lip, in others the
flowers do not open widely and form a cup
or bell shape. In Epipactis the lip is divided
into two parts. The inner section is known
as the hypochile; it is cup-shaped and often
contains nectar. The outer part, or epichile,
is flatter, more or less triangular in shape and
often reflexed (bent under) at the tip. At the
base of the epichile there are raised areas
known as bosses and in some species these
are contrastingly coloured. The hypochile and
epichile are connected by a narrow strip that
is rigid in most species.
Reproductive structures
The column is a robust structure which projects
from the ovary into the centre of the flower.
In Epipactis it is often conspicuous and can be
important for identification. It is topped by the
anther or anther cap, which is attached to the
remainder of the column by a short stalk or
narrow flexible hinge. The two pollinia develop
side-by-side within the anther. Shaped like a
fat sausage, each pollinium is in turn divided
into two segments by a fine groove, and the two
pollinia are joined together at one end in the
form of a wishbone.
Below the anther cap, the top of the column
has a system of grooves, the clinandria. These
form a platform on top of the main part of
the column. As the pollinia mature, the anther
splits open and the pollinia drop down to rest
in the clinandria, which are moulded to
their shape. The pollinia remain protected
from above by the anther cap and are often
largely hidden.
At the front of the column, separating
the clinandria from the stigma below, lies
the rostellum. (Literally the little beak,
the rostellum is actually a greatly modified
sterile third stigma.) The rostellum secretes
a detachable white cap, the viscidium. This is
a thin-skinned sack containing a sticky liquid
and the slightest touch will rupture the skin
and release the glue, which sets hard on contact
with the air. Below the rostellum lies the fertile
stigma. Unlike other flowers, where the stigma
stalk
(petiole)
ovary
column
75
anther cap
pollinia
viscidium
rostellum
bract
boss
hypochile
epichile
lip
MBroad-leavedHelleborineflowerinprofilewithsepals
andpetalsremoved.
is an obvious structure, in Epipactis and other

orchids it is just a region on the front of the
column that becomes sticky when it is receptive
to pollen.
Pollination
In cross-pollinated helleborines, the clinandria
or grooves on top of the column are relatively
deep and the pollinia lie there securely. The
rostellum functions as an additional barrier,
preventing the pollinia from falling onto the
stigma below. In these species the pollinia
remain intact and lie with their tips in contact
with the viscidium. A wasp visits the flower to
drink nectar from the cup-shaped hypochile
but to reach this it must rub its head or the top
of its thorax against the rostellum. In doing
so it ruptures the viscidium and the sticky
contents glue the pollinia to the insects head.
Thus, when the wasp flies off, it carries one
or both of the pollinia away with it. The wasp
goes on to visit another flower, deposits parts
of the pollinia on the wet surface of the stigma
and pollination takes place; once pollinated, the
stigma changes from whitish to brownish.
In self-pollinated helleborines, the
clinandria are shallow and do not hold the
pollinia securely, the rostellum is usually
reduced in size and the viscidium either never
develops or is present in the bud but disappears
when the flowers open; there is therefore little
to prevent the pollinia, or parts of them, from
falling onto the stigma below. The pollinia also
tend to be more friable and easily fall to pieces.
The pollinia may either disintegrate where they
29/1/09 12:11:01
76
GENUS EPIPACTIS
r
r
MBroad-leavedHelleborine, 28July, Norfolk.Theviscidiumandpolliniaareconspicuous.
lie (the pollen germinating in the clinandrium

and the pollen tubes growing down to the
stigma and ovary), or the pollen may fall on to
the stigma below and pollinate it. Indeed, in
some Green-flowered Helleborines the flowers
never open and pollination occurs within the
unopened bud. Contrastingly, it is probable that
the Dune Helleborine, although self-pollinating,
occasionally has a functional viscidium and is
therefore sometimes cross-pollinated.
In the self-pollinated species, crosspollination can occur from time to time, even
without a viscidium. Their flowers, although
typically small, dull and not opening widely,
are visited by insects in order to drink nectar.
The pollinia may stick to their bodies despite
the lack of the viscidium as glue and the insects
may carry the pollinia away with them.
Separation of cross-pollinated and

self-pollinated helleborines
r
MNarrow-lippedHelleborine, 12July, Buckinghamshire.
Thisflowerisfreshlyopened,butneverthelesslacksaviscidium,althoughthepolliniaarepresent.
M Narrow-lipped Helleborine, 22 July, Oxfordshire.The

polliniaarevisiblydisintegratingin situ, andthereisno
viscidium.
Four of the Epipactis helleborines are crosspollinated: Marsh, Dark-red, Broad-leaved

and Violet. When the flowers first open they
have neat, undamaged, creamy-yellow pollinia
and a well-formed whitish viscidium. Although
the pollinia are nearly concealed by the anther
cap, they can usually be seen poking out
slightly, like fat yellow sausages. After a short
while it is likely that a visiting wasp will remove
the viscidium and the pollinia, although the
yellow anther cap remains. The flower will
probably have been fertilised by this stage
and the whole column will start to look tatty
and brown.
Narrow-lipped, Dune, Green-flowered and
Lindisfarne Helleborines are self-pollinated.
They have no viscidium (if it is present, it is
almost always small and vanishes before the
flower opens). The pollinia are released by the
anther and sit on top of the column. By the
time the flower opens the pollinia have begun
to swell and crumble, fragments falling over the
edge onto the stigma below. The pollinia are
largely hidden by the anther cap but appear to
foam-out from under it. The flower, having
been fertilised, starts to go-over and the column
turns brown.
29/1/09 12:11:05
GENUS EPIPACTIS
Growth pattern
The aerial stem grows from a rhizome which
puts out numerous roots, each of which can
live for around three years. The growth pattern
of the rhizome is sympodial; the tip of the
rhizome grows upwards to form a flower spike
that withers and dies off once seed has been set,
while the rhizome continues to grow from one
or more buds that are formed at the base of the
aerial stem. These buds are formed at least a
year before the next stem appears above ground
but may remain dormant for one or more years,
in which case the plant remains underground
for a year or more. The probability of a plant
appearing above ground and blooming is
determined by growing conditions over the 12
months previously.
Fungal partners
There is disagreement about the extent to
which adult helleborines obtain nutrients from
fungi. Examination of rhizomes and roots led
to a consensus that some species were free
from fungi, including Violet Helleborine,
which thrives in deep shade. However, several
species occasionally throw up variants that
lack chlorophyll (including var. rosea of Violet
Helleborine). These cannot photosynthesise
but prosper nevertheless, so at least some
helleborines are able to gain substantial
quantities of nutrients from fungi. Recently,
sophisticated techniques have been developed
to identify fungi genetically and to assess their
contribution to the helleborines nutrition. The
evidence produced so far suggests that fungi
do play a very significant role. Several Epipactis,
including Broad-leaved and Dune Helleborines,
display consistent associations with one
group of fungi, the Ascomycetes. Some
Epipactis, such as Broad-leaved and Dark-red
Helleborines, may also have an association
with ectomycorrhizal fungi and therefore gain
nutrients from nearby trees via their fungal
partners, which they cheat.

Poorly known, as the subterranean seedlings
are difficult to find. The seeds have a well-
77
developed outer shell or carapace and this slows

the uptake of water and delays germination,
suggesting that they have a period of dormancy.
Seed probably germinates in spring, forming
first a protocorm and then a mycorhizome (the
earliest and most heavily infected stage in the
development of the rhizome). The first roots
develop in the autumn and, as the rhizome
grows, fungal activity is transferred to the roots
so the rhizome itself becomes infection-free.
This may take place via several mechanisms.
1. The roots develop buds, which go on to
form a secondary rhizome (recorded for Darkred Helleborine). 2. The rhizome produces
two flower spikes that each produce buds and
adventitious roots below ground, and these
buds go on to produce new rhizomes (recorded
for Dune Helleborine). 3. The rhizome
branches (as in Marsh Helleborine). In all these
examples, if the central mother plant dies off
or the rhizome is broken up in some other way,
two or more new plants may result. However,
vegetative reproduction is not recorded for
several of the helleborines and appears to be
relatively unimportant for most species.
Name
Rather obscure. The name Epipactis was
first given to a plant used to curdle milk by
Theophrastus (c. 370-285 BC; often regarded as
the father of botany). It is not certain what he
was referring to, but it may have been a Veratrum,
a genus now known as false helleborines but
formerly called hellebores. When the German
botanist Johann Zinn described the genus
Epipactis in 1757 he may have been struck by
the resemblance of Broad-leaved Helleborine to
a Veratrum, especially the broad, prominently
veined leaves, and therefore chose to use
Theophrastuss name for his new genus.
The word helleborine has been used in
English since the 16th century and means
a plant resembling a hellebore; presumably
again, a reference to its similarity to a Veratrum
(rather than the modern hellebores, genus
Helleborus).
29/1/09 12:11:05
78
GENUS EPIPACTIS
Marsh helleborine
Epipactis palustris
This is one of the most attractive orchids, and when examined closely the individual flowers
are simply stunning, being miniature versions of the gaudy hothouse hybrids. Marsh Helleborine occurs widely across England, Wales and Ireland, but due to its specialised marshy
habitats it is very local. It has declined significantly but can still occur in large numbers,
especially in coastal dune slacks.
Identification
Distinctive. Marsh Helleborine is easily
identified by its colourful, purple and white
flowers, habitat and mid-summer flowering
period. The flowers resemble those of the
other Epipactis helleborines in shape but
are relatively large and bright. The lip shows
striking purple veins at the base, and the tip

has a distinctive frilled edge.
Similar species
None.
Habitat
Marsh Helleborine is found in a wide variety
of wet, marshy habitats but requires neutral
to alkaline ground water and relatively short,
open, vegetation to thrive. The two most typical
habitats are dune slacks and spring-fed fens. In
dune slacks the ground water is calcium-rich
due to the presence of shell fragments in the
sand. In spring-fed fens the ground water is
both nutrient-poor and calcareous and such
fens can be found nestled in heathland valleys
or within more extensive acid bogs, as well
as on more obviously chalk or limestone-rich
soils. Marsh Helleborine may also be found in
meadows which are seasonally flooded with
chalky water, but it cannot compete with tall
vegetation and such habitats must be regularly
mown or grazed for it to survive. Sometimes,
it may be found growing among Common
Reeds, but these, too, are likely to take over and
eventually crowd it out. Marsh Helleborine
occasionally grows in other habitats. These
include wet, slumped, clay cliffs (as in Dorset
and the Isle of Wight), gravel pits and flyash pools. Very occasionally it is found in
small numbers on dry chalk grassland (as in
Bedfordshire, Kent, Surrey and Wiltshire),
especially where quarrying and excavations have
left a compacted surface prone to becoming
O 8July,Norfolk.Growingonclose-croppedturf,itispossibletoseethestructureofthewholeplant.
29/1/09 12:11:06
MARSH HELLEBORINE
waterlogged; it also grows in large numbers on

the floor of an old chalk pit in Norfolk where
there is standing water in the winter months.
79
1987-99
1970-86
pre 1970
Flowering period
Late June to early August, very exceptionally to
early September but mostly in July.
Range
Widespread in England, Wales and Ireland
but often very local and absent from large
areas (much of Kent, Sussex, Surrey,
Wiltshire, Devon, Cornwall, the Midlands,
Co. Durham, Cumbria, mid-Wales, Co. Kerry,
Co. Cork, Co. Waterford and Co. Wexford in
southern Ireland and Co. Antrim in Northern
Ireland). The strongholds are in Norfolk and
Hampshire, but even in these areas it has
declined significantly. Very rare in Scotland
and recently only recorded from a handful of
sites: Perthshire in the Central Highlands,
Argyll and the Inner Hebrides on Islay and
Colonsay. World range: Europe and Asia. In

Europe it is found north to Denmark, southern
Scandinavia and the Baltic States and south to
Portugal, northern Spain, southern Italy, central
Greece, Bulgaria, the Crimea and Caucasus,
M8July,Norfolk.Summerrainhascausedshallowfloodinginthisabandonedquarry.
29/1/09 12:11:08
29/1/09 12:11:11
MARSH HELLEBORINE
and also Corsica and Sicily, but absent from

the Mediterranean lowlands. Also found in
Turkey, northern Iran, Central Asia and extends
eastwards to Lake Baikal in eastern Siberia.
How to find it
This is a showy, conspicuous orchid that often
grows in large numbers, especially in coastal
dune slacks. Notable sites include Lindisfarne
in Northumberland, Sandscale Haws in
Cumbria, Ainsdale in Lancashire, Sandwich
Bay in Kent and Wells-Holkham, Beeston
Common and Buxton Heath in Norfolk.
DESCRIPTION
Height: Usually 20-45cm, occasionally up to
82cm. Dwarf forms, only 10cm high but with
normal-sized flowers, have been recorded,
especially in dry habitats.
Stem: Green, flushed with brownish-purple,
especially towards the tip, prominently hairy,
ridged towards the tip and with one or two
purplish sheaths at the base. Several stems may
grow from the same rhizome.
Leaves: Mid-green, sometimes washed purple
around the base or sheaths, with three to five
prominent veins. Of the four to eight leaves,
ovary
81
sepal
petals
stem
sepal
bract
sepal
hypochile
epichile
lip
the lowest one or two are short, egg-shaped

and formed into a cowl around the stem; the
remainder are large, pointed, broadly strapshaped with a distinct keel, papery in texture
but rather stiff; these are held erect, c. 45-60
above the horizontal, arranged in a spiral
around the stem and grade into narrower and
more bract-like non-sheathing leaves towards
the spike.
Spike: Up to 25 flowers, although usually
rather fewer, form a lax spike; all face more
or less to one side and are initially held
horizontally but slowly droop.
Bract: Green, lanceolate and pointed, the
lowest a little longer than the ovary, the upper
much shorter and also blunter.
Ovary: Brownish-purple, prominently hairy,
O 21July, Norfolk.Aparticularly richly coloured

plant.
P 8 July, Norfolk. The

large boss at the base of
the outer half of the lip
(epichile)andthedelicate
purpleveiningontheinteriorofthecup(hypochile)
areclearlyvisible.
29/1/09 12:11:13
82
GENUS EPIPACTIS
cylindrical, with six fine ribs, narrowing at each

end and tapering to a short, purple stalk that is
twisted through 180.
Flower: Purplish with a white, frilled lip
and prominent purple veins; the flowers
open widely and the lip is held projecting
horizontally outwards. The sepals are ovallanceolate, the lateral sepals asymmetrical.
On the outer surfaces they are sparsely hairy
and dull greenish-yellow, flushed, veined and
mottled purple, especially towards the base;
on the inner face they are more solidly washed
purple, especially towards the tip, with a neat
whitish border. The petals are slightly shorter
and blunter, white, washed pinkish-purple
towards the base on the outer surfaces (shining
through to the inner surface) and also delicately
veined with purple towards the base on the
inner face. The hypochile is formed into a
dish, the base of which is longer than wide and
yellow, variably blotched with reddish nectarproducing swellings. The sides, including the
erect, triangular side-lobes, are white with
prominent purple veins on the inner surface.
The epichile is white with a slight purple wash
towards the edges at the base; it is more or less
circular but with the sides turned upwards
and strongly frilled or crimped. At the base of
the epichile there is an irregular boss which is
whitish with yellow edges and bisected by a
deep, narrow groove; this also produces small
quantities of nectar. The hypochile and epichile
are connected by a narrow, flexible hinge. The
column is very pale yellowish-white with a
well-developed white viscidium, dull yellow
anther cap and primrose yellow pollinia.
Var. albiflora is similar but lacks the purple

veins. It is rare.
Subspecies
BIOLOGY
None.

Var. ochroleuca lacks brown and purple
pigments and is very pallid. The stem and ovary
are green, the sepals yellowish-white to pale
green, and the petals and lip are white, although
the interior of the hypochile still has purple
veins. It is uncommon, but where it is found it
occurs in large numbers.
M 9 July, Norfolk.Var. albiflora.This rare variety lacks

anthocyaninpigmentsandhasagreenstem, ovaryand
sepalsandunmarkedwhitepetalsandlip, lackingeven
purplebanding.

Marsh Helleborine is cross-pollinated but, despite
several investigations, there is disagreement as to
which insects are the most effective pollinators
and what role the unusually flexible outer part of
the lip plays in the mechanism.
The flowers are visited by a wide variety
of insects, including flies and beetles. Some of
them, such as hover flies and honeybees, groom
29/1/09 12:11:14
MARSH HELLEBORINE
themselves extensively and even if they pick up

pollinia are not effective in delivering them to
other flowers. Honeybees also feed their young
on pollen. Solitary bees and wasps are probably
the most efficient pollinators, the nectar on the
large boss on the lip acting as a guide to entice
the insect into assuming the correct position for
pollination. Ants also visit the flowers and may
cause self-pollination, or may carry fragments
of pollinia away and pollinate other flowers on
the same spike (geitonogamy). Self-pollination
may also occur spontaneously as pieces of
pollinia fall onto the stigma below. An average
of over 80% flowers set seed.
Charles Darwin suggested that the outer
part of the lip hinged downwards due to the
weight of the visiting insect, allowing it to
enter the flower without removing the pollinia.
Once the insect was within the hypochile,
however, the epichile hinged back up to its
original position. The insect, as it backed out
of the flower, was therefore forced upwards,
allowing the pollinia to become attached to its
head. Later authors suggested other functions
for the hinged lip, such as causing the insects
to struggle to keep their balance and this being
83
sufficient to bring their heads into contact with

the viscidium.
Vegetative reproduction may occur if the
rhizome breaks up into several sections.

The aerial stems grow from a relatively slender,
well-branched rhizome that creeps horizontally
near the surface of the soil. A single plant may
have an extensive rhizome and produce several
aerial stems. Indeed, it has been claimed that
over 100 flower spikes may grow from the
same plant (Davies et al. 1983). Roots are
produced at many points along the rhizome,
both horizontal roots that penetrate the more
organic surface layers and vertical roots that
often grow deep into the mineral soil. The
roots are reported to have little or no fungal
infection, except in soils deficient in nitrogen.
And indeed, recent isotope studies have shown
that Marsh Helleborine may acquire around
30% of its nitrogen from its fungal partner but
it does not appear to receive any carbohydrates
via that route.
The subterranean seedling stage has never
been observed in the wild and there is no information on the early development of this species.
Hybrids
None.

The specific name palustris means of swampy
ground.
HISTORY aND
CONSERVaTION
M 8July, Norfolk.Theflowerinprofile, showingthetwo

halvesofthelip,connectedbyaflexiblehinge.
The first published record was in the 1633

edition of Gerards Herball, where John Goodyer
recorded it: within a mile ofPeters-field, in a
moist meadow named Wood-mead, neere the
path leading from Peters-field towards Beryton
(a record in Mathias de Lobels Stirpium
Illustrationes (Illustrations of plants) dated from
1601, but this was not published until 1655).
Marsh Helleborine has declined
substantially and is now gone from 60% of its
historical range in Britain and 39% in Ireland.
The decline has affected all areas but perhaps
29/1/09 12:11:15
84
GENUS EPIPACTIS
especially those away from the coast. It is

extinct in the Channel Islands, Bedfordshire,
Huntingdonshire, Northamptonshire,
Worcestershire, Radnorshire, Dumfries and
Galloway, Roxburghshire, Berwickshire, the
Lothians and Fife.
Many of the losses occurred in the 19th
century due to the drainage and destruction
of marshes and fens. Drainage and the subtler
effects of water abstraction continued to cause
losses in the 20th century. More recently,
eutrophication, that is the enrichment of
ground water by fertiliser run-off or even the
discharge of sewage, has caused suitable fens
to become overgrown with more vigorous
vegetation. The abandonment of grazing or
mowing compounds this effect and has led to
the invasion of fens by scrub, in which case
they quickly become too overgrown for the

helleborine to survive.
Marsh Helleborine is specially protected in
Northern Ireland under Schedule 8 of the 1985
Wildlife Order (NI).
PastandpresentoccurrenceofMarshHelleborineinBritainandIreland(basedonpresenceorabsencein10km
squaresoftheNationalGrid;datafromthe New Atlas).
Britain
Ireland
450
160
180 (6.3%*)
98 (9.7%*)
53%
29.5%
% lost, 1970-1986
7%
9.5%
% lost, total
60%
39%

1500-1999
current range
% lost, 1500-1969
O 8July,Norfolk.Arather
dusty-pinkplant.
29/1/09 12:11:17
DARK-RED HELLEBORINE
85
Dark-reD helleborine
Epipactis atrorubens
This orchid is found very locally in open, rocky places in the north and west of Britain and
Ireland and is strongly associated with outcrops of limestone. Whether they are emerging
from the grykes in a limestone pavement or set against a grassy slope, the spikes of reddishpurple flowers and rather dusky-green foliage are very striking. In Europe it is often a
woodland plant, and the absence of trees in our denuded landscape may be a reason for its
very scattered and local distribution in the British Isles. Dark-red Helleborine is the county
flower of Banffshire.
Identification
With its attractive reddish-purple flowers,
Dark-red Helleborine is easy to identify. It has
very obvious yellow anthers and pollinia, and is
cross-pollinated.
Similar species
Broad-leaved Helleborine is the only other
helleborine to occur in the same rocky habitats,
albeit only occasionally. It may sometimes have
rather dark-reddish flowers and, conversely,
Dark-red Helleborine may rarely have paler
pinkish or greenish-red flowers, similar to
those of some Broad-leaved Helleborines.
However, Dark-red Helleborine can always be
distinguished by its leaves, which are darker,
more markedly folded and held in two opposite
rows. It also has larger and rougher bosses on
the lip and a very hairy ovary.
Habitat
Dark-red Helleborine is very strongly associated
with limestone, growing on cliff ledges, scree
slopes, rocky hillsides, in old quarries and in the
shelter of the grykes of limestone pavements.
It is usually found in the immediate vicinity
of bare rock but sometimes also on welldrained grassy slopes with scattered scrub
or even in meadows or on road verges. And,
although most sites are open and sunny, it is
also found in moderate shade on well-wooded
limestone pavements, in open ash woodland
or in pine plantations. Indeed, light woodland
P 17July, Co. Durham.Theleavesareclearlyarranged
intotwooppositeranks.
29/1/09 12:11:18
86
GENUS EPIPACTIS
29/1/09 12:11:20
87
and woodland edges may be the more typical

habitat, as in Europe; most British sites are
deforested and heavily grazed and the species
has a very fragmented, relict distribution. Most
sites are between sea level and 270m, but it
is present at 400m in eastern Cumbria, over
500m on Cronkley Fell in upper Teesdale and
at 610m in Gleann Beag in east Perthshire.
Flowering period
Early June to early August, but mostly late June
to late July.
Range
Very local and scattered. In North Wales
recorded from Caernarvonshire and Flintshire
(including the Great Orme), and in England
found in the Peak District of Derbyshire,
the Yorkshire Dales, in west Lancashire in
the Morecambe Bay area, in Cumbria in the
southern Lake District (around the Kent
estuary and at Hodbarrow), the north Pennines
(upper Eden Valley) and in Co. Durham. In
Scotland found in the eastern Highlands in east
Perthshire, in Banffshire and in the northwest
on the north coast of Sutherland, in west Ross
& Cromarty and on Skye. In Ireland confined
to the Burren region of Co. Clare and Co.
Galway. World range: Primarily a European
species, extending a little way into western
Asia. Occurs north to northernmost Norway
1987-99
1970-86
pre 1970
M 5 July, Lancashire. Dark-red Helleborine will grow in

quitewoodedsituations.
O 5July,Lancashire.Aclassicplantintheclassichabitat,
limestonepavement.
29/1/09 12:11:22
88
GENUS EPIPACTIS
and in Russia to the Arctic Circle, and south

to southern Spain, southern Italy, southern
Greece, Romania and patchily to the Caucasus
but absent from the Mediterranean lowlands.
Also found in western and northern Turkey,
northern Iran, and southern Siberia to
about 85E.
How to find it
Dark-red Helleborine is usually a relatively easy
species to locate when it is growing in the open.
Undoubtedly the best site to see the species
is Bishop Middleton Quarry in Co. Durham
where it is abundant, creating a great spectacle.
DESCRIPTION
Height: 11.5-60cm, sometimes to 100cm.
Stem: Dull green, variably washed purple,
especially towards the base (sometimes entirely
purple), with a dense covering of whitish hairs,
particularly on the upper part of the stem.
There are one to three funnel-shaped basal

sheaths, the uppermost often green towards the
tip. Stems usually grow singly but sometimes
two or three may arise from the same rhizome.
Leaves: Around five to ten, arranged more or
less in two opposite rows towards the base of
the stem. The leaves are dark green, variably
washed reddish-purple on the underside
and are sometimes purple at the base. They
are always distinctly longer than wide (the
lower elongated-oval, the upper tending to be
narrower and more lanceolate), strongly folded
and keeled, and held stiffly at about 30 above
the horizontal.
Spike: Rather lax, with six to 45 flowers
set loosely to one side of the stem. Plants
in sheltered localities tend to have the most
flowers but there are often fewer than ten in
more exposed situations. There is usually a
distinct gap between the uppermost leaf and
the lowermost flower.
M 17July,Co.Durham.Thecontrastingyellowanther-capisverynoticeable;thelarge,roughbossatthebaseofthelip
less so.
29/1/09 12:11:24
upper sepal
anther
petal
petal
pollinia
viscidium
stigma
ovary
sepal
hypochile
(cup)
boss
sepal
epichile
Bract: Lanceolate, green, sometimes washed

purple at the base. The lower bracts may be a
little longer than the flowers but they become
shorter towards the tip of the spike.
Ovary: Pear-shaped, six-ribbed, green, variably
washed purple (sometimes deep blackish-purple)
with abundant short, pale hairs. The flower stalk
is short and greenish to purplish-black.
Flower: Usually a distinctive, rich, wine red. The
flowers often do not open very widely and may
assume a slightly drooping, bell-like shape. The
sepals are oval with slightly pointed tips and
a downy outer surface. The petals are shorter
and broader, more oval-triangular in shape.
Both sepals and petals are deep purple, slightly
paler on the inner surfaces and towards the
base of the petals. The hypochile is dull green
on the outer surface of the cup, becoming rich
purple towards the front, with the interior pale
greenish-white and mottled purple. The epichile
is heart-shaped, broader than long and variably
turned under at the tip. It is rich purple with
two elaborate wrinkled bosses at the base which
sometimes merge into a V or heart shape. The
column is pale greenish-white, flushed purple,
with the anther cap contrastingly yellow with a
narrow brown stripe at the sides; the pollinia are
also yellow. There is a functional viscidium, and
the flowers have a vanilla-like scent.
Subspecies
None.

Occasionally the flowers may be paler red or
even greenish, especially in sites exposed to full
89
sunlight, but normal plants always have some

traces of dark red. Two aberrant varieties have
been named:
Var. albiflora has white or creamy flowers and
has been recorded in the Kishorn area of west
Ross and Cromarty.
Var. lutescens has yellowish or buff flowers and
has been found in Cumbria and The Burren in
Ireland.
BIOLOGY
This species is cross-pollinated. Both the colour
and the scent of the flowers may be important
in attracting suitable pollinators and wasps,
bees and hover flies visit to feed on the nectar,
removing the pollinia in the process.
Vegetative reproduction may take place, new
plants developing from buds on the roots.

The aerial stem grows from a short, thick,
hard rhizome that puts out 40-50 long,
slender, widely spreading roots. These
sometimes form irregular swellings from
which fresh rootlets grow.
The roots of this species are reported to
have a rather limited fungal infection, and it
was thought to be largely phototrophic as a
mature plant, depending on photosynthesis to
supply its nutrients rather than fungi. Recent
isotope studies have shown, however, that Dark
Red Helleborines acquire about 65% of their
nitrogen and 15% of their carbon from fungi.
And, unexpectedly, they have an association
with ectomycorrhizal fungi and thus, like
Coralroot and Birds-nest Orchids, may gain
nutrients from the roots of nearby trees via
these fungi (see p.8). These studies took place
in Europe, where Dark Red Helleborines are
often found in wooded environments. British
and Irish plants may grow well away from any
trees, and the absence of suitable host trees may
be a limiting factor in its distribution in the
British Isles.
There is no information on the length of the
period between germination and flowering.
29/1/09 12:11:24
29/1/09 12:11:28
Hybrids
E. x schmalhausenii, the hybrid with Broadleaved Helleborine, has been reported from
several parts of the range, notably Cumbria. It
is fertile and therefore very difficult to confirm
because infertility cannot be used to distinguish
potential hybrids from plants that are merely
aberrant.
91
Denbighshire, and old records for west

Gloucestershire and Herefordshire are not
always accepted as valid.

The specific name atrorubens means simply very
dark red.
HISTORY aND
CONSERVaTION
1677 in Rays Catalogus Plantarum Angliae
et Insularum adjecentium: On the sides of the
mountains near Malham 4 miles from Settle in
great plenty.
PastandpresentoccurrenceofDark-redHelleborinein
10kmsquaresoftheNationalGrid; datafromtheNew
Atlas).
Britain
Ireland
60
13
42 (1.5%*)
8 (0.8%*)
% lost, 1500-1969
18%
0%
% lost, 1970-1986
12%
38%
% lost, total
30%
38%

1500-1999
current range
Dark-red Helleborine is very local and

absent from many apparently suitable sites
within its restricted range. In Britain it is
Nationally Scarce. Most populations are small,
with many non-flowering plants, and many
plants that would flower are prevented from
doing so by sheep, deer or rabbits. Bishop
Middleton quarry in Co. Durham holds 2,000
or more plants, probably more than all the
other British sites put together.
Some sites have been lost due to quarrying,
and overgrazing threatens others; both
the British and Irish populations may still
be in decline. Extinct in Breconshire and
M 17 July, Co. Durham.The leaves have been largely

grazed off, but nevertheless it is able to flower well,
perhapsaidedbyitsfungalpartner.
O 17July, Co. Durham. Usuallyarisessingly, buttwoor
threespikesgrowingtogetherisnotuncommon.
29/1/09 12:11:30
92
GENUS EPIPACTIS
broaD-leaveD helleborine
Epipactis helleborine
The commonest and most widespread of the helleborines, this species is found in and
around woodland and, in the north and west, sometimes also in more open habitats. But,
although at heart a forest orchid, it is a species that you stumble upon in unexpected places
rather than set out to find; a shady lane, a road verge, an old railway cutting nowhere is too
humble for this adaptable helleborine. It is said to be commoner in the city of Glasgow than
anywhere else in Britain and has spread from coast to coast in North America since it was
introduced in 1879. Youngs Helleborine, described in 1982 and thought to be endemic to
Britain, is now known to be a minor variant of Broad-leaved Helleborine.
Identification
A very variable species. Broad-leaved
Helleborine can be a tall, robust, leafy plant or
a small, weedy specimen with just one or two
flowers. The flowers themselves can be almost
entirely green or almost completely purple
but are usually a mixture of pale green, pink
and purple. It is said that plants in deep shade
tend to be taller and greener whereas those in
sunnier locations are shorter with more red or
purple coloration in the flower, but the variation
often seems to be random.
Typically, Broad-leaved Helleborine has
several relatively large leaves which are a dull,
mid to dark green, sometimes washed purple
but lacking yellow tones. They are usually
obviously veined or pleated and placed all
around the stem, in three rows or in a spiral
pattern, but are sometimes arranged into two
opposite ranks. As the name suggests, the leaves
are broad, especially the lowest, which may be
roughly as wide as they are long. The upper part
of the stem and the ovaries are hairy, although
hairs can be sparse on the ovaries. The outer
part of the lip is heart-shaped, broader than
long and usually turned under at the tip. There
are two bosses at the base of the lip which are
usually brownish and rough or wrinkled but
can be smooth and pink. Importantly, the base
of the flower stalk is washed purple. Broadleaved Helleborine is cross-pollinated; when
O 28July, Norfolk.Atypicalplant, withanopen, rather
sparsely-floweredspike.
29/1/09 12:11:31
BROAD-LEAVED HELLEBORINE
freshly opened, the flowers have an obvious

and functional white viscidium, an important
distinction from Narrow-lipped, Dune and
Green-flowered Helleborines (see p.76 for a
discussion of the separation of cross-pollinated
and self-pollinated helleborines).
Similar species
Violet Helleborine may occur in the same
woods but is typically found in densely shaded
areas. It is usually distinctive. Its leaves are more
greyish-green with a distinctive purple wash
to the undersides, and also narrower, with the
lowest leaf longer than wide. Its flowers are
larger, brighter and cleaner; the sepals and petals
are pale greenish-white, lacking pink or purple
tones, and the lip is whitish with two smoothly
pleated, pink bosses. Like Broad-leaved
Helleborine, it is cross-pollinated.
Narrow-lipped Helleborine is a scarce
inhabitant of southern beechwoods. It has
green flowers, often with a delicate pink wash
to the petals and lip (but not the sepals). As
its name suggests, the tip of the lip is long,
narrow and pointed, and held projecting
prominently outwards. Its leaves are a paler and
more yellowish-green and usually held in two
opposite ranks. In combination these features
should be distinctive but the tip of the lip of
Broad-leaved Helleborine may not always be
reflexed, especially when the flower has just
opened. In Broad-leaved Helleborines with
largely green flowers, this can cause confusion
but the outer part of the lip (epichile) in Broadleaved Helleborine is always broader than long.
In case of any doubt, the base of the flower
stalk is greenish-yellow in Narrow-lipped
Helleborine and the flowers are self-pollinated
and lack an effective viscidium.
Dune Helleborine is only found on
Anglesey, in northern England and in southern
Scotland, usually on dunes but also at inland
sites; conversely Broad-leaved Helleborine is
occasionally found on open dunes or under the
pines that are often planted on coastal sand
hills. In its typical form, Dune Helleborine can
be separated from Broad-leaved by its more
yellowish-green, two-ranked leaves that are
93
held rigidly at about 45 above the horizontal.

Its flowers are smaller, duller and do not
open as widely. The petals and base of the lip
are variably washed pink, but it never shows
pink or purple tones to the sepals or a strong
purple wash on the lip. At inland localities the
variant of Dune Helleborine, known as Tyne
Helleborine, is found. This is very like Narrowlipped Helleborine and can be distinguished
from Broad-leaved by the forward-pointing
tip to its lip and a yellowish-green base to the
flower stalk. Dune Helleborine is normally
self-pollinated.
Green-flowered Helleborine typically has
green flowers which are held drooping and do
not fully open. Sometimes, however, its flowers
may be held more horizontally and may open
widely although they are still predominantly
green, with any pink tones restricted to a
delicate wash on the lip. Whatever, its upper
stem and ovaries are hairless or there are just
a few, sparse hairs on the stem, the base of the
flower stalk is green and it is self-pollinating.
Dark-red Helleborine is usually rather
distinctive but Broad-leaved is occasionally
found in the rocky habitats beloved of Darkred Helleborine and sometimes has rather
dark-reddish flowers, too, while Dark-red
Helleborine rarely has pinkish or greenish-red
flowers. If there is doubt, Dark-red Helleborine
can be distinguished by its leaves, which are
darker, more markedly folded and held in two
M 26July, Norfolk.Arichlycolouredflower.Thepollinia
havealreadybeenremoved.
29/1/09 12:11:33
94
GENUS EPIPACTIS
O 28 July, Norfolk. Two

plantssidebyside, onein
fullflower,theotherstillin
bud.
opposite rows. The bosses on its lip are also

larger and rougher and its ovary is densely hairy.
Habitat
Broad-leaved Helleborine is essentially a plant
of deciduous woodland. It favours the better-lit
areas along paths, rides and roadsides, in glades
and on the woodland fringe, but can grow in
deep shade. Like many of the helleborines it has
an affinity for Beech trees. It can also be found
in suitable shady conditions in scrub, along
well-grown hedges, banks, disused railways and
stream-sides. Broad-leaved Helleborine will

also grow in the open, on limestone pavements,
cliffs, scree and grassland, but only in the
cooler and damper conditions of the north and
west. In Ireland and south Wales it is found
in dune slacks. One of the most adaptable
of the helleborines, it is an opportunist
and can colonise newly available habitats,
such as mature birch scrub on spoil heaps,
willow and alder carr and conifer plantations
(especially where conifers have replaced
ancient woodland). In Glasgow and a few
29/1/09 12:11:35
1987-99
1970-86
pre 1970
other cities in southern Scotland and northeast

England, Broad-leaved Helleborine has
moved into mature gardens, as well as parks,
cemeteries, golf courses, playing fields, rubbish
tips, roadsides and railway embankments. It
is comparatively tolerant of soil pH and will
grow in slightly acid conditions but is usually
commonest on calcareous soils. Recorded up to
350m above sea level (Ystradfellte, Breconshire).
Flowering period
Early July to early September but mostly
from mid-July to mid-August. Like all the
helleborines, it flowers earlier and more sparsely
in dry summers.
95
Armagh, Co. Derry and Co. Antrim. World

range: Very widespread in Europe and
Asia and introduced to North America. In
Europe it occurs north to c. 71N in Norway
and to southern Finland, and south to the
Mediterranean, Crimea and Caucasus. It is
absent from the Mediterranean lowlands but
found on the Balearic Islands, Corsica, Sardinia
and Sicily. Also scattered in Turkey, Lebanon
and Israel. In Asia it also ranges through
southern Siberia east to Lake Baikal and south
through the mountains of Central Asia to the
Himalayas. Populations from North Africa and
the Far East are often now treated as different
species. Introduced to North America, perhaps
deliberately as a remedy for gout. It was first
recorded in 1879 near Syracuse, New York,
and is now widespread; it reached California by
1950 and is still spreading.
How to find it
The commonest of the helleborines but nevertheless often rather local, it is best looked for
in and around undisturbed ancient woodland,
especially along shady roads and tracks passing
through suitable habitats. It is frequently robust
but despite this the relatively small, dull flowers
can be surprisingly hard to see when it is
growing among brambles or Bracken.
DESCRIPTION
upper sepal
Range
Widespread in England and Wales, although
rather local in some areas, such as Norfolk,
Suffolk, Cambridgeshire and Lincolnshire, and
absent from west Cornwall and some upland
areas in northern England. In Scotland it is
well distributed in the Central Lowlands and
scattered in the Borders but to the north is
extremely local, with a few sites in southern
Perthshire, Angus, Banffshire, Sutherland, west
Ross & Cromarty, Argyll and Kintyre, and on
the Inner Hebrides on Skye. Widespread in
Ireland but mostly found in the west and the
north and rather local away from The Burren
(Co. Clare and Co. Galway), Co. Fermanagh
and the region around Lough Neagh in Co.
anther
petal
petal
pollinia
viscidium
stigma
ovary
sepal
hypochile
(cup)
boss
sepal
epichile
Height: 10-120cm but usually around 25-80cm.

Stem: Pale green, often washed purplish
towards the base, with short, whitish hairs on
the upper portion and two or three leafless
sheaths at the extreme base. Usually grows
29/1/09 12:11:36
96
GENUS EPIPACTIS
M 1August, Northumberland. A spike of rather purple

flowers.Broad-leavedHelleborines,wheninspectedclosely,alltoooftenappearratherdishevelled.
singly but clumps of two or three are quite

frequent and five or more may occasionally
grow from the same rhizome. Non-flowering
stems are frequently produced.
Leaves: Four to ten, more-or-less spirally
arranged around the stem. They are oval to
oval-lanceolate and less than twice as long
as wide; the lowest leaf is sometimes almost
rounded, but they become narrower and more
bract-like towards the flower spike. The leaves
are dull, mid to dark green, sometimes tinged
purple, with several prominent veins.
Spike: Up to 60 flowers, rarely as many as 100,
are arranged into a roughly one-sided spike that
is sometimes very dense but on other plants
rather lax.
Bract: Dull green, lanceolate, with a pointed
tip. The lowest are significantly longer than the
flower but they become shorter, roughly the
length of the ovary, towards the tip of the spike.
Ovary: Green, hairless or with a few short
hairs, boldly six-ribbed and rather pear-shaped,
tapering into a long, twisted stalk which is
washed purple at the base.
Flower: Very variable in colour but usually
with some purple tones and opening widely.
The sepals are oval, tapering to a point, with
their outer surface green mottled with variable
amounts of dull purple or pink (sometimes
none) and a prominent green midrib; their
inner face varies from pale green to dull, dusty
purple. The petals are similar but slightly
shorter and less tapering and tend to be paler,
cleaner and often pinker. They vary from pale,
dusty pink to purple and are often whiter
towards the centre; they also have a green
midrib on the outer face. The hypochile is pale
greenish-white, variably washed pink or purple,
with the interior of the cup purple to midbrown, glistening with nectar. The epichile is
heart-shaped, broader than long, with the major
part strongly curled down and under. There are
two bosses at the base of the epichile, usually
purplish-brown and wrinkled, but they may be
green or pink, and are sometimes smooth. The
epichile varies in colour from dull greenishwhite, washed pink, to pale pink or dull purple.
29/1/09 12:11:38
97
The column is greenish-white, the anther cap

dull pale yellow with brown stripes at the side,
and the pollinia are creamy-yellow. There is an
obvious and functional white viscidium. The
flowers usually have no scent.
Subspecies
None recognised.

Var. youngiana Youngs Helleborine
Youngs Helleborine was described as a new
species, E. youngiana, in 1982 from plants
found in Northumberland. It was subsequently
identified in Scotland and, more controversially,
in Yorkshire and south Wales. It was reported
to differ from Broad-leaved Helleborine in a
number of ways. Its wavy-edged, more-or-less
two-ranked leaves are on average paler and more
yellowish-green. The lowest leaf is usually longer
than wide, rather than wider than long. Its ovary
is sparsely hairy to hairless (only rarely hairless in
typical Broad-leaved Helleborines). The flowers
are similar to some Broad-leaved Helleborines
but relatively large, clean and bright. The critical
difference, however, was to be found in the
reproductive structures; in Youngs Helleborine
the viscidium is very small and disappears
rapidly. The pollinia remain in the flower
and disintegrate onto the stigma causing selfpollination. The column has a long rostellum (as
long as the anther cap) which, together with two
pointed bosses at the base of the stigma, forms a
distinctive three-horned shape.
Some botanists were, however, always
sceptical about the distinctiveness of Youngs
Helleborine and recent genetic studies have
shown that it does not exist as a distinct
entity. At each site investigated, the Youngs
Helleborines were genetically closer to the
local Broad-leaved Helleborines than they
were to Youngs Helleborines at other sites.
Also, rather than being self-pollinated, it has a
high level of genetic diversity and a population
genetic structure that indicates that it is crosspollinated. Plants matching the description of
Youngs Helleborine in terms of their leaves,
ovaries and flowers are found at its classic sites
M 1August,Northumberland.AYoungstype,withrather
yellowish-greenleavesandbrightflowers.
29/1/09 12:11:39
29/1/09 12:11:42
(for example, Settlingstones in Northumberland)

but almost all appear to have a large and fully
functional viscidium and are presumably crosspollinated, supporting the genetic studies. With
the claimed differences in reproductive biology
gone, the remaining distinctions between
Youngs and Broad-leaved Helleborine are very
subtle and it is hard to justify even the status
of variety for Youngs, especially given the wide
variation in Broad-leaved.
Var. neerlandica Dutch Helleborine Overall
deep green and rather short (15-40cm), with
short, stiff, rounded leaves that are held moreor-less erect and grouped at the base of the
stem, which they closely sheathe. The leaves
have a border of tiny teeth that are irregular
and fused at the base (use a 20x hand-lens;
typical Broad-leaved Helleborines have more
regular teeth). The spike is dense and the
flowers are dull purplish-pink, bell-shaped and
do not open widely. Found along the coast of
the North Sea from Pas-de-Calais in northeast
France to Denmark and on the Baltic coast
of northern Germany; similar plants growing
amongst Creeping Willow in the dune slacks
of south Wales have been identified as this
form. Although it is treated as a distinct species,
E. neerlandica, by some Continental authors,
genetic studies reveal little difference between it
and typical Broad-leaved Helleborines.
Var. monotropoides (also known as var. albifolia)
lacks chlorophyll and is pale pink or strawcoloured with white or rosy flowers. It is very rare.
Var. viridiflora lacks anthocyanins and has pale
green flowers with a greenish-white lip and
shows no trace of red or purple. It is rare.
Var. purpurea has especially dark purple or
reddish-violet flowers. It is rare.
BIOLOGY
Broad-leaved Helleborine is pollinated by
wasps, especially long-headed species of
the genus Dolichovespula. Wasps have short
O 28July,Norfolk.Abeautiful,richly-colouredplantwith
wine red petals and lip.The pollinia have already been
removedfromalltheflowers.
99
mouthparts but can nevertheless easily reach

the nectar produced in the hypochile; in the
process they rupture the viscidium and the
pollinia are stuck to their heads. Other insects,
including short-headed wasps, bees, hover
flies and beetles, may visit the flowers but
are the wrong size or shape to act as efficient
pollinators. Fermentation of the nectar in the
helleborines flowers may produce ethanol and
this can have a narcotic effect on visiting wasps,
which become slow and sluggish and may even
fall to the ground drunk.
The flowers are self-compatible and are
frequently pollinated by wasps carrying
pollinia from flowers of the same spike (i.e.
geitonogamy). It is sometimes stated that
Broad-leaved Helleborine may be self-pollinated
(i.e. fertilised by pollen from the same flower).
The evidence for this is contradictory but
experiments have shown that in normal
circumstances self-pollination either does not
occur or only takes place rarely when small
insects carry pieces of pollinia onto the stigma
below. It has also been reported that in drought
conditions the flowers shrivel without opening
and may be cleistogamous and self-pollinate in
the bud (Ettlinger 1997). Seed-set is usually
good and almost all the flowers may produce
ripe capsules, each containing up to 3,000 seeds.

The aerial stem grows from a small, woody
rhizome which sends numerous cordlike
roots deep into the soil. The degree of fungal
infection of the roots is reported to vary, being
high in plants growing in humus-rich soils
and negligible in mineral soils, but at least
some Broad-leaved Helleborines are heavily
dependent on their fungal partner. Recent
isotope studies have shown that they acquire
around 60% of their nitrogen and about 14%
of their carbon via fungi. Unexpectedly, Broadleaved Helleborine has an association with
ectomycorrhizal fungi and thus, like Coralroot
and Birds-nest Orchids, may gain nutrients
from the roots of nearby trees via these fungi
(see p.8). It is probably not very fussy about its
29/1/09 12:11:42
100
GENUS EPIPACTIS
Scotland and confirmed by genetic analysis.

This hybrid has no scientific name.

The specific name helleborine means like
a hellebore. The leaves of Broad-leaved
Helleborine do indeed look like those of
Veratrum album, a plant that was once called
White Hellebore but is now, rather ironically,
known as White False Helleborine. (Broadleaved Helleborine bears no resemblance to the
plants now called hellebores, quite unrelated
flowers belonging to the buttercup family.)
M 28 July, Norfolk.A very pallid plant, with just a hint
of purplish-pink in the sepals and lip.The pollinia and
viscidiumhavealreadybeenremoved.
fungal partners but might have a preference

for Ascomycetes, including ectomycorrhizal
fungi of the genus Tuber, better known by their
English name truffle.
Broad-leaved Helleborine, presumably
supplied by its fungal partner, may spend a
significant proportion of the time underground.
Plants may flower and then spend one or, rather
less often, two or even three years dormant
before appearing again. In a study in America,
25%-50% of the population appeared above
ground each year and around a third of these
flowered. Very few plants flowered every year,
however, although annual flowering may be more
frequent where the soil is reasonably moist.
The interval between germination and
flowering can be as little as 18 months, although
periods of eight or nine years, including several
years above ground as a non-flowering plant, are
also quoted.
Hybrids
E. x schmalhausenii, the hybrid with Dark-red
Helleborine, has been reported from several
parts of the range, notably Cumbria. It is fertile
and very difficult to confirm.
E. x schulzei, the hybrid with Violet Helleborine,
has been recorded quite frequently but, like
E. x schmalhausenii, is fertile and hard to
confirm.
x Dune Helleborine has been found in
HISTORY aND
CONSERVaTION
The first British record was in 1562 when
William Turner stated in his Herball: I have
seen itin England in Soffock.
PastandpresentoccurrenceofBroad-leavedHelleborinein
BritainandIreland(basedonpresenceorabsencein10km
squaresoftheNationalGrid;datafromtheNew Atlas).
Britain
Ireland
1,218
161
840 (29.5%*)
107 (9.7%*)
% lost, 1500-1969
19%
24%
% lost, 1970-1986
12%
9.5%
% lost, total
31%
33.5%

1500-1999
current range
The New Atlas states that the overall

distribution is stable but although the
boundaries of the range may be largely
unchanged there has been a significant decline,
with the loss of around a third of the historical
range. Losses have been concentrated in the
Midlands and northern England, where the
distribution is now rather fragmented, and
more recently in the Home Counties. The
clearance or coniferisation of woodland,
increase in dense shade due to a lack of
woodland management, ground disturbance by
machinery and horses, and grazing by deer may
all have contributed to the decline, and losses
seem to be ongoing.
29/1/09 12:11:44
VIOLET HELLEBORINE
101
violet helleborine
Epipactis purpurata
Thought to be a long-lived orchid, older and more mature plants may produce several flowering spikes from the same rootstock, and a group of Violet Helleborines blooming in the
cathedral-like gloom of a late summer beechwood certainly makes a dramatic sight. Among
the last orchids to come into flower, it is endemic to western and central Europe and in Britain
is confined to the southern half of England, but it is always rather local and uncommon.
Identification
Relatively distinctive. The flowers are pale green
or whitish with the lip faintly washed pink
and bearing two pink bosses; the petals and
sepals spread widely and the large, pale flowers
contrast strongly with the dark purplish stem
and the rather small, dark leaves. It is crosspollinated, and the flowers have an obvious
and functional viscidium, a useful distinction
from Narrow-lipped and Green-flowered
Helleborines (see p.76 for a discussion of
the separation of cross-pollinated and selfpollinated helleborines).
Similar species
Broad-leaved Helleborine is much commoner
than Violet Helleborine. It has broader leaves,
with the lowest more-or-less wider than long
(longer than wide in Violet Helleborine).
Its leaves are also a cleaner and brighter
green (duller, more greyish-green in Violet
Helleborine, with a distinctive purplish wash
to the underside). Its flowers are smaller and
often duller and darker with a purplish wash,
with the bosses on the lip usually rougher and
browner. In Violet Helleborine the flowers are
cleaner, brighter and paler, with two smoothly
pleated, pink bosses. Like Violet Helleborine, it
is cross-pollinated.
Narrow-lipped Helleborine is also found
in densely shaded beechwoods in southern
England but is easily separated by its long,
pointed lip and much paler stem, flower stalks
and leaves.
P 11August, Suffolk. Summer 2003 was hot and dry,
andthehelleborinesweresmallandfewinnumber.
29/1/09 12:11:45
102
GENUS EPIPACTIS
woodland relicts and in wild gardens. Although

it can tolerate acid soils, it is strongly associated
with areas of calcareous bedrock, especially
chalk, and its deep root system requires a
substantial thickness of soil. It can grow on
sands and gravels but is particularly associated
with clays, especially the clay-with-flints found
on plateaus and hilltops overlying chalk. Within
woods, it is often found in areas of deep shade
where little else can grow; it can flourish in
much darker situations than Broad-leaved
Helleborine but may share its dimly-lit haunts
with Narrow-lipped Helleborine.
Flowering period
Mid-July to early September, exceptionally late
June to late September, but typically peaking in
early August. It tends to flower earliest in more
open situations and will be especially early in a
dry season.
Range
Found in southeast England (including the Isle
of Wight) and the Midlands, occurring west
to Dorset, north Somerset, Gloucestershire,
Herefordshire and Shropshire, and north
to south Staffordshire, Leicestershire, south
Lincolnshire, Huntingdonshire, Hertfordshire
and southern Suffolk. Also recorded from
Denbighshire. The species is commonest in
Kent, Surrey and the Chilterns. World range:
M 11August,Suffolk.Growingonarelativelybarewoodlandfloorinoldhazelcoppice,atypicallyshadyspot.
1987-99
1970-86
pre 1970
Green-flowered Helleborine may be in

bloom late in the summer, like Violet Helleborine, but is easily separated by its more-or-less
hairless, green stem and ovaries, green flower
stalks and rather smaller flowers which are
usually held drooping and seldom open widely.
Habitat
Violet Helleborine is very much a woodland
orchid, favouring beech, hornbeam and oak
woods as well as overgrown hazel coppice. It
is occasionally found in hedgerows that are
29/1/09 12:11:47
VIOLET HELLEBORINE
103
Confined to western and central Europe where

it occurs north to Denmark, west to France (to
a line between Mont St Michel and Grenoble),
south to the Alps, Balkans and Transylvanian
Alps in Romania, and east to the Baltic States.
How to find it
Despite its size and showy flowers, this can be
a tricky species to find. Like other helleborines,
it is loosely colonial, and although the colonies
sometimes grow alongside paths and roads or
on the edges of glades and clearings, they are
equally at home in deep shade and can easily
be missed. It is best looked for in wet seasons,
as the species is affected by drought and in dry
summers the size and number of spikes may be
greatly reduced.
DESCRIPTION
upper sepal
anther
petal
petal
pollinia
viscidium
stigma
ovary
sepal
hypochile
(cup)
boss
sepal
epichile
Height: 20-90cm but usually less than 70cm.

Stem: Greyish-green, variably but often heavily
washed purple, with dense, short, grey hairs
on the upper part of the stem and one to three
small, purplish-brown sheathing scales at the
base (the uppermost scale often tipped green).
Stems usually grow singly but multiple stems
are fairly common, groups of six to eight not
unusual, and a cluster of 38 has been recorded.
Leaves: Well-spaced up the stem and arranged
spirally (or sometimes in two opposite rows),
the four to 14 leaves are relatively small, moreor-less oval in shape and taper to a point. They
are usually rather more than twice as long as
wide, with the upper leaves narrower and more
bract-like and the lowest leaf short and cowl-
M 10August, Suffolk. Of all the woodland helleborines,

Violethasthebrightestandcleanestflowers.
like. The leaf posture is variable; they may be

held horizontally with the tips slightly drooping
or at about 30 above the horizontal. They are
a rather dull, cold, greyish-green and may be
washed purple towards the tips. The undersides
have a diagnostic purple wash and the leaf
sheaths are also frequently tinged purple.
Spike: Slightly to moderately one-sided, there
are usually seven to 40 flowers but some wellgrown plants can carry over 100 blooms.
Bract: Green, variably washed purple. The
bracts are narrow, lanceolate, held roughly
horizontally and are longer than the flower in
the lower part of the spike, becoming shorter in
the upper part.
Ovary: Green, with six prominent ribs, which
29/1/09 12:11:48
104
GENUS EPIPACTIS
29/1/09 12:11:50
VIOLET HELLEBORINE
may be washed purple, and sparsely hairy. The

flower stalk is purplish and variably twisted.
Flower: Rather large, opening widely, and
overall greenish-white with a pinker lip. The
sepals are triangular-oval, rather large and
pale green, becoming paler towards the edges
and with a prominent green midrib on the
outer surface. The petals are smaller, whitish,
becoming slightly greener towards the centre,
with a fine green midrib. The hypochile is
translucent-whitish, slightly greener towards
the base of the cup, with the interior variably
washed pale purplish-rose to pale brown or
pale greenish. This colour shines through to
the outside. The epichile is short, triangular
or heart-shaped with the tip strongly folded
downwards. It is whitish with two prominent,
smoothly pleated, pink bosses at the base. The
column is whitish and the large, conspicuous
anther cap is very pale yellowish-white
with narrow brown stripes at the sides. The
viscidium is whitish and the pollinia are pale
yellow. The flowers are faintly scented.
Subspecies
105

Violet Helleborine grows from a rhizome
which lies more-or-less vertically in the soil
and has up to 50 fleshy roots, each up to 70cm
long (exceptionally 120cm), growing vertically
downwards. It is reported that the roots are
fungus-free and therefore the mature plant is
phototrophic, depending on photosynthesis
rather than fungi for nutrition. However, given
the dense shade in which it grows, it seems
much more likely that fungi contribute a large
part of its nutritional budget; the rare var. rosea
lacks chlorophyll but nevertheless is able to
flower and fruit successfully and must depend
entirely on fungi.
Violet Helleborine is long-lived and only
appears above ground when mature enough to
flower. Immature, non-flowering plants are very
rarely seen. A single-stemmed plant may be 30
years old, and it has been suggested that large,
multi-stemmed plants are probably hundreds of
years old.
There is no information on the duration of
the period between germination and flowering.
None.
Hybrids
E. x schulzei, the hybrid with Broad-leaved

Helleborine, has been recorded widely but this
hybrid is fertile and therefore very difficult to
confirm. (Infertility and the failure to produce
viable pollen and seed are a standard means of
confirming a hybrid.)
Plants may sometimes have variegated leaves,

and some may have the leaves very extensively
streaked violet. One variety has been named:
Var. rosea is rare but stunning. It lacks
chlorophyll and the entire plant is a rosy pink
with whitish flowers.
BIOLOGY
Routinely cross-pollinated, often by wasps
which are attracted to the nectar. As with
Broad-leaved Helleborine, the nectar is
reported to have a narcotic effect, with drunken
wasps falling to the ground. Pollination is
efficient and most or all of the flowers on a
spike will set seed. There are no reports of
vegetative reproduction.
O 11August,Suffolk.Amulti-stemmedplant(therecord
is38spikesgrowingtogether).

The specific name purpurata means purplish.
HISTORY aND
CONSERVaTION
This species was discovered by Rev. Dr. Abbot
parasitical on the stump of a maple or hazel
in a wood near the Noris farm at Leigh,
Worcestershire, in 1807 and was described
scientifically from English specimens in 1828
by Sir J.E. Smith in The English Flora. Notably,
the type specimen was probably the rare
variant rosea.
Violet Helleborine has declined steadily over
the last 150 years as ancient woodlands have been
29/1/09 12:11:51
106
GENUS EPIPACTIS
destroyed or replanted with conifers, although

most sites should now be safe from this particular
threat. It has vanished from the edge of the range
and is extinct in Devon, Cambridgeshire, Norfolk
and southwest Yorkshire, and is reduced to one
site in south Lincolnshire.
Favouring the dense shade of closed-canopy
woodland, this is perhaps the only species of
orchid to have benefited from the abandonment
of coppicing in many British woods during
the 20th century. Conversely, the great
storms of 1987 and 1990 devastated many
woods, opening up the canopy and leading
to a great reduction in the numbers of Violet
Helleborines in affected areas. Another threat
is deer, and whole populations of orchids can
be grazed off in some woods. The species can

be very persistent, however, and has even been
recorded pushing its way through newly
laid tarmac.
PastandpresentoccurrenceofVioletHelleborineinBritainandIreland(basedonpresenceorabsencein10km
total historical
range, 1500-1999
current range
Britain
Ireland
235
145 (5%*)
% lost, 1500-1969
28.5%
% lost, 1970-1986
10%
% lost, total
38.5%
O 10 August, Suffolk.
Violet Helleborine is
cross-pollinated,mainlyby
wasps.
29/1/09 12:11:53
NARROW-LIPPED HELLEBORINE
107
narrow-lippeD helleborine
Epipactis leptochila
Data Deficient
Other names: E. muelleri

This enigmatic orchid is confined to southern England with an outpost in south Wales. It
is a characteristic species of the beechwoods of the Cotswolds and Chilterns but is scarce
and local and has declined markedly in recent years. For a while, a case of mistaken identity
allayed fears about this decline, but it is now clear that this surprisingly attractive helleborine
is in need of friends.
Identification
Narrow-lipped Helleborine looks almost
uniformly green and has relatively large, clean,
pale-green flowers with purplish-pink confined
to a delicate wash around the base of the lip
and a variably obvious tinge on the petals. The
flowers are held drooping and the tip of the
lip projects forward rather than being turned
under as in most other helleborines; the lip
therefore appears long and pointed. The upper
stem and ovaries are hairy (use a hand-lens to
check this), and the base of the flower stalk is
greenish-yellow. The leaves are usually carried
in two opposite ranks and are fresh green,
sometimes tinged with yellow. The flowers
are self-pollinated and lack a viscidium, the
pollinia crumbling apart where they lie and
falling piecemeal onto the stigma. This is a
useful distinction from Broad-leaved and Violet
Helleborines (see p.76 for a discussion of the
separation of cross-pollinated and self-pollinated
helleborines). Rarely, when freshly open, the
flowers of Narrow-lipped Helleborine can be
temporarily cross-pollinating (facultatively
allogamous), with functional viscidia and intact,
cohesive pollinia; such plants must be identified
as Narrow-lipped Helleborine with caution.
Similar species
Broad-leaved, Violet and Green-flowered
Helleborines may be found in the same woods
although only Violet Helleborine is frequent
P 22 July, Oxfordshire. This can be a very stately
helleborinebutalltoooftenitsflowerspikesaregrazed
offbydeer.
29/1/09 12:11:54
108
GENUS EPIPACTIS
in the deep shade beloved of Narrow-lipped

Helleborine. Broad-leaved Helleborine can
have largely green flowers, but they are usually
extensively washed with dull pink or purple,
and the tip of the lip is almost always turned
down and backwards to give the lip a very
short, blunt-ended appearance. Even if the
lip is not reflexed, the heart-shaped epichile
is always broader than long. The base of the
flower stalk is tinged with purple, and, in most
Broad-leaved Helleborines, the leaves are a
darker and duller green and are usually carried
spirally around the stem. Violet Helleborine
has flowers that are closer in coloration to
Narrow-lipped, but they are held more erect
and face outwards, giving the spike a different
character. The flower stalk is purple and the
leaves are a far duller greyish-green and often
have a faint purple wash on the underside.
Green-flowered Helleborine is selfpollinated, like Narrow-lipped Helleborine,
and can be rather similar to it (especially var.
vectensis of Green-flowered), with drooping,
bell-like green flowers and a green flower stalk.
1987-99
1970-86
pre 1970
However, it has a hairless, or almost hairless,

upper stem and ovaries, and the interior of
the cup at the base of the lip is pale greenish.
In many cases, the flowers of Green-flowered
Helleborine are very distinctive as they often
hang down almost vertically and in some
populations hardly open at all.
M 22July,Oxfordshire.GrowingonatypicallybarewoodlandfloorbelowBeeches.
29/1/09 12:11:56
109
Habitat
Narrow-lipped Helleborine is always found on
calcareous soils derived from chalk or limestone,
especially on the steeper slopes where the soil is
very thin and skeletal. It is strictly a woodland
orchid and usually to be found in ancient
woodland. The classic habitat is a beechwood
on chalk, sometimes with a mixture of Yew, but
occasionally it is found under a variety of other
deciduous trees, including overgrown
ash-hazel coppice. Whatever the type of
woodland, Narrow-lipped Helleborines will be
found in areas of deep shade where the ground
cover is sparse or absent. It is intolerant of
direct sunlight.
Flowering period
Rather short, from the second week in July to
mid-August (exceptionally from late June) but
mostly in the last half of July.
Range
Highly localised. The strongholds are the
Cotswolds in Gloucestershire and the
Chilterns in Berkshire, Oxfordshire and
Buckinghamshire. Away from these areas
it is rare with just a few widely scattered
populations in Surrey, Hampshire, Dorset,
south Wiltshire, north Somerset, Shropshire
and Glamorganshire. The species is inconspicuous, however, and could still be found at
new localities. Delforge (1995) reported the
presence of an unidentified helleborine of the
leptochila group from the Burren in Co. Clare.
World range: Although once thought to be
endemic to Britain, the species has now been
found in Europe, north to Denmark, east to
Slovakia, Hungary and Yugoslavia, and south to
Italy and the Pyrenees.
How to find it
With a restricted distribution, specialised
habitat and short flowering period, this can
be a hard species to find. It is also superficially
rather anonymous, although well-grown plants
are very attractive. The apparent size of colonies
can vary dramatically from year to year but the
underground population may be much more
stable and rather larger. One of the places to see
M 12 July, Buckinghamshire. This species has largely

green flowers but with a pinkish-purple tinge to the lip
andsometimesalsothepetals.
the species at its best is the Warburg Reserve

at Bix Bottom in Oxfordshire where a strong
population is protected from deer.
DESCRIPTION
upper sepal
petal
anther
petal
pollinia
ovary
stigma
sepal
hypochile
boss
(cup)
epichile
sepal

Stem: Green, with the upper part hairy. Usually
grows singly but sometimes there are two
together and rarely up to five or six stems may
arise from a single rootstock.
Leaves: Three to seven, fresh green and arranged
in two opposite rows up the stem, although
29/1/09 12:11:57
110
GENUS EPIPACTIS
M 22July,Oxfordshire.Aparticularlycrowdedspikeona
robustplant.
sometimes not obviously so. The leaves are

elliptical and mostly more than twice as long
as wide, with the uppermost long, narrow and
grading into the lowest bracts. They are rather
floppy and held more-or-less horizontally but
the uppermost, bract-like leaves are pendant.
Spike: Usually rather lax, with four to 35
flowers, often facing to one side. Initially held
horizontally, they droop to a variable extent as
they age.
Bract: Pale green and lanceolate; in the lower

part of the spike the bracts are very long,
project well beyond the flower and often hang
downwards, but towards its tip they are shorter.
Ovary: Pale green, variably hairy, prominently
six-ribbed but not twisted; the flower stalk is
greenish-yellow.
Flower: Overall greenish, relatively large
and usually opening fairly widely, although
occasionally some flowers may not open fully,
especially towards the tip of the spike. The
sepals are oval-triangular and elongated into
pointed tips, pale green on the outer surface
and slightly paler and more whitish-green
on the inner. The petals are similar in shape
but slightly smaller. They are also paler, more
greenish-white, becoming even paler towards
the edges but greener towards the midrib
(which is prominent on the outer face) and
variably flushed pale pink with faint pink veins.
The hypochile is whitish-green on the outside
of the cup, variably flushed purplish-pink at
the sides, and similarly pale on the inside but
with a wine-red or chocolate-brown rear wall;
it contains nectar. The epichile is whitish-green,
sometimes delicately flushed pink, shaped like
an arrowhead, longer than wide, and its pointed
tip projects outwards. It has two relatively
small, smooth bosses at its base and these are
washed purple or purplish-pink; they flank
a longitudinal central groove connecting the
hypochile and epichile. The column is greenishwhite, the pollinia creamy-yellow and the anther
cap pale greenish-yellow with a narrow cream
and broad chocolate-brown stripe on either
side; it is attached to the column at the rear by
a projecting spur or stalk (and in profile looks a
little like the leaping jaguar motif of the famous
sports car, a distinction from Broad-leaved
Helleborine where the anther is unstalked). The
rostellum is reduced in size (less than half as
long as the anther) and the viscidium, although
present in the bud, has almost always withered
by the time the flower opens.
Subspecies
None.
29/1/09 12:11:58
111
lacking chlorophyll, with a lemon-yellow stem

and flowers, and pale greenish-yellow leaves,
has been recorded (Young 1962b). It apparently
thrived and flowered. This observation, together
with the dense shade of its normal habitats,
suggests that Narrow-lipped Helleborine must
be able to acquire a large proportion of its
nutrients from fungi.
Hybrids
None. (E. x stephensonii, the hybrid with Broadleaved Helleborine, has been reported, but not
confirmed, in southern England.)

M 22July, Buckinghamshire. Showingthepointedtipto
thelip,whichisnotbentbackunderneaththeflower,and
therelativelysmall,smoothbossesatitbase.

Var. cleistogama was described from plants
found in Gloucestershire on the steep western
escarpment of the Cotswolds near Wottonunder-Edge, but this population is apparently
now extinct. It had pendulous bracts, flowers
that did not open and, within the flower, a
greener lip. It has been treated as a distinct
species but the upper flowers of some normal
plants may never open, and it seems much more
likely to have been a minor variant.
Var. cordata is poorly-known. It was described in
1950 as having smaller, more bell-shaped flowers
with a broader lip and less spreading sepals.
BIOLOGY
Narrow-lipped Helleborine is routinely selfpollinated but occasionally the flowers may
have a functional viscidium. Visiting insects will
then carry away the pollinia, and some crosspollination is therefore possible. Seed-set is
good and each capsule forms 1,000-2,000 seeds.

The rootstock lies rather deeply in the soil
and numerous roots grow from it. Otherwise,
there is no information on this species. A plant
The specific name leptochila means with a

slender lip.
Initially confused with variants of Broadleaved Helleborine, it was not until 1921 that
Narrow-leaved Helleborine was described as
a distinct species. It was then confused with
Green-flowered Helleborine until the British
botanist Donald Young resolved the identity
and characteristics of the two species in the
1950s and early 1960s. Confusion then arose
again in the mid-1970s when plants resembling
Narrow-lipped Helleborine were found inland
in northern England (the Tyne Helleborine, see
p.97). In some places these Tyne Helleborines
were found together with Dune Helleborines
and intermediates were reported, leading
to the conclusion that Narrow-lipped and
Dune Helleborines must be one and the same
species. However, more recent genetic studies
have shown that all the plants in northern
England are Dune Helleborines, and that they
are genetically distinct from the true Narrowlipped Helleborine, which is once again
restricted to its classic habitat, the southern
beechwoods.
Narrow-lipped Helleborine is one of a
group of self-pollinating helleborines found in
Europe, all of which have evolved independently
from within the cross-pollinated Broad-leaved
Helleborine complex. Being self-pollinated,
they have a restricted genetic diversity but
some, such as Narrow-lipped Helleborine, are
successful species which have spread widely.
29/1/09 12:12:00
112
GENUS EPIPACTIS
These self-pollinating species have long caused

problems for both taxonomists and botanists,
and their identification and correct classification
is only now being resolved by genetic and
biochemical techniques. For example, it has
been suggested that Narrow-lipped and Dune
Helleborines are the same as Epipactis muelleri
of Continental Europe (most recently by Stace
2004). Recent genetic research shows that this
is not the case.
HISTORY aND
CONSERVaTION
The first British record comes from Woods
at Bosmere pool, Salop in 1841. This was
published in Leightons Flora of Salop and was
later identified as Narrow-lipped Helleborine.
It is Nationally Scarce, generally rather
localised and declining. Indeed, following the
clarification of the status of Narrow-lipped
and Dune Helleborines, it is now known to be
much scarcer than was reported in Scarce Plants
in 1994.
PastandpresentoccurrenceofNarrow-lippedHelleborine
inBritainandIreland(basedonpresenceorabsencein
10kmsquaresoftheNationalGrid; datafromtheNew
Atlas).
1500-1999
current range
Britain
Ireland
58
29 (1%*)
% lost, 1500-1969
26%
% lost, 1970-1986
24%
% lost, total
50%
The New Atlas records Narrow-lipped

Helleborine from just 29 10km squares from
1987 onwards, and it has vanished from 50%
of its historical range, with many of the losses
being comparatively recent. It is now extinct in
Devon, West Sussex and Monmouthshire, has
not been seen in Kent and Hertfordshire for
some years and was recorded for the first time in
Herefordshire in 1970 only to disappear again.
Direct threats include the destruction of
M 22July,Oxfordshire.Twostemsgrowingtogether.
its woodland habitat and the conversion of

suitable woods to conifer plantations, although
these practices have largely ceased. The loss of
suitable shaded woodland due to the opening
up of the canopy by severe gales, such as the
great storms of 1987 and 1990, is a subtler
threat. The compaction of woodland soils
by the use of heavy machinery in forestry
operations, horse riding, mountain biking and
the spread of wild boar are other potential
hazards. However, the most obvious is the
widespread damage to flowering plants by the
ever-increasing population of deer. Unlike
Violet Helleborine, it does not seem to have
benefited from the abandonment of coppicing
during the 20th century, perhaps because it is
slow to move into overgrown coppice.
29/1/09 12:12:01
DUNE HELLEBORINE
Dune helleborine
113
Data Deficient
Epipactis dunensis
Other names: E. muelleri
Often considered to be one of the less attractive helleborines due to its relatively small, dull
flowers, Dune Helleborine has the distinction of being endemic to the British Isles and has
a very curious distribution and ecology. Typical plants are found in damp dune slacks on the
coasts of north Wales and northwest England and at a few inland sites in northern England
and southern Scotland. A distinct subspecies, the Tyne Helleborine, is found at other inland
sites in northern England, usually on old spoil heaps and in areas contaminated with heavy
metals. It differs genetically, albeit only slightly, from typical dune slack populations. Dune
Helleborine is the county flower of Lanarkshire.
Identification
Dune Helleborine has yellowish-green, tworanked leaves which are held rather stiffly at
about 45 above the horizontal. The upper part
of the stem is distinctly downy, and the flowers
are relatively small, do not open very widely and
are yellowish-green with the petals and base of
the lip washed with pink. The outer part of the
lip (epichile) is heart-shaped, usually broader
than long, and its tip folds downwards to a
variable extent as the flower ages. The base of
the flower stalk has a violet tinge.
Tyne Helleborine is found at inland sites
and has greener flowers with an epichile that is
longer than broad and not folded downwards.
The base of its flower stalk is yellowish-green.
Both forms are usually self-pollinated (see
p.76 for a discussion of the separation of crosspollinated and self-pollinated helleborines).
Similar species
Broad-leaved Helleborine typically has broader,
darker and greener leaves, arranged all around
the stem and not held rigidly erect. It has
larger and more widely opening flowers, often
with pink or purple tones on the sepals and
a distinctly purple tinge to the lip. Unlike
Tyne Helleborine, the outer part of its lip is
almost always strongly bent down and under;
P 6July, Lancashire.AtypicalDuneHelleborineamong
CreepingWillow,withsmall,slightlysickly,yellowish-green
leavesthatareheldstifflyerect.
29/1/09 12:12:02
114
GENUS EPIPACTIS
usually self-pollinated, only very occasionally

showing a functional viscidium.
Green-flowered Helleborine is, like Dune
Helleborine, self-pollinating and potentially
more confusing. However, its upper stem and
ovaries are either hairless or have a few sparse
hairs. Its leaves are a clean apple green and often
short and rounded, and its flowers hang moreor-less downwards, both in bud and when
open. A feature to check in difficult cases is the
fringe on the edge of the leaves (use a 20x handlens). In Green-flowered Helleborine the tiny,
transparent, whitish teeth (cilia) are arranged
into irregular groups separated by gaps, but in
Dune Helleborine there is an even fringe of
minute teeth.
Narrow-lipped Helleborine is very similar
to Tyne Helleborine, and indeed, for almost
30 years after its discovery, Tyne Helleborine
was thought to be Narrow-lipped Helleborine.
Fortunately, separation of the two is academic,
as Narrow-lipped Helleborine is confined to
southern England, with a few colonies on the
Welsh Marches and in south Wales. The sepals
and petals of Narrow-lipped Helleborine are
on average larger and slightly broader, its lip is a
little more attenuated into a point, the bosses at
the base of the lip are washed pink and the slot
between them (the keyhole) is rather broader.
Habitat
M 17 July, Northumberland. Tyne Helleborine typically
appearsneatandclean-cut.
Tyne Helleborine can also be separated by

the yellowish-green base to its flower stalk
(washed violet in Broad-leaved). However,
Broad-leaved Helleborine is very variable and
in northern England and southern Scotland
some have more yellowish-green, two-ranked
leaves, although they still have large, widely
opening flowers (these are the so-called Youngs
Helleborine, see p.97). In tricky cases it is
worth checking the reproductive structures
within the flower as Broad-leaved Helleborine
is cross-pollinated and Dune Helleborine
Dune Helleborine is typically found in damp

dune slacks where it grows on the slightly
higher and drier areas among and through a
carpet of low, scrubby Creeping Willow. It has
also spread into nearby pine plantations. Tyne
Helleborine, on the other hand, is found at
inland sites, in light, regenerating woodland,
especially birchwoods, where the ground is
kept relatively open by the presence of heavy
metals, mine waste or clinker. For example, in
Northumberland most of the populations grow
along the River South Tyne on well-drained
gravel soils that are heavily contaminated by
P 17July,Northumberland.AgroupofTyneHelleborines
growing under birches and willows by the River South
Tyne.
29/1/09 12:12:03
DUNE HELLEBORINE
115
29/1/09 12:12:06
116
GENUS EPIPACTIS
habitats as all the inland sites are of recent

origin and are man-made. And, although
seemingly very different, adaptations which
allow the species to grow in dune slacks prone
to the stresses of summer drought and salt
spray may be similar to those needed in habitats
stressed by the presence of toxic metals.
Flowering period
Late June to mid-August, usually peaking in the
second week of July in the dune populations
but a little later in adjacent conifer plantations
and at inland sites. The flowers are short-lived
and can be badly affected by drought.
Range
Dune Helleborine is found on Anglesey, the
coasts of Merseyside and Lancashire and
by the Duddon Estuary at Sandscale Haws
in Cumbria. It has also been reported from
the coast of Co. Dublin. Tyne Helleborine
occurs in Northumberland, Cumbria, Co.
Durham and northwest Yorkshire. Scattered
populations of typical dune-type plants
have also been recorded at inland sites in
north Lincolnshire, southeast Yorkshire, Co.
Durham, Cumbria, and in southern Scotland
in Lanarkshire, Midlothian and West Lothian.
However, the features that separate these from
Tyne Helleborine (especially the colour of the
base of the flower stalk) have only recently
been clarified and all the records of Dune
1987-99
1970-86
pre 1970
M 6July,Lancashire.AtypicalDuneHelleborinegrowing
upandthroughthepinkflowersofRestharrow.
zinc and lead tailings, but it is also found on

wooded spoil-heaps contaminated with lead.
In Scotland Dune Helleborine is found on the
wooded slopes of old pit bings (the shale-rich
spoil heaps produced by coal mining), and at
Almond Bing (Falkirk) and Carlisle (Cumbria)
it is also found on and around old railway lines,
both in deep shade and in more open areas.
It seems likely that Dune Helleborine has
spread inland from its semi-natural coastal
29/1/09 12:12:07
DUNE HELLEBORINE
117
Helleborine at inland sites merit

re-examination. World range: Endemic to
Great Britain.
How to find it
Usually easy to find in dune slacks, although
often hidden among Creeping Willow. The
dune populations are all protected on reserves,
including Newborough Warren on Anglesey,
Ainsdale in Lancashire and Sandscale Haws
in Cumbria. Inland, the most accessible sites
for Tyne Helleborine are the tiny Williamston
Reserve and Beltingham River Gravels (both
Northumberland).
DESCRIPTION
upper sepal
anther
petal
petal
pollinia
ovary
stigma
sepal
hypochile
(cup) epichile
boss
sepal
Due to the significant differences between the

two subspecies, they are described separately.
M 6 July, Lancashire. A typical Dune Helleborine. The

outerpartofthelip(epichile)isbroaderthanlongand,in
thiscase,stronglywashedpink.
1. Typical Dune Helleborine

Height: 20-40cm, sometimes to 50cm, with the
tallest plants occurring in sheltered sites.
Stem: Pale green, tinged violet towards the base
and downy towards the tip, with fine, pale hairs.
Usually arises singly, although sometimes there
may be two or three together.
Leaves: Three to ten, arranged in two
opposite rows up the stem and held stiffly
at about 45. The leaves are oval-lanceolate,
mostly more than twice as long as wide and
become relatively narrower towards the flower
spike, but the lowest is very short, broad and
rounded and forms a cowl-shaped sheath.
They are yellowish-green, deeply veined and
their margins have fine, regular, whitish
teeth (cilia) 0.03-0.06mm wide. The leaves

are often damaged by wind or drought by
flowering time.
Spike: Rather lax, with six to 20 flowers,
sometimes as many as 30, set to one side of the
stem. The flowers are initially held horizontally
(patent) but often droop and become pendant
as they go over. At the Scottish sites the flowers
are characteristically held in a drooping position,
but this could be evidence of hybridisation.
Bract: Strap-shaped with a pointed tip; the
lower bracts are slightly longer than the flowers,
but they become shorter towards the tip of
the spike.
Ovary: Green, hairy, six-ribbed (but not
29/1/09 12:12:08
118
GENUS EPIPACTIS
distinct notch running between the epichile and

hypochile. The anther is yellowish-green and is
variably stalked. The pollinia are crumbling and
whitish. The viscidium, although present in the
bud, usually disappears as the flowers open.
M 6July,Lancashire.AtypicalDuneHelleborinewiththe
tipofthelipbentdownandunder.
twisted) and pear-shaped, tapering into a short

stalk that is washed violet at the base.
Flower: Relatively small, dull and not opening
widely, the flower is cup-shaped. The sepals
are yellowish-green, oval-triangular, relatively
short and blunt with a prominent midrib on
the outer surface. The petals are similar in
shape but slightly smaller and very pale green,
often washed pink. At the base of the lip, the
interior of the hypochile is reddish-brown to
dark brown and contains nectar; the exterior of
the hypochile is whitish, variably flushed pink
(the colour shining through from the inside).
The epichile is whitish, sometimes with a pink
flush in the centre, with a greener tip. Heartshaped, it is usually broader than long and the
tip is sometimes folded under, especially as the
flower matures; often the entire epichile is bent
downwards near the join with the hypochile. At
the base of the epichile there are two smooth,
pink or greenish bosses, and these frame a
2. Tyne Helleborine
Height: 17-52cm.
Stem: Green, with fine, whitish hairs on the
upper part. Usually grows singly but fairly
frequently two stems arise together.
Leaves: The three to seven fresh green leaves
are arranged in two opposite rows up the stem,
although sometimes not obviously so. They
are rather floppy and not held as rigidly erect
as in typical dune slack plants; indeed, the
uppermost leaves and lower bracts may be held
horizontally. The leaves are deeply veined but
not sharply folded, the lower elliptical, mostly
more than twice as long as wide, the uppermost
long, narrow and grading into the lower
bracts. The leaf margins have very fine, regular
teeth (cilia) 0.01-0.05mm wide, sometimes
imperceptible.
Spike: Usually rather lax, with five to 35 flowers
(mostly between ten and 25), often facing to
one side. Initially held horizontally, they droop
as they age.
Bract: Fresh green, lanceolate and several times
the length of the flower in the lower part of the
spike but shorter at its tip.
Ovary: Green, hairy, ribbed (but not twisted)
and tapering into a yellowish-green stalk.
Flower: Greenish and relatively small but
opening quite widely. The sepals are narrowly
oval in shape, elongated towards the bluntly
pointed tip. They are pale green on the outer
surfaces, paler and more whitish-green on the
inner surfaces with a diffuse whitish margin.
The petals are similar but slightly smaller
and proportionally shorter, paler and whiter.
The hypochile is transparent-whitish with its
interior washed dirty chocolate-brown at the
base and rear, this colour shinning through to
the exterior of the cup. The epichile is heartshaped, variably longer than broad, with a
pointed tip that projects forward. At its base
there are two small, smooth bosses that frame
29/1/09 12:12:09
DUNE HELLEBORINE
119
a narrow, longitudinal groove. The epichile,

including the basal bosses, is whitish with a
faint cream or greenish wash. The column is
greenish-white, the pollinia cream-coloured,
and the anther cap is dull pale ochre with a
brown stripe on either side. The rostellum is
short, around half as long as the anther. The
clinandrium (depression on the top of the
column) is much smaller than typical Dune
Helleborine, and the viscidium is always absent.
Subspecies
E. d. dunensis is found in coastal dune systems
on Anglesey and in northern England, and
at a handful of inland sites in Lincolnshire,
Yorkshire, Cumbria and southern Scotland.
E. d. tynensis occurs at inland sites in
northern England (Yorkshire, Cumbria and
Northumberland, with genetic analysis of three
populations in Northumberland confirming
subtle differences from dune-slack populations).
There is, however, a near-continuous
gradation between inland sites that hold
classic Tyne Helleborines (e.g. Williamston in
Northumberland) to those with plants with
intermediate characters and those holding
plants much like dune-slack populations listed
above under E. d. dunensis.

Among dune populations, those growing
in the open have a distinctive, almost sickly
yellow cast and often look rather wind-blasted,
whereas those growing nearby in the shelter of
plantations are taller, have more flowers and
both the leaves and flowers are greener and
healthier. Dune-slack type plants growing at
inland sites are also reported to be greener and
more robust.
BIOLOGY
M 17 July, Northumberland;Tyne Helleborine.The pale,

nearlywhitish,tipofthelipcontrastswiththesooty-brown
interiorofthecupatitsbase(thehypochile).

Dune Helleborine is self-pollinated and even
in bud, fragments of the pollinia may fall onto
the stigma and effect pollination. However, in
some plants the viscidium may persist until
the flower has opened, allowing the pollinia to
be removed by insects and cross-pollination to

take place. Even when no viscidium is present,
plants may be cross-pollinated, wasps or other
insects carrying away the pollinia that stick
to their bodies regardless. In one case, around
29/1/09 12:12:11
120
GENUS EPIPACTIS
P 17 July, Northumberland;TyneHelleborine.The
tip of the lip is pointed
and does not bend down
andunder;itisalsorather
narrowandusuallylonger
thanbroad.
two-thirds of the flowers in a population of

Tyne Helleborine had their pollinia removed
(Richards 1986). At a site near Glasgow,
genetic studies have shown that Dune
Helleborine is interbreeding with Broad-leaved
Helleborine to form a hybrid swarm.
Vegetative reproduction may also take place,
the rhizome producing two flower spikes and
these in turn producing buds and roots.

The rhizome is short, slender and woody. It lies
deeply buried in the soil and bears up to ten
thin, wiry roots. There is no information on the
Hybrids
Broad-leaved Helleborine has been found at
most (or all?) of the sites in Scotland. Although
confirmed by genetic analysis, this hybrid has
no scientific name. At Bardykes Bing near

Glasgow there is a hybrid swarm of Dune and
Broad-leaved Helleborines and the individual
plants are very difficult to identify.

The specific name dunensis means of dunes.
The taxonomy of Dune Helleborine is
complex and confusing, and is best understood
in seven stages:
Stage 1: In 1921 Dune Helleborine was first
recognised as something different and described
as a subspecies of Narrow-lipped Helleborine,
using the name E. leptochila dunensis.
Stage 2: In 1926 Dune Helleborine was
elevated to the status of a full species, E.
dunensis, known only from coastal dune systems
on Anglesey and in northwest England.
Stage 3: Colonies of plants identified as
Narrow-lipped Helleborine, a species hitherto
29/1/09 12:12:13
DUNE HELLEBORINE
known only from mature woodland on chalk

and limestone in southern England, were
found at inland sites in Yorkshire in 1967,
Northumberland in the early 1970s and
subsequently more widely in northern England.
(These would eventually become known as the
Tyne Helleborine.)
Stage 4: From the late 1970s onwards a few
colonies of Dune Helleborines were discovered
at inland sites, from Lincolnshire northwards.
Stage 5: The discovery at a few sites in northern
England of populations apparently intermediate
between Dune Helleborine and Narrow-lipped
Helleborine led to the conclusion that they
must be one and the same species, and that
Dune Helleborine should therefore be treated
as a variety of Narrow-lipped Helleborine,
named E. leptochila var. dunensis.
Stage 6: More recently, research, based in part
on DNA studies, has led to the conclusion
that all the populations in northern England,
both Dune and Narrow-lipped, are closely
related, but not the same species as Narrowlipped Helleborine in southern England. These
northern helleborines take the name
E. dunensis, a species endemic to Great Britain.
Stage 7: Tyne Helleborine is formally named
as a distinct subspecies, E. d. tynensis. There
is support for this from genetic studies, which
do show some differences between Tyne
Helleborine and typical dune slack plants.
However, they also indicate that they are
probably each others closest relatives, have the
same ancestor and have only recently separated
into two slightly different entities.
Note that the helleborines on Holy Island
off Northumberland, once thought to be
Dune Helleborine, have now been separated
as a distinct, endemic species, Lindisfarne
Helleborine (see p.122). Delforge (1995)
suggested that Tyne Helleborine, together
with Lindisfarne Helleborine, were the British
representatives of a European species, Mullers
Helleborine E. muelleri. The genetic evidence
shows that this is not the case, but this name
has nevertheless been used in some recent
publications.
121
HISTORY aND
CONSERVaTION
Dune Helleborine is Nationally Scarce and
recorded from just 26 10km squares in the
New Atlas.
Dune slack populations are confined to a
small number of sites but at these localities
Dune Helleborine is often common, for
example, at Sandscale Haws in Cumbria over
1,000 flowering plants have been counted. It
is, however, sensitive to the level of grazing,
especially by rabbits. If this is too intense the
spikes are all nipped off and the plants fail to
set seed. Conversely, if grazing is too light,
scrub invades the habitat and may eventually
shade it out.
The inland populations in northern England
were found in the 1960s and 1970s. Some were
large, with colonies of over 1,000 flowering
plants. All these sites are in young, secondary
woodland, however, and as this matures the
accumulation of humus seems to buffer the
effects of the toxic metals and the ground
cover increases; such changes may lead to the
disappearance of the helleborines.
PastandpresentoccurrenceofDuneHelleborineinBritainandIreland(basedonpresenceorabsencein10km
Britain
Ireland
26
1?
24 (0.8%*)
% lost, 1500-1969
0%
% lost, 1970-1986
8%
% lost, total
8%

1500-1999
current range
29/1/09 12:12:13
122
GENUS EPIPACTIS
linDisfarne helleborine
BaP
Epipactis sancta
For almost 50 years after their discovery, the helleborines on Holy Island in Northumberland
were considered to be Dune Helleborines. However, studies in the 1980s highlighted some
apparently minor differences, and recent genetic work has confirmed that they are not the
same as the helleborines growing in the dune slacks along the coasts of northwest England
and Anglesey. Rather they are a distinct species, endemic to this one small island.
Identification
Straightforward, due to its extremely limited
distribution, although this species is very
similar to Dune Helleborine, with yellowishgreen leaves held in two opposite ranks and
rather dull, greenish flowers which are normally
self-pollinated.
Similar species
Dune Helleborine is very close in appearance,
but typical dune slack plants have slightly
larger flowers and the base of the flower stalk
is washed violet (greenish in Lindisfarne
Helleborine); the details of the column also
differ. Lindisfarne Helleborine is even closer to
Tyne Helleborine, which also has a greenishyellow flower stalk and a similar column
structure, but the lip of Tyne Helleborine tends
to be slightly longer and narrower, and does not
turn under at the tip.
Habitat
Dunes and dune slacks, especially the slightly
raised and more steeply sloping zone around
the perimeter of the slacks. It grows among
Creeping Willow and various grasses or, just as
frequently, on bare sand among Marram grass.
It may be associated with the disturbed ground
around rabbit burrows.
Flowering period
Late June and the first three weeks of July but
usually at its best early in July. Self-pollinated;
the flowers go over quickly.
P 16July,Northumberland.Towardstheendofthefloweringperiod, withthelowerflowerswithered, butinthis
robustplanttheleavesareunusuallyundamaged.
29/1/09 12:12:15
LINDISFARNE HELLEBORINE
123
29/1/09 12:12:19
124
GENUS EPIPACTIS
Range
Only found on the island of Lindisfarne off
the coast of Northumberland, growing at the
Snook at the western end of the island. World
Range: Endemic to England.
How to find it
Found singly and in small groups scattered
through the dunes of the Snook, although
scarce and rather scattered, and absent from
much apparently suitable habitat. The species is
prone to drought and may not flower if it is too
dry, or the buds will shrivel before opening.
DESCRIPTION
upper sepal
anther
petal
petal
pollinia
ovary
stigma
sepal
hypochile
(cup) epichile
boss
sepal
Height: 6.5-42cm.
Stem: Yellowish-green, finely hairy towards
the tip.
Leaves: Yellowish-green, arranged in two
ranks on either side of the stem and held at
around 45. The lowest leaf is rather short,
broad and rounded and forms a cowl-shaped
funnel near the base of the stem. The two to
six higher leaves are elongated-oval in shape,
becoming narrower and more lanceolate
towards the spike. They are clearly veined
and longitudinally folded but not sharply
keeled. The leaf margins have tiny, very fine,
regular teeth (cilia), 0.01-0.05mm wide
(sometimes imperceptible). By flowering
time many leaves are wind-burnt, grazed or
otherwise damaged.
O 7 July, Northumberland. Two flower spikes growing
amongst CreepingWillow; note the blooms of the introducedPirri-pirri-burr,somethingofapestonLindisfarne.
M 16 July, Northumberland.The stem and ovary are

distinctly hairy, and the pollinia have apparently been
removed.
Spike: Fairly loose, with most flowers facing

to one side and held roughly horizontal or just
below the horizontal, although the flowers
droop as they age. The spike consists of up to
ten flowers, although large plants may have 27
or more.
Bract: Yellowish-green, lanceolate, longer than
the flowers in the lower part of the spike but
rather shorter than the uppermost leaves and
becoming shorter towards the tip.
Ovary: Green, ribbed, hairy and quickly
inflating as the flower self-pollinates. The flower
stalk is green or yellowish-green.
Flower: Rather small and drab but opening
fairly widely. The sepals are greenish and
roughly triangular with the petals smaller
and paler. The hypochile is whitish with a
chocolate-brown inner rear wall to the cup and
sometimes a pink wash to the exterior. The
epichile is heart-shaped with the tip variably
deflexed and whitish, washed green in the
29/1/09 12:12:22
LINDISFARNE HELLEBORINE
125
England and Wales. Delforge & Gvaudan

(2002) went on to propose the name Epipactis
sancta for the species. Genetic studies show that
not only are Holy Island plants distinct from
both classic dune slack populations and the
Tyne Helleborine but also that they probably
evolved independently of both of these.
HISTORY aND
CONSERVaTION
M 7July,Northumberland.Thelipistypicallywhitish,becominggreentowardsthetip.
centre towards the tip. The column is whitish,

the anther cap yellow with a narrow brown
stripe at the side, and the pollinia are ochreyellow. The clinandria (depressions on the
top of the column) are very reduced and the
rostellum is short, around half the length of
the anther.
These helleborines were first found on Holy

Island in 1958. In recent years there has been
an increase in the number of flowering plants
due to a reduction in the number of rabbits
(which nip off the stems) and around 150-300
spikes appear annually. The presence of rabbits
may be important, however, despite their
impact on flowering numbers; they prevent
scrub from invading the dunes and the bare
ground created by their scrapings may help in
the establishment of seedlings.
Subspecies
None.

None.
BIOLOGY
Self-pollinated.

No information.
Hybrids
None.

The specific name sancta means holy or sacred,
a reference to Holy Island.
Holy Island plants were initially identified
as Dune Helleborine, but the colour of their
flower stalks and details of the flower structure
led to suggestions from the 1980s onwards
that they were not the same as the Dune
Helleborines growing on the coast of northwest
M 16 July, Northumberland.The yellow pollinia are still

intactonthisplant.
29/1/09 12:12:24
126
GENUS EPIPACTIS
Green-flowereD helleborine
Epipactis phyllanthes
Formerly: E. pendula, E. vectensis. Other names: Pendulous-flowered Helleborine
This mysterious orchid has a habit of coming and going at its known sites and of popping
up unexpectedly in new places, so is always worth looking out for in almost any type of
woodland in England and Wales. It is also one of the most variable orchids, and this has
caused a great deal of confusion over the years.
Identification
Green-flowered Helleborines come in a
wide variety of sizes and shapes. Some are
diminutive plants with large, swollen, pearshaped ovaries and small flowers which hang
vertically downwards and never really open.
Others are rather robust, with wide-open,
saucer-shaped flowers that can be held facing
more outwards than downwards (with such
well-developed plants being commonest
in northern England). However, despite
this diversity, this species often has a fairly
distinctive feel.
Green-flowered Helleborine is typically
relatively slender with short leaves which are
a fresh apple-green colour. Leaf shape and
posture are variable, but some plants have
characteristically well-spaced leaves that are
very rounded and held stiffly horizontal. The
flowers are green with a whitish or sometimes
pinkish lip and often do not open widely or,
indeed, may not open at all; such plants appear
to be permanently in bud, although the large
ovaries swell conspicuously. The base of the
flower stalk is greenish, and in most plants
the flowers hang vertically downwards. The
lip shape is very variable. In some populations
it is almost identical to the petals (a feature
shown by no other British helleborine) and
in others the lips are fully formed and divided
O 1 August, Northumberland, var. pendula. A typically
small, few-flowered plant, with very rounded leaves held
horizontallyandflowersthatdonotopenwidely.
P 12July,Northumberland,var.pendula.Arobustplant,
withmanyflowers, butalmostallofthem, asusual, are
drooping.
29/1/09 12:12:25
GREEN-FLOWERED HELLEBORINE
127
29/1/09 12:12:28
128
GENUS EPIPACTIS
as normal into a cup-shaped hypochile and

heart-shaped epichile. The flowers are generally
self-pollinated, the pollinia crumbling within
the flower onto the stigma.
Whatever the appearance of the flowers,
three features will confirm an identification.
First, the upper part of the stem and the
ovaries are either hairless or, rather less
frequently, the upper stem may be sparsely
hairy (use a hand-lens). Second, the hypochile
is greenish-white, lacking an obvious dark
brown or purple lining. Third, in difficult
cases look at the edge of the leaves (a 20x
hand-lens is best for this). In all the other
helleborines there is an even fringe of tiny,
whitish, hair-like projections, but in Greenflowered Helleborine these cilia are unevenly
distributed in groups.
irregularly bunched cilia along leaf margin
Similar species
Narrow-lipped Helleborine is found in
beechwoods in southern England and south
Wales and is also greenish overall. At all times
it can be distinguished by its rather hairy upper
stem and ovaries.
Dune Helleborine occurs on coastal dunes
in Anglesey and northwest England, and as the
distinct variety known as Tyne Helleborine at
inland sites in northern England and southern
Scotland. It always has a hairy upper stem and
ovaries and a dark lining to the hypochile at
the base of the lip. In addition, typical dune
slack plants have a violet wash to the base of
the flower stalk (yellowish-green in Tyne and
Green-flowered Helleborines).
Broad-leaved Helleborine sometimes has
greenish flowers but always has a densely hairy
upper stem and slightly hairy ovary. It is also
cross-pollinated (see p.76 for a discussion of
M 26August,Norfolk,var.degenera.Verytypically,growing through a carpet of Ivy. In this season the flowers

hardlyopenedatall.
29/1/09 12:12:29
the separation of cross-pollinating and selfpollinating helleborines).
Habitat
Very varied, and although the species favours
alkaline soils, it is not confined to areas of
chalk or limestone and will grow on calcareous
to mildly acidic sands and clays and in silty
river valleys. It is mostly found in light to
moderate shade but can occur in densely
shaded situations, although it is often rather
small and slight in such places. Green-flowered
Helleborine frequents a wide variety of
woodlands but has a definite preference for
beechwoods in southern England and overall
favours smaller woods, copses, belts of trees
or tall hedges next to woodland, or the betterlit edges of larger woods; it is often found on
road verges. It prefer areas where the ground
cover is rather sparse or low, no taller than 1520cm, and is characteristically found growing
through a carpet of Ivy.
Rather than having an association with
ancient woodland, many of its sites are
relatively recent in origin, such as beech
and pine plantations and shelterbelts and,
in Northumberland, the maturing birch and
hawthorn scrub found on old waste tips
contaminated with zinc and lead. Another
favoured habitat is thickets of willows and
other trees alongside rivers and streams
that are subject to occasional flooding, the
helleborines growing on the better-drained
ridges and banks. Indeed, this may be the
natural habitat of the species and it has
even been found in Hampshire growing
among reeds and willows in the tidal part of
the River Itchen. Conversely, Green-flowered
Helleborines also occur in very dry woods.
At a few sites in Wales, northwest England
and Co. Dublin, it grows in the open on
sand dunes, coming up through a blanket of
Creeping Willow on the drier hummocks.
However, it tends to look yellow and sickly
in this habitat, appearing rather healthier
where it has spread into adjacent conifer
plantations.
129
Flowering period
Late June to early or even mid-September but
mostly from mid-July to mid-August, with
dune populations typically earliest.
Range
In England rather scattered and local but
found west to Dorset, Gloucestershire and
Herefordshire, east to Kent and Norfolk and
north to south Cumbria and Northumberland.
The bulk of the population is found in central
and northern Hampshire, Wiltshire, West
Sussex and Surrey, where Green-flowered
Helleborine can be locally frequent. In Wales
there are scattered sites in coastal dunes
in Glamorganshire (including Kenfig and
Whitford Burrows) and Merionethshire
(Morfa Dyffryn), but it is otherwise only found
in Flintshire. There are also a handful of sites
in Ireland, in Co. Dublin, Co. Leitrim and
Co. Ferrmanagh. World range: Restricted to
western Europe, from northern Spain through
France and Belgium to Denmark. May occur
more widely but the taxonomic confusion
surrounding the species and its close relatives
in Europe has clouded an understanding of its
true range.
1987-99
1970-86
pre 1970
How to find it
Its overall green coloration, relatively small size,
habitat and flowering period can make this a
29/1/09 12:12:30
130
GENUS EPIPACTIS
hard species to find, although the apple-green,

rounded leaves, which are held horizontally, and
the mass of large, drooping buds often catch the
eye. Colonies are often small, and numbers can
fluctuate, with fewer plants in dry years. Most
of the sites for the species in southern England
are anonymous and have little or no other
orchid interest, and the most accessible sites to
see the species at its best are the dunes on the
Lancashire and Cumbrian coasts.
DESCRIPTION
upper sepal
anther
petal
petal
pollinia
ovary
stigma
sepal
hypochile
(cup) epichile
boss
sepal
Height: 5-75cm but mostly 15-50cm; taller

plants are commonest in northern England.
Stem: Robust, purple-brown at the base
(where there are one to three similarly coloured
sheaths) but becoming apple-green for most of
its length. The stem, including the upper part, is
hairless or has sparse, short hairs. Usually grows
singly but occasionally two or three stems may
arise together. We have seen a photograph of
a group of nine (Killick et al. 1998), although
such a large cluster may have resulted from the
rhizomes of several plants growing together.
Leaves: There are three to seven, exceptionally
as many as 16, apple-green leaves which are well
spaced but often rather high up on the stem.
They are very variable in shape and posture
but usually relatively small. Many plants have
leaves that are broadly oval or egg-shaped but
on others they are longer, narrower and more
pointed. On all plants, the upper leaves are
usually narrower and more bract-like, and
the lowest green leaf is often rather small and
frequently funnel-like, partially sheathing the

stem. The leaves may be arranged in two ranks
on either side of the stem and on some plants
are held flat in a very characteristic horizontal
plane. On others they may be held lightly folded
and positioned nearer to 45. The margins
have tiny, whitish teeth (cilia) arranged into
irregular groups.
Spike: The buds may be held upright,
horizontally or drooping, but, once open, the
flowers usually hang down near to the vertical.
The spike contains between two and 25 flowers,
exceptionally as many as 35, and is often fairly
crowded. The flowers often face in the same
direction, and the uppermost buds in the spike
may fail to open.
Bract: Green and lanceolate; the lowest
bracts are much longer than the flowers but
they become progressively shorter, with the
uppermost a little shorter than the flowers.
M 12 July, Northumberland, var. pendula.The ovary is

boldly-ribbedbuthairless,andthebaseoftheflowerstalk
is green.
29/1/09 12:12:33
131
The bracts are usually horizontally, especially on

less densely crowded spikes.
Ovary: Swells rapidly and becomes large and
pear-shaped. It is shiny, virtually hairless but
with six prominent ribs, tapering into the short,
curved and twisted green stalk.
Flower: Var. pendula (see below for other
flower shapes). Overall greenish. The sepals are
elongated-ovals, variably tapering to a pointed
tip, and pale green with a prominent green
midrib on the outer surface. The petals are a
little smaller, paler and more greenish-white.
The hypochile is whitish or dull olive-white,
almost translucent, with the interior of the
cup pale greenish (sometimes lightly washed
brown). The epichile is heart-shaped with a
pointed tip which is strongly turned under, and
there are two rough bosses at the base with a
central groove between them. It is whitish and
tinged with green or sometimes pale pink in the
centre or towards the tip, especially the bosses.
The column is whitish and the anther cap dull
yellowish-white with narrow brown stripes
at the side. The pollinia are cream or whitish,
quickly crumbling and inconspicuous in the
anther. A small and not very sticky viscidium
may be present in the bud, but this withers by
the time the flower opens.
Subspecies
None.

Green-flowered Helleborine is very variable,
both in terms of the structure of the flowers
and in the overall size and shape of the plant.
The flowers range from those where the lip is
well developed, with a clear distinction between
the basal cup (hypochile) and the heart-shaped
tip (epichile), to those where the lip is simple
and petal-like. There are also parallel differences
in the shape of the anther. Between the two
extremes the range of variation is almost
continuous. There are four named varieties,
although due to the variation it can be difficult
to identify plants. Notably, there is no real
dividing line between var. pendula and var.
vectensis, and flowers with mixed characters can
M 1 August, Northumberland, var. pendula. The lip is

well-formedinthisvarietyand, onthisplant, theflowers
haveopenedextraordinarilywidely.
29/1/09 12:12:35
132
GENUS EPIPACTIS
be found. It would probably be better to unite

the two varieties as var. vectensis.
There are broad geographical trends in
lip shape, with populations in the north and
west having the most robust plants, the best
developed lip structure and widely opening
flowers (var. pendula), whereas those in the
south tend to be smaller, with incompletely
developed flowers that open only partially.
Even in the same region, however, there can
be a good deal of variation, and at a few sites
a mixture of lip shapes can be found in the
same colony.
Var. pendula The lip is large and fully
var. pendula
var. vectensis
var. degenera
var. degenera
var. phyllanthes
var. phyllanthes
(After Young 1952b)
M 17 July, Norfolk, var. vectensis. In this variety the lip is

well formed and it can, as here, have a pink tinge.
developed. The hypochile is well formed and

around 4mm long. The epichile is heartshaped, as long as the hypochile or only
slightly longer, with a pointed tip that is
normally strongly reflexed. The epichile is
wrinkled at the base or has two bosses. The
flowers open widely and cleistogamic flowers
are rare. This is the commonest variety in
north Wales and northern England but it has
been found further south and intermediates
with var. vectensis are not infrequent.
Var. vectensis As var. pendula but the
hypochile is smaller, 2.5-3.5mm long, and
more hemispherical so that it embraces the
column closely. The epichile is distinctly longer
than the hypochile, sometimes markedly so,
and the tip is generally not reflexed so that the
whole lip points forwards. The flowers open to
29/1/09 12:22:02
M 14August, Norfolk, var. degenera.The flowers have

openedasmuchastheyaregoingto,andtheovariesare
allswollen(theywerepresumablyself-pollinatedinbud).
133
a variable extent and are often cleistogamous.

Mostly found in southern England and the
Midlands but has been recorded north to
Yorkshire. E. cambrensis was described from a
few reportedly distinctive plants, delicate and
yellowish-green, with yellowish-white flowers,
found in the 1940s at Kenfig and Margam
Burrows in south Wales. Initially described
as a distinct species, it has more recently been
demoted to E. p. var. cambrensis, but was not
recorded again until similar plants were found
at Kenfig NNR in 2004. The most distinctive
feature of cambrensis appears to be the leaves,
which are narrow and sharply folded along the
midrib, but whether it is appropriate to treat
it as a named variety has still to be confirmed.
Var. phyllanthes The lip is not divided into
hypochile and epichile and is oval or lanceolate
with a central midrib, resembling a petal in
size and shape. The flowers rarely open widely
and are usually cleistogamous. This form is
commonest in southern England.
Var. degenera This is intermediate between
var. vectensis and var. phyllanthes. The
hypochile is reduced to a shallow depression
at the base of the lip and although there may
be small bosses at the sides of the base of the
epichile, the lip has either no waist at all or
only a rudimentary one and always lacks
the central groove between the two bosses.
The flowers rarely open widely and are
usually cleistogamous. It is mainly found
in southern England.
The size and shape of the plants and their
overall colour also varies, especially the size
and shape of the leaves. This variation seems
to be determined by the local environment
(for example, on exposed dunes the plants are
dwarfed and yellowish). In addition, individual
plants and whole populations vary in the
degree to which the flowers open and how
pendulous they are, although this can change
from season to season; plants in which the
flower buds never open are commonest
in dry seasons, whereas in very wet years
more plants than normal may show widely
opening flowers.
29/1/09 12:12:37
134
GENUS EPIPACTIS
BIOLOGY
Green-flowered Helleborine is always selfpollinated. In some plants this occurs when
the flowers are still in bud (the flowers are
cleistogamous); the buds may or may not open
subsequently. Following pollination the whole
column, together with the lip, withers rapidly
but the sepals and petals can remain intact for
a long time.

The aerial stem grows from a rhizome which
varies from nearly horizontal to nearly vertical,
with numerous thick, fleshy roots, both long
and short. There is no information on the
Hybrids
None.

The specific name phyllanthes may celebrate
Phillis Wood near Treyford in West Sussex,
the site from which the first specimens were
collected in the 1830s, but is more probably
from a compound of Greek words meaning
leaf-like flower.
This species was misunderstood for a
long time and widely confused with Narrowlipped Helleborine. It was not until 1952
that E. vectensis (described as a variety of
Narrow-lipped Helleborine in 1918 and then
as a distinct species in 1940) and E. pendula
(described in 1942) were shown by the British
botanist Donald Young to be the same as E.
phyllanthes, a species described in 1852 in the
Gardeners Chronicle but ignored for a hundred
years. Epipactis cambrensis (described in 1950)
has since been added to the pot-pourri.
in the number of records in recent years but

this is offset by the loss of known sites. It is
extinct in the Isle of Wight, Somerset, Suffolk,
Warwickshire, Derbyshire, Co. Antrim, Co.
Derry and Co. Wicklow.
Past and present occurrence of Green-flowered HelleborineinBritainandIreland(basedonpresenceorabsencein10kmsquaresoftheNationalGrid; datafrom
the New Atlas).
Britain
Ireland
134
86 (3%*)
4 (0.4%*)
% lost, 1500-1969
21%
44.5%
% lost, 1970-1986
15%
11%
% lost, total
36%
55.5%

1500-1999
current range
Losses may be due to the grubbing-out

or coniferisation of woodland, but some
of its habitats are ephemeral in nature and
become unsuitable as the woodland matures
and becomes more shaded. In general, there
has been a decrease in numbers in southern
England but some spread on the edges of the
range. For example, it was first found in Norfolk
in 1969 at Santon and subsequently a number
of colonies were discovered in the 1990s
along the banks of the Rivers Yare and
Wensum in Norwich.
HISTORY aND
CONSERVaTION
Nationally Scarce in Britain and specially
protected in Northern Ireland under Schedule
8 of the 1985 Wildlife Order (NI). A better
awareness of the species has led to an increase
29/1/09 12:12:38
Distribution
There are around five species in this genus, distributed in
Europe, Asia, Africa and through the Pacific Islands to
Australia. All are fully mycotrophic and depend on fungi
throughout their life.
Name
The generic name Epipogium derives from the Greek and
means overbeard, a reference to the position of the lip,
uppermost in the flower.
GENUS EPIPOGIUM
GHOST ORCHIDS
12/2/09 16:55:27
136
GENUS EPIPOGIUM
Ghost orchid
Extinct (?)
Epipogium aphyllum
This may be the rarest wild plant in Britain. It has only been recorded in two widely separated
regions: the Chilterns and around Herefordshire and Shropshire. Fully dependent on fungi
throughout its life, it only appears above ground to flower and fruit. There have, however, been
no documented sightings in England for almost 20 years and it has become the Holy Grail for
many orchidophiles. Once known as Spurred Coralroot, the English name Ghost Orchid much
more accurately reflects its elusiveness. Its small size and pallid, ethereal appearance make it
extremely hard to find in its gloomy woodland haunts truly ghostly qualities.
Identification
Very distinctive, it is pale and waxy with
relatively large, pinkish flowers and no green
leaves.
Similar species
None.
Habitat
In Herefordshire and Shropshire it has been
found in oak woodland on clay soils. The
Chiltern sites are in beechwoods on chalk or
clay-with-flints. Ghost Orchid often grows
where the soil is slightly deeper, in hollows
or on the top and sides of ditches (but not,
apparently, in the bottom of a ditch). The
margins of roads and tracks appear to be
favoured but this may reflect the fact that it is
easiest to search for the plants in such places. It
often grows in a deep mat of decaying leaves but
may also be found in areas with almost no leaf
mould, the rhizome lying in the mineral soil
itself. Several times it has been found growing
out of old tree stumps, and one has even been
recorded growing through a rotten mattress in a
roadside ditch.
Ghost Orchid is often said to grow in
heavily shaded woodland where the ground
is otherwise bare. In 1953, however, having
discovered the largest ever colony in Britain,
Rex Graham stated, The canopy, locally heavy,
is on the whole rather more open than what one
might have expected to be ideal for this orchid,
and a few plants grew in comparatively light
conditions and amongst the type of ground
M 23 July, Scwartzwald, Germany (Sean Cole). The spur

of the flower lies at the top and the lip faces downwards.
O 14 August, Buckinghamshire (Nigel Redman). A
ghostly apparition on the beechwood floor.
29/1/09 12:25:04
GHOST ORCHID
vegetation that invades these woods whenever

light gives a chance.
Flowering period
April to mid-October but mostly mid-July to
mid-September. Plants in Buckinghamshire
tend to flower from mid-July to the third week
of August and those in Oxfordshire a little later,
from mid-August to mid-September.
Flowering is said to follow a wet spring.
This allows the plant to store up water prior to
developing an aerial stem. The flower bud is,
however, initiated in the year prior to flowering,
so the conditions in two consecutive years may
be critical in determining whether the plant
is able to bloom. The Ghost Orchid may even
flower underground, buried in the leaf-litter.
1987-99
1970-86
pre 1970
137
to the Kamchatka Peninsula, Russian Far East,

Sakhalin, Japan, northeast China and Korea
and is also found to the south scattered
through the Himalayas and southern China,
including Taiwan.
How to find it
Ghost Orchid is probably the hardest plant
to find growing wild in Britain. Its small size
makes it extremely inconspicuous and very
difficult to see in the deep leaf-litter of the
woodland floor. Plants may even be hidden
under wind-blown leaves. Some ingenious
techniques have been used to try to locate it,
including night forays where a powerful torch
is shone parallel to the ground in the hope of
highlighting the tiny spikes. The prolonged
period during which it may bloom does not
help, nor does the fact that slugs so often
destroy the flower spikes; even if they survive,
the flowers do not last more than a few days.
Knowledge of the precise location of previous
records is of only limited use, as they seldom
reappear in exactly the same spot. Finally,
even in the right place at the right time few
flower spikes are produced, with one to four,
exceptionally seven, at any one site in any one
year. The total of 25 near Marlow in 1953 was
quite unique.
DESCRIPTION
Range
Confined to England, where it has been
found in Oxfordshire, Buckinghamshire,
Herefordshire and Shropshire. World Range:
Central and northern Europe and through
northern Asia to the Far East, with a few
scattered sites south to the Himalayas and
China. Occurs north to northern Scandinavia
and south to the Alps, central Italy, northern
Greece, Crimea and Caucasus; also found
on Corsica. Largely absent west of the Alps
and Massif Central, but there are odd sites,
including the Pyrenees. In Siberia it ranges east
Height: Usually 5-10cm but sometimes to

12.5cm and exceptionally to 24cm.
Stem: Swollen at the base, thick but fragile and
translucent-white, washed dull rose-pink or
pinkish-brown and variably streaked pinkish.
Above the uppermost flower the stem continues
as a short, narrow projection (like an aborted
flower stalk). Stems arise singly but occasionally
two spikes grow from the same rhizome.
Leaves: There are no green leaves, merely two
or three brown, sheathing scales at the base of
the stem and one or two longer, often darkedged, tightly clasping, scale-like leaves higher
on the stem.
Spike: Most plants carry one or two flowers,
but the more robust bear three or four blooms,
29/1/09 12:25:04
138
GENUS EPIPOGIUM
which are held pointing outwards or nodding

downwards on short stalks. Neither the stalk
nor the ovary is twisted, hence the spur points
more-or-less upward and the lip similarly lies at
the top of the flower.
Bract: Roughly equal in length to the flower
stalk and ovary together, papery and translucent
with a yellowish-straw wash.
Ovary: Bag-like, pale straw, veined or spotted
with pink or violet; the flower stalk is noticeably
slender and curves to hold the ovary hanging
downwards.
Flower: The spur and lip lie at the top of the
flower, with the lip facing downwards and
outwards. The lip is whitish or very pale pink
and divided into two sections: at the base the
hypochile is short with two broad, spreading,
triangular side-lobes flanking the entrance to
the spur; the epichile is longer, broadly tongueor heart-shaped and deeply concave with
crimped margins and several rows of raised
magenta ridges. The lip is strongly curved to
lie parallel with the spur, which it may touch.
The spur is relatively large, sack-shaped, slightly
curved, pale pink and filled with nectar. The
sepals and petals are very pale yellowish-straw
with fine reddish dots or streaks; they are
narrow and strap-shaped with their edges
curled inwards, making them appear even
narrower. The upper sepal and two petals hang
down below the rear of the flower, and the two
lateral sepals project downwards and variably
outwards at the sides. The column is pale yellow
and short with a well-developed rostellum; the
anther is pale yellow and the two pollinia are
connected by a stalk at their base to the two
distinct viscidia. The nectar is said to smell
of fermenting bananas or vanilla but it is also
described as foetid.
BIOLOGY
Possibly pollinated by bumblebees and wasps.
The insect lands on the lip and moves towards
the spur to access the nectar. As it backs out,
it ruptures the rostellum, pushes the anther
M 23 July, Scwartzwald, Germany (Sean Cole). At some

sites in Europe this enigmatic orchid is rather predictable
in its appearances and almost common, although never
commonplace.
cap aside, and the slender stalks of the pollinia

are glued to its back or head. However, it has
been suggested that bees, although they do
visit the flowers, are unlikely pollinators. They
are the wrong shape and size, especially given
that the flowers are upside-down (and all
references to bees as pollinators may originate
with the observations of a Dr Rohrbach in
the 19th century); if this is so, the mechanism
of pollination is a mystery. Self-pollination
in flowers in the normal pendant position is
prevented by the position of the stigma, above
the anther, because pollen cannot fall upwards.
However, self-pollination may be possible while
the flowers are upright when still in bud.
Whatever the mechanism, few flowers
produce capsules, perhaps because of the
lack of suitable pollinators in the dark woods
where Ghost Orchids are found, or because
of a lack of other simultaneously flowering
29/1/09 12:25:05
GHOST ORCHID
Ghost Orchids to cross-pollinate the flower.

The species certainly reproduces
vegetatively and this may be the major means
of propagation, although the literature is
contradictory about the mechanism. Some
sources state that new plants grow from buds
at the tip of the roots. Others state that some
of the branches of the rhizome become slender,
attenuated and thread-like, and function as
underground runners. Bulbils are formed at
intervals along these runners and these may
become detached and grow into independent
new plants. After flowering, the major part of
the rhizome usually dies, leaving the runners to
form new plants.

Throughout its life Ghost Orchid is entirely
dependent on fungi for its nutrition (that is,
fully mycotrophic). It is often stated that Ghost
Orchid is saprophytic and gets its nutrients
from decaying organic material, but this is
not correct, as it feeds by consuming a living
fungus. It seems likely that, like Birds-nest and
Coralroot Orchids, its fungal partner will prove
to be ectomycorrhizal and have a symbiotic
association with the roots of trees. This would
allow the Ghost Orchid to acquire nutrients, via
its fungal partner, from the trees.
The aerial stem grows from a whitish
rhizome which has many branches. In turn,
these are forked or tri-lobed. The lobes are
rounded at the tip and often spread fanwise.
The much-branched rhizome is said to look
like certain types of coral, hence the old name,
Spurred Coralroot. There are no roots, rather
the rhizome has a sparse covering of fine, long
hairs. Once the plant is mature enough to flower,
a bud appears in the autumn and swells as
the food and water reserves from the rhizome
are transferred to it. If the following season is
sufficiently wet, a flower spike may be produced.
Seed probably germinates in the autumn,
and up to ten years, possibly even twenty, may
elapse before flowering, although no one has
performed a long-term study of marked plants
to confirm this.
139
Subspecies
None.

None.
Hybrids
None.

The specific name aphyllum derives from the
Greek and means without leaves.
HISTORY aND
CONSERVaTION
A Red Data Book species that is classified as
Extinct, it is fully protected under Schedule 8
of the Wildlife and Countryside Act 1981.
The first British record dates from 1854,
when Ghost Orchid was found by the Sapey
Brook on the border of Herefordshire and
Worcestershire, as detailed in the Journal of
Botany: The discovery was communicated
on the 9th of this month [September], by the
Rev. W. Anderton Smith, of Tedstone, from
the Rectory, Delamere, Bromyard: - a few
weeks since, Mrs. Anderton Smith found a
specimen For some time we looked in vain
for other specimens; but, on the 23rd ult., I
was fortunate in detecting a considerable mass
of it. All were found at the foot of a very steep
woody bank, close to a brook; the soil very wet
and stiff. As the banks are very much trampled
on at present (timber and faggots being drawn
along), I decided on digging it up, and planting
it in a similar spot in our own grounds.
The next records of Ghost Orchid came
from Bringewood Chase near Ludlow
(then in Shropshire, the area is now in the
administrative county of Herefordshire). It was
found by a Miss Lloyd in 1876 and again in
the same wood by a Miss Peel in 1878, with a
third record from a different part of the wood
in 1892. A Ghost Orchid was also recorded
from the Wye Valley near Ross-on-Wye,
Herefordshire, in July 1910.
There was then a gap of 13 years before
the species was seen again in Britain, this time
29/1/09 12:25:06
140
GENUS EPIPOGIUM
in Oxfordshire. In June 1923 two plants were

found just north of Henley-on-Thames. After
a prolonged search, Dr. G. C. Druce, perhaps
the most eminent botanist of the day, succeeded
in tracking down a third plant in July. In late
May of the following year two more plants
were found there. Then, on 30 June 1931,
Ghost Orchid was found a few miles to the
west by Mrs Vera Paul (then a schoolgirl), in
beechwoods towards Stoke Row. There was a
single large spike, 24cm high with three flowers,
growing from the middle of an old tree stump.
A single spike was also seen nearby in 1933.
The trail went cold again for 20 years and it
was not until 1953 that Ghost Orchid appeared
again, this time in Buckinghamshire (around
ten miles from the Oxfordshire sites). Rex
Graham had been searching for Ghost Orchids
for 20 years and on 18 July 1953 was in woods
west of Marlow. He was lighting his pipe when,
over the bowl of it, he saw a Ghost Orchid
growing among the beech leaves. There proved
to be a scattered colony of 25 spikes belonging
to 22 plants. Three specimens were collected
and at least three more spikes were felled by
slugs. It was found there again the following
year but not in such great numbers. These
woods went on to become the most productive
area for the species, although the number of
plants never matched the 1953 total. Spurred
on by the events in Buckinghamshire, Vera
Paul searched the woods near Stoke Row in
1953 and two spikes were found. Plants were
seen there in 1954 and 1956, too, while on 17
September 1963 five spikes were found close
to the stump where the species had been first
found in 1931. Sightings in this complex of
woods have continued, with reports up to and
including recent years.
On 19 September 1982, after a gap of
62 years, the Ghost Orchid reappeared in
Herefordshire, when a single spike was found
in a cart track under oak and pine in a wood far
removed from its old localities. By 1 October the
stem had rotted, probably due to slug damage.
The Red Data Book states flatly that Ghost
Orchid has not been seen since 1986, but
there were reliable reports from near Marlow

until 1989 and several reports since from both
Buckinghamshire and Oxfordshire, covering
almost every year up to 2003. The evidence for
some of these is certainly limited, however, and
the finders have displayed a great reluctance
in coming forward and allowing the record to
be confirmed, probably to the detriment of
the plant. The relative lack of records in recent
years may merely be part of the natural cycle
of a small population or more likely reflects a
genuine decline. It has been suggested that the
opening of the canopy following tree falls after
the great storms of 1987 and 1990 may have
resulted in the exposure of the woodland floor
to too much sun, conditions becoming too dry
for the species (but see Habitat).
Ghost Orchid is on the very edge of its
range in England and is naturally rare. Its
position here has not been helped by the
unsympathetic management of ancient
woodland, which has been cleared or replanted
with conifers (for example, Bringewood Chase
in Shropshire has largely been coniferised,
with little ancient woodland remaining). Other
more immediate threats include horse riders
and cyclists who create paths through areas
where the plants have flowered in the past, and
forestry operations which involve the use of
heavy machinery and the dumping of materials.
Some of these problems could be avoided if the
responsible authorities knew the location of the
plants, but there is such secrecy surrounding
Ghost Orchid that many sightings are hushed
up. Humans can be a direct threat, too, with
photographers trampling the area around
flower spikes, and in 1978 and 1979 collectors
apparently dug up several plants. Finally, there
are natural hazards, with deer and slugs taking
a toll. Well-meaning naturalists have advocated
surrounding plants with slug pellets, but even a
moribund slug may cause damage and there is
no information on the effect of molluscicides on
the plant or its fungal partner.
P 16 June, Norfolk. Fen Orchid, a relatively robust plant.
29/1/09 12:25:07
Genus LIPARIS
Fen orchid
Distribution
This small, green-flowered orchid
grows from a pseudobulb, a structure
commonly found in tropical epiphytes
and, although Liparis has an almost
worldwide distribution, it is essentially
tropical. There are 300 or so species but
just one is found in Britain and Europe.
Name
The generic name Liparis is from the
Greek liparos and means fatty, greasy or
shiny, a reference to the appearance of
the leaves.
29/1/09 12:25:10
142
GENUS LIPARIS
Fen orchid
Endangered, BaP
Liparis loeselii
Other names: Loesels Twayblade (North America)
This small and rather drab orchid has always been very localised in Britain and is now confined
to just a handful of sites in Norfolk and South Wales. Indeed, it is threatened throughout its
range in Europe and has declined in much of North America, too. Surprisingly, studies of its
life history suggest that it is a weedy species, well-adapted to colonising and rapidly filling
newly available habitats, and that it thrives on disturbance.
Identification
Very rare and localised, this is an orchid highly
unlikely to be stumbled upon by chance. When
in flower, Fen Orchid is distinctive. The small,
pale green flowers appear to be a jumble of thin,
spidery projections and are unique. Importantly,
they are usually held facing upwards, with the
column vertical, and the three strap-shaped
sepals, two thread-like petals and the lip all
held more-or-less horizontally in a cross; two of
the sepals lie parallel, close together under the
tongue-like tip, to form one arm of the cross.
Similar species
Non-flowering plants may form the bulk
of the population. They could be confused
with Common Twayblade or Lesser Butterfly
Orchid but are much smaller and their leaves
sheathe the swollen base of the stem to form a
pseudobulb.
Habitat
M 18 June, Norfolk. The aerial stem grows from a pseudo

bulb, a swelling in the stem that is concealed by the base
of the leaves.
Fen Orchid is found in two distinct habitats;

marshes in the Norfolk Broads and damp dune
slacks on the coast of South Wales. Although
superficially very different, both offer the same
combination of bare ground which is kept
damp or wet by neutral or calcium-rich
ground water.
In the Norfolk Broads, Fen Orchid
occurs as a member of a particularly speciesrich community in areas of wet, peaty fen
dominated by Common Reed or Great Fen
Sedge. It grows on the mossy carpet at the
base of the reeds, on the sides and tops of
sedge tussocks, or on the bare peat itself. It is
29/1/09 12:25:12
FEN ORCHID
flowers
old
pseudobulb
early
spring
143
capsules
new
pseudobulb
new
pseudobulb
summer
late
summer
early
winter
roots
M Annual replacement of pseudobulbs in Fen Orchid.
extremely fussy, however, and an analysis of its

current and former sites in East Anglia reveals
that they have several features in common:
1. All are wet, with a water table which is
typically just below the surface and relatively
stable. Fen Orchid does not necessarily require
waterlogged conditions and, indeed, may
grow on tussocks up to 15cm above the water
level, but the presence of surface water could
be important in protecting it from slugs and
snails. 2. The ground water is neutral to mildly
alkaline and very low in nutrients. 3. Most
sites were cut for sedge (Great Fen Sedge was
used for thatching) or litter (hay). The cutting
and removal of reed and sedge, combined with
infertile ground water, slow the development of
scrub and keep the vegetation relatively sparse
and open. 4. Most of the sites in the Broads are
on abandoned turbaries where peat was cut as
fuel in past times. These cut-over areas then
flooded, and gradually the resultant turf ponds
were colonised by vegetation to form semifloating mats or hovers. These mats are able to
rise and fall with changing water levels, and this
may be important in keeping the Fen Orchids
at roughly the same level relative to the water
table. (As more peat accumulates, however,
and despite continued cutting or grazing, the
vegetation inevitably changes, and Fen Orchid
will eventually disappear.)
In South Wales (and formerly north Devon)
Fen Orchid is found in dune slacks, growing
with Creeping Willow, Marsh Pennywort and a

variety of mosses. As in the Norfolk Broads, the
species has very specific habitat requirements:
1. The slacks must be calcareous; calcium is
provided by fragments of seashells in the sand.
2. The slacks must be moist (although they may
be flooded in the winter, sometimes for as long
as five months, the water table can fall to more
than 50cm below ground level in August and
September). 3. The slacks must be young.
Fen Orchid can colonise newly formed
slacks very quickly, within a few years of the
slack developing from bare sand. Indeed,
establishment from seed seems to be most
successful in these young slacks, perhaps
because some bare ground remains, and
completely new populations of Fen Orchid
are only found in such new slacks. As the
vegetation becomes more established, however,
it is less able to compete; fewer new plants
appear and eventually there are no new recruits
to the population. Fen orchid will usually die
out in a slack about 50 years after its formation.
Flowering period
In south Wales, early June to late July,
exceptionally as late as mid-September. In
Norfolk, early or mid-June to early July, with a
few appearing later, to late July. In dry seasons
few plants flower, whereas flowering tends to
be retarded in wet years if surface water lingers.
The individual flowers are rather short-lived.
29/1/09 12:25:12
144
GENUS LIPARIS
29/1/09 12:25:15
FEN ORCHID
1987-99
1970-86
pre 1970
145
Non-flowering plants, which may be much

more numerous, are exceedingly hard to
spot. Visitors are asked to keep way from the
Norfolk sites but are welcome at Kenfig in
Glamorganshire. Detailed and up-to-date
directions are required to find Fen Orchid at
this site, and it is best to contact the visitor
centre in advance.
DESCRIPTION
petal
anther
lip
lateral
sepals
column
Range
Much reduced in Britain and now found at
only three sites in the Norfolk Broads and at
three sites in South Wales, in Glamorganshire
and Carmarthenshire, but only in significant
numbers at Kenfig. World range: Temperate
Europe, Asia and North America. Found
north to c. 61N in southern Scandinavia,
Finland and the Baltic States and south
to northernmost Spain, southern France
(including Corsica), northern Italy, Bosnia and
Romania (and formerly also Bulgaria). In Asia
the range extends eastwards to c. 80E in central
Siberia and also recorded in far eastern Siberia
and southeast Kazakhstan. In North America
widespread in the northeast, from Nova
Scotia and New England to Saskatchewan and
Minnesota, ranging south to North Carolina,
Illinois and Iowa. There are also other scattered
populations west to the Northwest Territories,
British Columbia and Washington, and south
to Alabama.
How to find it
This small, green orchid is hard to find. It
often grows in fairly dense vegetation and even
flowering plants can be very inconspicuous.
O 26 June, Norfolk. Flower spikes (or their remains) are
visible from three seasons growth; all presumably arise
from the same psudobulb.
upper
sepal
ovary
flower
stalk
petal
Height: 3-18cm, exceptionally to 30cm.

Stem: Green and three-angled, becoming
almost winged towards the tip. The base of
the stem is enlarged and this elliptical, green,
shiny swelling is enveloped by the bases of the
leaves to form a pseudobulb, the top of which is
sometimes visible.
Leaves: Two or very occasionally three, strapshaped, tapering both towards the base and
the pointed tip. They are held erect and nearly
opposite each other. The leaves are pale green,
shiny and prominently keeled. Immature,
non-flowering plants, with just one leaf, are
common.
Spike: Up to 12 flowers form a loose spike, but
there are usually rather fewer, sometimes just
one, although as many as 17 have been recorded.
Bract: Green, minute, lanceolate.
Ovary: Green, narrow and tubular, six-ribbed,
straight or slightly twisted at the base; the
flower stalk is around half the length of the
ovary, three-angled and twisted.
Flower: The flowers are entirely greenishyellow and face upwards so that all the petals
and sepals lie more-or-less horizontally and the
ovary and column point upwards. The sepals
29/1/09 12:25:16
146
GENUS LIPARIS
are long and narrow (c. 5.5mm long x 1mm

wide) with their edges rolled under, making
them appear even narrower; the lateral sepals
lie parallel below the lip and are often twisted.
The petals are even finer (c. 4.5mm x 0.5mm)
and are usually curved, both downwards in a
gentle bow and forwards in the direction of the
lip. The lip is tongue-shaped, rather broader
than the sepals (c. 5mm x 2.5mm), with a wide
longitudinal groove that is deepest towards the
base, and has slightly wavy or frilly margins. It
points upwards, parallel with the column, and
then bends sharply over at right angles to lie in
a horizontal plane. The column is pale green,

relatively large and prominent, slightly curved
towards the lip (like an erect cobra), with the
anther cap sitting on top and pointing forwards.
There is a minute rostellum and two waxy,
yellow pollinia, each of which is divided into
two flat plates and attached to one of the two
viscidia.
Subspecies
L. l. loeselii is found in Norfolk and has up to
12 flowers and pointed, relatively narrow leaves
which are at least four times as long as broad. It
is widespread but scarce and declining in fens in
continental Europe.
L. l. ovata occurs in South Wales and formerly
also in north Devon. On average it is shorter
and has fewer flowers (usually up to six,
rarely as many as ten). The leaves are rather
more broadly elliptical or egg-shaped, blunter
and more hooded at the tip and held more
consistently erect. It also occurs in dune slacks
in northern France and northern Brittany and
in the early 1900s was present in the Dutch and
German Waddenzee.
There has occasionally been speculation that
the two subspecies should be treated as distinct
species, but recent genetic analysis has shown
virtually no difference between them.

None. (Subspecies ovata is sometimes treated as
var. ovata.)
BIOLOGY
M 21 June, Norfolk.The wispy petals and upwardpointig

column and lip result in flowers that do not resemble most
other orchids and are hard to make sense of.
The Fen Orchid is probably routinely selfpollinating, the process being assisted by rain.
Raindrops hit the anther cap, which lies on top
of the column like a tiny lid. This in turn knocks
the pollinia towards the stigma. The upturned
lip may function to deflect raindrops towards
the anther. The prolific numbers of seedlings
recorded suggest that seed is the major means of
reproduction; in South Wales up to 128 shoots
have appeared in a 0.25m2 plot.
Vegetative reproduction also occurs but
its importance relative to seed is unknown.
29/1/09 12:25:17
FEN ORCHID
147
M 16 June, Norfolk.The flowers are self-pollinated and the lip is thought to deflect raindrops onto the column to knockout the pollinia.
Buds (detachable propagules) are formed on

the swollen stem of the pseudobulb, and these
are dispersed in the autumn, already carrying a
fungal infection.

The aerial stem grows from a pseudobulb,
the swollen tip of the rhizome which acts as
a storage organ and is surrounded by the leaf
sheaths. Each summer, the rhizome continues

its growth from a bud at the base of the existing
pseudobulb, and this new section swells in turn
as it stores nutrients. Normally, only the two
youngest sections of the rhizome, representing
the last two years growth, are alive at any one
time, and so there are two pseudobulbs lying
side-by-side.
29/1/09 12:38:18
148 l GENUS LIPARIS
P 16 June, Norfolk. Flow

ers from above, showing
the threadlike petals,
which, together with the
sepals and lip, form a cross
shape.
Fen Orchid remains partially dependent on

fungi as an adult plant, fungal infection being
concentrated in the rhizome; the pseudobulb
is always fungus-free. A few short, thick, hairy
roots, largely free of fungi, develop in the spring
from the base of the pseudobulb.
Seed germinates in the autumn and the
protocorm produces the first green leaf by the
following August. The first small pseudobulb
develops in the autumn of that year and the
protocorm starts to wither away. No roots are
developed in this first year and all the plants
requirements for water must be met by its
minute root hairs or by its fungal partner. In
the summer of the second year both roots and
small leaves develop and a new pseudobulb is
formed. Fen Orchid can flower in the fourth year
after germination (perhaps as early as the second
year), and plants reach their maximum size from
the seventh year onwards. However, there is a
high mortality among immature plants, and the
vast majority disappear before they can flower.
But, once mature, most flowering individuals
reappear the next season, and they may flower
for several years in succession; the average
lifespan of mature Fen Orchids is ten years.
Occasionally, plants may remain underground
for one year, very rarely two, although it is hard
to be sure in such cases that the leaves have not
appeared and been rapidly grazed off.
Hybrids
None.

The specific name loeselii celebrates Johann
Loesel (1607-1657), a Prussian professor of
medicine and a botanist.
HISTORY aND
CONSERVaTION
1660, when John Ray found it in the watery
places of Hinton & Teversham Moors in
Cambridgeshire (Catalogus Plantarum circa
Cantabrigiam nascentium; Catalogue of plants
found around Cambridge).
A Red Data Book species, it is considered
Endangered and included in Schedule 8 of the
Wildlife & Countryside Act 1981. It has always
been scarce and localised in Britain but was
formerly known from 34-36 sites in eastern
England, one in Devon and a further nine in
South Wales. Fen Orchid is now reduced to
three sites in the Norfolk Broads where the
overall population is stable: two in the Ant
Valley, one holding a few hundred plants, the
other about a thousand, and a much smaller
colony in the Bure Valley, typically with about
10 plants but up to 40 can appear in a very
good year. There are also three sites in South
29/1/09 12:25:21
FEN ORCHID
149
O 16 June, Norfolk. The

flower in profile, show
ing the erect column and
horizontal lip.
Wales where the population totalled over

10,000 plants until recently but has declined to
an unknown extent.
In Norfolk, always the stronghold of the
species, in addition to the three surviving
localities, there were formerly eight more sites
in the Broads, with a further eight colonies
scattered elsewhere in the county, especially the
Waveney-Ouse Valley on the Norfolk-Suffolk
border. In Suffolk there were four or five sites
but the species was last recorded in 1974 at
Thelnetham. In Cambridgeshire there were
eight sites, six of which were drained long ago,
but Chippenham Fen and Wicken Fen remain
intact, although Fen Orchid was last recorded
at these sites in 1928 and 1945 respectively.
In Huntingdonshire there were two sites, the
last record coming from Whittlesea Mere in
1849, but this too was subsequently drained.
Elsewhere in eastern England, Fen Orchid was
recorded from Ham Fen in Kent in 1802 and
was reported in 1884 near Lincoln.
Early losses in East Anglia were largely due
to the drainage of the fens and their conversion
to agriculture. Although drainage and water
abstraction continued to cause extinctions,
most of the more recent losses have been caused
by the decline of peat and turf-cutting, reed
and sedge harvesting and grazing, all traditional
land uses. Even in areas protected as reserves
(indeed, probably especially in protected

areas such as Wicken Fen) this resulted in the
development of rank vegetation, scrub and in
some cases the eventual transformation of the
fen into wet carr woodland. Thus, although
quite a few of the old sites still exist as fens,
changes in the vegetation have rendered
them totally unsuitable for Fen Orchid.
Collecting also took its toll, as demonstrated
by this account of a club outing to Burwell
Fen, Cambridgeshire, in 1835: We had very
good sport both in plants and insects. Ophrys
loeselii was found in great plenty. Between four
hundred and five hundred specimens were
brought home. It was growing in the grass and
moss among the pits where they cut turf. There
were two bulbs to each plant, and the bulbs
were scarcely in the ground at all, so that we
picked them out easily with our fingers (quoted
in Marren 1999). As with most orchids,
however, such wanton destruction was a minor
factor in its decline compared with changes in
its habitat.
In South Wales, Fen Orchid was first
recorded in 1897 from dunes at Pembrey in
Carmarthenshire and was eventually found
at a further eight Burrows (Kenfig, Margam,
Baglan, Crymlyn, Oxwich, Whiteford, Tywn
and Pendine/Laugharne). It was still present
in seven dune systems in the early 1970s but
29/1/09 12:25:23
150
GENUS LIPARIS
by 2001 was reduced to just three: Kenfig,

which still holds a substantial population
and, although there has been some decline,
concerted management efforts have stabilised
numbers; Whiteford, where it is on the point
of extinction, with just two plants in 2004;
and Pendine/Laugherne, where 76 plants were
counted in 1997 but just one was found in
2000. The other sites have either been buried
by sand, dried out, become overgrown or have
been reclaimed for heavy industry.
The dune slack subspecies ovata was also
found at Braunton Burrows in north Devon in
1966, when two flowering spikes were found in
a slack that had been virtually bare of
vegetation in 1953. Fen Orchid was last
seen there in 1987.
Fen Orchid has declined in South Wales
as the habitat has become unsuitable. Overstabilisation of the dunes has meant that few
new slacks have been formed and a lack of
grazing, exacerbated by a decline in the rabbit
population due to myxomatosis, has hastened
the ageing process in the existing slacks. At
Kenfig there was a large amount of mobile sand
after World War Two and many new slacks
were formed. It is this era of slack formation
which has provided the current supply of
suitable habitat: in 1946 40% of the area was
bare sand but this had declined to 2% by 1992
and inevitably there will soon be a sharp decline
in suitable habitat as the existing slacks mature
and are not replaced.
In both Norfolk and South Wales the
problems of Fen Orchid conservation are
similar and so are the solutions. Fen Orchid is
a weedy species adapted to grow in dynamic,
changing environments. It is able to colonise
wet, calcareous habitats with plenty of bare
ground and multiply quickly but is eventually
crowded out as the vegetation matures. There is
a high rate of turnover of individual plants, with
considerable ups and downs in overall numbers.
Many plants are short-lived but large numbers
of seedlings can be produced, and to maintain
a population conditions need to be right for
seedlings to become established.
A variety of conservation measures are being

employed, such as reed and sedge harvesting
in the Broads and scrub removal, grazing, close
mowing and turf stripping in South Wales.
These should provide the plants, both adults
and seedlings, with the appropriate niche.
Many of these techniques are only possible on
a relatively small scale, however, and are interim
measures which will slow but not stop the
inevitable succession to unsuitable habitats. The
only real hope for the Fen Orchids long-term
survival is more radical action: digging peat
from the Norfolk Broads to create fresh turf
ponds; and the destabilisation of the dunes in
South Wales to create a succession of young
slacks. If the right conditions were available,
reintroduction also becomes an option. There
have been small-scale experimental trials at
sites in Norfolk and Suffolk but so far no full
programme has been attempted.
Past and present occurrence of Fen Orchid in Britain and
Ireland (based on presence or absence in 10km squares
of the National Grid; data from the New Atlas).

1500-1999
current range
Britain
Ireland
26
7 (0.25%*)
% lost, 1500-1969
61.5%
% lost, 1970-1986
1.5%
% lost, total
63%
P 13 July, New Forest. Bog Orchids often grow from tiny

bulbils which develop on the edge of the leaf and can thus
form small groups.
29/1/09 12:25:24
Genus
HAMMARBYA
BoG orchid
Distribution
Bog Orchid is the only member of this
genus, which has a circumpolar distribution
in Europe, Asia and North America.
Name
The generic name Hammarbya has an
auspicious history. In the mid-18th century
the great Swedish naturalist Linnaeus gave
Bog Orchid the name Ophrys paludosa.
As the relationships between the various
orchids came to be better understood, Bog
Orchid was moved from Ophrys to its own
genus by Otto Kuntze in 1891. He named
the new genus Hammarbya in honour of
Hammarby, Linnaeuss summer residence
near Stockholm.
29/1/09 12:25:28
152
GENUS hAMMARbyA
BoG orchid
Hammarbya paludosa
Formerly: Malaxis paludosa. Other names: Bog Adders-mouth (North America)
This tiny green orchid is the exact opposite of the classic big, bright and blousy hothouse
hybrid. Indeed, it is the smallest orchid in the British Isles and one of the hardest to find.
But, despite its rather dull flowers and diminutive stature (or perhaps because of these),
it holds a particular fascination for botanists and is always a delight to see. It is still fairly
common, at least locally, in northwest Scotland but has declined markedly in the lowlands
and in England is now only frequent in the New Forest.
Identification
The small size, tiny green flowers and habitat
make this orchid distinctive when and if you
can find it.
Similar species
Of the plants found in its boggy habitat, Marsh
Arrowgrass is closest in general appearance but
has very different flowers.
Habitat
As its name suggests, this species is found
in bogs. It is associated with a good cover
of Sphagnum (bog-mosses) and grows
alongside sundews, butterworts, cottongrasses,
White Beak-sedge and Cross-leaved Heath.
Importantly, it requires a bog which has a flow
of water through the peat, and it does not like
stagnant conditions; as well as streams and
runnels, the slow flow of water may even be
evident on the surface of the peat. Bog Orchid
is often found in the vicinity of such moving
water and also close to the shores of lakes and
lochs. The ground water is usually moderately
acidic but in parts of Wales the species is also
recorded from areas flushed with alkaline water.
Another essential requirement is that the bog
must not dry out, even in a hot summer. Bog
Orchids often grow on carpets of Sphagnum
but can also be found on bare peaty mud or in
denser vegetation amidst sedges, grasses and
O 13 July, New Forest. The flower spike continues to
grow as the flowers open and becomes significantly taller.
This plant was 12cm high.
29/1/09 12:25:29
BOG ORCHID
small shrubs. Generally found in the lowlands

but it has been recorded up to 500m above sea
level (Llyn Anafon, Caernarvonshire).
153
1987-99
1970-86
pre 1970
Flowering period
Late June to mid or even late September but
perhaps most reliably from early July to midAugust. It tends to flower early in a hot summer
and later in a cool, wet season. Numbers are
erratic, with wide variations between years.
Flowering at any one site is not necessarily
synchronised, however, and the flowers
themselves are long-lasting, and thus larger
colonies may have at least some plants in bloom
over a lengthy period.
Range
Bog Orchid is widespread in Scotland. There
are a few sites in the southwest, in Dumfriesshire and Kircudbrightshire, and in the eastern
Highlands in Angus and south Aberdeenshire.
The majority are, however, in the northwest
where it is found north to Sutherland and
extends to the Inner and Outer Hebrides and
to Yell in Shetland. In Ireland, Bog Orchid is
known from relatively few, scattered localities;

in the west in Co. Clare and from Co. Mayo
to Co. Donegal; in the east in Co. Carlow,
Co. Kilkenny, Co. Wicklow and Co. Dublin,
and in Northern Ireland in Co. Armagh, Co.
Down and Co. Antrim. In Wales it is also
recorded at just a few, scattered sites, mostly in
the west, from Breconshire, Carmarthenshire
M 29 June, New Forest. Although often hard to find, Bog Orchids can be conspicuous at times, contrasting as here with
a carpet of russet Sphagnum.
29/1/09 12:25:30
154
GENUS hAMMARbyA
29/1/09 12:25:33
BOG ORCHID
and Pembrokeshire north and west to

Caernarvonshire, but often there are just one or
two sites in each county. In England the species
is now very local, occurring in Cumbria, at a
handful of sites in northwest Yorkshire and
Northumberland and at one site in Norfolk.
In southern England it is known from a
very few localities in Cornwall and Devon
(including Dartmoor, although it is very rare
there) but occurs much more commonly in the
Purbeck region of Dorset and especially in the
New Forest in Hampshire. World range: In
Europe it occurs north to the Faeroe Islands
and to c. 69N in Scandinavia, and south in
the mountains to France, the Italian Alps, the
former Yugoslavia and Romania. In Asia there
are scattered records from across southern
Siberia to Sakhalin and Japan. In North
America it occurs from Alaska eastwards to
western Ontario and south to Minnesota.
How to find it
This tiny green plant is one of the hardest
British orchids to find. It is so inconspicuous
that it is easy to tread on it unawares, and the
pseudobulbs, often only half-buried in the
moss, can easily be dislodged. All in all, it is best
not to look for it until you are an experienced
orchid hunter.
Faced with a large area of apparently
suitable habitat, the task of searching for Bog
Orchid may be daunting. It is best, however, to
concentrate on the most suitable areas. These
are likely to be along the edges of streams
and runnels or where there is some obvious
movement of surface water and away from
dense stands of shrubs such as Cross-leaved
Heath and Bog Myrtle. Like all our smaller
orchids, Bog Orchid is difficult to see from
a walking height and becomes much more
obvious when viewed from a low vantage point.
On open, flat Sphagnum carpets or lawns it
can then be fairly conspicuous, but plants may
grow among grasses and sedges and these are
extremely difficult to see; dense hummocks of
O 29 June, New Forest. Growing as usual on a carpet of
Sphagnum, minute bulbils are starting to appear on the
margins of the leaves.
155
Sphagnum are not favoured but the orchids can

be found around their edges, especially if they
are close to water.
Once a Bog Orchid has been seen, before
diving in for a closer look, it is wise to make
a very careful inspection of its immediate
surroundings. This species often grows in
scattered groups, and one or two relatively
obvious plants may be accompanied by several
others hidden in the vegetation at ones feet.
DESCRIPTION
lateral
sepals
ovary
lip
lip
column
petals
column
upper
sepal
flower
stalk
upper
sepal
bract

Stem: Yellowish-green and three to five angled.
The stem grows from a pea-sized pseudobulb
which is covered by the leaves and often only
half-buried in the moss.
Leaves: There are two or three (occasionally four)
oval to oblong, fleshy, pale-green or yellowishgreen basal leaves. They are prominently veined,
sheathe the pseudobulb at their base, and their
margins and tips are strongly curved inwards
giving them a hooded appearance. One to three
minute, triangular, scale-like leaves are scattered
higher along the stem.
Spike: Up to 25 flowers are carried in a spike
which is dense at first but elongates and
becomes much more open as it matures.
Bract: Green, narrow and pointed, about as
long as the ovary.
Ovary: Green, ovoid and just a little fatter than
the flower stalk, which is about three times its
length and twisted through 360 (therefore the
lip is held uppermost and the flower is hyperresupinate).
29/1/09 12:25:34
156
GENUS hAMMARbyA
BIOLOGY
The flowers are probably pollinated by gnats
and tiny flies, attracted to the nectar at the base
of the lip, and the pollinia are usually removed.
Seed-set is good. Perhaps more importantly
in terms of reproduction, the leaves are often
fringed with numerous, minute, protocorm-like
buds, called bulbils. These drop off and can
develop into new plants if they are infected
from the soil by fungi of the right species
(unlike the propagules of Fen Orchid, they do
not carry fungi from the mother plant). This
may account for the frequency with which Bog
Orchid is found in small groups.
M 7 July, Norfolk.The lip is uppermost (the flower stalk is
twisted through 360), and the lateral sepals wrap around
the back of the flower.
Flower: Greenish and tiny, about 2mm wide

by 4mm tall. The sepals are yellowish-green
and tongue-shaped; the dorsal sepal points
downwards and is slightly longer than the lateral
sepals, which point upwards. The petals are
rather smaller and narrower, strap-shaped and
green. They are held spreading horizontally but
curve sharply back around the sepals to clasp
the flower. The lip is dark green with paler green
longitudinal stripes and is rather shorter than
the petals, triangular and curled upwards at
the sides. It is held erect, pointing upwards and
forwards between the two lateral sepals and
clasping the column at its base. There is no spur,
and the very short, broad, green column projects
horizontally from the centre of the flower, with
the lid-like anther shrivelling to expose the
pollinia shortly after the flower opens. The two
pollinia are each made up of two thin plates of
waxy pollen, and the minute rostellum is topped
by a small, sticky mass. The flowers are said to
have a sweet, cucumber-like scent.
Subspecies
None.

None.

The rhizome lies almost vertically in the peat
or Sphagnum, and a swelling is formed each
year at its tip. In the spring leaves develop
around the base of this swelling and the
flower spike grows from a bud on the top;
the swollen stem and leaf sheaths combine
to form a pseudobulb 4-8mm in diameter.
After it has flowered the aerial stem dies off,
but the rhizome continues to grow from a
bud at the base of the pseudobulb and at the
end of the growing season again terminates
in a swollen internode. Although in theory
a whole string of pseudobulbs could be
produced (as seen in many epiphytic orchids),
in practice only the two most recent are alive;
the older, lower, pseudobulb is buried in the
moss and surrounded by the remains of the
previous years leaves. Unusually, the rhizome
of Bog Orchid grows vertically with the two
pseudobulbs one above the other. This vertical
growth pattern presumably allows the orchid to
adjust to the changing level of the bogs surface
due to, for example, the growth of Sphagnum.
There are no roots, merely root hairs, and
Bog Orchid is largely dependent on fungi
throughout its life cycle; the rhizome and leaf
bases are infected at all times. The pseudobulb,
which acts primarily as a storage organ, is
separated from the older parts of the rhizome
29/1/09 12:25:34
BOG ORCHID
157

The specific name paludosa means growing in
boggy ground.
Bog Orchid is sometimes placed in Malaxis,
a genus of rather similar orchids. It is also
closely related to Fen Orchid and could easily
be united with it in the genus Liparis, but the
current evidence is that it is sufficiently distinct
to justify its continued separation in its own
genus, Hammarbya.
HISTORY aND
CONSERVaTION
The first British records, from Hertfordshire
and Kent, were published in John Parkinsons
Theatrum Botanicum (The Theater of Plantes)
of 1640: Bifolium palustre. In the low wet
grounds between Hatfield and S. Albones; in
divers places of Romney Marsh.
protected in Northern Ireland under the
1985 Wildlife Order (NI) and in Eire under
the Flora (Protection) Order. Bog Orchid
is threatened and declining throughout its
European range due to loss of habitat, and some
of the largest populations are now found in
western Scotland and the New Forest.
M 13 July, New Forest.The lip, which is much smaller than
the sepals, is boldly striped with pale and dark green.
by a band of lignified tissue. This effectively cuts

off one years growth from the next and prevents
fungi from reaching the pseudobulb and its
store of nutrients. An internal root is produced
at the base of the new segment of the rhizome,
and this grows down through the lignified
barrier into the older segment, apparently
reabsorbing nutrients and water from the
decaying tissue and, bypassing the pseudobulb,
carrying the fungal infection into the new
segment.
Life expectancy and the period between
germination and flowering are unknown.
Hybrids
None.
Past and present occurrence of Bog Orchid in Britain and

Britain
Ireland
302
44
118 (4%*)
15 (1.5%*)
% lost, 1500-1969
47%
59%
% lost, 1970-1986
14%
7%
% lost, total
61%
66%

1500-1999
current range
Systematic searches have produced records

from many new localities in recent years but,
despite this, Bog Orchid has vanished from
61% of its historical range in Britain and
66% in Ireland. It is now extinct in most of
England, including Kent, East and West Sussex
29/1/09 12:25:35
158
GENUS hAMMARbyA
O 29 June, New Forest. As

in Fen Orchid, the aerial
stem grows from a pseudo
bulb, a swelling in the stem
concealed by the base of
the leaves; a second, non
flowering plant is growing
alongside.
(where it was formerly found in several areas,

especially Ashdown Forest, but was last seen
in 1956), Surrey, Hertfordshire, Bedfordshire,
Cambridgeshire, Suffolk, Lincolnshire,
Staffordshire, Cheshire, Lancashire and Co.
Durham. The dramatic decline in lowland
Britain started with the Enclosure Acts of
the late 18th and early 19th centuries and the
consequent reclamation of bogs and wet heaths.
Habitat destruction has continued to the present
day, and in the lowlands the remaining areas of
mire and heath have also suffered from a lack of
grazing, necessary to maintain the open sward
that the Bog Orchid requires. Conversely, in the
uplands, overgrazing may have caused suitable
habitats to be degraded.
Many of the remaining colonies are small,

with just a few plants. Even in its strongholds
Bog Orchid is uncommon to rare; in Cumbria
there are around ten sites, with just two
populations regularly exceeding 100 spikes.
In the New Forest there are around 30
populations, with over 200 spikes appearing
in good seasons at two or three of the largest,
and in Dorset there are nine recorded sites.
In Norfolk, which holds the only remaining
population in eastern England, there is now just
one site, and in recent years there have
been only one to four flowering plants in an
area where 117 were counted in 1910; to add
insult to injury all the flowering plants were
stolen in 2000.
29/1/09 12:25:37
Distribution
A genus of 11 species, exclusively found in North
and Central America with the exception of
Coralroot Orchid C. trifida, which extends to the
boreal zone in Europe and Asia. All Corallorhiza are
largely or totally dependent on fungi as a source of
nutrition throughout their life.
Name
The generic name Corallorhiza derives from the
Greek korallion coral and rhiza root, thus coralroot;
there are no roots, however, only a rhizome, and this
does not resemble coral in all species.
Genus CORALLORHIZA
coralroot orchids
29/1/09 12:25:40
160
GENUS CORALLORhIZA
coralroot orchid
Vulnerable
Corallorhiza trifida
Other names: Early Coralroot (North America)
This tiny orchid lacks green leaves and throughout its life is largely parasitic on its fungal
partner, living entirely underground apart from the brief period during which the flower
spikes are produced. It is found throughout the Northern Hemisphere, and its generally
northerly distribution is reflected in Britain where it is confined to Scotland and northern
England. Both the English name and the generic name Corallorhiza refer to the shape of the
underground rhizome, which is said to be coral-like. Coralroot Orchid is the county flower
of Fife.
Identification
With no leaves (merely scale-like sheaths on
the stem) and tiny greenish-white flowers, this
species is very distinctive.
Similar species
None. Birds-nest and Ghost Orchids also lack
green leaves but are very different. Birds-nest
Orchid is usually taller and always more robust,
with large, honey-coloured flowers, and Ghost
Orchid has a proportionally much larger flower
with the lip uppermost and is only found, very,
very rarely, in southern England.
Habitat
Coralroot Orchid is found on permanently
damp ground with a good layer of peaty organic
matter or moss, including Sphagnum. It can
grow both in full sunlight and in the shade,
even heavy shade, and favours mildly acidic soils
which are low in nutrients. It is commonest in
wet willow and alder carr growing on raised
bogs and around lochs. Its other favoured
habitat is a damp dune slack with a carpet
of Creeping Willow but Coralroot Orchid
appears to have rather exacting requirements
regarding the level of the water table and does
not like prolonged flooding (for example, on
P 7 June, Northumberland. Among Creeping Willows in
a dune slack; Coralroot Orchid extracts nutrients from the
willow, via its fungal partner.
O 7 June, Northumberland. In dune slacks Coralroot
Orchid usually has a stout, mahoganybrown stem. Like all
orchids growing in dunes, it often has a scorched look, in
this case with the tips of the sepals blackened.
29/1/09 12:25:41
CORALROOT ORCHID
161
Lindisfarne in Northumberland a slack that

regularly held 150 flower spikes now supports
only 20 after several wet years). Rather less
frequently, Coralroot Orchid is found in birch
and pine woodland, including plantations, and
in overgrown scrubby fens with a mixture of
Sphagnum, sedges and willows. It is found up to
365m above sea level (Braemar, Aberdeenshire).
Coralroot Orchid is parasitic on a group
of fungi that form mycorrhizal relationships
with birches and willows (also pines in North
America and presumably also in Scotland). The
presence of both the correct fungal partner and
one of these trees is essential and may explain
the very local occurrence of Coralroot Orchid.
Flowering period
May to August. In dune slacks it flowers in
May and early June, increasingly in recent
years from early May onwards, and emerging
earlier in drier slacks compared to wetter ones.
Woodland plants are mostly in bloom from
early June to late July.
Range
Widespread in eastern and central Scotland,
from the Borders north to Caithness. However,
it is absent from the west, apart from Ayrshire
and Renfrewshire. In England it is very
local, occurring in Northumberland and in
Cumbria, where it is found in dune slacks at
Eskmeals, North Walney and Sandscale. The
1987-99
1970-86
pre 1970
M 6 June, Cumbria.
southernmost records come from Ribblesdale

in Yorkshire. World range: This species has
a circumpolar distribution, occurring very
widely in northern Europe, Asia and North
America, ranging southwards locally where
mountains provide a suitable habitat. It occurs
north to Greenland, Iceland and northernmost
Scandinavia and south to the Pyrenees, Massif
Central, Alps, Apennines and Balkans, also
Corsica, the Crimea and Caucasus. In Asia,
primarily boreal but with isolated outposts in
the mountains of Central Asia and China. In
North America occurs from Newfoundland
east to Alaska and south in the mountains to
California, New Mexico and West Virginia.
How to find it
Small and leafless, this is an inconspicuous
orchid and very hard to spot. The number of
flowering spikes can also vary greatly from
season to season. As with all such diminutive
orchids, scanning from a low level can be
productive. Dune slack populations are the
easiest to localise, with one of the biggest at
Sandscale in Cumbria.
29/1/09 12:25:42
162
GENUS CORALLORhIZA
DESCRIPTION
Height: 5-30cm, usually 10-13cm.
Stem: Usually yellowish-green in woodland
plants but tends to be mahogany-purple in
dune populations. Frequently found in small
groups and up to ten spikes may develop from
one rhizome.
Leaves: There are no green leaves, merely two
to four long, membranous, sheathing scales on
the lower half of the stem, and these may be
brown, whitish or green.
Spike: There are four to nine flowers
(occasionally as many as 13) held pointing
outwards and slightly drooping in an open,
lax spike.
Bract: Green, minute, triangular and pointed.
Ovary: Green or mahogany-purple, spindleshaped, six-ribbed, on an extremely short,
twisted stalk.
Flower: The sepals and petals are strap-shaped,
with the petals slightly smaller; both are
greenish-yellow, often tinged reddish-brown
around the fringes and tip and thus apparently
browned off . The interior of the petals may
also be blotched with reddish-brown. The
lateral sepals curve inwards and are held
forward and slightly drooping on either side
of the lip. The upper sepal and petals form a
loose hood. The lip is tongue-shaped, shorter
and broader than the petals and sepals, with
a ruffled margin. A central groove, which
may produce nectar, runs between two raised
longitudinal ridges and there is a very short
spur. The lip is kinked downwards towards
the base and, although technically three-lobed,
the tooth-like side-lobes near the base are very
small. It is white, spotted with crimson at the
base, and these spots may occasionally coalesce
into a larger blotch. The column is long, green
and curved, the lid-like anther lies on the top
and there are four waxy yellow pollinia, a small
rostellum and two distinct viscidia. The flowers
are slightly scented, with the perfume reported
to b e musk-like.
M In woodland Coralroot Orchid has a green stem and
ovaries and is relatively slender (Paul Sterry/Nature
Photographers Ltd).
Subspecies
None.
29/1/09 12:25:44
CORALROOT ORCHID
163

Plants growing in dune slacks tend to be
shorter than those in woodland, with reddish
rather than greenish stem and ovaries, but there
are no named varieties.
BIOLOGY
The flowers are routinely self-pollinated. The
rostellum is small and degenerates quickly,
and the pollinia crumble apart, falling onto
the stigma below; 85-100% of flowers set
seed. Small insects, including flies, wasps
and beetles, visit the flowers, but the pollinia,
although easily detached, do not readily stick
to the insects and any cross-pollination is
purely accidental; visiting insects are probably
more effective in nudging fragments of the
disintegrating pollinia onto the stigma below.
Vegetative reproduction may also occur,
via fragmentation of the rhizome, the sidebranches elongating and producing new plants.

The aerial stems grow from a creeping,
horizontal, underground rhizome. This is a
much-branched mass of cream-coloured, fleshy,
coral-like knobs. There are no roots and water
must be absorbed either through tufts of root
hairs or via the fungal partner.
The rhizome is permanently infected
with fungi and, throughout its life, Coralroot
Orchid is almost completely dependent on
its fungal associate for nutrients. Like Birdsnest Orchid, it is very fussy and only forms a
relationship with the Thelephora-Tomentella
complex of fungi (family Thelephoraceae). This
group of fungi, apart from its relationship with
orchids, is exclusively ectomycorrhizal, forming
symbiotic relationships with the roots of trees
(see p.8). The fungi that Coralroot Orchid
partners are species that simultaneously attach
themselves to the roots of willows, birches and
pines. It has been shown in the laboratory that
the orchid obtains carbohydrates from the trees
via their mutual fungal partner, but the orchid
undoubtedly cheats in its relationship with the
M The tiny flowers are usually selfpollinated and most

flowers set seed (Bob Gibbons/Natural Image).
fungus-tree partnership, receiving nutrients but

giving nothing in return. It is therefore parasitic.
(Coralroot Orchid is not a saprophyte; it gets
its nutrients from a living fungus, not dead
organic matter.)
Coralroot Orchid does have a limited ability
to photosynthesise as the stem, ovary and scalelike leaves on the stem contain chlorophyll.
Even in diffuse daylight such photosynthesis
can contribute in a small way to its overall
nutritional budget.
Seed germinates from the spring onwards
but may remain dormant until the spring of the
second year, perhaps even longer. Germination
will, however, only take place when the seeds
have been colonised by the appropriate fungi.
The seedling is initially a globular protocorm
which develops scattered root hairs. It then
elongates and starts to branch; each time the
rhizome branches, the main rhizome bends
in one direction and the side branch goes off
in another, producing the coralloid growth
29/1/09 12:25:44
164
GENUS CORALLORhIZA
P 6 June, Cumbria. The

lip has a central groove
that may produce nectar;
this would attract insects
but despite this the flowers
are usually selfpollinated.
pattern. After as little as nine months, some

seedlings have developed into a branched
rhizome 15-25mm long with the bud for the
aerial shoot already well-developed. Flower
spikes may be produced two to five years after
germination but there is a little uncertainty
about the exact timing. Some related species
of Corallorhiza are monocarpic and die after
flowering once.
Hybrids
None.

The specific name trifida comes from the Latin
for split into three, a reference to the shape of
the lip. This is barely appropriate, as the lip is
only very slightly three-lobed.
HISTORY aND
CONSERVaTION
1777 by the Rev. John Lightfoot, curate of
Uxbridge. In Flora Scotica he noted: Ophrys
Corallorhiza In a moist hanging wood near
the head of Little Loch Broom on the western
coast of Ross-shire.
Nationally Scarce and classified as
Vulnerable. This small, inconspicuous orchid
is hard to find, and the true picture of its
status and distribution is still emerging. Efforts
have been made to seek it out in recent years
(especially for the New Atlas scheme), and these

have turned up many new sites. But, despite
these new records, there has been a net loss
of 46% of the total historical distribution and
many of the losses are relatively recent. Many of
its habitats seem relatively secure, however, and
importantly many are not affected by changes in
agricultural practices. It is possible that climate
change could be responsible as many northern
plants are currently in retreat.
Woodland populations are usually small,
scattered and hard to find but in some dune
slacks there can be large numbers; at Sandscale
in Cumbria at least 3,000 plants were counted
in five slacks in 1991, making it the largest
English population.
Past and present occurrence of Coralroot Orchid in Brit
ain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
102
55 (1.9%*)
% lost, 1500-1969
17%
% lost, 1970-1986
29%
% lost, total
46%
29/1/09 12:25:46
Genus GOODYERA
ladys-tresses (i)
Distribution
A genus of 40-100 species, nearly worldwide in
distribution although most are found in South-East
Asia. Some are called jewel orchids and are known
for their beautiful foliage. Only two species occur
in Europe, the widespread Creeping Ladys-tresses
and G. macrophylla, endemic to Madeira.
Name
The genus Goodyera is named in honour of John
Goodyer (1592-1664), a manorial steward at
Petersfield, Hampshire, who has been dubbed the
first amateur naturalist and the Cavalier botanist.
29/1/09 12:25:49
166
GENUS GOODyERA
creepinG ladys-tresses
Goodyera repens
Other names: Lesser Rattlesnake Orchid, Lesser Rattlesnake Plantain (North America)
This small, white-flowered orchid is almost always found growing under pine trees and is
locally common in Scotland, the home of the only native pine woodland in the British Isles.
A few colonies in old pine plantations in Norfolk represent an enigma. The ladys-tresses
could have been introduced accidentally with the pine seedlings, could be indigenous to the
area and have simply moved into a congenial new home, or could have arisen from windblown seed.
Identification
The spikes of small, densely hairy, white flowers
are distinctive. Creeping Ladys-tresses is
evergreen and can be found and identified all
year. Indeed, it is often easier to locate in winter
when much of the other vegetation has died
down. It forms small patches of rosettes which
are composed of small, oval, dark-green leaves
rather like Garden Privet in size, shape and
colour. Notably, the veins on the leaves form
a faint net over the surface; almost all other
British orchids have veins that are parallel.
Similar species
The other species of ladys-tresses belong to
the genus Spiranthes but are nevertheless rather
similar. They have small white flowers which
have a covering of glandular hairs, although
none of them are as densely hairy as Creeping
Ladys-tresses. Their flower structure is similar
but the Spiranthes have the tip of the lip broadly
frilled or crimped, rather than being a simple
unadorned wedge shape.
Autumn Ladys-tresses is very unlikely
to be found in the same pinewood habitat as
Creeping Ladys-tresses (although it did occur
until 1979 in a grassy woodland ride within
a few metres of Creeping Ladys-tresses at a
remarkable site at Holt in Norfolk). Once the
possibility of confusion is acknowledged, the
two species can be separated easily because
P 11 July, Norfolk. Typically grows under pines, some
times mixed with birch or, as here in Norfolk, oak.
O 10 July, Norfolk. Creeping Ladystresses.
29/1/09 12:25:51
CREEPING LADYS-TRESSES
Autumn Ladys-tresses has its flowers arranged

into a distinct row that is often spirally twisted.
Irish and Summer Ladys-tresses grow from
a rosette of long, strap-shaped leaves which
have parallel veins, and are rather taller than
Creeping Ladys-tresses; their flowers are also
arranged into a more definite pattern.
167
1987-99
1970-86
pre 1970
Habitat
Creeping Ladys-tresses is found in mature
pinewoods with a damp, well-shaded forest
floor, growing in the deep humus formed by
the accumulation of dead pine needles. It
can be found in areas with a relatively open
understorey of grasses and small shrubs, such
as Heather and Bilberry, or in places where
there is merely a carpet of moss over the needles
(sometimes even bog mosses, Sphagnum),
indicating a suitably moist microclimate. The
classic habitat is ancient Caledonian woodland
where Scots Pine is mixed with birches. It
has also spread into pine plantations that are
sufficiently mature to have a shaded, moist

open, forest floor. A balance of light conditions
is required; too little light and the orchid
will not flower, too much sun and a dense,
overwhelming shrub layer develops. Rarely,
M 10 July, Norfolk. Growing in an old pine plantation on sand dunes.
29/1/09 12:25:52
168
GENUS GOODyERA
Creeping Ladys-tresses is found on damp

dunes or among Heather and Bell Heather
on moorland, sometimes far from woodland
(perhaps most often in coastal areas; for
example on a peat moor at Auckengill Loch
in Caithness). Unlike most British orchids,
Creeping Ladys-tresses is found on acid as well
as neutral soils. Most sites are in the lowlands
but it is found up to 335m above sea level
(Morinsh, Banffshire).
Flowering period
Late June to late August but often at its best in
mid-July. In dry weather the individual flowers
are short-lived.
Holkham, although the colonies there are

scattered and hard to find.
DESCRIPTION
Height: 7-20cm, occasionally to 35cm.
Stem: Pale green and ridged, with dense
glandular hairs towards the tip.
Leaves: The flower spikes grow from a rosette
of three to nine dark-green (almost blue-green)
leaves which have a network of faint paler veins
(reticulations). The leaves are oval and taper
to a pointed tip and the leaf-stalk is short but
broad (winged). They are held more-or-less flat
to the ground, although there may be one or
Range
Northern and eastern Scotland, with an
isolated population in Ayrshire, also northern
England in Cumbria, Northumberland and
Co. Durham, with an outpost in Norfolk (see
History and Conservation). World range:
This species has a circumpolar distribution
and occurs throughout Europe, northern Asia
and northern North America. Found north
to 70N in Scandinavia and south to the
Pyrenees, Corsica, Alps, northern Apennines
in Italy, northern Greece, the Crimea and the
Caucasus, although confined to the mountains
in the south of the range. In Asia it ranges
across Siberia to the Kamchatka Peninsula,
Sakhalin and Japan and is also found in the
mountains of southern Asia in Turkey, Central
Asia, Afghanistan, the Himalayas and in a band
from southwest China to Korea. In northern
North America it occurs from Alaska east
to Newfoundland and south in the Rocky
Mountains to New Mexico and Arizona and in
the Appalachians to North Carolina.
How to find it
Often abundant where it is found, any site in
the Caledonian pinewoods of Scotland could
yield this species. In Norfolk the well-known
population at Holt Country Park was largely
destroyed by forestry operations in 2002
(but may recover in time), and the species is
best looked for in the coastal pines at Wells-
M 14 July, Norfolk. The stem, bracts and flowers have

abundant glandular hairs.
29/1/09 12:25:53
two leaves at the base of the flower spike, and

several very small, long and narrow sheathing
leaves further up the stem. Sterile, nonflowering rosettes are also produced and the
leaves are evergreen.
Spike: The 5-25 flowers are arranged spirally
around the stem but mostly face in the same
direction, and the spiral pattern is seldom
obvious.
Bract: Green, with scattered glandular hairs
towards the base. The bracts are lanceolate and
slightly longer than the ovary, which they clasp.
Ovary: Green, narrowly pear-shaped, tapering
towards the base, three-ribbed and with
glandular hairs. The ovary is held upright but
bends at the tip, so that the flowers lie either
horizontally or are slightly drooped, and is
either stalkless and slightly twisted or has a
short stalk, in which case it is the stalk that is
slightly twisted.
Flower: Small, creamy-white and densely hairy.
The sepals are oval and concave, with the upper
sepal a little narrower than the lateral sepals;
they may have a faintly greener midrib and
have many conspicuous long, glandular hairs
on their outer surface. The petals are similar
but slightly smaller and more spatulate. The
upper sepal and petals form a tight hood and
the lateral sepals are held drooped and slightly
spreading with their tips bent outwards. The
lip is white and is shorter than the sepals, oval
in shape with a pointed tip; the basal hypochile
is sometimes tinged pink and forms a deep,
rounded, bag-shaped pouch that contains
nectar. The epichile forms a narrow, blunttipped triangle, folded into a shallow groove and
bent downwards towards the tip. The column is
creamy and the rostellum is formed into short,
curved horns which enclose the single, roughly
circular viscidium. The anther cap is ochre with
a reddish-brown margin and the two pollinia
are yellowish. The flowers have a sweet scent.
Subspecies
None.

None.
169
BIOLOGY
The flowers are thought to be pollinated by bees
which are attracted to the nectar at the base
of the lip. Bumblebees of the genus Bombus
are usually mentioned as pollinators and have
certainly been seen carrying pollinia, but
smaller bees of the genus Lasioglossum may be
more important. The mechanism of pollination
is thought to be very similar to that of Autumn
Ladys-tresses and other orchids in the genus
Spiranthes. The flower tube is initially only wide
enough to allow an insects proboscis to enter;
this trips the mechanism and removes the
pollinia. The lip then moves slowly downwards,
allowing access to the stigma, which becomes
sticky and receptive. Fragments of pollinia,
picked up from other flowers and attached
to a visiting insects proboscis, can then effect
pollination. The mechanism is effective and
seed-set is good, with 77% of flowers setting
seed in one Scottish study.
Vegetative reproduction may be more
important than reproduction from seed. Buds
are produced on the tip of the rhizome in the
autumn, and these grow into slender runners
which grow horizontally through the cushion
of moss. At first these runners have merely
a few short, sheathing scales, but eventually
green leaves are produced from buds at or near
their tip (a period of five years was given by
Summerhayes (1968) for the production of
green leaves but this seems excessively long).
Once it has appeared above ground, the new
rosette will live for between two and eight years
before flowering and dying (and in Norfolk a
period of at least six years has been recorded
between the first appearance of rosettes and
the production of flowers). Each runner can
produce a separate plant; after flowering the
central mother plant dies off, leaving the
surrounding rooted runners as separate entities.
In this way large patches can form.

The aerial stem grows from a creeping rhizome
which is often only shallowly buried and which
29/1/09 12:25:54
170
GENUS GOODyERA
ground and assume an existence totally

dependent on fungi.
Seeds probably germinate in spring, but
the period between germination and the first
appearance above ground is not known.
Hybrids
None.

The specific name repens derives from the
Latin and means creeping. The American name
Lesser Rattlesnake Plantain probably derives
from it use, including a poultice of the chewed
leaves, in the treatment of snakebites.
HISTORY aND
CONSERVaTION
M 10 July, Norfolk. The flower structure is very similar to

the Spiranthes ladystresses, but the lip forms a simple
spoutshape rather than being crimped or frilled.
puts out a few short, thick, fleshy roots that

have numerous hairs. Both rhizome and roots
are heavily infected with fungi, and the species
is probably dependent on its fungal partner to a
significant extent throughout its life.
Creeping Ladys-tresses is the only British
orchid which is evergreen (some other species,
such as Autumn Ladys-tresses and Early
Spider Orchid, bear leaves through the winter
but are leafless and dormant for at least part
of the year). Unlike most other orchids, this
species shows no tendency to disappear below
The first published British record dates from

1777, when Rev. John Lightfoot (1735-1785),
curate of Uxbridge, noted in his Flora Scotica:
We found it in an old shady hanging birch
wood about two miles from the head of
Little Loch Broom Ross-shire. (The species
had, however, been found a few years before by
a Scotsman, James Robertson (c. 1745-1796),
in a wood called Cregenon, but this was not
published at the time.)
Nationally Scarce and the subject of a
Plantlife International Back from the Brink
project. It is common to locally abundant
in northeast and north-central Scotland,
especially in the relict Caledonian pinewoods
of the Strathspey and Cairngorm regions,
but it is scarcer in the west of Scotland and
northern England. Colonial, it may occur in
large numbers, and we have counted 620 flower
spikes in an area of only about 50m x 20m.
Although still locally common, there has
been a significant reduction in the range, and
it has now gone from 44% of its historical
distribution. It is extinct in Dumfries-shire,
West Lothian, Peebles-shire and Orkney
(where recorded in the 1950s). There are
also old records from southeast Yorkshire
(1888, Houghton Hall Woods, near Market
Weighton) and east Suffolk (1932-1935,
Stuston Common, a site with no pine trees).
29/1/09 12:25:55
Past and present occurrence of Creeping Ladystresses

in Britain and Ireland (based on presence or absence in
Atlas).

1500-1999
current range
Britain
Ireland
186
104 (3.6%*)
% lost, 1500-1969
29.5%
% lost, 1970-1986
14.5%
% lost, total
44%
With its requirements for a certain

minimum level of shade in order to thrive,
Creeping Ladys-tresses is vulnerable to
woodland management. Colonies can survive
even if a wood is felled and replanted, but it
takes many years for the number of flowering
plants to recover in these circumstances and
the woods must be replanted with pines;
many sites have been lost when restocked
with alien conifers such as spruce and fir. The
extensive thinning of plantations can also have
a very adverse effect, although perhaps not a
permanent one.
The status of the species in Norfolk has
been the subject of controversy since its
discovery in 1885 at Westwick. Subsequent
records came from Holt and adjacent
localities from 1890, Beeston Regis from
1900, Sheringham from 1909, Cawston from
1910, Wells from 1952, Horsford from 1958,
Holkham from 1962 and Cranwich from
1965.
Scots Pine is not native to Norfolk, but
there are plantations of conifers dating from
at least the 1830s. It has been widely assumed
that Creeping Ladys-tresses was accidentally
introduced with pine seedlings brought in from
Scotland, but there is little evidence to support
this. Investigations by W.H. Burrell in 1909
established that pines supplied by a selection
of nurseries in Scotland came bare-rooted
and thus free from weeds and that Creeping
Ladys-tresses was unknown as a weed in these
171
nurseries. The trees were, however, sometimes

packed in dry Bracken, Heather and moss
collected from the woods and this packing
could have been a source of seeds or plants.
An alternative theory is that Creeping
Ladys-tresses was present in Norfolk before
the advent of pine plantations and merely
moved in when they became suitable. There
is some support for this, in that it was found
at Sheringham in 1909, concealed among
Heather on open heathland, and similarly it
was found on heathland at Beeston Regis in
1900. Alternatively, the Norfolk populations
may originate from wind-blown seed, either
from Scotland or the Continent. The timing
of records is interesting, as Creeping Ladystresses were recorded for the first time from
some of the plantations many years after they
were established; pines were first planted at
Wells and Holkham in the 1850s, largely with
seedlings produced in a nursery at Holkham
itself, but the ladys-tresses were not found
until 1952.
At some sites there have been documented
introductions by well-meaning naturalists,
as at Horsford in Norfolk in 1955-58
(although there were probably already natural
populations there) and Warwick Moor Wood
in Cumbria in 1931 (where the plantation
was clear-felled in 1987). Other Cumbrian
populations may well have been accidentally
introduced with conifers when plantations
were established.
Whatever their origin, the populations
in Norfolk are extremely interesting from a
scientific point of view, yet the controversy
over their status has compromised their
conservation. After showing keen and active
concern in the 1950s and 1960s, the attitude
of the Nature Conservancy and Nature
Conservancy Council (predecessors of English
Nature) to the largest and best-known
population at Holt changed, and in 1986 its
status as part of an SSSI was removed. As the
New Atlas maps all the British populations as
native, attitudes may now change.
29/1/09 12:25:55
GENUS SPIRANTHES
LADYS-TRESSES (II)
12/2/09 16:56:48
GENUS SPIRANThES
173
These small orchids have distinctive small, tubular white flowers and swollen, tuber-like
roots. Sadly, one member of the genus, Summer Ladys-tresses, is the only orchid that is
extinct in Britain and Ireland.
Distribution
The genus contains 45-300 species (the taxonomy is very uncertain), found in Europe, Asia,
Australia and South America, with the majority
in North and Central America. Four species
occur in Europe, with three in the British Isles.
Floral structures
The column is held horizontally within the
tubular flower with the circular stigma on its
underside, facing downwards. The rostellum
is well-developed and deeply forked with the
single torpedo-shaped viscidium held between
the two prongs of the fork. There are two
pollinia, each composed of two leaf-like plates,
and both are attached to the viscidium. Once
shed by the anther, the pollinia lie on top of
the column with the viscidium below their tips,
waiting to be removed by a visiting insect.
period as a single tuber and the rhizome

does not die off. New, swollen roots develop
each year and these push the roots from the
preceding season, which are shrunken and
depleted, out towards a horizontal position;
this may give the impression that there are
two types of roots: vertical, fleshy roots and
horizontal, fibrous roots.
The aerial stem dies off once seed has been
set, but a bud develops on the side of the base
of the stem and this will go on to form the new
leaves and aerial stem.
Additional lateral buds may develop in the late
summer or autumn at the base of the aerial
stem. These can produce their own roots and
leaves and may eventually become separate
daughter plants.
Growth pattern
Name
The aerial stem grows from a very short

rhizome which puts out two to six fleshy,
tuberous roots. Unlike the tubers of Orchis
and Dactylorhiza, the roots are heavily infected
with fungi. And, again unlike Orchis and
Dactylorhiza, the plant does not spend a resting
The generic name Spiranthes means twisted

flowers and derives from the Greek speira
meaning twisted or coiled and anthos meaning
flower. The English name probably refers to the
flower spikes of Autumn Ladys-tresses, which
look like plaited hair.
new
tuberous
root
January
rosette
dies off
March
July
remains
of
flower
spike
new
rosette
August
December
M Growth pattern of Autumn Ladystresses (after Wells 1981).

O 18 August, Norfolk. Autumn Ladystresses.
29/1/09 12:25:58
174
GENUS SPIRANThES
irish ladys-tresses
BaP
Spiranthes romanzoffiana
Other names: Hooded Ladies-tresses (North America)
This delicate, white-flowered, late summer orchid has one of the most unusual distributions
of any British plant, occurring in North America, Ireland, western Scotland and southwest
England but nowhere else in Europe. It is also one of the hardest orchids to find, partly due
to the remote location of the colonies and partly to the unpredictable nature of its flowering.
The biology of this species is a mystery. It is fairly widespread, with many recorded sites,
but very rarely sets seed in Britain and Ireland and has no obvious method of dispersal or of
maintaining its numbers.
Identification
Distinctive, with long, grass-like leaves and a
compact spike of small white flowers, usually
arranged into three spirally twisted rows. The
upper stem, ovaries, bracts and sepals have
numerous glandular hairs. These features,
combined with its usual damp grassy habitat,
restricted range and late flowering, are unique.
Similar species
Creeping Ladys-tresses is very occasionally
found on moorland and also has glandular
hairs on the flowers, but it is usually smaller,
with a different flower structure and very
different leaves.
Autumn Ladys-tresses also flowers in August
and September but is much commoner, has a
predominantly southerly distribution and is
almost always found on short, dry, calcareous turf.
Its flowers are arranged into an obvious single row.
Summer Ladys-tresses is now extinct in
Britain. Like Irish Ladys-tresses, it has long,
lanceolate leaves on the stem and is found
in wet, acid grassland but the flowers, as
well as being arranged into a single row, are
significantly smaller; 6-8mm long rather than
10-14mm, with the bracts 6-9mm long rather
than 10-20mm, sometimes even 30mm.
Other white-flowered orchids, such as Small
White Orchid and some Heath and Common
Spotted Orchids, may be found in the same
habitats and in the same geographical area
O 7 August, Colonsay, Inner Hebrides (Richard Gulliver).
29/1/09 12:25:59
IRISH LADYS-TRESSES
but flower earlier in the season. They all lack

glandular hairs on the upper stem and flowers
and can also be excluded by looking at the
details of the flower structure.
Flowering period
Habitat
Range
Frequently found in open grassy areas where

the soil is low in nutrients and permanently
damp or wet, either because it is flushed with
ground water or, more usually, because it is
close to a river, stream or lake and is flooded
from time to time. Often grows close to the
shore of a lough and has even been recorded
flowering with the leaves submerged. At most
sites the water and soil are mildly acidic to
neutral but Irish Ladys-tresses is sometimes
recorded from more alkaline, base-rich flushes.
Suitable conditions are provided by damp
meadows and rushy pastures, flushed grassy
slopes, wet heathland, and bogs, where it may
grow amongst Sphagnum on the disturbed
ground around old peat workings. In the
Hebrides it may particularly favour the band
of marginal land (known locally as blackland)
that is found at the transition between the limerich machair and the acid moorland inland.
And, although often growing in damp habitats,
it has also been found on heather moorland.
Old lazy beds is often given as a prime habitat
for Irish Ladys-tresses (lazy beds are used to
grow potatoes). Recent studies on Coll and
Colonsay have, however, comparatively rarely
found the species in this habitat.
Most of the sites in Britain and many in
Ireland are subject to extensive grazing, which
helps to keep the sward comparatively short
and reduce competition from other vegetation.
Disturbance by grazing animals may also
stimulate dormant plants, and it can favour
areas where cattle are fed in the winter or where
the ground has been broken up by ditching or
fencing. Generally found at low altitudes but
recorded up to 240m above sea level in Ireland.
In North America the Irish Ladys-tresses is
found in similarly damp habitats, such as bogs,
marshes, wet meadows and stream-sides, but in
California it is also found in coastal grasslands
and pine forests that are baked dry in summer.
175
Early July to early September, depending on

weather conditions and water levels, but mostly
in late July and early August.
In Ireland, it is very local in Eire but there are
some good populations in the lakes area of
western Co. Galway and Co. Mayo, especially
around Loughs Conn, Cullin, Corrib and Mask.
Otherwise, it occurs at a few sites in the far
southwest, in Co. Kerry and Co. Cork, and in
the northwest, in Co. Donegal. In Northern
Ireland the main concentration of populations
lies in Co. Antrim, around Lough Neagh and
the streams and rivers that flow into it. It is
also present along the River Bann through
Lough Beg and as far north as Coleraine in
Co. Derry and around the southern shores of
Lough Neagh in Co. Armagh. There are also
some scattered sites elsewhere in Co. Antrim,
including Gortnagory on the Garron Plateau,
and it has recently been found at Aird on the
north coast near the Giants Causeway. There
are also a very few sites in Co. Fermanagh, Co.
Down and probably also still Co. Tyrone.
In Scotland the Irish Ladys-tresses is
widespread but scattered on Coll and Colonsay
in the Inner Hebrides and has been recorded
from around 30 sites on each island. Other
concentrations are found in the southern
1987-99
1970-86
pre 1970
29/1/09 12:25:59
176
GENUS SPIRANThES
California, New Mexico and Arizona in the

west and Illinois and Pennsylvania in the east.
Irish Ladys-tresses is found in North
America and on the western fringe of Europe
in the British Isles. This is an extreme example
of a pattern of distribution known as amphiAtlantic and is shared by just a few other
plants, including Blue-eyed Grass, Pipewort
and American Pondweed. A variety of
theories have been advanced to account for
the distribution of Irish Ladys-tresses but the
simplest and most likely is that it arrived in
Britain and Ireland either via the long distance
dispersal of wind-blown seed or by pieces
of root and rhizome being carried across the
Atlantic by birds.
How to find it
The grass-like leaves are very hard to see,
making this species effectively invisible when
not in flower. The number of plants flowering
varies from year to year and can be very low
following a dry spring, while many that do
flower are grazed off. A visit to the Hebrides,
Lough Corrib (Co. Galway) or Lough
Mask and Lough Con (Co. Mayo) would,
nevertheless, be worthwhile.
DESCRIPTION
M 31 July, Co. Londonderry (Sean Cole) The flowers are

arranged more or less obviously into three rows.
Outer Hebrides on Vatersay, Barra, South

Uist and Benbecula. Otherwise it is rare but
has been recorded from Islay, Mull and Tiree.
On the mainland the species is found in three
small areas of Inverness-shire close to its
Hebridean strongholds: Moidart, Morvern
and Ardnamurchan (where small numbers
have been found around the western end of
Loch Shiel). There are also old records from
Kintyre. In England it is known only from
one site in Devon, to the east of Tavistock
on the southwest fringe of Dartmoor. World
range: Widespread in North America, from
the Atlantic to the Aleutian Islands, south to
Height: 10-35cm.
Stem: Yellowish-green with scattered glandular
hairs towards the tip.
Leaves: There are three to five (sometimes up
to eight) yellowish-green leaves. The lower are
long, narrow and parallel-sided with a hooded
tip and are held very erect. The upper leaves
are short, pointed and loosely sheathe the
stem. The margins of the leaves may be rolled
inwards, making them even narrower and more
grass-like, especially in Northern Ireland. The
shoots appear in the autumn, around October,
but the leaves may not expand until the spring;
in North America this species is wintergreen.
Spike: Five to 40 flowers are arranged in three
rows up the stem (rarely only two rows) and
each row is variably twisted; this arrangement is
most obvious on plants with numerous flowers.
Bract: Narrow and pointed, sheathing the
29/1/09 12:26:01
IRISH LADYS-TRESSES
177
ovary, with glandular hairs on its outer surface.

The lower bracts are as long as the flowers, the
upper bracts shorter.
Ovary: Cylindrical, three-ribbed, on a very
short stalk and with a few glandular hairs. It is
held vertically but bends at the tip so that the
flowers lie horizontally.
Flower: White and tubular. The sepals and
petals are creamy-white, washed green towards
the base. The sepals are narrowly triangular
and blunt-tipped, with glandular hairs on their
outer surface and three greenish veins; the
petals are narrower and more strap-shaped. The
sepals and petals form a hood that encloses the
column and the basal half of the lip, with the
tips of the sepals distinctly turned outwards.
The lip is creamy-white with fine green veining
and is fiddle-shaped, with the waist nearer the
tip. The sides of the larger and broader basal
portion are turned upwards to form a trough or
gutter, with two small nectar-producing bosses
at the extreme base. The smaller distal portion
is tongue-shaped or square-ended, frilled
and toothed along the edges and sharply bent
downwards. The flowers have, at most, a faint
scent (earlier reports of a strong vanilla-like
scent are presumably errors).
Subspecies
None.

Plants in Northern Ireland are said to be
on average taller and leggier than those in
southern Ireland. They have creamy rather
than white flowers, a looser spike, narrower
lip and leaves that are often in-rolled at the
edges, giving them an especially grass-like
appearance. Plants from Scotland and Devon
are intermediate. At one time Northern Irish
plants were treated as the same species as in
North America, although separated as a distinct
variety (S. romanzoffiana var. stricta), with
southern Irish plants separated as a distinct,
endemic, species (named S. gemmipara). Two
distinct subspecies are still recognised by some
authors, but examination of the variation across
the entire range in North America has shown
M 16 August, Colonsay, Inner Hebrides (Richard Gulliver).

The lip is frilled and spoutlike as in all the Spiranthes; this
plant has rather widelyspaced flowers.
that in large populations plants resembling

both of these extremes as well as a range of
intermediates are present, and no subspecies
or varieties are usefully recognised in the
British Isles.
Genetic studies have shown that there is a
split between a northern group of populations,
from Coll, and from Barra and Vatersay in the
Outer Hebrides and a southern group from
Colonsay and Ireland (no plants from mainland
Scotland were examined). There has been little
recent contact or gene flow between these two
groups, but this division is not reflected in any
known differences in appearance or ecology.
29/1/09 12:26:02
178
GENUS SPIRANThES
O 4 August, Benbecula,
Outer Hebrides (Richard
Gulliver). In a small
minority of cases two plants
grow together, presumably
the product of vegetative
reproduction. These are
very fewflowered spikes.
BIOLOGY
The reproductive biology of Irish Ladys-tresses
in the British Isles is a mystery. In North
America, medium-sized, long-tongued bees
act as pollinators but until very recently bees
had only very rarely been recorded visiting the
flowers in the British Isles. Even more curiously,
mature seed capsules had never been found in
Scotland and only once or twice in Ireland and it
was assumed that little or no seed was produced
(studies in North America have shown that
the column structure of Irish Ladys-tresses
prevents automatic self-pollination). But, given

the scattered distribution of Irish Ladys-tresses
in the British Isles, wind-blown seed is the
only plausible mechanism for dispersal and
subsequent propagation.
Concerted research efforts have now
produced more records of bumblebees visiting
the flowers in Scotland, and at the largest
colony in Ireland the orchids are frequently
visited by medium-tongued and long-tongued
bumblebees (Bombus pascuorum and B.
hortorum respectively), and occasionally by
short-tongued honeybees. Only bumblebees
29/1/09 12:26:03
IRISH LADYS-TRESSES
were observed to pick up pollinia, however, and

presumably these act as the primary pollinators.
Careful examination of plants from Colonsay
in the Inner Hebrides revealed that c. 40 %
of the flowers had their pollinia removed and
c. 70 % had pollen on their stigmas, while at
the Irish colony pollinia were found on the
probosci/thorax of visiting bumblebees and
on the stigmas of randomly checked flowers.
Not surprisingly, in view of these observations,
it has been shown that a small but consistent
proportion of flowers on Colonsay do set seed
(although much lower than the proportion of
flowers that are pollinated, and the number
of seeds per capsule is very low). What is
surprising is that examination of more than
1000 flowers at the Irish site found no ripe
capsules. The reason for the very low levels of
seed production is not clear, but the populations
on Colonsay and in Ireland (the southern
group) have been shown to have a low level of
genetic diversity, perhaps indicating a genetic
bottleneck during which numbers fell to a very
low level. High levels of inbreeding may cause
problems such as impaired stigma receptivity
and low pollen viability, but examination
of flowers on Colonsay showed high levels
of pollen germination on the stigma, with
pollen tubes growing down the style and some
penetrating an ovule as normal. The northern
group of populations show a high level of
genetic diversity and it would seem logical that
they produce seeds much more consistently.
Searches in 2007 failed to confirm this, but
it was poor year for the species; when flowers
are few and scattered, or the weather poor, the
chances of insect pollination are much reduced.
British and Irish populations can reproduce
vegetatively through the development of an
additional bud at the base of the stem (rarely
two or very occasionally three). Two buds can
produce two aerial stems in the next growing
season, and these may eventually separate to
form two plants. Extra buds are, however, only
produced by a small percentage of plants each
year (less than 5% in the Hebrides), and it seems
that many of these extra buds disappear for one
179
reason or another and relatively few develop.

Nevertheless, the incidence of twinned orchids
in a population, presumably the product of
vegetative reproduction, varies from very low
to over 25% of plants. Genetic studies do not,
however, indicate that groups of clones are at
all common. It is possible that new plants may
develop from fragments of root, perhaps broken
off by cattle or sheep, but this yet to be recorded.
Irish Ladys-tresses is an enigma. Seed
is apparently rarely produced. Vegetative
reproduction is uncommon and does not account
for long-distance dispersal. Clearly, much more
remains to be learned about this beautiful orchid,
especially if it is to be effectively conserved.

Poorly understood. The aerial stem grows from
a cluster of two to six thick, fleshy, tuberous
roots, more-or-less vertical in the soil and
connected at the top by a very short rhizome.
A lateral bud develops at the base of the aerial
stem (or on the rhizome if the plant is dormant
underground) between July and October and
overwinters, going on to form the leaves in the
spring of the following year and, if the plant is
to flower, the stem appears in early June. The
species may become dormant underground,
with up to six years absence recorded.
Little is known about the development from
seed to flowering plant, but the species possibly
spends five years growing underground before
the first leaves appear.
Hybrids
None recorded in the British Isles.

The specific name romanzoffiana honours
Nicholas Romanzof, a Russian minister of state.
The species was discovered in Alaska around
1828 when it was still a Russian territory.
HISTORY aND
CONSERVaTION
8 of the 1985 Wildlife Order (NI) and in Eire
29/1/09 12:26:03
180
GENUS SPIRANThES
under the Flora (Protection) Order. It is the

subject of a UK Biodiversity Action Plan and
a priority for conservation because Britain and
Ireland hold the only European populations.
The first Irish record was published in
1828 by Sir J.E. Smith in the English Flora:
Near Castletown opposite to Bearhaven on
the northern side of Bantry Bay, County of
Cork, Mr Drummondcommunicated to me
in August, 1810. Irish Ladys-tresses was first
found in Northern Ireland in 1892, at Brackagh
Bog near Lough Neagh, Co. Armagh, on an old
dug-out peat bog.
Past and present occurrence of Irish Ladystresses in Brit
Britain
Ireland
21
44
17 (0.6%*)
17 (1.7%*)
% lost, 1500-1969
5%
36%
% lost, 1970-1986
14%
25%
% lost, total
19%
61%

1500-1999
current range
The first positive identification for Scotland

came from Colonsay in 1930 (a ladys-tresses
had been found on Coll in August 1921 but
was not certainly identified until 1939). The
first record for the Scottish mainland was as
recent as 1954.
Irish Ladys-tresses was found on the
southwestern edge of Dartmoor, Devon, in July
1957, when there were seven plants on a heavily
grazed lawn of Purple Moor-grass and on the
adjacent bog. It has but not been seen since c.
1993, although the site is essentially unchanged.
Most populations of Irish Ladys-tresses
are small and scattered but some of the largest
produce from 100 to 200 spikes annually,
with 400 spikes recorded recently from one
area in Co. Mayo. The largest site in Scotland
was recently estimated to hold 1,100 plants
(flowering and vegetative combined).
Irish Ladys-tresses is very hard to survey
and the population trends are not clear. On

the one hand, it is known from an increasing
number of sites, both in Ireland and Scotland,
largely due to particular efforts being made to
find it; it was found for the first time on Mull
in 1990 and on Tiree in 2002. On the other
hand, it tends to vanish unpredictably and often
rapidly from known localities. It is, however,
effectively impossible to find unless in flower
and the most likely explanation for its erratic
appearances is that populations are relatively
stable but grazing and, perhaps sometimes,
the weather prevent many or most plants from
flowering. Sheep, cattle or rabbits often graze-off
the flower spikes and whole colonies can appear
to vanish overnight if sheep are turned out in
the vicinity. As with many orchids, slugs can also
be a particular problem, grazing-off both the
flower spikes and the leaves. Recent experiments,
where sheep have been excluded from large
populations, have produced a profusion of
flowers, and careful long-term monitoring on
Barra has indicated that the population there is
comparatively stable and long-lived.
The impact of grazing on Irish Ladystresses is of conservation concern, but the
situation is not clear-cut. Colonies, sometimes
large, have been recorded on sites with a
variety of grazing regimes, and indeed, it may
be tolerant of heavy grazing. It is becoming
clear that many orchids do best when grazing
reduces competition and breaks up the sward,
providing suitable sites for the establishment
of seedlings, even if this means that many or
most of the flowers are grazed-off before setting
seed. This may apply to Irish Ladys-tresses.
However, some well-established Irish sites are
only lightly grazed or even ungrazed.
The species is certainly vulnerable to
changes in management, such as drainage, and
relatively few sites are protected as SSSIs or
reserves.
29/1/09 12:26:03
AUTUMN LADYS-TRESSES
autumn ladys-tresses
181
Near Threatened
Spiranthes spiralis
This delicate little orchid blooms in August and September and is the last species to flower
in the orchid season. It is confined to short turf, often near the sea, and is one of the orchids
that has taken happily to lawns. If conditions are right, hundreds or even thousands of
flower spikes may appear, but it can exist undetected for many years if the grass is cut too
often.
Identification
Distinctive. The delicate, slender spike is
decorated with a row of small, tubular, white
flowers that are usually arranged in a spiral
pattern. A few tiny, bract-like leaves clasp the
stem and there is a rosette of short, oval leaves
lying flattened to the ground a little to one
side.
Similar species
Irish Ladys-tresses is confined to Ireland
and northwest Scotland with an outpost on
Dartmoor. When in flower it has prominent,
long, narrow leaves on the stem. Habitat is
also a good distinction, as it is found in wet
grassland and bogs.
Summer Ladys-tresses is extinct in Britain
and, like Irish Ladys-tresses, has long, narrow
leaves along the stem when in flower.
Creeping Ladys-tresses is found in
coniferous woodland (rarely moorland or
dunes), usually on acid soils. It is found in
northern England and Scotland with a few
populations in Norfolk. Soil conditions can
change over a very short distance, however,
and prior to 1979 Autumn and Creeping
Ladys-tresses could be found growing within
a few metres of each other at Holt in Norfolk.
Creeping Ladys-tresses is easy to distinguish
as it grows from horizontal rhizomes that
form irregular patches of leaves with scattered
flower spikes. Its little bell-like flowers are
exceptionally hairy and its leaves are faintly
net-veined.
P 18 August, Norfolk. The flowers can be arranged into
either clockwise or anticlockwise spirals.
29/1/09 12:26:04
182
GENUS SPIRANThES
29/1/09 12:26:07
183
animals may knock-over or bite off the flower

spikes but ignore the leaf rosettes, and mowing
also leaves the rosettes unharmed. Conversely,
the lessening or abandonment of grazing or
improvement with fertilisers will promote
more vigorous vegetation that will swamp the
orchid. In the past, Autumn Ladys-tresses
was also found in damp meadows in the drier
climate of eastern England, but these are almost
all now drained and improved and the orchids
have gone.
Flowering period
Early August to the end of September or even
early October.
Range
M 18 August, Norfolk.The new rosette of leaves appears

alongside the current flower spike but is hidden in the
grass.

O 30 August, Norfolk.
Habitat
Found on short, dry, nutrient-poor turf in
sunny situations, often near the sea. It usually
grows on calcareous soils on chalk, limestone,
dunes, shingle banks or in the grykes of
limestone pavements. Ancient earthworks are
favoured, as are lawns; some old lawns were
made with turf from nearby pastures and
the orchids may have come with the turves,
but they can certainly also colonise lawns
via wind-blown seed. It is also sometimes
found on old tennis courts, road verges and
reservoir embankments and has occasionally
been recorded from grassy places on lessacid heaths. The critical factor is the lack of
competition from taller and more vigorous
herbs and grasses, and the correct conditions
may be provided by grazing or mowing; grazing
Southern England from the Isles of Scilly

to Kent, now mostly south of a line from
Bristol to London and rare to the north of
this, although there are some scattered sites as
far north as Norfolk, southeast Lincolnshire,
Yorkshire and in Cumbria around the northern
shores of Morecambe Bay. Also found in
Wales, especially the southeast and coastal
regions, the Isle of Man and Ireland, where
it is scattered in the west and on the south
and east coasts, and absent from the northern
third of the island. World range: Almost
confined to Europe, with outposts in North
Africa and around the Caucasus. Found north
1987-99
1970-86
pre 1970
29/1/09 12:26:08
184
GENUS SPIRANThES
to Denmark and southernmost Sweden, east

to the Baltic States and western Ukraine
and south to the Mediterranean, including
most of the Mediterranean islands from the
Balearics to Cyprus. Also Algeria and Tunisia,
northern Turkey, Israel, Syria, the Caucasus and
northwestern Iran.
How to find it
The slender spikes of small greenish-white
flowers do not grab the attention and can easily
be missed from a walking height. They become
much more obvious from a low vantage point,
and squatting down to scan along the ground
is often the easiest way to see the plants. Nonflowering ladys-tresses are almost impossible
to find, however, as the rosettes are only 2.57cm in diameter, lie hidden in the grass and
are easily overlooked as plantains. But, from
nothing, flower spikes may appear and start to
bloom in just a week.
The number of flowering spikes may
fluctuate very widely from year to year, but
the total number of plants present, including
non-flowering rosettes or those dormant
underground, is much more stable. The
proportion of plants producing flower spikes
is probably related to the weather, but the
optimum set of conditions is not known.
Some populations may not be able to flower
for several years but will then bloom en masse
when grazing or mowing ceases. In a dry
summer lawns may be left uncut, allowing
the orchids to flower, and until this happens
Autumn Ladys-tresses may grow unseen for
many years; suitable lawns may be picked out
later in the autumn as they often support a
good variety of fungi, including the colourful
waxcaps.
DESCRIPTION
M 22 August, Norfolk. The tiny flowers are arranged in a

spiral around the stem, resembling a ladys braided hair.
Height: 3-15cm, sometimes to 20cm high, but

in the wet summer of 2004 a population in
Norfolk averaged 15cm high, with the tallest
28.5cm.
Stem: Pale green, densely covered towards
the spike with fine, white glandular hairs.
The flower spike grows from the centre of the
29/1/09 12:26:09
previous seasons rosette, with or without the

remains of the dead leaves at its base. The next
seasons rosette grows beside it.
Leaves: The flowering spike has three to seven
small, narrow, lanceolate, bract-like leaves that
tightly sheathe the stem. They are greenish with
a narrow, whitish or translucent fringe. Just to
one side of the base of the spike there is a tight
rosette of up to ten leaves. These are around
3cm long, dark, shiny green with a faint blue
tone, oval or elliptical in shape, taper to a point,
have a thick keel and broadly sheathe the stem
at their base. This rosette of leaves emerges in
August or September and overwinters, dying
off in late May or early June.
Spike: 3-21 flowers (the average is 9-11) are
arranged in a row up the stem. In most plants
this is twisted so that the flowers form a spiral
pattern, but in some the twist is so slight that
the flowers simply form a line along one side of
the spike. The spiral can be either clockwise or
anti-clockwise and is mostly twisted through
less than 360, although in some plants it may
be through three full turns.
Bract: Pale green with scattered glandular hairs
towards the base and a narrow transparent-
185
whitish fringe. The bracts are lanceolate, taper

abruptly to a fine point and are a little less than
twice the length of the ovary, which they clasp.
Ovary: Green, three-ribbed, stalkless, with fine
glandular hairs. The ovaries are held upright
but bend at the tip so that the flowers are held
more-or-less horizontally.
Flower: A small, white, trumpet-shaped tube.
The sepals and petals are white, often washed
green towards the base. The sepals are oblong
but taper slightly to a blunt tip and have
glandular hairs on the outer surface. The petals
are slightly shorter, rather narrower and more
strap-shaped. The upper sepal and the petals,
together with the lip, form a long, narrow tube
with the tip of the upper sepal curved upwards.
The lateral sepals are held slightly drooped and
spread horizontally away from the tube. The
lip is pale green, becoming whiter towards the
edges, with two small, globular nectaries at
the base. It can be oval or even heart-shaped
with a slightly squared-off tip, and the sides
curve upwards to form a trough or gutter; for
its entire length the lip also bends downwards
to resemble the lip of a china teapots spout,
and the extreme tip is rolled downwards
and crimped. The greenish column projects
horizontally into the tube made by the petals
and lip. The flowers are honey-scented.
Subspecies
None.

None.
BIOLOGY
M 22 August, Norfolk. The lip has a spoutlike tip with

crimped edges, and there are abundant glandular hairs
on the stem, ovaries and sepals.
The flowers are pollinated by bumblebees.

When the flower opens the longboat-shaped
rostellum lies close to the lip at the bottom of
the tubular flower with the sticky viscidium
facing downwards. In this position the
rostellum blocks access to the stigma so that the
flower cannot be self-pollinated. A bumblebee
lands on the lip and inserts its proboscis in
search of the nectar that is produced at its base.
There is just enough room for the proboscis to
29/1/09 12:26:11
186
GENUS SPIRANThES
all flowers. The production of abundant seed

is probably a factor is the ladys-tresses success
in colonising new sites (although the capsules
contain an average of only 850 seeds each, a
relatively low figure).
The species also reproduces vegetatively
from lateral buds at the base of the stem. This
leads to the formation of clusters of two or
three plants (clusters of up to 12 have been
recorded). Vegetative reproduction is, however,
thought to play only a minor role in the
turnover of a population of Autumn
Ladys-tresses.
M 22 August, Norfolk. Despite their small size, the flow

ers are pollinated by bumblebees.
reach the nectaries but in the process it pushes

past the rostellum, and the pollinia are attached
to the proboscis by the fast-drying glue of the
viscidium. The bee then moves on, carrying the
pollinia with it. Over the following 24 hours
or so the column and lip move apart, creating
enough space for a visiting bumblebee to insert
its proboscis, with the pollinia attached, into
the flower, where they rub against the stigma
(which is now exposed and has become much
stickier). The pollinia are brittle and break
off in small pieces. A single bee can pollinate
several flowers. Due to the 24-hour interval,
older flowers are always pollinated with pollinia
from a younger flower. The lowest flowers in
a spike open first and, as bumblebees work
upwards from the bottom of a spike (i.e. from
older to younger flowers), they visit the older
flowers first and cannot pollinate them with
pollinia taken from the same spike. In this way
cross-pollination is virtually guaranteed. The
mechanism is efficient and seed is set by almost
The aerial stem arises from a very short

rhizome that is almost concealed by two or
three (rarely up to five) thick, fleshy, tuberous
roots that are very much like miniature parsnips
in shape and have a few short, transparent hairs.
The roots that supported the current years
leaves and flower spike eventually shrivel when
their store of food is exhausted and are replaced
by new ones. The protocorm that is produced
by the germinating seed is heavily infected with
fungus, but by the time the roots appear the
rhizome is free of infection. However, the roots
are each infected in turn as they develop.
Plants may spend one or possibly more
years underground with no aerial leaves and
still flower the following year. This indicates
that the fungal partner plays a significant
part in the nutrition of the mature plant. In a
study in Bedfordshire the half-life averaged
6.9 years and varied from 4.6-9.2 years (the
half-life is a measure of the life expectancy
of the orchid after its first appearance above
ground. It marks the point at which 50% of
the population that emerged in any given year
have died).
Early researchers concluded that Autumn
Ladys-tresses took many years to go from seed
to flower. However, in the laboratory green
leaves have been produced six months after
germination, and flowering plants in five years.
Hybrids
None.
29/1/09 12:26:14
187

The specific name spiralis means twisted or
spiral (as does the generic name).
HISTORY aND
CONSERVaTION
In 1548 William Turner wrote in his Names
of Herbes: Satyrion is very commune in
Germany, and a certeyne ryghte kynde of the
same groweth besyde Syon [Sion, Middlesex],
it bryngeth furth whyte floures in the end of
harueste and it is called Lady traces. With this
statement the Autumn Ladys-tresses became
the first species of wild orchid to be recorded
in Britain, an honour it shares with Common
Twayblade.
Past and present occurrence of Autumn Ladystresses
Atlas).

1500-1999
current range
% lost, 1500-1969
Britain
Ireland
668
117
302 (11%*)
34 (3.4%*)
50%
62%
% lost, 1970-1986
5%
9%
% lost, total
55%
71%
In the 450 years since William Turners

account, the species has not fared too well in
the British Isles and has disappeared from
55% of its historical range in Britain and 71%
in Ireland; it is classified as Near Threatened.
On the edge of its range in Britain and Ireland,
Autumn Ladys-tresses has retreated to its
core habitats, and the losses are concentrated
in the northern and eastern parts of its former
range and at inland sites in general. Sadly,
it has almost gone from northern England,
the Midlands, East Anglia and Kent. It is
extinct in Northamptonshire, Warwickshire,
Staffordshire, Nottinghamshire, Derbyshire,
Cheshire and, needlees to say, Middlesex.
With its requirement for short, nutrient-
M September, Norfolk. Autumn Ladystresses can occur

in large numbers and, although the number flowering
each year is very variable, the total population is relatively
stable.
poor grassland the Autumn Ladys-tresses

cannot tolerate any sort of improvement by the
addition of fertlisers. Ploughing of downland
and pastures and their conversion to arable or
reseeding with vigorous grasses are even more
directly destructive processes. Conversely, the
abandonment of grazing leads to the invasion
of grassland by scrub. These practices have been
taking their toll since the 19th century, but the
losses are mitigated to a small extent by gains
as the species is able to colonise new sites and
has appeared in large numbers on the lawns of a
few favoured housing estates.
29/1/09 12:26:15
188
GENUS SPIRANThES
summer ladys-tresses
Extinct
Spiranthes aestivalis
The delicate, pure white flower spikes of Summer Ladys-tresses once graced a few favoured
bogs on the Channel Islands and in the New Forest. Sadly, it is the only orchid to have
become extinct in the British Isles and was last recorded in about 1952. Changes to the
habitat were important factors in its decline, but it may have been given the final death knell
by collectors. It has also declined sharply throughout northwest Europe and the chances of
it reappearing naturally in England appear to be slim.
Identification
The small, white, trumpet-shaped flowers,
arranged on the stem in a spiral pattern,
identify this species as one of the ladys-tresses.
When in flower it has several long, narrow
leaves at the base of the stem.
Similar species
Autumn Ladys-tresses is relatively common on
short, dry grassland but has also been found,
albeit rarely, in damp meadows and in grassy
places on less acid heaths. When in flower it has
a few small, bract-like leaves on the stem but
the basal rosette has already died off; the new
rosette appears a little to the side and the leaves
are always much shorter and blunter than in
Summer Ladys-tresses.
Irish Ladys-tresses is largely confined to
Ireland and western Scotland but has been
found on Dartmoor and could conceivably
turn up elsewhere in southern England. It too
favours wet, boggy habitats but it has larger
flowers than Summer Ladys-tresses, usually
arranged into three spiral rows. Each flower has
a longer bract, 10-20mm long, rather than just
6-9mm.
Habitat
In the New Forest the species was confined
to wet, peaty, valley bogs with bog mosses
(Sphagnum spp.). It favoured areas that were
slightly less acidic and where the vegetation
P The flowers are very similar in shape to Autumn Ladys
tresses, but this species grows in acidic bogs, in this case
with Crossleaved Heath (Bob Gibbons/Natural Image).
29/1/09 12:26:16
SUMMER LADYS-TRESSES
was relatively low and open. On the Channel

Islands it was found in a Sphagnum bog on
Guernsey and on wet sandy ground on the
margin of St. Ouens Pond on Jersey. In Europe
it is also found on moist heathland, damp dune
slacks and other damp, base-rich areas with
short, open vegetation.
Flowering period
Mid-July to mid-August.
Range
Extinct in the wild in Britain. Formerly found
in the New Forest in Hampshire and on
Guernsey and Jersey in the Channel Islands.
World range: Central and southern Europe,
north to Germany and northwest France,
east to the Czech Republic and the former
Yugoslavia, and south to the Mediterranean,
including the Balearic Islands, Corsica and
Sardinia. Also found in North Africa in
Morocco and Algeria, and possibly also in
Turkey. It is rare and declining in much of
Europe due to habitat destruction, especially
towards the northern edge of the range, and is
extinct in Holland and Belgium.
How to find it
On the surface there seems little chance of
finding this species. From time to time there are
rumours that it has appeared again in the New
Forest, but these have never come to fruition.
DESCRIPTION
Height: 10-20cm, sometimes to 40cm.
Stem: Yellowish-green, with fine glandular
hairs towards the tip.
Leaves: Yellowish-green, glossy, narrow and
strap-shaped; three to six leaves are held erect at
the base of the stem and there are one to three
smaller, bract-like sheathing leaves above these.
The leaves emerge in spring and are retained all
summer.
Spike: Five to 20 flowers are arranged into a
single row which is twisted spirally around the
stem, sometimes several times.
Bract: Lanceolate, finely hairy at the base. The
bracts are rather longer than the ovary, which
they tightly clasp.
189
Ovary: Yellowish-green, finely hairy, six-ribbed

and slightly twisted. The ovaries are stalkless
and held upright but bend at the tip so that the
flowers lie horizontally.
Flower: White and trumpet-shaped. The sepals
and petals are glandular-hairy on their outer
surfaces and strap-shaped, with the petals a
little shorter and narrower than the sepals.
They form a tight tube around the column with
their tips splayed outwards at the mouth of the
tube. The lip is tongue-shaped with two small,
nectar-secreting glands at the base. Towards the
rear, the sides of the lip curve upwards to form a
gutter shape and at the tip the lip turns sharply
downwards and has a conspicuously frilled
or crimped margin. The flowers are slightly
scented.
Subspecies
None.

None.
BIOLOGY
The structure of the flower is very similar to
Autumn Ladys-tresses, and the flowers are
presumably pollinated in the same manner
but there is no specific information on this
species. The flowers are said to be fragrant in
the evening, which may indicate that they are
pollinated by night-flying moths. The species
also reproduces vegetatively from additional
lateral buds at the base of the stem.

The aerial stem grows from a cluster of two to
six thick, fleshy roots. Little is known about the
period between germination and first flowering.
Development may be relatively rapid, however,
because, following germination, protocorms
appear by the spring, the first root forms in
July and the first tiny leafy shoot appears above
ground in the late summer.
Hybrids
None.
29/1/09 12:26:17
190
GENUS SPIRANThES
29/1/09 12:26:20
SUMMER LADYS-TRESSES

The specific name aestivalis means of the
summer.
HISTORY aND
CONSERVaTION
The first British record dates from 1840.
At a meeting of the Linnean Society on 17
November of that year, Mr. Janson exhibited a
specimen discovered in August last by himself
and Mr. Branch near Lyndhurst, Hampshire
(Proceedings of the Linnean Society).
In England this species was always confined
to the region southwest of Lyndhurst in the
New Forest. In this area it occurred regularly,
sometimes in large numbers, in at least five sites
until the end of the 19th century. As with many
species of orchid, the number of flowering
plants varied tremendously from year to year.
Thus, in 1901, E.D. Marquand noted that he
once saw half an acre of bog perfectly white
with these flowers, but the following year only
a few spikes of bloom appeared. At another site
200 flowering spikes were noted around 1900.
There was, however, a marked decline in the
20th century, and the species had vanished from
most sites by 1940. It persisted in the original
1840 location, a bog north of the A35, just
north of New Forest Gate and east of Highland
Water, until 1952 (with a possible sighting
nearby in 1959).
The decline was partly due to habitat
destruction, especially drainage and
afforestation (thus a site west of Brick Kiln
Inclosure was unwittingly destroyed when
drained by the Forestry Commission). Some
of the original sites are still very wet, however,
and superficially unchanged, although shading
by trees and scrub and subtle changes to
the vegetation through a process of natural
succession may have made them unsuitable.
Summer Ladys-tresses may also be one of the
few cases where collecting was a genuine cause
191
of decline and extinction; rare and on the edge

of its range, it was perhaps especially vulnerable.
In the 19th and 20th centuries all-toonumerous collections were made of flowering
plants, often complete with roots, both for
private and public herbaria.
Summer Ladys-tresses was also found on
the Channel Islands. The first record was on
24 July 1837, from the banks of St. Ouens
Pond on Jersey. This was its only locality on the
island and it was always scarce at this site. Due
to overcollecting it quickly became even scarcer
and was last recorded in 1926, when just a
single plant was seen (and even this last plant
was possibly later collected). On Guernsey it
was found on boggy ground around the lake
of Grande Mare in around 1841 and was
fairly common to start with, but collecting and
drainage led to its demise by 1914.
There are occasionally rumours that
Summer Ladys-tresses has been refound in
the New Forest or reports that it has been
discretely reintroduced, without official
sanction, to one or more of its former sites.
Nothing has ever come of these stories, but the
possibility of an undocumented reintroduction
will cast a permanent shadow over any natural
reappearance.
O Extinct in Britain, Summer Ladystresses is also on

the decline throughout northwest Europe (Bob Gibbons/
Natural Image).
29/1/09 12:26:20
Distribution
Primarily Asian, with 30 species in total, but just
one ranging west to Britain and Europe.
Name
The origin of the generic name Herminium is
uncertain. It may derive from the Greek and
means buttress or foot of the bed. This is
supposedly an allusion to the pillar-like tubers
but these are spherical in Musk Orchid, the first
species to be given the name. Alternatively, it may
derive from Hermes, the messenger of the gods.
Genus HERMINIUM
musk orchid
29/1/09 12:26:24
MUSK ORCHID
musk orchid
193
Vulnerable, BaP
Herminium monorchis
Despite its name, this small orchid does not smell of musk. It is confined to a relatively
few sites in southern England, where it is found in very short turf on chalk or limestone,
occasionally in large numbers. Surprisingly, and despite its status as a Nationally Scarce
orchid that has been lost from around 70% of its former range, it seems to have received
remarkably little attention from scientists and conservationists.
Identification
This diminutive orchid is easily overlooked but
when found is not hard to identify. The flower
spike is crowded with tiny greenish-yellow
flowers. Unless examined closely, the lip is
hardly different in appearance to the petals and
sepals, and the flower therefore appears to be
made up of six almost identical narrow petals
that form a little bell.
Similar species
Bog Orchid is also very small and greenishyellow in colour but is strictly confined to acid,
boggy ground, and the structure of its flower is
completely different.
Habitat
Musk Orchid is found exclusively on short,
well-drained grassland on chalk or limestone
soils. Its small stature means that it cannot
compete if the vegetation is tall, so thin or
compacted soils that restrict plant growth
are favoured. It particularly likes the narrow
terracettes formed on steep downland slopes
by soil creep, as well as ancient earthworks,
abandoned quarries, chalk and lime pits, and
spoil heaps. It has been recorded up to 215m
above sea level.
Flowering period
Early June to early July, sometimes to early
August. There can be large variations from
O 29 June, Hampshire. The colour of the flower varies a
little, from slightly yellower to slightly greener.The smallest
plants may be just 2cm high.
P 29 June, Hampshire.The small, slender spikes of Musk
Orchid are often hard to see from walking height.
29/1/09 12:26:26
194
GENUS hERMINIUM
29/1/09 12:26:29
MUSK ORCHID
year to year in the number of flowering

spikes, with the temperature and amount of
rainfall over the previous summer being the
determining factors. A hot dry summer can
cause a big drop in the number of flowers the
following year.
1987-99
1970-86
pre 1970
195
How to find it
This small, greenish-yellow orchid is hard
to spot. This is especially so on anything but
the shortest turf or if the spikes are few and
scattered. In those situations, scanning carefully
from a low-level vantage point, sitting or
squatting, can help. Once one spike has been
seen, however, others will usually be found
nearby. It can be easier to tread on than to
see, so it is good practice to keep to whatever
paths are available. On the other hand, in
a few favoured sites Musk Orchid occurs
in large numbers and can form dense and
obvious stands. One of the best sites for this
species is Noar Hill (Hampshire), and others
include St Catherines Hill (Hampshire) and
Malling Down (Sussex), with small numbers
at Tottenhoe Knolls (Bedfordshire), Park Gate
Down (Kent), Box Hill (Surrey) and Ham Hill
(Wiltshire).
Range
Confined to the chalk of the North and
South Downs in Kent, Sussex and Surrey,
also Hampshire, Dorset (just two sites),
Wiltshire and Berkshire, the Cotswolds
in Gloucestershire and the Chilterns in
Buckinghamshire, Hertfordshire and
Bedfordshire. It is very local and is absent
from large areas of apparently suitable habitat.
World range: Widespread in Europe and
Asia. In Europe it is found north to c. 61N
in Scandinavia, southern Finland and the
Baltic States, and it ranges south to southeast
France, Italy (the Apennines) and the Balkans,
with isolated populations in northwest Turkey
and the Caucasus. The species reaches its
western limit in northern and eastern France,
but there is an isolated population in central
Spain. It ranges in a narrow band across Siberia
to the Russian Far East, North Korea and
northeast China, and is also found in another
narrow band across northern China, with
scattered records from northern Japan and the
Himalayas.
M 29 June, Hampshire. In the tiny, belllike flowers the

lip appears similar to the petals and sepals, with only its
narrow central lobe visible.
O 28 June, Hampshire.
29/1/09 12:26:31
196
GENUS hERMINIUM
M 29 June, Hampshire.The flowers are scented but smell

of honey, not musk.
DESCRIPTION
Height: 2-30cm but usually 3.5cm-15cm and
rarely more than 20cm.
Stem: Yellowish-green to dark green, distinctly
ridged towards the tip.

Leaves: There are two mid-green basal leaves
(rarely three or four), strongly keeled and
oblong to oval-oblong in shape, and one to
three small, lanceolate, bract-like leaves further
up the stem. The leaves emerge from early May
onwards and persist until mid-September but,
in a dry season, they may not appear until early
June and will wither early.
Spike: The flowers appear densely packed on
the spike due in part to the relatively large
ovary. There are 20-30 flowers on most plants
but over 70 on the very largest. The spike is
often one-sided.
Bract: Green, lanceolate and roughly the length
of the ovary.
Ovary: Relatively large and inflated, even
when the flowers are fresh, greenish-yellow,
prominently ribbed and slightly twisted. The
ovary is upright but narrows at the top into
a stalk-like base for the flower which is bent
through more than 90, thus holding the flower
pendant.
Flower: The whole flower is greenish-yellow
and very small, about 2.5mm wide x 3mm from
front to back. It does not open widely and the
sepals, petal and lip all point forward and are
more-or-less parallel, giving the flower a spiky,
tubular or bell-like shape. The sepals are oval,
with the lateral sepals slightly smaller than the
upper sepal. The petals are slightly paler and,
rather unusually, longer than the sepals. They
are spear-shaped, with variable small side-lobes
towards the base and a narrower tip. The lip
has a long and narrow central lobe and rather
shorter side-lobes. To the rear the lip narrows
into a short, blunt chamber (a rudimentary
spur) which secretes nectar. There are two oval
pollinia, each attached by a very short, elastic
caudicle (stalk) to a saddle-shaped viscidium
that is almost the same size as the pollinia. The
viscidia each have a delicate skin but are not
enclosed in a bursicle (pouch). The flowers have
a sweet, honey-like scent.
Subspecies
None.
29/1/09 12:26:32
MUSK ORCHID

None.
BIOLOGY
The flowers are pollinated by a variety of tiny
insects such as flies, parasitic wasps, gnats and
beetles, typically just 1-1.5mm long. As it feeds
on the nectar at the rear of the lip, the insect
ruptures the skin of the relatively large viscidia,
and these stick the pollinia to the insects legs.
Once the insect leaves the flower, the pollinia
rotate forward to be in the correct position to
make contact with the stigma of the next flower
to be visited; this can be on the same plant.
There are conflicting reports on the possibility
of self-pollination. A study in Sweden found
that it did not take place, but it has also been
stated that the flowers can be self-pollinated,
the anther withering and the pollinia dropping
onto the stigma immediately below. Whatever
the mechanism, 70-95% of flowers set seed.
The species also reproduces vegetatively and
197
this may be the major means of recruitment

to the population; two to five daughter tubers
are produced at the ends of slender rhizomes
up to 20cm from the parent tuber. In this way
extensive clones can develop.

The leaves and flower spike grow from a single
spherical tuber which starts to wither away
by flowering time, and there are a few short,
thin roots. The tip of the tuber and usually
also the roots are infected with fungi. Two
or more new tubers are formed each growing
season, the larger of which provides for the
next years leaves and flower spike, the smaller
ones are daughter tubers. Musk Orchid is
sometimes described as migratory because the
replacement tuber grows at the end of a short
rhizome and therefore the aerial shoot appears
in a slightly different place each season. After
flowering once, plants may appear again merely
as vegetative rosettes or even remain dormant
underground for one or two years before they
flower again.
M 29 June, Hampshire. Musk Orchid can reproduce vegetatively to form extensive patches of clones.
29/1/09 12:26:33
198
GENUS hERMINIUM
Musk Orchid is reported to flower after

a period of immaturity lasting several years,
but in cultivation plants have flowered within
two years of seed being sown. Individuals can
be long-lived, with an age of 27 years being
recorded.
Hybrids
None.

The specific name monorchis means onetesticle (i.e. one-tuber). When flowering this
species appears to grow from a single tuber,
unlike members of genera such as Orchis and
Dactylorhiza, which have two obvious tubers
at the base of the stem. In fact, Musk Orchid
has several, but the replacement and daughter
tubers are formed at the end of thread-like
rhizomes and easily missed. The specific name
is therefore a misnomer.
HISTORY aND
CONSERVaTION
Musk Orchid was first recorded in Britain in
1663 by John Ray in his Catalogus Plantarum
circa Cantabrigiam nascentium (Catalogue
of plants found around Cambridge): Orchis
pusilla odorataIn the chalk pit close at
Cherry Hinton.
Past and present occurrence of Musk Orchid in Britain

1500-1999
current range
Britain
Ireland
104
32 (1.1%*)
% lost, 1500-1969
52%
% lost, 1970-1986
17%
% lost, total
69%
Not surprisingly, it is classified as Vulnerable.

The ploughing of chalk grasslands from the
late 18th century onwards caused the first
wave of losses, especially in East Anglia, and
many sites had gone by 1930, including John
Rays in Cambridgeshire. Subsequently, the
usual suspects of agricultural intensification,
the scrubbing-over of grassland (especially
following the outbreak of myxomatosis)
and overgrazing have taken their toll. It is
now extinct in Somerset, Glamorganshire,
Oxfordshire, Rutland, Cambridgeshire, Suffolk
and west Norfolk, where three 18th or 19th
century records included a site at Heacham, the
northernmost locality for the species. However,
it can, at least occasionally, colonise new areas.
Many sites are now protected as SSSIs and
reserves, and a few colonies hold hundreds or
even thousands of plants.
Despite its predilection for dry chalk
grassland, the species is vulnerable to
summer drought, which can lead to the early
withering of the leaves and thus a reduction
in the amount of carbohydrates stored in the
tubers for the following season. In some cases
there may then be a big fall in the number of
flowering spikes, the plants appearing merely
as non-flowering rosettes, but the population
can recover rapidly following a good growing
season. Prolonged droughts can, however,
devastate the species; the long, hot summer of
1976, the worst drought in southern England
for 250 years, caused big losses in Sussex from
which it has not yet recovered.
Musk Orchid has been lost from 69% of its

total historical range and is Nationally Scarce,
having been recorded from just 32 10km
squares in the New Atlas period of 1987-99.
P 14 May, Kent, Lady Orchid.
29/1/09 12:26:34
Genus ORCHIS
typical orchids
29/1/09 12:26:38
200
GENUS ORChIS
This genus contains some of the rarest and most spectacular orchids in the British Isles.
Many species have flowers that recall tiny human figures, with lips that are divided into
arms and legs. All grow from tubers and can spend part of the year underground in a safe
resting state. This allows the orchid to survive unfavourable periods, such as the summer
drought in the Mediterranean or sub-zero winter temperatures.
Distribution
There are about 50 species, most of which are
found in Europe and adjacent areas in North
Africa and the Middle East. Just five species
occur in the British Isles.
Floral structures
There are two pollinia, each narrowing to a
caudicle (stalk) which is attached to one of
the two viscidia. The viscidia are each in turn
contained within a bursicle, a pouch-like cavity
on the column.
Pollination
Members of this genus are pollinated by insects,
but only a few species produce nectar to reward
their pollinators. The majority rely instead
on deceit; insects are attracted to the brightly
coloured and scented flowers but receive
no reward.
Growth pattern
At flowering time all Orchis species have two
almost spherical tubers side by side at the
base of the aerial stem. The tubers are more
August
October
December
February
accurately termed root-tubers or root-stem

tubers and are essentially roots that have
been modified to become specialist storage
organs.
To follow the pattern of growth through
the year it is best to start in the late summer
when the orchid is resting and consists only of
a single tuber with one terminal bud; the leaves,
rhizome and roots die off once the orchid has
flowered.
In the autumn the bud on the tuber
produces a short rhizome and from this a few
roots develop. The roots serve two functions.
First, their infection with fungi provides the
plant with nutrients. Second, they supply the
plant with water and this becomes particularly
important once the leaves have appeared. In
some members of the genus the leaves develop
in the autumn, in others they do not expand
until the spring.
Once the roots have developed, they begin
to produce nutrients. Some of these are stored
in a new tuber that has started to form on the
April
May
August
M Annual cycle of growth and replacement of tubers in the genus Orchis (after Wells, 1981).
29/1/09 12:26:39
GENUS ORChIS
rhizome, side by side with the old; due to its

relationship with fungi, the orchid can produce
nutrients even before its leaves appear. The
rhizome goes on to form the aerial stem and
when the flowers open in the spring this has
two tubers at its base. The older of these has
supplied the current seasons growth, including
the leaves, stem and flower spike. This tuber is
starting to shrink and will have vanished by the
late summers resting period. The newer tuber is
plump and swollen and continues to grow until
the leaves die down. It will go on to overwinter
and form the flower spike in the following year.
Fungal partners
The tuber does not have a fungal infection
although fungi are found in its epidermis (and
in hot, dry, climates the fungus may only be
able to survive the summer drought in the
skin of the orchids tubers). Fungal activity is
concentrated in the roots and sometimes also
the rhizome. All Orchis species are able to spend
one or more years underground, presumably
sustained by fungi.

In all species the seed germinates in late
summer or autumn and forms a protocorm.
This rapidly forms the first root, which is
infected with fungi and supplies the protocorm
with nutrients. The following summer, a small
Additional tubers may be formed at the base
of the aerial stem and these will go on to form
separate plants as the connecting stem dies off
in the autumn.
Name
The generic name Orchis derives from the
Greek orkhis and means testicle. The two
rounded tubers have long been considered to
resemble male genitalia.
replacement
tuber
bud
spring
201
rhizome develops and the first tuber develops

on this rhizome, the root and protocorm dying
away in the late summer to leave only the tuber.
In the autumn, a new rhizome grows from this
tuber and produces one or more roots and in
the spring a leafy shoot is produced and a new
tuber forms at its base during the summer; the
pattern of growth is then similar to that of the
adult plant. Some species spend several years
as an underground seedling, producing a new
tuber each year.
The pattern of development of the seedling
apparently varies slightly between species and
even between populations. Some may produce
a tuber directly from the protocorm, without
any roots developing beforehand, whereas
others develop an aerial stem and root in the
first season after germination and then go on to
produce a tuber.
protocorm
root
tuber
summer
autumn
spring
YEAR 2
summer
autumn
YEAR 3
M Development from seed in the genus Orchis (after Rasmussen 1995).
29/1/09 12:26:40
202
GENUS ORChIS
man orchid
Endangered, BaP
Orchis anthropophora
Formerly: Aceras anthropophorum
The narrow flower spikes of Man Orchid do not command attention from a distance but on
closer acquaintance they reveal how apt the English name is. Just why do they look so much
like tiny, hooded, human figures, and what benefit does the plant gain by having flowers of
this particular size and shape? As with so many questions about orchids, we do not know
the answer. Man Orchid is relatively widespread in southern England, occurring north to
Lincolnshire and west to Gloucestershire, but away from Kent and Surrey it is extremely
local and either uncommon or rare; in the long term the future of the few remaining isolated
populations must be in doubt.
Identification
The long, narrow spike and very man-like
flowers are distinctive. The sepals and petals
form a hood or cowl and the tiny figure faces
downwards, concealing its face. The lip is deeply
lobed to form the arms and legs. The flowers are
yellowish to greenish, variably washed with red;
plants in full sun may be on average the reddest
and some may even have bright foxy-red lips.
Similar species
Frog Orchid may be similarly coloured but is
usually rather smaller, and the lip is not divided
into arms and legs.
Common Twayblade is also vaguely similar
but the tiny green flowers are rather different in
shape and it has only two large, rounded leaves.
Habitat
Typically found on well-drained grassland
on chalk or limestone, often on or at the foot
of a slope, with a predilection for abandoned
quarries and pits. Roadside verges, churchyards,
field margins and stabilised dunes or shingle
can also provide suitable habitat. It frequently
grows in relatively long, rank grass and among
scrub and it will sometimes spread under the
eaves of nearby woodland. It is vulnerable to
O 2 June, Northamptonshire. The spikes are often tall
and very slender; the basal leaves may be hidden (or nib
bled by rabbits).
P 2 June, Northamptonshire. Although classified as
Endangered, Man Orchid can still be found in good num
bers at a few favoured sites.
29/1/09 12:26:43
29/1/09 12:26:48
204
GENUS ORChIS
competition, however, and dense scrub will

crowd it out; conversely, populations can be
eliminated by heavy grazing.
Flowering period
Early May to late June, sometimes from late April,
but usually at its best in late May. The flowers
open in slow progression from the bottom of the
spike, so flowering can be protracted.
Range
Southern and eastern England, with the bulk of
the population on the North Downs in Surrey
and Kent. It is rare and very local elsewhere
and confined to the region south and east of
a line from Bristol to the Humber. There are
four or five sites in Hampshire, around four in
Suffolk, mostly on protected roadside verges,
and between two and four sites in Essex. Two
each remain in Sussex, Northamptonshire
and Lincolnshire, with just one in Wiltshire,
Gloucestershire, Oxfordshire, Bedfordshire and
Norfolk; there is a single site in Warwickshire
but it was apparently introduced there, in 1968.
World range: Western and southern Europe,
North Africa and the Levant. It ranges north
to Holland, southern Germany, Switzerland,
the former Yugoslavia and Greece and south
to the Mediterranean, including most of the
Mediterranean islands from the Balearics to
Cyprus. In North Africa it is found in Tunisia,
M 9 June, Northamptonshire. The flowers are truly

manlike.
Algeria and Morocco, and it also occurs in

southwest Turkey and Lebanon.
How to find it
A very localised species, nowadays only likely to
be found on reserves or protected road verges.
It can be hard to spot if it is growing among
long grass. Two of the best sites for this species
are Barnack Hills and Holes (Peterborough)
and Wye (Kent). Box Hill (Surrey) is another
classic locality, although it has declined there in
recent years.
DESCRIPTION
1987-99
1970-86
pre 1970
Height: 15-65cm but usually 20-30cm and

rarely over 45cm.
Stem: Pale green with some membranous
sheaths at the extreme base.
Leaves: Green, dull or slightly bluish, distinctly
veined, keeled and narrowly oval-lanceolate to
strap-shaped. There is a basal rosette of three or
four leaves, some lying flat and some held at about
45, and higher on the stem one or two smaller
and more lanceolate sheathing leaves. In some
areas the leaves appear in spring, in others in
November or December, becoming fully formed
by January or February. In all plants the tips of
the lower leaves are often scorched by May and
they all die off after flowering; this pattern of
29/1/09 12:26:50
MAN ORCHID
growth is probably an adaptation to the mild wet

winters and hot dry summers in the core area of
its distribution around the Mediterranean.
Spike: Tall, narrow, more-or-less cylindrical
and dense, with up to 50 or even 90 flowers.
Bract: Green, lanceolate and half the length of
the ovary.
Ovary: Pale green, long, cylindrical, boldly
ribbed and twisted.
Flower: Green or yellow, variably tinged red, and
very man-like. The sepals are oval and various
shades of yellowish-green, often with a distinct
maroon fringe and midrib. The petals are pale
green, slightly shorter, much narrower and more
strap-shaped. Both the sepals and petals form
a hood over the column, with the petals fully
concealed. The lip is variably green or yellow,
often strongly washed red or reddish-brown,
especially around the edges, but can be pure red
or yellow. It hangs almost vertically downwards
and has three lobes: two long, narrow side-lobes
at the base (the arms) and a terminal lobe that is
itself divided half way to the base into two lobes
(the legs). There is sometimes a tiny projecting
tooth between the legs. There is no spur, rather
there are two shiny, whitish swellings on either
side of the base of the lip which curve round to
join the column and enclose a shallow pit with
two small, nectar-secreting depressions. The
flowers have a faint, unpleasant smell.
205
moderate to good, but despite this it is thought

that most reproduction is vegetative via the
production of additional tubers.

Once they have appeared above ground for
the first time, plants may live for up to 14
years although they may not flower every year
or even appear above ground. Conversely,
some may flower for five years in a row. Man
Orchids only rarely die after flowering just
once, and in a study in Bedfordshire the
half-life averaged 5.8 years and varied from
4.0-7.8 years (the half-life is a measure of
the life expectancy of the orchid after its
first appearance above ground and marks the
Subspecies
None.

Var. flavescens lacks red pigments (anthocyanins)
and has a green hood and contrasting yellow lip.
It is rare.
BIOLOGY
Little is known about the pollination of this
species. However, the numerous hybrids with
Lady, Monkey and Military Orchids found
in Europe would suggest that it shares a suite
of pollinating insects with those species; in
England ants and hover flies have been seen
with pollinia on their heads. Seed-set is
M 8 May, Kent. Many flowers have largely red arms and

legs.
29/1/09 12:26:51
206
GENUS ORChIS
point at which 50% of the population that

emerged in any given year have died). There
is no information on the period between
germination and flowering.
Hybrids
In Europe, hybrids with Monkey, Military and
Lady Orchid are common but in England such
hybrids have only been found twice.
O. x bergonii, the hybrid with Monkey Orchid,
was found in Kent in 1985, although it has
been suggested that this may be the result of
inadvertent hand pollination.
O. x macra, the hybrid with Lady Orchid,
was found in Kent in 1998, when two plants
were seen.

The specific name anthropophora means manbearing.
Man Orchid was formerly placed in the
genus Aceras as Aceras anthropophorum. Indeed,
it was the only species in that genus, which was
distinguished from the genus Orchis by the lack
of a spur. Recent DNA studies have confirmed
that this difference is purely superficial and that
Man Orchid is a perfectly good Orchis. The
generic name Aceras derived from the Greek
and meant without-a-horn, a reference to this
absence of a spur.
have also been lost to scrub encroachment. Extinct

in Somerset, the Isle of Wight, Hertfordshire,
Buckinghamshire, Cambridgeshire, Leicestershire
and also Derbyshire (where it was introduced at
Ashover but not seen there recently).
Past and present occurrence of Man Orchid in Britain and

1500-1999
current range
Britain
Ireland
109 (1.7%*)
48
% lost, 1500-1969
46%
% lost, 1970-1986
10%
% lost, total
56%
HISTORY aND
CONSERVaTION
The first British record dates from 1690, when
John Ray published his Synopsis Methodica
Stirpium Britannicarum (Methodical synopsis
of British plants): Found by Mr. Dale in an
old Gravel-pit at Dalington (Ballingdon) near
Sudbury.
Nationally Scarce and still decreasing in
numbers, Man Orchid has been lost from 56% of
its historical range and is classified as Endangered.
It was formerly much commoner in East Anglia
but vanished as pastures were ploughed from
the late 19th century onwards. Sites were also
destroyed when quarries and pits were used as
landfill, field margins were sprayed or subject to
drift from nearby operations and road verges cut
or sprayed unsympathetically. Conversely, sites
M 2 June, Northamptonshire. Some flowers are much

yellower, with little or no red.
29/1/09 12:26:53
MONKEY ORCHID
monkey orchid
207
Vulnerable, BaP
Orchis simia
One of our rarest orchids, the exotic, monkey-like appearance of the flowers, combined with
the inevitable secrecy that surrounded such a rarity, has excited botanists for generations. It
is confined to three sites in the Chilterns and Kent, two of which welcome visitors.
Identification
To see a Monkey Orchid you will almost
certainly have to visit one of its two public sites,
so identification is straightforward. The flowers
are distinctive and definitely resemble a Spider
Monkey, with the hood forming the head and
the lobes of the lip, the slender, curved arms
and legs.
Similar species
Military Orchid is also very rare and the two
species are not found together anymore in
England. However, should a new colony of
plants be discovered, identification would be an
issue. In Military Orchid the legs are straight,
distinctly broader than in Monkey Orchid and
widen towards the tip, and the hood forms a
longer, neater helmet. The shape of the flower
spike is rather different, too, being taller and less
crowded, with the flowers opening in sequence
from the bottom of the spike upwards.
Habitat
Monkey Orchid prefers south-facing slopes
on open, grazed chalk grassland. It probably
favours the interface between grassland and
woodland or scrub, benefiting from the shelter
which scattered trees and shrubs provide from
desiccating winds and grazing animals. The
slightly moister conditions in light shade are
also likely to be beneficial and the bare ground
under scrub can provide suitable conditions
for seedling establishment. Should the shade
become too dense, however, it ceases to flower
or even to appear above ground. Monkey
Orchid may be badly affected by drought; for
example at Faversham in Kent the population
crashed after the hot, dry summers of 1975
and 1976 and was slow to recover.
P 28 May, Kent. The rosette of shiny, green basal leaves

is usually hidden in the grass.
29/1/09 12:26:54
208
GENUS ORChIS
Range
Currently confined to Hartslock in Oxfordshire
and two sites in Kent (a confidential site near
Faversham and Park Gate Down, where it was
introduced). World range: Southern Europe
and the Mediterranean region, including
the Balearics, the Aegean islands, Crete and
Cyprus, north to Holland (very rare), southern
Germany, Hungary and Romania. Ranges east
to Syria, Turkey, the Crimea, Caucasus, Iran
and Turkmenistan, and also found in North
Africa in Algeria, Tunisia and Libya.
1987-99
1970-86
pre 1970
How to find it
M 28 May, Kent. At Parkgate Down in Kent, Monkey
Orchid grows on open downland, but it can also grow in
light scrub.
Park Gate Down and Hartslock reserves are the

places to go, and Monkey Orchids are easy to
find at both.
Flowering period
DESCRIPTION
Late May to early June; flowering peaks

around a week earlier in the Chilterns than
in Kent. In most orchids the flowers open
from the bottom of the spike upwards, but in
Monkey Orchid the flowers often open rapidly
from the top downwards, although they may
open synchronously (or even from the base
upwards in Kentish plants). Overall, it is the
rapidity with which all the flowers open that is
characteristic of Monkey Orchid, rather than
the direction of opening. The spike is at its best
for a rather brief period.
Height: 10-30cm tall, occasionally to 45cm.

Stem: Green, usually washed brownish-purple
towards the tip, angled and with two or three
sheaths at the base.
Leaves: There are three or four shiny green
basal leaves, oval-oblong, often keeled and
blunt-tipped, and two or three sheathing leaves
higher on the stem. The leaves may appear
above ground as early as November but more
usually do so in January or February.
Spike: Roughly globular in shape with the
flowers crowded together, and the whole thing
29/1/09 12:26:56
MONKEY ORCHID
looking a bit scruffy. Most spikes have ten

to 20 flowers but there are up to 30 on welldeveloped plants and as many as 42 on the
most robust.
Bract: Very small, around a third of the length
of the ovary, triangular, chaffy and whitish.
Ovary: Green, heavily washed purple, boldly
six-ribbed, twisted and curved.
Flower: Monkey-like, white or pink, with
darker spots and variably redder extremities.
The sepals are lanceolate, and the petals are
slightly shorter, much narrower and more
strap-shaped; together they form a hood that
encloses the column with the sepals slightly
splayed outwards at the tips. The sepals are
whitish on their outer surface, variably washed
pink and with irregular violet-purple dots,
blotches and streaks. The inner surfaces are
more heavily blotched and streaked, sometimes
almost solidly so. The petals are similar but
sometimes more solidly washed purplish-pink.
The lip has two lobes forming the arms and
two lobes forming the legs, with an additional
small projection between the legs (more
phallic than tail-like); the tips of the arms and
legs are curved forwards and upwards. The
lip is whitish in the centre, variably flushed
violet-purple and spotted violet (the spots are
formed by tufts of papillae), becoming violetpurple towards the extremities of the arms
and legs. The spur is pale pink, about half to
three-quarters the length of the ovary, slightly
down-curved and blunt-tipped. The column
is reddish. The flowers have a faint scent of
vanilla.
Subspecies
None.

The Kentish Monkey Orchids are on average
taller than those in the Chilterns, with stouter
stems and, more significantly, more and bigger
leaves; their largest leaf is up to twice as long.
Their flowers have a more extensive area of
purple spotting on a more purple-washed
ground, and the spots are larger. The sepals
have larger and darker markings on their inner
209
M 28 May, Kent. With long arms, legs and a tail, it is

not hard to see the monkey in a Monkey Orchid.
surfaces, making the hood appear pale pink

rather than white on the outer surface, and
the spur is pale pink to purple (white to pale
pink in the Chilterns). However, there is much
overlap, and the differences do not warrant the
recognition of distinct varieties. Var. macra,
with a violet hood and purplish-red arms and
legs, was recorded many years ago and may
have been more prevalent in large populations
but probably falls within the normal range of
variation and may not be worthy of recognition.
BIOLOGY
Some natural pollination does occur in
England, with the flowers visited by flies,
bees and butterflies. Although the spur is not
thought to contain nectar, two swellings near
its mouth may contain sugars which can be
extracted by insects. Seed-set has been poor in
England and some populations have been handpollinated. Pollination rates appear to improve
significantly, however, once a population of
Monkey Orchids is above a certain threshold;
29/1/09 12:26:58
210
GENUS ORChIS
O 28 May, Kent. Kentish

Monkey Orchids are, on
average, taller and more
richlycoloured than those
in the Chilterns.
the spectacle of large numbers of flowers may

be more attractive to potential pollinators. In a
study in Holland, vegetative reproduction was
found to be rare and only to occur following
good growing seasons.

Much of the information about the life cycle of
the Monkey Orchid comes from observation
of a small population in the Netherlands. This
originated with a single founder plant and may
not be entirely typical. A period of three or

four years elapses between germination and the
appearance of the first aerial leaf and a further
three to six years before flowers are produced,
by which time the plants have at least four basal
leaves. A similar time scale has been recorded
for Kentish Monkeys. Young plants occasionally
disappear underground after they have
produced their first leaves and then reappear
again after one or two years.
29/1/09 12:27:01
MONKEY ORCHID
Individual plants can be long-lived, flowering

for up to 19 consecutive seasons although this
may be exceptional. Plants often rest between
bouts of flowering as vegetative rosettes and may
even be dormant underground for one or two
years. If absent for three years, however, they
are almost certainly dead. Severe winters will
inhibit flowering and may result in many plants
dying, as the tubers typically lie in shallow soil.
Similarly, if the leaves are grazed off the plant
will not flower the following season, presumably
because without leaves it is unable to build up
sufficient resources.
Hybrids
O. x. beyrichii, the hybrid with Military
Orchid, occurred in the Thames Valley until the
middle of the 19th century, after which the two
parent species were not found together.
O. x. bergonii, the hybrid with Man Orchid, was
recorded in 1985 at Faversham in Kent (but it
has been suggested that this may have been the
result of hand pollination gone wrong).
The hybrid with Lady Orchid has appeared
in small numbers since 2006 at the Hartslock
reserve.
211
Oxfordshire and East Yorkshire but there

are now only three sites, one in Oxfordshire
and two in Kent. A Red Data Book species,
it is classified as Vulnerable and is specially
protected under Schedule 8 of the Wildlife and
Countryside Act 1981.
Monkey Orchid has apparently always
been very local in England, but in the 18th and
early 19th centuries it was frequent in south
Oxfordshire, in the area between Wallingford,
Reading and Henley. Most records came from
the slopes overlooking the north bank of the
Thames between Goring and Caversham
(the latter now on the northern outskirts of
Reading) where the river cuts through the
southern outliers of the Chilterns. The species
then declined dramatically from about 1840,
due to the ploughing of downland and the
collection of specimens for both herbaria
and gardens. However, the main cause of its
demise may well have been a big increase in
the number of rabbits. By the mid-1920s there
was only one substantial colony remaining in

The specific name simia means of the ape or of
the monkey.
HISTORY aND
CONSERVaTION
In 1666 Christopher Merrett noted both
Monkey and Military Orchids in his Pinax
Rerum Naturalium Britannicarum (A picture of
British natural history): on several Chalkey
hills neer the highway from Wallingford to
Redding on Barkshire side the river This
reference to the Berkshire Downs was apparently
the first record of both species in England
and originated with information supplied by a
William Brown of Magdalen College, Oxford.
Old records of Monkey Orchid are often
difficult to assess because for a long time the
species was confused with Military or even
Lady Orchid. Monkey Orchid went on to
be recorded from Kent, Surrey, Berkshire,
M 28 May, Kent.The monkeys often look as if they have

been thrown into a heap.
29/1/09 12:27:02
212
GENUS ORChIS
the Chilterns, at Hartslock, but such was the

secrecy surrounding this site that the species
was generally thought to have become extinct in
Britain.
Past and present occurrence of Monkey Orchid in Brit

1500-1999
current range
Britain
Ireland
10
2 (0.07%*)
% lost, 1500-1969
60%
% lost, 1970-1986
20%
% lost, total
80%
The Hartslock colony was stable at over 100

flowering plants in the 1920s and increased
to 200 spikes in the years leading up to World
War Two; in 1933 over 30 were picked. But,
following the war, the increasing mechanisation
of agriculture allowed steeper and steeper
slopes to be cultivated and in 1949 and 1950
the field at Hartslock that held the Monkey
Orchids was ploughed. Fortunately, the upper
part of the slope (the area where the orchids
now grow) escaped as it was steeper, scrubbier
and protected by a thick hedge. There may
already have been a few Monkey Orchids
growing in this refuge, and some tubers rescued
from the lower part of the field were replanted
there but with unknown results.
Just one Monkey Orchid flowered at
Hartslock in 1950-52 and numbers remained
painfully low for many years. It was not until
1968 that the population reached even the
modest total of eight flower spikes, and even
in 1977 there were just eight plants. In 1975
BBOWT bought Hartslock but, as on most
downland sites, scrub encroachment was a
problem. This led to a substantial programme
of scrub removal, and the area is now grazed in
the autumn and winter by sheep.
Despite the initial management efforts,
the orchid population increased very slowly
to around 60 plants in the late 1980s, with

roughly a third flowering each year. In the early
1990s the number of flower spikes actually
dropped, to as few as five in 1990, although
the total number of plants was more constant.
Then, in 1994 the population at Hartslock
started to expand rapidly. Three factors may
have helped; first, a run of mild winters, which
probably reduced mortality and encouraged
plants to flower. Second, systematic hand
pollination, which began in 1977; this has now
ceased, as rates of natural pollination and seed
production are high. Third, in 1992 the main
colony was fenced against rabbits. Over the
next three years the number of plants trebled
and by 1995 there were 123, including 47 new
individuals, of which 72 flowered. It is thought
that prior to the erection of the fence rabbits
ate most of the first orchids to emerge; they can
appear above ground as early as January, well
before the seasonal warden puts wire netting
cages on the orchids in April.
By 1999 there were 200 plants with 100
of them flowering, and from 2000 Hartslock
held around 300 plants, with one to two-thirds
flowering each year. Despite this apparently
healthy population, however, DNA studies have
shown that the genetic variability at Hartslock
is very low, as the population had gone through
a genetic bottleneck when it was reduced to a
tiny handful of individuals. In the long term, it
may be better to introduce more variability by
hand pollinating again using pollen from other
Chiltern plants, Kent or even France. In the
last few decades Monkey Orchid has appeared
sporadically and in very small numbers at other
sites in the Chilterns. In 1966 it was found
near Pangbourne and in 1965 and 1971-74 at
Aston Rowant.
In Kent, Monkey Orchid was first recorded
in 1777 from near Faversham and then again
in the early 1800s but it was not seen again
until 1920-23 when a few plants flowered at
Bishopsbourne (near Canterbury). At the other
end of the county, in west Kent, from 1952
onwards a single Monkey Orchid appeared
on the rough grass of a disused tennis court
29/1/09 12:27:02
MONKEY ORCHID
at a vicarage at Otford. Every year, until 1955,

the vicar took the seed capsules and scattered
seed onto nearby downland. In 1956 there
was a single robust spike and a further six
non-flowering plants at the vicarage. However,
on the retirement of the botanically minded
vicar the new incumbent would not guarantee
to safeguard the colony. All the orchids were
moved to nearby private land, where the largest
flowered once only, in 1957, and then the
Monkeys vanished.
In 1955 a Monkey Orchid appeared near
Faversham in Kent (at the current native site),
but the single plant was eaten, perhaps by a
horse. It may well have been able to bloom
due to the outbreak of myxomatosis which
resulted in the mass-flowering of orchids in
Kent in that year. More were found in the
following years, with up to 38 plants, ten of
which flowered, in 1957-58, growing partly on
open downland and partly in a nearby hazel
copse. From 1958 until at least 1985 the plants
in this colony were hand-pollinated to make
sure that seed was produced. Numbers steadily
increased, with 246 plants and 162 flowering
spikes in 1964 and 205 flowering plants in
1965. The 1975-76 drought badly affected
the colony, however, with none flowering but
in 1977 plants reappeared and by the mid1980s there were again 30-50 plants, with ten
or so flowering. The population here is now
stable at over 200 plants, although only a small
proportion flower.
From 1958 onwards seed from the
Faversham site was scattered at several other
places in Kent, and a population became
established at Park Gate Down. The first three
plants flowered in 1965, seven years after seed
was sown, but then not again until 1976 when
two spikes appeared. This population stabilised
at about 100 plants in the mid-1990s but has
increased significantly since then.
A few plants appeared in dune grassland at
Spurn Point in southeast Yorkshire in 1974,
over 250km from the nearest known source of
seed. This colony increased to a maximum of 25
plants, with nine flowering, but only persisted
213
M 28 May, Kent.The colour of the flowers is very variable.

Some appear pale pink at a distance, others are darker
and redder.
until 1983, after which the site was washed

away in a storm.
The Sainsbury Orchid Project organised by
the Royal Horticultural Society has become
involved in the conservation effort. Each year a
few seedpods are sent to Kew from Hartslock.
Progress with propagating Monkey Orchid
has been slow because cultivation has been
restricted to sterile, asymbiotic techniques, and
germination and growth rates are low. By 1996,
ten plants had been raised to a size where
they could be planted out, and in September
1996 the tubers were placed in two clumps on
the edge of the existing colony. In 1997 two
of these came up, but since then only one has
appeared.
29/1/09 12:27:03
214
GENUS ORChIS
military orchid
Vulnerable
Orchis militaris
The Military Orchid combines two of the qualities which make orchids so alluring; great
rarity and great beauty. It is one of Britains most attractive species and is found regularly
at just three sites (two of which are, happily, open to the public). Thought to be extinct
in Britain by the early part of the 20th century, it was dramatically rediscovered in 1947,
although for a long time the site was kept a closely guarded secret. Like its cousin, the
equally rare Monkey Orchid, it has been monitored, managed and mollycoddled since its
rediscovery, but we are still far from unlocking all its secrets.
Identification
One of the so-called manikin orchids in which
the flower resembles a tiny human figure,
Military Orchid brings to mind a soldier. The
sepals and petals form a helmet, purple-striped
on the interior, and the lip has four lobes, two
for the arms and two for the legs. The rows
of purple spots down the centre of the lip are
reminiscent of buttons on a soldiers tunic.
The allusion to the military was coined before
soldiers habitually wore red uniforms and may
refer to the resemblance of the hood to an
ancient coal-scuttle helmet.
Similar species
Monkey Orchid resembles this species in the
general structure of the flower, but its legs are
kinked, narrower and do not broaden towards
the tip, and the hood formed by the sepals and
petals is more open. Also, its flower spike is
not only shorter and more crowded but also
more jumbled and disarrayed, lacking military
precision, and all the flowers open at roughly
the same time.
Habitat
Military Orchid is found in grassland, scrub,
woodland glades, on woodland edges and,
formerly, rough fields. It always grows on chalk.
The species does best in light scrub on old
pastures and in the shelter of woodland edges.
It favours some shade and needs bare ground
for seedling establishment (rather than a closed
grass sward), but it does not do well if there is
too much shade. Recorded up to 183m above
sea level in the Chilterns.
M 31 May, Suffolk. The colony at Mildenhall has thrived

since the overshadowing trees were removed.
29/1/09 12:27:04
MILITARY ORCHID
1987-99
1970-86
pre 1970
Flowering period
Mid-May to mid-June, the flowers are at their
best in late May and early June. Once a flower
has been pollinated it usually shrivels within
a day.
Range
Currently confined to two sites in the Chilterns,
in Buckinghamshire and Oxfordshire, and
one in Suffolk. Plants have been introduced
into industrial waste-ground at Bolton in
Lancashire (apparently from outside the United
Kingdom) and to sites in Cambridgeshire and
Kent. World Range: Europe and Siberia. In
Europe occurs north to Holland, northern
Germany, southeast Sweden and Estonia and
south to northern Spain, central Italy, the
former Yugoslavia, Bulgaria, Romania and
European Turkey. Confined to the mountains
in the southern parts of the range. In Siberia it
extends east to the Altai Mountains and Lake
Baikal.
How to find it
Homefield Wood in Buckinghamshire and the
Rex Graham Reserve at Mildenhall in Suffolk
are the two public sites for the species.
DESCRIPTION
Stem: Green, variably tinged purple towards
the tip.
215
Leaves: Bright shiny green, prominently keeled,

strap-shaped or, on more robust plants, broader
and more oval, and slightly hooded at the tip.
Two to five leaves are carried in a basal rosette
at around 45 above the horizontal, and there
are two or three sheathing leaves higher on
the stem, although the upper part of the stem
is bare. In Suffolk the tip of the shoot appears
above ground in late December or more usually
early January and the leaves start to unfurl by
early March.
Spike: Oval or conical, becoming cylindrical
as all the flowers open. There are two to 25
flowers but up to 57 have been recorded on a
particularly robust spike.
Bract: Triangular to oval, very short (much
shorter than the ovary) and green, strongly
washed purple or rose.
Ovary: Green, washed purple, boldly ribbed
and strongly twisted.
Flower: Man-like, whitish, with purple spots
and purple extremities. The sepals are oval
with pointed tips, and the petals are narrower
and more strap-shaped; they form a hood
with the tips of the sepals swept upwards and
the mouth of the hood very open. The sepals
are pale dove grey, their outer surfaces lightly
washed with lilac, becoming more purplish at
the base; the unopened buds are therefore pale
pinkish-grey. The inner surfaces have bold,
longitudinal purplish lines and an irregular
purplish wash. The petals are more uniformly
pale purplish. The lip has two narrow, strapshaped lobes forming the arms and two shorter
and broader lobes forming the legs, with an
additional small pointed projection between
the legs. It is whitish, flushed pink, with the
arms and legs more-or-less solidly purple.
There are two rows of purple spots down the
centre of the lip (the spots are formed by tiny
papillae) and solid dark purple lines along the
centre of the arms. The spur is purple, short
(about half the length of the ovary), cylindrical
and slightly down-curved. The flowers are
faintly vanilla-scented.
Subspecies
None.
29/1/09 12:27:05
216
GENUS ORChIS

Var. militaris is on average taller, with seven
to 42 flowers. It has a relatively lax, open spike
and paler flowers. The Suffolk colony has been
assigned to this variety which is widespread in
Europe.
Var. tenuifrons is on average shorter, with two
to 26 flowers, a denser spike, darker flowers
and narrower leaves. Endemic to England, the
Chilterns plants belong to this variety
BIOLOGY
Military Orchid is pollinated by hover flies
and bumblebees. It produces no nectar but
insects may be attracted to the sugary sap in
the wall of the spur. It has always been thought
that few flowers are pollinated in Britain; rates
of 3-11% are given for plants in the Suffolk
colony and 2-28% of flowers were recorded
as setting seed by Summerhayes (1968). At
Homefield Wood, however, 40% and 24% of
flowers were naturally pollinated in 1999 and
2000 respectively. Vegetative reproduction is
probably important for the British populations,
maintaining numbers when recruitment from
seed is low.
M 31 May, Suffolk.
This is a relatively long-lived orchid. Many

plants will live for ten years after their first
appearance above ground and a significant
proportion live for at least 17 years. In a study
in Suffolk the half-life varied from 2.2-7.8 years
(the half-life is a measure of the life expectancy
of the orchid after its first appearance above
ground and marks the point at which 50% of
the population that emerged in any given year
has died).
The period between germination and the
appearance of the first aerial shoot is three to
five years and between 30% and 50% of plants
flower in their first year above ground. Once
they have appeared, most plants flower at least
once and many do so every year, but intervals
of 11 years between flowering have been noted.
Perhaps a third of plants retreat underground
29/1/09 12:27:11
MILITARY ORCHID
217
during their life, usually spending only one year

dormant, but in some cases up to three years
and absences of up to eight years have been
recorded in Suffolk. In any given year, 5-15%
of the adult population can be underground.
However, dormancy is not a good option for
the plants as underground plants have the
highest probability of dying.
Hybrids
O. x. beyrichii, the hybrid with Monkey
Orchid, occurred in Oxfordshire until the
middle of the 19th century, after which the two
parent species were not found together.

The specific name militaris means military.
HISTORY aND
CONSERVaTION
A Red Data Book species that is classified as
Vulnerable and specially protected under
Schedule 8 of the Wildlife & Countryside
Act 1981.
Gerards Historie of Plants, first published
in 1597, contains a description of Souldiers
Satyrion or Soldiers Cullions (literally
testicles) which presumably pertains to the
Military Orchid: Souldiers Satyrion bringeth
forth many broad large and ribbed leaves,
spread upon the ground like unto those of the
great Plantaine: among the which riseth up a fat
stalke full of sap or juice, clothed or wrapped
in the like leaves even to the tuft of flowers,
whereupon doe grow little flowers resembling
a little man, having a helmet upon his head, his
hands, and legs cut off; white upon the inside,
spotted with many purple spots, and the backe
part of the flower of a deeper colour tending
to redness. The rootes be greater stones than
any of the kinds of Satyrions. Gerard gives,
however, no localities for his Souldiers Satyrion
and may have copied his description from a
European herbal.
The earliest localised record for the British
Isles was in 1666 when Christopher Merrett
noted both Military and Monkey Orchids in
his Pinax Rerum Naturalium Britannicarum
M 31 May, Suffolk. A fewflowered spike with very lightly

marked flowers.
(A picture of British natural history): on

several Chalkey hills neer the highway from
Wallingford to Redding on Barkshire side the
river This reference to the Berkshire Downs
was apparently the first record of both species
in England and originated from information
supplied by a William Brown of Magdalen
College, Oxford.
Military Orchid was often confused with
Monkey and Lady Orchids, and many older
records, unless supported by identifiable
specimens, are open to doubt. It was recorded
reliably from Belchamp Walter near Sudbury in
Essex in 1729, and there are also old records for
the North Downs of Surrey, including Box Hill
in the 1830s, and Kent (a herbarium specimen
dated 1836 from Cobham near Rochester).
Like Monkey Orchid, the stronghold of
the species was always the Chilterns, where
it ranged from just west of the Thames in
the region of Streatley and Basildon, along
the Chiltern Hills through Oxfordshire and
29/1/09 12:27:12
29/1/09 12:27:14
MILITARY ORCHID
Buckinghamshire, to the neighbourhood of

Tring in Hertfordshire and to Harefield in
Middlesex. It seems to have been particularly
abundant around High Wycombe and between
this town and Great Marlow on the Thames
(Summerhayes, 1968). Once fairly common
within this limited area, it declined dramatically
from around 1850 and was thought to be
extinct after 1929 when the last specimen was
collected from Hertfordshire. Several factors
contributed to the decline, including the
ploughing-up of downland and the collection
of specimens for both herbaria and gardens,
but the main cause may well have been a big
increase in the number of rabbits. There have
been no subsequent records from Berkshire,
Hertfordshire or Middlesex.
The Military Orchid became something
of a Holy Grail for British botanists. Indeed,
it was the title and main subject of Jocelyn
Brookes semi-autobiographical The Military
Orchid, published in 1948. In May 1947 it was
dramatically rediscovered by J. E. Lousley at
Homefield Wood in Buckinghamshire, an area
from which it had not previously been recorded.
The excursion was intended as a picnic, so
I had left my usual apparatus at home and
took only my note-book. But I selected our
stopping places on the chalk with some care,
and naturally wandered off to see what I could
find. To my delight I stumbled on the orchid
just coming into flower (Lousley 1969). There
were 39 plants, with 18 flower spikes, although
five had been bitten off, probably by rabbits.
Lousley noted that the orchids growing in
shade were either flowerless or had small spikes
and thought that the increase in available light
after trees were felled during World War Two
had probably prompted the appearance of
the plants.
Lousley kept his discovery a closely guarded
secret, fearing that the colony would be wiped
out by collectors, and many others tried to track
down this hidden treasure but to no avail. At
last, in 1956, after a long and systematic search
O 31 May, Suffolk.
219
M 31 May, Suffolk. A richlycoloured spike.
of all likely sites, the colony at Homefield Wood

was found by Richard Fitter and Frances Rose.
They dispatched a postcard to Lousley with
the simple but cryptic message The soldiers are
at home in their fields. The discovery was not
made public even then, but Homefield Wood
came to be managed by BBOWT in 1969
and they eventually announced to the press in
1975 that the Military Orchid had returned.
This resulted in headlines like The Beauty that
must blossom in secret. It was not until the end
of the 1980s that the location was made truly
public, and visitors are now welcomed.
Homefield Wood was originally largely
composed of Beeches with some areas of
chalk grassland on a south-facing slope. The
wood was largely clear-felled in the winter
of 1947, and the Military Orchid was found
in the spring of that year on chalk grassland
with scattered trees and scrub. The Forestry
29/1/09 12:27:16
220
GENUS ORChIS
Commission has owned the wood since 1955,

and most of the felled areas were replanted
with conifers in the 1950s. The area around
the colony was planted with Beech in 1960-61,
and other trees regenerated naturally to form a
mixed woodland; trees also invaded
the grassland.
When first found, there were two distinct
colonies at Homefield Wood. Colony A held 31
plants in 1947 and peaked at 35 in 1949, but
the numbers fell steadily until the last few were
seen in 1958, with a single plant reappearing
in 1961. Myxomatosis had decimated rabbit
populations in 1955, and this probably caused
a dramatic increase in scrub which, combined
with disturbance during tree-planting and
the spread of Rosebay Willowherb, led to the
extinction of this colony. Colony B presumably
held just eight plants when found, but this
is the area where the orchid survives to the
present day. This colony was fenced off in 1968,
brambles invaded and a thick understorey
developed. The population of Military Orchids
slowly fell to a low point of 28 plants in 1984,
with just five flowering.
Active habitat management started in
1981 with the removal of scrub and later on
large, overshadowing Yews. This may have led
to a recovery in numbers in this area to over
50 plants (with over 30 flowering) by 1989.
In 1985 a small adjacent area was clear-felled
and the orchids eventually spread into this,
flowering for the first time in 1995; this clearing
held half the flowering plants by 2003. Military
Orchids also appeared over 100m away through
the wood in a third open area in 1983. Overall,
the number of plants at Homefield Wood
steadily increased to about 80 in 1995, with 45
in flower.
Since 1995 there has been a dramatic
upturn, with over 200 plants, at least half
of which flowered, in 2003. In recent years
management has included fencing the colonies
to exclude rabbits and deer during the growing
season. These enclosures are grazed by sheep
in the autumn and winter, before the orchids
appear above ground. They are also mown and
raked, and scrub is controlled. In addition,

up to 20% of the plants at Holmfield Wood
were hand-pollinated from 1986-98. Handpollination can result in almost 100% seed set
and does not weaken the plants (as had been
suggested); hand-pollinated individuals have
flowered every year for a decade.
Military Orchid was found at a second site
in the Oxfordshire Chilterns in 1970. The
number of flowering plants did not exceed five
until 1999 and there were none in 1984-87.
Since 1999, however, there has been a rapid
increase, with 25 flowering plants in 2003,
together with a further 25 vegetative rosettes.
This population was also hand-pollinated for at
least 10 years from 1988.
As part of the conservation programme,
plants propagated asymbiotically at the
Royal Botanic Gardens were planted out at
Homefield Wood and the nearby Warburg
reserve in 1996 (a total of 231 tubers, both
one and two years old). Survival was poor but
those that remained first flowered in 2000 and
2002 respectively. In addition, 25 wild plants
were transplanted from Homefield Wood in
2000 to a site around 25km away. Survival of
these mature plants has been better than young
seedlings, with some flowering and setting
seed.
On 2 June 1955 the Military Orchid was
found at Mildenhall in Suffolk, a region from
which there were no previous records. The
colony was in an old chalk pit where there were
at least 500 plants, mostly on a heap of pure
chalk among birch and Wild Privet. There were
over 100 flowering spikes, but within a short
time the majority had been nibbled off, perhaps
by deer, and only 16 fruiting spikes remained
by mid-August. The colony was within a
Forestry Commission plantation and at the
time of its discovery the surrounding pines
were only 1.5m tall, thus the site was open
and sunny.
By 1958 the number of plants in the Suffolk
colony had risen to 2,854 and remained at this
level until the late 1960s, although only about
10% flowered. However, the population then
29/1/09 12:27:16
MILITARY ORCHID
221
declined rapidly to 252 plants in 1971, with

perhaps only 100 in the following years. To
protect the orchids the Forestry Commission
had erected a tall wire fence around the pit in
the 1960s. This excluded both deer and people
and within a few years Sycamore and Wild
Privet had taken over. These were cleared in
autumn 1972 and the colony increased slightly
and stabilised at 300-400 plants, with about
100 flowering. Scrub clearance continued and
the overshadowing Corsican Pines were finally
removed in 1985-86, allowing in much more
light, and from 1987 there was a dramatic
increase in the population. In 1990 there were
279 flowering spikes and 1,115 plants and in
2000 there were 748 flowering plants and too
many non-flowering plants to count.
The story of the two Military Orchid
colonies highlights the need for the appropriate
management of rare orchids. For example,
in both cases the colonies were fenced in
the 1960s, and although this excluded deer
and rabbits it came close to destroying them
because scrub took over. The recent increases
in Military Orchid are likely to be directly
attributable to better management rather than
a more favourable climate. This species seems
well-adapted to hard winters and is therefore
unlikely to have benefited from recent mild
years, unlike Monkey Orchid.
Genetic fingerprinting has shown that the
three English colonies are distinct and may
represent independent colonisations from
Europe. Each colony is thus valuable and
worthy of conservation in its own right.
Past and present occurrence of Military Orchid in Brit

1500-1999
current range
M 7 June, Suffolk. A large spike, wellout.
Britain
Ireland
19
3 (0.1%*)
% lost, 1500-1969
79%
% lost, 1970-1986
5%
% lost, total
84%
29/1/09 12:27:18
222
GENUS ORChIS
lady orchid
Endangered
Orchis purpurea
The sight of a group of Lady Orchids in a woodland glade in May is always a delight. Even
those lucky enough to see them regularly cannot fail to be impressed by the spectacular show
put on by this stately orchid. It is largely confined to Kent where it is relatively common in
chalky woodlands on the North Downs.
Identification
Straightforward. It is usually rather large
and statuesque, and the unopened buds are
dark reddish-purple. Lady Orchid is one of
the manikin orchids, and the flowers form a
miniature human figure. The sepals and petals
form a dark bonnet which, with the unopened
buds, contrasts strongly with the whitish, darkspotted lip. The lip itself is divided into several
lobes to form the arms and the skirt of the lady.
Similar species
Burnt Orchid is superficially similar, with dark
buds, a dark hood and a white, purple-spotted
lip but it is very much smaller, seldom more
than 15cm tall.
Habitat
Lady Orchid is found in woodland, both
ancient woodland and secondary woods, but
almost always on thin, well-drained chalky soils
(rarely also on limestone or other calcareous
substrates). It favours beechwoods and often
grows on south-facing slopes, frequently on
banks or on the terraces formed by the root
plates of the trees, either among a carpet of
Dogs Mercury or on bare leaf-litter. However,
its preferred habitat may be scrub or coppice,
and it does not flower so freely in shade, being
happier in open, well-lit situations, such as
along paths and rides, in clearings and along
the lower edges of woods. Indeed, it may cease
to flower and disappear if the shade becomes
too dense, only spectacularly to reappear after
coppicing, tree falls or felling opens up the
canopy. Conversely, although it is often found
P 14 May, Kent. The leaves are shiny green and rather
flaccid.
29/1/09 12:27:20
LADY ORCHID
just outside a wood, it rarely occurs in the full

sun of open downland. Lady Orchid also likes
shelter from the wind, and in exposed situations
the leaves and flowers may be scorched in a cold
spring, the leaves often turning yellow. It occurs
up to 200m above sea level.
223
1987-99
1970-86
pre 1970
Flowering period
Early May to early June, exceptionally from mid
to late April, varying from season to season,
but plants are generally at their best in mid
to late May.
Range
Lady Orchid is locally frequent on the
North Downs in Kent with more than 100
sites in two areas. First, the downs on either
side of the Medway Valley and sporadically
eastwards towards the Stour Valley. Second,
the downs between the eastern slope of the
Stour Valley and Dover. Elsewhere, there are
two current sites in Oxfordshire (see History
and Conservation) and it has recently been
recorded in north Hampshire. World range:
Essentially Europe, with outlying populations

in North Africa, Asia Minor, the Crimea and
the Caucasus (the last sometimes treated as
a distinct species). In Europe it occurs north
to Holland and Denmark, east to western
and southern Poland and Ukraine and south
M 14 May, Kent.There are around 100 sites in Kent, some with over 1,000 flowering plants.
29/1/09 12:27:22
224
GENUS ORChIS
to central Spain, Italy, northern Greece and

Bulgaria, also found on Corsica and Sardinia
and, in North Africa, in Algeria and Tunisia. In
the south of the range Lady Orchid is confined
to the mountains.
How to find it
This large and conspicuous species is easy to
spot when in flower, but as with all orchids,
non-flowering plants can be hard to find and
the proportion of a population in flower varies
from year to year. Especially good sites in
Kent include Yockletts Bank and it also occurs
at Wye.
DESCRIPTION
M 14 May, Kent. Although Lady Orchid is associated with

woodland, it does best where there is some sun.
Height: 20-50cm, exceptionally to 100cm.

Stem: Green, becoming purplish-brown
towards the tip.
Leaves: Green and shiny or almost greasy.
There are three to five, sometimes seven, broad,
oval to oval-oblong leaves. The lower leaves
are blunter and form a basal rosette, the upper
leaves becoming successively more pointed,
keeled and clasping. They appear above ground
between mid-January and mid-February.
Spike: Oval to oblong in shape and lax or
densely flowered. Robust plants have up to
50 flowers.
Bract: Greenish to purple, tiny, elongated and
scale-like.
Ovary: Bright green, sometimes washed purple
along the six ribs, and distinctly twisted.
Flower: Whitish, finely spotted with reddishpurple and with a contrastingly dark hood.
The sepals are oval, and the petals are shorter,
much narrower and more strap-shaped but
broaden to a spear-shaped tip; together they
form a hood (the bonnet). They are pale green,
irregularly blotched on both surfaces with
dark purple or purplish-brown, the blotches
becoming more numerous and coalescing
towards the tip, base and sides; the unopened
buds are therefore very dark. The lip is pale
pink to white, washed violet or rose around the
edges, variably spotted pink to reddish-purple
(the spots are formed by tufts of tiny papillae).
29/1/09 12:27:23
LADY ORCHID
225
hood may also vary from paler to darker shades

of purple.
Plants in west Kent differ slightly from
those in east Kent (from the eastern side of the
Stour Valley to Dover). They are, on average,
shorter (20-38cm tall rather than 30-76cm)
with a shorter, denser flower spike, shorter
ovaries (13-19mm rather than 19-25mm) and
lips that are more heavily spotted and washed
rose to purple (rather than salmon to brownishred); their lips are also blunter and less deeply
lobed. These two groups of populations have
not, however, been given names.
Var. albida lacks anthocyanin pigments and
thus has a pure white lip and a distinctive white
or straw-coloured hood with green veins. It is
scarce.
Var. pseudomilitaris has narrower and more
reddish lobes on the lip, resembling the arms
and legs of a Military Orchid. The hood is,
however, the normal reddish-purple. It is rare.
M 14 May, Kent. The spots on the lip are composed of
tufts of tiny papillae.
BIOLOGY
It is deeply lobed with two long, narrow arms

and two broad terminal lobes, often with frilled
edges, which form the ladys skirt (the latter
usually have a tiny tooth between them). The
spur is cylindrical, from a quarter to a half of
the length of the ovary, curved and pale green
blotched with purple. The column is pale green
washed pinkish or purplish, and the pollinia
are blotched purple. The flowers are variably
reported to smell of vanilla or bitter almonds or
to be unscented.
The flowers are pollinated by small flies and

bees, including small digger wasps. Seed-set
is variable, sometimes very low, with only
3-10% of flowers producing ripe capsules.
However, in some years and at some sites it
can be good. Most reproduction is by seed
although vegetative reproduction also takes
places, and clumps of plants can be formed in
this way.
Subspecies
None.

Relatively variable. Plants growing in the
open tend to be shorter and darker-flowered
than those in woods with a tendency towards
brown rather than purple markings. There is
also much variation in the shape, colour and
markings of the lip; this may be white and even
unspotted in some plants and heavily washed
pink with dark purple spots in others. The

From germination to first flowering takes eight
to ten years and the pattern of development
is typical of the genus Orchis. Plants may live
for at least another ten years, flowering at
least three times in that period but seldom
every year; the remains of the previous years
dried, dead spike is not often seen next to the
current flowers. There are therefore usually
large numbers of non-flowering plants in any
population.
Hybrids
O. x wilmsii, the hybrid with Early Purple
29/1/09 12:27:24
226
GENUS ORChIS
O 14 May, Kent. A flower

with a relatively pale hood
and fine markings.
Orchid, has been reported very rarely in Kent.

O. x macra, the hybrid with Man Orchid, was
found in east Kent in 1998.
The hybrid with Monkey Orchid has
appeared in small numbers since 2006 at the
Hartslock reserve.
Hybrids with Military Orchid have also
been reported (Summerhayes, 1968).

The specific name purpurea means simply
purple.
HISTORY aND
CONSERVaTION
First recorded by Christopher Merrett in 1666
in his Pinax Rerum Naturalium Britannicum (A
picture of British natural history), On Gadshill in Kent.
Since its first discovery, the Lady Orchid has
been largely restricted to Kent, where there are
many sites. Currently, there are also two sites
in south Oxfordshire. In the Thames Valley
area a single plant was discovered growing
29/1/09 12:27:26
LADY ORCHID
227
under Beeches in mixed woodland in 1961.

It flowered again in 1962 and 1964 but then
vanished. Seven plants were found at the same
site in 1986 (the Beeches and nearby Yews
having been felled in the late 1970s) and this
colony has subsequently flourished, with at least
46 plants and 29 flower spikes in 1997. Also in
south Oxfordshire, there have been up to three
plants (mostly non-flowering) at Hartslock
since at least 2000.
Past and present occurrence of Lady Orchid in Britain and

1500-1999
current range
Britain
Ireland
37
16 (0.6%*)
% lost, 1500-1969
46%
% lost, 1970-1986
11%
% lost, total
57%
Classified as Endangered, Lady Orchid is

Nationally Scarce and has been lost from 57%
of the historic range. It was formerly found
in Surrey, mostly prior to 1930, but the last
site was at Coulsdon, where it was present
until 1959. Also extirpated in West Sussex,
where there were at least five sites, the most
recent record being in 1976; there is an old,
unconfirmed record for East Sussex. Elsewhere,
recorded from Herefordshire in a disused
quarry in 1967, at Leigh Woods in the Avon
Gorge, Somerset, in 1990 (although this plant
quite possibly originated from Lady Orchids in
cultivation at the University of Bristol Botanic
Gardens a short distance away), and formerly
occurred in the Channel Islands. A widely
quoted 1738 record from Essex has been shown
to have involved Military Orchid.
Past declines were due to the loss of
woodlands and the cessation of coppicing. The
population in Kent is now stable and some
colonies are large (over 3,000 plants were
counted at one site in recent years) although
M 14 May, Kent. The spike is pyramidal at first but

becomes cylindrical as more flowers open.
others are declining, and it does not easily

colonise new areas. It has long been noted that
Lady Orchids are subject to the depredations of
rabbits and deer which nip off the flowers and
attack the leaves. Slugs are also attracted to the
species. Deer are now commoner in England
than at any time for a thousand years, making
this more of an issue. Feral wild boars are also a
potential problem as they can root out and eat
the tubers.
29/1/09 12:27:28
228
GENUS ORChIS
early purple orchid

Orchis mascula
As its name suggests, this is the first orchid to appear in the spring in most of Britain and
Ireland. It heralds the coming season and gladdens the heart on an early spring day, whether
as splashes of purple among a carpet of Bluebells or scattered among Cowslips and violets on
a roadside bank. One of the commonest and most widespread orchids, Early Purple Orchid
is nevertheless very local in many areas and has largely vanished from farmland habitats in the
lowlands where it is now confined to woods, roadsides, churchyards and nature reserves.
Identification
Early-flowering, purple flowers and spotted
leaves are a distinctive combination. Unspotted
leaves are not uncommon, however, and there is
a scarce white-flowered variant; these could be
more problematic but a look at the structure of
the flower should prevent any confusion.
Similar species
Green-winged Orchid is easily separated by the
parallel green stripes on its sepals. The hood of
its flower is formed by all the sepals and petals,
so that it lacks the erect wings of Early Purple
Orchid. In addition, Green-winged Orchid
always has unspotted leaves and usually also a
smaller, fewer-flowered spike, and it does not
grow in woodland.
Marsh orchids have flowers of various
shades of purple and some have spotted leaves,
but the flowers are usually more extensively and
more heavily marked with black dots, lines and
squiggles. Their spur is short and often sacklike and is either straight or curves downwards,
whereas it curves slightly upwards in Early
Purple Orchid. The spots on the leaves tend to
be regular and often elongated sideways rather
than irregular and often elongated lengthwise as
in Early Purple Orchid.
Habitat
M 23 April, Norfolk. In most places this is the first orchid to

bloom, with rich purple flowers; wellnamed indeed.
Very variable. It can occur in both grassland

and woodland and on a variety of soils,
although it does have a definite preference
for calcareous soils on chalk, limestone or
boulder clay and avoids acid conditions. It
is found in a wide variety of old grasslands,
29/1/09 12:27:29
EARLY PURPLE ORCHID
both dry chalk downland and damp hill

pastures, as well as meadows, rocky mountain
ledges, railway embankments and cuttings,
road verges, grass-covered dry-stone walls
and limestone pavements. It also grows in
deciduous woodland, usually in the better-lit
areas along rides, tracks and woodland edges.
Early Purple Orchid is particularly associated
with coppice woodland, where there is usually a
great increase in the number of flowering plants
in the second or third year after coppicing but
a decline thereafter as the canopy closes again.
The species does not colonise new sites easily
and is usually found in ancient woodland rather
than relatively recently established plantations
or secondary woodland (unless, of course,
woodland and scrub have invaded old orchidrich grassland). Conversely, many colonies on
road verges and banks may be relicts of longgone woods. Recorded up to 880m above sea
level (Caenlochan, Angus).
Flowering period
Early April to early June in the south,
exceptionally from mid-March, although
229
most are in flower from late April to late May.

Flowering is on average a little later in upland
areas and in Scotland, the season occasionally
lasts until early July. Plants in sunny, sheltered
spots flower earliest, whereas those in cool,
shaded, wet areas will be last. Once it has
set seed, the tall spikes with numerous dark
purplish-brown capsules are conspicuous well
into the summer.
Range
Found throughout the British Isles, including
the Channel Islands, Isle of Man, Hebrides,
Orkney and Shetland. Although generally
fairly common or even abundant, it may be very
local in areas of acid soils and is largely absent
from some regions, such as the Fens, south
Lancashire and mid-west Wales. It is also very
scattered in the Borders, northeast Scotland,
Orkney, Shetland, the Outer Hebrides and
southeast Ireland. World range: Western
Europe, extending to North Africa and Asia
Minor. Occurs north to the Faeroe Islands, c.
70N in Norway, central Sweden and the Baltic
States. The southern and eastern limits are
M 24 May, Co. Clare. Early Purple Orchid is very conspicuous in The Burren and sometimes forms beautiful rock
gardens with Primroses and other flowers.
29/1/09 12:27:31
230
GENUS ORChIS
29/1/09 12:27:35
EARLY PURPLE ORCHID
1987-99
1970-86
pre 1970
poorly known due to confusion with closelyrelated species, but it is found south and east
to Italy, Greece, Turkey and Bulgaria and is
probably absent from much of Central Europe.
Also present in the Canary Islands, Balearics,
Corsica, Sicily and Malta, and Tunisia in
North Africa.
231
Spike: Oval or cylindrical but often rather

irregular in shape and rather open, especially
in the lower half, with ten to 50 flowers,
occasionally more.
Bract: Green, usually strongly washed purple.
The bracts are lanceolate and about as long as
the ovary, which they clasp.
Ovary: Green, variably washed purple and
clearly ribbed.
Flower: Various shades of purple, occasionally
pale rose-pink or white. The sepals are
lanceolate, the lateral sepals asymmetrical, and
the petals are smaller and more arrow-shaped.
The upper sepal and petals form a hood over
the column and the two lateral sepals are held
erect and pushed backwards, resembling angels
wings (they may almost touch at the rear).
The lip points downwards and outwards and
is three-lobed with the central lobe usually the
How to find it
Generally common and easy to find. In some
areas, such as the Peak District, Yorkshire
Dales and The Burren in western Ireland,
Early Purple Orchid occurs in large, extensive
colonies but it is more usually found in small
and often scattered groups.
DESCRIPTION
Stem: Stout, pale green, angled and usually
flushed purple towards the tip.
Leaves: The three to eight basal leaves are
variably oblong-lanceolate in shape and often
blunt-tipped. They are glossy green, usually
marked with large, irregular, rounded or
elongated blackish-purple spots on the upper
surfaces and rarely also on the undersides. They
are held close to the ground, either spreading
upwards and outwards or in a flatter rosette.
There are two or three rather smaller and more
pointed sheathing leaves higher on the stem and
these may have a few spots or a purple wash.
M 2 May, Norfolk. The shape of the lip and, to a lesser

extent, the colour of the flower, is variable.
O 2 May, Norfolk.
29/1/09 12:27:37
232
GENUS ORChIS
largest. The side-lobes are folded downwards,

and the entire lip is sometimes folded or
creased along its centre line. The terminal lobe
is shallowly notched and the edges of all three
lobes are crinkled (crenate). The base of the
lip and mouth of the spur are much paler and
whiter, sometimes with some yellow tones,
and usually spotted with purple, the spots
composed of dense tufts of short papillae. The
spur is long and narrow, at least as long as the
ovary, broadens a little towards a blunt tip
and curves upwards. The column is variably
greenish or purple, and the pollinia are dark
green. The flowers are initially sweet-smelling,
recalling honey or Lily-of-the-valley, but this
quickly changes to a rank smell, usually said
to be like tomcats urine and to be especially
pungent at night.
Subspecies
None recognised. In the Outer Hebrides, far
north of mainland Scotland, Shetland and a
few places in western Ireland plants are just
5-10cm tall and flower from early June to early
July. These populations are sometimes named
O. m. ebudium.

As in many orchids, plants growing in the open
are often shorter and stockier than those in
woodland. These are sometimes referred to as
M 25 May, Co. Clare.The white flowered variety alba has

an unmarked lip and yellow pollinia. It is scarce.
the grassland and woodland forms.

Var. alba lacks anthocyanin pigments and has
white, unmarked flowers with yellow pollinia,
an entirely green stem and bracts, and no leaf
spots. It is rather scarce. An even scarcer variant
has white flowers which retain purple spots at
the base of the lip and sometimes has spotted
leaves too. A broken-coloured variant has also
been recorded very rarely and this has a pale
pink lip, copiously flecked with tiny purplish
markings.
BIOLOGY
M 12 May, Norfolk.The long spur contains no nectar; the
bright colours of the flower are therefore a deceit to lure
potential pollinators.
The flowers are pollinated by bumblebees and

to a lesser extent cuckoobees and a variety
of solitary bees. A visiting insect touches the
29/1/09 12:27:40
EARLY PURPLE ORCHID
233
column and the pollinia are stuck to its head

by the sticky viscidia. Once the bee has left
the flower, the pollinia on the insect rotate
forward in about 30 seconds so that they will
make contact with the stigma in the next flower
visited. Early Purple Orchid is self-compatible
and is sometimes self-pollinated.
The flowers of Early Purple Orchid have
no nectar and offer no reward to a pollinator,
although it has been suggested that a sugary sap
is produced inside the wall of the spur. Rather,
it is thought that the orchid takes advantage of
the naivety of the bees which, newly emerged
from hibernation in the spring, have yet to learn
which flowers are genuine sources of nectar.
These naive bees are attracted, at least for a
while, by the bright colours and scent of the
flowers. The various species of bee emerge from
hibernation at different times, and therefore
the orchid can take advantage of a succession of
pollinators as the season progresses. Seed-set
is variable, with the lowest, earliest-opening
flowers most likely to be pollinated. Vegetative
reproduction may occur occasionally via the
production of additional tubers.

The first aerial leaves are usually produced in
the fourth year after germination, and more
and bigger leaves appear in successive seasons
until enough reserves have been accumulated
to produce a flower spike; up to eight years
may elapse between the first appearance above
ground and the first flowers. Around 60%
of plants will flower in successive years; the
remainder either appear as a rosette of leaves
or, in about 17% of cases, spend a year dormant
underground before appearing again. Up to
12 years dormancy has been noted in Europe.
A maximum lifespan of 13 years after the first
appearance of leaves has been recorded.
Hybrids
O. x wilmsii, the hybrid with Lady Orchid, has
been reported very rarely in Kent.
P 22 May, Co. Clare.The influence of the different flower
colours on potential pollinators is unknown.
29/1/09 12:27:42
234
GENUS ORChIS

The specific name mascula means male and may
be a reference to its vigour, early flowering or is
perhaps another allusion to the masculinity of
the tubers.
HISTORY aND
CONSERVaTION
The first British record dates from as long ago
as 1562 when William Turner noted in his
Herball, There are divers kindes of orchis
one kinde hath many spottes in the leafe and
is called adder grasse in Northumberland.
Locally common in many areas, Early
Purple Orchid is the third most widespread
species of orchid in the British Isles (after
Common Spotted and Heath Spotted
Orchids). However, it has vanished from 28%
of its historical range in Britain and 21% in
Ireland, and the decline appears to be ongoing
in Britain. Losses are due to the destruction
or coniferisation of woodland and, perhaps
more importantly in recent years, the loss of
permanent grasslands as pastures and meadows
have been ploughed and reseeded. In most of
lowland Britain, away from reserves, the species
is now largely confined to ancient woodland
and to marginal sites, such as road verges and
churchyards, which have escaped agricultural
improvement. In the north and west
overgrazing may be a problem as the species
is tolerant of light grazing only.
Past and present occurrence of Early Purple Orchid in
Atlas).

1500-1999
Britain
Ireland
1,971
475
1,416 (50%*)
377 (37%)
M 24 April, Norfolk. Hazel coppice is the classic habitat.
current range
Intergeneric hybrids
% lost, 1500-1969
17%
17%
O. x morioides, the hybrid with Greenwinged Orchid, has been recorded rarely and
sporadically in England and Wales. As Greenwinged Orchid is no longer in the genus Orchis,
a new name is needed for this hybrid.
% lost, 1970-1986
11%
4%
% lost, total
28%
21%
29/1/09 12:27:43
The single species in this genus is

found in Europe and Greenland,
with just a toe-hold in North
America, in eastern Canada.
Taxonomy
Sometimes Pseudorchis is united
with the fragrant orchids in the
genus Gymnadenia. It differs in
having a shorter spur, all three
sepals arranged with the petals to
form the hood of the flower (not
just the upper sepal) and a tuber
that is so deeply divided that it may
appear to be several cylindrical
tubers.
Name
The generic name Pseudorchis
derives from the Greek pseud
meaning false and is a reference to
its relationship to the genus Orchis.
GenuS PSEUDORCHIS
Small white Orchid
29/1/09 12:31:31
236
GENUS PSEUDORCHIS
Small white Orchid
Vulnerable, BAP
Pseudorchis albida
Other names: Leucorchis albida, Gymnadenia albida; in North America Pseudorchis straminea
Newfoundland Orchid
This delicate and unassuming orchid is a boreal species whose generally northern distribution
worldwide is reflected in Britain and Ireland. In this respect it is similar to Lesser Twayblade
and Creeping Ladys-tresses. Like many other northern plants, it has declined drastically in
the southern parts of its range and disappeared from much of England, Wales and Ireland.
Identification
The combination of its small stature, dense
spike of creamy-white flowers and deeply threelobed lip is distinctive. The individual flowers
are bell-shaped and very small, just 2-4mm
across (smaller even than the ovary). The spike
is carried on a long stem with a cluster of shiny
green leaves at the base.
Similar species
Creeping Ladys-tresses is superficially similar
but flowers later in the year, on short, dry
turf (although occasionally on moorland). Its
flowers also have glandular hairs but its lip is
rather different, being spout-shaped rather than
three-lobed.
Irish Ladys-tresses flowers even later in
the summer than Creeping Ladys-tresses and
barely overlaps with Small White Orchid. It
has bigger flowers arranged in three columns
around the spike. The lip is also formed into a
spout.
Dense-flowered Orchid is superficially
similar to Small White Orchid, and the
flowering periods may just overlap, but the lip is
very different and the hood is tightly closed.
White-flowered varieties of Pyramidal,
spotted and fragrant orchids have been
mistaken for Small White Orchid, but their
flowers are larger and differ in many other
details.
M 6 June, Cumbria. Despite its white flowers, this petite
orchid can be hard to see in long grass.
O 6 June, Cumbria.
Habitat
Small White Orchid grows in rough grassland
on poor, well-drained soils, both mildly acidic
and base-rich. It is found on hill pastures, hay
29/1/09 12:31:32
SMALL WHITE ORCHID
237
meadows, road verges, banks, streamsides and

grassy ledges. It will grow in the partial shade
of shrubs and bushes and is sometimes found
on recently burnt moorland among short
Heather (disappearing again once the Heather
regenerates to form a closed community).
Very rarely, the species has been found in oak
woodland on acid soils and on stabilised coastal
dunes. Generally it occurs in the uplands at over
165m above sea level and has been recorded as
high as 500m at Alston Moor in Cumbria and
550m at Ben Chaisteil in Argyll, but it is also
found near sea level in western Scotland.
Flowering period
Late May to mid-July, depending on altitude
and latitude. It is latest at higher altitudes in the
north and earliest in Ireland but is generally at
its best around mid-June. The flowers tend to
wither quickly.
Range
Small White Orchid is commonest and most
widespread in northern and western Scotland,
including the Inner Hebrides and Orkney. It is
now almost absent from the central lowlands
and southern uplands of Scotland, merely
clinging on at a few isolated sites in Dumfriesshire, Ayrshire and Roxburghshire. The
species is similarly much reduced in northern
England and is now found rather rarely in
1987-99
1970-86
pre 1970
M 7 June, Perth and Kinross.The sepals are whiter than the

petals and lip, which tend to be washed greenish-yellow.
29/1/09 12:31:34
238
GENUS PSEUDORCHIS
over the Urals into northwest Siberia). In

the south it is found in the Pyrenees, Massif
Central, Corsica, Alps, Apennines, Balkans,
Carpathians and Transylvanian Alps east
to Ukraine. Between and around these two
centres of distribution there are scattered
records, e.g. in Bulgaria, northern Greece and
recently from Crete. But, as in the British Isles,
the species has vanished from many lowland
areas and is extinct in Belgium, Luxemburg
and Holland. It is also found in southern
Greenland and in North America, where
it is restricted to a small area in northwest
Newfoundland and western Quebec.
How to find it
Small and inconspicuous, this orchid can be
hard to spot, especially as it is often found
singly or in small numbers. Sites in Scotland
include Keltneyburn (Perth & Kinross), Glen
Cova (Angus) and Feoch Meadows (Ayrshire).
DESCRIPTION
M 6 June, Cumbria. The three-lobed lip can only be seen
on close inspection.
Cumbria, Northumberland, Co. Durham

and mid-west and northwest Yorkshire. In
Wales it is found at a few scattered sites in
Breconshire, Merionethshire, Denbighshire
and Caernarvonshire. In Ireland it is similarly
very scattered, with odd sites in Co. Kerry,
Co. Limerick, Co. Clare, Co. Cavan and Co.
Donegal. There are small clusters of sites in
northern Co. Tipperary and central Galway but
the main concentration lies in the northwest,
in Co. Sligo, Co. Leitrim and Co. Fermanagh.
Elsewhere there is another cluster of sites in
Co. Tyrone, Co. Derry and Co. Antrim in
Northern Ireland. World range: Small White
Orchid has an amphi-Atlantic distribution
and is found in Europe and northeast North
America. In Europe it occurs in two discrete
areas. To the north it is found in Iceland
(where it is abundant), the Faeroe Islands,
Denmark, Scandinavia (north to northernmost
Norway) and northern Russia (just creeping
Height: 8-40cm but usually less than 20cm and

only very exceptionally to 40cm.
Stem: Greenish, slightly angled towards the tip,
with two or three whitish or brownish sheaths
at the base.
Leaves: There are four to six shiny, green, oval
to oval-lanceolate, keeled sheathing leaves at the
base of the stem and one or two narrower and
more bract-like leaves above them.
Spike: Dense, cylindrical and often rather onesided, with 20-40 flowers (exceptionally as few
as ten or as many as 70).
Bract: Green, lanceolate with a pointed tip, and
as long as or just longer than the ovary, which
it clasps.
Ovary: Green, slightly twisted, with three
obvious ridges. The ovary is strongly curled over
towards the tip so that the flower faces moreor-less downwards.
Flower: The flowers are small (rather smaller
than the ovary and bracts) and very pale, with
the sepals whitish or creamy and the petals, lip
and spur washed more greenish or yellowish.
The sepals are elliptical and blunt-tipped and
29/1/09 12:31:36
SMALL WHITE ORCHID
form a loose hood that encloses the similarly

shaped petals and the column. The lip is short,
broader than long and deeply three-lobed. The
central lobe is triangular, usually longer, wider
and blunter than the side-lobes, which are
narrower and more lanceolate in shape. The
spur is short (2-3mm), tubular or sack-shaped,
blunt-tipped and down-curved and contains
abundant nectar. The flowers have a delicate
scent of vanilla.
239
taper gradually to a long, pointed tip and are

often deeply divided into several long fingers
that diverge widely. There are also long, fleshy
roots that lie horizontally close to the surface
of the soil. The roots and the slender tips of the
tubers have a heavy fungal infection. The first
aerial stem is reported to appear four years after
germination.
Subspecies
Small White Orchid is divided into two
subspecies. The nominate subspecies P. a. albida
is found in the British Isles, central Europe and
lowland Scandinavia. Subspecies straminea,
with larger and yellower flowers, is found in the
mountains of central Europe and Scandinavia,
Faeroe Islands, Iceland, Greenland and North
America. The latter is often treated as a distinct
species, Pseudorchis straminea.

British plants belong to the nominate
subspecies, P. a. albida, which in turn is
divided into two varieties.
Var. albida is found on more acid soils and has
the lateral lobes of the lip clearly shorter than
the central lobe.
Var. tricuspis favours calcareous soils and has
the lateral lobes almost as long as the central
lobe. The distribution and abundance of the
two varieties in the British Isles has not been
studied.
BIOLOGY
The flowers produce nectar and are visited by
butterflies, day-flying moths and solitary bees.
The specific pollinator has not been identified,
but the narrow entrance to the spur suggests
that it may be butterflies. Some self-pollination
also occurs, as the pollinia eventually fall onto
the stigma if an insect has not removed them.
Seed may be set by over 90% of flowers.

The aerial stem grows from paired tubers that
M 6 June, Cumbria.The flowers are small smaller than

the ovary. The specific pollinatior is unknown but seed-set
is good.
29/1/09 12:31:37
240
GENUS PSEUDORCHIS
X Pseudadenia schweinfurthii, the hybrid with
fragrant orchid (presumably Heath Fragrant
Orchid), has been recorded from several places
in Yorkshire and Scotland, where it is fairly
frequent in the northwest.
X Pseudorhiza bruniana, the hybrid with
Heath Spotted Orchid, was recorded from
Orkney in 1977 and Skye in 1994.

The specific name albida means white.
HISTORY AND
CONSERVATION
The first British record was in 1670 when it was
discovered by John Ray on Mount Snowdon
in North Wales: This we found on the back
of Snowdon-hill by the way leading from
Llanberis to Carnarvan (Catalogus Plantarum
Anglicum et Insularum adjecentium).
Classified in Britain as Vulnerable, the
species is specially protected in Northern
Ireland under Schedule 8 of the 1985 Wildlife
Order (NI) and in Eire under the Flora
(Protection) Order.
Past and present occurrence of Small-white Orchid in Britain and Ireland (based on presence or absence in 10km
Britain
Ireland
385
110
132 (4.6%*)
33 (3.3%*)
% lost, 1500-1969
52%
63.5%
% lost, 1970-1986
13.5%
6.5%
% lost, total
65.5%
70%

1500-1999
current range
Small White Orchid is commonest in

northern and western Scotland and becomes
rarer further south; in the remainder of
Scotland it has almost vanished from the area
south of a line from the Clyde to Aberdeen
and has not been recorded recently from
Kirkcudbrightshire, Lanarkshire, Selkirkshire,
Berwickshire, the Lothians, Fife and

Stirlingshire. It is extinct in Shetland, where
there has only been one record, from Bressay
prior to 1845.
In England, Small White Orchid is now
rare even in Cumbria, a former stronghold that
still holds the bulk of the English populations,
although most colonies comprise just a handful
of plants. It is now extinct in Kent (with just
one old record, from near Lyminge), East
and West Sussex (where present until at least
1913, with three localities in West Sussex
and ten in East Sussex, notably Ashdown
Forest), Gloucestershire (last recorded 1899),
Herefordshire, Staffordshire, Shropshire
(last recorded 1856), Derbyshire, Cheshire,
Lancashire, southwest and northeast Yorkshire
(where it has recently gone from the North
York Moors National Park). The situation
in Wales is probably even worse, as numbers
there have collapsed in recent years, with no
records from Monmouthshire, Glamorganshire,
Carmarthenshire, Cardiganshire, Montgomeryshire and Flintshire. In total, Small White
Orchid has been lost from 66% of the historical
range in Britain. In Ireland there has also been
a very sharp drop in numbers. It has been
lost from 70% of the historical range and is
extinct in Co. Cork, Co. Waterford, Laois,
Co. Wicklow, Co. Dublin, Co. Mayo, Co.
Monaghan and Co. Down.
Populations tend to be small and scattered
and even in Scotland many vice-counties have
just one or two sites for the species. Colonies
were lost throughout the 20th century due
to habitat destruction, forestry, agricultural
improvement and overgrazing. As with many
grassland orchids it requires both a short
sward and areas of bare soil so that seedlings
can become established. It therefore needs a
certain level of grazing to thrive; too little and
it is swamped by coarse grasses and scrub, too
much and it can never flower and is eventually
eliminated.
P 14 June, Kent. Greater Butterfly Orchid.
29/1/09 12:31:38
GENUS
PLATANTHERA
BUTTERFLY
ORCHIDS
12/2/09 17:01:41
242
GENUS PLATANTHERA
Distribution
There are 80-200 species in this genus. They are
scattered almost worldwide, although most are
found in the temperate regions of the Northern
Hemisphere, with eight in Europe and two in
the British Isles.
Floral structures
The column is short and there are two pollinia.
They taper into slender caudicles (stalks), each
attached to one of the two viscidia, which are
naked (there is no pouch-like bursicle). There is
a single flat stigmatic zone.

in the autumn.
Name
The generic name Platanthera derives from
the Greek plat meaning broad, wide or flat
and antherus meaning flowery or anther. It is
usually taken to mean flat-anthers, a reference
to the shape of the anther, which has a hollow
at its base to hold the pollinia.
column
Growth pattern
The aerial stem grows from a pair of spindleshaped tubers which taper to a long, narrow
point, and there are also a few slender roots
that spread into the surface layers of the soil. As
with the genus Orchis, one of the tubers forms
the summer before and has supplied the current
years growth, while a new tuber, complete
with a bud ready for next years flower spike,
develops beside it.
Fungal partners
All the roots are infected with fungi.

Seed probably germinates in the spring, usually
not far below the surface of the soil. The
protocorm produces a large bud in the first
autumn and this develops into a leafy shoot
the following spring. Following the emergence
of this shoot, a root is produced. These die off,
presumably in the late summer, and the seedling
overwinters as a short, two-segmented rhizome.
Similarly, a leafy shoot and one or two roots
are produced the following summer, and it is
not until the third year that the first tuber is
produced and the protocorm finally vanishes.
A full-sized tuber appears after the fourth
summer. (This chronology is based on work
in the 1920s and, as with many other orchids,
development may be rather faster.)
pollinium
stigma
mouth of the spur
viscidium
lip
column
pollinium
stigma
viscidium
mouth of
the spur
lip
of the aerial stem. These will go on to form
M Columns of butterfly orchids showing position of

pollinia (sepals and petals removed).
29/1/09 12:31:41
LESSER BUTTERFLY ORCHID
leSSer Butterfly Orchid
243
Vulnerable, BAP
Platanthera bifolia
The more delicate and daintier of the two butterfly orchids, this species has a northerly
and westerly bias to its distribution and has vanished from much of central and eastern
England. It is found on rough pastures, damp heathland, bogs and, rather less frequently, in
woodland. The delicate flowers are highly scented at night in order to attract hawkmoths;
as the moth sips nectar from the orchids long spur the pollinia are glued to its proboscis and
it becomes the unwitting servant of the orchid.
Identification
The two butterfly orchids are distinctive. They
have two oval, shiny leaves, placed at the base
of the stem and often hidden away in the grass,
and exquisite white flowers. The lip of the
flower is long, narrow and undivided, and the
spur is extremely long and slender and projects
prominently to the rear. There are two subtly
different forms of Lesser Butterfly Orchid,
heathland and woodland (see Variation and
varieties).
Similar species
Greater Butterfly Orchid can easily be separated
by the size and shape of the pollinia. In Lesser
Butterfly Orchid the two pollinia are placed
close together and parallel, while in Greater
Butterfly Orchid the bases of the pollinia are
well separated and they lean inwards towards
the tip. Other differences between the two
species are subtler and less consistent. Lesser
Butterfly Orchid is generally smaller and
daintier (averaging around two-thirds the size
in most dimensions), with fewer, smaller flowers
in a narrower spike; the mouth of the spur is
smaller and the spur tends to be straighter.
Habitat
The heathland form of Lesser Butterfly Orchid
is by far the commoner and grows on heathland
in the south and east and on moorland and
damp pastures in the north and west. On both
heathland and moorland it usually occurs in
the damper areas, and these are frequently
P 19 June, Norfolk. A robust plant with numerous flowers in a compact spike.
29/1/09 12:31:43
244
GENUS PLATANTHERA
open deciduous woodland and scrub, often

beechwoods, on calcareous soils, especially
clay-with-flints. It can also occur on open chalk
downland and among Bracken on the less acid
areas of dry grassy heaths. Lesser Butterfly
Orchid occurs up to 365m above sea level
(Glenfeshie, Inverness-shire).
Flowering period
Late May and June in woodland populations,
June to July for heathland forms. On average it
flowers latest in the north of Scotland.
Range
Lesser Butterfly Orchid occurs throughout
Britain and Ireland, including the Isle of Man,
Inner and Outer Hebrides and Orkney. There
is, however, a distinct western and northern
bias to the distribution, and the species was
always scattered and local in much of England,
eastern Scotland and in southern and southeast
Ireland. It is now extinct in most of East Anglia,
the Midlands and northern England, although
it is still widespread in Cumbria. World range:
Widespread in Europe and Asia and also found
in North Africa. In Europe it occurs north
to the Faeroe Islands and c.70N in northern
Scandinavia. It ranges south to Spain, Italy,
northern Greece, the Crimea and Caucasus.
It is found on the Balearic Islands, Corsica,
Sardinia, Sicily and the Aegean islands. In
1987-99
1970-86
pre 1970
M 20 June, Norfolk. A rather small, few-flowered plant in

the early morning dew.
P 19 June, Norfolk.
marked out by the dusty grey foliage of Crossleaved Heath. It can often be found around
the margins of valley bogs or growing on drier
tussocks and other raised areas amidst sodden,
boggy ground. It grows on neutral or mildly
acidic soils; on moorland and around bogs it
favours the areas where springs and flushes
buffer the general acidity to some extent. The
woodland form is much scarcer and largely
restricted to southern England. It is found in
29/1/09 12:31:45
29/1/09 12:31:48
246
GENUS PLATANTHERA
North Africa it is found in Algeria and Tunisia.

In Asia it occurs in northwest and northeast
Turkey and northern Iran. The range extends
across southern Siberia to at least Lake Baikal.
How to find it
Often rather small and slender, this can be a
hard plant to find wherever the vegetation is tall.
However, in other areas it can be very obvious,
especially when found in large numbers.
DESCRIPTION
Stem: Pale green, more-or-less triangular
and ribbed towards the tip, with two or three
whitish or brownish sheaths at the extreme base.
Leaves: The two pale green, slightly shiny or
greasy looking leaves are held opposite each
other, one just above the other, at the base of
the stem. They are variable in shape, from oval
to narrower and more strap-shaped, and taper
at the base to a whitish, winged stalk. There
are also one to five small, lanceolate, bract-like
Spike: Variable, the five to 25 flowers
(occasionally more) form a compact cylindrical
spike in moorland plants but are more widely
spaced in woodland plants.
Bract: Green, narrow, pointed and slightly

shorter than the ovary.
Ovary: Pale green, long, narrow, clearly ribbed
and twisted and also curled into a C-shape to
hold the flowers pointing outwards and slightly
downwards.
Flower: Whitish, with a long, strap-shaped
lip and extremely long spur. The sepals are
white, washed greenish towards the tip and
bluntly lanceolate in shape. The upper sepal
is slightly broader and more triangular and
may be bent upwards at the tip. The petals are
creamy and variably washed greenish. They are
smaller, narrower and more strap-shaped than
the sepals. The upper sepal and petals form a
loose hood over the column. The lateral sepals
are held spreading and slightly drooped. The
lip is creamy, sometimes greener towards the
tip, narrow, strap-shaped and 6-12mm long. It
projects forwards and downwards. The spur is
long and slender (1mm wide x 13-23mm long,
occasionally to 27mm), sometimes slightly
curved and it may be washed with green.
The pollinia are whitish, about 2mm tall and
lie parallel, 1mm apart, at the front of the
column. The flowers emit a heavy, sweet scent,
sometimes likened to carnations, especially
at night.
Subspecies
None.
M 20 June, Norfolk.The pollinia lie parallel to each other,

above the mouth of the spur.
In both Greater and Lesser Butterfly Orchids

plants growing in the open tend to be shorter
and more compact than those in shaded
woodland localities. In Lesser Butterfly Orchid
the heathland form has leaves that are eggshaped and a relatively dense flower spike
whereas the woodland form has narrower
and less pointed, more tongue-shaped leaves.
The heathland form is found on acid soils
throughout Britain and Ireland whereas the
woodland form is most frequent in southern
England, often growing on calcareous soils,
and becomes rare to the north, although it has
been found in southern Scotland. Intermediates
occur and the differences between the two
29/1/09 12:31:49
LESSER BUTTERFLY ORCHID
247
forms may be caused solely by the different

environments in which the plants are growing.
Var. trifolia has three main leaves rather than
two. It is not rare.
Var. quadrifolia has four leaves.
BIOLOGY
The flowers are pollinated by insects attracted
by the copious nectar in the spur (nectar can
be seen filling the tip of the spur, which is
translucent). However, the nectar can only
be reached by an insect with a suitably long
proboscis and Lesser Butterfly Orchid is
pollinated by night-flying moths, especially
hawkmoths such as Elephant, Small Elephant
and Pine Hawkmoths. The flowers scent,
which is particularly pungent around dusk,
and white coloration (they almost glow
in the dark) help the moths to find them.
Hawkmoths hover in front of the flower to
feed, resting their forelegs on the lateral sepals,
whereas other moths land on the flower itself.
The two pollinia, which are relatively small,
extremely short-stalked and placed parallel
to each other at the mouth of the spur, have
small sticky pads, the viscidia, at their base. As
the moth inserts its proboscis into the spur,
the viscidia glue the pollinia to it. The moth
continues on its way, visiting other flowers,
and after a short while the pollinia rotate
forwards and in this new position will make
contact with the stigma in the next flower
visited. The mechanism is fairly effective and
seed-set is moderate to good.

Grows from a pair of underground tubers.
There is no information on the period between
germination and flowering in the wild but in
cultivation plants may flower in three or
four years.
Hybrids
P. x hybrida, the hybrid with Greater Butterfly
Orchid, has been reported from scattered
localities but is rare; hybrids are intermediate
in the positioning of the pollinia and in spur
M 19 June, Norfolk. The flowers are pollinated by moths,

especially hawkmoths, and a long proboscis is required to
access the nectar in the spur, visible here.
length. The differences in habitat, flowering time

and flower structure make hybridisation less
likely than it seems; in Lesser Butterfly Orchid
the pollinia are attached to the proboscis of
a visiting moth, whereas in Greater Butterfly
Orchid they are typically attached to the eyes.
In these different positions the chances of the
pollinia making contact with the stigma of the
wrong species are greatly reduced. Hybrids
have the pollinia in an intermediate position,
reducing the effectiveness of the pollination
mechanism and making such hybrids less likely
to reproduce successfully.
29/1/09 12:31:50
248
GENUS PLATANTHERA
Past and present occurrence of Lesser Butterfly Orchid

Atlas).

1500-1999
current range
Britain
Ireland
950
308
342 (12%*)
159 (16%*)
% lost, 1500-1969
48%
36%
% lost, 1970-1986
16%
12.5%
% lost, total
64%
48.5%
M 13 July, Norfolk.The parallel pollinia are clearly visible.

The specific name bifolia means two leaves. The
two butterfly orchids, although easily separated
in the field, are extremely close genetically.
This suggests that they have only very recently
separated into two species.
HISTORY AND
CONSERVATION
The first reference to the Lesser Butterfly
Orchid was in the second edition of John Rays
Synopsis Methodica Stirpium Britannicarum
(A methodical synopsis of British plants),
published in 1696, which simply noted its
presence i n pastures.
Lesser Butterfly Orchid has declined
greatly, with a loss of 64% of the historical
range in Britain and 48.5% in Ireland. It is

classified as Vulnerable and is one of the most
rapidly declining wild flowers in the British
Isles. It has vanished from much of southern
and eastern England and is now extinct in
Essex, Suffolk, Hertfordshire, Middlesex,
Berkshire, Northamptonshire, Worcestershire,
Lincolnshire, Nottinghamshire, Derbyshire,
Cheshire, Denbighshire, Roxburghshire and
the Lothians. It is also seriously reduced in
many other areas. For example, in Norfolk it
was locally common on wet heaths in 1914 and
fairly frequent on bog and wet heath, always on
acid soils in 1968, when 12 localities were listed,
but by 2008 was present at only two sites, with
just a handful of plants at each. It has now gone
altogether from some classic heathland area in
southern England such as Woolmer Forest in
Hampshire and Ashdown Forest in Sussex.
Some losses are due to the outright
destruction of heathland, with both agriculture
and urban development being responsible. Even
where heathland remains, the losses continue,
probably due to the lack of grazing and resulting
transformation of heathland into scrub and
woodland. Notably, Lesser Butterfly Orchid is
still common in the New Forest, which continues
to support large numbers of grazing animals.
Away from heathland sites the improvement of
pastures and hay meadows and the clearance or
coniferisation of woodland are responsible for the
decline. Paradoxically, overgrazing is a problem in
the uplands of western and northern Britain.
29/1/09 12:31:51
GREATER BUTTERFLY ORCHID
249

Platanthera chlorantha
Near Threatened
Found locally throughout mainland Britain and Ireland but with a distinctly southern
bias, this species favours both grassland and woodland. The exquisite white flowers, often
held on a tall, stately spike, have evolved to glow in the dark as the pollinators are nightflying moths.
Identification
The two butterfly orchids are distinctive, with
a pair of oval, shiny green leaves at the base of
the stem and an open spike of beautiful waxy
white or greenish-white flowers. The lip is long,
narrow and undivided, and the extremely
long slender spur projects backwards from
the rear of the flower across the width of the
flower spike.
Similar species
Lesser Butterfly Orchid is distinguished by
the shape and position of its pollinia, which lie
close together and are parallel for their entire
length. In Greater Butterfly Orchid the pollinia
are well-separated at the base but lean inwards
so that their tips almost touch. There are other,
subtler differences between the two species
but none of these can be taken as diagnostic.
Greater Butterfly Orchid is on average taller
and sturdier, with a broader flower spike (the
ovaries are longer, holding the flowers further
away from the stem). It has greener flowers with
a larger and more obvious mouth to the spur,
and the spur itself is usually slightly expanded
at the tip.
Habitat
Rather variable but it almost always grows on
calcareous soils: chalk, limestone and baserich clays. It is found in deciduous woodland
(where it is strongly associated with ancient
woodland) and has a preference for hazel
coppice. It grows in light, dappled shade
and is usually found in the more open areas
around the edge of a wood and in clearings and
P 14 June, Kent. In woodland the spike is often very
attenuated.
29/1/09 12:31:53
250
GENUS PLATANTHERA
populations may be relicts of deciduous

woodland previously on the site.
In the north and west of Britain and Ireland
Greater Butterfly Orchid is most frequently
found in grassland such as old pastures and
hay meadows, again usually on calcareous
soils; it has been found on rare occasions on
mildly acidic soils on moorland, wet heath
and pastures but is clearly less tolerant of acid
conditions than Lesser Butterfly Orchid. It was
probably frequent in pastures and meadows in
southern England prior to their improvement
but in the south, away from woodland sites, is
now only likely to be found on chalk grassland,
often in long grass with variable amounts of
scrub. It is sometimes also found in calciumrich dune slacks and on railway embankments.
Occurs up to 460m above sea level (Harwood
Dale, Northumberland).
Flowering period
Late May to late July, being earliest in the south,
where it is sometimes in flower from mid, or
even early, May. Usually it is at its best in early
June in much of England but it may not be
in full flower in parts of Scotland until July.
Typically it flowers one or two weeks earlier
than Lesser Butterfly Orchid.
1987-99
1970-86
pre 1970
M 15 June, Kent. Greater Butterfly Orchid will grow on

open grassland as well as in woodland. This plant has
three leaves rather than the usual two.
along rides. It is tolerant of both dry and wet

conditions but in woodland on chalk may have
some preference for the heavier and wetter soils
found at the foot of slopes. Greater Butterfly
Orchid has occasionally been found growing
in rides through conifer plantations but such
Range
Found throughout Britain and Ireland except
Orkney and Shetland, although rather scattered
29/1/09 12:31:55
in the southern half of Ireland and absent

from large areas of the Midlands, northern
England, southern and eastern Scotland and
the Outer Hebrides. The centre of gravity
of the distribution lies in southern England
and Wales, in contrast to the northern and
western bias shown by Lesser Butterfly Orchid.
World range: Almost confined to Europe,
occurring north to c. 63N in Scandinavia,
southern Finland and the Baltic States. It
ranges south to the Mediterranean, Crimea and
Caucasus, including Corsica, Sicily, the Aegean
Islands, Cyprus and possibly Sardinia. Occurs
eastwards to central European Russia. It is also
found in North Africa in Tunisia and possibly
Morocco and in Turkey, northeast Syria and
northwest Iran.
251
Bract: Green, narrow, pointed and as long as

the ovary.
Ovary: Pale green, long, narrow, clearly sixribbed and twisted. It is also curved to hold
the flowers pointing outwards and a little
downwards.
Flower: White, washed green, with a long
strap-shaped lip and extremely long curved
spur. The sepals are white, washed greenish
towards the tip; the lateral sepals are ovaltriangular, asymmetrical and even sickle-shaped
whereas the upper sepal is rather shorter,
broader and more triangular or heart-shaped.
The petals are white, variably washed greenish
How to find it
Woodland plants are often easy to spot,
being rather tall, but those in the open
among grass and scrub can be well hidden.
A large proportion of most populations are
non-flowering, and when heavily shaded in
overgrown coppice or very dark woods plants
can remain in a vegetative state for decades.
They reappear and flower again following
coppicing, tree-falls or other changes that let
more light in. In woods with a regular coppicecycle, flowering is most prolific two or three
years after coppicing.
DESCRIPTION
Stem: Pale green, more-or-less triangular
and ribbed towards the tip, with one to three
brownish sheaths at the extreme base.
Leaves: Two, oval to elliptical, keeled, pale
green (often slightly bluish) and rather shiny.
They lie opposite each other at the base of the
stem, one just above the other, and are held
variably erect. There are also one to six small,
lanceolate, bract-like leaves higher on the stem.
Spike: Variable. The ten to 30 flowers
(occasionally up to 40) form a loose, open spike
in woodland plants, but in full sun the spike
tends to be more compact.
M 14 June, Kent. The flower spike is extraordinarily

graceful.
29/1/09 12:31:56
252
GENUS PLATANTHERA.
in the open tend to be shorter and more

compact and to have shorter leaves than those
in shaded woodland localities. It seems likely
that these differences are produced by the
local environment rather than by any genetic
difference. Plants may occasionally have just
one leaf or sometimes three or four.
BIOLOGY
M 14 June, Kent. The extraordinarily long spur contains

nectar that can only be reached by insects with an equally
long proboscis.
and are shorter, narrower and more strapshaped than the sepals. The upper sepal and
petals form a loose hood over the column, and
the lateral sepals are spread horizontally and
are often a little twisted or wavy-edged. The
lip is creamy but becomes greener towards the
tip and is narrow and strap-shaped, 10-16mm
long and projects forwards and downwards. The
spur is long and slender, 19-35mm long x 1mm
wide, broadens towards the tip and is often
strongly curved; it is washed green. The pollinia
are conspicuously yellow and relatively large,
3-4mm tall including the long caudicle (stalk).
The viscidia at the base of the pollinia lie about
4mm apart on either side of the foot of the
column, and the pollinia lean inwards towards
each other. The flowers emit a heavy scent,
especially at night, and it is said that people with
a sensitive nose can smell them from several
hundred metres on a still summers evening.
The pollination mechanism is very similar to

that of Lesser Butterfly Orchid, and nightflying moths are the primary pollinators. But
although hawkmoths are involved, members
of the large family of Noctuid moths are
more important as pollinators. In the Greater
Butterfly Orchid the sticky viscidia at the base
of the pollinia face inwards on either side of,
and just above, the entrance to the spur. Due to
this more widely spread position, the pollinia
usually become attached to the large compound
eyes of the visiting moth rather than its
proboscis. The mechanism is effective and seed
is set in 70-90% of flowers.
Subspecies
None.

As in Lesser Butterfly Orchid, plants growing
M 8 July, Perth and Kinross. The lip is longer than wide

and often only obscurely lobed, and the wings are oval,
pointed and usually held more or less drooped.
29/1/09 12:31:59
Vegetative reproduction can occur via the

formation of additional tubers but is of little
importance in the dynamics of a population.
253
meadows, have been responsible for the decline.

The species will also vanish from woodland if it
becomes too intensely shaded but may reappear
if the canopy is opened again.
The process of development from seed to

flowering plant is very similar to Lesser
Butterfly Orchid. The adult plant is reported
to be entirely independent of fungi, but this
seems highly unlikely given that it can become
dormant in heavy shade for long periods.
Hybrids
See Lesser Butterfly Orchid.

The specific name chlorantha means green
flowers. The two butterfly orchids, although
readily separated by the shape of the pollinia,
are genetically almost indistinguishable,
suggesting that they have only very recently
separated into two species.
HISTORY AND
CONSERVATION
The first British record dates from 1597 when
John Gerard noted this species in his Herball:
in the wood belonging to a worshipfull
gentleman of Kent named Master Sedley of
Southfleete.
Past and present occurrence of Greater Butterfly Orchid in
Britain
Ireland
1,163
251
626 (22%*)
117 (11.6%*)
% lost, 1500-1969
32%
41.5%
% lost, 1970-1986
14%
12%
% lost, total
46%
53.5%

1500-1999
current range
It has been lost from 46% of the historical

range in England and 53.5% in Ireland and is
classified as Near Threatened. The destruction
and coniferisation of woodland, as well as
agricultural improvements to pastures and hay
M 28 May, Norfolk. The spike may be relatively fewflowered and can be inconspicuous.
29/1/09 12:32:01
GENUS GYMNADENIA
FRAGRANT ORCHIDS
12/2/09 17:03:12
Distribution
Europe and Asia, with about 14 species,

five of which occur in Europe and three in
Britain. Until recently all three were treated
as subspecies of a single species, the Fragrant
Orchid.
Early observations suggested that seeds

probably germinate in the spring. The
resulting protocorm produces roots in the
second autumn, the first leaves in the third
spring (i.e. when it is two years old) and the
first tuber the following season. Given the dry
habitats, however, in which fragrant orchids
often grow, it seems unlikely that protocorms
could survive for over two years, and the
period prior to the development of the first
tuber may be very much shorter. The sequence
protocorm-root-aerial stem-tuber is shared
with other genera, including some of the
Orchis and Dactylorhiza orchids.
Floral structures
The column is short and erect with a long
rostellum that projects forward between the
two viscidia. The two club-shaped pollinia
each narrow into a caudicle (stalk) that is
attached to one of the two long, narrow
viscidia, but there is no bursicle (the flap or
pouch-like structure found in many orchids
which covers and protects the viscidia). The
two stigmas lie on the side-lobes of
the column.
Growth pattern
The aerial shoot grows from a pair of
flattened, deeply divided tapering tubers.
Roots develop in the autumn and penetrate
the upper layers of the soil. The roots and the
root-like tips of the tubers are infected with
fungi. The annual cycle of growth and the
replacement of tubers are similar to those of
the genus Orchis (see p.200).
of the aerial stem. These will go on to form
in the autumn.
Name
The generic name Gymnadenia originates from
the Greek gymnos naked and aden gland and
means quite literally naked-gland, a reference to
the lack of a bursicle (see Floral structures).
12/2/09 17:03:17
256
GENUS GYMNADENIA
cOmmOn fraGrant Orchid

Gymnadenia conopsea
Other names: Chalk Fragrant Orchid
This attractive orchid is found in species-rich grassland on chalk and limestone soils
throughout England and Wales but is rather rare in Scotland and Ireland. It sometimes
occurs in large numbers, and the fragrance emitted by a big colony can be quite overpowering,
especially in the evening.
Identification
The fragrant orchids are easy to separate from
other orchids. The pink flowers are held in a
tall, narrow, spire-like spike. Each flower has
two wings (the lateral sepals) that are held
roughly horizontally, a flat, three-lobed lip and a
long, slender, down-curved spur.
Similar species
Pyramidal Orchid has flowers that are a similar
colour, size and shape to those of Common
Fragrant Orchid and also have a long slender
spur. Pyramidal Orchid has, however, an
obviously shorter, more conical and more
closely-packed flower spike. Its flowers are a
deeper pink with a more markedly three-lobed
lip that has two diagnostic parallel raised ridges
at the base. Pyramidal Orchid also flowers later
in the summer, and there is usually relatively
little overlap between the two species.
Fragrant orchids are difficult to separate
from each other, although they are usually
found in distinct habitats. Common Fragrant
Orchid favours old, species-rich grassland on
chalk or limestone and is just occasionally
found in fens. Its scent is sickly-sweet but has
a slightly acid, rancid, or musty background.
Its main flowering period is June and early July.
However, even in its preferred dry grassland
habitat, Marsh Fragrant Orchid or Heath
Fragrant Orchid could also occur, and a
careful examination of the flower is necessary
to be reasonably certain of the identification.
O 14 June, Kent. At some sites Common Fragrant Orchid
occurs by the thousand.
P 14 June, Kent.
Intermediates also occur, and it is best to

examine several plants in a population to gain
an overall impression. The criteria used to
separate the three fragrant orchids are still
being developed, and some plants or even whole
populations may not yet be identifiable.
In Common Fragrant Orchid the flowers are
medium sized, the lip is about as wide as long,
lacks shoulders and is distinctly three-lobed.
The central lobe is the longest. The wings are
angled downwards and are pointed, narrow and
parallel-sided (with the lateral sepals rolled into
a tube to produce this linear shape). See also
p.258.
Habitat
Found in dry, species-rich grassland on
calcareous soils, mostly chalk downland in
the south and limestone pastures in northern
England but sometimes also on stabilised
dunes, road verges, railway banks and in old
quarries on suitable calcium-rich soils. It
occasionally occurs in base-rich fens (together
with Marsh Fragrant Orchid) or on alkaline
Leblanc waste in Lancashire. Recorded up to
365m above sea level (near Whitely Shield,
Northumberland).
Flowering period
Late May (sometimes mid-May) to late July,
but most are in flower during June.
Range
Widespread in England, although confined
to areas with suitable habitat, such as
the North and South Downs, the chalk
districts of Hampshire, the Berkshire
Downs and Chilterns, Dorset, and the
29/1/09 12:32:06
COMMON FRAGRANT ORCHID
257
29/1/09 12:32:09
258
GENUS GYMNADENIA
Separation of Common, Marsh and Heath Fragrant Orchids.

Common
Fragrant Orchid
Marsh
Fragrant Orchid
Heath
Fragrant Orchid
Width of florets*
10-11mm (7-13mm)
11-13mm (10-14.5mm)
8-10mm (7-12mm)
Lip: width
5.5-6.5cm (4.5-7mm)
6.5-7mm (5.5-8mm)
3.5-4mm (3-5mm)
Lip: length
5-6mm (4-6.5mm)
3.5-4mm (3-4.5mm)
4-4.5mm (3.5-5mm)
Lip: shape
fractionally broader
than long
much broader
than long
longer than
broad
Lip: lobes
distinctly 3-lobed,
central lobe longest
distinctly 3-lobed,
side-lobes largest
obscurely lobed,
longer than wide
Lip: shoulders**
absent
distinct
narrow
Lateral sepals: width
c. 1mm
c. 1mm
c. 2mm
Lateral sepals: length
c. 5-6mm
c. 6-7mm
c. 4-5mm
Lateral sepals: shape
linear, pointed at tip
linear, blunt at tip
oval-lanceolate,
pointed at tip
Lateral sepals:
position
deflexed at c. 30
horizontal
deflexed
Length of spur
12-14mm (11-17mm)
14-16mm (13-17mm)
11-14mm (8-15mm)
Height
20-40cm, sometimes
more
30-60cm, less in
dry habitats
15-25cm, rarely
more
Fragrance
sickly-sweet with acid

or rancid overtones
spicy sweet, clovelike; no acid overtones
spicy sweet,
recalling cloves
Flowering period
early June-mid July
early July-mid August
June-August
Habitat
chalk & limestone

grassland
fens; rarely chalk

grassland on northfacing slopes

mildly acidic to
base-rich grassland;
base-rich heathland
flushes; very rarely
chalk grassland
Largely after Francis Rose in Rich & Jermy (1998).

* The width of the floret is measured from tip to tip of the lateral sepals
** The shoulders are formed by the abrupt angle at the base of the lip where it narrows towards the mouth of the spur.
limestone areas of the Cotswolds, Mendip

Hills, Northamptonshire, Lincolnshire, the
Derbyshire and Yorkshire Dales, Cumbria and
Co. Durham. Widespread in Wales, but in
Scotland it is rare and local and information
about its distribution is incomplete; it may
have been recorded from Dumfries-shire,
Roxburghshire, Banffshire and a few areas in
the far north, in Sutherland, Caithness and
the Outer Hebrides. Its status in Ireland is
also obscure due to confusion with the other
fragrant orchids and with hybrids between
them. It may be most frequent in the central
counties, but it is probably scarce or rare.
World range: The Fragrant Orchid, i.e. an
aggregate of all three species, is found in Europe
1987-99
1970-86
pre 1970
Common, Heath
and Marsh
Fragrant Orchids
combined
29/1/09 12:32:10
259
and Asia east to Japan and Korea. In Europe

it ranges north to northernmost Scandinavia
and south to the Mediterranean (although it
avoids the Mediterranean lowlands), Turkey
and the Caucasus. Further east it occurs in
the Himalayas and in central China. Records
seldom distinguish the three species of fragrant
orchid, and their exact ranges have yet to be
defined. Fragrant Orchid has been found once
in North America, in Connecticut in 1887.
How to find it
This is usually an easy species to find at suitable
sites, now often reserves or SSSIs. It sometimes
appears in large numbers but the number of
flowering spikes can vary widely from year
to year.
DESCRIPTION
Height: 10-60cm but usually15-30cm and
seldom over 40cm.
Stem: Green, becoming more purplish towards
the flower spike.
Leaves: Mid-green. The three to five narrow,
strap-shaped basal leaves, keeled and with
pointed tips, are held loosely erect. These grade
into two or three narrow, lanceolate, bract-like
Spike: More-or-less cylindrical and moderately
densely packed with 20-50 flowers, rarely more.
The spike becomes looser as more flowers open.
Bract: Green, sometimes tinged purple, strapshaped, narrowing around the mid-point into
a finer pointed tip and roughly as long as
the ovary.
Ovary: Green, variably washed purple, long,
narrow, prominently three-ribbed and twisted.
Flower: Pink with a hint of purple, varying
in the exact shade. The sepals are elongatedoval in shape, the lateral sepals being slightly
irregular but with their upper and lower
margins rolled backwards so that face-on they
appear as parallel-sided oblongs with a short
pointed tip. The petals are a little shorter, more
oval-triangular in shape and asymmetrical,
with one side squared off. The upper sepal and
petals form a hood over the column, whereas
M 9 June, Northamptonshire. A full, densely-flowered spike.
the lateral sepals are held to the side, about 30

below the horizontal and pressed backwards.
The lip is flat and about as wide as long, with
three well-developed, rounded terminal lobes;
the central lobe is longer than the lateral lobes
and often broader. The spur is a darker, more
purplish-pink, very slender, down-curved and
rather long, around twice the length of the ovary.
It is filled with nectar and this can sometimes
be seen through its semi-translucent walls. The
flowers have a strong sickly-sweet scent.
29/1/09 12:32:12
260
GENUS GYMNADENIA
Subspecies
None.

Var. albiflora has white flowers and is fairly
frequent.
Var. crenulata has broad lateral lobes to the
lip which are narrowly serrated at the edges.
It is rare.
BIOLOGY
Nectar is produced in the bottom of the
long spur, and only insects with a sufficiently
long proboscis can reach this. Pollinators
include butterflies and both day- and nightflying moths, including Large Skipper, Sixspot Burnet and hawkmoths. Night-flying
moths may, however, be the most important
pollinators; the scent becomes more pungent
towards dusk, and, with its white flowers, var.
albiflora may only be attractive to nocturnal
moths. As the visiting insect advances to sip
nectar from the spur, the pollinia are fixed by
their sticky viscidia (which lie just above the
mouth of the spur) to the insects proboscis.
The pollinia are then carried to another flower
or another plant, having in the meantime swung
forward into a position ready to make contact
with the stigma. Pollination is very efficient and
seed is set in large quantities.
Vegetative reproduction may also occur via
the production of additional tubers.

The interval between germination and flowering
is usually around five years but may be as short
as three.
Hybrids
Opposite is a list of all the hybrids listed
officially for Fragrant Orchid by Stace (2004),
which does not separate the three fragrant
orchids into distinct species. The names of these
hybrids may well need to be revised to reflect the
new status of the fragrant orchid parent:
O 14 June, Kent. The white-flowered variety albiflora is
fairly frequent.
29/1/09 12:32:14
261
P 9 June, Northamptonshire. The lip is distinctly

three-lobed and about as
long as it is wide, while
the wings formed by the
lateral sepals are narrow,
parallel-sided and held
slightly drooped.
X Gymnanacamptis anacamptis, the hybrid

with Pyramidal Orchid, has been recorded
rarely (presumably x Common Fragrant
Orchid).
X Pseudadenia schweinfurthii, the hybrid
with Small White Orchid, has been recorded
from Yorkshire and Scotland, where it is fairly
frequent in the northwest (presumably x Heath
Fragrant Orchid).
X Dactylodenia jacksonii, the hybrid with Frog
Orchid, has been noted rarely but widely.
X Dactylodenia st-quintinii, the hybrid with
Common Spotted Orchid, is uncommon but
has been found widely through Britain and
Ireland; all three species of fragrant orchid may
be involved.
X Dactylodenia legrandiana, the hybrid
with Heath Spotted Orchid, has been found
widely scattered through Britain and Ireland
(presumably x Heath Fragrant Orchid).
X Dactylodenia vollmannii, the hybrid with
Early Marsh Orchid, has been recorded from
Cornwall and Cumbria.
X Dactylodenia wintonii, the hybrid with

Southern Marsh Orchid, has been found in
Devon and Hampshire.
X Dactylodenia varia, the hybrid with
Northern Marsh Orchid, has been found in
northern England, Scotland and Co. Down in
Ireland.
Common Fragrant Orchid can also
hybridise with Marsh Fragrant Orchid and
Heath Fragrant Orchid, but these hybrids do
not yet have names.

The specific epithet conopsea means gnat-like
or cloudy but the reason for this reference is
obscure.
Since 1997 the Fragrant Orchid has been
separated into three distinct species, Common,
Marsh and Heath Fragrant Orchids. Long
treated as separated varieties or subspecies, new
molecular and genetic evidence has shown that
they merit specific status, even though they can
be hard to identify with certainty.
29/1/09 12:32:15
262
GENUS GYMNADENIA
HISTORY AND
CONSERVATION
The first British record of Fragrant Orchid
dates from 1634 when Thomas Johnson
published the Mercurius Botanicus (Botanical
Mercury) and recorded the Orchis palmata
minor calcaribus oblongis. In montosis
(lesser palmate Orchis with oblong spurs In
mountains). In 1660 John Ray stated that the
Fragrant Orchid could be found in meadows
everywhere in Cambridgeshire.
Although the species is still common or
even abundant at suitable sites, it declined
significantly in the 19th and 20th centuries due
to the conversion of downs and pastures into
arable land and the improvement of grazing
land. Even where the habitat still remains,
like all grassland orchids this species requires
a specific level of grazing; too little and scrub
invades, too much and the diversity of species
is lost.
The identification and recording of the
three fragrant orchids is still in its infancy, and
it is hard to identify trends for the individual
species. But, taken together, the Fragrant
Orchid has vanished from 39.5% of the total
historical range in Britain and 30.5% in Ireland.
Past and present occurrence of the fragrant orchids (all
three species combined) in Britain and Ireland (based on
presence or absence in 10km squares of the National
Grid; data from the New Atlas).
Britain
Ireland
1,341
365
810 (28%*)
253 (25%*)
% lost, 1500-1969
26.5%
24.5%
% lost, 1970-1986
13%
6%
39.5%
30.5%
total historical
range, 1500-1999
current range
% lost, total
M 29 June, Hampshire. A rather open, few-flowered spike.
29/1/09 12:32:17
HEATH FRAGRANT ORCHID
263
heath fraGrant Orchid

Gymnadenia borealis
Formerly: Gymnadenia borealis subspecies densiflora
Until very recently this species was, like Marsh Fragrant Orchid, considered to be merely
a subspecies of the composite Fragrant Orchid. It favours a variety of heathy and grassy
habitats and is the commonest fragrant orchid in northern and western Britain with scattered
outposts in southern England. It may yet be discovered elsewhere.
Identification
This is a relatively small, delicate, few-flowered
cousin of Common Fragrant Orchid. The
conical spikes of pink flowers, each with a
vaguely tri-lobed lip and long slender spur,
should nevertheless identify it as one of the
three fragrant orchids. These can be difficult
to separate from each other and, although
they are usually found in distinct habitats, can
occur together. Unlike Common and Marsh
Fragrant Orchids, Heath Fragrant Orchid does
not require alkaline soils and can be found in
neutral grassland in the uplands of northern
and western Britain and also very locally in
boggy hollows on grassy heaths in southern
England. Its status in Ireland is not certain
due to confusion with the other two fragrant
orchids. The scent is very sweet and carnationor clove-like and its main flowering period is
from mid-June to late July.
Similar species
Marsh and Common Fragrant Orchids are
similar, and where there is a possibility that
these could also occur a closer look is required
to be certain of the species. In Heath Fragrant
Orchid the flowers are relatively small, around
8-10mm across. The lip is longer than wide
and often rather obscurely lobed, with small
side-lobes and a longer central lobe. The wings
(lateral sepals) are oval, taper to a pointed
tip and are held angled downwards to about
four and eight o-clock. Intermediates occur,
however, and it is best to examine several
plants in a population and take an average.
See table on p.258.
M 15 June, New Forest. Genetic evidence has recently

confirmed that Heath Fragrant Orchid is distinct.
29/1/09 12:32:18
264
GENUS GYMNADENIA
Habitat
Flowering period
In the north and west it is found on unimproved

hill pastures, roadside verges and grassy
moorland, often in areas flushed with ground
water. In such places it grows on tussocks of
grass and heathers, sometimes with Lesser
Butterfly Orchid and Heath Spotted Orchid.
It is also found on machair, dunes and in old
quarries. It is tolerant of a wider range of pH
than Common and Marsh Fragrant Orchids
and will grow on soils that are both mildly acidic
and alkaline, on sands, limestones and clays. In
the New Forest and Ashdown Forest it is found
in marl bogs (flushes and hollows on grassy
heaths on base-rich clays). Very rarely it has been
recorded from chalk grassland, as near Lewes in
Sussex. Occurs up to 610m above sea level (Ben
Lawers, Perthshire).
June to August in Scotland and Ireland and

early June to early July in the New Forest
(around the same time as Common Fragrant
Orchid or just a little later); anomalously, the
population on chalk grassland in Sussex flowers
from late July to early August.
Range
Poorly understood as it has only recently been
separated from the other fragrant orchids. So
far it is known to occur in southern England
in Cornwall (including the Lizard Peninsula
and Bodmin Moor), Devon, Dorset, the New
Forest in Hampshire (about 12 relatively small
populations of 50-100 flowering plants), East
and West Sussex (Ashdown Forest and three
sites on the South Downs), and in Shropshire
O 15 June, New Forest. Growing

with Heath Spotted Orchid on a
large tussock.
29/1/09 12:32:20
HEATH FRAGRANT ORCHID
265
(one site), Derbyshire, Northumberland and

Cumbria (just two sites). In Wales it is known
only from Breconshire and Cardiganshire.
Scattered through much of Scotland, including
the Inner and Outer Hebrides, Orkney and
Shetland (although very rare there and now
found only on Unst). Occurs in western
and northern Ireland but there is very little
information on the distribution of Heath
Fragrant Orchid in Ireland. World Range:
Apparently more widespread in Europe than
Common Fragrant Orchid and found in France
as far south as the Dordogne. See Common
Fragrant Orchid.
How to find it
Widespread and locally common in Scotland
but very local in England and Wales. The
M 8 July, Perth and Kinross. The lip is longer than wide

and often only obscurely lobed, and the wings are oval,
pointed and held more or less drooped.
New Forest, where it is locally frequent,

is undoubtedly the best area to look for
this species in southern England. It can be
surprisingly elusive among tussocks of long
grass, and the number of flowering spikes can
vary widely from year to year.
DESCRIPTION
M 8 June, Dunbartonshire.The spike can be relatively squat.
Height: 10-25cm, rarely to 30cm.

Stem: Green, becoming more purplish-brown
towards the tip.
Leaves: Mid-green, the three to five narrow,
strap-shaped basal leaves are keeled and have
pointed tips. They are held loosely erect, often
in two ranks, and grade into two or three
narrower and more lanceolate bract-like leaves
higher on the stem.
Spike: More-or-less cylindrical, although often
slightly irregular in shape, with 20-30 or more
flowers in a fairly lax spike.
Bract: Green, tinged purple (especially on
the edges), strap-shaped, narrowing abruptly
around the mid-point into a finer pointed tip,
and about one-and-a-half times the length of
the ovary.
Ovary: Green, variably washed purple, long,
narrow, three-ribbed and twisted.
29/1/09 12:32:21
266
GENUS GYMNADENIA
Flower: Dark pink to lilac. The sepals are

oval or drop-shaped with the margins of the
lateral sepals rolled back to give them an ovallanceolate shape with a pointed tip. The petals
are shorter, more oval-triangular but less regular
in shape. The upper sepal and petals form a
hood over the column, whereas the lateral sepals
are held to the side, a little below the horizontal
and pressed backwards. The lip is flat and longer
than wide, lobed at the tip with the side-lobes
shorter than the central lobe; the lobes are
usually poorly developed with the incisions
between them reduced or absent. The spur is a
darker, more purplish-pink, very slender, downcurved and rather long, around twice the length
of the ovary; it is filled with nectar. The flowers
have a powerful, sweet scent, recalling cloves,
which is strongest when they are newly opened
and lacks the rancid overtones of Common
Fragrant Orchid. See also table on p.258.
Subspecies
None.

Var. albiflora has white flowers and is fairly
frequent.
BIOLOGY
No specific information, although the
differences in the size and shape of the flower
suggest that a different suite of pollinators is
involved for each of the three fragrant orchids.
See Common Fragrant Orchid.

No specific information.
Hybrids
Hybrids have been recorded with both
Common Fragrant and Heath Fragrant
Orchids. See also Common Fragrant Orchid
(p.260).

The specific name borealis means northern, a
reference to the generally northerly distribution.
This species was originally described as
a variety of Fragrant Orchid by the British
botanist G.C. Druce in 1918. It was raised to
the status of a subspecies in 1991 before finally
achieving specific status (yet to be universally
recognised) a few years later. Despite the
difficulties in identification, DNA evidence
strongly supports the separation of Heath
Fragrant Orchid from Marsh and Common
Fragrant Orchids.
HISTORY AND
CONSERVATION
M 8 June, Dunbartonshire. The spikes are variable in

shape and in the density of flowers.
This species is the most poorly known of the

three fragrant orchids and has only really been
brought to the attention of botanists since
1988 (when it was treated in the Botanical
Society of the British Isles Plant Crib). Its
history and conservation is obscure, but there
has undoubtedly been a considerable decline,
with overgrazing of upland grasslands being a
particular problem.
29/1/09 12:32:22
MARSH FRAGRANT ORCHID
267
marSh fraGrant Orchid

Gymnadenia densiflora
Formerly: Gymnadenia conopsea subspecies densiflora
Until recently this orchid was treated as a subspecies of Fragrant Orchid, a composite
that encompassed Common, Marsh and Heath Fragrant Orchids, but genetic evidence
has recently confirmed that all three are good species. It has been recorded from scattered
localities throughout the British Isles and is rather scarce, being mostly confined to alkaline
fens, a habitat that has suffered badly from drainage and water abstraction. Although
coverage was undoubtedly incomplete due to confusion with its cousins, the New Atlas
mapped it from just 58 10km squares in Britain during the period 1987-99, and it probably
qualifies for the status of Nationally Scarce.
Identification
With spire-like spikes of pink flowers, each
with a distinctly three-lobed lip and a long,
slender, curved spur, this species is easy to
identify as one of the three fragrant orchids.
Similar species
The fragrant orchids can be hard to separate
from each other, although they usually grow in
different habitats. Marsh Fragrant Orchid is
found in meadows and fens where the ground
water is distinctly alkaline and sometimes
occurs in large numbers, often together with
Marsh Helleborine. It is occasionally recorded
from chalk downland and can sometimes occur
together with either Common or Heath Fragrant
Orchids. Marsh Fragrant Orchid is nevertheless
relatively distinctive, being a tall, robust plant
with numerous broad basal leaves that are
usually held noticeably erect and a good-sized
spike of relatively large, dark pink flowers. To
clinch the identification, a careful examination,
preferably of several plants, is necessary. In
Marsh Fragrant Orchid the flowers are around
11-13mm across and the lip is broader than long,
prominently lobed (with the side-lobes larger
than the central lobe) and has distinct shoulders.
The wings (lateral sepals) are long and narrow
with parallel sides and a blunt tip and are
held roughly horizontal. See table on p.258.
P 11 July, Norfolk. The robust fragrant orchids of fenland
habitats have long been recognised as distinct.
29/1/09 12:32:23
268
GENUS GYMNADENIA
Intermediates and hybrids do occur, however,

and some populations do not fit any of the
three species (see photograph on p.270).
Flowering period
Habitat
Rather poorly known because until recently

many records were given merely as Fragrant
Orchid. In England found in a belt of counties
along the south coast from Cornwall to Kent
(including the Isle of Wight), at a handful
of sites in Berkshire and Hertfordshire, in
Norfolk and adjacent parts of Suffolk, and
then through the Midlands and northern
England north of a line from Cambridgeshire
to Herefordshire, although very scattered.
In Scotland the species is similarly local and
has been recorded from Peebles, Midlothian,
Banffshire, a few islands in the Inner and Outer
Hebrides, Ross & Cromarty, Sutherland and
Orkney. Similarly scattered in western and
northern Wales. Apparently the commonest
of the three fragrant orchids in Ireland but
there are few specific records. World range:
Widespread in Europe and especially common
in the southeast, particularly the Balkans.
Usually found in fens and meadows flushed

with calcareous, base-rich water, also dune
slacks and, on the Isle of Wight, slumped clay
cliffs. Occasionally occurs on chalk grassland
on north-facing slopes, as at Ditchling Beacon
and Heyshott Down in Sussex. Recorded up to
310m above sea level (Tomintoul, Banffshire).
Late June to mid-August.
Range
How to find it
The tall plants in fens and calcareous meadows
are easy to find, although this habitat is very
local. Chalk grassland populations can very easily
be overlooked, however, and any population of
fragrant orchids is worth careful examination.
DESCRIPTION
M 17 July, Norfolk. The deep, purplish-pink colours of

the fragrant orchids are extremely hard to reproduce in
print.
P 11 July, Norfolk. A typical tall, spire-like spike.
Height: 30-60cm, exceptionally to 90cm; plants

in chalk grassland are the smallest.
Stem: Green, washed purple towards the tip.
Leaves: There is a rosette of several erect,
lanceolate and relatively broad leaves at the base
of the stem and several smaller, narrow and
more bract-like leaves higher up.
Spike: Up to 100 flowers form a rather tall,
narrow spike.
Bract: Green, washed purple (especially towards
the edges) and narrowly oval, tapering to a point;
the bracts are roughly the length of the ovary.
Ovary: Green, variably and sometimes heavily
washed purple; the ovary is long, narrow,
curved, prominently three-ribbed and twisted.
29/1/09 12:32:24
269
29/1/09 12:32:27
270
GENUS GYMNADENIA
Flower: Deep pink, variably tinged purple

and often whiter around the base of the lip.
The sepals are elongated-oval in shape with
the upper and lower margins of the lateral
sepals rolled back so that they appear to be
parallel-sided with a blunt tip. The petals are
a little shorter and more oval-triangular in
shape. The upper sepal and petals form a hood
over the column whereas the lateral sepals are
held horizontally to the side. The lip is flat,
broader than long and has distinct shoulders
(abrupt angles on each side at the base where
it narrows towards the mouth of the spur).
It has three well-developed rounded lobes,
with the side-lobes larger than the central
lobe. The spur is relatively dark, very slender,
down-curved and long. The scent is spicy
sweet, recalling cloves, and lacking the rancid
overtones of Common Fragrant Orchid.
See also table on p.258.
M 11 July, Norfolk. The lip is wider than long, with large

side lobes and distinct shoulders, and the wings are long,
narrow, blunt and usually carried horizontally.
Subspecies
None.

The dune-slack population at Kenfig in south
Wales has recently been assigned to var.
friesica, described from Dutch plants and said
to intermediate in appearance between Marsh
and Heath Fragrant Orchids. The validity of
var. friesica and its relationship to the three
species of fragrant orchid has, however, yet to
be clarified.
BIOLOGY
No specific information but the differences in the
size and shape of the flower, notably the long spur
and distinct scent in Marsh Fragrant Orchid,
suggest that a different suite of pollinators is
involved for each of the three fragrant orchids.
See also Common Fragrant Orchid.

Hybrids
M 9 July, Norfolk.The shape of the flowers is variable.
Hybridises with Heath Fragrant Orchid in

Ireland and sometimes with Common Fragrant
Orchid. See also Common Fragrant Orchid.
29/1/09 12:32:30
271

The specific name densiflora refers to the
densely packed flower spike, although the spike
is not necessarily more crowded than Common
Fragrant Orchid.
Originally described as a distinct species
in 1806, Orchis densiflora was quickly reduced
to being merely a variety of Fragrant Orchid
(although it did appear as a distinct species in
Francis Roses excellent Wild Flower Key as
long ago as 1981). Recently, DNA evidence has
shown that it, together with Heath Fragrant
Orchid, merits full specific status.
HISTORY AND
CONSERVATION
Confusion with Common Fragrant Orchid
means that its past history in Britain is obscure.
Although var. densiflora warranted some
attention, the identification criteria used were
sometimes dubious, notably the width of the
lower leaves and the length of the flower spike,
and many records merit re-examination.
Marsh Fragrant Orchid has undoubtedly
undergone a serious decline in much of Britain.
The direct drainage and destruction of fens
and marshes has caused some losses. However,
subtler effects, including eutrophication and
the lowering of water tables, have also caused
declines which reserve or SSSI status could
not protect against. The few chalk grassland
populations are also subject to losses due to
improvement or abandonment. The species
is now much reduced in Norfolk, one of its
strongholds, and has withdrawn to just five
sites in Suffolk. There are no recent records
from Somerset, Wiltshire, Gloucestershire,
Bedfordshire, Lincolnshire, Co. Durham,
Lanarkshire, Berwickshire, Perthshire or
Angus.
P 3 July, Norfolk.The flowering period can be somewhat

variable in this small population by two to three weeks
and is of limited use in identification or in delineating
the various forms.
29/1/09 12:32:31
GenuS DACTYLORHIZA
marSh OrchidS and
SpOtted OrchidS
29/1/09 12:32:39
GENUS DACTYLORHIZA
273
This genus, which includes the marsh and spotted orchids, presents the toughest identification
challenge among British and Irish orchids. The species are closely related and individually
quite variable. Indeed, there is disagreement as to how the various populations should be
classified into species, subspecies and varieties. Hybridisation is also relatively common,
making identification even more difficult. The situation can be particularly complex in
Ireland, west Wales and western Scotland due to the number of possibilities that have to
be considered. If in doubt it may be necessary to take a statistical approach and carefully
measure a selection of plants in order to reach a satisfactory identification. On the other
hand, a field full of Southern Marsh Orchids or the scatter of Heath Spotted Orchids across
a heather moor are straightforward to identify and, indeed, spectacular.
Distribution
Approximately 75 species are found in
Europe and temperate Asia. The range of one
species extends into Alaska via the Aleutian
Islands, and Frog Orchid has a circumpolar
distribution.
Classification
Dactylorhiza is one of several closely related
genera that all have well-developed tubers. It
is probably most closely related to the fragrant
orchids Gymnadenia and butterfly orchids
Platanthera and only more distantly to the
typical orchids in the genus Orchis. This is
despite the fact that until the 1930s and 1940s
the marsh and spotted orchids were placed in
the genus Orchis.
The Dactylorhiza orchids can be divided
into two groups depending on the number of
chromosomes they possess. The diploid group
has a chromosome count of 2n = 40. This
includes Early Marsh Orchid and Common
Spotted Orchid. The tetraploid marsh orchids
have a chromosome count of 2n = 80. This
group includes Southern, Pugsleys, Northern,
Irish and Hebridean Marsh Orchids. The
chromosome count is significant for two reasons.
The first is hybridisation as it has some influence
on the fertility of hybrids. Second, the tetraploid
marsh orchids are thought to have evolved from
a cross between the ancestors of two members
of the diploid group, Early Marsh Orchid
and Common Spotted Orchid, followed by a
doubling of the chromosomes (in which 2n =
40 became 2n = 80). This allowed the hybrid to
become reproductively isolated from its parents.

This hybridisation event occurred several times
and with slightly differing parents, producing the
closely similar species we see today.
On genetic evidence the Frog Orchid has
recently been moved into Dactylorhiza. It is
thought to be a primitive member of the genus
and differs from the other British species not
only in its general appearance but also in having
a rudimentary spur and bursicle, by producing
nectar, and by having the hood of the flower
made up from all of the sepals and petals (in
the other species the lateral sepals are held away
from the hood).
Identification
The most useful tools in the identification of
marsh and spotted orchids are a small ruler, a
notebook and pencil. The ruler is useful because
for many species the width of the flowers lip
is a critical measurement, as well as its length.
Sometimes the width of the widest leaf and
the total height of the plant are important too.
The notebook and pencil are useful because it
is almost always necessary to take the details of
several plants. The following are worth noting:
1. Lip size: the width across the widest point of
the flattened lip and also the length (from the
mouth of the spur to the tip of the central lobe).
2. Lip shape: the lip is often folded or has the
sides turned down. To judge its shape it is
necessary to flatten it. This can easily be done
by sliding the ruler or a finger below the lip and
gently pushing upwards. The degree to which
the side-lobes are turned downwards in its
29/1/09 12:32:43
274
GENUS DACTYLORHIZA
upper
sepal
lateral
sepal
lateral
sepal
petal
petal
mouth of
the spur
side-lobe
bract
non-sheathing
leaf
double loop
markings
central lobe
non-sheathing
leaf
sheathing
leaf
sheathing
leaf
basal
leaf
basal
sheath
tuber
(divided into
finger-like lobes)
M A typical Dactylorhiza orchid.
O 1 June, Norfolk. Southern Marsh Orchid: The Dactylorhiza may produce spectacular displays, but the colours
of the marsh orchids are extremely hard to reproduce in
print.
sinus
natural position may also be important. The

lip is almost always lobed but it may also have
small cuts or incisions (sinuses) separating the
lobes; see figure above.
3. Lip markings.
4. Flower colour: only useful in some cases,
as the tetraploid marsh orchids (Southern,
Northern, Pugsleys, Irish and Hebridean) are
all similar shades of purplish-pink.
5. Position of lateral sepals: these may be held
horizontally at the side of the flower, vertically
over the flower or at any angle in between.
6. Lateral sepal markings.
7. The shape of the spur.
8. The number of flowers: surprisingly hard to
count and only really useful in the identification
of Pugsleys Marsh Orchid. If there are too
many to count, it cannot be that species.
9. The number of leaves: the leaves are divided
into sheathing and non-sheathing. Sheathing
leaves are found on the lower part of the stem
and at their base they completely encircle the
stem with a short, unbroken, tubular sheath.
Non-sheathing leaves are found higher on the
stem, are narrower and more bract-like. They
have a clearly defined base that may encircle the
stem but never have an unbroken sheath.
Importantly, the basal leaf is not included
in any counts. This can range from a very short
sheath at ground level with a green tip to a
29/1/09 12:32:44
GENUS DACTYLORHIZA
fully-formed leaf up to half the length of the

sheathing leaf immediately above it (see figure
opposite).
10. Leaf markings.
11. Leaf hooding: the tips of the leaves in some
species are described as hooded but this is a
subtle and very variable character and is of
limited use (see figure below).
hooded leaf-tip
Floral structures
There are two pollinia, each narrowing into a
caudicle (stalk) and attached by a basal disc
to one of the viscidia. A two-lobed bursicle
is present and encloses the two viscidia. The
stigma is more-or-less two-lobed and situated
at the roof of the entrance to the spur.
Pollination
Dactylorhiza are cross-pollinated by bees but
most members of the genus produce no nectar.
The insects receive no reward and are perhaps
just attracted to the mass of brightly coloured
flowers which are often found in large numbers.
The attraction is therefore a deceit. The exception
is Frog Orchid, which does produce nectar.
Growth pattern
The aerial stem grows from a pair of tubers.
These are flattened and each divided into two to
five finger-like lobes which taper to a fine point.
They may be long in dry habitats, extending
well down into the soil, but in wet areas may
bend upwards towards the surface of the soil,
perhaps to avoid becoming waterlogged. There
are also several long, fleshy roots growing near
the surface of the soil.
275
As in the genus Orchis, the next seasons

tuber develops alongside the current one and
the system of annual replacement is similar (see
p.200).
Fungal partners
In the adult plant only the roots and sometimes
the tips of the tubers are infected with fungi.
Members of this genus have well-developed
foliage, and fungi may play a relatively minor
role in their nutritional budget.

Seed probably germinates in the autumn
to produce a tiny, conical or turnip-shaped
protocorm about 2mm long. The first root and
leafy shoot are produced the following spring,
and the first tuber develops from a bud at the
base of this leafy shoot. This tuber is rodshaped with just one finger-like extension and
in the late summer the protocorm, root and
leafy shoot disappear, leaving just the tuber.
In the autumn a short rhizome grows from a
bud at the tip of the tuber. This produces roots
and, in the spring, elongates to produce the
next leafy shoot. The sequence of events is very
similar to that of the genera Orchis, Gymnadenia
and Platanthera (see p.201). A regular annual
cycle now begins. The tuber that produced
the current years growth gradually shrivels
away, and a new tuber forms beside it from
a bud on the short rhizome. This swells over
the summer as it stores up nutrients produced
by photosynthesis. The period between
germination and flowering is four or five years
in most species of Dactylorhiza.
Additional tubers may develop at the end of
short shoots growing from the base of the stem,
and these become separate plants when the
central stem dies away in the autumn.
Name
The generic name Dactylorhiza originates
from the Greek daktulos finger and rhiza root
and means finger-like root, a reference to the
distinctive shape of the tubers.
29/1/09 12:32:44
276
GENUS DACTYLORHIZA
early marSh Orchid

Dactylorhiza incarnata
Formerly: Orchis latifolia, Dactylorchis latifolia
This is the most widespread of the marsh orchids, occurring throughout Britain and Ireland
in a wide variety of habitats. However, it is always rather local, often uncommon and indeed
has disappeared from many areas. It is also the most variable marsh orchid with five named
subspecies and flowers that range in colour from deep red to creamy. Because the subspecies
are so different and occur in different habitats, they are discussed separately in the text.
Identification
M 6 July, Lancashire. Subspecies incarnata. A very

stout plant.
O 13 June, Norfolk. Subspecies incarnata. A slender, petite plant with few
flowers.
The five subspecies in brief:

1. D. i. incarnata has flowers that are usually
very pale pink but sometimes purplish-pink. It
is found in alkaline fens and marshy meadows
throughout the British Isles.
2. D. i. coccinea has flowers that are deep red
and often looks like a fat little hyacinth. It
occurs scattered on both the east and west
coasts in coastal dune slacks. There are also
some inland sites, especially in Ireland.
3. D. i. pulchella has flowers that are purplishpink. It is largely confined to bogs on acid
heathland in southern England.
4. D. i. cruenta Flecked Marsh Orchid has
flowers that are mid to dark pink. It also
frequently has bold spots on the leaves and
bracts. It is confined to alkaline fens in western
Ireland and northwest Scotland.
5. D. i. ochroleuca has unmarked creamy
flowers and is found in alkaline fens in East
Anglia but is now very rare.
Early Marsh Orchid is probably the most
variable species of orchid in the British Isles,
both in flower colour and in stature. It may
be very petite and just 5cm tall or a robust
giant at 60cm. Nevertheless, it is relatively
distinctive because the individual flowers have
a characteristic size and shape: They are always
small, indeed disproportionately so on robust
plants compared to the stem, bracts and ovaries.
The lip is no more than 9mm wide and usually
rather less (push a finger gently upwards from
below to flatten the lip before measuring it).
Furthermore, the flowers appear even narrower
29/1/09 12:32:45
EARLY MARSH ORCHID
277
M 19 June, Norfolk. Subspecies coccinea.

This subspecies appears in many different
shades of red, from orange-red to purplishred.
O 27 June, Norfolk. Subspecies coccinea.
Grows in dune slacks and also on slumped
clay cliffs.
because the sides of the lip are often folded

downwards, sometimes sharply so, especially as
the spike matures. This narrow appearance is
further accentuated by the lateral sepals, which
are held vertically above the flower as if it is
holding its arms up in surrender. And, whatever
their coloration, the flowers have the same
basic pattern of markings (with the exception
of the unmarked creamy flowers of subspecies
ochroleuca). The centre of the lip has a scatter
of wriggly lines of varying length and these are
all bounded by a solid dark line that forms a
double loop, with few, if any, dark markings
outside this loop.
Similar species
The pink-flowered subspecies D. i. incarnata
is distinctive as no other British marsh orchid
shows this coloration. Similarly the deep red
D. i. coccinea is characteristic, but the various

purple-flowered forms can be more confusing.
Southern Marsh Orchid always has larger
flowers with a broader lip. The sides of the
lip are quite frequently folded downwards
but never as sharply as in some Early Marsh
Orchids, and the lip is often held flat or even
dished. It typically lacks prominent double loop
markings and although Leopard Marsh Orchid
(var. junialis of Southern Marsh Orchid) does
have loop markings it also has ring-shaped
spots on its leaves.
Northern Marsh Orchid is quite similar
to purple-flowered Early Marsh Orchids but
its flowers are on average slightly larger with
a broader, distinctively diamond-shaped lip
which is usually held flat. It is often an intense
reddish-pink, lacks the double loop markings,
29/1/09 12:32:48
278
GENUS DACTYLORHIZA
extend to the edge of the lip and it often lacks

the double loop of Early Marsh Orchid. In
Scotland, Lapland Marsh Orchid (subspecies
lapponica of Pugsleys Marsh Orchid) has been
found in the same localities as D. i. cruenta,
the spotted-leaved subspecies of Early Marsh
Orchid, but although they share spotted leaves
and bracts and heavily marked lips, they can be
separated as above.
Irish Marsh Orchid is usually stouter
than Early Marsh Orchid with the leaves
neither as rigidly upright nor hooded. It has a
proportionally larger flower spike and a flatter,
broader lip that has more evenly rounded lobes.
Flowering period
Late May to late June, occasionally from
mid-May or to late July. There is considerable
variation both within individual colonies and
between sites but late May and early June is
probably the peak for many. Beyond a tendency
for plants in the south and west to flower
earlier and plants in wetter sites and at higher
altitudes to flower later, there seems to be no
real difference between the flowering times of
the various subspecies.
Habitat
M 15 June, New Forest. Subspecies pulchella. Grows in

acid bogs with bog-mosses Sphagnum.
and its lateral sepals are not usually held as

near to the vertical. Subspecies cambrensis of
Northern Marsh Orchid has boldly spotted
leaves and a slightly more three-lobed lip than
typical plants, but the lip is still broader than in
Early Marsh Orchid.
Pugsleys Marsh Orchid has narrower leaves
than Early Marsh Orchid, a rather few-flowered
spike and relatively large flowers. The lip is
usually over 9.5mm wide and distinctly threelobed, often with a prominent projecting central
lobe. The dark markings are variable but usually
Subspecies incarnata is found on calcareous

or neutral soils on damp to wet grassland.
It is especially characteristic of unimproved
wet meadows on floodplains. It is also found
in spring-fed fens and flushes, even in the
mountains of Scotland. Paradoxically, although
it is usually found in wet areas, the species
cannot survive a prolonged submergence.
Other habitats include dune slacks (where it
may be found with the red-flowered subspecies
coccinea), old fly-ash tips, occasionally old chalk
quarries where soil compaction may cause
seasonal water-logging and, very rarely, chalk
grassland. Recorded up to 440m above sea level
(Ben Lawers, Perthshire; plants not referred to
any subspecies have been found at 610m in the
Atholl District in Perthshire and also at 610m
at Caenlochan in Angus). Subspecies coccinea
is found in damp dune slacks, the machair
grasslands of the Hebrides, the flushed slopes
29/1/09 12:32:49
EARLY MARSH ORCHID
279
vegetated areas. Suitable sites were created

when peat was cut for fuel and the workings
subsequently flooded. Such turf ponds slowly
fill with vegetation which eventually forms a
floating mat or hover but in the long run the
continued build-up of material dries them out
and they become unsuitable.
Range
M 15 June, New Forest. Subspecies pulchella. The flower

is purplish-pink.
of slumped clay cliffs in east Norfolk and damp

lake shores inland in Ireland. It is also recorded
from alkaline Leblanc waste in Lancashire and
pulverised fly-ash waste. Subspecies pulchella
is found in valley bogs and acid marshes on
heathland, often growing among Sphagnum and
sundews. Subspecies cruenta Flecked Marsh
Orchid is found in the limestone districts of
Ireland in calcium-rich fens around loughs and
turloughs, frequently with Black Bog-rush;
indeed, it often grows in the hollows amongst
slabs of limestone. Its only three Scottish sites
are on roughly neutral flushes and mires up to
450m above sea level. Subspecies ochroleuca
is found in calcareous spring-fed fens where it
grows on a mossy carpet in the more sparsely
Subspecies incarnata is locally frequent in

England and Wales but is very scattered in
Scotland and Ireland. Subspecies coccinea
is very locally common on the west coast
from Devon north to the Inner and Outer
Hebrides and Shetland (South Mainland
only; it is absent from Orkney). On the east
coast it is scattered in southeast Scotland,
northeast England and north Norfolk. It has
been recorded inland in Hertfordshire and
Derbyshire, and in Ireland is very scattered but
occurs both inland and on the coast. Subspecies
coccinea is often found in very large numbers.
Subspecies pulchella occurs in Cornwall,
Dorset, the New Forest, the heaths around
the Surrey/Hampshire border, Ashdown
Forest in Sussex and perhaps also Dartmoor
in Devon. Similar plants, perhaps subspecies
pulchella (rather than subspecies incarnata with
pinkish-purple flowers), may be found in acid
habitats elsewhere in Britain, especially Ireland,
but the exact status of these populations is
1987-99
1970-86
pre 1970
29/1/09 12:32:51
280
GENUS DACTYLORHIZA
through the mountains of Central Asia to the

Himalayas. Subspecies coccinea and subspecies
pulchella are endemic to Britain. Subspecies
cruenta is widespread in Scandinavia and the
Alps and is also found in Poland, the Baltic
States and through central and northern Russia
to eastern Siberia. Subspecies ochroleuca is
local and rare but occurs widely in central
Europe from Haute-Savoie in southeast France,
Switzerland, Austria, Hungary and Romania
north to Denmark, southern Scandinavia,
Poland, the Baltic States and perhaps northwest
Russia; it has also been recorded from Spain.
But as in Britain, confusion with pale-flowered
variants of the other subspecies has obscured its
precise distribution.
How to find it
M 23 May, Co. Clare. Subspecies cruenta. Grows in calcareous fens. Not all plants have spotted leaves.
unclear. Subspecies cruenta occurs in western

Ireland in Co. Galway, Co. Mayo and the
southeastern portion of The Burren in Co.
Clare. In Scotland there are a couple of tiny
populations near Ullapool in Ross & Cromarty
and one in Assynt in Sutherland. Subspecies
ochroleuca is confined to East Anglia. World
range: Europe and Asia. Subspecies incarnata
is found throughout Europe away from the
Mediterranean lowlands, north to northern
Scandinavia and south to Spain, Italy, Greece,
the Crimea and Caucasus. In Asia it occurs
from northern Turkey to northern Iran and
in Siberia east to Lake Baikal and south
With the exception of dune populations, Early

Marsh Orchid tends to occur in small numbers,
unlike the massed ranks of Southern, Northern
or Irish Marsh Orchids, and is easily overlooked.
A search of any rough-looking riverside
meadows, with a good growth of rushes, is
recommended. In the London area, the Lee
Valley Park is an excellent site for this species.
The red-flowered form can be found in good
numbers at most of the larger protected dune
systems, and the purple-flowered subspecies
pulchella is common in many of the New Forest
bogs and also at Thursley Common in Surrey.
A visit to western Ireland is necessary to see
Flecked Marsh Orchid, subspecies cruenta.
DESCRIPTION (incarnata)
Height: 7-65cm but typically 20-40cm and
rarely more than 20cm in Scotland.
Stem: Bright yellowish-green or apple green
and usually hollow; in larger plants the thick,
hollow stem is conspicuous.
Leaves: Bright yellowish-green or apple green.
There are three to five sheathing leaves, often
crowded towards the base of the stem and
rather broad (the largest more than 2cm wide,
sometimes as much as 3.5cm), with up to two
non-sheathing leaves higher on the stem. The
leaves are typically held rather erect at c. 45, are
strongly keeled and often hooded at the tip.
29/1/09 12:32:52
EARLY MARSH ORCHID
Spike: Crowded, with ten to 70 flowers.

Bract: Apple green, sometimes flushed rosepink or purplish, lanceolate and fairly long, up
to twice as long as the ovary and projecting
well beyond the flowers in the lower part of
the spike.
Ovary: Green, six-ribbed and twisted, also bent
through approximately 45.
Flower: The flowers are usually pale pink but
at some sites there may also be plants with
flowers of various shades of purplish-pink.
The sepals and petals are off-white, variably
washed rose-pink. The lateral sepals are oval
to strap-shaped, rather like donkeys ears,
and slightly asymmetric; they are held erect
and often have dark pink spots or sometimes
ring-shaped markings. The upper sepal and
petals are more oval in shape (the petals a little
smaller) and form a tight hood. The lip is offwhite washed rose-pink, especially towards
the edges, with irregular dark pink dots and
lines in a central zone enclosed within two
complete or near-complete loops. It is usually
less than 8.5mm wide and 7mm long and is
slightly to moderately lobed, with the sides
usually strongly turned downward; indeed,
this deflexing may be so marked that the
lip is almost folded in two. The spur is pale
pink, stout, slightly tapering and slightly to
moderately decurved (rarely straight); it ranges
from half as long to nearly as long as the ovary
(but less than 7.5mm long).
281
Marsh Orchids can be found growing together.

This variation in flower colour appears to be
commoner in Wales, Cumbria, Scotland and
Ireland, but even in these areas mixed colonies
are still mostly or always found in neutral or
base-rich habitats.
(Note that D. i. gemmana has been recorded
from east Norfolk and east Galway with similar
plants found elsewhere in England, Wales and
Ireland. Ignored or forgotten for many years
it has recently been resurrected but it may be
best to treat it as a large, late-flowering variant
of subspecies incarnata. It is robust, growing to
50cm (sometimes 80cm), has six or more leaves
and is large flowered, with the lip more than
8.5mm wide x 7mm long and the spur usually
longer than 7.5mm. The flowers are either pink
or purple and the lip is marked with fine dots
rather than a double loop, suggesting perhaps a
hybrid origin.)
D. i. coccinea is often relatively small at 5-20cm
but may grow to 30cm; it can appear stout,
squat and apparently stemless. The flower is a
Subspecies
Each of the five subspecies is sometimes treated
as a distinct species by European authors,
but recent genetic analyses show that they
are very similar indeed, with the exception
of subspecies cruenta, which does show some
genetic differences. The dividing lines between
the various subspecies are not hard and fast
and intermediates occur. See also Habitat and
Range.
D. i. incarnata is the typical subspecies and is
described above.
In southern England the vast majority
of plants have pale pink flowers but at a few
sites both pale pink and purplish-pink Early
M 23 May, Co. Clare. Subspecies cruenta. The flower is

mid to dark pink.This is a heavily marked plant.
29/1/09 12:32:53
282
GENUS DACTYLORHIZA
29/1/09 12:32:58
EARLY MARSH ORCHID
distinctive deep red with even darker markings

although these are less obvious than in other
subspecies due to the reduced contrast with the
dark red ground colour; the markings on the lip
are rather narrow and appear quite faint. The
bracts are rather long and are washed and edged
purple, and the upper stem and ovaries are also
strongly washed purple. The spur is stout and
bag-like or conical. The leaves are broad-based
but sharply tapering, dark green and unspotted.
In most dune colonies there are also plants with
flowers which are paler and pinker, intermediate
with subspecies incarnata, and often some
typical pale pink incarnata too.
D. i. pulchella has purplish-pink flowers that
fade to white around the mouth of the spur;
they are rather similar in colour to Southern
Marsh Orchid although slightly more purple.
The upper stem, bracts and ovaries are variably
washed purple, and as in subspecies incarnata
the lateral sepals may have ring-shaped
markings. Apart from the colour of the flower
283
it differs from subspecies incarnata in having

thicker, bolder and more complete double loop
markings on the lip, which is less obviously
lobed. The sides of the lip either do not fold
downwards at all or do so only as the flower
gets older, and the lip may therefore be flat
or even slightly dished. On average the leaves
are narrower and a darker and deeper green.
The bracts are shorter and less prominent.
There is some variation in flower colour and
in some colonies very pale yellowish flowers
are relatively common; these are subspecies
pulchella, var. ochrantha.
The subspecific name pulchella is often
used willy-nilly for any purple-flowered Early
Marsh Orchid wherever it grows. We prefer
to reserve the name pulchella for the heavily
marked, purple-flowered populations growing
in acid bogs and to treat the purple Early Marsh
Orchids that grow with pale pink-flowered
plants in alkaline habitats as colour variants of
subspecies incarnata.
M 6 July, Lancashire. Subspecies incarnata among Creeping Willow. On the edges of dune slacks subspecies incarnata
and subspecies coccinea can be found growing together.
O 25 May, Norfolk. Subspecies incarnata.
29/1/09 12:32:59
284
GENUS DACTYLORHIZA
M 1. 19 June, Norfolk. Subspecies incarnata. 2. 19 June, Norfolk. Subspecies incarnata. 3. 17 June, Norfolk. Subspecies
coccinea. 4. 15 June, New Forest. Subspecies pulchella. 5. 23 May, Co. Clare. Subspecies cruenta. 6. 14 June, Suffolk.
Subspecies ochroleuca.
D. i. cruenta Flecked Marsh Orchid has

mid to dark pink flowers that lack the purple
tones of subspecies pulchella. Around 30-65%
of plants have dark purplish-brown spots on
one or both surfaces of the leaves, the spots
becoming denser towards the tips of the leaves
and sometimes merging. If present, the spots
on the underside of the leaves are paler, smaller
and sparser. The stem, bracts and ovaries are
more strongly washed purple than in subspecies

pulchella and are sometimes spotted or flecked
darker. There are sometimes ring markings
on the bracts, too (almost never spotted in
pulchella), and the lateral sepals more often
have ring markings. The lip is more distinctly
three-lobed, about 4.5-7.5mm long x 4.5-9mm
wide, the side-lobes are moderately reflexed
and the margins of the lip are often slightly
29/1/09 12:33:10
EARLY MARSH ORCHID
285
wrinkled (crenate). The lip has broader, bolder

loop markings but few dashes within the loops
and these markings often spread over most of
the lip (extending over no more than two-thirds
of the lip in pulchella). The stem is slender,
the leaves are held stiffly erect, and the spike
is looser and less crowded than in subspecies
incarnata, especially in Scottish plants.
Controversy surrounds this subspecies and
doubts are sometimes expressed as to whether
British and Irish plants are true cruenta as
found in Europe. Bizarrely, it is sometimes
suggested that while Scottish plants are the
real thing, Irish plants are not, being merely
spotted-leaved variants of subspecies pulchella.
D. i. ochroleuca is often relatively large,
sometimes as much as 70cm tall, with a broad
stem and large leaves that are usually held very
erect. The bracts are large, the lowest usually
larger than 30mm x 20mm. The flowers are
creamy to pale yellow with a relatively large lip,
around 9mm wide x 7mm long, that is deeply
three-lobed with notches on the side-lobes.
This subspecies has often been confused with
pale-flowered variants of other subspecies, i.e.
with var. ochrantha, but it has larger and more
distinctly lobed flowers.

As well as the various subspecies, several
varieties have been named. In principle, each
different subspecies could be found as each
different variety, giving around 15 different
combinations.
Var. punctata is usually rather small with a
few small dots on the leaves towards the tip. It
is rare but has been recorded from Yorkshire,
the New Forest and the Isle of Coll, among
subspecies incarnata and pulchella.
Var. leucantha has unmarked pure white
flowers but is very rare.
Var. ochrantha has unmarked pale creamyyellow flowers with the lip pale yellow or rarely
green at the base (it is similar to subspecies
ochroleuca but has smaller and less distinctly
three-lobed flowers). It is scarce and usually
found among subspecies pulchella, although
M 13 June, Norfolk. Subspecies incarnata. Early Marsh

Orchid can be robust, with many proportionally small
flowers.
it has also been recorded among incarnata,

coccinea and cruenta.
A hyperchromic variant with an excess of
pigmentation and a solidly dark lip has also
been recorded, albeit rather rarely.
BIOLOGY
There is no nectar, and pollination is by
bumblebees that are duped into visiting the
flowers. The process is efficient and seed-set
is good.
29/1/09 12:33:11
286
GENUS DACTYLORHIZA
Hybrids
Most or all of these hybrids are sterile and
only found as isolated individuals or in small
groups.
D. x kernerorum, the hybrid with Common
Spotted Orchid, has been found scattered
throughout Britain and Ireland. It is sterile.
D. x carnea, the hybrid with Heath Spotted
Orchid, has been recorded scattered in Britain
and Ireland but is rare. It is probably sterile.
D. x wintoni, the hybrid with Southern Marsh
Orchid, has been found scattered throughout
the range of the latter but is rare. It is sterile.
D. x latirella, the hybrid with Northern Marsh
Orchid, has been recorded throughout the
range of the latter but is scarce.
D. x dufftii, the hybrid with Pugsleys Marsh
Orchid, has been found in northwest Wales,
Yorkshire and Co. Wicklow. It is rare.
D. x aschersoniana, the hybrid with Irish Marsh
Orchid and Hebridean Marsh Orchid, has
been recorded in Co. Limerick and the Outer
Hebrides respectively. This name is no longer
valid for both species due to changes in the
classification.
X Dactylodenia vollmannii, the hybrid with
fragrant orchid (in the broad sense), has been
recorded twice from Cornwall.

M 14 June, Suffolk. Subspecies ochroleuca. Grows in calcareous, spring-fed fens.

Seed probably germinates in the spring, and the
first leaves appear a year later. Early estimates
of the period between germination and first
flowering were as long as 16 years, but these
were made by counting growth marks in tubers
and other structures that were thought to
represent a years development. This technique
is probably unreliable, and in cultivation plants
flower aged four or five years. Individual plants
have been recorded living for up to 25 years
after their first appearance above ground.
The specific name incarnata means fleshcoloured. Most of the subspecific names also
refer to the flower colour; pulchella is Latin for
beautiful, coccinea means crimson or scarlet (as
in the dye produced from galls on the Kermes
Oak), cruenta is Latin for blood-coloured and
ochroleuca comes from the Greek for yellowishwhite.
The Early Marsh Orchid and its relatives are
often known as the diploid marsh orchids. This
is because their chromosome count is 2n = 40
compared with a count of 2n = 80 in the
tetraploid marsh orchids, a group that includes
all the other British and Irish marsh orchids.
Although of little use to the orchid-lover in the
field, it represents a fundamental division.
29/1/09 12:33:12
EARLY MARSH ORCHID
HISTORY AND
CONSERVATION
For a long time this species was amalgamated
with the other marsh orchids. The first British
record that distinguished the Early Marsh
Orchid was in the Flora of Shropshire published
in 1841.
Early Marsh Orchid is frequently found
in small numbers and is often the first of the
marsh orchids to vanish if its habitat dries out
due to drainage or other changes. Overall, the
species has gone from 43.5% of its historical
range in Britain and 39% in Ireland. In lowland
areas the species declined significantly as
riverside meadows were ploughed, drained
or otherwise improved, the water table fell
due to abstractions, or indeed meadows were
abandoned to scrub.
287
Past and present occurrence of Early Marsh Orchid (all

subspecies) in Britain and Ireland (based on presence or
absence in 10km squares of the National Grid; data from
the New Atlas).
Britain
Ireland
1,192
331
671 (23.5%*)
202 (20%*)
% lost, 1500-1969
29.5%
30.5%
% lost, 1970-1986
14%
8.5%
43.5%
39%

1500-1999
current range
% lost, total
Similar factors have led to the near

extinction of the creamy-flowered subspecies
ochroleuca, which is treated in the British Red
Data Book and is a BAP priority species. It was
first recorded from Roydon Fen (near Diss) in
Norfolk in 1936 and was subsequently found
at several fens in the Waveney Valley on the
Norfolk-Suffolk border and at Chippenham
Fen in Cambridgeshire. It is now extinct at all
sites with the exception of Chippenham Fen,
where it has declined from around 30 plants in
1990 to a single clump, and one of the Waveney
Valley fens in Suffolk, where it was rediscovered
in the late 1990s, and there are currently
around a dozen flowering plants.
The red-flowered dune subspecies
D. i. coccinea has suffered declines due to the
scrubbing-over of sand dunes and also coastal
development but still occurs in very large
numbers at some favoured sites. The spottedleaved Flecked Marsh Orchid D. i. cruenta is
also treated in the British Red Data Book and
is classified as Endangered. It is only found
at three sites in Scotland which, although
remote, may suffer from overgrazing. It is
rather commoner in Ireland but with the rapid
changes occurring there, including The Burren
region, it could be vulnerable to development
and improvement.
M 14 June, Suffolk. Subspecies ochroleuca. A well as

being a pale cream colour, the lip is very distinctly threelobed.
29/1/09 12:33:14
288
GENUS DACTYLORHIZA
frOG Orchid
Vulnerable, BAP
Dactylorhiza viridis
Formerly: Coeloglossum viride; Other names: Bracted Green Orchis (North America)
The English names of many orchids make sense, and it is easy to see the man in a Man
Orchid or the monkey in a Monkey Orchid. Not so this species, for although the name has
been in use since the 17th century it is hard to see the frog in a Frog Orchid; the two-lobed
lip could resemble hind legs, and the hood may look like a frogs body, but to anyone except
the most imaginative any real resemblance to a frog is fanciful. Frog Orchid grows in short
grassland, frequently on chalk or limestone, but is inconspicuous and often hard to find. It
has undergone a serious decline and is now missing from much of its former range.
Identification
The small size, generally greenish or reddishpurple flowers, tight hood and rather plain,
strap-shaped lip are distinctive.
Similar species
Of the vaguely similar species, Man Orchid has
long and narrow arms and legs on its lip and
Common Twayblade has a smaller flower, again
with arms and legs.
Habitat
In southern Britain this species is confined
to well-drained short grassland on chalk or
limestone, especially the slopes of ancient
earthworks, abandoned quarries, old chalk
and lime pits and spoil heaps. It is tolerant of
grazing and trampling but cannot compete
with rank vegetation. In the past it was also
found in damp or wet permanent pastures and
meadows in southern England, but this habitat
has almost entirely vanished, taking the Frog
Orchid with it. In the north and west it is found
in a wider range of short-grass habitats, often
damp, and on both calcareous and neutral soils,
including limestone pavements, rocky ledges,
road verges, railway embankments, upland
flushes, mountain pastures, coastal grassland,
machair and dune slacks. It occurs from sea
level to 915m (Glen Doll, Angus).
O 29 June, Hampshire. Frog Orchid is shy and the flowers typically face downwards without showing their face.
29/1/09 12:33:15
FROG ORCHID
289
Flowering period
Early June to early August, sometimes from
late May or even until early September, but
usually peaking in late June and early July. Once
pollinated, the flowers persist for a long time,
although the lip will have withered.
Range
Frog Orchid is widely distributed in Britain
and Ireland, including the Inner and Outer
Hebrides, Orkney and Shetland, but has
declined severely in much of lowland Britain.
In southern England it is now almost confined
to the chalk districts of Hampshire, northeast
Dorset and Wiltshire, extending very locally
into the Berkshire Downs, Chilterns and the
South Downs in Sussex. In the Midlands it is
more-or-less restricted to the Peak District. It
is very local in Wales, lowland Scotland and
the southern half of Ireland (and absent from
southernmost Ireland from Co. Cork to Co.
Wicklow) but more widespread in northern
England, upland Scotland and the northern
half of Ireland. World range: Frog Orchid
has a circumpolar distribution and is found in
Europe, Asia and North America. It is mostly
confined to the temperate regions but extends
north to Iceland, the Faeroes, northernmost
Scandinavia and Alaska. It ranges furthest
south in the mountains, to Spain, Italy, Greece,
the Crimea and Caucasus (and adjacent
1987-99
1970-86
pre 1970
M 13 July, Dorset. Frog Orchid varies from overall green

to generally very reddish.
northeast Turkey); in Asia to Central Asia and

the eastern Himalayas; in North America to
Washington, New Mexico, Iowa and North
Carolina.
How to find it
Although variable, this species is often small,
green, rather inconspicuous and easy to tread
on. It can easily be overlooked; the first record
in Cornwall for over 50 years was when over
4,000 flower spikes were found near Colliford
Reservoir in 1986. It is also rather local in
29/1/09 12:33:17
290
GENUS DACTYLORHIZA
29/1/09 12:33:20
FROG ORCHID
occurrence, and the number of flowering plants

can vary greatly from year to year. As with
Musk Orchid, another short-turf specialist, it
can be easiest to spot when scanning from a
low-level vantage point. In southern England
small numbers are found at several chalk
grassland reserves from Dorset east to Sussex.
Notable sites in Scotland include North
Berwick and Crail golf courses which hold
populations of many thousands.
DESCRIPTION
Height: 4-45cm but mostly 5-15cm and only
very exceptionally over 25cm.
Stem: Green, on some plants washed reddishpurple towards the flower spike. There are
one or two brown, leafless sheaths at the
extreme base.
Leaves: There are three to five dark green,
strap-shaped sheathing leaves, sometimes rather
broad, with the two lowest leaves being the
largest and bluntest. They are held at around
30-45 above the horizontal. Higher on the
stem there are also several narrower and more
lanceolate non-sheathing leaves. The rosette
appears in the autumn and overwinters, and
there may be a relatively large number of nonflowering plants; these have fewer leaves.
Spike: Rather loose and often irregular in
shape, with the ovaries variably twisted and
the flowers therefore pointing in different
directions. Mostly there are 5-25 flowers but
sometimes only two, and particularly large
plants may have up to 50.
Bract: Green, variably washed reddish-purple
and lanceolate in shape; the lower bracts may
be rather longer than the flowers (up to a
maximum of twice the flowers length), but
towards the tip of the spike the bracts
become shorter.
Ovary: Green, variably washed reddish-purple,
spindle-shaped, six-ribbed and twisted.
Flower: Green or yellowish-green, variably
291
washed brown or reddish-purple, in some the

whole flower may be reddish-purple. The sepals
are dark green variably washed reddish-purple,
roughly oval to oval-triangular in shape. The
upper sepal is a little smaller than the lateral
sepals. The petals are pale green, washed
reddish-purple around the edges and strapshaped, much smaller and narrower than the
sepals. The sepals and petals form a tight hood
over the column, with the petals often showing
as much paler slots between the sepals. The lip
is usually paler and greener than the rest of the
flower and is even paler and yellower towards
the base, with reddish tones usually confined
to the tip and edges. It is tongue-shaped and
may broaden slightly towards the tip, with two
small terminal lobes that have a notch between
them and a third smaller lobe within this
notch. There is also a thickened ridge down the
centre of the lip, and nectar is secreted from
two hollows formed by the raised and curved
margins of the lip at its base. The lip may hang
downwards or be folded backwards below the
ovary. The spur is very short (1-2mm), almost
O 29 June, Hampshire.
P 29 June, Hampshire. It is not easy to see the frog in a
Frog Orchid.
29/1/09 12:33:21
292
GENUS DACTYLORHIZA
hemispherical in shape and rather colourless.

The anthers are washed purple, and the pollinia
are club-shaped and pale yellowish with a
very lobed surface. The flowers are faintly
honey-scented.
Subspecies
None.

On average shorter, stouter and more reddishbrown in the dry grassland habitats of southern
England and taller and greener in the north.
Some plants, including the flower spike, may be
entirely yellowish-green.
Var. longibracteatum has unusually long bracts
and is rather taller and more robust. It has been
found in northern England.
BIOLOGY
Unlike the other Dactylorhiza orchids, the
flowers produce nectar, and this is secreted
into the short spur. A variety of insects visit
the flowers, including small beetles and wasps,
especially ichneumons. These alight on the lip
and are directed by its central ridge to either
side; as they approach the spur, the pollinia
are stuck to their head by the viscidia which lie
on either side of the spur. Once on the insects
head it takes about 20 minutes for the pollinia
to rotate forward. They are then in position to
make contact with the stigma of the next plant
to be visited. Self-pollination is also reported
to occur. Seed-set is variable, with the capsules
maturing rapidly and containing around 1,2505,000 seeds. This species only occasionally
reproduces vegetatively.

The aerial stem grows from a pair of tubers
that are forked into three or four finger-like
lobes. They are typical of the genus Dactylorhiza
although rather small.
The period between germination and
flowering is short. The first green leaves appear
one to three years after seed has germinated
and a flower spike is usually produced in that
first year above ground. Frog Orchid is shortlived with a rapid turnover of the population.
Many plants are monocarpic and die after just
one year above ground, although some may live
and flower for at least seven years. In a study in
Holland the half-life of a population averaged
1.5 years and varied from 1.0 to 2.4 years (the
half-life is a measure of the life expectancy
of an orchid after its first appearance above
ground and marks the point at which 50% of
the population which emerged in any given year
have died). Occasionally, plants may be dormant
underground but not for more than one year.
Hybrids
D. x mixtum, the hybrid with Common
Spotted Orchid, is rare but has been found at
widely scattered localities.
D. x conigerum, the hybrid with Heath Spotted
Orchid, has been found rarely (the two parent
species do not occur together as frequently).
D. x viridella, the hybrid with Northern Marsh
Orchid, has been recorded from Co. Durham
and the Inner and Outer Hebrides.
X Dactylodenia jacksonii, the hybrid with
fragrant orchid (in the broad sense) has been
noted sporadically but widely in England.

The specific name viridis means fresh-green or
youthful.
Until recently the Frog Orchid was named
Coeloglossum viride, and indeed the genus
Coeloglossum contained just this one species.
However, genetic and biochemical techniques
have shown that Coeloglossum is sufficiently
similar to the genus Dactylorhiza for the Frog
Orchid to be transferred to it. This explains
why the Frog Orchid has such a propensity
to form hybrids with the other members of
the genus Dactylorhiza, the spotted orchids
and marsh orchids. The chromosome number
is 2n=40, and Frog Orchid is one of the
more primitive Dactylorhiza. (The old name
Coeloglossum means hollow-tongue, a reference
to the short conical spur at the base of the lip.)
29/1/09 12:33:21
FROG ORCHID
293
Generally commoner in the north, especially

in Scotland, Frog Orchid is abundant at a
few favoured localities with hundreds or even
thousands of flower spikes. It is rather local,
however, even in Scotland and very much so in
the rest of Britain.
Past and present occurrence of Frog Orchid in Britain and

1500-1999
current range
Britain
Ireland
964
214
381 (13%*)
99 (9.8%*)
% lost, 1500-1969
49%
47%
% lost, 1970-1986
11.5%
6.5%
% lost, total
60.5%
53.5%
M 13 July, Dorset. Frog Orchid has recently joined the

spotted and marsh orchids in the genus Dactylorhiza but
any similarity is not obvious.
HISTORY AND
CONSERVATION
Despite its inconspicuous nature, the Frog
Orchid was known to early naturalists and
was first recorded in Hertfordshire in 1650
by William How in his Phytologia Britannica
natales exhibens Indigenarum Stirpium sponte
emergentium (A British botany presenting
the origins of native wild plants): Orchis
BatrachitesBy Barkway, Dr. Johnsons MS.
The species was then listed in John Rays flora
of Cambridgeshire in 1660 and noted by
Christopher Merrett growing near Lewes in
Sussex and, in many places about Oxford in his
Pinax of 1666.
This species has declined throughout

almost the entire range in the British Isles
and in much of Europe too. The biggest losses
have been in England, especially the Midlands
and East Anglia, where the ploughing-up of
old pastures on neutral or chalky boulder-clay
soils appears to have been a major factor as
well as the development of scrub and rank
vegetation on what had been short grassland.
Frog Orchid is extinct in Essex, Norfolk,
Middlesex, Huntingdonshire, Worcestershire,
Warwickshire, Lancashire, Monmouthshire,
Breconshire, Radnorshire, Pembrokeshire,
Montgomeryshire, the Isle of Man,
Dumfries-shire, Wigtownshire, Berwickshire,
Kincardineshire and Dunbartonshire. It was
last seen in Kent in 1988, and just one site
each remains in Suffolk, Cambridgeshire,
Lincolnshire and Herefordshire. Not
surprisingly, it has recently been classified
as Vulnerable by the UK governments
conservation advisers.
29/1/09 12:33:22
294
GENUS DACTYLORHIZA
cOmmOn SpOtted Orchid

Dactylorhiza fuchsii
Formerly: Orchis fuschii, Dactylorchis fuschii
Aptly named, this species is both common and has boldly spotted leaves. Indeed, it is the
commonest and most widespread orchid in the British Isles and is found in a very wide
variety of habitats from open grassland to woodland and fen. An opportunist, it can colonise
new areas relatively easily and quickly and can sometimes appear in very large numbers;
the massed ranks of thousands of Common Spotted Orchids on a road verge or in an
abandoned quarry or chalk pit has to be one of the most spectacular sights of the summer.
Common Spotted Orchid is the county flower of West Lothian and Linlithgowshire and
the Hebridean Spotted Orchid (subspecies hebridensis) is the county flower of the Western
Isles.
Identification
Identification is usually straightforward. The
leaves are marked all over with solid dark
spots or bars, the flowers are various shades of
pink or lilac (and are often very pale or almost
white), and the lip is divided into three roughly
equal lobes and decorated with bold dark lines
and loops. It is, however, very variable: the
leaves may be heavily or lightly marked; plants
with unspotted leaves are not uncommon;
and the flowers vary from white to rather dark
pinkish-purple. It tends to be tall, attenuated
and pale-flowered in shady woodland sites
but shorter and more compact in the open. It
thrives best in damp or wet habitats; plants on
dry chalk grassland can be very petite while
those in exposed, windswept sites may be short
and squat.
M 6 July, Norfolk.The leaves are typically well-spotted.
Similar species
Early Purple Orchid has spotted leaves, but
the lip is a very different shape and does not
have bold darker lines and loops. It also flowers
rather earlier in the spring.
Heath Spotted Orchid can be very similar
to Common Spotted Orchid but is separated
by its fewer, rather narrower, more obviously
keeled and more pointed leaves, with the lowest
leaf not significantly shorter or blunter than
the remainder. The spots on its leaves are often
small and rounded, compared to the larger and
29/1/09 12:33:23
COMMON SPOTTED ORCHID
bolder marking of Common Spotted Orchid,

and plants with unspotted leaves are more
frequent. The lip of Heath Spotted Orchid
is broader and the lobes are very unequal in
size. The central lobe is rather small and barely
longer than the side-lobes, and the incisions
(sinuses) between the lobes cut much less than
halfway to the base of the lip. The lip is often
more faintly marked, with dots and fine dashes
rather than bold loops and lines, and the spur is
longer and rather finer.
Marsh orchids mostly have darker and
more purple flowers, although there is some
overlap, and often have unspotted leaves. Both
Common and Heath Spotted Orchids can
be distinguished from the marsh orchids by
having a spur that is slender and parallel-sided
rather than short, fat and bag-like (but beware
hybrids). The stem of the spotted orchids is
usually solid and unsquashable, whereas that of
some of the marsh orchids is obviously hollow.
However, Common Spotted Orchids growing
in wet habitats may have hollow stems, so this
is not always a safe distinction. The number of
non-sheathing leaves on the upper part of the
stem may also be helpful; often two to six in
Common Spotted (but sometimes just one) and
one or two in most marsh orchids.
Hybrids are common, particularly with
Southern and Northern Marsh Orchids.
Identification of hybrids requires a good
knowledge of the range of variation of both
species and even then is not always certain.
First generation hybrids usually show hybrid
vigour and may be obviously large, sometimes
ridiculously so. Hybrids with marsh orchids
often have the fat, conical spur of the marsh
orchid parent rather than the thin spur of the
spotted orchid but in subsequent generations
back-crossed with the parent species, or in hybrid
swarms, these distinctions will break down.
Habitat
Common Spotted Orchid favours a very wide
range of habitats. It occurs on dry grassland,
from heavily grazed downland swards to ranker
and scrubbier sites, and also thrives in damper
295
M 22 June, Norfolk. A petite, few-flowered plant.
conditions in wet meadows, the machair of

northwest Scotland, marshes, fens and dune
slacks, including alkaline flushes in otherwise
unsuitable moorland. The species is also found
in woodland, usually in the better-lit areas
along rides, clearings and edges, although it
can tolerate quite deep shade where it may
persist in a non-flowering state. Common
Spotted Orchid is usually found on alkaline or
neutral soils but it can grow in slightly acidic
conditions, such as among grass and Heather
on less acid areas of heathland. An opportunist,
it is capable of colonising new sites, including
29/1/09 12:33:24
296
GENUS DACTYLORHIZA
man-made habitats such as abandoned gravel

and chalk pits and industrial waste sites (e.g.
fly ash tips and alkaline Leblanc waste). It is
often found on roadside verges and cuttings,
including motorways, on railway embankments
and sometimes on lawns, and can quickly build
up in numbers to form substantial populations.
Recorded up to 530m above sea level in
Cumbria (Garrigill) and 600m in Perthshire
(Loch na Lairige).
Flowering period
Mid-May to early August, being earliest in
open, sheltered localities in the south (where
generally best around mid-June) and later in
woodland sites and in the north.
Range
Found throughout Britain and Ireland,
including the Isle of Man, Inner and Outer
Hebrides and Shetland, and also the Channel
Islands. However, it is absent from most of
Cornwall, much of north Devon and from
large areas of northern and northeast Scotland.
Formerly recorded from Orkney and the Isles
of Scilly. World range: Widespread in Europe
away from the Mediterranean lowlands, south
to Spain, northern Italy and Greece, north to
Scandinavia and Finland and east to Russia
and Siberia. The precise boundaries of the
distribution are obscure due to confusion with
Heath Spotted Orchid.
1987-99
1970-86
pre 1970
M 20 June, Norfolk. Typical coloration.
How to find it
In much of Britain and Ireland this is the
commonest orchid, and it is often found in large
numbers. Nevertheless, there are large tracts of
countryside that are not graced by its presence.
DESCRIPTION
Stem: Pale green, sometimes lightly washed
with purple towards the tip, grooved and
usually solid (but can be hollow, especially in
larger plants in wetland habitats).
Leaves: Green, usually marked all over with
elongated solid dark spots and blotches but
sometimes unmarked. There is a relatively
small, oval, blunt-tipped leaf at the base of the
29/1/09 12:33:27
297
green, often with a violet tinge.

Flower: Various shades of pale lilac, pink or
purplish-pink or sometimes almost white. The
sepals are oval-lanceolate with the lateral sepals
asymmetrical, marked with lines and spots
and usually held spreading horizontally (but
they may be closer to 45). The upper sepal
and the petals, which are a little shorter, form
a hood over the column. The lip is a flattened
oval in shape, 8-12mm wide, divided into three
lobes by deep wedge-shaped incisions (sinuses)
that extend about halfway to the base. The
side-lobes have broad rounded tips that are
sometimes toothed with roughly parallel sides.
The central lobe is slightly narrower and more
triangular in shape but as long or longer than
the side-lobes. The lip is held flat or with the
side-lobes lightly depressed. It is usually paler
and whiter towards the centre and is marked
with bold dashes and broken dark pink or dark
reddish-purple lines which often form a pattern
of concentric double loops (occasionally just
dots and dashes). The spur is about half as
long as the ovary to roughly the same length,
narrow, cylindrical, slightly tapering and either
straight or very slightly curved. The pollinia are
pale brownish-pink to purple, or sometimes
yellowish. The flowers are faintly scented.
M 6 July, Norfolk. A relatively pale spike.
stem and the three to six (sometimes two to

seven) sheathing leaves are crowded together
above this; they are broad, up to 4cm or even
5.5cm wide, and lanceolate. The two to six
(sometimes one to nine) non-sheathing leaves
higher on the stem become narrower and more
bract-like towards the spike.
Spike: Variable in shape but often pyramidal or
conical as the flowers begin to open, becoming
longer and more cylindrical later on, with 20-70
flowers, exceptionally 150.
Bract: Green, sometimes washed purple,
lanceolate and finely pointed; the lowest are
longer than the ovaries but higher on the spike
they are about equal in length.
Ovary: Cylindrical, six-ribbed, twisted and
Subspecies
D. f. fuchsii is the commonest and by far the
most widespread subspecies.
D. f. hebridensis Hebridean Spotted Orchid is
characterised by dark flowers and a broad lip.
It is found on the machair and similar coastal
habitats in the Outer Hebrides, northwest
Scotland, Shetland and western Ireland (Co.
Donegal, Co. Galway and Co. Kerry). It tends
to flower relatively late and often occurs in large
numbers. In some areas overlaps and mixes
with subspecies fuchsii.
Hebridean Spotted Orchid is small and
stocky, 8-20cm tall (range 6-40cm), with
the upper stem usually washed purple and
the leaves heavily spotted. The spike is often
pyramidal or conical and densely packed, the
lip is usually over 9.5mm wide (8-15mm), and
29/1/09 12:33:28
29/1/09 12:33:31
299
the flowers are often deep rose-pink or reddishpurple (rarely white) with the spur relatively
long at 7-8.5mm. It is sometimes treated as
a separate species by European authors but
its distinctiveness has been exaggerated and,
conversely, it is also sometimes treated as merely
var. hebridensis.

Woodland and grassland plants are sometimes
separated as distinct varieties but the
differences between the tall leafy plants found
in the shade of woodland and scrub and the
shorter plants of sunny grassland (the so-called
var. trilobata) seem to be purely a product of
their different growing conditions.
Var. alpina is a dark-flowered variant of
subspecies fuchsii. The dark flowers recall
those of subspecies hebridensis but they are
smaller, with the lip usually less than 9.5mm
wide and with narrower side-lobes. It is found
in Scotland, both at inland sites and on the
machair on the Hebrides and northwest coast
(including some populations in the Inner
Hebrides traditionally called hebridensis). It is
also present at inland sites in northern England
and possibly also Wales and western Ireland.
Var. cornubiensis is very similar to subspecies
hebridensis and essentially differs only
statistically. It is a little bigger and more
robust, with a larger and looser flower spike,
longer bracts (often over 8mm long) and a
narrower lip. The lateral sepals are rarely held
horizontally, and the spur is stouter. Confined
to Cornwall, where it is found on the north
coast on stabilised dunes at Lelant golf course
and cliffs near St Ives and at Tintagel.
Var. albiflora has unmarked white flowers
(rarely washed cream or pink) and unspotted
leaves. It is widespread but rather uncommon.
Var. okellyi has narrow leaves and flowers
which are sometimes a little smaller or broader
than typical plants and often almost white or
creamy with very faint pink or lilac markings,
although in some areas a substantial proportion
(often the majority) have darker and heavier
markings; the leaves are spotted or unspotted.
M 10 June, Norfolk. A particularly richly-coloured spike.

O 11 July, Norfolk.
It is found along the western seaboard of

Ireland, on the Isle of Man and in northern
Britain; individual white-flowered plants within
these populations are well-nigh identical to var.
albiflora (and vice-versa).
A lot of controversy surrounds okellyi with
some European authors treating it as a distinct
species and other botanists treating it merely
as a poorly defined variety. The classic okellyi
of the literature always has white, almost unmarked flowers and is said to be confined to
29/1/09 12:33:33
300
GENUS DACTYLORHIZA
western Ireland (especially The Burren region of

Co. Clare and Co. Galway), the Isle of Man and
the west coast of Kintyre in western Scotland
(with single records for Tiree and Sutherland).
However, in The Burren and elsewhere these
classic white-flowered okellyi are just part of a
population of plants with a variable flower colour.
Var. rhodochila is an attractive hyperchromic
variant with an excess of pigmentation. The
lip is solidly reddish or purple, usually with a
narrow paler pink or white border. The leaves
may be more heavily spotted or in extreme cases
entirely washed purple. It is widespread but rare.
BIOLOGY
A wide variety of insects has been recorded
carrying off pollinia from this species, but
studies in Austria and Poland have identified
beetles, especially longhorn beetles, as the major
pollinators. In common with other members
of the genus Dactylorhiza, Common Spotted
Orchid does not produce nectar. Insects may
therefore be deceived into visiting the flowers,
although Charles Darwin suggested that
visiting insects feed on sap sucked from inside
the wall of the spur. This hypothesis has since
been disputed, and it has been suggested that
they feed instead on tiny sugar-rich papillae
that are present in a zone from the middle of
the lip down into the spur. A third option is
that bumblebees receive no reward and are
deceived into pollinating the flowers but that
honeybees are able to access a sugary liquid
produced on the surface of the stigma.
Whatever the mechanism, pollination is
efficient and around 50-90% of flowers produce
seed. Notably, twice as many of the lower
flowers, which open first, produce ripe capsules,
compared with the upper flowers, which open
later on. This suggests that as the season
progresses insect pollinators become scarcer
or lose interest as there is little or no reward.
Vegetative reproduction is also possible.

The length of the period between germination
and flowering in the wild is unknown but
in cultivation it can be as little as two years
although it is more often four or five years.
It is thought that the plant can flower for
several years in a row or remain as a dormant
non-flowering rosette if conditions become
unsuitable.
Hybrids
M 6 July, Norfolk. A few-flowered spike.
Sterile hybrids are likely to be found either

singly or in very small numbers, whereas even
partial fertility can result in large numbers of
hybrids which in turn can show a great deal of
individual variation.
29/1/09 12:33:34
301
D. x transiens, the hybrid with Heath Spotted

Orchid, is widespread but scarce and can be
hard to identify with certainty. It is sterile. See
also Name and classification.
D. x kernerorum, the hybrid with Early Marsh
Orchid, has been found scattered throughout
Britain and Ireland. It is sterile.
D. x mixtum, the hybrid with Frog Orchid, is
rare but there are widely scattered records.
D. x venusta, the hybrid with Northern Marsh
Orchid, occurs throughout the range of the
latter and can be common. It is partially fertile
and can back-cross with either parent species
to produce a range of characters in a hybrid
swarm.
D. x grandis, the hybrid with Southern Marsh
Orchid, can be found not only where both
species occur together but also in the absence
of one or even both parents. The pollen is
highly sterile and seed production is very low.
Nevertheless this partial fertility allows backcrossing with both parents, creating hybrid
swarms which may be very persistent. It is
probably the commonest orchid hybrid in
southern Britain.
D. x silvae-gabretae, the hybrid with Pugsleys
Marsh Orchid, has been recorded from
northwest Wales, Yorkshire and Ireland. The
pollen is highly sterile.
D. x braunii, the hybrid with Irish Marsh
Orchid, has been recorded in Co. Clare.
(The name may be changed following the
elevation of Irish Marsh Orchid to the status
of a full species.)
X Dactylodenia st-quintinii, the hybrid with
fragrant orchid, is uncommon and scarce but
has been found widely through Britain and
Ireland. All three species of fragrant orchid may
be involved.

The specific name fuchsii commemorates
Leonard Fuchs (1501-66), a German botanist
who is also celebrated in the name Fuchsia.
Common Spotted Orchid is closely related
to Heath Spotted Orchid and although they
M 10 June, Norfolk. Presumably Common Spotted Orchid

x Southern Marsh Orchid. Such hybrids can sometimes
occur where one of the parents is absent.
29/1/09 12:33:36
302
GENUS DACTYLORHIZA
between the two are not absolute. Common

Spotted Orchid is generally found on calcareous
or neutral soils and Heath Spotted Orchid
on acid soils. However, there is some evidence
that spotted orchids growing on intermediate,
neutral or slightly acidic soils do tend to be
intermediate in appearance.
HISTORY AND
CONSERVATION
Although spotted orchids had been known
from the time of Turners Herball of 1562,
this species was not distinguished from the
Heath Spotted Orchid until 1915 when it
was described by the English botanist George
Claridge Druce.
This is the most widespread species of
orchid in the British Isles and is currently
present in 67% of the total available 10km
squares in both Britain and Ireland. There
has been some decline but Common Spotted
Orchid is so versatile that it can usually hangon somewhere within any particular 10km
square. The New Atlas figures are thus likely
to conceal a significantly greater fall in the
number of actual populations. Notably, much
of the decline in Britain appears to be recent
and it could be that a fall in the number of
populations is increasingly expressing itself as
a decline in the overall range. At least some of
the losses are compensated for, however, by the
colonisation of new areas, especially in recent
man-made habitats.
M 30 June, Norfolk. A densely-flowered spike; as all the
flowers open the spike typically becomes more cylindrical.
are clearly distinct in Britain, France and

Scandinavia they are rather similar in central
Europe and are considered to be the same
species by many European authors. They are
usually quoted as differing in the number
of chromosomes, with Common Spotted
Orchid having 2n=40 and Heath Spotted
Orchid 2n=80, but various studies have given
chromosome counts of 2n=40, 2n=60 and
2n=80 for both species (i.e. diploid, triploid
and tetraploid). Even in Britain the differences
Past and present occurrence of Common Spotted Orchid

Atlas).
Britain
Ireland
2,219
774
1,913 (67%*)
673 (67%*)
% lost, 1500-1969
7%
12%
% lost, 1970-1986
7%
1%
% lost, total
14%
13%

1500-1999
current range
29/1/09 12:33:37
HEATH SPOTTED ORCHID
303
heath SpOtted Orchid

Dactylorhiza maculata
Formerly: Orchis ericetorum, Dactylorchis ericetorum
Closely related to Common Spotted Orchid, this species replaces it in more acid, heathy
habitats. It is similarly very widespread but has a northern and western bias to its range and
is now absent from much of the Midlands and southeast England. The flowers are usually
rather pale pink with delicate markings, and the lip is broad, resembling a frilled skirt. It can
occur in large numbers and gives a real splash of colour to otherwise sombre tracts of moor
or bog.
Identification
A dainty and very attractive orchid, variable in
stature but often just 10-20cm high, especially
in exposed or relatively dry sites. The flowers
are very variable but are often a pale shade of
pink or pinkish-lilac and marked with fine dots
and dashes, the ground colour frequently fading
to whitish as they age. The broad lip resembles
a voluminous petticoat and its side-lobes are
much larger than the small, tooth-like central
lobe and often have serrated or frilly edges.
The leaves are usually marked with numerous
dark spots.
Similar species
Common Spotted Orchid is rather similar
but can be separated by its broader and flatter
leaves with the lowest leaf usually significantly
shorter than the rest with a broad, rounded tip.
In both species the leaves are usually spotted
but in Common Spotted the spots are often
larger, bolder and usually elongated into short
bars rather than being rounded. In Common
Spotted the lip is more deeply lobed with the
incisions (sinuses) between the lobes cutting
around halfway to the base of the lip and the
three lobes more equal in size; the central lobe
is at least half the width of the side-lobes and
usually a little longer too. Finally, in Common
Spotted Orchid the lip is marked with loops
and lines rather than dots and the spur is
shorter, thicker and tapers slightly towards
the tip.
P 5 July, Norfolk.
29/1/09 12:33:39
304
GENUS DACTYLORHIZA
Marsh orchids are usually easily separated

as most have darker and more boldly marked
flowers than is typical in Heath Spotted
Orchids. Sometimes, however, Heath Spotted
may have darker and more purplish flowers
and, especially if the leaves are unspotted,
may superficially resemble a marsh orchid.
In case of doubt check the spur: long, slender
and roughly parallel-sided in Heath Spotted
and squat and sack-like in marsh orchids.
Also check the number of non-sheathing
leaves: often two to five in Heath Spotted (but
sometimes just one) and one or two in most
marsh orchids. Finally, the stem of the spotted
orchids is usually solid and unsquashable and
that of at least some of the marsh orchids is
obviously hollow.
Hybrids are common, particularly with
Northern and Irish Marsh Orchids. First
generation hybrids usually show hybrid vigour
and may be obviously large. Hybrids with
marsh orchids often have the fat, conical spur
of the marsh orchid parent rather than the thin
spur of the spotted orchid. Later generations
of hybrids, if back-crossed with one or other of
the parent species, become harder and harder
to determine.
Habitat
Heath Spotted Orchid is found in a wide
variety of habitats. It favours heathland and
moorland, especially the damper, more peaty
areas on the margins of valley bogs, flushes and
mires and the raised and slightly drier areas
within Sphagnum bogs; suitable areas are often
picked out by the greyish foliage and dusty-pink
flowers of Cross-leaved Heath. It is also found
on acid grassland (for example rhos pastures
in Wales and culm grassland in southwest
England), damp unimproved meadows and hill
pastures. It usually grows on mildly acidic soils
and in areas of chalk or limestone bedrock;
suitably acidic conditions may be provided by
pockets or blankets of peat or areas of drift
(superficial deposits of sands and gravels).
It may sometimes grow in neutral or even
slightly alkaline marshes and fens and in such
habitats can be very robust. Heath Spotted
M 15 June, New Forest. A petite but richly marked plant.
Orchid is not as tolerant of shade as Common

Spotted and is much less commonly found in
light woodland and even then usually on the
edge or along rides rather than in dense shade.
Recorded from sea level up to 915m (Ben
Lawers, Perthshire).
Flowering period
Mid-May to July, sometimes from the second
week of May or rarely to early August. Typically
it flowers earliest in drier situations and later
in wetter habitats. The flowering time relative
to Common Spotted Orchid varies in different
29/1/09 12:33:40
1987-99
1970-86
pre 1970
parts of the country and can be later or earlier,

perhaps depending on their respective habitat
in the area.
305
leaves crowded towards the base of the stem.

These are long, narrow, keeled and pointed,
the largest up to 20mm wide (more rarely
to 25mm) with the tips sometimes hooded.
There are also two to five (one to seven) nonsheathing leaves and these become more bractlike towards the flower spike. All the leaves are
green, variably marked with solid dark spots
which are rounded or only slightly elongated;
the leaves are fairly frequently unspotted.
Spike: Usually rounded or pyramidal, even
when all the flowers are open, with 5-20
flowers, sometimes up to 60 on robust plants.
Bract: Green, often washed purple around the
edges and towards the tip, lanceolate, tapering
to a fine point, one-and-a half to two times
longer than the ovary on the lower flowers but
around equal in length higher up.
Ovary: Green, cylindrical, six-ribbed and
twisted.
Range
Found throughout Britain and Ireland,
including the Scottish Islands, Isle of Man and
the Channel Islands, but with a predominantly
northern and western bias to the distribution.
World range: Western Europe, north to
Iceland, the Faeroes, Scandinavia, Finland and
the Baltic States, east to Russia and south to
northern Portugal and Spain, Italy, the former
Yugoslavia and Bulgaria. As with Common
Spotted Orchid, dispute over the exact
definition of the species makes it hard to be
precise about the boundaries of the range.
How to find it
Usually common in suitable habitats, the delicate
pale pink blooms stand out among the grasses
and heathers that typically surround it.
DESCRIPTION
Height: 4-50cm but usually 10-25cm. Plants
have been noted up to 75cm tall but such
robust individuals are probably hybrids.
Stem: Pale green, washed purple towards the
tip and slightly ridged.
Leaves: There are one or no basal leaves (or
sheaths) and two to four (one to five) sheathing
M 21 July, Norfolk. A relatively few-flowered spike, with

sparse but bold markings.
29/1/09 12:33:43
306
GENUS DACTYLORHIZA
that are usually rather fine and form a pattern

of concentric loops or circles but are sometimes
bolder and form the clear double loops more
typical of Common Spotted Orchid. The spur
is slender, cylindrical or slightly tapering, about
half as long as the ovary to roughly the same
length and straight or slightly decurved. The
flowers are faintly scented.
Subspecies
M 21 July, Norfolk. The flower often has a blousy look

and the central lobe of the lip is much smaller than the
side lobes.
Flower: Usually whitish to pale pink or

pale lilac, sometimes purplish-pink, with
dark reddish markings. The lateral sepals
are lanceolate, asymmetrical, marked with
darker spots and lines and held horizontally
or drooping. The upper sepal and petals are
slightly shorter and form a hood over the
column. The lip is shorter than wide (around
5-9.5mm long x 6.5-13mm wide), roughly
circular or circular-diamond shaped. It is
divided into three lobes by relatively shallow,
triangular notches (sinuses). The side-lobes
are much larger than the central lobe and their
edges may be wavy or toothed. The central lobe
is small, triangular, rather tooth-like and does
not usually project beyond the side-lobes (if
it does, it rarely projects by more than 1mm).
The lip is held flat, slightly dished or with
the side-lobes slightly deflexed. It is whitish
to pink, sometimes deep lilac-pink, variably
marked all over with darker dots and dashes
British and Irish plants are conventionally

treated as belonging to subspecies ericetorum
which also occurs along the Atlantic seaboard
of Europe from northern Portugal to northern
Scandinavia and Finland. The typical
subspecies, D. m. maculata, is found in the
remainder of continental Europe.
Within the British Isles there is much
variation but it does not show any consistent
pattern, and no other subspecies can usefully be
recognised, even though some European authors
state that subspecies maculata and/or subspecies
elodes occur here (and some split the spotted
orchids into many different species and treat
subspecies maculata and ericetorum as different
species, stating that both occur in Britain).
Plants from Rhum in the Inner Hebrides
with dark flowers intermediate in shape
M 21 July, Norfolk. A boldly marked plant.
29/1/09 12:33:44
307
between Heath and Common Spotted Orchids

were described as subspecies rhoumensis. But
the population on Rhum is very variable and
such dark, three-lobed flowers occur elsewhere.
It is not worthy of recognition.

Var. concolor has an excess of pigmentation.
It has a solidly dark reddish-purple lip and
the leaves may also be washed purple. It is
extremely rare, with just one or two records.
Var. leucantha has pure white flowers. It is
widespread but rare.
BIOLOGY
A variety of insects have been observed carrying
pollinia, especially bees and flies, and seed
production is good (45% of flowers set seed
in one Scottish study). The mechanism is
probably similar to Common Spotted Orchid,
although the pollinia are relatively small,
possibly an adaptation to smaller and less
robust pollinating insects.

The pattern of development is probably similar
to Common Spotted Orchid, with seedlings
appearing above ground less than two years
after germination and the first flower spike
three years later.
Hybrids
Sterile hybrids are likely to be found either singly
or in very small numbers, whereas even partial
fertility can result in large populations of hybrids
displaying a great deal of individual variation.
D. x transiens, the hybrid with Common
Spotted Orchid, is widespread but scarce. It is
highly sterile.
D. x carnea, the hybrid with Early Marsh
Orchid, has been recorded scattered in Britain
and Ireland but is rare. It is probably sterile.
D. x conigerum, the hybrid with Frog Orchid,
has been found rarely but widely.
D. x hallii, the hybrid with Southern Marsh
Orchid, occurs infrequently as the parent
species favour different habitats. It is fertile and
M 20 June, Norfolk. The white flowered var. leucantha is

widespread but rare.
can therefore create hybrid swarms.

D. x formosa, the hybrid with Northern
Marsh Orchid, occurs throughout the range
of Northern Marsh Orchid. It is fertile and
frequently back-crosses with the parents to
form hybrid swarms. This is probably the
commonest hybrid orchid in northern Britain
and Ireland.
D. x jenensis, the hybrid with Pugsleys Marsh
Orchid, has been recorded from northwest
Wales, Yorkshire and Ireland. It appears to be
sterile.
D. x dinglensis, the highly fertile hybrid with
Irish Marsh Orchid, has been found widely.
29/1/09 12:33:46
308
GENUS DACTYLORHIZA
of disagreement over the number of species

involved. Some European authors treat them
both as the same species, and there is some
support for this argument as the differences
between the two are not as clear cut in parts
of Europe as they are in the British Isles. Even
in Britain the differences are not as absolute as
some field guides suggest (see also discussion
under Common Spotted Orchid).
HISTORY AND
CONSERVATION
M 20 June, Norfolk. Hybrid Heath Spotted Orchid x

Common Spotted Orchid. Surprisingly infrequent, such
hybrids are typically very robust.
X Pseudorhiza bruniana, the hybrid with
Small White Orchid, was recorded from
Orkney in 1977.
X Dactylodenia legrandiana, the hybrid with
fragrant orchid (presumably x Heath Fragrant
Orchid), has been found widely scattered
through Britain and Ireland.

The specific name maculata means spotted or
blotched.
Heath and Common Spotted Orchids are
members of a complex group of Dactylorhiza
orchids which have generated a great deal
The first British record of a spotted orchid

dates from 1562 when the second part of
William Turners Herball noted: There are
divers kindes of orchisone kindehath many
spottes in the leaf and is called adder grasse in
Northumberland.
One of the commonest and most
widespread orchids in the British Isles. There
has been some decline, however, especially in
lowland areas in England, and the species now
occupies only around three-quarters of its
historic range (and many of the losses in Britain
are relatively recent). Agricultural changes,
leading to the ploughing and improvement
of meadows and pastures and the destruction
of heaths have caused some losses, as has
the abandonment of heathland causing it to
scrub-over and turn into woodland; extensive
heath fires can also be damaging. The species
tolerates, and even benefits from, some grazing
but in northern Britain it has, like much native
vegetation, suffered from overgrazing.
Past and present occurrence of Heath Spotted Orchid in

1500-1999
current range
% lost, 1500-1969
% lost, 1970-1986
% lost, total
Britain
Ireland
2,025
687
1,587 (56%*)
531 (53%*)
12.5%
20.5%
9%
2%
21.5%
22.5%
29/1/09 12:33:47
SOUTHERN MARSH ORCHID
309
SOuthern marSh Orchid

Dactylorhiza praetermissa
Formerly: Orchis praetermissa, Dactylorchis praetermissa
This species often occurs in large numbers and can produce a spectacular show in riverside
meadows and pastures. Locally common and widespread in much of England (south and
east of a line from the Ribble to the Humber) and in south and southwest Wales, it can
tolerate a wide variety of conditions, not always marshy, and is even found on certain
favoured brownfield sites where a variety of chemical wastes have been dumped. In size and
flower colour it is rather variable, and it also hybridises readily with the spotted orchids,
making identification difficult at times.
Identification
In its typical form Southern Marsh Orchid is
distinctive. It is usually a robust orchid with a
stout stem, numerous rather broad, unmarked
green leaves and a spike of purplish-pink
flowers. The lip is broad and rounded with a
tooth-like central lobe; to judge the shape of the
lip it needs to be flattened out from below, using
a finger or a ruler. In its natural pose the lip may
be held flat, slightly dished or with the sides
turned downwards, and it is marked in a central
zone with fine dots and short dashes.
Southern Marsh Orchid is, however, rather
variable. Most plants have flowers that appear
a little dusty or washed-out, but they may be a
darker and more intense colour or have bolder
and darker markings on the lip. In addition,
two varieties of Southern Marsh Orchid are
markedly different: var. junialis, the so-called
Leopard Marsh Orchid has large ring-shaped
spots on the leaves and a boldly marked flower
and var. bowmanii has a very prominently threelobed lip in various shades of pink with bold,
dark dashes and loops (recalling Common
Spotted Orchid in lip shape and pattern).
Similar species
Early Marsh Orchid is usually easily
distinguished by flower colour alone (very pale
pink, deep red or creamy). The various purpleflowered forms of Early Marsh Orchids can be
very similar in colour to Southern Marsh but
always have smaller flowers with a narrower
M 22 June, Norfolk. Southern Marsh Orchid can be tall,

slender and stately.
29/1/09 12:33:48
310
GENUS DACTYLORHIZA
lip that usually has the side-lobes clearly

folded downwards and prominent double-loop
markings.
Northern Marsh Orchid usually has darker
and redder flowers (deep magenta), a smaller,
more angular, diamond-shaped lip with heavier,
more blotched or looped dark markings and
often fewer non-sheathing leaves.
Pugsleys Marsh Orchid has a more slender
stem, fewer, narrower leaves, and a rather loose,
one-sided spike composed of a few, relatively
large flowers.
Hybrids are a major identification headache.
Southern Marsh Orchid frequently hybridises
with Common Spotted Orchid, and the hybrids
are usually tall and robust with the sheathing
leaves shorter and blunter than Southern
Marsh Orchid and the non-sheathing leaves
narrower and more pointed. Importantly,
the leaves are usually faintly spotted, and
the flowers are intermediate between the
two parents. These hybrids can be harder to
distinguish from Leopard Marsh Orchid as
this, too, has spotted leaves (see below for more

details). Southern Marsh Orchid also freely
hybridises with Pugsleys Marsh Orchid, and as
these hybrids are fully fertile, back-crossing can
lead to a range of plants intermediate between
the two parent species.
Habitat
Found in a variety of habitats on calcareous
to neutral or even acidic soils that are usually
(but not always) moist or wet. Typical habitats
include damp meadows, fens, marshes, the less
acid parts of bogs and wet heathland, dune
slacks, marshy gravel pits, road verges and also
old industrial sites, especially in northwest
England and the West Midlands (waste alkali,
colliery and fly-ash tips). It is also found in old
chalk quarries where compacted ground may
lead to water-logging, and it occasionally grows,
often in a dwarf form, on dry chalk grassland
and downs. Perhaps, surprisingly, Southern
Marsh Orchid cannot tolerate being submerged
for long periods, and winter floods lasting more
M 21 June, Suffolk. Often occurs in large numbers.

P 6 June, Norfolk.
29/1/09 12:33:50
311
29/1/09 12:33:54
312
GENUS DACTYLORHIZA
O 17 June, Norfolk. Southern Marsh

Orchid can be short and squat.
than a month may cause a severe decline or even

kill all the plants.
Flowering period
Late May to early July, occasionally from
mid-May to mid-July or even to early August.
1987-99
1970-86
pre 1970
North America, where it has been present for

many years at Tilt Cove, Newfoundland. It was
probably accidentally imported with mining
equipment from Europe in the early 1900s.
How to find it
Southern Marsh Orchid is fairly common and
sometimes occurs in large numbers, although
in the modern agricultural landscape it is
necessarily local. Big colonies stand out but
odd plants in marshy meadows may be easy to
miss; the presence of quantities of rushes is a
good clue.
DESCRIPTION
Range
Widespread in England roughly south of a
line from the Ribble to the Humber with a few
scattered colonies a little to the north of this. It
is also found in south and southwest Wales, the
Isles of Scilly and the Channel Islands. World
range: A European endemic found along the
Atlantic fringe of the continent from northern
France through Belgium, Holland, Germany
and Denmark to southwest Norway (and
possibly also Sweden and Finland); reported
recently from northern Italy. Introduced to
Height: 20-50cm, occasionally to 70cm or even

95cm (the largest plants possibly show some
hybrid influence). Dwarfed plants, just 10cm
high, may occur in marginal habitats such as
chalk grassland.
Stem: Stout (usually over 5mm in diameter),
hollow and green.
Leaves: Mid-green, sometimes slightly tinged
greyish-green. There are three to nine sheathing
leaves (typically four to six), held erect at 45 or
more and often slightly more crowded towards
the base of the stem. Oblong-lanceolate, the
leaves are more-or-less flat, lack a marked keel
and may be slightly hooded at the tips. The
largest are usually 2-3.5cm wide but they can
be as narrow as 1.5cm. There are one to three
narrower and more pointed non-sheathing
Spike: Rather crowded and occupies around
20% of the stem with 20-60 flowers, although
29/1/09 12:33:55
313
M 22 June, Norfolk. The markings on the lip are very variable but tend to be relatively fine, short dashes and dots
concentrated in the central zone.
large plants may have over 100. Conical as the

first flowers open, becoming cylindrical later on.
Bract: Green, often washed purple. The bracts
are around twice the length of the ovary and
project beyond the flowers (although usually
not as prominently as in Early Marsh
Orchid and less obviously so once the spike
is fully open).
Ovary: Green, variably washed purple,
cylindrical, six-ribbed and twisted.
Flower: Purplish-pink but variable in its exact
shade; sometimes a deep, intense colour but
usually a little washed-out. The lateral sepals
are lanceolate, asymmetrical and occasionally
blotched darker. They are variably positioned,
from nearly vertical over the flower to nearly
horizontal but usually held at around 45. The
upper sepal and petals are lanceolate, the petals
slightly shorter than the sepals, and together
they form a loose hood over the column. The
lip is 9-12mm long x 8.5-14mm wide (but
mostly more than 10mm wide). Overall it is
roughly circular in shape (if slightly squashed)
and subtly three-lobed, with broad, rounded
side-lobes and a small, often tooth-like central
lobe which usually projects slightly; there are

shallow incisions (sinuses) between the lobes.
The side-lobes turn downwards at the base but
are often level or turn upwards towards the
tip of the lip, which therefore appears dropshaped or like a well-rounded triangle in its
natural position. It is purplish-pink, becoming
paler towards the base and then white around
the mouth of the spur, and is marked with
fine dark spots and short dashes, usually
concentrated in the centre. The spur is twothirds to four-fifths the length of the ovary,
stout, tapering slightly to a blunt tip and may
be slightly down-curved.
Subspecies
None.

Var. junialis Leopard Marsh Orchid
has leaves marked with large oval spots or
sometimes bars which are hollow, and its
lip is marked with unbroken dark loops or
horseshoe-shaped marks rather than just dots
and dashes. It is found most often in the south
and southeast but is uncommon.
29/1/09 12:33:57
314
GENUS DACTYLORHIZA
M 22 June, Norfolk. A Leopard Marsh Orchid with bold

loop markings on the lip and ring-spotted leaves; it is easy
to confuse this variety with hybrid Southern Marsh Orchid
x Common Spotted Orchid.
At one time, Leopard Marsh Orchid was

thought to be a variation of the hybrid between
Southern Marsh Orchid and Common Spotted
Orchid which is commonly found whenever
the two species grow together. The occurrence
of relatively uniform populations of Leopard
Marsh Orchid, lacking hybrid vigour and often
without Common Spotted Orchids growing
nearby, suggests, however, that junialis is a

variety of Southern Marsh, and this has been
supported by genetic evidence.
Both Leopard Marsh Orchid and hybrids
can have hollow, ring-shaped spots on the
leaves, and the presence of such annular spots
is not diagnostic of Leopard Marsh Orchid,
although hybrids are more likely to have solid
dark spots. The following characters are useful
in distinguishing Leopard Marsh Orchid from
ring-spotted hybrids:
Size: Relatively small and slender, with fewer
leaves (averaging smaller and more slender than
normal Southern Marsh Orchids). Hybrids are
often robust, with many leaves, especially many
non-sheathing leaves.
Lip size: On average narrower, up to 10mm
wide. Hybrids have broader lips, 10mm or more
sometimes much more.
Lip shape: Often more obviously three-lobed
than typical Southern Marsh Orchids but not
as distinctly three-lobed as the hybrid.
Lip markings: Confined to the central zone,
within the heavy double loop, with just one or
two faint markings outside this area. In hybrids
heavy markings extend over most of the lip.
Var. bowmanii is relatively slender, with slightly
fewer, narrower leaves than typical Southern
Marsh Orchid (three or four sheathing leaves
and one to three non-sheathing leaves). The
flowers have a prominently three-lobed lip with
a very long central lobe and heavy line and loop
markings. Recorded from marshy areas or dry
grassland on acid soils in Devon, Dorset and
Hampshire. Initially described as a variety of
Pugsleys Marsh Orchid but clearly outside the
range of that species and genetic evidence places
it with Southern Marsh Orchid.
Var. macrantha has a lax flower spike and large
flowers (the lip usually much more than 7.5mm
long x 9.5mm wide) with an obvious central
lobe that exceeds the lateral lobes by more than
1mm. It is rare and found mostly at sites where
Southern Marsh and Pugsleys Marsh Orchids
occur together; it may be an intermediate
between the two.
Var. albiflora has white flowers but is very rare.
29/1/09 12:33:58
BIOLOGY
Pollination is efficient and large quantities of
seed are produced. Vegetative reproduction,
although far less important, may sometimes
produce groups of plants.

The period between germination and flowering
is two or three years.
Hybrids
D. x grandis, the hybrid with Common Spotted
Orchid, is rather frequent and can usually be
found where both species occur together (even
on dry chalk grassland and in the absence
of one or both parents). It is probably the
commonest hybrid orchid in southern Britain.
The pollen is highly sterile, and seed production
very low but even such partial fertility allows
back-crossing and the creation of long-lived
hybrid swarms.
D. x hallii, the hybrid with Heath Spotted
Orchid, occurs much less frequently than
D. x grandis as the two parent species favour
different habitats. It is fertile and can form
hybrid swarms.
D. x wintoni, the hybrid with Early Marsh
Orchid, has been recorded north to Lancashire
but is rare. It is sterile.
D. x insignis, the hybrid with Northern
Marsh Orchid, is rare and has only been
recorded from Wales. Despite their superficial
similarity and complementary distributions
in northern and southern Britain, Northern
and Southern Marsh Orchids may not be very
closely related.
x Pugsleys Marsh Orchid has been recorded
from Norfolk, Suffolk and Cambridgeshire.
Importantly this hybrid is fully fertile and
through a process of back-crossing with its
parents can form a full range of intermediates
(see also var. macrantha opposite).
X Dactylodenia wintoni, the hybrid with
fragrant orchid (in the broad sense), has been
found in southern England.
315

The specific name praetermissa means
overlooked and refers to the fact that the
Southern Marsh Orchid was not recognised
until 1914.
This species is one of the tetraploid marsh
orchids and the chromosome number is
2n = 80.
HISTORY AND
CONSERVATION
This species was overlooked (due to confusion
with Early Marsh Orchid) until 1914 when it
was described by the eminent English botanist,
George Claridge Druce.
Past and present occurrence of Southern Marsh Orchid
Atlas).

1500-1999
current range
% lost, 1500-1969
Britain
Ireland
1,014
812 (28.5%*)
12%
% lost, 1970-1986
8%
% lost, total
20%
Southern Marsh Orchid has inevitably

suffered quite significant declines as agricultural
changes and development, especially the
draining and ploughing of flood-plain meadows
and pastures, have destroyed its habitats. It has
vanished from 20% of the former range with
much of the loss relatively recent. This statistic
is likely to conceal a much greater decline,
however, as the number of populations within
each 10km square can fall significantly without
it registering as a loss. This problem applies to
all orchids but is perhaps most acute with those
species found in meadows and unimproved
pastures, habitats that were once so much a part
of the British landscape. The colonisation of
derelict industrial sites is small compensation
for such losses in the wider countryside.
29/1/09 12:33:59
316
GENUS DACTYLORHIZA
puGSleyS marSh Orchid

Dactylorhiza traunsteinerioides
Formerly: Narrow-leaved Marsh Orchid Dactylorhiza traunsteineri; also Dactylorhiza majalis
subspecies traunsteineri
The scarcest, most enigmatic and most localised of the marsh orchids (with the exception
of Hebridean Marsh Orchid), this species is endemic to Britain and Ireland. It occurs at
scattered localities throughout the British Isles, almost always in wet fens that are formed
by the upwelling of lime-rich but nutrient-poor springs. It hybridises freely, especially with
Southern Marsh Orchid, making certain identification difficult at times. Lapland Marsh
Orchid, a heavily marked plant found at a few localities in Scotland, has recently been shown
to be a variant of Pugsleys Marsh Orchid.
Identification
This delicate marsh orchid can be identified by
a combination of features which taken together
give it a subtle but distinctive appearance. It
is typically slender with a thin, weak stem,
and some plants even appear to need to be
supported by the surrounding vegetation. It
has a few narrow leaves that are often held
erect. The spike is made up of a few relatively
large flowers and is usually rather open and
one-sided, with all the flowers facing roughly
the same way. The upper stem and especially
the bracts are washed purple. The flowers
themselves are variable in colour but often
purplish-pink. The lip is usually obviously
three-lobed with the side-lobes turned
downwards and the central lobe projecting as
a prominent tooth (itself also sometimes
turned downwards). It is well-marked with
dark dots, loops and squiggles, the markings
often extending right to the edge of the lip.
Habitat is also a good clue, as the species is
almost always found in alkaline fens, often in
relatively low or sparse vegetation.
In order to confirm the identification it is
necessary to look carefully and in detail at the
orchids. It is best to examine a selection of
plants to get a feel for the range of variation
and work out some average measurements. It is
M 25 May, Norfolk. Typically slender, with narrow leaves
and a few, relatively large flowers (in this plant there are
11).
29/1/09 12:34:00
PUGSLEYS MARSH ORCHID
unlikely that a Pugsleys Marsh Orchid would

occur as a single isolated individual among a
colony of commoner species such as Southern
or Northern Marsh Orchid, and there should
always be a few candidates to look at. To be
Pugsleys Marsh Orchids, plants must have the
following characteristics:
1. Number of flowers: Relatively few, usually
just five to 14, but sometimes as many as 18,
particularly on the robust plants found in some
parts of Ireland.
2. Width of lip: At least 7mm wide and usually
wider, up to a maximum of around 13mm
(the average of several plants should be at least
9mm). The individual flowers are relatively
large.
3. Lip shape: Distinctly three-lobed, usually
with prominent incisions (sinuses) between
the lobes and with the central lobe longer and
rather narrower than the rounded side-lobes;
flatten the lip from below with a finger in order
to measure the width and accurately judge the
shape. In a natural position the lip often has
a characteristic triangular or deltoid shape,
narrower at the base and broadest at the tip; a
shape accentuated by the side-lobes being more
strongly reflexed at the base and flatter towards
the tip.
4. Total number of leaves: Three to five,
including up to one, rarely two, bract-like nonsheathing leaves; do not count the short basal
leaf hidden at the bottom of the stem. The
leaves are more-or-less evenly spaced, rather
than being gathered into a rosette.
5. Width of leaves: Relatively narrow, with
the second leaf from the bottom, which is
usually the widest, 6-15mm across (rarely to
18mm and usually averaging no more than
12mm wide).
6. Leaf spotting: Mostly unmarked but at a few
sites in Yorkshire and Ireland the plants have a
scatter of fine dark spots, and in Scotland the
subspecies lapponica (Lapland Marsh Orchid)
has very boldly marked leaves and bracts
Similar species
Early Marsh Orchid is usually easy to separate
from Pugsleys Marsh Orchid because most
317
M 28 May, Norfolk. The lip is usually marked with dots,

loops and squiggles, the markings often extending right to
the edge, and is typically three-lobed with the central lobe
projecting as a little tooth.This plant has 12 flowers.
Early Marsh Orchids in similar habitats will

have pale pink flowers. The purple-flowered
forms of Early Marsh Orchid are, however,
much more similar in colour and sometimes
have few flowers on a spike. Also, in Scotland
the rare spotted-leaved subspecies cruenta of
Early Marsh Orchid has been found in the
same localities as the similarly spotted Lapland
Marsh Orchid. But, whatever the colour of
the flower, Early Marsh Orchid always has a
more crowded flower spike and the individual
flowers have a rather smaller lip, less than
9mm wide. The markings are also different on
Early Marsh Orchid, with dots and dashes in
a central zone enclosed by bold double loops
and few if any markings outside this area. Their
leaves are broader and their spur is slightly
29/1/09 12:34:01
318
GENUS DACTYLORHIZA
tapering and slightly to moderately decurved.

Southern Marsh Orchid is typically taller
and more robust than Pugsleys Marsh Orchid.
It has a rather thicker stem and more leaves
which are broader and frequently more crowded
towards the base of the stem; there are often
two or more non-sheathing leaves. The spike
is larger and usually has many more flowers,
and these have a less obviously three-lobed
lip which is often held flat or slightly dished.
Small, petite Southern Marsh Orchids are more
similar to Pugsleys Marsh Orchid but still
have relatively unmarked flowers with less
prominent lobes.
Northern Marsh Orchid is rather closer
in size and stature to Pugsleys Marsh Orchid
but again has more leaves which are broader
and more crowded at the base of the stem. The

spike usually has more than 20 flowers and
the flowers themselves often have a slightly
smaller lip (sometimes less than 8mm wide
and averaging less than 9mm), which is usually
roughly diamond-shaped and held flat or
slightly dished. The flowers are also usually a
dark, velvety reddish-pink, a rather different
colour to those of Pugsleys Marsh Orchid.
Irish Marsh Orchid can be of a similar
size to Pugsleys Marsh Orchid and has a
three-lobed lip (although not so deeply lobed),
often with similar markings. It has, however,
more leaves which are often broader and more
crowded at the base of the stem and a denser
spike with more flowers.
Hybrids with Common and Heath Spotted
Orchids and Early Marsh Orchid should show
clearly intermediate characters. Much more
problematical is the hybrid Pugsleys Marsh
Orchid x Southern Marsh Orchid which is
apparently fully fertile. Through a process of
back-crossing this can form a complete range
of intermediates between the two species, and
at some localities in southern England there are
hybrid swarms and it is difficult to find any plants
that are completely pure Pugsleys Marsh Orchid.
Habitat
M 30 June, Norfolk. Tends to flower early, from mid-May

to mid-June, but as with all orchids, this can vary (12
flowers).
This species has very specific habitat

requirements and is only found where these are
met. It needs wet fens and flushes where the
ground water is base-rich due to the influence
of chalk or limestone. It grows in the wettest
areas among a relatively open community of
sedges, rushes and scattered reeds, rooting
into the mossy layer at the base of the taller
vegetation. It is almost always associated with
Black Bog-rush. Some of the sites are fairly
extensive but at others the correct conditions
are restricted to small patches within a larger
area of marshland. In Ireland it has been
recorded rarely from dune slacks. Pugsleys
Marsh Orchid is only exceptionally found in
very slightly acid conditions and in Scotland
Lapland Marsh Orchid may spread from its
base-rich flushes into nearby wet heathland,
which is slightly more acid. It is found from
29/1/09 12:34:02
319
sea level to 370m (Carperby, Wensleydale,

northwest Yorkshire), with subspecies lapponica
recorded up to 310m (Knapdale, Kintyre).
Flowering period
Plants come into flower in late May in the south
and west, exceptionally in early May, and some
may still be in flower in late June. The onset
of flowering can be a little later, towards early
June in the north, with Lapland Marsh Orchid
sometimes in flower until July.
Range
Found across the British Isles but the colonies
are very scattered and it is absent from most
areas. The strongholds are in Norfolk (and
formerly adjacent parts of north Suffolk),
northern Yorkshire, Anglesey and the Llyn
Peninsula in Caernarvonshire. Elsewhere
there are a few sites (often just one) in north
Somerset, north Hampshire, Berkshire,
Cambridgeshire, Co. Durham, Cardiganshire
and Pembrokeshire. Pugsleys Marsh Orchid
is similarly scarce and local in Ireland: in Co.
Carlow, Co. Wicklow and Co. Kildare in the
east and from Co. Limerick north to Co. Mayo
in the west, with just three sites in Northern
Ireland, all in Co. Antrim. In Scotland it is rare,
and both the typical form and Lapland Marsh
Orchid have a broadly similar distribution
in the west, in Kintyre, west Inverness-shire,
1987-99
1970-86
pre 1970
M 28 May, Norfolk. Although it had 16 flowers and a

total of 5 leaves, this well-marked plant was part of a
larger population of classic Pugsleys Marsh Orchids.
Ross & Cromarty, west Sutherland, the Inner

Hebrides (Rum, Raasay, Tiree) and Outer
Hebrides (Harris). World Range: Endemic to
Britain and Ireland.
How to find it
Pugsleys Marsh Orchid is easily overlooked.
This species is small, slender and often partially
concealed among relatively long grasses and
sedges. At some localities it can be abundant
but it may also be scarce and hidden amongst
hybrids. At many sites, especially in Scotland,
there may only be small numbers of flower
spikes. Two good clues for locating the species
are flowering period and habitat. Pugsleys
Marsh Orchid is one of the first marsh orchids
to come into flower and is almost always found
with Black Bog-rush. This is a rather large,
29/1/09 12:34:03
320
GENUS DACTYLORHIZA
tussock-forming member of the sedge family

that through the winter and into the spring
forms stands of distinctively grey vegetation.
Many of the sites for this species are rather
small and vulnerable to excessive disturbance
or, as at Beeston Common in Norfolk, have become dominated by hybrids. The best areas to
see it include Buxton Heath in Norfolk and
the Anglesey fens. In Ireland sites include
Slyne Head (Co. Galway) and Blacksod Bay
(Co. Mayo).
DESCRIPTION
Height: 6-40cm but usually 10-30cm with the
most robust plants in Ireland.
Stem: Green, usually washed purple towards
the flower spike, slightly hollow and rather
slender. The stem is often rather floppy or
bendy, averaging only 2.3-3.5mm in diameter
and is rarely more than 5mm across.
Leaves: Green, unspotted or occasionally with a
few, well-scattered, faint, purplish-brown spots
or transverse bars about 1mm in diameter;
spotting is only common in Yorkshire and
Ireland (see also Subspecies). The uppermost
non-sheathing leaf may be washed purple.
There is one short basal leaf and two to four
(rarely five) sheathing leaves. These are narrow,
strap-shaped, moderately keeled, sometimes
slightly hooded at the tip, well spaced along
the stem and often held erect at around 45.
The longest leaf is rarely more than 12cm long
(17cm in robust Irish populations) and the
broadest leaf is usually 6-15mm wide (rarely
to 18mm). There are also up to one (rarely
two) bract-like non-sheathing leaves on the
upper part of the stem, and the total number of
sheathing and non-sheathing leaves is three to
five (this count excludes the short basal leaf ).
Spike: Short and distinctly loose or lax with
rather few flowers, usually just six to 14,
occasionally as few as two or as many as 18 on
the largest plants. The spike is rather irregular
in shape but roughly cylindrical or tapering and
the flowers all face in more-or-less the same
direction (the spike is thus described as secund
or subsecund).
M 28 May, Norfolk.The spike is often very one-sided (11

flowers).
Bract: Lanceolate and long (the lower bracts

longer than the flowers), the bracts are green,
variably but often strongly washed reddishpurple.
Ovary: Green, strongly washed purple,
cylindrical and six-ribbed.
Flower: Often purplish-pink, similar in colour
to Southern and Irish Marsh Orchids, but
variable in intensity from rather pale to dark
and deep. The sepals are oval-lanceolate and
the petals slightly smaller and more oval.
The lateral sepals are asymmetrical, very
occasionally have darker markings and are
held at an angle that varies between 45 and
vertical but is usually closer to the latter. The
upper sepal and petals form a hood over the
column. The lip is relatively large, 6.5-9.5mm
long x 7-13mm wide (usually at least 7.5mm
long x 9.5mm wide). It is very variable in
shape but is usually a flattened oval divided
29/1/09 12:34:04
into three obvious lobes with the central lobe

rather narrower and variably longer than the
side-lobes. The lobes are separated by incisions
(sinuses) of various depths but sometimes
there are no obvious sinuses, just a prominent
central projection. The side-lobes are folded
downwards to a variable extent (and the tip
of the central lobe may also turn downwards),
giving the lip a rather drop-shaped outline,
narrowest at the base. The lip is a variable
shade of purplish-pink, becoming whitish at
the mouth of the spur, with dark lines, dots and
blotches in an irregular pattern that extends
more-or-less to the edges (sometimes forming
a double loop as in Early Marsh Orchid).
There is no correlation between the ground
colour of the lip and the extent and colour of
the markings; dark plants in Yorkshire may
have relatively small dot and dash markings
whereas paler flowers on Anglesey may have
long, heavy and contrastingly dark lines and
blotches. Conversely, pale flowers in southern
England may have relatively light markings.
The spur is long, straight and thick but tapers
slightly to a blunt tip.
321
Hebrides on Harris. It has also been identified

in Northern Ireland, on the Garron Plateau in
Co. Antrim. In at least one site Lapland Marsh
Orchid is found with typical Pugsleys Marsh
Orchids, and intermediates have been recorded.
Lapland Marsh Orchid was first discovered
at Knapdale (Kintyre) in 1967 and initially
identified as Pugsleys Marsh Orchid. This
identification was then revised and it became
Western Marsh Orchid (the form cambrensis,
now itself a subspecies of Northern Marsh
Orchid). Later still, in 1988, the plants
were identified as Lapland Marsh Orchid
D. lapponica a species found in the Alps,
Scandinavia and northern Russia which was
new to the British Isles. Ironically, the wheel
has now turned full circle and Scottish Lapland
Marsh Orchids have become a subspecies
of Pugsleys Marsh Orchid, confirming in a
roundabout way the initial identification.
Lapland Marsh Orchid may, in fact, be best
treated as a variety of Pugsleys Marsh Orchid,
Subspecies
D. t. lapponica Lapland Marsh Orchid has
bold, dark purplish-brown spots, blotches, bars
and rings on the upperside of the leaves and
bracts, which may also be edged with purple
(and the bracts are often spotted below). There
are up to two non-sheathing leaves which may
also have a few small marks on the underside.
The lip has intense dark purple or crimson
lines, rings and spots on the lip, sometimes
merging to form a dark patch in the centre, and
the lateral sepals are marked with dark rings,
elongated spots and dots. It averages slightly
shorter than the typical form at 6-18cm tall,
sometimes to 24cm, but Scottish populations
of typical plants are also small.
Occurs very locally in western Scotland,
in west Sutherland, Ross & Cromarty, west
Inverness-shire (including Ardnamurchan and
Morvern), Kintyre and, in the Inner Hebrides,
on Skye, Rum and Raasay and on the Outer
M 11 June, Kintyre. Lapland Marsh Orchid. A beautifullymarked plant, with bold loops and blotches on the lip and
lateral sepals.
29/1/09 12:34:06
322
GENUS DACTYLORHIZA
O 8 June, Kintyre.Lapland
Marsh Orchid. In classic
plants the leaves, bracts
and flowers are boldly
marked, but may occur in
mixed colonies with more
typical Pugsleys Marsh
Orchids; there appears to
be a north-south gradation
in the degree and depth
of markings, from relatively lightly-marked plants
with unspotted leaves in
southeast England to boldmarked plants in northern
and western Scotland.
rather than a subspecies; the Lapland Marsh

Orchid of Scandinavia remains, however, a
distinct species, D. lapponica.

Pugsleys Marsh Orchid is rather variable.
The size and shape of the leaves, presence or
absence of leaf spotting, number of flowers
and their colour and markings all vary, both
between colonies and within them. Yorkshire
plants with spotted leaves are sometimes
named var. eborensis. However, as there is
disagreement about the other characteristics
of this variety and as spotted leaves are also

found in Irish populations it does not seem
worthwhile to use this name. Rather small
Scottish plants from near Loch Maree (Ross
& Cromarty) with very pale flowers and
contrasting dark markings have been named
var. francis-drucei but may represent just one
small aberrant population. (Note that var.
bowmanii, originally described as a variety
of Pugsleys Marsh Orchid, is treated under
Southern Marsh Orchid.)
Var. albiflora has white flowers. It is very rare.
29/1/09 12:34:08
BIOLOGY
There is little specific information. Pollination
is probably undertaken by bees and although
other insects such as flies may remove the
pollinia they do not go on to pollinate other
orchids successfully. In studies of Pugsleys and
Lapland Marsh Orchids in western Scotland,
only 17-35% of flowers set seed, reflecting low
levels of pollination, probably due to a lack of
suitable pollinators and the poor weather. Each
ripe capsule that was produced contained about
3,000 viable seeds. Vegetative reproduction may
also be possible.

Hybrids
D. x silvae-gabretae, the hybrid with Common
Spotted Orchid, has been recorded from
Ireland, Anglesey and Yorkshire. The pollen is
highly sterile.
D. x jenensis, the hybrid with Heath Spotted
Orchid, has been recorded from northwest
Wales, Yorkshire and Ireland. It is probably
sterile.
D. x dufftii, the hybrid with Early Marsh
Orchid, has been found in northwest Wales,
Yorkshire and Co. Wicklow but is rare.
x Southern Marsh Orchid has been recorded
from Norfolk, Suffolk and Cambridgeshire.
Fully fertile, it can be common where the two
species occur together.

The specific name traunsteinerioides is derived
from traunsteineri, the scientific name of
Narrow-leaved Marsh Orchid, a European
species that is, in turn, named in honour of
Traunsteiner, a pharmacist who lived in Austria
from 1798-1850.
This species is a tetraploid marsh orchid
with the chromosome number 2n = 80. All
of the tetraploid marsh orchids are thought
to have originated from ancient hybridisation
events involving the ancestors of Early Marsh
Orchid and Common or Heath Spotted
323
Orchids. Pugsleys Marsh Orchid is intriguing

in that the same parent species have crossed on
several occasions, in the British Isles, the Alps,
Scandinavia and perhaps elsewhere, and each
hybridisation event has given rise to a similar
plant. After much debate, it was generally
accepted that British and Irish plants were the
same species as those in Europe and should
be known as Narrow-leaved Marsh Orchid
D. traunsteineri. However, recent genetic
research has shown that, although very
similar, they each have a distinct ancestry
and represent separate lineages and are best
treated as separate species. British and Irish
plants therefore become an endemic species, D.
traunsteinerioides, and should generate greater
interest among scientists and conservationists.
Note that Delforge (2005) gives the range of
this species as Britain, Ireland and northern
France but his concept of Pugsleys Marsh
Orchid appears to be rather different; for
example, he gives the number of flowers as
10-25, with extremes of 5-30.
HISTORY AND
CONSERVATION
This species was first found in Ireland by H.W.
Pugsley, who realised that it was something
different from herbarium specimens and plants
sent to him from Co. Wicklow. But, from
the start, the exact status of Pugsleys Marsh
Orchid was the subject of much debate. In
1936, Pugsley described his plants as a new
subspecies of a widespread European marsh
orchid and called it D. majalis subspecies
traunsteinerioides. In 1940, however, Pugsley
changed his mind and declared it to be an
entirely new species, D. traunsteinerioides. Then,
in 1953, the controversial British botanist
J. Heslop-Harrison demonstrated that it
belonged with another European species, D.
traunsteineri. This determination generated a
large and sometimes controversial literature,
but the consensus eventually emerged that it
was indeed the same as the European plant. Just
as the question seemed to have been settled,
however, genetic evidence showed that Pugsley
29/1/09 12:34:09
324
GENUS DACTYLORHIZA
had been right all along and that it should

be treated as a separate, endemic species, D.
traunsteinerioides. For more details of Lapland
Marsh Orchid see Subspecies.
Pugsleys Marsh Orchid is Nationally
Scarce in Britain and is specially protected in
Wildlife Order (NI). In addition, Lapland
Marsh Orchid (until recently treated as a
distinct species) is in the Red Data Book and is
considered Near-threatened in Great Britain
and fully protected under Schedule 8 of the
Wildlife and Countryside Act 1981.
Past and present occurrence of Pugsleys Marsh Orchid
(including Lapland Marsh Orchid) in Britain and Ireland
(based on presence or absence in 10km squares of the
National Grid; data from the New Atlas).
Britain
Ireland
85
40
63 (2.2%*)
13 (1.3%*)
% lost, 1500-1969
8.5%
37.5%
% lost, 1970-1986
17.5%
30%
26%
67.5%

1500-1999
current range
% lost, total
Lapland Marsh Orchid is mapped in the

New Atlas for 18 10km squares (seven of which
it shares with typical Pugsleys Marsh Orchid).
Pugsleys Marsh Orchid has declined
by 26% in Britain, with most of the
decline relatively recent and concentrated
in East Anglia. It is probably extinct in
Huntingdonshire and Northumberland. Losses
have been even greater in Ireland, at 67.5%,
and its numbers appear to have fallen there
more than almost any other orchid. There are
no recent records from Co. Cork, Co. Kerry,
Co. Tipperaray, Co. Offaly, Westmeath, Meath,
Co. Leitrim, Co. Cavan, Co. Louth and Co.
Fermanagh.
In the lowlands, Pugsleys Marsh Orchid
has disappeared due to the destruction and
drainage of its habitat. Even where wetlands
remain, often protected as SSSIs and reserves,
a general lowering of the water table (due
to drainage and abstraction) can cause a

degradation of the habitat. This has been
especially noticeable in East Anglia where the
species has declined sharply in the WaveneyOuse fens along the Norfolk-Suffolk border. A
more insidious threat is posed by hybridisation.
The hybrid with Southern Marsh Orchid is
fully fertile and at some sites hybrids have
come to dominate the population and Pugsleys
Marsh Orchid has been hybridised out (e.g. at
Beeston Common in Norfolk there were several
hundred Pugsleys Marsh Orchids in the late
1980s but by 2000 it was difficult or impossible
to find a single pure plant). At other sites,
hybrids apparently involving this species are
found where Pugsleys Marsh Orchid has never
been recorded, indicating a more extensive
distribution in the past. Similar threats face
the species throughout Europe as its highly
specialised habitat is degraded or destroyed.
In the uplands, sheep or deer graze most
sites. This can keep the vegetation open and
prevent a succession to scrub but it limits
the number of flowers produced, sometimes
severely so. When sites are not grazed the
number of flowering plants has increased,
although this may not reflect the number of
seeds that are able successfully to germinate.
More direct threats include forestry and
drainage, and only a few of the 30 or so
populations of Lapland Marsh Orchid are
protected in any way at all.
Any changes in the overall status of
Pugsleys Marsh Orchid are hard to assess. It is
a difficult species to identify with certainty and
also easy to overlook and is therefore still being
found and identified at new sites. These new
records do not indicate, however, an expansion
or increase as the species has presumably
always been present at these sites. This applies
particularly to Lapland Marsh Orchid, which
has been actively searched for since its discovery.
Conversely, Pugsleys Marsh Orchid is probably
recorded from some sites by mistake (i.e. it is
overrecorded).
29/1/09 12:34:09
HEBRIDEAN MARSH ORCHID
heBridean marSh Orchid
325
Vulnerable
Dactylorhiza ebudensis
Formerly: Dactylorhiza majalis subspecies occidentalis
This is one of four species of orchid endemic to the British Isles (the others being
Lindisfarne Helleborine and Pugsleys and Irish Marsh Orchids). It is known only from
Berneray and North Uist in the Outer Hebrides where it is locally abundant in the flowerrich machair.
Identification
Usually rather short and squat with a
distinctive purple tone to the flowers. The leaves
vary from more or less plain to solidly blackish
purple, but most are marked with rings or fine
blotches. In general, plants with darker leaves
have a darker ground colour to the flower.
Similar species
Only likely to be confused with Northern
Marsh Orchid, but distinguished as follows:
l Flower spike looser, less symmetrical and
often one-sided (typically symmetrical and
compact in Northern Marsh).
l Lip relatively larger, more spreading, and
more obviously three-lobed; the side lobes
seldom bend upwards at the edges in the

manner that gives the lip of Northern Marsh its
characteristic diamond shape.
l Flowers usually a clearer, purer purple (when
freshly opened the flowers of Northern Marsh
have a distinct crimson tone; on older flowers
the colour become more purple, but the dark lip
markings retain a deep crimson lustre).
l Leaves often (but not always) more heavily
marked, with ring-spots and blotches, and
also finely edged with purple (especially the
uppermost leaf ), even when not spotted.
Northern Marsh may have fine, regular spotting
on the leaves, but never extensive dark markings
(subspecies cambrensis of Northern Marsh
M 14 June, North Uist.The flowers are distinctly purple, and the leaves variably spotted.
29/1/09 12:34:11
326
GENUS DACTYLORHIZA
Orchid, which has heavily spotted leaves, has

been recorded elsewhere in the Outer Hebrides,
but the spots tend to be smaller, more even in
size and more evenly distributed over the leaf
surface than in Hebridean Marsh).
l Whole plant averages smaller and more petite.
stretching for c. 4 km south-westwards from

Newtonferry (Port nan Long), with a couple of
very small satellite populations a few km to the
west, and to the north, on Berneray.
Habitat
Height: 4.5-20cm.
Stem: Green, heavily washed with dark purple
towards the tip and hollow.
Leaves: There are two or three, sometimes
four, keeled, lanceolate sheathing leaves that
are relatively broad (10-14mm wide) and are
crowded towards the base of the stem with
the lower leaves tending to be held nearer the
horizontal than the vertical; the lowest leaf
is often rather shorter than the rest and the
single non-sheathing leaf is lanceolate. On most
plants the leaves are green, moderately marked
with brownish-purple rings or fine blotches
(especially the non-sheathing leaves). Markings
are concentrated towards the tips of the leaves
and on more heavily marked plants they begin
Found on the machair, a unique, species-rich,

coastal grassland habitat which develops on
low calcareous dunes in the wet and windy
environment of western Scotland and Ireland.
Flowering period
Late May to late June.
Range
Confined to North Uist and Berneray in the
Outer Hebrides. World range: Endemic to
Scotland.
How to find it
Only found on the north coast of North Uist,
where there is effectively a single metapopulation
DESCRIPTION
M 14 June, North Uist.Two plants, showing some of the range of variation in leaf markings.
29/1/09 12:34:13
HEBRIDEAN MARSH ORCHID
327
to coalesce; there is a continuous gradation

from plants with unmarked green leaves to
those with leaves that are solidly blackishpurple; spotting is occasionally also present
on the underside of the leaves. The leaves are
also finely rimmed with purple (especially the
uppermost), even when not spotted.
Spike: Short, compact and cylindrical, with
5-20 flowers.
Bract: Lanceolate, the lower slightly longer than
the flowers but becoming shorter towards the
top of the spike. They are green, heavily washed
purple or sometimes entirely purple and often
spotted darker.
Ovary: Ribbed, twisted and very heavily
washed purple.
Flower: Deep purple-magenta with darker
markings. The sepals and petals are elliptical,
with the lateral sepals slightly asymmetric and
held at 45 above the horizontal, or even higher,
whereas the upper sepal and petals (which
are shorter) form a hood over the column.
The lip is broader than long (6-8.5mm long x
8-12mm wide) and distinctly three-lobed with
the central lobe longer than the side-lobes. It is
flat or dished when the flower first opens but
the side-lobes may become slightly deflexed
with age. The lip is rich purple-magenta,
whiter towards the throat of the spur, and
heavily marked with dark purple spots and
dashes which are bounded within one or two
concentric dark loops. The spur is a little
longer than the ovary, thick, cylindrical and
slightly tapering.
Hybrids
Subspecies
This distinctive orchid first attracted attention

in the 1930s, but it was not until 1976 that
it was given a name. The first to be used was
scotica but this was not published in the correct
manner according to the International Code
of Botanical Nomenclature and was eventually
replaced by ebudensis.
Much of the range of the species lies within
the Machairs Rabach and Newton SSSI and
there seems to be no immediate threat to the
species, but such a limited distribution means
that Hebridean Marsh Orchid will inevitably
always be vulnerable.
None.

A hyperchromic variant has occasionally been
found. This has an excess of pigmentation
producing a solidly dark lip with a paler border.
BIOLOGY

No information.
x Northern Marsh Orchid? Plants showing

characters intermediate between these two
species have been found but may be merely
variant Northern Marsh Orchids; they are
fertile.

The specific name ebudensis is from the Latin
ebudes, Hebridean Islands.
The taxonomy of this species is confusing.
Until recently Hebridean Marsh Orchid was
grouped together with Irish Marsh Orchid
and the majaliformis form of Northern Marsh
Orchid into one species named Western Marsh
Orchid D. majalis. Thus, in many recent books,
the populations on North Uist will be found
under the heading Western Marsh Orchid
Dactylorhiza occidentalis and either listed as
subspecies scotica or combined with Irish
plants as subspecies occidentalis. Unexpectedly,
genetic studies have shown that Hebridean
and Irish Marsh Orchids should be treated
as distinct species (and thus Western Marsh
Orchid D. occidentalis does not occur anywhere
in Britain or Ireland). Hebridean Marsh Orchid
is one of the tetraploid marsh orchids and has
a chromosome number of 2n = 80. Genetic
evidence suggests that it originated in a cross
between Early Marsh Orchid, subspecies
coccinea, and Common Spotted Orchid,
subspecies hebridensis.
HISTORY AND
CONSERVATION
29/1/09 12:34:13
328
GENUS DACTYLORHIZA
nOrthern marSh Orchid

Dactylorhiza purpurella
Formerly: Orchis purpurella, Dactylorchis purpurella
This is the most commonly encountered magenta-flowered orchid of damp, marshy ground
in northern Britain and can sometimes be found in large numbers, giving a wow factor to
the landscape, even sometimes to derelict industrial sites. As its name suggests, this species
has a northerly distribution and occurs widely in northern England, northern Ireland
and Scotland, as well as in Wales. Although the intense reddish-pink flowers are usually
distinctive, the presence of hybrids with the spotted orchids can be a complication.
Identification
Relatively easy to identify. The flowers are
a deep velvety magenta with a distinctive
crimson tone that is most obvious when they
are freshly opened, while the dark markings
on the lip are always a lustrous dark crimson.
The lip typically appears diamond-shaped
with straight sides, especially to the base of the
diamond (the lip is actually three-lobed but
the side lobes are folded upwards at the edges
to form the diamond). The leaves are either
unspotted or have a few small spots, except
in parts of west Wales, northern England
and northwest Scotland where the subspecies
cambrensis has dark spots all over the leaves.
M 6 June, Cumbria. As with all the marsh orchids,

Northern Marsh Orchid is variable in size and structure
and can look quite stocky.
Similar species
Early Marsh Orchids with purple flowers are
quite similar to Northern Marsh Orchid, but
their flowers are, on average, slightly smaller,
with the lip rather narrower and usually
obviously three-lobed but with the side-lobes
sharply turned downwards (it may be unlobed,
but is never the distinctive diamond shape
of Northern Marsh); their lateral sepals are
usually held vertically over the flower.
Southern Marsh Orchid has more purplishpink flowers that are usually a little washed out
and lack the deep crimson tones of Northern
Marsh. Its lip is usually roughly circular, with
rather broad rounded side-lobes and a small
tooth-like central lobe. The dark markings are
often small, fine and confined to the centre of
the lip. Leopard Marsh Orchid (var. junialis
29/1/09 12:34:18
NORTHERN MARSH ORCHID
329
of Southern Marsh) has bolder, more looped

markings on the lip but also has heavily spotted
leaves, with the spots usually in the form of
hollow rings.
Irish Marsh Orchid has a distinctly
rounded, three-lobed lip that is on average a
little broader than Northern Marsh. It is more
purplish-pink and often also paler and less
intensely coloured. Up to 50% of Irish Marsh
Orchids have spotted leaves.
For distinctions from Hebridean Marsh
Orchid, see p. 325.
Hybrids with both Heath Spotted and
Common Spotted Orchids are relatively
common and can be abundant where the two
parent species grow together. Both hybrids
are very variable. They can have either spotted
or unspotted leaves and flowers intermediate
between the two parent species.
Habitat
Northern Marsh Orchid is found in damp or
wet sites on calcareous, neutral or slightly acid
soils, such as marshy fields, roadside verges,
lake margins, fens, marshes, flushes, seepages
along coastal cliffs, dune slacks, machair and
M 6 June, Cumbria. The leaves may have some fine dark

spots.
N 7 June, Northumberland. Northern Marsh Orchid can
be common in dune slacks.
29/1/09 12:34:27
330
GENUS DACTYLORHIZA
sometimes in less acidic peat bogs or in open,

damp woodland. It is an opportunist and has
colonised old quarries and urban brownfield
sites, including derelict industrial areas and
old waste tips, such as alkaline Leblanc waste
in Lancashire. Northern Marsh Orchid is
sometimes found on drier substrates such
as rubble and has been found in gardens.
Recorded up to 610m above sea level (Creag
Dhuba, Loch Ericht, west Inverness-shire).
Flowering period
Late May to late July (exceptionally from midMay) but mostly early June to mid-July.
Range
Widespread in north and west Wales, northern
England and Scotland, including the Isle of
Man, Inner and Outer Hebrides, Orkney and
Shetland. Occurs north of a line from Swansea
to Hull, but the population is sparser and more
scattered towards this southern boundary.
Also found in Northern Ireland, especially in
Co. Fermanagh, Co. Tyrone, Co. Derry and
Co. Antrim, and in adjacent Co. Donegal, but
otherwise rare and local in Eire, with almost
all the current sites on the east coast or on the
coasts of Co. Waterford, Co. Cork, Co. Galway
and Co. Mayo. There are a few isolated records
from southern England. In Oxfordshire a single
specimen was found near Wychwood in 1981
1987-99
1970-86
pre 1970
M 6 June, Cumbria.The lip is usually a distinctive diamond

shape.
and in Hampshire it was discovered in 1955 at

Moorgreen near Southampton where two small
colonies held up to 100 flowering plants at
times. It was thought that a cutting for the M27
in 1986 had destroyed these colonies but plants
were found again in the area in 1999. (It has
been suggested, however, that this species was
introduced to Hampshire.) World range: A
European endemic found along the northwest
fringe of the continent in western Norway
(north to around Trondheim), Sweden,
Denmark and the Faeroe Islands.
How to find it
Quite common and sometimes found in large
numbers.
DESCRIPTION
Stem: Green, washed purple towards the tip,
ribbed, often stout and slightly hollow.
Leaves: The four to eight lanceolate sheathing
leaves are slightly crowded towards the base of
the stem, broad (the largest 1.5-2.5cm wide),
hooded at the tip and held at up to 45 above
the horizontal. There are also one or two nonsheathing leaves higher on the stem (sometimes
none, rarely up to four) and a small basal leaf.
29/1/09 12:34:29
331
O 13 June, North Uist. Despite their names, Northern and Southern Marsh
Orchids are not each
others closest relatives and
are not analogous. Perhaps
as a consequence, hybrids
between these two species
are surprisingly scarce.
The leaves are mid-green to dark green,

unspotted or sometimes with a few very small
spots near the tip.
Spike: Dense, with ten to 40 flowers (sometimes as many as 80), oval to cylindrical in
shape and often flat-topped when fully open.
Bract: Green, often flushed purple, especially
towards the edges. The bracts are narrow and
lanceolate, the lower longer than their flowers,
the upper shorter.
Ovary: Green, washed purple (and sometimes
spotted on the ribs), cylindrical, six-ribbed and
twisted.
Flower: Deep velvety magenta, whiter around
the mouth of the spur. The sepals are roughly
oval and marked with irregular dark reddishpurple rings and lines. The petals are a little
shorter and unmarked. The lateral sepals are
held at around 45 and the upper sepal and
petals form a loose hood over the column. The
lip is relatively small, 5-8mm long x 6-10mm
wide. Although variable in shape, it is usually
shallowly three-lobed with a small central lobe
and and held flat or dished, with the margins
of the side lobes folded upwards to give the
lip a diamond shape. Heavy dark crimson
lines, dots and swirls cover much of the lip in
a concentric pattern. The spur is thick, conical,
slightly downward pointing and shorter than
the ovary.
29/1/09 12:34:33
332
GENUS DACTYLORHIZA
lobes slightly reflexed. It tends to be taller than

typical Northern Marsh Orchids with longer
and narrower leaves but with the spike smaller
in proportion to the whole plant.
This subspecies grows near to the sea in
western Scotland, often on damp grassy slopes,
in the Outer Hebrides, Kintyre, Ross and
Cromarty, Sutherland, Caithness and Orkney.
In west Wales it has been found rather rarely
in dune slacks and flood plain meadows on the
coast from Cardiganshire to Anglesey. It has
also been recorded from southeast Yorkshire
and recently from several sites in Cumbria, but
is otherwise only known from Denmark. It
has a chequered history and both Scottish and
Welsh plants started life as forms of Western
Marsh Orchid (a conglomerate of cambrensis
with Irish and Hebridean Marsh Orchids).
M 6 June, Cumbria. The lip markings are variable but

there tend to be heavy lines, dots and swirls more or less
all over the lip, often in a concentric pattern.
Subspecies
D. p. cambrensis (formerly majaliformis) has
leaves that are boldly spotted dull purple,
and bracts that are also spotted or washed
purple. The spots are distributed over the
whole surface of the leaf and can be sparse
or occasionally so numerous that they
coalesce into a solid dark patch; spotting
is absent in a small minority of plants. The
flowers average slightly paler and pinker than
normal and therefore the dark markings are
more contrasting. The lip is a little larger at
6-8.5mm long x 9-11mm wide and is rather
distinctly three-lobed, often with incisions
(sinuses) between the lobes and with the side-
Var. albiflora has white flowers.

Var. atrata is a hyperchromic variant with an
excess of pigmentation. The lip is solidly dark
magenta, lacking spots or lines but sometimes
with a well-defined paler margin. The leaves are
heavily spotted (rarely there is an overall purple
wash) with a few small spots on their lower
surfaces too. It is known only from damp fields
near an industrial area at Hartlepool which may
be contaminated with heavy metals.
Var. crassifolia has broad, fleshy leaves and a
large lip. It is rare.
Var. maculosa has many small dots all over the
leaves and sometimes a paler lip. Rare, it has
been recorded in southeast Scotland.
Var. pulchella has the lip marked with spots
and dashes rather than bold loops. It has been
recorded from Scotland.
BIOLOGY
No specific information, but it is probably
pollinated by bees, as bumblebees, especially
queens of Bombus terrestris, have been recorded
as frequent visitors. Pollination is efficient and
seed-set is good (52% of flowers setting seed in
one Scottish study).
29/1/09 12:34:37

Hybrids
Hybrids are frequently found, especially with
Common and Heath Spotted Orchids.
D. x formosa, the hybrid with Heath Spotted
Orchid, occurs throughout the range and is
probably the commonest orchid hybrid in
northern Britain and Ireland.
D. x venusta, the hybrid with Common
Spotted Orchid, occurs throughout the
range and can be common. It is variously
noted that the pollen is highly sterile and
little seed is produced or that this hybrid is
sometimes partially fertile. Whatever the case,
it is certainly capable of backcrossing with
either parent species or crossing with itself to
produce a range of intermediate characters in a
hybrid swarm.
D. x latirella, the hybrid with Early Marsh
Orchid, has been recorded throughout the
range but is rare.
D. x insignis, the hybrid with Southern
Marsh Orchid, has only been recorded in
Cardiganshire and Merionethshire. Despite
the relatively extensive area of overlap between
the two species in Wales and northern England
they tend to maintain a separate identity, even
in mixed colonies.
D. x viridella, the hybrid with Frog Orchid, has
been recorded from Co. Durham and the Inner
and Outer Hebrides.
x Hebridean Marsh Orchid? Plants showing
characters apparently intermediate between
the two species have been recorded but these
may be variant Northern Marsh Orchids; they
are fertile.
333
HISTORY AND
CONSERVATION
Northern Marsh Orchid was not distinguished
from the other marsh orchids until 1920
when it was described from plants found near
Aberystwyth by T. and T.A. Stephenson.
Inevitably there have been losses due to
habitat destruction (drainage, ploughing-up
of pastures, etc.) but the overall boundaries of
the range have remained stable. It has declined
by 21% in Britain, with many of the losses
comparatively recent, and by 46% in Ireland.
Past and present occurrence of Northern Marsh Orchid in
Britain
Ireland
1,202
151
946 (33%*)
81 (8%*)
% lost, 1500-1969
11%
33%
% lost, 1970-1986
10%
13%
% lost, total
21%
46%

1500-1999
current range
X Dactylodenia varia, the hybrid with one
of the fragrant orchids, has been found in
northern England, Scotland and Ireland.

The specific name purpurella means simply
purple. This species is a tetraploid marsh orchid
(2n = 80).
M 6 June, Cumbria.The lip shape can vary.This plant has

a projecting central lobe.
29/1/09 12:34:41
334
GENUS DACTYLORHIZA
iriSh marSh Orchid

Dactylorhiza occidentalis
Formerly: Dactylorhiza majalis subspecies occidentalis; Dactylorhiza comosa subspecies
occidentalis; Orchis occidentalis; Dactylorchis occidentalis
An Irish endemic, this species is fairly widespread and locally common in marshy or grassy
habitats although it has suffered some decline, especially more recently. As with all the marsh
orchids, it can be tricky to identify with certainty but its relatively early flowering season is
a great help.
Identification
The compact spike of purplish-pink, wellmarked flowers identifies this species as one
of the marsh orchids. It is rather variable, and
the dwarf plants of coastal grasslands can look
rather different to the robust populations in
damper and more sheltered spots. The leaves
may be spotted or unspotted. The lip has three
rounded lobes and is usually heavily marked,
often with double loops.
M 22 May, Co. Clare. Some have relatively paler and

pinker flowers and at least 50% of plants have unmarked
leaves.
Similar species
Separation of the marsh orchids can be difficult
at times, and in Ireland three species need to
be eliminated: Early, Northern and Pugsleys
Marsh Orchids.
Early Marsh Orchid is very variable and
includes the purple and pink-flowered subspecies
pulchella and cruenta, the latter often showing
bold spots on the leaves. But, whether their
leaves are spotted or unspotted, all Early Marsh
Orchids have rather smaller flowers with a
narrower lip that is usually less than 9mm wide
and narrower leaves that are held stiffly erect.
Northern Marsh Orchid tends to flower a
little later, has unspotted leaves or only a few
small spots, and flowers that are a darker and
deeper reddish-pink. Characteristically it has
an unlobed, straight-sided, diamond-shaped
lip that, on average, is a little narrower, no more
than 10mm wide.
Pugsleys Marsh Orchid is reasonably
distinctive, with a slender stem, a few relatively
large flowers held in an open, one-sided spike
and a lip that appears longer than wide.
29/1/09 12:34:46
IRISH MARSH ORCHID
Early Purple Orchid flowers at the same

time as Irish Marsh Orchid, sometimes in
the same grassy habitats, and also has heavily
spotted leaves. It is generally tall and slender,
however, with the leaves arranged in a floppy
rosette at the base of the stem. Its flowers are
much less heavily marked, with just a few dark
spots in the centre of the lip.
335
1987-99
1970-86
pre 1970
Habitat
Varied but mostly on neutral or slightly alkaline
soils and including wet meadows and pastures,
road verges, lough shores, the edges of acid
bogs, dune slacks, damp hollows on short,
closely grazed grassland and dry grassy slopes
near the sea.
Flowering period
Mid-May to mid-June, sometimes to July.
How to find it
Range
Generally fairly common and sometimes found

in large numbers.
In Eire there are concentrations in Co. Donegal

in the northwest, from Co. Galway south to
Co. Cork in the west and in Co. Wexford.
Otherwise, it is absent from much of Eire,
especially the east, apart from a cluster of
records from Co. Longford. Although mapped
for Northern Ireland, its occurrence there is
disputed. World range: Endemic to Ireland.
DESCRIPTION
Height: 10-30cm, rarely to 40cm.
Stem: Green, variably washed purple, ridged
towards the tip and slightly hollow.
Leaves: The four or five (sometimes only three)
lanceolate sheathing leaves are fairly broad,
M 25 May, Co. Clare. Irish Marsh Orchid can still be found in large numbers, even on roadside verges.
29/1/09 12:34:49
336
GENUS DACTYLORHIZA
29/1/09 12:34:52
IRISH MARSH ORCHID
M 21 May, Co. Clare. Some plants (perhaps especially in

coastal grassland) are short and squat.
O 24 May, Co. Clare.
usually 15-25mm wide (extremes 12-30mm).

They are grouped towards the base of the stem
and are usually held rather spreading, closer to
the horizontal than the vertical. There are also
two, sometimes three, non-sheathing leaves
higher on the stem. The leaves are green and in
about 50% of plants spotted with brownishpurple. The spots are generally concentrated
onto the outer half of the leaf and are variable
in shape; they may be rings or elongated into
transverse bars. Rarely, there are spots on
the lower surface of the leaf, too, or it has a
purple rim.
Spike: Crowded, with about 20 flowers
occupying 25-50% of the stem length.
Bract: Fairly long, green, variably washed
purple (sometimes strongly so) but only
occasionally spotted.
Ovary: Slender, twisted and ridged, green,
often heavily washed purple.
Flower: Purplish-pink with a well-marked
three-lobed lip. The lateral sepals are oval,
asymmetrical, variably marked darker with dots,
squiggles, or rings (sometimes unmarked) and
usually held closer to the horizontal than the
vertical. The upper sepal is oval, and the petals
are slightly shorter, narrower and more pointed.
Together they form a hood over the column.
337
The lip is purplish-pink, often rather deep but

becoming paler towards the centre and white
around the mouth of the spur (near to Southern
Marsh Orchid in colour but slightly deeper,
more intense and closer to purple). It is usually
wider than long, 7-11mm long x 9-14mm wide
and broadest around the middle. It has three
distinct rounded lobes with noticeable incisions
(sinuses) between them. The side-lobes are
variably reflexed, especially as the flower gets
older, and sometimes notched or with wavy
edges. The central lobe is smaller and sometimes
tooth-like and often projects a little beyond the
side-lobes. The lip is marked with heavy dots
and dashes, often enclosed within double loops
(sometimes concentric double loops), and there
may also be markings outside the loops. The
spur is roughly conical and straight.
Subspecies
None.

Var. kerryensis is found in southwest and
western Ireland, sometimes in pure colonies,
M 24 May, Co. Clare. The lip has three well-defined

rounded lobes marked with heavy dots and dashes, often
both within and without double loop markings.
29/1/09 12:34:54
338
GENUS DACTYLORHIZA
Marsh Orchid, which it resembles in flower

colour and pattern although the dark spots on
the lip are more numerous and more extensive.
A hyperchromic variant with an excess of
pigmentation and a solidly dark lip has, rarely,
been recorded.
BIOLOGY

Hybrids
D. x aschersoniana, the hybrid with Early
Marsh Orchid, has been recorded in Co.
Limerick. This name may be changed following
taxonomic changes.
D. x dinglensis, the highly fertile hybrid with
Heath Spotted Orchid, has been found widely.
D. x braunii, the hybrid with Common Spotted
Orchid, has been found in Co. Clare.
M 24 May, Co. Clare. The coloration varies from a deep

magenta to paler purple or deep pink.
but is scarce. It always has unspotted leaves and

bracts. Its flowers are, on average, paler and less
intense and the lip is often flat and marked with
dots and dashes that do not form a pattern of
loops. It also tends to be slightly shorter and
stockier and is said to flower a little later, into
July.
This form is sometimes treated as a distinct
species, D. kerryensis, due to its floral characters
and later flowering, and allied with Southern
The specific name occidentalis derives from

occidental, i.e. western, while the older name
majalis means in May.
Until recently this species was part of a
conglomerate of marsh orchids that took the
name Western Marsh Orchid Dactylorhiza
majalis. The conglomerate included D.
majalis from mainland Europe as well as Irish
Marsh Orchid, Hebridean Marsh Orchid
and the cambrensis subspecies of Northern
Marsh Orchid. Recent research, including the
analysis of DNA, has suggested that Irish and
Hebridean Marsh Orchids are unique endemic
species and that cambrensis from west Wales
belongs with Northern Marsh Orchid. This
means that D. majalis of Europe does not occur
anywhere in the British Isles.
This species is a tetraploid marsh orchid and
the chromosome number is 2n = 80.
HISTORY AND
CONSERVATION
Irish plants were recognised as being distinctive
29/1/09 12:34:56
IRISH MARSH ORCHID
339
O 25 May, Co. Clare. The

sides of the lip can be
sharply folded downwards,
and the markings are very
variable.
in the 1930s and were first given the name

occidentalis in 1935 by the British botanist
H.W. Pugsley.
Irish Marsh Orchid is vulnerable
to agricultural changes, especially with
European Union money funding agricultural
improvement in Eire, but so far losses appear
to be modest. The difficulties of marsh orchid
identification and the relatively low level of
botanical recording in Ireland may be obscuring
any changes in its status.
Past and present occurrence of Irish Marsh Orchid in Britain and Ireland (based on presence or absence in 10km
Britain
Ireland

1500-1999
159
current range
130 (13%*)
% lost, 1500-1969
9%
% lost, 1970-1986
9%
% lost, total
18%
29/1/09 12:34:58
Distribution
Growth pattern
Six species, found in central and southern

Europe, North Africa and the Middle East.
The aerial stem grows from a pair of tubers

and the annual cycle of the replacement of
tubers is similar to that of the genus Orchis
(see p.200).
Classification
Until recently Dense-flowered Orchid was
the only species in this genus, but recent
genetic studies have seen Neotinea enlarged
to embrace five more from the genus Orchis,
including Burnt Orchid. Paradoxically,
Neotinea is probably as close to the bee
orchids Ophrys, tongue orchids Serapias and
Lizard Orchid as it is to the genus Orchis.
Floral structures
There are two pollinia, each narrowing to
a short stalk which attaches it to one of the
viscidia, which in turn are enclosed in a
bursicle. There are two stigmas, placed on
each side of the mouth of the spur but joined
at the base.
of the stem and these will form separate plants
when the connecting stem dies off in the
autumn.
Name
The genus is named neo (= new) plus tinea,
after Vicenzo Tineo, a Sicilian botanist
(1791-1856). An alternative origin has been
suggested, New-Tinnea, due to its supposed
similarity to Tinnea, a genus of shrubby herbs
from Africa named after Dutch explorers
Henrietta Tinne and her daughters.
GenuS NEOTINEA
denSe-flOwered
and Burnt OrchidS
29/1/09 12:35:01
DENSE-FLOWERED ORCHID
341
denSe-flOwered Orchid
Neotinea maculata
This small and rather inconspicuous orchid is confined to western Ireland, although for a
while it also occurred on the Isle of Man. It has a very peculiar world range, being essentially
a Mediterranean species adapted to the mild wet winters and hot dry summers of that
region and seemingly ill-fitted to the damp, windy, oceanic climate of western Ireland. It
is not unique, however, as several other Mediterranean species, such as Strawberry-tree,
occur in Ireland. Various theories have been advanced to explain this distribution, including
land-bridges between Ireland and the Continent or glacial refuges off western Ireland, and
the subject has generated a good deal of controversy. On balance, colonisation of Ireland via
wind-blown seed would seem the most likely explanation.
Identification
The tiny off-white flowers are distinctive.
Similar species
Superficially similar to Small White Orchid,
but the lip is a very different shape.
Habitat
Typically found on short turf on pastures,
road verges, limestone pavements and around
loughs and turloughs. It has occasionally been
recorded on dunes or from ash and hazel
woods in the hills. Although mostly confined
to calcareous soils on limestone it has been
found growing on gravels and also on light,
peaty soils overlying more acidic rocks. Occurs
up to 300m above sea level but most are found
below 100m.
Flowering period
Late April to early June, usually peaking in midMay. The flowers go over very quickly.
Range
Confined to Ireland, where most populations
are in The Burren region of Co. Clare and
adjacent Co. Galway, including the Arran
Islands. Otherwise there are a few very scattered
sites in eastern Co. Cork, Co. Limerick and
O 29 May, East Sussex. Burnt Orchids.
P 21 May, Co. Clare. Var. alba is commonest in Ireland
and has unspotted leaves and unmarked creamy flowers.
29/1/09 12:35:03
342
GENUS NEOTINEA
How to find it
1987-99
1970-86
pre 1970
Small and inconspicuous, this is a hard orchid

to spot, especially from walking height. It is
usually found in small scattered colonies with
1-20 flowering plants. Locally frequent in The
Burren, a mid-May excursion to this beautiful
area is well worthwhile.
DESCRIPTION (var. alba)
Co. Donegal and in Northern Ireland in

Co. Fermanagh. Extinct in the Isle of Man.
World Range: The Mediterranean basin
from Portugal and Spain east to Cyprus,
Turkey, Syria and Israel; ranges north to
southwest and southern France, the former
Yugoslavia, Greece, Romania and the Ukraine,
and south to Morocco, Algeria and Tunisia
in North Africa. Also found on most of the
Mediterranean islands, the Canary Islands and
Madeira.
Height: 4-40cm but mostly 6-15cm and very

rarely over 30cm.
Stem: Green, with one or two brown
membranous sheaths at the extreme base.
Leaves: There are three to six rather dark green
leaves, of which two or three (sometimes four)
are broadly oblong-elliptical in shape and held
rather spreading at the base of the stem; the
remainder are narrower and more pointed and
loosely sheathe the stem with the uppermost
even smaller and more bract-like. The leaves
appear in October and are wintergreen.
Spike: Cylindrical and dense, composed of 1520 tiny flowers (occasionally as many as 35), all
facing more-or-less in the same direction.
Bract: Green, becoming whitish towards the
tip. The bracts are lanceolate with a pointed
tip and sometimes a tooth at the side and are a
little shorter than the ovary, which they clasp.
M 21 May, Co. Clare.With Birds-foot Trefoil in the rocky landscape of The Burren.
29/1/09 12:35:04
DENSE-FLOWERED ORCHID
343
M 21 May, Co. Clare. The tiny flowers have a distinctly

three-lobed lip and vaguely resemble human figures.
Ovary: Pale green, fat, cigar-shaped and

twisted, with three diffuse ribs. The ovary is
rather larger than the flower and stands upright.
It narrows and bends at the tip, however, to
hold the flower facing outwards.
Flower: Small and whitish, resembling a tiny
man with a rather oversized helmet. The sepals
are white, washed greenish and with green
veins. They are oval-lanceolate to lanceolate,
pointed at the tip and fused at the base to
form a tight elongated hood. The petals are
greenish-white with green veins, very narrow,
strap-shaped and pointed; they are enclosed
within the hood. The lip is small in relation to
the hood and is held at an angle forwards and
downwards. It is three-lobed with the central
lobe strap-shaped and notched or shallowly
forked at the tip (sometimes with a small tooth
in the notch). The side-lobes are positioned
near the base of the lip and are rather shorter
and narrower. The spur is very short and
conical. The flowers are reported to smell faintly
of vanilla.
Subspecies
None.

Var. alba has unspotted leaves and whitish or
creamy flowers (rarely more greenish). This is
the common variety in Ireland.
Var. maculata has spotted leaves with the
M 21 May, Co. Clare. Like most Mediterranean orchids,

the leaves appear in October and are wintergreen, dying
off after flowering.
29/1/09 12:35:07
344
GENUS NEOTINEA
produced in good quantities (c. 1350 seeds per

capsule).

Hybrids
None known.

The specific name maculata means spotted or
blotched.
HISTORY AND
CONSERVATION
M 24 May, Co. Clare. The typical variety, which is scarce

in Ireland, has spotted leaves, purple-veined flowers and
often spotted ovaries, too.
reddish or purple spots arranged in parallel

longitudinal lines. The lip is pale pink or
marked with a longitudinal pink stripe and,
the sepals and petals have pink or brown
veins. Occasionally the stem is spotted and
sometimes also the bracts, ovaries and even
sepals and petals. This is the typical variety (in
that it takes its name from the species) but it is
relatively scarce in Ireland.
Var. luteola has primrose-yellow flowers. It is
very rare and perhaps extinct in Ireland.
BIOLOGY
The spur contains traces of nectar, suggesting
that the flowers are attractive to small insects.
Although cross-pollination is possible, however,
most or all plants are self-pollinated, sometimes
even before the buds have opened. Whether
self-pollinated or cross-pollinated, seed is
First found in Ireland in May 1864 by Miss

Frances More at Castle Taylor in Co. Galway.
It was discovered on the Isle of Man in 1966
in an area of dunes on the north coast at The
Ayres, Ballaghennie. This colony persisted
until 1986 but Dense-flowered Orchid is now
extinct there.
In Ireland the species is scarce and local
and has been lost from 46% of the historical
range. Many of the losses are comparatively
recent, although some of the decline has
been offset by the discovery of new sites.
It is extinct in Co. Offaly, Co. Roscommon
and perhaps Co. Mayo (where noted from
Lough Mask by Summerhayes, 1968). The
improvement of pastures and overgrazing are
likely causes for the losses. With the current
rapid pace of development in Eire there is
cause for concern about the future of such a
localised orchid.
Past and present occurrence of Dense-flowered Orchid
Atlas).
Britain
Ireland

1500-1999
24
current range
13 (1.3%*)
% lost, 1500-1969
0%
21%
% lost, 1970-1986
100%
25%
% lost, total
100%
46%
29/1/09 12:35:09
BURNT ORCHID
Burnt Orchid
345
Endangered, BAP
Neotinea ustulata
Formerly: Orchis ustulata; Other names: Burnt-tip Orchid
This delicate species is one of our most attractive and delightful orchids but, sadly, is also
probably the orchid that has suffered the greatest decline in the last 50 years or so. It is now
confined to around 75 sites in England and just one in Wales, mostly on well-managed
chalk grassland reserves in southern England but with a few colonies still surviving on the
limestone of northern England. The unopened buds are deep purple, giving the top of the
spike a scorched or burnt appearance, hence the name. Burnt Orchid is the county flower
of Wiltshire.
Identification
Burnt Orchid is one of the group of orchids
in which the flower resembles a tiny human
figure, with the lip divided into arms and
short, stumpy legs. It is easy to identify, with
a distinctive combination of small size, dark
reddish-purple buds and a white lip marked
with fine reddish-purple spots. The contrast
between the dark buds at the top of the spike
and the white lips of the lower flowers is very
striking. There are two forms of Burnt Orchid,
early and late flowering, but they do not differ
significantly in appearance.
Similar species
Lady Orchid resembles Burnt Orchid in general
flower structure and colour but is very much
larger, and its lip differs in the details of shape
and coloration.
Habitat
Burnt Orchid is found on ancient short
grassland on chalk and limestone soils, often on
south- or west-facing slopes, although the lateflowering populations are not so fussy about
the aspect. In southern England the species
favours the narrow terracettes that follow the
contours of the slope on the chalk downs and
also Bronze and Iron Age earthworks where
the ground has been undisturbed for centuries.
Only on rare occasions will it colonise new sites.
For example, at Martin Down in Hampshire
P 13 July, Hampshire. In the late flowering var. aestivalis
the hood does not fade after the flower has opened.
29/1/09 12:35:11
29/1/09 12:35:14
BURNT ORCHID
it appeared in the 1980s on grassland that

had been under the plough until 1957. Burnt
Orchid is also sometimes found in alluvial
water meadows where the silt is derived from
chalk, e.g. in Lincolnshire and formerly at sites
in Hampshire and Oxfordshire. This habitat
was always uncommon in southern England but
in the past Burnt Orchid was found relatively
frequently in riverside pastures in the north. It
is predominantly a lowland species but has been
recorded up to 275m above sea level.
347
1987-99
1970-86
pre 1970
Flowering period
There are two varieties that differ in their
flowering period. The early-flowering form
appears from the second or third week of May
to mid-June in southern England and may, on
average, be just a few days later in the north; it
is usually at its best in the last ten days of May.
The late flowering form flowers from the end of
June through July to early August.
Range
Once found throughout the chalk and
limestone areas of England, the species has
undergone a major decline and is now extinct
in many areas. In southern England, Burnt
Orchid can still be found in Dorset, the Isle of
Wight and West Sussex but the strongholds are
the downs of Wiltshire, Hampshire and East
Sussex. Outlying relict populations cling on
in Gloucestershire, Berkshire, Hertfordshire,
Bedfordshire and Kent. The decline has
been even more marked in the Midlands and
northern England, and the species is now only
found in Derbyshire, Lincolnshire, northern
Yorkshire and at a single site on the coast of
Co. Durham (the most northerly in Britain). In
1993, bucking the trend of range contraction
and decline, Burnt Orchid was recorded for the
first time in Wales, on limestone grassland at
Llanmadog, Glamorganshire.
The late-flowering variety aestivalis (see
Variation and varieties) is only found in
southern England. It has been recorded from
O 29 May, East Sussex. In the early flowering form the
hood fades once the flower has opened.
Wiltshire (four sites), Hampshire (five sites,

plus one now destroyed) and East Sussex (14
sites, mostly between Lewes and Eastbourne,
one holding nearly 1,000 spikes in 2002).
World range: Confined to Europe, although it
extends just slightly into western Siberia (to c.
70E). It occurs north to Denmark, southern
Sweden, Estonia and the St Petersburg region
of Russia and south to northern Spain, Italy,
northern Greece, Bulgaria and the Ukraine. It
is also found in the Caucasus. In the south of
the range it is mostly found in the mountains
and is absent from the Mediterranean lowlands.
Extinct in Holland.
How to find it
Sadly, the chances of coming across this lovely
little orchid unexpectedly have become slim but
it is nevertheless still worthwhile looking for in
suitable undisturbed short swards. Otherwise,
a visit to a good chalk grassland reserve is
necessary to see it but even then it can be hard
to find. The colonies can be very localised, with
hundreds of plants in one area and few or none
elsewhere, even on seemingly suitable ground.
Being so small, it can be surprisingly hard to
spot among the various short downland herbs
such as thyme and milkwort. As with many
orchids, the number of flowering spikes varies
greatly from year to year and in extremely dry
seasons there may be none at all. One of the
29/1/09 12:35:16
348
GENUS NEOTINEA
M 13 July, Hampshire. The flowers look like tiny human

figures; this is the late flowering var. aestivalis.
best sites is Mount Caburn (East Sussex).

Others include Seven Sisters (also East Sussex),
Martin Down (Hampshire) and Pewsey
Downs, Parsonage Downs and Clearbury
Rings (Wiltshire).
DESCRIPTION
Height: 2.5-15cm, very exceptionally to 30cm.
In southern England it is usually around 4-10cm,
with the very shortest plants found at heavily
grazed sites. In northern England it is a little
taller, usually around 9-12cm, even to 15-25cm
when growing among taller vegetation.
Stem: Yellowish-green, slender and ridged
towards the tip, with two or three membranous
white sheaths at the very base.
Leaves: Green with a faint blue tone, fading to
pale green by flowering time. There is a rosette
of two to five elliptical-oblong keeled leaves
at the base of the stem and one or two bractlike leaves higher up. The leaves appear in the
autumn, are wintergreen and start to wither
as the plant comes into flower (by which time
non-flowering plants have already vanished).
Spike: The 15-50 flowers are densely packed
to form a spike that is initially conical but

lengthens and becomes cylindrical when all the
flowers have opened.
Bract: Reddish-purple, lanceolate and rather
short, at around two-thirds the length of the
ovary.
Ovary: Green, twisted and obscurely six-ribbed.
Flower: The sepals are oval, the lateral sepals
asymmetrical with their outer surfaces dark
reddish-purple (thus the unopened buds are
very dark) and their inner surfaces greenish.
The petals are paler and more strap-shaped.
The sepals and petals form a compact hood that
embraces the column; this is reddish-purple
when the flower is fresh although it quickly
fades (and see var. aestivalis below). The lip
is white with a few scattered reddish-purple
spots (formed by minute papillae) and is deeply
lobed to form two broad chubby arms and two
short stumpy legs, sometimes with a tiny tooth
between them. There is a groove in the centre
of the lip towards the base that leads into the
short, conical, down-curved spur. The column
is short and whitish and the pollinia are yellow.
The flowers of the early-flowering variety have a
strong, sweet, honey-like scent said to be similar
to Heliotrope, and var. aestivalis has a weak,
citron-like scent that is vaguely unpleasant.
Subspecies
None.

Var. ustulata and var. aestivalis Very unusually
there are two varieties of the Burnt Orchid
which differ in their flowering time. The earlyflowering var. ustulata is much the commoner
and blooms in late May and early June. The
late-flowering var. aestivalis is only found in
southern England and flowers around July. The
two varieties are rarely if ever found together,
but at a site in East Sussex a colony of 200 lateflowering plants merges with a much smaller
group of early-flowering plants on a northfacing chalk slope.
The two varieties differ slightly in
appearance. In the late-flowering aestivalis the
reddish-purple of the hood is on average darker
29/1/09 12:35:17
BURNT ORCHID
349
and the colour does not fade once the flower

has opened. There is also a distinct rose-purple
wash to the edges of the lip and this wash
may spread across the lip. The early-flowering
variety never shows the persistent dark hood or
the coloured fringe to the lip. Other differences
are rather subtler. On average, var. aestivalis is
taller, although this may simply be a product
of the plants having to compete with the taller
vegetation found later in the season. It has a
more open spike of slightly smaller flowers and
the lip is also shorter, with a slightly narrower
waist and shorter legs which have a deeper
cleft between them. Also, the spots on the lip
are larger.
Similar late-flowering populations in
Europe have been described as a distinct
subspecies, N. u. aestivalis, and it has even
been suggested that they should be treated as
a different species. Genetic analysis shows that
there is far too small a difference between the
early- and late-flowering varieties to justify this.
Var. albiflora has unmarked white flowers. It is
very rare but has been recorded in Derbyshire,
Kent, Berkshire, Hampshire and Wiltshire.
BIOLOGY
Poorly known. Early-flowering plants are
pollinated by flies that feed on nectar and
sugary plant juices. Presumably the flies are
initially attracted by the combination of
colour and scent, but this species does not
produce nectar to reward them and they must
be satisfied in some other way. Once on the
flower, the flies work from the uppermost,
unopened buds downwards and insert their
proboscis in a head-down position. They
taste with their feet beforehand and there
may be some sugary secretion from the flower
to guide them into the correct position; the
groove at the base of the lip may in addition
act as a leading line. The mouth of the spur is
narrow and rather like a keyhole; this shape
and the design of the column may be related
to the unconventional upside-down position
of the pollinating fly. Butterflies also visit the
M 13 July, Hampshire. The late flowering var. aestivalis

differs slightly in appearance from typical plants as well
as in its flowering period, but genetic research shows no
appreciable difference between the two.
flowers but have not been recorded carrying

pollinia. With its different scent and later
flowering, var. aestivalis may attract a different
suite of species and, in Europe, beetles have
been recorded as pollinators. Whatever its
mechanism, pollination is not very efficient in
England and seed-set is relatively poor, with
around 20% of flowers producing ripe capsules.
Nevertheless, most new plants are recruited to
the population from seed.
Vegetative reproduction takes place but is
thought to be relatively unimportant. Groups of
four to six spikes are not uncommon, however,
and we have seen a compact cluster of 12 (see
29/1/09 12:35:18
350
GENUS NEOTINEA
photo); such groups may well be the product of

vegetative reproduction.

The degree of fungal infection is unclear,
with both high and low rates being reported.
Dormancy is common, however, and Burnt
Orchids are able to spend up to three years
(sometimes even four) underground without
producing aerial parts. During this period the
fungal partner must play a major role (although
dormant plants have a significantly reduced
chance of surviving). Most plants flower for one
to four seasons in succession (rarely for up to
seven successive years) and then either die or
retreat into a period of dormancy underground
or as non-flowering rosettes.
Seed germinates to produce a protocorm
which grows to a length of 20-30mm before the
first root is produced (the longest protocorm
known among orchids with a similar pattern
of development). Both in cultivation and in the

wild Burnt Orchids can flower within about
three years of germination, but older estimates
of up to 16 years are still quoted. This figure
is based on the pioneering work of Fuchs and
Ziegenspeck in the 1920s. They estimated the
length of time the protocorm spent developing
underground by examining it for constrictions,
each of which was thought to represent a
years growth, like annual rings on a tree. This
technique is now known to be unreliable.
Hybrids
None are known.

The specific name ustulata means scorchedlooking. Until recently Burnt Orchid was placed
in the genus Orchis, but genetic studies have
resulted in its transfer to the genus Neotinea.
There it joins the superficially rather different
Dense-flowered Orchid.
M 13 July, Hampshire.This group of 12 plants (var. aestivalis). is probably the product of vegetative reproduction.
29/1/09 12:35:20
BURNT ORCHID
351
HISTORY AND
CONSERVATION
The first British record was in 1634 in Thomas
Johnsons Mercurius Botanicus (Botanical
Mercury): ...in montosis pratis (mountain
pastures). The first specific locality was given in
1650 in William Hows Phytologia Britannica
natales exhibens Indigenarum Stirpium sponte
emergentium (A British botany presenting the
origins of native wild plants): On Scosby-lease,
Mr Stonehouse. Therefore the first localised
record was from near Doncaster in northern
England rather than the chalk downs of the
south where it was presumably much commoner.
Formerly locally frequent over much of
England, the Burnt Orchid has suffered one
of the most catastrophic declines of our wild
orchids. It is now greatly reduced, both in terms
of numbers and distribution, and is classified
as Endangered. Out of a total historical range
covering 265 10km squares, the New Atlas
records a post-1987 presence in just 55,
representing a 79% decline. It is now Nationally
Scarce.
Past and present occurrence of Burnt Orchid in Britain

1500-1999
current range
Britain
Ireland
265
55 (1.9%*)
% lost, 1500-1969
72%
% lost, 1970-1986
7%
% lost, total
79%
It is possible that subtle changes in climate

in the 19th and 20th centuries may have
caused some decline, but most losses can be
directly attributed to agricultural changes. The
species vanished from some counties at the
edge of the range long ago (e.g. last recorded
in Norfolk at the end of the 18th century),
mainly due to the ploughing of grasslands
following the Enclosures. The decline continued
M 29 May, East Sussex. The reddish-purple buds and

hood give the flowers a scorched or burnt appearance.
through the 19th and 20th centuries due to

the continued loss of downland and pastures
(perhaps especially during World War Two),
while in the latter half of the 20th century
the improvement of grasslands with artificial
fertilisers and pesticides continued to take a toll.
The remaining grassland sites are vulnerable
to scrub invasion due to a lack of grazing,
especially in southern England, where first
rabbits were decimated by myxomatosis and
latterly livestock has become less economically
viable. Paradoxically, overgrazing is also a
problem, especially in northern England, where
losses due to building and other development
have also occurred. Sites were even wilfully
destroyed, as at Cottondale in southeast
Yorkshire, ploughed out in 1965 to counter the
attention shown it by botanists.
There were around 350 historical localities
in southern England but only around 75
colonies survived into the 1990s. Most sites
29/1/09 12:35:21
352
GENUS NEOTINEA
have fewer than 50 flowering plants and,

indeed, at some just one or two spikes make
sporadic appearances; fewer than ten regularly
hold more than 200 plants. The strongholds
are now in Wiltshire, Hampshire and East
Sussex, and notable concentrations include
Parsonage Down in Wiltshire where there
may be 30,000 flower spikes spread over 95ha
in one continuous mega-colony, probably
the largest surviving population in northwest
Europe. In other counties where Burnt Orchid
still survives, many or most of the sites have
been lost. Thus in Dorset only two out of 12
known sites remain and the equivalent figures
for the Isle of Wight are one of 12, for West
Sussex one of six, for Kent seven of 18, for
Berkshire one of 17, for Hertfordshire one of
11, for Bedfordshire one of ten and for east
Gloucestershire one of 21.
Of over 120 sites recorded from northern
England around 30 survive. There are a few
in Derbyshire and northern Yorkshire; it
reappeared in mid-west Yorkshire in 1988 but
the single plant was promptly dug up. In the
north and west of Lincolnshire there are three
small colonies and just one remains in Co.
Durham. Only two of the sites in the north, in
Derbyshire and northwest Yorkshire, hold more
than 200 spikes, and the total population in
northern England is probably fewer than 1,000
flowering plants.
Fortunately many sites are now protected
as SSSIs, nature reserves or are on Ministry
of Defence land and managed for the benefit
of their chalk flora and fauna. As the chalk
landscape was created and maintained by
hundreds or thousands of years of grazing, it is
important to continue this practice. For Burnt
Orchid the ideal regime is light spring grazing
until late April with the return of sheep from
late July until late September (when the new
leaves start to appear); cattle are unsuitable as
they can trample and destroy the plants and
also damage the turf.Distribution
ExtINCt
the Burnt Orchid is extinct in the following vicecounties (the number of recorded sites, followed
by the date of the last record, is given in parentheses). Source: Foley (1987, 1990, 1992).
South Devon (four sites, last seen in 1932)

North Somerset (seven, 1961)
West Kent (three, late 19th century)
Surrey (twelve, 1966)
North Essex (two, 19th century)
Middlesex (one, pre 1737)
Oxfordshire (two, 1982)
Buckinghamshire (five, 1961)
East Suffolk (but this 19th century site might
have been in Norfolk)
West Suffolk (three, 1939)
West Norfolk (one, 18th century)
Cambridgeshire (twelve, 1955)
Huntingdonshire (one, 1890)
Northamptonshire (four, 1956)
West Gloucestershire (eight sites)
Herefordshire (two, 19th century)
Worcestershire (five, early 20th century)
Staffordshire (two, early 20th century)
Shropshire (six, 19th century)
Leicestershire (two doubtful old records)
Rutland (three, around 1915)
Nottinghamshire (three sites)
West Lancashire (two, 1940s)
Southeast Yorkshire (eleven, 1973)
Southwest Yorkshire (four, prior to 1888)
South Northumberland (one, prior to 1868)
Westmorland (seven, 1970s)
Cumberland (ten, early 1970s)
29/1/09 12:35:22
GENUS HIMANTOGLOSSUM
LIZARD ORCHID
353
Distribution
There are five closely related
species in Europe, North Africa
and in the Middle East from
Turkey and Lebanon to Iran.
Perhaps surprisingly, their closest
relatives include the bee and
spider orchids Ophrys, tongue
orchids Serapias and pyramidal
orchids Anacamptis.
Name
The generic name
Himantoglossum derives from
the Greek himas leather strap or
thong and glossa tongue, thus
strap-tongued, a reference to the
long, narrow lip.
12/2/09 17:06:31
354
lizard Orchid
Near Threatened
Himantoglossum hircinum
The bizarre flowers of this robust orchid bear a fanciful resemblance to a lizard and smell
strongly of billy goat. It has always been rare in Britain but its range and numbers have ebbed
and flowed, probably due to subtle changes in climate; it once occurred north to Yorkshire
but retreated southwards and is now confined to a few sites in southern England, although it
is currently on the increase again and could turn up almost anywhere in the south and east.
Identification
Very distinctive. The tall spikes of large greyishgreen flowers have an untidy, straggly, ragged
appearance and on close inspection their
structure is unique. The long central lobe of
the lip resembles a lizards tail, and the shorter
side-lobes form the back legs. The hood of the
flower is said to recall the head and body of
the lizard, but it seems much more reasonable
to say that the fore-quarters of the lizard have
been swallowed and vanished into the throat of
the flower.
Similar species
None.
Habitat
M 30 June, East Sussex. By flowering time the lower

leaves, which appear in the autumn, have withered.
O 25 June, Cambridgeshire. Smelling of billy goat and
with such extravagant flowers, this is a truly bizarre orchid.
P 27 June, Kent.
The Lizard Orchid usually grows in open sunny

situations among fairly rank grass, although
sometimes it also occurs on shorter and more
closely-cropped swards. It can appear among
scrub or on the edge of woodland but would
eventually be shaded out if the canopy developed
too much. It is always found on well-drained
soils, usually on chalk but sometimes also on
limestone and occasionally on boulder clays,
sands or gravels; it has even been recorded
growing through broken tarmac. Whatever the
soil, it will probably be calcium-rich, although
some recent records have been on neutral
soils. Suitable sites include road verges, railway
embankments, ancient earthworks, field margins,
old chalk pits and dunes, and it even grows on
lawns on one favoured housing estate. At least six
of the 19 recent populations are on golf courses,
and it has been suggested that golfers may have
carried the seeds from one course to the next on
29/1/09 12:35:27
LIZARD ORCHID
355
29/1/09 12:35:32
356
their shoes or equipment. Found from sea level

to about 200m in Gloucestershire and on the
North and South Downs.
1987-99
1970-86
pre 1970
Flowering period
Early June to late July, sometimes from late May.
Traditionally peaking in the first week of July it
has, like many other orchids, tended to flower
earlier in the last few years, in mid-June.
Range
The distribution has ebbed and flowed, but
all the records have been southeast of a line
from the Severn to the Humber apart from
one old site further north in Yorkshire and a
doubtful record from Lancashire. In recent
years there has been a very large established
colony at Sandwich Bay in Kent and a smaller
permanent colony at Newmarket in Suffolk, as
well as small colonies in East Sussex, Dorset
and Suffolk. Other recent records, either of
sporadic appearances or tiny populations,
come from north Somerset, north Hampshire,
West Sussex, Surrey, Oxfordshire and west
Gloucestershire. It has also been recorded from
Jersey and Guernsey in the Channel Islands.
World range: Southwest and southern Europe,
north to Holland, Germany, Austria and the
Czech Republic, south to northern Spain (and
very scattered in central and southern Spain),
Italy and Sicily and east to Greece, also North
Africa in Morocco, Algeria and Tunisia. (The
lizard orchids further to the east and in the
Balkans are often treated as separate species.)

The species is commonest in France, where it is
a roadside weed in the wine-growing regions.
How to find it
Despite their size, the greyish-green spikes can
be hard to see among long grass. Undoubtedly
the best site is at Sandwich Bay in Kent where
there have been up to 3,000 flower spikes in
recent years, with 5,000 in 2000 (when the total
population was estimated at 27,500 plants over
an area of 0.5 square km). Another well-known
colony is on the Devils Dyke that runs across
Newmarkets July racecourse, with around 200250 flowering plants.
M 27 June, Kent. Lizard Orchid is an adaptable species and a prestigious colonist on a lawn.
29/1/09 12:35:34
LIZARD ORCHID
357
M 28 June, Cambridgeshire. As wih many orchids, the function of the elaborate lip is obscure.
DESCRIPTION
Height: 25-90cm but usually below 60cm and
commonly around 30cm.
Stem: Pale green with faint purple blotches,
ridged towards the tip and with two or three
scale leaves at the extreme base.
Leaves: Numerous, greyish-green and
sometimes blotched with purple. There are
four to ten large, oval-oblong, keeled basal
leaves forming an untidy rosette and three to
five smaller, narrower and more pointed leaves
which loosely clasp the stem up to the flower
spike. The leaves are wintergreen, and the basal
leaves have yellowed and often withered by
flowering time, especially in dry years.
Spike: Tall, occupying almost 50% of the stem,
cylindrical and rather dense, with 15-80 flowers.
Bract: Pale green to off-white, variably but
often strongly washed reddish, narrow and
pointed, and up to twice the length of the ovary.
Ovary: Pale green, cylindrical, tapering both
towards the flower and at the base into a short
stalk, twisted and with three ridges.
Flower: Overall greyish-green. The sepals
are oval and are fused at the base to form

a loose hood. They are pale greyish-green,
variably flushed or rimmed purple with a paler
interior that has lines of reddish-purple spots
and dashes along the veins (which may show
through to the exterior). The petals are very
narrow, strap-shaped and spotted reddishpurple. The lip is divided into three lobes.
The central lobe is 2.5-6cm long, narrow,
parallel-sided and ribbon-like, with a deeply
notched tip. It is twisted through two or three
turns and is whitish at the base, becoming pale
lilac-brown for the outer 80%. The side-lobes
are very much shorter and a little narrower,
pointed and pale lilac-brown. The sides of the
lip at the base are boldly folded or corrugated,
and this corrugation continues on to the outer
edges of the side-lobes. The centre of the base
of the lip is trough-like with a dense fur of
tiny white papillae and with variable bright
purplish-red tufted spots and blotches. The
lip is coiled like a spring in bud and slowly
unrolls to project outwards and downwards at
about 45 (sometimes nearer the horizontal on
29/1/09 12:35:36
358
few-flowered spikes). The spur is short, curved

downwards and bluntly conical. The anther is
greenish-white, and the two greenish pollinia
are fixed to a single viscidium that is concealed
in the purple bursicle. The stigmatic zone is
purple. The strong scent recalls a billy goat and
is most pungent in the evening.
Subspecies
None.

None.
BIOLOGY
The flowers are visited by a variety of insects,
including hover flies, various bees, wasps, ants,
beetles, butterflies and moths. Of these, bees
are probably the main pollinators. There is no
nectar (although glucose has been detected
in the spur), and the flower seems to offer its
pollinator no reward; despite this the pungent
scent may serve to attract flies and nightflying moths. Self-pollination is physically
possible but is thought not to occur. The
species is self-compatible, however, and solitary
isolated plants do set seed; for this to happen
either self-pollination (autogamy) or crosspollination from other flowers on the same
spike (geitonogamy) must occur. Whatever
the mechanism, it does not seem to be very
efficient and only around 30% of flowers are
pollinated, although each capsule contains up
to 1,200 seeds, perhaps more. It is possible for
the species to reproduce vegetatively by forming
extra tubers, but this seems to be a rare event.

The Lizard Orchid spends the summer resting
period underground as a tuber. The aerial
shoot appears from late August onwards, with
the lower leaves unfolding and some short and
rather thick roots growing from the base of the
stem. These leaves will overwinter and by early
spring may start to blacken at the tips. By then a
new tuber has started to form from a bud at the
base of the stem and by May the roots and older
tuber, as well as the leaves, are often withering.

At flowering time, therefore, the plant has
two large, egg-shaped oval tubers (one old and
withering, one new). After flowering and setting
seed the plant will disappear, leaving only the
new underground tuber to survive the summer.
Individual plants can be long-lived, surviving
for up to at least 19 years after first emergence,
but they may not flower every year and can
remain dormant as underground tubers for
a year or appear merely as a rosette of leaves,
sometimes for many years. An individual plant
in Sussex flowered in 1984 and then not again
until 1995, and a colony in Suffolk of between
16 and 40 plants did not produce any flowers
over a ten-year period.
Seed germinates in the autumn up to
two years after the seeds have ripened and
dispersed. Although some of the seed may blow
away, it is common for mature flowering plants
to have large numbers of seedlings close by, and
many seeds stay in the pods until the dead spike
falls over. With the aid of fungi, the seedling
initially develops into a protocorm the size of a
small pea. In the spring following germination,
the protocorm produces the first tuber and
then withers away, leaving just the tuber to
oversummer. In the autumn, a short rhizome
grows from this tuber and develops a root and
a second tuber which is fully developed by the
following spring. This establishes the annual
cycle of replacement tubers. When the seedling
is more than two years old the first leafy shoot
appears above ground. The seedling usually
remains in the one-leaf stage for two or more
years and then moves on to two leaves, again for
several years, and then three leaves. Although
in very favourable conditions a plant with two
or three leaves can flower, it is only when it
has grown four leaves that it can be considered
mature. Even then it may not flower every
year. The period between germination and first
flowering may therefore be a minimum of six
years.
Hybrids
None.
29/1/09 12:35:36
LIZARD ORCHID
359
P 28 June, Cambridgeshire.The long central lobe

of the lip uncoils like a
spring.

The specific name hircinum means of goats, a
reference to the scent.
HISTORY AND
CONSERVATION
Thomas Johnson published the second part of
his Mercurius Botanicus (Botanical Mercury)
and recorded it Nigh the highway betweene
Crayford and Dartford in Kent.
The Lizard Orchid is a Red Data Book
species that is classed as Near Threatened and

fully protected under Schedule 8 of the Wildlife
and Countryside Act 1981. Despite this it has
been targeted by collectors, and plants have
been dug up at Sandwich Bay in Kent (where
a 24-hour guard was maintained in the 1970s
and 1980s) and as recently as 2003 from a road
verge in Dorset.
The Lizard Orchid has an intriguing history
in England. From its discovery until about 1850
the species was restricted to the Dartford area
of Kent, although the original 1641 colony
29/1/09 12:35:38
360
Past and present occurrence of Lizard Orchid in Britain


1500-1999
current range
% lost, 1500-1969
% lost, 1970-1986
% lost, total
Britain
Ireland
115
20 (0.7%*)
76.5%
6%
82.5%
had been destroyed due to road-widening.

There was then a decline in that area and just
odd, isolated records in southeast England,
mostly in Kent; by 1900 just four sites were
known in England and it was thought that
the Lizard Orchid was almost extinct. Then,
during World War I and the 1920s there was
a marked expansion northwards with the
species turning up unpredictably, often just a
single plant but sometimes in larger numbers,
north to Yorkshire and west to Devon. A
remarkably large proportion of the new sites
were found by schoolgirls. This expansion
peaked around 1927 with 36 populations, but
the record is slightly clouded as Lizard Orchids
had been dug up and transplanted to gardens
(sometimes from as far away as France) and
at least two sites in this period are thought to
have originated with garden escapes. Adding a
further complication, plants were sometimes
transplanted by well-meaning naturalists, often
for safe-keeping, to new sites or even new
counties. Whatever their origin, in addition to
the counties detailed in Range there are older
records from north Devon, Wiltshire, Isle of
Wight, north Essex, Hertfordshire, Berkshire,
Buckinghamshire, Norfolk, Bedfordshire,
Worcestershire, Lincolnshire and northeast
Yorkshire.
In another change of fortunes, there was a
sudden drop in the number of colonies after
1934, although the geographical spread was
maintained. From the mid-1940s to 1993 the
total number of populations remained stable at
nine to 11, although some sites were lost and

others were gained. During this period all the
colonies were small and even the most thriving
lasted only about 20 years. A further increase
started around 1994 with plants appearing west
to Somerset and Gloucestershire. There were
16 sites in 1996 and 19 by 2000. As usual some
of the new sites are ephemeral with the orchid
flowering once and vanishing, e.g. by 2000 four
of the 1996 sites had disappeared but there
were seven new ones (thus a net increase of
three). With protection, some colonies, such as
the very large population at Sandwich Bay, have
expanded.
Lizard Orchid is on the edge of its range
in England, and it seems that subtle shifts in
climate have a big impact on its distribution.
The species is wintergreen, and the amount
of rainfall during the growing period of
September-April has a marked influence on
the flowering success, with a dry autumn and
winter limiting flowering. Furthermore, flower
buds are initialised and formed over two
seasons so the rainfall over a two-year period
is important in determining flowering success.
Conversely, severe drought may kill many plants
and wipe out small populations.
Lizard Orchid favours areas of long but
sparse grass that is cut once or twice a year.
None of the English sites are grazed but the
species is tolerant of grazing, at least by cattle,
although rabbits have caused severe damage at
times, especially during hard winters, as have
slugs. Some sites are managed to open up the
vegetation and prevent scrub invasion (with the
Newmarket site, for example, being burnt every
fifth year during the winter), but those on dune
grassland do not need much management.
29/1/09 12:35:39
Genus ANACAMPTIS
Loose-fLowered,
PyramidaL and
Green-winGed orchids
29/1/09 12:40:29
362
GENUS ANACAMPTIS
This genus formerly included just the Pyramidal Orchid. It is now a rather heterogeneous
combination of superficially different species, three of which occur in Britain.
Distribution
Pollination
The genus contains about 20 species, in

northern, central and southern Europe, North
Africa and southwest Asia, east to Afghanistan
and south to Yemen in the Arabian Peninsula.
The flowers do not produce nectar and may

rely on deceit to attract bees as pollinators.
Pyramidal Orchid, and perhaps also Greenwinged Orchid, do, however, produce a sugary
sap in the walls of the spur and this may
function as a reward for insect visitors.
Classification
From 1817, when it was described and named,
until 1997, this genus contained just one
species, Pyramidal Orchid. However, genetic
studies then led to the transfer into Anacamptis
of 19 species from the genus Orchis, including
Green-winged and Loose-flowered Orchids.
Surprisingly, the genus Anacamptis is closer to
the tongue orchids Serapias, bee orchids Ophrys
and lizard orchids Himantoglossum than to the
genus Orchis from which those additional 19
species came.
Floral structures
In Loose-flowered and Green-winged Orchids
the two pollinia narrow to a caudicle (stalk) and
each is attached to their own viscidium which,
in turn, is enclosed in a bursicle. The two-lobed
stigma lies on the roof of the mouth of the
spur. In Pyramidal Orchid the two pollinia are
attached to a single strap-shaped viscidium.
Growth pattern
The aerial stem grows from a pair of spherical
tubers, as in the genus Orchis (see p.200).
of the aerial stem and these will go on to form
separate plants when the connecting stem dies
off in the autumn.
Name
The generic name Anacamptis derives from the
Greek anakampto to bend back. It is a reference
to the structure of the flower of Pyramidal
Orchid but there is disagreement as to which
particular bit of the flower is being referred to.
It may be the long spur of the flower, the guideplates at the base of the lip or the position of
the pollinia.
O 10 May, Norfolk. Sadly, although relatively common

not so long ago, Green-winged Orchid has been banished
from around half its range.
29/1/09 12:40:29
LOOSE-FLOWERED ORCHID
363
Loose-fLowered orchid
Anacamptis laxiflora
Formerly: Orchis laxiflora; Other names: Jersey Orchid
This stately orchid provides a real splash of colour to wet meadows in the spring. In Britain
it is confined to the Channel Islands, but it is widespread in the Mediterranean region,
although everywhere it is threatened by drainage and agricultural improvement.
Identification
A typical orchid with a tall, loose spike of
purple flowers. The leaves are narrow and held
erect. The lip is purple with a long white central
zone that is unspotted or has some darker
purple markings. The central lobe is shorter
than the side-lobes and the sides of the lip are
very sharply folded downwards; the lateral
sepals are held vertically upwards. This gives the
lip a tall but flattened profile.
Similar species
Early Purple Orchid is superficially similar but
has broader leaves that are usually spotted and
held in a rosette flat on the ground. Its flower
spike is more compact, and the centre of the lip
is spotted. The lip itself is distinctly three-lobed
with the central lobe larger and longer than the
side-lobes.
Habitat
Damp meadows and marshy fields flushed with
base-rich water.
Flowering period
Early May to mid-June but mostly in late May.
Range
Confined to Guernsey and Jersey in the
Channel Islands. There are old records from
Co. Durham, including one dating from
1872 when it was found growing north of
Hartlepool. These plants may have originated
as casual imports with wool shoddy. World
range: Essentially Mediterranean, found from
Portugal east to Greece, Cyprus and Turkey,
and including most of the Mediterranean
islands. Occurs north to north-central France
P 27 May, Sussex (Wakehurst Place).
29/1/09 12:40:34
364
GENUS ANACAMPTIS
1987-99
1970-86
pre 1970
(Seine-Maritime and Marne), Switzerland and

the former Yugoslavia. It was discovered for the
first time in Holland relatively recently but is
extinct in Belgium.
How to find it
The tall purple flower spikes are conspicuous.
The best site on Jersey is Le Noir Pr and on
Guernsey the Orchid Fields at Les Vicheries.
broad white stripe down the centre of the lip.

The sepals are oval to oblong and blunt-tipped;
the lateral sepals are asymmetrical and held
erect above the rest of the flower (sometimes
touching), and the upper sepal is held over the
petals, which are slightly smaller and form a
loose hood over the column. The lip is broader
than long with two large, rounded side-lobes
and a rather shorter blunt central lobe (the
central lobe is sometimes absent altogether).
The sides of the lip are so sharply folded
downwards that the side-lobes may touch at
their tips. The spur is long and slender but
broadens towards the tip, which is usually
notched, and either straight or slightly upcurved. The centre of the lip may have small,
dark, vertical streaks within the white central
zone, extending into the purple borders.
Subspecies
None.

Var. rosea has pale rose-pink flowers but is rare.
Var. albiflora has white flowers but is rare.
DESCRIPTION
Height: 15-60cm.
Stem: Green, usually strongly washed reddishpurple towards the tip, with two to four loose,
brownish scales at the base.
Leaves: Three to eight, keeled, pointed and
strap-shaped, well-spaced along the stem and
held rather erect, with the upper leaves small
and bract-like. The leaves appear in autumn and
are wintergreen.
Spike: The six to 20 flowers are held in a very
loose spike.
Bract: Green, washed reddish-purple or
sometimes entirely reddish-purple. The bracts
are strap-shaped, narrow, pointed and roughly
the length of the ovary (which they clasp) or
longer.
Ovary: Green, washed reddish-purple or
sometimes entirely reddish-purple. The ovaries
are long, slender, cylindrical and twisted.
Flower: Dark purple to reddish-purple with a
M 27 May, Sussex (Wakehurst Place). The more or less

two-lobed, unspotted lip is sharply folded, and the spur is
often upturned.
29/1/09 12:40:37
LOOSE-FLOWERED ORCHID
365
P 27 May, Sussex (Wakehurst Place). The lateral

sepals are held upwards,
and the upper sepal and
petals form an untidy
hood.
BIOLOGY

No specific information, but in cultivation
the period between germination and the
production of the first green leaves is short.
Hybrids
A. x alata, the hybrid with Green-winged
Orchid, has been recorded occasionally on
Jersey and once on Guernsey.

The specific name laxiflora derives from the
Latin laxus wide or loose and flos flower and
means loose-flowered.
Until very recently this species was
considered to be a member of the genus Orchis,
and its overall similarity to some of the other
members of that genus, such as Early Purple
Orchid, certainly supported this. Recent genetic
studies have shown, however, that Looseflowered and Green-winged Orchids should join
the Pyramidal Orchid in the genus Anacamptis,
although this change is not universally accepted.
HISTORY AND
CONSERVATION
Loose-flowered Orchid was first found on
Guernsey around 1788 by Joshua Gosselin,
a local naturalist, and discovered on Jersey in
1837 by C.C. Babington.
There have been declines due to habitat loss,
but the species is still locally common or even
sometimes abundant on Jersey and especially
Guernsey. Most of the populations are in reserves,
although water abstractions for agriculture
and golf courses can affect even protected
areas. Loose-flowered Orchid is naturalised at
Wakehurst Place in Sussex (the Royal Botanic
Gardens country annexe). In 1987, 350 plants
grown from seed collected in Crete were planted
out and this population is still thriving.
29/1/09 12:40:38
366
GENUS ANACAMPTIS
PyramidaL orchid
Anacamptis pyramidalis
This widespread and very colourful orchid of open grassland is commonest on chalk and
limestone but could be found almost anywhere that has a hint of lime in the soil. When they
first appear, the flower spikes form a pyramid or cone, hence the name, but as more flowers
open the shape of the spike changes to more of a globe or cylinder. Pyramidal Orchid is the
county flower of the Isle of Wight.
Identification
The dense, conical spikes of unmarked
bright cerise pink flowers are distinctive.
On close examination the flowers have a
deeply three-lobed lip and a very long, thin,
down-curved spur.
Similar species
The fragrant orchids flower a little earlier in the
season (apart from Marsh Fragrant Orchid) and
have a looser, taller, thinner and more tapering
flower spike. Their flowers are slightly more
purplish-pink and their lip is not so deeply cut,
with the three lobes shorter, broader and more
uneven in size and shape. If there is any doubt,
Pyramidal Orchid can always be identified by the
two prominent raised ridges or guide-plates at
the base of the lip, unique to the species.
Habitat
Pyramidal Orchid grows on dry, well-drained
grassland on chalk, limestone or other calciumrich soils, such as boulder clay. It is found on
both close-cropped turf and in taller, ranker
swards. Suitable habitats also include the
grykes of limestone pavements, cliff tops,
among Marram and Lyme-grass on sand dunes
and, very locally, the coastal machair of the
Hebrides. The species is sometimes still found
in old meadows, although this habitat has
almost vanished in most areas. It also grows
among scrub; rarely in open woodland with a
broken canopy. Pyramidal Orchid takes readily
to man-made habitats, such as road verges and
roundabouts, churchyards, old quarries, disused
O 28 June, Cambridgeshire. An opportunist, it can do well
on the verges of main roads or even on roundabouts,
29/1/09 12:40:39
PYRAMIDAL ORCHID
railway lines and abandoned industrial sites,

and is one of the first orchids to move into such
newly available habitats. Recorded up to 350m
above sea level (Brough, Cumbria).
367
1987-99
1970-86
pre 1970
Flowering period
Early June to mid-August, very exceptionally
from May, but mostly mid-June and early to
mid-July.
Range
Mostly found south and east of a line from
south Devon to North Yorkshire and in
southern and especially central Ireland. The
species clearly enjoys a mild maritime climate
and ranges further north and west into coastal
districts of north Cornwall, south and north
Wales, northern Ireland and northeast England
(north to Lindisfarne). It is also found on
the Isle of Man, Isles of Scilly and Channel
Islands. In Scotland it is rare and confined
to a few sites in the Borders, Lothian, Fife,
Angus, Sutherland, Kintyre and Dumfries and
Galloway, as well as the Inner Hebrides on
Arran, Islay, Colonsay, Tiree, Gunna and Coll

and on the Outer Hebrides on South Uist,
Funday and Barra. World range: Essentially
Europe but extends into North Africa and
the Middle East. Occurs north to Denmark,
southern Sweden and the Baltic States, east
M 5 July, Northamptonshire.Troops of Pyramidal Orchids add a splash of colour to high summer.
29/1/09 12:40:41
368
GENUS ANACAMPTIS
O 22 June, Norfolk. The

spike is conical to pyramidshaped when the flowers
begin to open.
to Poland and the Ukraine and south to the

Mediterranean, Crimea and Caucasus. Also
found on the Mediterranean islands, in North
Africa in Morocco, Algeria and Tunisia, and in
the Middle East in Israel, Turkey, northern Iraq
and northwest Iran.
How to find it
Generally common in suitable habitats, indeed
often abundant.
DESCRIPTION
Stem: Green, slightly angled towards the tip
and often rather slender and flexuous (bendy);
the stems frequently have a distinct kink
somewhere along their length. There are two or
three brown sheaths at the extreme base.
Leaves: Green, tinged grey, keeled and strap-
shaped with a pointed tip. The three or four

sheathing leaves grade into five or six nonsheathing leaves that decrease in size up the
stem and become bract-like towards the spike.
The lower leaves may arch gracefully outwards.
The sheathing leaves emerge in the autumn and
are wintergreen, but they have often withered
by flowering time. The spike and non-sheathing
leaves do not appear until the spring.
Spike: Very dense, the 30-100 flowers are
packed so closely that they often conceal the
stem. When the first flowers break bud the
spike is distinctly conical but as more flowers
open it elongates to become domed or oval in
shape and sometimes eventually cylindrical.
Bract: Green, sometimes flushed purple,
narrow, a little longer than the ovary and
tapering to a fine point.
29/1/09 12:40:43
PYRAMIDAL ORCHID
Ovary: Cylindrical, twisted and green, often

washed reddish-purple.
Flower: Usually vivid pink, sometimes paler
pink or reddish-pink, and fading a little as the
flower ages. The sepals are oval-lanceolate with
a variably pointed tip and dished sides, and the
petals are a little shorter and blunter. The lateral
sepals are held horizontally or slightly drooping,
to about 15 below the horizontal, and the upper
sepal and petals form a hood over the column.
The lip is white towards the base, wedge-shaped
and held projecting forwards and downwards.
It is deeply divided into three virtually equal
strap-shaped lobes. The outer lobes broaden
slightly towards their blunt, shovel-like tips,
and the central lobe is more parallel-sided with
the extreme tip pinched in and upwards. On
either side of the base of the lip there are two
prominent narrow raised ridges or guide-plates
that are extensions of the column. The ridges,
although roughly parallel, converge towards the
mouth of the spur. The spur is slender, 12-14mm
long (often longer than the ovary) and downcurved. The column is white, variably washed
pink. The rostellum partially blocks the entrance
to the spur, and the two stigmas lie low down on
either side of the column and both pollinia are
joined to the same strap-shaped viscidium. The
flowers are scented, the perfume most obvious in
the evening and variously described as sweet or
as slightly unpleasant and foxy.
Subspecies
None.

Var. albiflora has pure white flowers but is rare
(very pale pink flowers are rather commoner).
Var. sanguinea has blood-red flowers. It is
scarce and known only from the Hebrides and
northwest Ireland.
Var. emarginata has an unlobed lip that
resembles a scallop shell. It is very rare.
Var. angustiloba has a very deeply lobed lip.
Var. fundayensis has a taller and more
cylindrical flower spike. It was described from
Funday in the Hebrides but is probably extinct,
if it ever existed at all.
369
M 22 June, Norfolk. On a close inspection, the raised

guide-plates at the base of the lip are characteristic.
BIOLOGY
Pyramidal Orchid is pollinated by moths, both
day- and night-flying, and by butterflies. Dayflying moths and butterflies are attracted by the
flowers vivid coloration, and the various burnet
moths may additionally be attracted by its
similarity to the bright red spots on their wings.
At night the flowers scent would assist its
detection. The pollination mechanism is highly
evolved. The walls of the spur contain a sugary
sap but only an insect with a suitably long
proboscis can reach this. The ridges or guide
plates on either side of the mouth of the spur
act as a guide correctly to position the insects
proboscis, not only to access the spur and the
sap within but also to trigger the mechanism.
The rostellum hangs over the entrance to the
spur and when it is touched by an insect the
protective, flap-like bursicle is pushed aside and
the strap-shaped viscidium, complete with the
two pollinia, sticks to the insects proboscis.
29/1/09 12:40:45
370
GENUS ANACAMPTIS
protocorm
bud
tuber
root
replacement
tuber
spring
summer
autumn
spring
YEAR 2
summer
autumn
YEAR 3
M Development from seed in Pyramidal Orchid (after Rasmussen 1995).
As soon as the viscidium is exposed to the air

it contracts like a watch-spring and coils itself
around the proboscis. This not only helps
to attach it more firmly but also moves the
pollinia apart so that they become separated
by 90. Then, a few seconds later in a second
contraction, the pollinia swing forward so as
to be in a perfect position to strike the sticky
surfaces of the two stigmas on the next flower
visited and deposit packets of pollen. The
mechanism is efficient, with 65-95% of flowers
setting seed.
Vegetative propagation is also possible via
the production of additional tubers, which
often develop at the end of short rhizomes.

The aerial stem grows from a pair of rounded
or roughly elongated tubers and there are a few
slender fleshy roots growing almost horizontally
near the surface of the soil; it is the roots that
are infected with fungi. The basal leaves appear
in autumn and die down in summer and after
flowering the orchid spends the late summer
and autumn resting as a tuber. The Pyramidal
Orchid is clearly adapted to a Mediterranean
climate of mild, wet winters and hot dry summers
and its overwintering leaves are probably
vulnerable to hard frosts. The generally southerly
distribution in Britain, with colonies further
north concentrated near the coast, emphasises its
preference for a mild winter climate.
It is not known when germination occurs

but initially a protocorm is developed. Early
researchers suggested that the seedling
remained at this stage for three years but it may
well be just a few months before the first tuber
develops. Following this the protocorm dies off
in the late summer to leave just the small tuber
in a resting state. In the autumn a bud on the
tuber produces a short rhizome and from this
one or two roots develop and sometimes also
the first leafy shoot. The new rhizome and roots
have to be reinfected with fungi from the soil as
the tuber itself did not carry any infection. This
plant, including the leafy shoot, overwinters,
and in the following spring a new tuber starts
to develop from a bud on the rhizome and the
old one starts to shrivel away. Eventually the
shoot and rhizome also wither, and the plant
will again spend the late summer resting as a
tuber. In this way the annual cycle of growth is
established. Again, it has been estimated that
several years elapse between the first appearance
above ground and the first flowers, but this is
probably far too long and the period between
germination and flowering could be as little as
three years.
x Gymnanacamptis anacamptis, the hybrid with
fragrant orchid (presumably Common Fragrant
Orchid), has been recorded very rarely from
Hampshire, Gloucestershire and Co. Durham.
29/1/09 12:40:45
PYRAMIDAL ORCHID
371

The specific name pyramidalis means simply
pyramidal or conical. Until recently the
Pyramidal Orchid was the only member of the
genus Anacamptis, but it has now been joined
by Green-winged and Loose-flowered Orchids.
HISTORY AND
CONSERVATION
John Ray recorded it in his Catalogus Plantarum
circa Cantabrigiam nascentium (A catalogue of
plants found around Cambridge): In many
places, as in a chalkie close at Hinton near
where they burn lime.
Past and present occurrence of Pyramidal Orchid in Britain and Ireland (based on presence or absence in 10km
Britain
Ireland
849
350
681 (24%*)
241 (24%*)
% lost, 1500-1969
13.5%
25%
% lost, 1970-1986
6.5%
6%
% lost, total
20%
31%

1500-1999
current range
The boundaries of the range are stable. There

have been local declines, and Pyramidal Orchid
has been lost from 20% of the historical range
in Britain and 31% in Ireland. Some of the
declines are due to the loss of old meadows and
pastures and although it is tolerant of grazing,
overgrazing may also be a threat, perhaps
especially at the few sites on the machair of
the Hebrides and western Scotland. On the
other hand, there have been some marked
increases in recent years, mostly in recently
created man-made habitats such as road verges,
and the species is a pioneer colonist of such
places. Artificial habitats have their drawbacks,
however, as brownfield sites are developed and
old pits and quarries used for landfill.
P 21 June, Suffolk. Once all the flowers have opened the
spike is cylindrical, sometimes markedly so.
29/1/09 12:40:47
372
GENUS ANACAMPTIS
Green-winGed orchid
Near Threatened
Anacamptis morio
Formerly: Orchis morio; Other names: Green-veined Orchid
This beautiful orchid once graced pastures and other areas of undisturbed grassland
throughout much of England, Wales and Ireland, and the sight of a field splashed with
the purples and yellows of thousands of Green-winged Orchids and Cowslips was
commonplace. Tragically, agricultural changes have led to a dramatic decline in most areas,
and the species has become rather local, although it can still be found in very large numbers
at some favoured sites. It now largely depends on the sympathetic management of the few
remaining old pastures as well those churchyards blessed by its presence. Green-winged
Orchid is the county flower of Ayrshire.
Identification
A dainty and usually petite orchid with
unspotted leaves. It has a few relatively large
and very attractive flowers that are extremely
variable in colour, from deep violet-purple to
rose-pink or whitish, although most are a shade
of purple. Whatever the colour, the flowers are
distinctive. The sepals and petals form a hood
marked with the green or bronze veins that give
the species its name. The sides of the lip turn
downwards to form a broad and often colourful
skirt, and the long straight spur projects
conspicuously back from the flower.
Similar species
Early Purple Orchid is superficially similar but
usually has spotted leaves and also tends to have
a longer flower spike and more flowers that are
a paler purple. On close examination only the
upper sepal and petals form the hood because
the two lateral sepals are held upright as wings.
Early Purple Orchid also lacks green or purple
veins on the sepals and finally, although both
species are found in grassland, only Early
Purple Orchid is found in woodland.
Habitat
M 20 May, Norfolk.The flowers are usually a deep violetpurple, a colour that is extremely hard to reproduce in
print.
Green-winged Orchid favours unimproved

grassland on neutral or calcareous soils where
grazing, mowing or other factors keep the
grass relatively short and the sward open. It is
very strongly associated with old species-rich
grassland and is slow to colonise new sites.
29/1/09 12:40:48
GREEN-WINGED ORCHID
The optimum habitats are damp pastures on

clay soils, but it is also found on dry chalk or
limestone grassland and sometimes on sands
and gravels, including pockets of neutral
grassland on otherwise acidic heathland. The
species can tolerate both mowing for hay and
grazing. Because unimproved damp meadows
and pastures are now rare Green-winged
Orchid is almost as likely to be found on
more marginal sites such as stabilised dunes,
lime-rich eskers, old railway cuttings and
banks, village greens (especially those which
are commons), churchyards and golf courses.
Occasionally it is found on lawns or on old
industrial sites, for example, alkaline waste
produced in the manufacture of washing soda.
It is intolerant of shade, and although it can be
found among scrub it is seldom if ever found in
woodland. Recorded up to 305m above sea level
(Co. Roscommon).
Flowering period
Mid-April to mid-June but mostly in May. It is
earliest in the west and exceptionally has been
recorded flowering in mid-March.
Range
Widespread in England, formerly north to a
373
1987-99
1970-86
pre 1970
line from Morecambe Bay to Holy Island in

Northumberland, although absent from much
of Devon and Cornwall and the Pennines.
Now it is greatly reduced and absent from
much of northern England, the Midlands, East
Anglia and the Home Counties. In Wales it
is largely confined to coastal regions, and in
Ireland to the central limestone belt, from Co.
Dublin west to Co. Galway and from southern
Co. Tipperary north to Co. Longford, with
M 10 May, Norfolk. Green-winged Orchid can occur in large numbers, especially on clay soils, on old commons and
greens, and on unimproved pastures and meadows
29/1/09 12:40:50
374
GENUS ANACAMPTIS
to flowering as the leaves are wintergreen.

Nowadays most colonies are on protected sites,
such as reserves and SSSIs or on private land
with a sympathetic landowner.
DESCRIPTION
M 12 May, Norfolk. The bold, parallel, green veins on the

sepals are diagnostic.
a few sites in Co. Cork and just one site in

Northern Ireland, on the coast of Co. Down.
It is very rare in Scotland, where it is found on
the coast of Ayrshire with an old record from
near Tomintoul in the Grampians. World
range: Predominantly European, extending
marginally into North Africa and the Middle
East. It ranges north to southern Norway,
southern Sweden (Gotland) and Estonia, east
to westernmost Russia and eastern Ukraine
and south to the Mediterranean, including
many of the Mediterranean islands, the Crimea
and Caucasus. Also Lebanon, Israel, Turkey,
northwest Iran and Morocco.
Height: 5-50cm but usually 7.5-15cm and

rather compact, and seldom more than 30cm.
Stem: Yellowish-green, washed purple
towards the tip and slightly angled, with
two or three thin whitish sheaths at the
extreme base.
Leaves: There are up to seven bluish-green
basal leaves, varying from roughly elliptical
to lanceolate, keeled and held semi-erect, and
two or three leaves higher on the stem that are
more pointed and become bract-like below the
spike. The leaves are wintergreen, emerging in
September-October and dying down by
mid-June.
Spike: Rather open and loose with four to 14
well-spaced flowers.
Bract: Green, typically washed purple, and
lanceolate in shape, two-thirds to twice the
How to find it
In favoured localities Green-winged Orchid
can still be found in thousands or even tens
of thousands; a site in Sussex, for example,
holds up to 50,000 flower spikes. Odd plants
can, however, be inconspicuous and are easily
missed, especially if they are very short. The
number of plants producing flowers varies
greatly from year to year, with a wet winter
and spring probably being most conducive
M 20 May, Norfolk. All the sepals and petals are arranged together to form a tight hood over the base of the lip.
29/1/09 12:40:56
GREEN-WINGED ORCHID
length of the ovary, which they curl around and

sheathe.
Ovary: Green, variably tinged purple,
cylindrical, strongly curved (through around
90), ribbed and twisted.
Flower: Mostly various shades of purple but
a few are rose-pink or whitish. The sepals are
oblong-oval, the upper a little narrower than
the lateral sepals. They are generally whitish
with a variable violet-purple wash, deepest
on the upper sepal and the upper margins of
the lateral sepals, which have three to seven
bold, green or bronze parallel veins that are
visible on both their upper and underside. The
petals are coloured as the upper sepal but are
shorter, narrower and more strap-shaped. The
petals and sepals form a hood that encloses
the column. The lip is rather broader than long
and divided into three lobes. The side-lobes are
rather larger than the central lobe, rounded,
often with crinkled (crenate) edges and folded
downwards. The central lobe is roughly equal
in length to the side-lobes. The lip is typically
violet-purple, whiter in the centre and at the
mouth of the spur, and there are usually some
violet-purple blotches and spots in this paler
central area. The column is whitish, washed
purple around the pollinia. The spur is coloured
as the lip, around half the length of the ovary,
narrow but flattened towards the tip, where it
is often notched and slightly curved upwards.
At least some flowers have a vanilla-like scent,
and the white-flowered forms may, on average,
have a stronger perfume although some people
cannot smell it at all.
Subspecies
None.

Var. alba has white flowers, sometimes very
faintly tinged pink, but still with green veins.
It is usually scarce (0.1-1% of plants) but
sometimes makes up as much as 15% of a
population.
Var. bartlettii is small with very small flowers
and var. churchillii is tall; neither seems well
defined.
375
M 24 May, Norfolk. Var. alba. A variable but usually very

small proportion of plants are white; the stripes on the
sepals are retained.
BIOLOGY
Bees, especially bumblebees, pollinate this
species. It produces no nectar and so is thought
to rely upon deceit to attract insects, although as
with Early Purple and Pyramidal Orchids it has
also been suggested that potential pollinators
can access sugary sap within the walls of the
spur. When the queen bees first emerge from
hibernation in the spring they are naive. They
have yet to learn which flowers offer a genuine
reward of nectar and are easily attracted by the
brightly coloured orchids. The bees usually visit
29/1/09 12:40:57
376
GENUS ANACAMPTIS

It is sometimes stated that Green-winged
Orchid is monocarpic and therefore usually
dies after it has flowered once. However, in one
population in Cambridgeshire some individual
plants flowered for 17 out of 18 years and
almost all flowered several times, interspersed
with seasons in which they merely produced a
rosette of leaves or more rarely were dormant
underground. At Iron Latch Meadow in
Essex, Green-winged Orchids reappeared and
flowered after an absence of about 30 years
when closed-canopy hawthorn scrub was
removed, showing just how long-lived this
species can be.
Seeds are thought to germinate in the
late summer or autumn, and the protocorm
probably produces the first roots and even the
first small leafy shoot during its first winter or
spring. The plant goes on to develop its first
tuber at the base of the leafy shoot, and it is this
tuber alone that will persist through the late
summer resting period, establishing the annual
cycle. It is thought that plants can flower within
three years of germination, and in cultivation
they have flowered after just two.
Hybrids
M 16 May, Norfolk. Some are very petite, with a few,

relatively large flowers.
a single flower on each of several plants before

they realise there is no reward on offer and move
on to more profitable species. On each orchid
visited, they start with the lower flowers of the
spike and therefore these are the ones that tend
to set seed. A cold spell in May, which disrupts
the bees routine, benefits the orchid because
the bees have to relearn their foraging routes
once the weather warms up again. Pollination
is variably efficient; in a study in Sweden only
5-30% of flowers set seed but in Britain almost
all the flowers may be pollinated, with each
capsule producing 4,000 individual seeds. The
species may also reproduce vegetatively via the
formation of additional tubers.
A. x alata, the hybrid with Loose-flowered

Orchid, has been recorded occasionally on
Jersey and once on Guernsey.
Orchis x morioides, the hybrid with Early
Purple Orchid, has been recorded rarely and
sporadically in England and Wales. As these
two species are now in different genera a new
name will be needed for this hybrid.

The specific name morio means dull or stupid.
The species was once known as fool stones.
Until very recently this species was
considered to be a member of the genus Orchis.
Its overall similarity to some of the members
of that genus, such as Early Purple Orchid,
certainly seemed to support this. Appearances
can be deceptive, however, and recent genetic
studies have shown that the Green-winged
29/1/09 12:40:58
GREEN-WINGED ORCHID
Orchid, together with Loose-flowered Orchid

of the Channel Islands, should join the
Pyramidal Orchid in the genus Anacamptis.
HISTORY AND
CONSERVATION
Thomas Johnson published his Mercurius
Botanicus (Botanical Mercury) with the first
localised record being published in 1660 when
John Ray noted the female Fool-stones from
Cambridgeshire in his Catalogus Plantarum
circa Cantabrigiam nascentium (A catalogue of
plants found around Cambridge).
Once common and widespread, Greenwinged Orchid has declined dramatically in the
last 50 years. It has been lost from many sites
and has gone from 49% of the historical range
in Britain and 60% in Ireland. It is classified
as Near Threatened in Britain and is specially
8 of the 1985 Wildlife Order (NI) and in Eire
under the Flora (Protection) Order.
377
decline significantly if fertilisers are applied to

grassland, probably due to competition with
the increasingly vigorous grass and herbs, and
the phosphorus in fertilisers may actually be
toxic to the orchids. If anything, habitat losses
have accelerated in the last 20 years as the last
few pristine pastures have been improved;
overall the decline in the amount of suitable
habitat must approach 99%. A typical example
is Cambridgeshire, where prior to 1900 at
least 36 sites held Green-winged Orchids.
In the period 1901-85 this figure fell to 13
and between 1986 and 1990 the population
collapsed to just five sites.
Past and present occurrence of Green-winged Orchid in

Atlas).
Britain
Ireland
939
124
479 (17%*)
50 (5%*)
% lost, 1500-1969
38%
55%
% lost, 1970-1986
11%
5%
% lost, total
49%
60%

1500-1999
current range
The decline is almost entirely due to

agricultural changes. In the 19th century its
favoured damp meadows and pastures were
ploughed and converted to arable. In the 20th
century, especially since 1945, tractors have
replaced horses, and permanent pastures
have become less and less a feature of most
farms. Those that have survived have been
drained, ploughed and reseeded. Research
has shown that Green-winged Orchid will
M 10 May, Norfolk. This delicate rose-pink is rather

unusual.
29/1/09 12:40:59
Genus SERAPIAS
TonGue orchids
29/1/09 12:41:03
GENUS SERAPIAS
379
This genus has only a toehold in Britain and Ireland, with two species recorded in recent
years from the coast of southwest England. We consider that a natural origin, via windblown seed, is the most likely explanation for their occurrence, although they have also been
treated, albeit without any evidence, as deliberate introductions.
Distribution
Serapias is basically a Mediterranean genus,
distributed from the Caucasus Mountains
through the Mediterranean basin west to the
Canary Islands and Azores and extending north
to Brittany. All the species are closely related
and thus there is considerable disagreement
as to the exact number. Between seven and 27
species are recognised in recent literature and
consequently there are considerable problems in
reaching a correct identification.
Classification
Genetic evidence shows that the closest
relations of the tongue orchids are the bee and
spider orchids Ophrys, the Pyramidal Orchid
group Anacamptis and the lizard orchids
Himantoglossum.
The genus Serapias is often divided into
three groups: the self-pollinating Smallflowered Tongue Orchid S. parviflora and allies;
the vomeracea group (the largest) and the lingua
group, which includes Greater Tongue Orchid,
and is distinguished by having a single dark
boss at the base of the lip (the other groups
having two bosses).
Floral structures
The column is slender, directed forwards and
tapers to a long, thin point. There are two
pollinia, each narrowing down into a caudicle
(stalk). Both pollinia are attached to the single
viscidium, which in turn is housed in a bursicle.
There is no spur.
Pollination
The vomeracea and lingua groups are crosspollinated. The flowers have no nectar and
are thought to lure insects, especially bees and
beetles, by providing them with shelter, both
overnight and during bad weather. The dark
O 27 April, Extremadura, Spain. Greater Tongue Orchid.
colour of the flower may help it to absorb heat,

and its internal temperature may be 1-3C
higher than that of its surroundings. It seems
likely, however, that there is more involved in
the pollination process than merely the offer
of shelter.
The tongue orchids are closely related and
genetically very similar. The various species
need to be isolated reproductively from one
another to have evolved in the first place and to
continue to maintain their separate identities.
A mechanism that involved their pollinators
would be one way for this to happen but if
they merely offer shelter there would be no
mechanism, apart from the size of the flower,
to stop insects from carrying pollen from the
flowers of one species to another. Indeed, this
might be why there are so many hybrids that
involve the various tongue orchids. But, so far
Greater Tongue Orchid S. lingua is the only
species where additional isolating mechanisms
have been observed. Small bees, apparently
attracted to the flower by pheromones, attempt
pseudocopulation with the flower (see also the
notes on the genus Ophrys, p.388). In these
cross-pollinated species pollination is apparently
inefficient and although their flowers are selfcompatible, self-pollination does not occur.
The parviflora group, which includes Smallflowered Tongue Orchid, is self-pollinated.
Growth pattern
At flowering time there are two to five moreor-less spherical tubers at the base of the aerial
stem, the one to three youngest at the end of
long, slender rhizomes.

Following germination, the protocorm produces
its first root in the autumn and the first leaf
appears shortly afterwards. The following
spring the first tuber starts to grow from a bud
29/1/09 12:41:04
380
GENUS SERAPIAS
at the base of the leafy shoot, and during the

summer resting period the leaf and root die
off, leaving just the tuber. From now onwards
the usual sequence of annual tuber replacement
takes over. In cultivation plants may flower
three years after seed is sown, occasionally after
just two.
Additional daughter tubers are formed at the
base of the aerial stem, usually at the end of
slender rhizomes. These will go on to form
separate plants as the connecting rhizome dies
off in the summer.
Name
The name of the genus, Serapias, is derived
from Serapis (or Osirapis). This is a composite
Graeco-Egyptian god combining the attributes
of Apis (the scared bull) and Osiris (ruler of
the underworld). Serapis attracted an orgiastic
cult and in about AD 64 the Greek scholar
Dioscorides, considered one of the fathers of
botany, gave the name Serapias to an orchid that
was reputed to be an aphrodisiac. The orchid in
question was probably actually a member of the
genus Orchis.
upper sepal
petal
upper sepal
petal
bosses
hypochile
side-lobe
lip
epichile
M Small-flowered Tongue Orchid (After Delforge 2005).
lateral
sepal
boss
lateral
sepal
hypochile
side-lobe
lip
epichile
M Greater Tongue Orchid.
29/1/09 12:41:04
SMALL-FLOWERED TONGUE ORCHID
381
smaLL-fLowered TonGue orchid

Serapias parviflora
This species is one of the most recent additions to the British flora, having been found for
the first time in 1989. It is confined to just one site in Cornwall, where a handful of plants
flower most years, but it could turn up almost anywhere on the south coast.
Identification
The Serapias or tongue orchids are very
distinctive, with the sepals, petals and base of
the lip forming a tight tube from which the
outer part of the lip, usually coloured a rustybrown, sticks out like a pointed little tongue. In
Small-flowered Tongue Orchid the flowers are
relatively small (indeed, they are the smallest
of the Serapias), with a small lip, the projecting
tip of which is 6-11mm long x 3.5-5mm wide.
There are two small dark parallel raised bosses
at the base of the lip, and the flowers are selfpollinated in bud, so that when they open the
pollinia have already disintegrated onto the
stigma.
Islands, and including Crete, Sicily, Corsica,

Sardinia and the Balearic Islands, and also
Morocco, Algeria and Tunisia in North Africa.
In France it occurs on the Mediterranean
littoral with a separate centre of distribution on
the Atlantic coast north to Brittany.
How to find it
The British population comprises just a few
plants that are very vulnerable to disturbance
and damage; indeed, flowers have already been
unwittingly broken off by admirers. This is one
orchid that is best left alone, at least until a
more substantial population exists.
Similar species
Greater Tongue Orchid has larger flowers with
a single dark boss on the lip.
Habitat
In Cornwall it is found in short, rabbit-grazed
grassland among scattered gorse on a southfacing slope near the sea. In Europe it is found
in dry and wet grassland, dune slacks, open pine
woods, scrub and olive groves, on calcareous to
mildly acidic soils.
Flowering period
Early May to early June.
Range
Known from a single site near Rame Head
in southeast Cornwall. World range: The
Mediterranean basin from Cyprus and the
Aegean Islands west through Greece, Italy,
France, Spain and Portugal to the Canary
P 7 May, Cornwall (Mike Frost). Although currently restricted to one site in Britain, this species can be expected
to appear elsewhere.
29/1/09 12:41:05
382
GENUS SERAPIAS
DESCRIPTION
BIOLOGY
(based on European populations)

Stem: Green, blotched or streaked with red.
Leaves: At the base of the stem there is a
cluster of four to seven erect, long, narrow,
strap-shaped leaves with a distinct keel. Higher
on the stem there are one to three smaller
bract-like leaves. In Europe the leaves appear
in autumn, overwinter and die down after
flowering.
Spike: Three to eight flowers (rarely as many
as 12) form a narrow spike with an indistinct
spiral arrangement, with each flower held
horizontally or angled upwards to 45.
Bract: Pinkish-brown washed grey and with
distinct dark veins; the bracts are lanceolate
and held erect, clasping the ovary. They are as
long as the ovary and hood put together or just
a little shorter, although the lower bracts are
sometimes much longer than the flower.
Ovary: Green, long, narrow, and held
upright.
Flower: The sepals are lanceolate and a pale
dusty-pink washed grey with distinct dark
veins. The petals are a little shorter, tapering
abruptly to a long narrow point, and purplishbrown at the base. The sepals and petals form
a tight hood over the base of the lip and the
column. The lip is 14-19mm long (extremes
13-22mm) and reddish-brown. The inner half
(the hypochile) is heart-shaped with two small,
raised, parallel, dark purple bosses at the base.
The sides, which are almost black, curl up and
round to form a gutter that is totally enclosed
by the hood. The floor of the lip has short,
sparse, whitish hairs or is sometimes almost
hairless. The outer half of the lip (the epichile)
is lanceolate, much narrower than the base
and bent down and under the remainder of
the flower.
Subspecies
The flowers are cleistogamous, that is selfpollinated in bud. The yellow pollinia have
already disintegrated onto the stigma by the
time the flowers open.

No specific information. Stems usually arise
singly, and vegetative reproduction may be
relatively unimportant.
Hybrids
None.

The specific name parviflora means simply small
flower.
HISTORY AND
CONSERVATION
First recorded in May 1989 near Rame Head in
Cornwall, when two spikes were found. Three
spikes appeared in 1990, with five each in 1991
and 1992, although photographers squashed
two spikes in the latter year. There was then
a gap of at least four years during which no
plants were found, but subsequently two or
three spikes have appeared most years up to
and including 2004. Seedlings germinated at
the Royal Botanic Gardens, Kew, from Cornish
seed were planted at two nearby sites in 1993
but did not survive.
It has been suggested that the plants in
Cornwall were deliberately introduced, but
there is no evidence for this whatsoever.
The plants were found hidden among gorse
scrub, well away from any footpaths and in
a locality highly unlikely to be chosen for an
introduction. It seems much more likely that
the plants arose naturally from wind-blown
seed. Whether this tiny population persists
or the species appears elsewhere, only time
will tell.
None.

None.
P 1 June, Cornwall (Robin Chittenden www.harlequinpictures.co.uk)
29/1/09 12:41:07
29/1/09 12:41:10
384
GENUS SERAPIAS
GreaTer TonGue orchid

Serapias lingua
This species has only the most tenuous claim to be British, having been found on Guernsey
in 1992 and in south Devon in 1998. These may be the vanguard of further discoveries,
however, if the climate becomes more Mediterranean due to global warming.
Identification
The tongue orchids are distinctive, with the
sepals, petals and base of the lip forming a tight
tube from which the tip of the reddish lip sticks
out like a tongue. The identification of the
various species is often rather difficult as they
are very similar, and there are also many hybrid
populations. It often requires the dissection
of a flower, but the conspicuous single dark
purple boss at the base of the lip, resembling a
coffee bean, easily identifies the Greater Tongue
Orchid and its allies. All the other Serapias
have two bosses. Within the lingua group,
identification depends on observation of the
precise shape of the boss, the width of the lip
and the overall colour of the flower.
Similar species
Small-flowered Tongue Orchid has a smaller
flower with two bosses at the base of the lip and
is self-pollinated.
Habitat
On Guernsey it was found on dry natural
grassland on the edge of a golf course. In south
Devon it was found in a meadow that had been
sown with a grass mixture in the 1970s and was
normally cut for hay in May or June. In 1998,
when the Greater Tongue Orchid was found,
it was not mown. In Europe it is found in poor
grassland, wet meadows, dune slacks, scrub,
olive groves and open woodland.
Flowering period
May to June.
Range
Found at single sites on Guernsey and in Devon.
World range: The Mediterranean basin, from
Greece and the former Yugoslavia west to
Portugal, also the Aegean Islands, Crete, Sicily,
Malta, Corsica, Sardinia and the Balearics. In

North Africa recorded from Morocco, Algeria
and Tunisia. In France it occurs north to a line
between Nantes and Lyon.
How to find it
Like the Small-flowered Tongue Orchid this
species is confined to just one site in Britain,
with just a few plants that are vulnerable to
disturbance and damage. It is best left alone.
DESCRIPTION
(from European plants)
Stem: Green, blotched or streaked with red.
Leaves: At the base of the stem there is a cluster
of four to eight erect, long, narrow, strap-shaped
leaves that have a distinct keel. Higher on the
stem there are one to three smaller bract-like
leaves. In Europe the leaves appear in autumn,
overwinter and die down after flowering.
Spike: The two to six flowers, rarely as many
as eight, are arranged around the stem in a
compact spike.
Bract: Lanceolate, held erect and clasping the
ovary, they are shorter than the ovary and hood,
put together. The bracts are reddish or, more
rarely, green tinged red towards the tip, with
distinct purple veins.
Ovary: Green, long, narrow and held upright.
Flower: The sepals are lanceolate and pale lilacgrey with distinct purple veins. The petals are a
little shorter, tapering abruptly to a long narrow
point, and are more purplish-brown at the base.
The sepals and petals form a tight horizontal
hood over the base of the lip and the column.
The lip is 22-29mm long (sometimes only
P 27 April, Extremadura, Spain. The origin of the English
name tongue orchid for the members of the genus
Serapias is obvious.
29/1/09 12:41:10
GREATER TONGUE ORCHID
385
29/1/09 12:41:13
386
GENUS SERAPIAS
16mm), variable in colour from yellowish to

fleshy-pink or reddish and only sparsely hairy.
The basal portion (hypochile) is kidney-shaped
with the side-lobes dark purple, rather angular
and curling up and around to form a gutter that
is totally enclosed by the hood. There is a single
shining, dark purplish-brown raised boss at the
base of the hypochile, often crescent-shaped
with the two horns of the crescent pointing
backwards and elongated into two fine ridges. It
is ungrooved or, at most, only slightly grooved.
The outer half of the lip (epichile) is broadly
lanceolate and slightly longer and narrower
than the hypochile. It is usually 13-18mm long
x 7-12mm wide but can be as small as 8mm
x 4mm. It is held pointing downwards and is
variably bent backwards below the rest of the
flower. The pollinia are greenish-yellow and the
flowers are strongly scented.
Subspecies
Disputed. More conservative taxonomic
treatments consider that Greater Tongue
Orchid has several subspecies, including S. l.
durieui. More radical treatments, including
Delforge (2001), treat durieui as a synonym
of a different species, S. strictiflora, that is
found from eastern Algeria to Morocco and in
southern Portugal and southwest Spain. This is
of some relevance because the plants in south
Devon were identified as subspecies durieui on
the basis of a narrow lip and longer and more
elegant sepals and petals. If this identification
was correct and if the more radical classification
is accepted, Serapias strictiflora would be on the
British list rather than S. lingua.

None in Britain.
BIOLOGY
It is thought that most Serapias are pollinated
by small bees and other insects which use
the flowers for shelter. However, the Greater
Tongue Orchid may take this strategy one stage
further. The flower mimics a nest hole and
lures small male bees, which emerge before the
females, with its powerful scent. In addition,

the large shiny boss at the base of the lip also
stimulates pseudocopulation, during which the
pollinia are attached to the head of the bee as it
attempts to copulate with the flower. This twopronged strategy enables the orchid to attract
bees both at night and on sunny days.
Vegetative reproduction may be an
important means of reproduction and clusters
of flower spikes are frequent.

Hybrids
None in Britain.

The specific name lingua means tongue.
HISTORY AND
CONSERVATION
A single plant was found on the western
side of Guernsey in May 1992 but it has not
reappeared at this site. In June 1998 three
flower spikes were found near the coast in the
vicinity of Kingsbridge in south Devon. They
were identified as the subspecies durieui and it
is speculated that the seed could have arrived
with Saharan sand from North Africa.
As with all the Serapias orchids that have
been found in Britain, some botanists are
sceptical as to its origins. They presumably
believe that the plants originated from seed that
came from an artificial source, such as a nearby
plant collection or agricultural seed mix, or that
deliberate fraud could be involved, with seed
being sown or even mature specimens planted
out. An artificial source of seed is certainly
possible (see Heart-flowered Tongue Orchid,
p.416) but in both Greater and Small-flowered
Tongue Orchids this seems highly unlikely,
while there is no evidence of fraud. Given the
occurrence of Greater Tongue Orchid in much
of southern France, including the Atlantic
coast south of Vende, a natural arrival in the
Channel Islands or southern England seems
eminently likely. Whether the species becomes
established is, however, a different matter.
29/1/09 12:41:14
Genus OPHRYS
Bee and sPider orchids
29/1/09 12:41:18
388
GENUS OPHRYS
This genus includes the most exotic and intriguing of British and Irish orchids, Bee and Fly
Orchids and Early and Late Spider Orchids. Their flowers have evolved to mimic insects,
predominantly bees (but definitely not spiders). In addition to their overall appearance, the
pheromones produced by the flowers and the distribution of hairs on the lip are critically
important in perpetuating the deception. Male bees are attracted to the orchids and attempt
to mate with the flowers in a process known as pseudocopulation. In doing so they carry
pollinia from one flower to the other and become unwitting pollinators.
1
Distribution
Pollination
Containing more than 250 species, this is

undoubtedly the largest genus of European
orchids. Its distribution is centred on the
Mediterranean basin, but Ophrys can be found
from the Canary Islands in the west to the
Caspian Sea in the east, Fly Orchid ranges
north to Scandinavia and other species extend
into North Africa. However, just four species
occur in Britain and Ireland.
The number of species recognised in this
genus has risen sharply; only around 60 were
listed in the 1980s. This reflects the increasing
appreciation of the importance of the highly
specialised pollination mechanism in the
evolution of the various species.
All species in this genus are cross-pollinated

with the exception of one, the Bee Orchid,
which is routinely self-pollinated. The Bee
Orchid had clearly evolved to use the same
mechanism as the other species, with an
elaborate pattern and texture to its lip, but
has largely abandoned cross-pollination,
presumably relatively recently.
In the genus Ophrys the flowers have no
nectar and offer their visitors no reward.
Instead, the orchid exploits the sex-drive of
insects with a combination of visual, olfactory
and tactile deceits. The pollinators are usually
bees and they are initially attracted to the
flowers by chemical signals released by the
lip of the orchid that mimic the pheromones
produced by virgin female bees. These
pheromones are complex cocktails of volatile
compounds that stimulate mating behaviour in
the male bees. Not only are the flowers version
of the bees pheromones an exact copy of the
appropriate compounds but also the orchid
produces such large quantities that the male
bee may even prefer the flowers deceit to the
real thing.
Male bees emerge from hibernation earlier
in the spring than females, and it is thought
that the orchid exploits this gap. In the absence
of females the orchid is able to lure the males to
Classification
DNA evidence suggests that the closest
relatives of the genus are the tongue orchids
Serapias, the lizard orchids Himantoglossum and
the Pyramidal Orchid group Anacamptis.
Floral structures
The stiff and often elaborately designed lip
renders the members of this genus distinctive.
It is usually prominently hairy in places and
velvety in others with a smooth, hairless central
patch, the speculum. This is sometimes shiny,
reflecting the light and probably therefore
simulating an insects wings. As in most orchids
the lowermost flowers open first but the spike
continues to elongate, sometimes substantially
so, as more flowers open.
The column is long and erect. There are two
pollinia, each attached by a caudicle (stalk) to a
separate sticky viscidium. These are concealed
and protected within separate bursicles.
P 1. O. levantina; 2. O. umbilicata; 3. O. flavomarginata;

4. O. (f.) iricolor; 5. O. lutea galilaea; 6. O. omegaifera
israelitica; 7. O. aesculapiiformis; 8. O. (f.) fusca; 9. O.
mammosa; 10. O. argolia elegans; 11. O. bormuelleri;
12. O. kotschyi. All Cyprus, March. The intimate relationship with a specific insect pollinator has powered the evolution of numerous species.
O 14 June, Kent. Late Spider Orchid
10
29/1/09 12:41:19
10
11
12
29/1/09 12:41:23
390
GENUS OPHRYS
its flowers but once the females have emerged

the orchids are ignored in preference to the real
thing. In at least some species, however, this is
not the case and the flowers continue to receive
visits from the males even after the female bees
have emerged.
The male bees home in on the orchids
pheromones, and as they get closer they catch
sight of the flower and land on the lip. All three
deceits now come into play. The male bee is
still stimulated by the scent of the flower and
the patterning on the lip but now also by the
texture of the various hairy, velvety and smooth
portions of the lip. This stimulates the bee to
orientate himself into the correct position,
extend his genital apparatus and attempt
copulation. Of course, the correct position is
the one in which he will pick up pollinia in
the first flower visited and deposit the pollinia
on the stigma of the second flower and so on.
Eventually this process of pseudocopulation is
interrupted and the bee leaves. The male bees
may show signs of exhaustion but ejaculation
of sperm has not been observed, although it
has been recorded in a similar process involving
an ichneumon wasp and the Australian orchid,
Cryptostylis leptochila.
The orchid is totally dependent on the bee.
If there is no pollination, there will be no seed
and therefore no more orchids; a powerful
engine to drive the evolution of the orchid.
Against a noisy background of multitudes
of chemical signals, each species of bee must
be able to produce a unique pheromone in
order to attract potential mates and reproduce
successfully. It may do this by having a unique
combination of relatively common chemicals
in its sex pheromones or by using chemicals
that are in themselves unique. Bees use both
systems and the various Ophrys orchids are able
to mimic both systems, too. For the orchid,
the better and more precisely it mimics its
pollinator the better its chance of successful
pollination. Each species of orchid therefore
evolves to mimic the pheromones of a particular
species of bee in order to maximize its chances
of attracting pollinators. In this way the genus
Ophrys has evolved many species, reflecting the

many species of potential pollinator. Each one is
precisely matched.
As a further complication, differences
in the patterning and shape of the lip and
the distribution of hairs allow two distinct
species of Ophrys to share the same species
of pollinating insect. The position and
direction of the hairs on the flowers are
critical in the positioning of the insect during
pseudocopulation. One species of orchid may
stimulate the bee to have its head uppermost,
facing the column and so picking up the
pollinia on its head. A second species of
orchid stimulates the bee to have its abdomen
uppermost, and the pollinia are attached to the
tip of its abdomen. In the respective flowers, the
pollinia and stigma are positioned according to
the orientation of the bee.
The orchid-pollinator match in Ophrys
is very close but not always perfect. All the
members of the genus are interfertile, and
there are a large variety of hybrids. It is a
reflection of how good the system is, however,
that these hybrids tend to occur rarely and in
very small numbers.
Growth pattern
At flowering time all Ophrys species have two
more-or-less spherical tubers side by side at
the base of the aerial stem together with some
short, thick roots. The roots appear in the
autumn, simultaneously with the leaves, and
are infected with fungi. The pattern of growth
and the annual replacement of tubers are very
similar to that of the genus Orchis (see p.200).
Fungal partners
In common with the other tuberous orchids,
fungal activity takes place in the roots and
rhizome but not in the tuber (indeed, the tuber
contains the orchids food reserve and may
even be protected from fungal infection by
fungicide, see p.9). As the orchid passes the
summer resting period as a tuber this presents
a problem. How can the fungal infection be
carried from one year to the next? In damper
climates there is a good chance that the fungi
29/1/09 12:41:23
GENUS OPHRYS
can persist in the soil but in the hot, dry,

Mediterranean summers adult fungi cannot
live in the parched soils and must survive as
spores. To guarantee continuity the members
of the genus Ophrys probably carried their
fungal infection from one year to the next in
the surface layers of their tuber. Observations
of Late Spider Orchid show that the terminal
bud on the tuber starts to grow at the same
time as the onset of wet weather in the autumn.
Simultaneously the fungal mycelium starts
to grow outwards from certain points on the
tubers surface. The orchids new roots grow
outwards through this fungal mycelium and
the fungus probably enters the roots through
the root hairs. As the next seasons tuber starts
to develop it too is infected (and will carry the
fungus through the next resting period) via the
soil, through minute, hair-like projections on
its surface.
391
Greek for eyebrow, perhaps an allusion to the

hairy lip (or the hairy fringe to the petals).
Alternatively, Ophrys was the name given in
Ancient Greece by Pliny the Elder to a small,
two-leaved plant used to dye eyebrows and hair.
The plant in question was probably Common
Twayblade, and Plinys name Ophrys may have
derived from a Greek word for snake due to the
similarity of the flowers of Common Twayblade
to the head and tongue of a snake.

Seed probably germinates in the spring
to produce a rounded protocorm. In Late
Spider Orchid and probably other species the
protocorm produces a root in the first autumn
and then a small tuber the following spring.
In the summer resting period the protocorm
and root die off, leaving just the tuber. In
the autumn the bud on the top of the tuber
produces a short rhizome and roots. The first
leafy shoot appears in the second spring. In
cultivation flowering plants can sometimes be
produced within three years of germination.
Additional tubers can be formed at the base
of the aerial stem, perhaps often at the end
of slender rhizomes, and these will go on to
form separate plants as the connecting stem
dies off in the autumn. It seems, however, that
vegetative reproduction is either rare (Late
Spider Orchid) or uncommon (Early Spider
Orchid) among the British species.
Name
The origin of the generic name Ophrys is
contentious. It may derive from ophrus, the
M Unlike all the other members of the genus Ophrys,

Bee Orchid is self-pollinated. Being independent of insect
pollinators, it has done well in Britain, and is one of the
most widespread and successful species.
29/1/09 12:41:25
392
GENUS OPHRYS
fLy orchid
Vulnerable, BAP
Ophrys insectifera
Formerly: Ophrys muscifera
Fly Orchid shows the most wonderful example of insect mimicry among British orchids, but
rather than flies it has evolved to lure male digger wasps to act as inadvertent pollinators as
they attempt to copulate with its flowers. For human observers the small flowers are often
hard to spot but once seen have a magnetic attraction, drawing the eye back again and again.
Fly Orchid is relatively common but inconspicuous in woodland and scrub on chalk in
southern England and occurs at scattered localities in a variety of lime-rich habitats elsewhere
in Britain north to Cumbria and Anglesey. It is also found in central and western Ireland.
Identification
Very distinctive, the individual flowers are
indeed like little flies. The purplish-brown lip
forms the body and the lustrous slate-blue
speculum shines like folded wings. The two
glistening depressions at the base of the lip are
the eyes and above these the dark, wire-like
petals look just like little antennae.
Similar species
None.
Habitat
Very varied, although it usually grows on
calcareous soils over chalk and limestone. It
is found in open deciduous woodland and in
southern England is particularly associated
with beechwoods. It favours the better-lit areas
in glades, rides and along the edge of woodland,
as well as shaded road banks and open scrub,
but sometimes grows in deeper shade such as
overgrown hazel coppice. In southern England
Fly Orchid is infrequently found on open
grassland in old pits, quarries and on spoil
heaps, but when it does grow in such habitats
it may occur in large numbers; it has been
recorded rarely from slumped coastal cliffs. In
northern England and Ireland it is probably
more frequent in open areas and in addition to
wooded sites is found on limestone pavements
and rocky hillsides. It grows in alkaline fens and
O 23 May, Co. Clare. In the west of Britain and Ireland
Fly Orchid will grow in the open, on limestone pavements
and in fens.
29/1/09 12:41:26
FLY ORCHID
393
on the margins of turloughs in western Ireland

and also in fens on Anglesey, often among
tussocks of Black Bog-rush. Recorded up to
390m above sea level (Helbeck Wood, Brough,
Cumbria).
Flowering period
Late April to early July but mostly in late May
and early June.
1987-99
1970-86
pre 1970
Range
It is widespread but very local. In southern
England Fly Orchid is more-or-less confined to
the North and South Downs in Kent, Sussex
and Surrey, the chalk of Hampshire, Dorset,
Wiltshire and the Chilterns, and also the
Cotswolds. There are scattered records away
from these areas in Somerset, Warwickshire,
Northamptonshire, Suffolk, Nottinghamshire,
Derbyshire and more widely in Yorkshire and
the Morecambe Bay area of north Lancashire
and Cumbria. The northernmost sites are in
the upper Eden Valley of the north Pennines
in Cumbria. In Wales it is now confined to
Anglesey and in Ireland to a central belt from
southern Co. Mayo, Co. Galway and Co.
Clare, through north Tipperary, Co. Offaly
and Westmeath to Laois and Co. Kildare.
P 27 May, Hampshire. Fly Orchid can grow in heavy
shade but visits by suitable pollinators are probably more
likely where it catches the sun for part of the day.
29/1/09 12:41:28
394
GENUS OPHRYS
29/1/09 12:41:31
FLY ORCHID
395
World range: Confined to Europe, where it has

the most northerly distribution in the genus
Ophrys, ranging north to c. 67 in Norway
and to central Sweden, southern Finland and
the Baltic States. Occurs south to Spain, the
Balearic Islands, Italy, northern Greece and
Romania and east to the Moscow region.
How to find it
The tall slender spikes of Fly Orchid, with their
small, well-spaced flowers, can be very hard to
see among other vegetation and even on a bare
woodland floor they can vanish with ease. But,
if you can get your eye-in where one Fly Orchid
is found, there may be plenty more. Within the
range in southeast England it can be found at
many woodland and scrubby sites on chalk or
limestone. In northern England sites include
Whitbarrow Scar (Cumbria) and Castle Eden
Dene (Co. Durham) and in Ireland it is locally
common in The Burren (Co. Clare).
DESCRIPTION
Height: 15-60cm.
Stem: Pale green and slender, with one or two
basal sheaths. Groups of up to ten plants may
grow together.
Leaves: There are two to five shiny, dark green
or bluish-green leaves. The lower are narrow
and strap-shaped, flaccid but keeled and usually
pointed at the tip, and the upper one or two
are narrower, more pointed and loosely sheathe
the stem. The leaves emerge in autumn and are
wintergreen.
Spike: Although initially bunched, the one to
ten flowers (exceptionally as many as 20) are
well-spaced along the stem by the time the
uppermost has opened.
Bract: Dark green or bluish-green and
lanceolate, often with the edges rolled inwards;
the lower bracts are rather longer than the
ovary but towards the tip of the spike they are a
little shorter.
Ovary: Pale green, slender, cylindrical, sixribbed and held upright but curving at the tip
to hold the flower facing outwards.
Flower: The sepals are yellowish-green and are
M 23 May, Co. Clare. The flower has evolved to mimic

digger wasps.The petals form the antennae and the bluish
speculum on the lip resembles the glint of the wasps
folded wings.
oval-oblong with a blunt tip but look narrower

because their edges are rolled back and the
inner face is concave. The lateral sepals are held
horizontally and the upper sepal is vertical but
arches forward over the column to a variable
extent. The petals are dark purplish-brown
with short, fine, downy hairs and are much
smaller than the sepals (less than half as long);
their edges are rolled back to give them a fine,
filiform appearance and they point forward.
The lip is longer than wide and hangs down
almost vertically. It is divided into three lobes,
with two relatively short, narrow side-lobes
spreading outwards at the base and a broad
terminal lobe that is in turn notched or forked
at the tip. The lip is velvety in texture (the
side-lobes are hairier), rich dark reddish-brown
29/1/09 12:41:32
396
GENUS OPHRYS
P 10 May, Kent. The

male digger wasp, excited
by the pheromones produced by the flower, as
well as by its appearance
and texture, attempts
pseudocopulation with
the orchid and in the
process acts as pollinator.
or purplish-brown, becoming a little paler

towards the tip and duller with age. A more-orless square, shining, pale slate-blue band across
the centre forms the speculum, and at the base
of the lip there are two shining pseudoeyes.
The column is short and reddish-brown with
a small, circular stigmatic cavity at its base and
a short, blunt beak. The two yellow pollinia
have their viscidia enclosed in two off-white
bursicles.
Subspecies
None.

There is considerable variation in the colour
and markings of the lip.
Var. ochroleuca has a lip that is pale yellowishgreen with a white speculum. It has been
recorded from Kent, Hampshire, Wiltshire and
Hertfordshire.
Var. flavescens is a little darker than var.
ochroleuca, with a yellowish-brown lip, pale blue
or whitish speculum and greenish-brown petals.
It has been recorded from Gloucestershire.
Var. subbombifera has a very broad, rounded
central lobe, notched as usual at the tip. Very
29/1/09 12:41:34
FLY ORCHID
rare, it has been recorded from Surrey and

Hampshire.
Var. parviflora has flowers that are around half
the normal size. It is very rare.
Var. luteomarginata has a broad yellow border to
the central lobe of the lip, often also a yellow stain
or tip to the side-lobes and yellow or green tips to
the petals. It has been recorded rarely in Surrey,
Hampshire, Gloucestershire and Anglesey.
BIOLOGY
Pollination is by male digger wasps Argogorytes
mystaceus and also A. fargeii, a species that is rarer
and emerges a little later in the season. The wasps
are attracted by pheromones emitted by the
orchid and by the shape and texture of the flower.
They attempt to copulate with it and during this
pseudocopulation the pollinia are attached to
their heads. Pollination rates are low, with less
than 20% of flowers setting seed (Summerhayes,
1968 quotes rates of 2.1% and 7.5% for two
samples of approximately 1,000 plants).

The aerial stem grows from a pair of tubers, the
younger of which is often stalked. The first leaf
appears in the winter after germination and the
first tuber is formed in the second year.
Hybrids
x Bee Orchid has been recorded at two sites
in the Avon Gorge in Somerset and one site in
West Sussex, but it is very rare. It is sometimes
called O. x pietzschii but this is an invalid name.
O. x hybrida, the hybrid with Early Spider
Orchid, has occurred occasionally in Kent.

The specific name insectifera derives from the
Latin insecta meaning insect and fero meaning
bearing, hence insect-bearing.
HISTORY AND
CONSERVATION
John Gerard published The Herball, or General
397
Historie of Plantes in which he noted: The

Bee, the Fly and the Butterfly Satyrions grow
upon barren chalky hilladjoining to a village
in Kent named Greenhithe, upon Longfield
downs by Southfleet. likewise in a field
half a mile from S. Albons.
There have been considerable losses
since 1597 with the species now occupying
only 42% of its historical range in Britain,
and it is classified as Vulnerable. Much of
the decline took place long ago, especially in
East Anglia. However, in Ireland, where Fly
Orchid is now found in just over 50% of its
historical range, rather more of the losses
have been recent. Causes include woodland
clearances and coniferisation but perhaps
equally important has been the maturation
of woodland and scrub due to changes in
forest management or its abandonment
altogether. In such cases increasing shade
probably means that the flowers are seldom
pollinated and the population declines and
eventually disappears. Fly Orchid is either
extinct or has not been reported recently from
Devon, the Isle of Wight, Middlesex, Essex,
Norfolk, Cambridgeshire, Huntingdonshire,
Worcestershire, Herefordshire, Shropshire,
Staffordshire, Leicestershire, Lincolnshire and
Co. Durham. Similarly in Wales there have
been no recent reports from Monmouthshire,
Glamorganshire and Denbighshire. There is an
old, unconfirmed record from near Killiecrankie
in Perthshire in Scotland.
Past and present occurrence of Fly Orchid in Britain and
Britain
Ireland
264
31
110 (3.9%*)
16 (1.6%*)
% lost, 1500-1969
47.5%
26%
% lost, 1970-1986
10.5%
22.5%
58%
48.5%
total historical
range, 1500-1999
current range
% lost, total
29/1/09 12:41:35
398
GENUS OPHRYS
Bee orchid
Ophrys apifera
Looking, even feeling exotic and very special, the Bee Orchid is a standard bearer for orchids
and orchid conservation. Even for those with just a passing interest in wildlife, the discovery
of a Bee Orchid can be a special event, and people take great pride in their local Bee Orchids.
An opportunistic and adaptable species, it often turns up in unexpected places; indeed, it
has even taken to garden lawns in recent years. Well named, the flowers look very much like
a bee and have evolved to attract male bees as pollinators. The mechanism has, however,
been abandoned, and almost all plants are now self-pollinated, a process that produces
copious seed, no doubt part of the secret of its success. It is widespread in England and parts
of Wales and Ireland, occurring in a wide variety of open grassland sites. Bee Orchid is the
county flower of Bedfordshire.
Identification
Unmistakably a bee orchid, the flower looks
like a bumblebee. It has large pink sepals and
slender, parallel-sided, greenish or pinkishbrown petals that form the bees antennae. The
lip is a velvety maroon-brown with a pattern of
creamy markings and noticeably hairy rounded
side-lobes.
Similar species
In most of Britain and Ireland it is the only
bee orchid and therefore distinctive, but in
southern England care should always be taken
to distinguish it from Early and Late Spider
Orchids.
Late Spider Orchid is superficially very
similar but usually has a broader, more
rectangular lip with a distinctive forward or
downward pointing yellowish nib at its tip. In
the Bee Orchid the tip of the lip is almost always
tucked under and out of sight. In addition, the
petals of Late Spider Orchid are always pinkish,
short and very distinctly triangular or conical
in shape. Late Spider Orchid is not usually selfpollinating and its pollinia never dangle loose.
Early Spider Orchid has yellowish sepals
and petals and a shiny slate-blue speculum on
its lip. It lacks any creamy or yellow markings.
O 24 June, Norfolk. As successive flowers open, the spike
lengthens. The pointed tip of the lip is normally tucked
under and out of sight.
29/1/09 12:41:36
BEE ORCHID
Habitat
Very varied and defying any easy
generalisations, although Bee Orchid is
essentially a species of open grassland. Most
sites are on light, well-drained soils that are
low in nutrients, but it also grows on heavy
clays, in areas that may have standing water
in the winter, or on flushed, slumped clay
cliffs. It can occur on closely cropped swards,
in areas with much bare ground or in ranker
grassland among scrub. It is usually supposed
to favour calcium-rich soils overlying chalk
and limestone as well as chalky boulder
clay, but it is much more widespread and
any poor, free-draining soil may be suitable,
although there may well be some factor
that makes the soil locally more alkaline.
Many habitats are obviously man-made,
such as road verges, railway embankments
and cuttings, old quarries, pits, spoil-heaps,
gravel pits, brownfield industrial sites and
garden lawns. Bee Orchids are also found in
more natural areas such as sand dunes, dune
slacks, limestone pavement, eskers (ridges of
399
glacial debris) and, especially in Ireland, fens.

The species is tolerant of heavy grazing and
trampling. Although it is usually found in open
sunny places it has been recorded growing in
shady woodland with little other vegetation.
In woods, it flowers a month later than nearby
plants in the open (Summerhayes, 1968).
Bee Orchids often behave like weeds,
colonising areas of bare or disturbed ground.
They increase rapidly in numbers until a
closed sward develops and then, being poor
competitors, they disappear. On the other
hand, they can thrive in permanent grassland
and such colonies may last for many years. It
occurs from sea level up to 335m (Parsley Hay,
Derbyshire).
Flowering period
Early June to late July, sometimes from late May,
but at least in southern England most plants
will have finished flowering by the end of June.
Range
It is widespread in England north to Cumbria
and Co. Durham, although sparse or absent in
M 16 June, Norfolk. Despite an aura of rarity, Bee Orchid is an adaptable and successful species and can be found in
large numbers, often at new sites.
29/1/09 12:41:37
400
GENUS OPHRYS
29/1/09 12:41:40
BEE ORCHID
1987-99
1970-86
pre 1970
401
flowering plants, but the total population,

including vegetative plants and those dormant
underground, is much more stable.
DESCRIPTION
north Devon and Cornwall, and the number

and density of colonies declines steadily
north and west of a line from Bristol to
Hull. In Wales it is largely confined to areas
on or near the north and south coasts. It is
also found on the Channel Islands and the
Isle of Man, and occurs throughout Ireland
but is very local and absent from large areas.
Very rare in Scotland, with a 1908 record
from Kircudbrightshire, but found on an
old industrial site in east Ayrshire in 2003.
World range: Europe, the Middle East and
North Africa. Found north to Holland,
northwest Germany, the Czech Republic and
the Ukraine and south to the Mediterranean
Islands, Crimea, Caucasus, Turkey, Lebanon
and Israel. In North Africa occurs in
Morocco, Tunisia and Algeria.
How to find it
Throughout its range Bee Orchid is rather
local and as likely to be found on man-made
sites as in old, species-rich grassland or scrub.
It is worth looking for in any area that has
been heavily disturbed to produce lots of bare
ground, especially if it is on poor, chalky or
sandy soils, which are slow to be recolonised
by vegetation. In established colonies
there are huge variations in the number of
O 18 June, Norfolk.

Stem: Pale green with two scale leaves at the
base.
Leaves: There are up to six pale green basal
leaves that are clearly veined, keeled and strapshaped but become narrower, more pointed
and loosely clasping higher on the stem. There
are also one or two bract-like non-sheathing
leaves towards the flower spike. The leaves
appear in September-November and are
often scorched or otherwise damaged by the
summer.
Spike: Loose, with two to seven flowers,
sometimes as many as 12.
Bract: Pale green, lanceolate, pointed at the tip
and much longer than the ovary.
Ovary: Green, boldly ribbed but not twisted,
held upright but slightly curved; the tip bends
further over to hold the flower facing outward.
Flower: The sepals are oval, tapering slightly
towards the tip, concave and often hooded.
They are various shades of pink, from pale rose
to a deep pink tinged lilac, and have three to
five variably obvious green veins. Occasionally
the sepals are white. The lateral sepals are held
horizontally or a little below the horizontal
and are swept backwards, whereas the upper
sepal is held upright but very frequently bends
backwards to lie almost horizontally behind
the flower. The petals are much shorter and
are strap-shaped with their margins rolled
back to make them appear even narrower; they
are greenish through pinkish-brown to pink
with fine white hairs. The lip is tongue-shaped
with the sides and front strongly moulded
downwards and two relatively small conical
sides lobes at the base that are conspicuously
hairy on the outer side. The tip of the lip has
two lobes with a pointed nib in the shallow
notch between them but appears rounded
because the entire tip is curled up underneath.
At the base of the lip there is an elongated,
29/1/09 12:41:41
402
GENUS OPHRYS
semi-circular, hairless, dull orange area that is

bordered by narrow maroon-brown and pale
yellow bands. The speculum radiates from these
and is a broader band of dull purple that is in
turn bounded with a pale yellow band. The
markings form a U or H-shape below the basal
area, sometimes irregular and asymmetrical.
The side-lobes are also bounded by dull purple
and pale yellow bands. The remainder of the
lip is velvety maroon-brown. The column is
greenish and held more-or-less at 90 to the
lip. In profile it is said to resemble the head of
a duck with the anther at the tip forming an
elongated beak. The pollinia are yellow and
their caudicles (stalks) lie in parallel grooves
M 26 June, Norfolk. The column is said to resemble the

head and beak of a duck. In these flowers the pollinia
have already become stuck to the stigmas.
until they are released, with the viscidia at their

bases enclosed in pale yellowish-green bursicles.
At the base of the column the circular stigmatic
cavity is yellowish with a horizontal band of
orange-brown.
Subspecies
None.

As with the other species in the genus Ophrys,
the flowers are rather variable in structure,
colour and pattern.
Var. trollii Wasp Orchid has a long narrow
central lobe to the lip that tapers to a point. The
side-lobes are often longer and narrower than
normal, being held away from the central lobe.
The lip is marbled asymmetrically with yellow
and rusty-brown and the speculum is either
distorted or absent. It occurs regularly at a few
sites in the West Country but is otherwise rare.
The tip of the lip in normal plants sometimes
fails to fold under and they then have an
elongated, pointed lip but retain the normal
markings. For a time, Wasp Orchid was
considered to be a distinct species.
Var. belgarum has an oval lip, lacking welldefined side-lobes but with hairy shoulders. It
has symmetrical markings with a horizontal
yellow band across the middle and smaller
vertical yellow bands; there is no speculum.
First described in 1998 it has been identified
widely in southern England.
Var. bicolor has the outer half of the lip dark
brown, grading to pale, unmarked greenish or
pale brown at the base. There is no speculum. It
is very rare but has been recorded from Dorset,
Essex/Suffolk and Anglesey.
Var. chlorantha lacks anthocyanin pigments
and has whitish sepals, yellow petals and a
bright greenish-yellow lip with a ghost pattern.
It is rare but occurs widely, especially in the
south and east. Var flavescens has also been
described. This is a less extreme version of
var. chlorantha with a pale brown lip that has a
normal but faded pattern. It is apparently very
rare and easily confused with the old, faded
flowers on normal plants.
29/1/09 12:41:42
BEE ORCHID
trollii
belgarum
bicolor
403
of a few hours the pollinia drop downwards on

their stalks and dangle in front of the stigma. In
this position the slightest breeze will waft them
on to its sticky surface and pollination will take
place. This mechanism is efficient and a large
proportion of flowers are pollinated. Each pod is
estimated to contain 6,000-10,000 seeds. Rarely,
the flowers may be cross-pollinated. Pseudocopulation involving bees has been recorded and
Bee Orchid has hybridised with several other
Ophrys; hybridisation is only possible if it is
cross-pollinated at least occasionally.
Vegetative propagation may occur and
in cultivation several small daughter tubers
were noted growing at the end of long, slender
rhizomes.
chlorantha
friburgensis
atrofuscus
Var. friburgensis has oval petals similar in

shape to the sepals but not quite as large. They
are pink or very rarely greenish. It is very rare
but has occurred widely; Bee Orchids with
slightly enlarged petals occur more commonly.
Var. atrofuscus has an unmarked, dark,
chocolate-coloured lip. It was found in Sussex
in 2001.
BIOLOGY
Bee Orchid is usually self-pollinated. Soon
after the flower opens the anther releases the
pollinia, which have unusually long, thread-like
flexible stalks. At their base these stalks are
each in turn attached to a viscidium, and these
remain in their bursicles (protective pouches),
effectively anchoring the pollinia. In the space
M 26 June, Norfolk. Soon after the flower opens the pollinia are released and dangle like little balls on the end
of their flexible stalks, waiting for a breeze to blow them
onto the stigma.
29/1/09 12:41:44
404
GENUS OPHRYS

It used to be thought that Bee Orchid was
monocarpic and therefore flowered just once
before dying. This was given as the explanation
for the large fluctuations in the number of
flowering plants. The opposite is the case,
however, as it is relatively long-lived. In a tenyear study in Cambridgeshire the half-life of
successive generations in a population of Bee
Orchids averaged 6.6 years and varied from
5.8-11.2 years (the half-life is a measure of
the life expectancy of the orchid after its first
appearance above ground and marks the point
at which 50% of the population that emerged in
any given year have died). Some plants live for
at least ten years after their first appearance and
may flower for three years in a row.
Bee Orchids can spend one or two years
dormant underground but do not flower again
when they next emerge. They seem to need at
least a years growth as a non-flowering rosette
to build-up enough reserves to flower again.
At any stage in their life they can only flower if
they have reached a minimum leaf area, and if
they do not have a big enough spread of leaves
the rosette will wither early, in May. Damage
to the plants by snails and slugs tends to be
slight but rabbits and deer can cause severe
problems.
Seed probably germinates in the spring but
there is no clear picture of the period between
germination and the first appearance above
ground. A 19th century author noted that
germinating seeds, protocorms and plantlets
with small leafy shoots were found around
adult plants in March. Young Bee Orchids
often appear above ground in the late winter as
non-flowering plants with a single leaf but these
frequently wither and die off rather quickly.
The specific name apifera derives from the Latin

apis bee and fero to bear or to carry and means
bee-bearing.
Hybrids
O. x albertiana, the hybrid with Late Spider
Orchid, was recorded from Kent in 1828.
x Fly Orchid is very rare but has been recorded
in Somerset and West Sussex (it is sometimes
called O. x pietzschii but this is an invalid name).
HISTORY AND
CONSERVATION
John Gerard published The Herball, or General
Historie of Plantes in which he noted: The
Bee, the Fly and the Butterfly Satyrions grow
upon barren chalky hilladjoining to a village
in Kent named Greenhithe, upon Longfield
downs by Southfleet likewise in a field
half a mile from S. Albons.
The Bee Orchid is specially protected in
Wildlife Order (NI)
Past and present occurrence of Bee Orchid in Britain and

1500-1999
current range
% lost, 1500-1969
% lost, 1970-1986
% lost, total
Britain
Ireland
940
182
785 (27.5%*)
93 (9%*)
10.5%
37%
6%
12%
16.5%
49%
There has been a modest decline in Britain,

with just 16.5% of the total range lost as sites
have been destroyed by ploughing, spraying
or, in the case of quarries and old pits, are
either filled with rubbish or become overgrown
with scrub. The losses have to some extent
been compensated for by the colonisation of
new sites and the expansion of the range to
the north and west, especially in man-made
habitats: Bee Orchid is one of the more
opportunistic and adaptable British orchids.
There has been a much more significant decline
in Ireland, especially in the period before 1930.
The explanation for this is not clear.
29/1/09 12:41:44
LATE SPIDER ORCHID
LaTe sPider orchid
405
Vulnerable
Ophrys fuciflora
A close relative of the Bee Orchid, this is one of the rarest and most localised species in Britain
and is restricted to a handful of sites in east Kent. A large proportion of the population is
confined to cages as protection from grazing rabbits, and a grassy slope scattered with wiremesh boxes is a strange introduction to such a rare plant. The flower has evolved to trick
one particular species of bee into acting as an inadvertent pollinator, but the appropriate
bee does not occur in England. The mechanism is therefore largely defunct, few flowers
are pollinated and little seed is produced. Nevertheless, this species manages to cling on,
perhaps because the individual plants are relatively long-lived.
Identification
Unmistakably a bee orchid, this species has
pink sepals and petals and a broad, dark, velvety
lip. The size and colour of the petals and the
colour of the sepals are variable, however, as is
the shape and pattern of markings of the lip,
but it usually has furry shoulders and always
has a projecting nib at the tip.
Similar species
Bee Orchid is rather similar but has longer,
narrower petals that are strap-shaped rather
than triangular and more often greenish than
pink. It has a narrower lip that is distinctly
three-lobed and never square and shouldered
(on the other hand, the lip of Late Spider
Orchid can be three-lobed). Diagnostically,
the pointed tip of the lip in Bee Orchid
normally curls back and under out of sight
and it therefore lacks the projecting nib. The
column in Bee Orchid is a little longer with a
slightly more prominent projecting beak, and
the pollinia often hang loose over the stigma, a
feature never seen in Late Spider Orchid.
Early Spider Orchid always has green sepals
and long, narrow, strap-shaped petals that are
much less downy and also usually greenish
rather than pink. The speculum on its lip is
normally H-, X- or -shaped rather than
incorporating broken rings and circles. As in
Bee Orchid, its lip lacks a forward-pointing
nib and the flowering periods of the two spider
orchids do not normally overlap.
M 12 June, Kent. The petals are always short and

triangular and usually pinkish, resembling little horns.
29/1/09 12:41:45
406
GENUS OPHRYS
Habitat
Well-drained grassland on infertile chalky soils,
grazed to produce a reasonably short sward
and some bare ground and also ideally facing
south. Ground disturbance of some sort may
be important for the establishment of new
populations. Current sites are on old spoil
heaps and areas which were ploughed in the
past or heavily disturbed by rabbits prior to the
outbreak of myxomatosis. The individual plants
are relatively long-lived, and colonies can persist
when conditions are no longer suitable for
seedlings to become established. Notably, the
existing colonies tend to be very discrete and do
not expand into adjacent superficially similar
grassland.
Flowering period
Late May to late July, exceptionally to August,
but mostly in early to mid-June. At Wye NNR,
colonies that face west to southwest flower three
or four weeks later than colonies facing south.
1987-99
1970-86
pre 1970
Range
M 12 June, Kent. As the flowers open the spike elongates;

these flowers have very well-defined side lobes to the lip.
Confined to the North Downs in Kent between

Folkestone and Wye. There are old records from
other counties in southern England but all of
these are probably errors. World range: Europe,
from France eastwards to Romania; it extends
north to southern Belgium, central Germany,
Austria, Hungary and the Czech Republic and
29/1/09 12:41:47
LATE SPIDER ORCHID
407
south to Italy and the former Yugoslavia. It is

extinct in Holland. Closely related species are
found in adjacent areas.
How to find it
Around 50% of the British population is
found in six discrete colonies at Wye NNR,
many plants are caged for protection. As with
Bee Orchid, the number of flowering plants
varies widely between seasons but the overall
population is much more stable.
DESCRIPTION
Height: 5-30cm, sometimes to 37.5cm.
Stem: Grey-green.
Leaves: Grey-green and prominently veined.
The three to five lanceolate basal leaves form a
loose rosette with the lowest flat to the ground
and the remainder held up to 30 above the
horizontal. There are one to three narrower and
more pointed leaves that loosely clasp the stem.
The rosette appears above ground in October
and withers while the plant is still in flower,
usually disappearing by early July.
Spike: The two to nine flowers (rarely as many
as 14) are well-spaced along the stem.
Bract: Grey-green, lanceolate, about one to
one-and-a-half times the length of the ovary,
which they clasp at the base.
Ovary: Grey-green, slim and cylindrical, boldly
six-ribbed, slightly twisted, and also curved
through about 90 to hold the flowers facing
outwards.
Flower: The sepals are broadly oval with their
edges rolled back and under and a blunt tip.
They vary from very pale pink to a rich, dark
pink and have a prominent green midrib on
the outer surface and one to three green veins
on the inner face. The lateral sepals are held
horizontally or slightly drooping at the sides
of the flower, whereas the upper sepal may be
vertical or project out horizontally over the lip
but more often curves forwards in a graceful
arch. The petals are much shorter, triangular,
velvety-hairy and pink (often a distinctly deeper
pink, sometimes almost flame-coloured, at the
base). They may have swellings on each side
M 14 June, Kent. The flowers are similar to Bee Orchid

but the pointed tip of the lip is not tucked out of sight.
at the base, called ears or auricles. The lip is

almost square in shape but usually broadens
towards the tip. The edges are strongly moulded
downwards apart from the tip, where there is a
prominent forward or downward projecting nib
set into a notch. This nib is usually yellowish
and sometimes three-lobed. The lip is a rich,
dark chestnut-brown or maroon-brown, usually
paler around the edges and velvety in texture.
On either side of the base of the lip there are
swellings or shoulders that are variable in size;
in some plants these swellings are large, welldefined and rounded and thus more obviously
lateral lobes. The lip is particularly hairy on the
shoulders and along the sides below them. At
the base of the lip below the column is a moreor-less semicircular, smooth, orange-brown
29/1/09 12:41:50
408
GENUS OPHRYS
patch that is usually bordered by a narrow,

creamy necklace. Radiating from this necklace
the speculum is a complex and extremely
variable pattern of smooth maroon or lilacbrown markings bounded by creamy-yellow
lines that often forms an irregular star or other
geometric shape around a central dark circle.
The column is held at about 90 to the lip and
is yellowish-green, becoming distinctly greener
towards the tip. The stigmatic cavity is dark with
a small, round, black pseudoeye on either side.
The projecting rostellum or beak is small and
the pollinia are yellow.
Subspecies
None.

There is a great deal of variation in the shape
and colour of the lip and colour of the petals
and sepals. One named variety is worth
mentioning:
Var. flavescens lacks anthocyanin pigments
and has whitish sepals and petals and a pale
greenish lip with a very faint pattern. It is
very rare.
BIOLOGY
This species is pollinated by insects, and in
Europe bees of the genus Eucera are the main
pollinators. The appropriate bees do not occur
in England, however, and insect-pollination
has not been recorded, although it must occur;
pollen-beetles are possible alternative vectors.
Very few ripe seedpods have been found in
Kent but despite this enough viable seed is
produced to maintain the populations. Selfpollination may occur occasionally, even in the
bud (Summerhayes, 1968).
Vegetative reproduction is either very rare
or does not occur at all. This statement is
based on the observation that very few new
plants appear within 10cm, or even 30cm, of
existing orchids.
O 12 June, Kent.The column, as in Bee Orchid, resembles
a ducks head but is a little shorter.
29/1/09 12:41:53
LATE SPIDER ORCHID

It had been thought that Late Spider Orchid
was monocarpic and flowered just once before
dying. The opposite is the case, as it is rather
long-lived. In studies in Kent the half-life of a
population averaged 12.5 years and varied from
7.1 to 16.8 years (the half-life is a measure of
the life expectancy of the orchid after its first
appearance above ground and marks the point
at which 50% of the population that emerged
in any given year will have died). Periods of
underground dormancy are frequent with
about 13% of plants dormant each season,
and plants may spend one or two years
underground before re-emerging.
There is no information on the length of
the period between germination and the first
appearance above ground but it is probably
three or four years. Many plants flower in their
first year above ground.
409
with plants appearing regularly at just five

sites although there are probably still sporadic
appearances at several more. The low point
was in the mid-1980s, and there has been
a substantial recovery since then, perhaps
due in part to the weather but certainly to
better management. There is now a total of
approximately 500 plants, almost half of which
are at Wye NNR. The population there rose
from about 50 in 1987 to around 220 in 1998.
Habitat has been lost because of agricultural
changes but also due to changes in grazing
Hybrids
O. x albertiana, the hybrid with Bee Orchid,
has been recorded rarely from Kent.
O. x obscura, the hybrid with Early Spider
Orchid, has been recorded rarely in Kent. The
parent species seldom occur together and the
flowering periods do not normally overlap.

The specific name fuciflora derives from fucus
drone (i.e. a male bee but also meaning paint
or dye, especially red or purple colours) and flos
flower and thus presumably bee-flowered.
HISTORY AND
CONSERVATION
when the Rev. Gerard E. Smith noted it, on
the southern declivities of chalky downs near
Folkestone (The English Flora).
A Red Data Book species, it is classified
as Vulnerable and specially protected under
Schedule 8 of the Wildlife and Countryside
Act 1981. Late Spider Orchid has only ever
been reliably recorded from Kent, with about
20 historic localities. It is now much reduced,
M 14 June, Kent. Ophrys x albertiana, the hybrid with Bee

Orchid. Under suspicion for some years, the identity of this
plant as a hybrid was finally confirmed by DNA analysis.
29/1/09 12:41:54
410
GENUS OPHRYS
patterns and the reduction in rabbit numbers

following myxomatosis, with grassland
reverting to scrub. All the regular sites are now
within SSSIs and are managed with the species
in mind, but at its former localities grazing was
abandoned many years ago. The Late Spider
Orchid is probably best adapted to a dynamic
system. It needs ground disturbance and some
bare ground to provide suitable conditions
for seedlings to become established, as well
as grazing and sometimes hand-mowing to
maintain a suitable short sward for the adult
plants to thrive.
The preferred pollinator is not present
in Britain and few flowers produce seed.
To counter this, hand-pollination has been
undertaken at Wye. Fortunately, with a very low
level of mortality in adult plants (less than 5%
per annum), only small numbers of seedlings
need to survive and flower in order to maintain
the population.
Past and present occurrence of Late Spider Orchid in Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
4 (0.14%*)
% lost, 1500-1969
33.3%
% lost, 1970-1986
0%
% lost, total
33.3%
O 14 June, Kent. The lip shape in the small English

population is extraordinarily variable; in this plant the
sepals are largely greenish.
29/1/09 12:41:55
EARLY SPIDER ORCHID
411
earLy sPider orchid

Ophrys sphegodes
Just about the earliest orchid to flower in spring (vying with Early Purple Orchid for the
honour), this species is almost entirely restricted to the south coast between Dorset and
Kent. Although Nationally Scarce it can be locally common and has appeared in vast
numbers at Samphire Hoe, Dover, on spoil excavated from the Channel Tunnel. Indeed, the
prospect of such a rare plant dotted in hundreds or even thousands across the downland
turf is almost a mockery of the term itself. The flowers may look like a spider to human eyes
but have in fact evolved to trick solitary bees into acting as pollinators.
Identification
Unmistakably a bee orchid, the circular deep
brown lip resembles the body of a large garden
spider and is marked with a lustrous bluish
speculum. The sepals are green, and the petals
are yellowish-green, narrow and strap-shaped
with wavy edges.
Similar species
Bee and Late Spider Orchids both have similar
large brownish lips but their specula are
bordered by a narrow creamy line and their
sepals and petals are pink.
Habitat
It is found on old species-rich grassland on
chalk or limestone, growing both in short
closely-cropped turf (even lawns) and in slightly
ranker swards. It has some preference for
previously disturbed areas, such as old quarries,
spoil heaps and tracks; as with Late Spider
Orchid heavy ground disturbance may aid the
successful establishment of new colonies. It has
been recorded rarely from shingle and almost all
the current sites are near the sea.
Flowering period
Late March to early June. Tends to flower
earliest in Dorset, from early or mid-April to
early or mid-May and may average a few days
later in Sussex and Kent. Flowering times are
very variable, however, both from year to year
and between colonies, even in the same area.
P 14 May, Kent. Many Early Spider Orchids are very
small.
29/1/09 12:41:57
412
GENUS OPHRYS
For example, plants at Samphire Hoe in Kent

bloom perhaps two weeks earlier than those at
Lydden, just 6km away.
1987-99
1970-86
pre 1970
Range
Largely confined to the coasts of Dorset, East
Sussex and Kent; in Dorset there are good
populations between St Aldhelms Head and
Durlston; in East Sussex now found mostly
between Beachy Head and Seaford and at
Castle Hill near Brighton; in Kent there are
around 30 colonies, most along the coast
but including one or two inland sites such
as Queendown Warren. Away from these
strongholds there have been isolated recent
records in west Gloucestershire (from 1975),

south Wiltshire (1988), Northamptonshire
(2001, the first record for 230 years) and west
Suffolk (where a single plant appeared from
1991 at Lakenheath, the first record since
1793). Introduced and naturalised near
Bishops Stortford in Hertfordshire. World
Range: Mainland Europe from Spain eastwards
to Greece and north to Belgium, Luxemburg,
central Germany, Austria and the Czech
Republic.
How to find it
One of the best and most accessible sites is
Samphire Hoe below Shakespeare Cliff at
Dover. This was created using five million cubic
metres of marl dug from beneath the seabed
during the construction of the Channel Tunnel.
In 1998 there were 61 plants but by 2004 this
has increased to 9,000. In East Sussex up to
50,000 plants have been recorded at Castle Hill,
and it is also found at Cuckmere Haven and the
Seven Sisters, and in Dorset the most accessible
site is Durlston Head.
DESCRIPTION
M 14 May, Kent. The greenish sepals and petals are distinctive.
Height: Mostly five to 15cm, sometimes to

20cm or rarely 35cm, but at Samphire Hoe
plants may be as tall as 45cm.
Stem: Yellowish-green, thick and fleshy.
29/1/09 12:41:58
EARLY SPIDER ORCHID
Leaves: Grey-green to green and prominently

veined. The three or four lower leaves are short
and broadly strap-shaped with a blunt tip; the
lowermost is held nearly horizontally but the
remainder are more upright. The upper leaves
are narrower, more pointed and loosely sheathe
the stem. Some plants are wintergreen but in
most the leaves appear in the early spring. All
have withered by the end of June.
Spike: Most have two to seven flowers,
exceptionally as many as ten, but at Samphire
Hoe some tall plants may have as many as 17
blooms widely spaced along the stem.
Bract: Pale green, lanceolate, blunt-tipped and
slightly longer than the ovary, which they moreor-less sheathe at their base.
Ovary: Green and boldly six-ribbed.
Flower: The sepals are oval-oblong with blunt
tips and their margins rolled under. They are
usually pale yellowish-green in coloration but
rarely are more whitish. The petals are more
strap-shaped, shorter and narrower, with a
squared-off tip and edges that are frequently
waved or undulate. They are often a slightly
deeper yellowish-green and their margins
may be more brownish with some fine hairs.
Very rarely the petals are pinkish. The lateral
sepals are held horizontally and the upper sepal
vertically although the latter usually arches
forward over the column. The lip is almost
circular in shape with the edges moulded
downwards to give it a convex profile. It is
deep purplish-brown, usually paler and more
yellowish around the lower edge and velvety in
texture with a smooth, slightly lustrous, slategrey, lead-coloured or bluish mark in the centre
or towards the base (the speculum), often in
the form of an H, X or the Greek letter . On
either side of the base of the lip there are two
roughly conical swellings or haunches that
are variable in size, and the lip is particularly
hairy around these and along the edges below
them. The column is greenish-white and held
at about 90 to the lip with the rostellum at
its tip forming a short blunt beak. At the base
of the column the stigmatic cavity is circular,
maroon-brown on the lower half and pale green
413
M 14 May, Kent.The speculum is usually a lustrous blue

or slate-grey, and the flower also has a pair of pseudoeyes on either side of the stigma.
on the upper with two pseudo-eyes, one on

each side; it has been reported to contain sugar
(Summerhayes, 1968). The pollinia are yellow.
The flowers fade rapidly, the lip becoming a dull
pale yellowish-brown or grey-brown.
Subspecies
None.

The shape and coloration of the lip is very
variable and rarely the speculum may be red
rather than blue.
Var. flavescens lacks anthocyanin pigments and
has a greenish or golden lip, sometimes brownish
towards the edges, with merely a shadow
speculum. It is rare but can easily be confused
with a normal flower that has faded with age.
BIOLOGY
It is cross-pollinated by bees. The specific
pollinator is the male solitary bee Andrena
nigroaenea, and the pollinia are attached to
the front of its head as it attempts pseudo-
29/1/09 12:41:59
414
GENUS OPHRYS
copulation. The mechanism can be successful

with about 25% of flowers recorded as setting
seed at Samphire Hoe, but pollination rates
can be very much lower with a lack of suitable
pollinators being the most likely explanation for
the low seed-set. It has also been suggested that
British populations are probably mostly selfpollinated. Indeed, the pollinia are occasionally
released to dangle in front of the stigma as in
Bee Orchid, making self-pollination possible.
The overall low rates of pollination suggest,
however, that self-pollination is not routine.
Vegetative reproduction is thought to
be uncommon or rare and as Early Spider
Orchid is a short-lived orchid, dependent on
seed to maintain its numbers, sufficient seed
M 8 May, Dorset. The lip fades rapidly to a dull brown

and thus as successive flowers open only one or two at a
time will be in good condition.
is presumably being produced to sustain the

current colonies despite the apparently low
rates of pollination.

The aerial stem grows from a pair of tubers, the
younger of which is often stalked.
Early Spider Orchid is relatively short-lived.
Few plants survive for more than three years
after their first emergence above ground and
the majority appear just once, flower and then
die (i.e. they are monocarpic), although a tiny
minority can live for at least ten years. The
number of plants above ground varies widely
from year to year, largely correlated with the
amount of rainfall over the previous winter, and
plants may spend one or two years dormant
underground.
Early Spider Orchid has a complex life cycle.
This complicates attempts to understand its
population dynamics and life history on the
basis of an annual census. In a study at Castle
Hill, Sussex, over a winter season, plants began
to appear above ground in early September
and continued to emerge through the winter,
with peaks in November-December and more
especially in March-May. By flowering time
therefore some plants had been above ground
for just two months and some for six. Those
that appeared early in the winter included
most of the older plants that had flowered the
year before and all these early plants suffered
grazing damage. Previously unrecorded plants,
presumably seedlings emerging above ground
for the first time, appeared throughout the
season but especially from March onwards.
However, around 75% of all the plants recorded
up till March did not survive above ground
to be counted in the annual census in May,
with grazing being the likely cause of their
disappearance.
There is no information on the period
between germination and the first appearance
above ground, but it is probably in the region
of one to three years. Early Spider Orchids first
appeared on spoil from the Channel Tunnel
four years after it had been spread.
29/1/09 12:42:00
EARLY SPIDER ORCHID
Hybrids
O. x hybrida, the hybrid with Fly Orchid, has
occurred occasionally in Kent.
O. x obscura, the hybrid with Late Spider
Orchid, has been recorded rarely in Kent. The
parent species seldom occur together and the
flowering periods do not normally overlap.

The specific names sphegodes derives from the
Greek sphex wasp or sphekeion small, wasp-like
spider and means wasp-like.
HISTORY AND
CONSERVATION
when William How published his Phytologia
Britannica natales exhibens Indigenarum
Stirpium sponte emergentium (A British botany
presenting the origins of native wild plants) in
which he noted it in Northamptonshire, upon
an old Stone pit groundhard by Walcot a
mile from BarnackDr. Bowle.
A Red Data Book species and although
no longer classified as Near-threatened it
remains specially protected under Schedule
8 of the Wildlife and Countryside Act 1981.
The Early Spider Orchid has vanished from
at least 73% of its historical range but the
majority of the losses occurred long ago, in
the 19th century. This was probably largely
due to the cultivation of grasslands following
the Enclosures, although the retreat towards
the coast of southeast England suggests that
climatic factors may also have been involved.
It is now extinct in the Channel Islands, west
Cornwall (where recorded at Upton Towans in
1961), the Isle of Wight (a few records, the last
in 1992), Hampshire (very few records, the last
near Lepe in 1978), West Sussex, Surrey (last
recorded from Titsey in 1942 but this record
probably originated with plants transplanted
from Queendown Warren in Kent), Essex,
Oxfordshire (1920, with an unconfirmed record
in 1975), Bedfordshire, Cambridgeshire and
Denbighshire (a 19th century record).
Many sites for the Early Spider Orchid are
now protected but its grassland habitat also
415
needs careful management to maintain the

populations. Grazing by sheep is beneficial,
as long as they are removed during the period
when the orchids flower and set seed. Any
future spread may depend upon the creation
of suitable bare ground and the sort of drastic
disturbance that created such ideal conditions
at Samphire Hoe.
Past and present occurrence of Early Spider Orchid in Britain and Ireland (based on presence or absence in 10km

1500-1999
current range
Britain
Ireland
63
17 (0.6%*)
% lost, 1500-1969
71.5%
% lost, 1970-1986
1.5%
% lost, total
73%
*current range as a % of the total number of 10km squares
M 14 May, Kent. Early Spider Orchids can have just one

flower; this plant has especially brownish petals.
29/1/09 12:42:01
416
OTHER SPECIES
oTher sPecies
Several other species of orchid have been recorded from time to time in Britain and Ireland.
This is a selection:
Short-spurred Fragrant Orchid
Gymnadenia odoratissima
Recorded from Black Hall Rocks in Co.

Durham in 1912, but doubt has been cast on
the identity of the specimen.
Frivalds Fragrant Orchid

Gymnadenia frivaldii
Found and photographed on wet heathland in

Dorset in 1972 or 1973, its normal range is the
Balkans and its occurrence here is a mystery
(Ettlinger 1997).
False Musk Orchid (Dwarf Alpine

Orchid)
Chamorchis alpina
A single plant was reported from the New

Forest in Hampshire in 1976. Among the
smallest of European orchids, this is a species
of damp, calcareous soils in the mountains
of Scandinavia and the Alps. It was probably
planted.
Pale-flowered Orchid
Orchis pallens
Heart-flowered Tongue Orchid

Serapias cordigera
Found in 1996 at Monkton on the Isle of

Thanet in Kent, with two or three plants in
1997. They were on a south-facing bank at
the bottom of a cliff in an abandoned chalk
quarry. This is a common and widespread
species around the Mediterranean and occurs
north to Finistere. Despite apparently excellent
credentials, these plants almost certainly
originated from wind-blown seed coming from
plants growing in pots on a patio just 300m
upwind of the nature reserve.
Bertolonis Mirror Orchid

Ophrys bertolonii
A single plant found in Dorset in 1976

generated much controversy as to its origins,
but it eventually emerged that a holidaymaker,
returning from Italy, had planted it. Ophrys
bertolonii has since been divided into several
species, and the precise identity of the Dorset
plant cannot be confirmed.
A few specimens were recorded in the 1920s

in Hampshire but there are no further details
(Ettlinger 1997).
Scarce Tongue Orchid

Serapias neglecta
Found in 1918 in a cornfield on the Isle of

Wight, the roots were dug up and taken into
cultivation. Rather localised in coastal regions
around the central Mediterranean, this species
may have been introduced with imported
cereal seed.
29/1/09 12:42:01
ORCHID FAMILY TREE
417
Orchid Family Tree

Cypripedium (ladys-slippers)
Subfamily
Cypripedioideae
tribe Neottieae
Cephalanthera (helleborines i)
Neottia (Birds-nest Orchid etc.)
FAMILY
Subfamily
Epidendroideae
Epipactis (helleborines ii)

tribe Gastrodieae
ORCHIDACEAE
tribe Calypsoeae
Epipogium (Ghost Orchid)

Corallorhiza (Coralroot Orchid)
tribe Malaxideae
Liparis (Fen Orchid)

Hammarbya (Bog Orchid)
tribe
Goodyera (Creeping Ladys-tresses)

Spiranthes (ladys-tresses)
Cranichideae
subtribe
Herminium (Musk Orchid)
Habenariinae
Subfamily
Orchidoideae
Dactylorhiza (marsh orchids)

tribe
Gymnadenia (fragrant orchids)
Orchideae
Platanthera (butterfly orchids)

Pseudorchis (Small White Orchid)
Orchis (typical orchids)
subtribe
Orchidinae
Neotinea (Dense-flowered Orchid etc.)

Himantoglossum (Lizard Orchid)
Anacamptis (Pyramidal Orchid etc.)
Serapias (tongue orchids)
Ophrys (bee orchids)
All orchids are members of the family Orchidaceae. Its most fundamental subdivisions are the
subfamilies, three of which are represented in Britain and Ireland: the Cypripedioideae, with just one
species Ladys-slipper; the Epidendroideae, a heterogeneous group, many of which are woodland
orchids that grow from rhizomes; and the Orchidoideae, a more homogenous group, all of which
grow from a swollen underground tuber. The subfamilies are further divided into tribes, genera and,
finally, individual species.
This family tree follows Bateman (2006) and has been constructed using the latest evidence from
genetic research. It gives a much better idea of relationships than is possible in a simple list. For
example, Bog Orchid and Creeping Ladys-tresses lie next to each other in the field guide, but it can
be seen that they are members of separate subfamilies and not at all closely related.
29/1/09 12:49:20
418
VICE-COUNTIES
vice-cOunTies
112
111
109
108
107
110
105 106
104
94
95
96
92
97
89
103
87
101
40
36
27
26
16
18
2
4
32
67
22
66
69
65
71
31
60
59
49 50
58
52
19 21
20
13
11 12
48
47
46
45
44
S
2
1
34
54
57 56
53
28 27
55
32
31
33 23 24
30
20
38
21
22
12
8
9
63
37
36
40
35
61
39
43
42
62
64
51
41
68
70
37 38
14
10
8
30
24
23
17
15
29
25
33
28
81
79 80
72
74 73
39
84
83 82
77 78
75
100
35
34
85
86
102
91
90
88
98
93
17
13
11
29
26 25
19
18
16
15
14
10
29/1/09 12:49:21
VICE-COUNTIES
England
1
West Cornwall
2
East Cornwall
3
South Devon
4
North Devon
5
South Somerset
6
North Somerset
7
North Wiltshire
8
South Wiltshire
9
Dorset
10 Isle of Wight
11 South Hampshire
12 North Hampshire
13 West Sussex
14 East Sussex
15 East Kent
16 West Kent
17 Surrey
18 South Essex
19 North Essex
20 Hertfordshire
21 Middlesex
22 Berkshire
23 Oxfordshire
24 Buckinghamshire
25 East Suffolk
26 West Suffolk
27 East Norfolk
28 West Norfolk
29 Cambridgeshire
30 Bedfordshire
31 Huntingdonshire
32 Northamptonshire
33 East Gloucestershire
34 West Gloucestershire
36 Herefordshire
37 Worcestershire
38 Warwickshire
39 Staffordshire
40 Shropshire
53 South Lincolnshire
54 North Lincolnshire
55 Leicestershire
56 Nottinghamshire
57 Derbyshire
58 Cheshire
59 South Lancashire
60 West Lancashire
61 Southeast Yorkshire
62 Northeast Yorkshire
63 Southwest Yorkshire
64 Mid-west Yorkshire
65 Northwest Yorkshire
66 Co. Durham
67 South Northumberland
68 Cheviot
(north Northumberland)
69 Westmorland
70 Cumberland
Isle of Man
71 Isle of Man
Wales
35 Monmouthshire
41 Glamorganshire
42 Breconshire
43 Radnorshire
44 Carmarthenshire
45 Pembrokeshire
46 Cardiganshire
47 Montgomeryshire
48 Merioneth
49 Caernarvonshire
50 Denbighshire
51 Flintshire
53 Anglesey
Scotland
72 Dumfries-shire
73 Kirkcudbrightshire
74 Wigtownshire
75 Ayrshire
76 Renfrewshire
77 Lanarkshire
78 Peebles-shire
79 Selkirkshire
80 Roxburghshire
81 Berwickshire
82 East Lothian
83 Midlothian
84 West Lothian
85 Fife
86 Stirlingshire
87 West Perth
88 Mid Perth
89 East Perth
90 Angus
91 Kincardineshire
92 South Aberdeen
93 North Aberdeen
94 Banffshire
95 Moray
96 Easterness (Inverness-shire)
97 Westerness
(Inverness-shire)
98 Main Argyll
99 Dunbarton
100 Clyde Islands (Buteshire)
101 Kintyre
102 South Ebudes
(Inner Hebrides)
103 Mid Ebudes
(Inner Hebrides)
104 North Ebudes
(Inner Hebrides)
105 West Ross
(Ross & Cromarty)
106 East Ross
(Ross & Cromarty)
419
107 East Sutherland

108 West Sutherland
109 Caithness
110 Outer Hebrides
111 Orkney
112 Shetland
Channel Islands
S
Channel Islands
Ireland
H1 South Kerry
H2 North Kerry
H3 West Cork
H4 Mid Cork
H5 East Cork
H6 Co. Waterford
H7 South Tipperary
H8 Co. Limerick
H9 Co. Clare
H10 North Tipperary
H11 Co. Kilkenny
H12 Co. Wexford
H13 Co. Carlow
H14 Laois
H15 Southeast Galway
H16 West Galway
H17 Northeast Galway
H18 Offaly
H19 Co. Kildare
H20 Co. Wicklow
H21 Co. Dublin
H22 Meath
H23 Westmeath
H24 Co. Longford
H25 Co. Roscommon
H26 East Mayo
H27 West Mayo
H28 Co. Sligo
H29 Co. Leitrim
H30 Co. Cavan
H31 Co. Louth
H32 Co. Monaghan
H33 Fermanagh
H34 East Donegal
H35 West Donegal
H36 Tyrone
H37 Co. Armagh
H38 Co. Down
H39 Co. Antrim
H40 Co. Londonderry
(Some alternative names are added in parentheses to allow easier
reference to modern or historical
counties.)
29/1/09 12:49:21
Orchid FlOwering periOds

Apr
May
Jun
Jul
Aug
Sept
Apr
May
Jun
Jul
Aug
Sept
Ladys-slipper
Red Helleborine
Sword-leaved Helleborine
White Helleborine
Lesser Twayblade
Common Twayblade
Birds-nest Orchid
Marsh Helleborine
Dark-red Helleborine
Broad-leaved Helleborine
Violet Helleborine
Narrow-lipped Helleborine
Dune Helleborine
Lindisfarne Helleborine
Green-flowered Helleborine
Ghost Orchid
Fen Orchid
Bog Orchid
Coralroot Orchid
Creeping Ladys-tresses
Irish Ladys-tresses
Autumn Ladys-tresses
Musk Orchid
Man Orchid
Monkey Orchid
Military Orchid
Lady Orchid
Early Purple Orchid
Small White Orchid
Common Fragrant Orchid
Heath Fragrant Orchid
Marsh Fragrant Orchid
Early Marsh Orchid
Frog Orchid
Common Spotted Orchid
Heath Spotted Orchid
Southern Marsh Orchid
Pugsleys Marsh Orchid
Hebridean Marsh Orchid
Northern Marsh Orchid
Irish Marsh Orchid
Dense-flowered Orchid
Burnt Orchid
Lizard Orchid
Loose-flowered Orchid
Pyramidal Orchid
Green-winged Orchid
Fly Orchid
Bee Orchid
Late Spider Orchid
Early Spider Orchid
29/1/09 12:49:21
SITE GUIDE
421
Site Guide
INTRODUCTION
The following is a selection of sites in Britain and Ireland where it is possible to find wild orchids.
Of course, orchids can be found in many places, but sadly more and more of the countryside is bereft
of their beauty and it is increasingly necessary to search out special places in order to experience the
magic of wild orchids.
We see this selection as a starting point and would encourage everyone to explore and discover
new places, to record the orchids that they find and to pass this information on to the local botanical
recorder. But, in the meantime, you may be in an unfamiliar area, on holiday, or just keen to find
somewhere that offers a good chance of success.
For each site we have included brief details of access (including a grid reference), habitat and
the species that may be found. Many orchid sites are accessed down a maze of lanes, and it is very
difficult to give concise and understandable written directions. To locate many of these sites you
will need a large-scale road atlas, especially one that marks the National Grid and therefore allows
you to pinpoint grid references or, even better, an OS map at 1:50,000 or 1:25,000 scale. If the site
is a reserve (and most are) further information can often be found on the internet by visiting the
managing organisations website. It is also worth trawling the net for information on nearby reserves
or Roadside Nature Reserves which are often also good for orchids.
This selection includes sites for some of the rarer orchids but these are all accessible and welcome
visitors. On many reserves, guided walks are organised from time to time by the managing body and
these can be a good introduction to the site. We have not tried to include localities for every species;
one or two are so rare and vulnerable that visitors are not welcome, and some species, for example
Green-flowered Helleborine, occur in small, scattered, hard-to-find colonies in places that have no
other orchid interest. Rather, we have tried to select areas that hold a good selection of species, or have
large and spectacular displays of one or two orchids. We have also endeavoured to choose sites that
are interesting and attractive, and almost all will have a great deal of other wildlife interest, be it other
plants, butterflies, dragonflies or whatever. All have public access throughout the orchid season, with
the exception of one or two where entry is restricted to a limited number of open days. Unfortunately,
we have not been able to give details of disabled access, but it is generally rather limited.
Orchid hunting requires very little equipment. As well as a good map, a hand-lens is useful, as is
a small plastic ruler and, of course, a field guide to all the other wonderful plants that will be seen.
Finally, it should be remembered that many orchids are scarce, and some are so rare that they are
protected by law. All orchids are valuable, as indeed are the other plants around them. Please look
where you are going, stay on paths where possible, and leave the site as you would wish to find it.
Inevitably there are errors in the accounts, for which we take full responsibility. We would be
delighted to hear of any corrections or updates, or indeed any additional sites that could be included
in future editions of the book (the authors can be contacted via their website: www.norfolknature.
co.uk or via the publishers).
29/1/09 12:49:21
422
SITE GUIDE
useFul addresses
An Taisce (National Trust for Ireland), Tailors Hall, Back Lane, Dublin 8, Ireland
Tel: +353 (0)454 1786. Email: info@antaisce.org Website: www.antaisce.org
CEH Monkswood, Abbots Ripton, Huntingdon, Cambridgeshire PE28 2LS
Tel: 01487 772400. Email: monkswood@ceh.ac.uk Website: www.ceh.ac.uk
The Countryside Agency, Head Office: John Dower House, Crescent Place, Cheltenham,
Gloucestershire GL50 3RA
Tel: 01242 521381. Email: info@countryside.gov.uk Website: www.countryside.gov.uk
Countryside Council for Wales, Maes-y-Ffynnon, Penrhosgarnedd, Bangor, Gwynedd LL57 2LQ
Tel: 01248 385500. Email: enquiries@ccw.gov.uk Website: www.ccw.gov.uk
Environment and Heritage Service, Commonwealth House, 35 Castle Street, Belfast BT1 1GU
Tel: 028 90 251477. Website: www.ehsni.gov.uk
Irish Wildlife Trust, 107 Lower Baggot Street, Dublin 2, Ireland
Tel: +353 (0)1 676 8588. Email: enquires@iwt.ie Website: www.iwt.ie
Joint Nature Conservation Committee, Monkstone House, City Road, Peterborough PE1 1JY
Tel: 01733 562626. Email: comment@jncc.gov.uk Website: www.jncc.gov.uk
National Parks and Wildlife Service, Department of the Environment, Heritage & Local
Government, 7 Ely Place, Dublin 2, Ireland
Tel: +353 (0)1 888 2000. LoCall: 1890 202021 (within Republic of Ireland only).
Email: natureconservation@environ.ie Website: www.environ.ie or www.npws.ie
National Trust, 36 Queen Annes Gate, London SW1H 9AS
Tel: 020 72229251. Website: www.nationaltrust.org.uk
National Trust (Northern Ireland), Rowallane, Saintfield, Ballynahinch, Co. Down BT24 7LH
Tel: 028 97 510721. Website: www.ntni.org.uk
National Trust for Scotland, Wemyss House, 28 Charlotte Square, Edinburgh EH2 4ET
Tel: 0131 2439300. Website: www.nts.org.uk
Natural England, 1 East Parade, Sheffield, S1 2ET.
Tel: 0845 600 3078. Email: enquiries@naturalengland.org.uk Website: www.naturalengland.
org.uk
Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire SG19 2DL
Tel: 01767 680551. Website: www.rspb.org.uk
Scottish Natural Heritage, 12 Hope Terrace, Edinburgh EH9 2AS
Tel: 0131 447 4784. Email: enquiries@snh.org.uk Website: www.snh.org.uk
Scottish Wildlife Trust, Cramond House, Off Cramond Glebe Road, Edinburgh EH4 6NS
Tel: 0131 312 7765. Email: enquires@swt.org.uk Website: www.swt.org.uk
The Wildlife Trusts, The Kiln, Waterside, Mather Road, Newark NG24 1WT
Tel: 0870 036 7711. Email: info@wildlife-trusts.cix.co.uk Website: www.wildlifetrusts.org
29/1/09 12:49:22
BOTANICAL ORGANISATIONS
423
BOTanical OrganisaTiOns
Botanical society of the British isles
Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road,
London SW7 5BD. Website: www.bsbi.org.uk
The BSBI is a learned society of professional and amateur botanists dedicated to the study of and
interest in the British and Irish flora. It publishes a journal, Watsonia, as well as the less formal (and
eminently readable) BSBI News. In addition, the BSBI organises the system of county plant recorders,
runs field meetings and has a panel of referees to advise on plant identifications (including, of course,
orchids). BSBI members provided the vast majority of the data used to produce the New Atlas. In
short, the BSBI is the society for anyone with a keen interest in wild plants.
Hardy Orchid Society

Website: www.hardyorchidsociety.org.uk
Dedicated to the study of European orchids, including field study, photography and conservation,
with an emphasis on cultivation and propagation. Publishes a quarterly Journal with articles on all
these subjects and organises field meetings and exhibitions of cultivated orchids.
Plantlife International
14 Rollestone Street, Salisbury, Wiltshire SP1 1DX
Tel: 01722 342730. Email: enquires@plantlife.org.uk Website: www.plantlife.org.uk
The wild plant conservation charity. Plantlife International (formerly Plantlife) has an increasingly
international focus but still does much useful work in Britain. It has a small number of reserves,
runs back from the brink projects for many declining species (including Sword-leaved Helleborine),
organised the county flowers poll and promotes various surveys to raise awareness of wild flowers, as
well as gathering much useful information. Publishes a quarterly magazine, Plantlife, and a newsletter.
Wild Flower Society

c/o Mike Hooper, 24 Muirfield Drive, Astley, Manchester, M29 7QJ.
E-mail: thewildflowersociety@yahoo.co.uk Website: www.thewildflowersociety.com
Established in 1886 for amateur botanists and wild flower lovers in the UK. Organises meetings
throughout the summer months to see and photograph British wild plants in their natural habitats.
aBBreviaTiOns
BBOWT: Berkshire, Buckinghamshire and
Oxfordshire Wildlife Trust
BC: Butterfly Conservation
BCNPWT: Bedfordshire, Cambridgeshire,
Northamptonshire and Peterborough
Wildlife Trust
BSBI: Botanical Society of the British Isles
CCW: Countryside Council for Wales
CEH: Centre for Ecology and Hydrology
FC: Forestry Commission
JNCC: Joint Nature Conservation Committee
LNR: Local Nature Reserve
LRWT: Leicestershire and Rutland Wildlife

Trust
NE: Natural England
NNR: National Nature Reserve
NPWS: National Parks and Wildlife Service
NT: National Trust
RSPB: Royal Society for the Protection of
Birds
SNH: Scottish Natural Heritage
SSSI: Site of Special Scientific Interest
WT: Wildlife Trust (with prefix, e.g. Sussex WT)
WT: Woodland Trust (without prefix)
29/1/09 12:49:22
424
SITE GUIDE
Southeast England
56
54 55 57
58
53
52
44
43
41
51
42 47
60
50
46
45
59
61
62
49
48
63
38
40
39
36
33
28
24
26
31
65
10
3
9
8 7 5
4
6
22
20
19 18
16
17 15
12
11
13
14
32
Kent
1 Trosley Country Park
2 Queendown Warren
3 Sandwich and Pegwell
Bays
4 Langdon Cliffs
5 Lydden/Temple Ewell
Down
6 Samphire Hoe
7 Park Gate Down
8 Bonsai Bank
9 Wye
10 Marden Meadows
East Sussex
11 Rye Harbour
12 Fore Wood
13 Wild Woods at Friston
Forest
14 Cuckmere Haven and
the Seven Sisters
Country Park
15 Mount Caburn
16 Malling Down
17 Castle Hill
64
21
23
30
66
34
27 25
29
37
35
18 Ditchling Beacon
West Sussex
19 Devils Dyke
20 Cissbury Ring
21 Furnace Meadow and
Brick Kiln Rough
22 Levin Down
23 Kingley Vale
Hampshire
24 Noar Hill
25 Butser Hill
26 Chapetts Copse/Hen
Wood
27 Old Winchester Hill
28 St Catherines Hill
29 Martin Down
30 New Forest
The Isle of Wight
31 Tennyson Down
32 Compton Down
Surrey
33 Thursley Common
34 Wallis Wood
35 Box Hill
36 Ranmore Common,
Denbies Hillside and
White Downs
37 Ashtead Common
51 Bernwood Meadows
52 The Ashridge Estate and
Ivinghoe Beacon
Berkshire
38 Hurley Chalk Pit
39 Greenham and
Crookham Commons
40 Basildon Park
41 Lardon Chase, the Holies
and Lough Down
Bedfordshire
53 Dunstable and
Whipsnade Downs
54 Totternhoe Knolls
55 Sharpenhoe Clappers,
Smithcombe and
Sundon Hills
56 Barton Hills
Oxfordshire
42 Hartslock
43 Dry Sandfort Pit
44 Parsonage Moor
45 Aston Rowant
46 Watlington Hill
47 Warburg Reserve
Hertfordshire
57 Hoo Bit
58 Hexton Chalk Pit
59 Therfield Heath and Fox
Covert
60 The Meads
61 Cheshunt Lock
Buckinghamshire
48 Homefield Wood.
49 Grangelands and Pulpit
Hill
50 Dancersend and the
Crong Meadow
Greater London
62 Rammey Marsh
63 The Ripple
64 West Kent Golf Course
65 Downe Bank
66 Hutchinsons Bank
29/1/09 12:49:23
SOUTHEAST ENGLAND
KEnT
1. Trosley Country Park
TQ 642614
An area of woodland and chalk downland bordering the North Downs Way. Part of a much larger
conservation area, there is an interesting chalkland
flora that includes Bee, Man, Common Fragrant and
Pyramidal Orchids.
The car park is off the A227 between Meopham and
Wrotham, south of Gravesend. (Kent County Council)
2. Queendown Warren
TQ 827629
Excellent species-rich chalk grassland on the site of
an old rabbit warren. Specialities include Early Spider Orchid and Man Orchid (although the former
is declining here); other species include Common
Fragrant, Common Spotted, Bee, Pyramidal, Early
Purple, Fly, Lady Orchids and Autumn Ladys-tresses;
also White and Broad-leaved Helleborines in the
fringing woodland.
Near Hartlip off the A2, southeast of Gillingham. Park
in Warren Lane, off Mount Lane, accessed from the
Lower Hartlip Road. (Kent WT, Plantlife, Swale Borough Council)
3. Sandwich and Pegwell Bays
TR 341632
A large reserve of mixed coastal habitats including
shingle, rich dune grassland, and coastal scrub. Good
for Southern Marsh Orchid, Pyramidal and Bee Orchids, Marsh Helleborine and the biggest colony of
Lizard Orchids in Britain (more than 4,000 were
counted in 2001).
Access from Sandwich Town via the Sandwich Bay Estate, where a substantial toll operates on the private
road.There are many footpaths along the beach, dunes
and golf courses. Park on sea-front near Princes Old
Clubhouse at Sandwich Bay. (Kent WT)
4. Langdon Cliffs
TR 335422
Lying to the east of Dover, this is part of the famous
white cliffs. With fantastic views, the chalk downland
on top of the cliffs holds many interesting plants
with Bee, Common Fragrant, Pyramidal and Common Spotted Orchids, as well as some Early Spider
Orchids and Autumn Ladys-tresses.
There are footpaths to the cliffs from Dover town centre and the port. Signposted from the M20 and Dover
town centre. Park past the Castle beside the Visitor
Centre. (NT)
5. Lydden/Temple Ewell Down
TR 277453
One of Kents best examples of chalk downland,
which is of international importance. There is also
some woodland and scrub. The speciality of the
steep slopes is Early Spider Orchid, and there are
also Common Fragrant, Bee and Pyramidal Orchids,
Common Twayblade, Autumn Ladys-tresses, and a
very few Burnt Orchids.
425
Close to Dover on the minor road between Lydden

and Temple Ewell; access is from the south side of the
railway bridge. Park behind the George and Dragon
Centre in Temple Ewell, from where the reserve is signposted. (NNR, Kent WT)
6. Samphire Hoe
TR 298395
Lying just west of Dover, this area of grassland
bounded by a sea wall was created with waste
material left over from the digging of the Channel
Tunnel. The spoil was deposited at Lower Shakespeare Cliff then landscaped and sown with wildflower seeds collected locally. A wide variety of
plants now grows here, including one of the largest
colonies of Early Spider Orchid in Britain (10,000
spikes in 2008).
Access by car is via a tunnel through the cliffs. You
can reach the tunnel from the A20; driving towards
Folkestone from Dover, follow the signs for Samphire
Hoe. (Samphire Hoe is owned and managed by
several bodies in partnership with the White Cliffs
Countryside Project.)
7. Park Gate Down
TR 168457
A chalk grassland in a dry valley fringed by woodland, this reserve has an exceptionally rich orchid
flora. It is best known for its colony of Monkey
Orchids (introduced here from seeds collected
elsewhere in Kent). These have increased in recent
years and put on a spectacular show. Other species include masses of Common Fragrant Orchids
as well as Early Purple, Bee and Pyramidal Orchids,
Common Spotted Orchid, Common Twayblade,
Greater Butterfly Orchid, a few hard-to-find Musk
Orchids and, in some seasons, a few Late Spider
Orchids. Fly, Lady and Man Orchids also occur in
small numbers.
From Sloane Street (B2068) go through Stelling Minnis and turn left (i.e. straight on) at the right-angle
bend, then take the next three right turns and carry on
through the wood; the reserve lies to the left after c.
1.5km. (Kent WT)
8. Bonsai Bank
TR 105510
Part of the Denge Wood complex, with Beech and
Sweet Chestnut mixed with conifer plantations, this
wood is named after its rather stunted crop of conifers and is an excellent site for Lady Orchid, as well
as Early Purple Orchids.
From the cross roads in Sole Street, follow Penny
Pot Lane north towards Thruxted. On entering the
woods the car park is signed to the right and the reserve lies 1/2 mile along the forest track. (Forestry
Commission)
9. Wye
TR 074469
The habitats on this steep chalk downland include
grassland, scrub and woodland, and an exceptional
29/1/09 12:49:23
426
SITE GUIDE
19 species of orchid occur here. The speciality is

Late Spider Orchid with around six distinct colonies,
but they are hard to find and visitors should contact the warden for advice. Early Spider Orchid and
Burnt Orchid also occur but are similarly elusive.
Not so Common Fragrant Orchid, Common Spotted Orchid, Common Twayblade and Pyramidal, Bee
and Man Orchids (the last favouring the western
flank of the escarpment around Dentons Field). The
woods may yield Birds-nest, Lady, Fly and Greater
Butterfly Orchids.
Northeast of Ashford on the minor road between Wye
and Hastingleigh; there are several car parks and
many footpaths. (NNR, NE)
10. Marden Meadows
TQ 763445
These three unimproved hay meadows have several
pools and are surrounded by ancient hedgerows.
Traditionally managed, there is an amazing display of
Green-winged Orchids in May, when up to 500,000
have been counted.
Just west of Staplehurst off the A229. The meadows
are sandwiched between the minor road to Marden
and the Tonbridge-Ashford railway line. Park on the
roadside. (Kent WT)
EaST SuSSEx
11. Rye Harbour
TQ 925185
A large coastal site with managed grassland, ditches,
marsh, scrub and woodland. Botanical interest is focused mainly on the ditches and sandier areas, with
orchids such as Common Spotted, Pyramidal and
Bee. Common Twayblade, Autumn Ladys-tresses
and Green-winged Orchid can also be found.
There is parking at Castle Water and a Visitor Centre
southeast of Rye along Harbour Road. (East Sussex
County Council)
12. Fore Wood
TQ 756126
A hornbeam and hazel coppice with some oak and
alder. There are wide grassy rides and a good spring
flora. Orchids include colonies of Early Purple and
Common Spotted Orchids, Broad-leaved Helleborine, Common Twayblade and Birds-nest Orchid.
Beside the village of Crowhurst, off the A2100 Battle to
Hastings road. (RSPB)
13. Wild Woods at Friston Forest TV 555995

This area of forestry includes extensive Beech plantations with many wide grassy rides. A rich flora is
developing, and includes a colony of several thousand White Helleborines.
Park in the Seven Sisters Country Park Forest car park
north of the A259 or off minor roads to the north;
there is a Visitor Centre at the car park, and waymarked walks. (FC)
14. Cuckmere Haven and Seven Sisters

Country Park
TV 519995
Lying just east of Seaford, the cliff-top chalk downs
have Common Fragrant, Common Spotted, Early
Purple, Pyramidal and Bee Orchids, Autumn Ladystresses and, in a few places, Early Spider, Frog and
Burnt Orchids (including the late-flowering variety
of Burnt).
Park in the Seven Sisters Country Park Riverside car
park on the south side of the A259. Paths lead east
along the cliff tops all the way to Beachy Head. (Sussex Downs Conservation Board)
15. Mount Caburn
TQ 445090
Just southeast of Lewes, Mount Caburn is an important Bronze Age hill fort; its south-facing chalk slopes
hold Common Fragrant and Pyramidal Orchids and
a large population of Burnt Orchids (over 6,000
plants in some years; look in the dry valley to the
northwest of the hill fort).
Access is from the village of Glynde, following the steep
public footpath to the fort from roughly opposite the
post office in Ranscombe Lane. (NNR)
16. Malling Down
TQ 423107
Old chalk pits on a steep north-facing slope, with
a rich flora including a selection of orchids. Musk
and Frog Orchids can be found in small numbers,
with good displays of Pyramidal and Common Fragrant Orchids and stunning numbers of Common
Spotted Orchids. There are also Bee Orchids and
Common Twayblades.
Easily reached by walking from Lewes town centre
(access in Wheatsheaf Gardens); alternatively, park in
the Ringmer Road lay-by on the B2192. (Sussex WT)
17. Castle Hill
TQ 371070
East of Brighton, on the South Downs between
Woodingdean and Kingston, this traditionally managed chalk grassland has Britains largest colony of Early
Spider Orchids, with up to 50,000 plants recorded.
Accessible along an un-signed track off the B2123 just
north of Woodingdean (park at TQ 356063 and walk
northeast for 1/2 mile). Access without a permit is
restricted to rights of way. (NNR)
18. Ditchling Beacon
TQ 330130
Lying to the north of Brighton on the South Downs
Way long-distance footpath, this Iron Age hill
fort has chalk grassland and scrub with Common
Fragrant Orchid, Common Spotted Orchid and
Common Twayblade (and also Marsh Fragrant
Orchid on north-facing slopes towards Plumpton).
The site lies a little off the South Downs Way to the
northwest of the National Trust car park (accessed
on minor roads south from the B2116 at Ditchling).
(Sussex WT)
29/1/09 12:49:23
SOUTHEAST ENGLAND
WEST SuSSEx
19. Devils Dyke
TQ 260110
On the South Downs to the north of Brighton, the
Devils Dyke is the largest dry chalk combe in Britain.
With some woodland and scrub, the Dyke has a rich
flora. The list of orchids includes Common Spotted,
Bee and Frog Orchids and Common Twayblade.
Well signposted on roads north of Brighton; park in the
large car park on Devils Dyke Road. (NT)
20. Cissbury Ring
TQ 130078
A large and impressive Iron Age hill fort on the
South Downs with Pyramidal, Bee and Frog Orchids
among its varied chalk flora.
Three miles north of Worthing, off the A24 at Findon.
Footpaths lead from Worthing and Findon. (NT).
21. Furnace Meadow and Brick Kiln Rough
Su 977277
A flower-rich meadow on clay soils, with an adjacent
ancient semi-natural woodland. The varied habitats
here hold a good orchid flora, with Early Purple,
Common Spotted, Greater Butterfly and Birdsnest Orchids, and Broad-leaved Helleborine among
others.
Three miles from Petworth on the minor road to Ebernoe. Park beside the church. The meadow is next to
the Sussex WT reserve of Ebernoe Common. (Sussex
WT, Plantlife)
22. Levin Down

Su 884138
A very steep chalk downland with Juniper and
patches of acidic chalk heath. Orchids include Pyramidal Orchid and Autumn Ladys-tresses.
Access on footpaths leading north from the minor road
between Singleton and Charlton, which is reached via
the A286. Park in the lay-by at the crossroads in Charlton. (Sussex WT)
23. Kingley Vale
Su 824088
The finest Yew forest in the world according to
English Nature. There is much of archaeological interest too, as well as some scrub and chalk grassland.
Species include Fly, Bee, Frog, Common Fragrant and
Pyramidal Orchids and Autumn Ladys-tresses.
On the South Downs, 41/2 miles northwest of Chichester. There is a car park and waymarked walks.
(NNR)
HaMPSHIRE
24. noar Hill
SY 740320
Medieval chalk pits that now form one of the best
examples of chalk grassland and scrub in Britain.
Rich in flowers and invertebrate life, 11 species of
orchid can be found, including a nationally important
427
colony of Musk Orchid. Also Early Purple, Frog,

Bee, Pyramidal, Common Spotted and Common
Fragrant Orchids, plus Common Twayblade and
Autumn Ladys-tresses.
Leave Selborne south on the B3006 and immediately
turn west towards Newton Valence. Then take the first
left, signposted Noar Hill. There is limited parking by
the footpath sign at SU 738323; follow this track to
the reserve. (Hampshire WT)
25. Butser Hill
Su 714198
This is one of the largest areas of chalk grassland in
Hampshire. It contains a variety of habitats, including Yew woodland and chalk heath. There is a good
orchid flora with Early Purple, Bee and Common
Spotted Orchids, and Broad-leaved Helleborine.
Part of the larger Queen Elizabeth Country Park, south
of Petersfield and north of Portsmouth on the A3.
There is a Visitor Centre and car parks. (Hampshire
County Council, FC)
26. Chapetts Copse/Hen Wood Su 654230

A beechwood, probably a plantation that was
established on the site of an ancient woodland,
with a very rich orchid flora. The star species is
Sword-leaved Helleborine; the largest colony in
Britain by far is found here, with more than 3,400
flowering plants. It even grows along the road bank
outside the wood. Other species include White
Helleborine, and Birds-nest, Fly and Common
Spotted Orchids.
Off the A32 between West Meon and East Meon; turn
south at the sign for Meon Springs Fly Fishery into
Coombes Lane, and park in the ride on the left after
about 450 yards. (Hampshire WT)
27. Old Winchester Hill
Su 645209
A spectacular hill fort at the west end of the South
Downs, comprising chalk grassland with mixed scrub,
Yew and broad-leaved woodland with 14 species of
orchid. On the south-facing slopes, large colonies of
Greater Butterfly, Frog, Common Fragrant Orchids
and Autumn Ladys-tresses can be found, with Man,
Bee and Fly Orchids, and Common Twayblade in
the scrub. White Helleborine, and Early Purple and
Birds-nest Orchids occur in the woodland.
From the A32 at Warnford, take the minor road southeast towards Clanfield. The car park is on the right
after 2 miles. (NNR)
28. St Catherines Hill
Su 485275
An ancient hill fort near Winchester city centre, with
a rich chalk flora including Musk and Frog Orchids,
and Autumn Ladys-tresses.
Access off the B3335 south of Winchester centre onto
Garnier Road. There is a large car park. (Hampshire
WT)
29/1/09 12:49:24
428
SITE GUIDE
29. Martin Down

Su 061192
A vast area of chalk grassland with wonderful displays of wildflowers, as well as other habitats such
as chalk heath and scrub. Various earthworks crisscross the relatively flat downland. Specialities include Burnt Orchid, which can be found around the
prominent Bokerley Dyke. Other species include
Common Fragrant, Pyramidal, Bee, Early Purple,
Greater Butterfly and Common Spotted Orchids,
hard-to-spot Frog Orchids, Southern Marsh Orchid,
and a very few Man Orchids.
There is open access on foot from the village of Martin,
south of Salisbury off the A354. (NNR, NE)
30. new Forest
Su 298081
The New Forest is one of the greatest biological treasures in Britain. An ancient system of grazing rights,
still widely exercised, has produced a rich and varied
landscape. Much of the forest is heath or woodland, with relatively few orchids, but the acid bogs
and valley mires are of exceptional interest. Heath
Spotted Orchid is very common, and the purpleflowered subspecies pulchella of Early Marsh Orchid
is fairly frequent. Bog Orchid occurs in the more
acidic bogs, such as Holmsley and Wilverley Bogs
(on the northern side of the Avon Water around
SU 230000 and SU 245005), the western fringe of
Long Beech Inclosure (north off the A31 at Stoney
Cross), Matley Bog (north of the B3056 21/2 miles
east of Lyndhurst), the area south and southwest of
Hatchet Pond (on the B3054 11/2 miles southwest
of Beaulieu), and at Acres Down. Bog Orchid can
be very hard to find, requiring patience and great
care, because it is easy to tread on them; some of
these sites are very wet with deep peat beware!
Another local speciality is Heath Fragrant Orchid,
found in a few of the more calcareous mires, such as
Stony Moors (north of Holmsley Camp Site), Dibden Bottom (west of Dibden) and Boundway Hill
(near Sway). Lesser Butterfly Orchids are relatively
frequent on the higher, drier ground in and around
the bogs and on adjacent wet heathland. Finally, the
New Forest was once home to Summer Lady'stresses, which, although probably extinct in the wild
since 1952, is reported to have been occasionally
reintroduced to its old sites, secretly.
THE ISLE OF WIGHT

31. Tennyson Down
SZ 327854
On land that drops steeply to the sea to end in
the famous Needles, this site is a combination of
coast and downland. The chalk grassland has large
colonies of Early Purple, Bee, Pyramidal and Greenwinged Orchids.
Take the B3322 out of Freshwater towards the Needles, where there is car parking. (NT)
32. Compton Down

SZ 375853
The most southerly area of chalk in the country, this
faces the sea. In spring there are Early Purple and
Green-winged Orchids, and later Pyramidal and Bee
Orchids amongst a rich chalk flora.
On the south coast of the Isle of Wight, east of Freshwater Bay and north of the A3055. (NT)
SuRREY
33. Thursley Common
Su 900417
Thursley is the largest remnant of the Surrey heaths,
with bog, dry and wet heath, and woodland. Southern Marsh Orchid and the purple-flowered subspecies of Early Marsh Orchid (pulchella) occur on the
wet heath (the creamy-flowered variety ochrantha
can also be found at Thursleynot to be confused
with the very rare creamy-flowered subspecies
ochroleuca!).
Access is from south of Godalming alongside the A3;
park between Elstead and Thursley. (NNR)
34. Wallis Wood
TQ 121388
Oak-hazel coppice with a stream, ponds and pasture.
There is a rich ground flora with Broad-leaved and
Violet Helleborines and Common Spotted Orchid.
On the minor road north of Walliswood village, just
east of the A29 south of Ockley. (Surrey WT)
35. Box Hill
TQ 180510
An outstanding area of woodland and chalk downland, Box Hill has long been famous as a destination
for day-trippers from London. Surprisingly extensive,
there are many walks and views towards the South
Downs. On the summit there is an Information
Centre and a nineteenth-century fort. There is an
impressive array of orchids, with Broad-leaved and
White Helleborines, Common Twayblade, Bee, Pyramidal, Common Fragrant and Common Spotted
Orchids, and Autumn Ladys-tresses in reasonable
numbers. Harder to find are Green-winged, Man,
Musk, Early Purple, Fly, Frog and Greater Butterfly
Orchids, and Violet Helleborine.
Box Hill is east of the A24 between Dorking and
Leatherhead. There are several car parks around the
perimeter and also along Zig Zag Road. (NT)
36. Ranmore Common, Denbies Hillside and

White Downs
TQ 145510
A cluster of National Trust sites on the North

Downs comprising a wooded common and chalk
downland. White Helleborine, and Birds-nest, Fly, Pyramidal and Bee Orchids occur among the scrub and
trees.There are also some Man Orchids and Autumn
Ladys-tresses.
The North Downs Way links Denbies Hillside and the
White Downs, with car parking two miles northwest of
Dorking off the A25. (NT)
29/1/09 12:49:24
SOUTHEAST ENGLAND
37. ashtead Common
TQ 175592
A relict woodland pasture with more than 2,300
pollarded oaks. Southern Marsh and Common
Spotted Orchids, Common Twayblade and Broadleaved and Violet Helleborines can be found here.
Near Junction 9 of the M25; access is from Woodfield
Road in Ashtead. Park at Ashtead Station. (NNR)
BERKSHIRE
38. Hurley Chalk Pit
Su 813822
A tiny reserve in a disused chalk pit with a surprisingly rich flora amidst its grassland and beechwood.
Common Spotted, Common Fragrant, Pyramidal
and Bee Orchids are all abundant. There are also
White Helleborines under the Beeches.
South of the B4130 Henley to Maidenhead road. Access from the bridleway south of the Black Boy Public
House. (BBOWT)
39. Greenham and Crookham Commons
Su 520643
Some Early Purple, Bee, Green-winged, Pyramidal,
and Heath Spotted Orchids, and Autumn Ladystresses can be found at these sites. Also visit nearby
New Greenham Park, where some very poor soils
and the removal of runways from the former airfield
have created good habitats for plants.
Around 3 miles southeast of Newbury; access to the
commons is off the A339 through New Greenham
Business Park, or from the minor road between Greenham and Thatcham.
40. Basildon Park

Su 611782
Woodland with Yew and Box, and open grassland. Bee
Orchid, and Narrow-lipped, Broad-leaved and Violet
Helleborines are among the species to search for.
Between Pangbourne and Streatley, northwest of Reading, on the west side of the A329. (NT)
41. Lardon Chase, the Holies and Lough Down
This area of National Trust-owned downland has
great views over the Thames Valley, where the
river divides the Chilterns from the North Wessex
Downs.The slopes form one of the largest remaining
areas of chalk grassland in the county and support
a wide range of invertebrates and flowers, including
Bee, Common Fragrant and Pyramidal Orchids.
Just north of Streatley, west of the A417. (NT)
OxFORDSHIRE
42. Hartslock
Su 616796
An area of rich chalk downland and scrub overlooking the Thames. The grassland is the sole Oxfordshire site for Monkey Orchid, while a handful
of Lady Orchids are also present, together with the
429
hybrid Monkey x Lady Orchid, as well as Common

Spotted, Bee, and Pyramidal Orchids, Common
Twayblade and White Helleborine.
One mile south along the minor road from Goring Station;
park by the cart track or in Goring village. (BBOWT)
43. Dry Sandford Pit

Su 467997
Based around an old quarry next to Sandford Brook,
with Common Spotted and Early Marsh Orchids,
Marsh Helleborine and Common Twayblade.
In Cothill village, 2 miles from Abingdon, off the B4017
Wootton Road. There is a small car park and marked
footpaths. (BBOWT)
44. Parsonage Moor
Su 462998
Part of the Cothill Fen NNR and one of the best
fenland sites in the county. Excellent woodland, fen,
carr and mire communities with Southern Marsh,
Pugsleys Marsh and Marsh Fragrant Orchids. Keep
to the footpaths because the fen is deep in places.
Use the same car park as for Dry Sandford Pit (site
43). The footpath to the reserve is opposite the Merry
Miller Restaurant. (BBOWT)
45. aston Rowant
Su 740968
Chalk grassland, scrub and beechwoods, now bisected by the M40, where White, Narrow-lipped and
Violet Helleborines occur amongst other species.
Between Stokenchurch and Lewknor. There is a car
park for the northern section of the reserve signposted
off the A40, with a Forestry Commission car park at
Cowleaze Wood serving the southern section. (NNR)
46. Watlington Hill
Su 702935
Fine chalk grassland and scrub with some stands of
Beech and Yew. There are Frog, Bee and Pyramidal
Orchids at the site.
One mile southeast of Watlington, east of the B480.
There is a small car park. (NT).
47. Warburg Reserve
Su 720879
This incredibly rich site contains woodland, scrub and
grassland in the dry valley at Bix Bottom. A network
of footpaths provides access to Broad-leaved, White,
Violet and Narrow-lipped Helleborines, Early Purple,
Birds-nest and Fly Orchids. Both Greater Butterfly and
Lesser Butterfly Orchids are present, along with Common Twayblade, and Bee, Common Spotted and Pyramidal Orchids. BBOWTs largest reserve is a real gem.
Northwest of Henley-on-Thames. Follow signposts for
Bix Bottom off the A4130 at Bix. The reserve is at the
end of a narrow two-mile road; there is a car park and
Visitor Centre. (BBOWT)
BuCKInGHaMSHIRE
48. Homefield Wood
Su 814867
Although some of the original beechwood has been
cleared and planted with conifers, there is still much
29/1/09 12:49:24
430
SITE GUIDE
mixed beechwood and open chalk grassland at this

site. It is traditionally managed and has a rich flora,
with 11 species of orchid including Birds-nest, Common Spotted, Fly and Bee Orchids and Broad-leaved
and White Helleborines, and it is one of only three
British sites for Military Orchid.
Off the A4155 west of Marlow. Turn right before the
Dog and Badger Public House.The reserve is on the left
between Bockmer and Lower Woodend. (BBOWT)
49. Grangelands and Pulpit Hill

SP 827049
These two sites hold a range of habitats, including
chalk grassland, scrub and beechwood, with Bee,
Common Fragrant, Musk, Frog, and Birds-nest Orchids and Violet, White and Narrow-lipped Helleborines.
Northeast of Princes Risborough on the A4010, on
the minor road from Askett to Great Missenden. Park
in the lay-by on the left-hand side of the road. Access
by the footpath going north. (Grangelands: BBOWT;
Pulpit Hill: NT)
50. Dancersend and the Crong Meadow
Su 616796
Woodland and meadows in the Chiltern Hills. The
chalk grassland has been cleared of scrub and has
a rich flora, with Common Spotted, Bee, Common
Fragrant and Pyramidal Orchids; Greater Butterfly
and Fly Orchids can also be found.
Off the B4009 Wendover to Tring road. Turn towards
St Leonards just before the junction with the A41; park
opposite the pond. (BBOWT)
51. Bernwood Meadows

SP 606111
This traditionally managed old hay meadow has a
large colony of Green-winged Orchids.
Northeast of Oxford, off the minor road between
Stanton St John and Oakley. Car parking is available beside the reserve on the south side of the road.
(BBOWT)
52. The ashridge Estate and Ivinghoe Beacon
SP 970131
Stretching along the northeastern edge of the
Chiltern escarpment on the HertfordshireBuckinghamshire border from Berkhamstead to
Ivinghoe Hills, this area is rich in archaeological remains. Bronze Age barrows survive around Ivinghoe Beacon and there is an impressive Bronze Age
hill fort at the top. The flora is excellent. There are
woodlands on the chalk scarp with Fly and Birdsnest Orchids, and Violet and White Helleborines,
wooded commons with Early Purple Orchids,
Green-winged Orchids, and hard-to-find Narrowlipped and Green-flowered Helleborines, and chalk
grassland with Bee, Pyramidal, Common Fragrant,
Frog and Common Spotted Orchids.
Just north of Berkhamsted, on both sides of the B4506

between the A41 and the B489. The main car park
is by the Visitor Centre, just off the B4506 between
Berkhamsted and Northchurch. (NT)
BEDFORDSHIRE
53. Dunstable and Whipsnade Downs
TL 000190
The extensive chalk grasslands here have fine views.
The grazed slopes are plant-rich, and Bee, Pyramidal
and Frog Orchids can be found.
Southwest of Dunstable, 4 miles northwest of Ashridge
between the B4540 and B4541. There are car parks
on the B4541. (NT)
54. Totternhoe Knolls

SP 979220
Part old chalk quarry (the Little Hills), part earthworks with grass and scrub, and part beech plantation, these varied habitats hold Common Spotted
and Common Fragrant Orchids, Common Twayblade and a few Musk Orchids.
Between Dunstable and Tring on the B489. Park in the
picnic site car park on Castle Hill Road. (BCNPWT)
55. Sharpenhoe Clappers, Smithcombe and
Sundon Hills
TL 065295
Sharpenhoe Clappers is the most prominent section
of chalk escarpment in Bedfordshire.This whole area
is interesting; there are beechwoods on the hill tops
with unimproved chalk grassland on both east- and
west-facing slopes. White Helleborine and Pyramidal,
Common Fragrant, Bee and Fly Orchids occur.
Reached via minor roads 1 mile southwest of Bartonle-clay. Park in the National Trust car park in Sharpenhoe village. (NT)
56. Barton Hills
TL 093295
A large area of chalk grassland, scrub, beechwoods
and ash-maple woodland in the Chilterns. Many species of orchid can be found including Bee, Common
Spotted and Common Fragrant Orchids.
On minor roads immediately south of Barton-le-clay.
Footpaths cross the area; park in Barton-le-clay. (NNR,
NE)
HERTFORDSHIRE
57. Hoo Bit
TL 117290
On the Icknield Way, this reserve has mixed broadleaved woods with a good shrub layer and some
chalk grassland. Adjacent to both Pegston Hills and
the Herts and Middlesex WT reserve of Telegraph
Hill, the whole area has a rich and interesting flora
which includes Fly, Bee and Common Spotted Orchids and White Helleborine.
Between Luton and Hitchin. Park at Treasures Grove
on the minor road off the A505 between Lilley and
29/1/09 12:49:25
SOUTHEAST ENGLAND
Hexton, and walk northeast along the Icknield Way.

(BCNPWT)
58. Hexton Chalk Pit
TL 107299
An old grassed-over chalk quarry with a good chalk
flora, and five species of orchid. The whole area is
rich in wildflowers, walks and good views.
Between Hitchin and Barton-le-Clay at Hexton. Park in
Hexton off the minor road to Lilley, and walk a short
distance south to the reserve. (Herts and Middlesex
WT)
59. Therfield Heath and Fox Covert
TL 335400
Therfield Heath is not a heath but a fine hillside
of unimproved grassland, chalk downland and
hawthorn scrub. Alongside the grassland is a small,
mature beechwood called Fox Covert. The flora
includes Bee and Common Fragrant Orchids, with
White Helleborine in the woodland.
At the roundabout on the A505 just west of Royston turn
south past the Little Chef and then right onto the minor
road towards Therfield. Access is from the small lay-by at
the top of the hill. (Herts and Middlesex WT)
60. The Meads
TQ 348141
One of the largest wetlands remaining in Hertfordshire, with flood meadows, pools and some chalk
grassland where Pyramidal Orchid can be found.
Between Hertford and Ware on the A119. Park in
Ware and walk west along the canal towpath to the
reserve. (Herts and Middlesex WT)
61. Lee Valley Park
This is the best site in the region to see spectacular numbers of Early Marsh Orchids. They thrive on
waste ash from coal-fired power stations; covering
an area the size of a football pitch, they are well
worth seeking out in June. There are also Southern
Marsh and Common Spotted Orchids. The main areas of interest are at Cheshunt Lock (TL 367023)
(an excellent boardwalk through a large colony of
Early and Southern Marsh and Common Spotted
Orchids); Bowyers Water (TL 367020) (near Cheshunt station) with Common Spotted, Early Marsh,
Southern Marsh Orchids, and Common Twayblade;
and Fishers Green (TL 367032) (close to Lee Valley
Information Centre), where small numbers of Bee,
Southern Marsh and Early Marsh Orchids surround
a large artificial pond, with Pyramidal Orchids close
to the nearby electricity substation.
The main access from the west is from the car park
close to Cheshunt station; from the east, head to the
Lee Valley Information Centre at Hooks Marsh (off the
B194 at Holyfield). The Centre can provide detailed
maps and directions to all these sites and information about orchid flowering times. (Lee Valley Regional
Park Authority)
431
GREaTER LOnDOn
62. Lee Valley Park: Rammey Marsh
TQ 370987
This area, along the outflow from King Georges
reservoir, has a magnificent display of Bee Orchids
in most years up to 15,000 have been counted.
There are also a few Pyramidal Orchids.
Near Enfield Lock. Park by the Swan and Pike Pool and
walk north towards the M25 along the canal. (Lee
Valley Regional Park Authority)
63. The Ripple
TQ 467824
Once a dumping ground for pulverised fuel ash,
this reclaimed industrial wasteland at Barking Reach
now holds a stunning display of hundreds of Southern Marsh and Common Spotted Orchids. May and
June are the best months to go.
Entrance to the site is on Thames Road/Renwick Road
in Barking. (London WT)
64. West Kent Golf Course
TQ 425612
This small reserve is close to Down House, which
was once the home of Charles Darwin. The chalk
grassland, scrub and woodland has a wonderful flora
and fauna with swathes of orchids.
Access is via a footpath from the village of Downe,
along West Hill and Milking Lane. (London WT)
65. Downe Bank
TQ 438609
Lying on the North Downs, this reserve, famous for
its association with Charles Darwin as his Orchis
Bank, has mixed woodland, coppice and chalk grassland. Many species of orchid occur such as Man, Bee,
Fly, Common Spotted and Pyramidal Orchids and
White and Broad-leaved Helleborines.
Downe Bank is situated midway between the villages
of Downe and Cudham, just south of Downe village,
where it is best to park. Formerly a closed reserve, the
southern half is now open for visitors. Contact the Kent
WT for access to the northern half. (Kent WT)
66. Hutchinsons Bank
TQ 381616 or TQ 386607
One of the largest areas of chalk grassland remaining in the Greater London area. The grassy slopes
hold Pyramidal, Common Spotted and Man Orchids. Common Twayblade and White Helleborine
occur on Chapel Bank, an area of ancient woodland,
scrub and grassland.
Southwest of New Addington, Croydon. Access is from
Featherbed Lane, with Hutchinsons Bank to the east,
via footpaths off Farleigh Dean Crescent (TQ 381616)
and Chapel Bank to the south and west (TQ 386607).
(London WT)
29/1/09 12:49:25
432
SITE GUIDE
Southwest England and Channel Islands

48
51
49
46
47
50
45
44
39
26
43
40
38
36 35
41
37
42
34
30 31
32
33
29
27
25
24
5
10
12
9
3
7
15
21
19
18
17
28
22
23
20
16
14
13
11
2
53
52
Cornwall
1 The Lizard Peninsula
2 Rame Head and Penlee Point
3 Sylvias Meadow (St Anns Chapel)
4 Creddacott Meadows
Devon
5 Dunsdon
6 Braunton Burrows
7 Andrews Wood
8 Berry Head - Sharkham Point
9 Dawlish Warren
Dorset
10 Kingcombe Meadows
11 Isle of Portland
12 Tadnoll and Winfrith Heath
13 Durlston Country Park
14 Ballard Down and Nine
Barrow Down
15 Corfe Mullen Meadow
16 Badbury Rings
17 Hambledon Hill
18 Fontmell and Melbury Downs
Wiltshire
19 West Wiltshire Downs
20
21
22
23
24
25
26
Pepperbox Hill
Ham Hill
Joness Mill
Pewsey Downs
Parsonage Down
Marlborough Downs
Clattinger Farm Reserve
Somerset
27 Long Sutton Plantations,
28 Barrington Hill Meadows
29 West Sedgemoor
30 Porlock Bay
31 Berrow Dunes
32 Ash Priors Common
33 Great Breach Wood
34 Cheddar Gorge and Black Rock
Nature Reserve
avon
35 Blagdon Lake Pumping Station
36 Hellenge Hill
37 Walborough
38 Netcotts Meadow
39 Ashton Court Meadow
40 Avon Gorge/Leigh Woods
41 Browns Folly
42 Folly Farm
Gloucestershire
43 Lower Woods
44 Minchinhampton and
Rodborough Commons
45 Strawberry Banks
46 Elliot (Swifts Hill)
47 Plump Hill Dolomite Quarry
48 Betty Daws Wood
49 Painswick Hill
50 Cotswold commons and
beechwoods
51 Greystones Farm and
Salmonsbury Meadows
Channel Islands
52 Le Noir Pr, Jersey
53 The Orchid Fields at Les Vicheries,
Guernsey
29/1/09 12:49:25
SOUTHWEST ENGLAND
CORnWaLL
1. The Lizard Peninsula
SW 687133 (Kynance)
SW 667163 (Predannack)
The Lizard is the most southerly point of the British Isles and has many rare native plants. Orchids,
however, are of interest more from a local viewpoint than a national one. Around 10 species occur,
including Green-winged and Early Marsh Orchids,
which occur virtually nowhere else in Cornwall.
Heath Fragrant Orchid can be locally common on
the Lizards heaths; others present include Common
Twayblade and Autumn Ladys-tresses.
Many orchids can be seen from the coastal footpath
between the car parks at Kynance and Predannack
Wollas. (NNR, NT)
2. Rame Head and Penlee Battery Sx 436492

An area of coastal grassland with extensive gorse
scrub. An interesting area, it holds Cornwalls main
population of Bee Orchid and is the only known site
in southeast Cornwall for Pyramidal Orchid. Early
Purple and Southern Marsh Orchids can be found
along the footpaths, as well as a few Autumn Ladystresses.
Parking is available at Penlee Battery Cornwall WT reserve and at Rame Head, from which coastal paths
are easily accessed.
3. Sylvias Meadow (St anns Chapel)
Sx 413707
This small reserve of unimproved herb-rich grassland is one of Cornwalls best single sites for orchids,
with eight species recorded. Heath Spotted, Lesser
Butterfly and Early Purple Orchids can be abundant,
and there may be a few spikes of Greater Butterfly
Orchid. Southern Marsh Orchids and their hybrids
with Heath Spotted Orchid also occur. Common
Spotted Orchid is a rarity in Cornwall but a few can
be found here, along with Autumn Ladys-tresses
and Broad-leaved Helleborine.
South of Kit Hill and west of Gunnislake, with parking
available at Honicombe. A public footpath runs south
from the A390 alongside the reserve. (Cornwall WT).
4. Creddacott Meadows
Sx 234963
One of the last remnants of species-rich damp culm
grassland in Cornwall. Lesser Butterfly Orchid and
Heath Spotted Orchid occur, among others.
Park off the B3254 Launceston to Bude road, at Week
St Mary. (Cornwall WT, Plantlife)
DEVOn
5. Dunsdon
SS 302080
One of the best examples of culm grassland in
433
Devon. There are good displays of Lesser Butterfly,

Heath Spotted and Southern Marsh Orchids.
Between Holsworthy and Bude, off the A3072 to Pancrasweek. The entrance to the reserve and car park is
just before Gains Cross. (Devon WT)
6. Braunton Burrows
SS 464326
The largest sand dune system in the UK, with a very
diverse plant community. There are vast numbers
of Pyramidal Orchids, and also Southern Marsh
Orchids and large colonies of Marsh Helleborines
in the dune slacks. Recently named the UKs first
Biosphere Reserve.
On the north bank of the Taw/Torridge estuary,
west of Barnstaple. There are two car parks with
information.
7. andrews Wood
Sx 707515
A fascinating reserve of ancient meadows and
woods, over an old field system. The flora benefits
from traditional management; there are Southern
Marsh and Heath Spotted Orchids, with Broadleaved Helleborine and Common Twayblade in the
woodland.
Close to Loddiswell north of Kingsbridge, there is a
track to the reserve from the car park just beyond
Coldharbour Cross. (Devon WT)
8. Berry Head - Sharkham Point Sx 945567
A large coastal area comprising two headlands
and a smaller promontory, Durl Head. There is an
important flora on the cliff-tops, and in the grassland
and scrub areas, with eight species of orchid. These
include Early Purple and Green-winged Orchids,
Common Twayblade, Bee, Common Spotted and
Pyramidal Orchids, and Autumn Ladys-tresses.
The site lies at the southern end of Torbay past Brixham. (Berry Head Country Park, NNR)
9. Dawlish Warren
Sx 983787
A range of coastal habitats around a double sand
spit that extends across the mouth of the River Exe.
The dune grassland is of great botanical interest and
has Autumn Ladys-tresses in good numbers.
Off the A379; the car park is through the entrance
tunnel under the railway in Dawlish Warren village.
(Devon WT, NNR)
DORSET
10. Kingcombe Meadows
SY 555992
A large area of species-rich old grassland, hedgerows, woodland and common land a real ancient
landscape. Heath Spotted, Early Purple and Bee Orchids are abundant.
The site is at Lower Kingcombe near Toller Porcorum.
There are small car parks, footpaths and a Visitor Centre at Pound Cottage, off the B356. (Dorset WT)
29/1/09 12:49:26
434
SITE GUIDE
11. Isle of Portland

SY 685700
Famous for its limestone quarries, there is still much
of interest on the cliff-tops and in the abandoned
quarry workings. Two are worthy of visits to search
for orchids:The Broadcroft Quarry Reserve and Tout
Quarry. There is also a large car park at Portland
Bill, and a variety of orchids to be found in the surrounding grasslands. Bee and Pyramidal Orchids and
Autumn Ladys-tresses are particularly abundant.
Access to the Isle of Portland is on the A354 from
Weymouth. Carry on to Southwell and the large car
park at the lighthouse for the Bill. (BC)
Access is from the car park on the north side of the

B3082 Wimborne to Blandford road. (NT)
12. Tadnoll and Winfrith Heath
ST 884176, ST 885187
An outstanding area of chalk downland, scrub and
woodland, which includes Melbury Beacon. Autumn
Ladys-tresses and Green-winged, Common Fragrant, Greater Butterfly, Pyramidal, Bee, Early Purple
and a few Frog Orchids can all be found there
Access is via footpaths from the National Trust viewpoint car park at the top of Spread Eagle Hill. (NT,
Dorset WT)
SY 795876
A large example of Dorset heath, bogs and meadows, part of Thomas Hardys Egdon Heath, with
typical flora and fauna. Heath Spotted and Southern
Marsh Orchids occur, along with the hard-to-find
Bog Orchid, and Autumn Ladys-tresses.
From the A352, the Tadnoll entrance is along Redbridge Road, while the Winfrith Heath entrance is on
Gatemore Road (with limited parking). (Dorset WT).
13. Durlston Country Park

SY 032773
Limestone grassland occurs along the south coast
of Dorset and its speciality, Early Spider Orchid, can
be found anywhere from Durlston Head west to
Worth Matravers. Other species in the area include
Green-winged, Early Purple and Bee Orchids.
Durlston Country Park lies on a minor road south from
Swanage.
14. Ballard Down and nine Barrow Down
SY 050820
On the Isle of Purbeck chalk ridge, these downs
contain steep slopes and cliff-tops, with Common
Fragrant and Early Purple Orchids present.
Between Ballard Head and Old Harry Rocks, the
downs are easily accessible on foot from car parks at
Studland (NT) and Swanage.
15. Corfe Mullen Meadow
SY 980967
A small but flower-rich meadow with a stunning
colony of Green-winged Orchids.
Northwest of Poole on the B3074 at Corfe Mullen,
with roadside parking. (Dorset WT)
16. Badbury Rings

ST 964029
An Iron Age hill fort on the Kingston Lacy Estate.
A wonderful site with a rich downland flora and a
great variety of orchid species. On the hill fort itself Frog Orchid is a speciality; other species include
Common Fragrant and Common Spotted Orchids,
Common Twayblade, Early Purple, Green-winged,
Pyramidal and Bee Orchids, Greater Butterfly Orchid and Autumn Ladys-tresses, with White Helleborine and Birds-nest Orchid under the roadside
Beech trees.
17. Hambledon Hill

ST 845125
Dramatic chalk grassland with an impressive Iron
Age earthwork. A rich chalk flora includes Early
Purple and Pyramidal Orchids.
Northwest of Blandford Forum off the A350, there is
limited parking off Duck Street and Shaftsbury Road
near Child Okeford. Plenty of footpaths give good access to the site. (NNR)
18. Fontmell and Melbury Downs
WILTSHIRE
19. West Wiltshire Downs
A wonderful landscape of natural and archaeological features, southwest of Salisbury. There are many
places to visit here is a selection.
Coombe Bisset Down (Su 111256) Rich chalk grassland with scrub and beechwoods. There are many
Bee Orchids, with the very local Burnt Orchid.
Clearbury Rings (Su 152245), with over 1,000
Burnt Orchids on the gentle, south-facing slopes of
Clearbury Down.
Middleton Down (Su 043252), with Early Purple
Orchids and Autumn Ladys-tresses, among others.
20. Pepperbox Hill

Su 212248
Rough chalk grassland and mixed scrub with a good
selection of orchid species. Common Spotted, Frog,
Common Fragrant and Pyramidal Orchids, White
Helleborine, Common Twayblade and Autumn
Ladys-tresses all occur.
Off the A36 southeast of Salisbury, with parking on the
minor road between West Grinstead and West Dean.
(NT)
21. Ham Hill
Su 334616
A small, very steep chalk downland reserve with
abundant wildlife and good views. The flora includes
Common Twayblade and Common Fragrant Orchid,
and also a few Musk Orchids.
Off the A336 Burbage to Hungerford road, between
Shalbourne and Ham on the road to Buttermere, with
parking on the left-hand verge. (Wiltshire WT)
29/1/09 12:49:26
SOUTHWEST ENGLAND
22. Joness Mill

Su 168613
Wet woodland and fen meadows with Southern
Marsh and Common Spotted Orchids.
Just north of Pewsey off the B3087 to Burbage; the
reserve entrance is close to a small lay-by, just over the
railway bridge. Alternatively, walk from Pewsey along
the Kennet & Avon towpath. (Wiltshire WT)
23. Pewsey Downs
Su 120640
Perhaps the finest site on the Marlborough Downs,
comprising Milk Hill, White Horse Hill, Walkers Hill
and Knapp Hill, with some of the best habitats the
region can offer. Common Spotted, Burnt, Bee, Frog,
Common Fragrant, Lesser Butterfly, Green-winged
and Pyramidal Orchids and Autumn Ladys-tresses
all occur here.
In the Vale of Pewsey between Devizes and Pewsey,
southwest of Marlborough. Access is via the unclassified road between the A361 and A345. Car parking is
at Knapp Hill. (NNR)
24. Parsonage Down
Su 055415
Some wonderful downland and pastures with a
history of traditional grazing management. There
are many interesting species here; orchids include
Green-winged, Frog and perhaps as many as 30,000
Burnt Orchids spread over 95 hectares, forming the
largest colony in Northwest Europe.
The site is at Cherry Lodge Farm, Shrewton, on the
south edge of Salisbury Plain, between Shrewton and
Winterbourne Stoke on the B2083. (NNR)
25. Marlborough Downs
An impressive area surrounding Marlborough, mostly consisting of gently undulating chalk grassland.Traditional grazing has ensured a rich flora and fauna,
with many areas now designated nature reserves.
Here are two great places to visit.
Morgans Hill (Su 025672) and Kingsplay Hill (Su
006658) On the western edge of Marlborough
downs with chalk grassland and scrub; both have a
good orchid flora.
Cherhill Down and Oldbury Castle (Su 046694)
The chalk grassland on the southern slopes has a
rich flora, including Lesser Butterfly Orchid. (NT)
26. Clattinger Farm Reserve

Su 017937
This site consists almost wholly of unimproved
species-rich alluvial grasslands, with spectacular displays of hay-meadow flowers. Green-winged Orchid,
Common Twayblade, Common and Heath Spotted
Orchids, Early and Southern Marsh Orchids and
Burnt Orchid (outside its normal downland habitat)
are present.
Off the B4040 between Malmesbury and Cirencester,
with parking on the roadside. (Wiltshire WT)
435
SOMERSET
27. Long Sutton Plantations, Burnt House
and Mondays Court Lane
ST 461269
This is a wonderful mix of lanes, plantations and beech
copses, with White Helleborine, Greater Butterfly
Orchid and Common Twayblade in the woods and
Bee and Pyramidal Orchids in the open areas.
Northwest of Yeovil, off the B3165 between Long Sutton and Mortock. Park on the road verge before the
railway bridge.
28. Barrington Hill Meadows

ST 290169
Several species-rich unimproved meadows on the
southern edge of the Blackdown Hills, with old
hedges and ponds.There is a large colony of Greenwinged Orchids.
Northwest of Ilminster on the minor road off the A358
between Bickenhall and Broadway. Park beside the
road along Folly Drive. (NNR)
29. West Sedgemoor
ST 365255
Woodland and traditionally managed grazing meadows, often flooded in winter. The wet meadows
contain a wide variety of flowers; one notable field,
slightly higher than the rest, holds cowslips and
Green-winged Orchids.
East of Taunton on the A378 Langport road. Car parking is east of Fivehead. (RSPB)
30. Porlock Bay
SS 885482
A fine area of species-rich dunes and foreshore that
is particularly good for orchids, with a large colony
of Autumn Ladys-tresses.
Porlock is west of Minehead on the A39. Minor roads
lead to the coast and the coastal footpath.
31. Berrow Dunes
ST 299515
An area of reedbeds and dunes beside Berrow Golf
Course. The dune slacks hold Southern Marsh Orchids and Marsh Helleborines in good numbers, and
Heath Spotted and Early Marsh Orchids also occur,
with Lizard and Bee Orchids (including var. belgarum
of Bee) on the drier areas..
Just north of Burnham-on-Sea. Access is on waymarked
trails from the car park on Coast Road, Berrow (opposite Sandy Glade Caravan Park). (LNR, Sedgemoor
District Council)
32. ash Priors Common
ST 151289
Scrub, grassland, woodland and heath habitats, with
a good variety of orchids: Heath Spotted, Common
Spotted, Southern Marsh and Early Purple Orchids,
and Common Twayblade all occur.
Northwest of Taunton off the A358 at Ash Priors. Car
parking is available beside the common. (Taunton and
District Borough Council)
29/1/09 12:49:26
436
SITE GUIDE
33. Great Breach Wood

ST 506325
One of Somersets largest reserves, with mixed
woodland and grassland. In some areas orchids are
plentiful; there are Early Purple, Greater Butterfly
and Pyramidal Orchids on the steep south- and
west-facing slopes, for example. There are many
rides and paths through the site.
South of Glastonbury, between Street and Somerton.
Park near Butleigh Park on the minor road from the
B3153. (Somerset WT)
34. Cheddar Gorge and Black Rock
ST 482545
Limestone grassland, plantation and woodland at
the head of Cheddar Gorge. Part of the Cheddar
complex, the area is botanically rich; orchids to be
found include Broad-leaved and Narrow-lipped
Helleborines.
Park at Black Rock Gate, on the B3135 just northwest
of Cheddar. (Somerset WT)
aVOn
35. Blagdon Lake Pumping Station ST 503598
An excellent if unusual site for Green-winged

Orchids.
Bristol Water have many open days for public access.
Contact them for details of orchid flowering times.
(Bristol Water. Tel: 0117 966 5881)
36. Hellenge Hill

ST 345572
Calcareous grassland and scrub on the south side
of the Mendips with great views. Autumn Ladystresses are abundant in some areas.
At Bleadon, south of Weston-Super-Mare. (Avon WT)
37. Walborough
ST 316579
An area of limestone grassland beside the sea wall
at Uphill. There are Green-winged and Early Purple
Orchids and Autumn Ladys-tresses.
South of Uphill village. Park by the sluice gates near the
beach and walk along the sea wall. (Avon WT)
38. netcotts Meadow
ST 476696
An area of semi-improved neutral, damp grassland
with Green-winged, Early Purple, Southern Marsh
and Bee Orchids present in large numbers.
Close to Nailsea, Bristol, with parking after the railway
bridge. (Avon WT)
39. ashton Court Meadow
ST 545720
Part of the Ashton Court Estate and close to the
centre of Bristol. In May the meadow is full of
Off the A369 Bristol to Portishead road. Park in the layby on Beggar Bush Lane, the B3129. (Avon WT)
40. avon Gorge
ST 553731
A fantastic area, the Avon gorge cuts through the
limestone only 2 miles from the centre of Bristol.

Clifton and Durdham Downs are large areas of
limestone grassland with abundant Bee and Common Spotted Orchids. There are smaller numbers
of Pyramidal and Green-winged Orchids. On the
west side of the gorge Leigh Woods (ST 555730)
hold Birds-nest and Fly Orchids, and Broad-leaved
Helleborines.
The area is easily accessed from Bristol on the A4018
and the A4. Leigh Woods are on the A369 Portishead
to Bristol road.
41. Browns Folly

ST 795665
Calcareous grassland and mature secondary woodland with some ancient semi-natural areas. Broadleaved Helleborine together with Early Purple, Bee,
Pyramidal and Common Spotted Orchids can be
found on and around the grassland. Elsewhere,
Birds-nest Orchid and White Helleborine occur
within the wood.
East of Bath, off the A36 at Bathampton opposite the
golf course. (Avon WT)
42. Folly Farm
ST 607605
Species-rich neutral grassland and ancient seminatural woodland. There are Common Spotted and
Heath Spotted Orchids, as well as hybrids between
the two. Common Twayblades are also present
within the wood, with Early Purple Orchids and Violet Helleborines.
East of Bishop Sutton, south of Bristol; the site is signposted off the A368, to the east of its junction with the
A37. (Avon WT)
GLOuCESTERSHIRE
43. Lower Woods
ST 743876
One of Englands largest stands of ancient oak-ash
woodland. It contains a diversity of habitats, including species-rich grassland, coppiced woodland and
wooded common. Early Purple, Greater Butterfly
and Common Spotted Orchids and Violet Helleborine can all be found.
Between Hawksbury and Wickwar. Park by the
Lodge on the minor road off the B4509 at Wickwar.
(Gloucestershire WT)
44. Minchinhampton Common
and Rodborough Common
SO 850010
SO 850038
Some of the best areas of limestone grassland on
the edge of the Cotswolds. There are great views,
and Minchinhampton has much of archaeological
interest. Many species of orchid have been recorded among the varied flora.There are Common
Spotted, Pyramidal, Early Purple, Green-winged,
Common Fragrant, Bee, Frog and Greater Butterfly Orchids, as well as Autumn Ladys-tresses,
Common Twayblade and White, Broad-leaved and
29/1/09 12:49:27
SOUTHWEST ENGLAND
Narrow-lipped Helleborines. Both commons are

worth exploring.
Off the A46 between Stroud and Nailsworth. (NT)
45. Strawberry Banks
SO 910033
This limestone grassland on a west-facing slope has
a rich flora. There is also some mixed scrub and a
small stream. Greater Butterfly, Bee, Common Spotted, Green-winged, Early Purple and Pyramidal Orchids all occur. It is reached by walking through Three
Groves Wood Nature Reserve, where Broad-leaved
Helleborine can be found.
East of Chalford between France Lynch and Oakridge villages.There is a small car park. (Gloucestershire WT)
46. Elliot (Swifts Hill)
SO 877067
This area of limestone grassland and old quarry
workings is carefully managed by grazing, and 11
species of orchid regularly occur. These include
Pyramidal, Common Fragrant, Early Purple and Fly
Orchids, and Autumn Ladys-tresses.
Just northeast of Stroud, on the minor road to Elcombe.
(Gloucestershire WT)
47. Plump Hill Dolomite Quarry
SO 661171
An interesting site with steep exposed rock faces
and a limestone flora on the quarry floor. There is a
good colony of Autumn Ladys-tresses.
South of Mitcheldean beside the A4136. Park in the
lay-by. (Gloucestershire WT)
48. Betty Daws Wood
SO 696284
An ancient Sessile Oak woodland, part of a larger
complex, with a good ground flora that includes
Birds-nest Orchid and White Helleborine.
West of Gloucester, northwest of Newent, at Four
Oaks. Park and walk from the village. (Gloucestershire WT)
49. Painswick Hill
SO 869120
Excellent chalk grassland on a steep-sided hill fort,
where Bee, Fly, Musk, Frog and Greater Butterfly
Orchids, and Autumn Ladys-tresses can be found.
South of Gloucester on the B4073 to Painswick. There
are several footpaths and two long-distance paths, the
Wysis Way and the Cotswold Way, meet here.
437
Broad-leaved, White, Narrow-lipped, and Greenflowered Helleborines.

Access to Buckholt Wood at various points from minor
roads east off the A46, 3 miles northeast of Painswick.
(NNR)
51. Greystones Farm and Salmonsbury

Meadows
SP 173209
With traditional hay meadows, grazing land and

streams, this is an attractive place to visit. Early
Marsh and Southern Marsh Orchids are abundant
in the meadows.
Close to Bourton-on-the-Water; park and follow the
Oxfordshire Way, off Moor Lane. (Gloucestershire
WT)
THE CHannEL ISLanDS

52. Le noir Pr, Jersey
Two adjoining wet meadows, the speciality here is
Loose-flowered Orchid. There are also Southern
Marsh, Heath Spotted and Common Spotted Orchids.
The entrance is at Le Chemin de LOuziere, a minor
road off Le Grand Route de Mielle (Five Mile Road).
There is an open day in late May, and unrestricted
access for several weeks in the summer. (National
Trust for Jersey)
53. The Orchid Fields at Les Vicheries,

Guernsey
An area of low-lying wet meadows in southwest
Guernsey. The rich flora includes Loose-flowered
Orchid as well as Heath Spotted, Common Spotted
and Southern Marsh Orchids.
In the St Peter in the Wood region. The reserve lies
close to the main coast road along the Rue du Douit
du Moulin and the Rue des Vicheries, where there is a
La Socit Guernesiaise information board.
50. Cotswold commons and beechwoods

The finest Cotswold beechwoods occur on the
scarp and dip slopes between Gloucester and
Stroud around Cranham, Sheepscombe and Painswick. The whole area contains important examples
of ancient woods, grassland and streams and there
are many designated nature reserves. Buckholt
Wood (SO 894131) is one of the finest with an
interesting shrub layer and good ground flora. Orchids typical of beech woodland occur here, such
as Birds-nest and Greater Butterfly Orchids and
29/1/09 12:49:27
438
SITE GUIDE
East anglia
12
10
13
11
14
16
15
9
17
8
18
23
7
6
5
4
22
20
25
24
19
21
27
26
28
29
30
32
Cambridgeshire
1 Hayley Wood
2 Thriplow Meadows
3 Beechwood
4 Fulbourn Educational Reserve
5 Devils Dyke
6 Wicken Fen
7 Soham Meadows
8 Upwood Meadows
norfolk
9 Narborough Railway Line
10 Holme Dunes
11 Holkham
12 Wells Woods
13 Beeston Regis and Sheringham Commons
14 Overstrand
15 Foxley Wood
16 Buxton Heath
17 Thompson Common
18 New Buckenham Common
31
Suffolk
19 Reydon Wood
20 Winks Meadow
21 Redgrave and Lopham Fens
22 Market Weston Fen
23 Rex Graham Reserve
24 Bradfield Woods
25 Chippenhall Green
26 Wolves Wood
27 Groton Wood
Essex
28 West Wood
29 Hatfield Forest
30 Roding Valley Meadows
31 Langdon
32 Grays Chalk Quarry
29/1/09 12:49:27
EAST ANGLIA
CaMBRIDGESHIRE
1. Hayley Wood
TL 294534
An ancient woodland with a long recorded history
of management.There are fine displays of woodland
flowers in spring, with Early Purple, Common Spotted and Birds-nest Orchids.
On the B1046 between Cambridge and Sandy, about
3 miles from Gamlingay. Park on the road verge.
(BCNPWT)
2. Thriplow Meadows
TL 445470
A small area of traditionally managed unimproved
grassland. Bee Orchids are among the rich flora on
the dry ground, with good populations of Early and
Southern Marsh Orchids in the wetter areas.
Parking is by the Village Hall in Thriplow, south of Cambridge off the B1368. A footpath leads to the reserve
from the road junction. (BCNPWT)
3. Beechwood LnR
TL 486548
A beech plantation on thin chalky soils, with a fine
colony of White Helleborines (3,800 were counted
in 1998) and a few Common Twayblades. Greenflowered Helleborine has been recorded here.
From the roundabout on the A1307 southeast of Cambridge turn northeast onto the minor road towards
Cherry Hinton; then turn right towards Fulbourn. Park
in the lay-by opposite the wood. (BCNPWT)
439
in plant life with Southern Marsh and Early Marsh

Orchids (in a variety of colour forms).
Signposted from Wicken village, on the A113 northeast
of Cambridge. With nature trails and a Visitor Centre.
(NNR, NT)
7. Soham Meadows
TL 608722
An area of traditionally managed unimproved
grassland with Bee and Common Spotted Orchids,
Common Twayblade and a small number of typically
elusive Frog Orchids.
Alongside the A142 Soham bypass, with parking in the
lay-by. (BCNPWT)
8. upwood Meadows
TL 251825
A permanent pasture on calcareous boulder clay.
The ridge and furrow landscape indicates an ancient arable history, but the meadows have been
grassed and traditionally managed for at least 300
years. They have a rich flora with large numbers of
Green-winged Orchids in May.
Two and a half miles southwest of Ramsey, which is
on the B1040. Go down Meadow Road and park in
Upwood. (BCNPWT)
nORFOLK
9. narborough Railway Line
TL 528560
Ancient wet meadows with alder woods and scrub;
Common Twayblade, Early Marsh and Southern
Marsh, Common Spotted and Bee Orchids can be
found in the rough pasture.
On the minor road southeast out of Fulbourn leading
to Balsham. Park on the roadside. (BCNPWT).
TF 750118
This sunny railway embankment was built with chalk
ballast, and now supports a rich chalk flora. Common Twayblade, and Southern Marsh and Common
Spotted Orchids occur along the path, with Marsh
Helleborines in the pits on the sides.
Take the A47 from Narborough towards Swaffham;
there is a small car park beside the railway line. (Norfolk WT)
5. Devils Dyke
10. Holme Dunes
4. Fulbourn Educational Reserve
TL 616616
This massive 8-mile long earthwork, a ditch and
rampart about 60 feet high, is thought to have been
built in the sixth century. It bisects Newmarkets July
race course and continues towards Reach. It supports important areas of chalk grassland notable for
colonies of Lizard Orchids (especially to the southeast of the A45). Other species include Pyramidal
and Common Spotted Orchids.
A footpath runs along the dyke from the roundabout at
the junction of the A1303 and A1304, 2 miles southwest of Newmarket. The Dyke can also be accessed
from the B1102 between Swaffham Prior and Burwell.
6. Wicken Fen
TL 554702
One of Britains oldest nature reserves, Wicken Fen
lies nine miles northeast of Cambridge, and contains
reedbeds, sedge-fen, ditches, open water, peat diggings, and birch and sallow carr. The open fen is rich
TF 697438
This site includes a range of coastal habitats, with
good dune and saltmarsh communities. Marsh Helleborine, Early Marsh Orchid (including the redflowered coccinea), and Southern Marsh Orchid are
common.
Access is from the A149 east of Hunstanton, through
Holme village. There are car parks at the beach and
at the reserve centre one mile further east. (NNR,
Norfolk WT)
11. Holkham
TF 892441
The westward continuation of Wells Woods (site
12), but with more extensive dune slacks on the
seaward side of the pines. Marsh Helleborine and
Southern Marsh Orchid can be abundant, with scattered Common Spotted, Pyramidal and Bee Orchids. Creeping Ladys-tresses occurs very locally in
the pinewoods.
Access is from Lady Annes Drive (north off the A149,
29/1/09 12:49:28
440
SITE GUIDE
opposite the entrance to Holkham Hall); the areas to

the west of the car park, including the dunes past the
west end of the pines, are best. (NNR)
12. Wells Woods
TF 913454
These coastal dunes are planted with Corsican
Pines, but contain some relict areas of dune slacks,
especially at The Dell, where Southern Marsh and
Common Spotted Orchids and Marsh Helleborine
are abundant, with a few Common Twayblades and
Bee Orchids as well. Creeping Ladys-tresses occurs
very locally in the pinewoods.
Access is from Wells beach car park (signposted from
the town centre);The Dell lies to the north of the path
450 yards west of the kissing gate.
13. Beeston Regis and Sheringham Commons
TG 166425
A small but very diverse site with rough grassland, alkaline mires, acid heath and a little woodland. Marsh Helleborine and Common Spotted
and Marsh Fragrant Orchids are abundant, as are
a confusing mixture of marsh orchids (many Pugsleys Marsh Orchids and intermediates, plus a few
pure Southern Marsh). Common Twayblade and
Bee, Pyramidal and Heath Spotted Orchids occur
in small numbers; Early Purple and Lesser Butterfly
Orchids are also present but hard to find.
Access is from the lay-by on the south side of the A149,
1 mile east of Sheringham.
14. Overstrand
TG 246411
The slumped clay cliffs at Overstrand have many
flushes and seepages, and hold large numbers of the
delightful red coccinea form of Early Marsh Orchid,
as well as Southern Marsh, Common Spotted and
Bee Orchids and Marsh Helleborines.
Access is from the cliff-top car park at Overstrand, on
the coast east of Cromer. Take the path to the beach
and walk north, scrambling up the slopes from time to
time (it can be very slippery).
15. Foxley Wood
TF 049229
The largest area of ancient woodland in Norfolk,
with wide, grassy, flower-rich rides. Early Purple,
Common Spotted and a very few Greater Butterfly
Orchids can be seen here.
Fifteen miles northwest of Norwich at Foxley village, off
the A1067 Norwich to Fakenham Road. There is a car
park at the wood. (Norfolk WT)
16. Buxton Heath
TG 172216
An area of heath and mire with a small alkaline
stream. The mix of vegetation is very interesting
with Marsh Helleborine, Marsh Fragrant Orchid,
Common and Heath Spotted Orchids and Southern and Pugsleys Marsh Orchids (and some confusing intermediates).
Just east of the B1149 Norwich to Holt Road northwest of Heavingham.There is a small car park.
17. Thompson Common
TL 934967
A fascinating place to visit, with open water (Thompson Water), carr, grassland, scrub and woodland as
well as a number of pingos (shallow ponds formed
by glacial activity). Early Marsh (subspecies incarnata) and Southern Marsh Orchids can be found, and
possibly Pugsleys Marsh Orchid.
Accessed from Watton on the A1075 to Thetford. Car
parking is available just before Stow Bedon at the
Great Eastern Pingo Trail car park. (Norfolk WT)
18. new Buckenham Common

TM 090906
This traditionally managed common has never been
ploughed. The sites speciality is a large colony of
Four miles south of Attleborough on the B1113 at New
Buckenham, with parking available beside the common. (Norfolk WT)
SuFFOLK
There are many well-managed Roadside Nature Reserves in Suffolk that hold some interesting and local
orchids; look out for the wooden marker posts. In particular, the Claydon and A140 roundabouts along the
A14 hold huge colonies of Pyramidal Orchids.
19. Reydon Wood

TM 480788
This small ancient wood is managed as a Community Wood by local people. There is hazel coppice,
ancient Ash stools, and many rides. Several orchid
species occur here, including Early Purple, Common Spotted and Birds-nest Orchids and Common
Twayblade.
On the B1126 Southwold to Wangford road. Park at
the end of Wood Lane. (Suffolk WT)
20. Winks Meadow
TM 303799
This is a traditionally managed meadow with an excellent flora. Species to be found include Common Twayblade, Common Spotted, Green-winged, Early Purple,
Pyramidal and Bee Orchids, and the only colony of
Frog Orchids in Suffolk (they are hard to find).
Accessed via Metfield on the B1123. Take the road
opposite the garage, parking on the concrete pad near
the meadow entrance. (Suffolk WT, Plantlife)
21. Redgrave and Lopham Fens
TM 046797
This large reserve lies on the border between
Suffolk and Norfolk in the Waveney Valley. The internationally important valley fen is flanked by woodland, open water, reed and sedge beds and wet
heath. Southern and Early Marsh Orchids, Common
Twayblade, and a few Marsh Fragrant Orchids are
present.
29/1/09 12:49:28
EAST ANGLIA
Signposted between Redgrave and South Lopham on

the B1113; there are car parks and a Visitor Centre.
(NNR, Suffolk WT)
22. Market Weston Fen
TL 981789
A fine remnant of a valley fen dominated by Great
Fen Sedge, with adjacent heath and ponds. Species include Marsh Fragrant, Common Spotted and
Southern Marsh Orchids. An array of hybrids can
usually be found too.
West of Diss on the B1111, off the minor road between
Hopton and Coney Beeston Road. (Suffolk WT)
441
ESSEx
28. West Wood
TL 624332
An ancient woodland on chalky boulder clay, with
a rich ground flora that includes Early Purple and
Greater Butterfly Orchids.
Midway between Thaxted and Great Sampford on the
B1051. Park beside the road. (Essex WT)
29. Hatfield Forest
This Suffolk WT reserve protects one of the few

remaining British sites for Military Orchid. The Trust
opens the reserve to the public on 1-2 days each
year in late May or early June, to allow viewing of
this very special orchid. Contact the Trust for details.
(Suffolk WT)
TL 547202
One of the last medieval royal forests to retain
much of its original character and composition. The
site contains mixed habitats of ancient woodland,
pasture, glades and marshy ground, separated by
ditches and banks. The ground flora includes Violet
Helleborine and Birds-nest, Bee and Pyramidal Orchids in the woods and glades, and Early Marsh and
Common Spotted Orchids in the fens.
Park off the A120 at Takely near the M11 Junction 8.
(NNR, NT)
24. Bradfield Woods
30. Roding Valley Meadows
23. Rex Graham Reserve
TL 935581
An ancient wood, one of the richest in the country,
comprising coppice-with-standards. Parts are known
to have been traditionally managed since 1252.
There is a diverse ground flora, and Early Purple
Orchid can be found in spring, with Green-flowered
Helleborine elusive in mid-summer.
On the road between Bradfield St George and Felsham,
12 miles southeast of Bury St Edmunds. (NNR, Suffolk
WT)
25. Chippenhall Green

TM 285757
A very impressive colony of Green-winged Orchids
grows on this area of common land enclosed by
cattle grids.
Southeast of Fressingfield on the B1116. Take the minor road to Chippenhall Green at Rackhams Corner.
26. Wolves Wood
TM 054440
An ancient woodland with hazel and hawthorn coppice and scrub. The wide rides have a good ground
flora with several species of orchid, including Violet
Helleborine under the coppice.
On the A1071, 2 miles east of Hadleigh and 8 miles
west of Ipswich. (RSPB)
27. Groton Wood
TL 976428
A stand of ancient woodland with old lime coppice,
traditionally managed. The rides have an interesting
flora with Early Purple Orchid and Violet Helleborine, with a particularly good colony of helleborines
under the old hazel coppice on the west side of
the wood.
North of the A1071; along a maze of lanes 3 miles
west of Hadleigh, between Castlings Heath and Kersey
Tye. (Suffolk WT)
TO 430943
A traditionally managed river valley with hay meadows, scrub and woodland beside the River Roding.
The wet meadows hold many Southern Marsh
Orchids.
Park next to the David Lloyd Centre, Roding Lane,
Chigwell. There are other footpaths on the Loughton
side of the River. (Essex WT)
31. Langdon
TQ 659874
An unusual site with much social history. The abandoned plotlands are patchworks of old gardens and
grassland bordered by hedgerows and orchards.
Green-winged Orchids can be seen in their thousands in the Lincewood area, and Common Spotted, Green-winged, Pyramidal and Bee Orchids can
be found in the area of Marks Hill.
West of Basildon off the B1036. There is a car park
and Visitor Centre. (Essex WT)
32. Grays Chalk Quarry
TQ 611787
A long-disused quarry now with woodland, a small
area of chalk grassland and a pond. There are many
orchid species present, with good numbers of Man,
Pyramidal and Bee Orchids.
Park in the public car park on London Road near Grays
town centre. The entrance to the site is opposite, via a
short path. (Essex WT)
29/1/09 12:49:28
442
SITE GUIDE
Central England
12
13
14
8
1
7
3
16
2
19
11
6
9
21
17
10
20
18
28
29
27
31
26
30
22
36
25
35
39
40
37
38
23
15
42
46
44
45
43
41
24
34
32
33
Cheshire
1 Witton Lime Beds
Staffordshire
2 Rod Wood
3 Coombes Valley
Derbyshire
4 Hilton Nature Reserve
5 Rose End Meadows
6 Derbyshire Dales
7 Deep Dale
8 Millers Dale Quarry
nottinghamshire
9 Bentinck Banks
10 Wilwell Farm Cutting
11 Eaton Wood
Lincolnshire
12 Messingham Sand Quarry
13 Kingerby Beck Meadows
14 Donna Nook and
Saltfleetby Dunes
15 Candlesby Hill Quarry
16 Heaths Meadows
17 Ancaster Valley
18 Moulton Marsh
Shropshire
19 Wem Moss
20 Granville Country Park
21 Llynclys Common
22 Wenlock Edge
Warwickshire
23 Snitterfield Bushes
CHESHIRE
1. Witton Lime Beds
SJ 660746
These lime beds were produced by the chemical
and salt industries near Northwich. The Astons
Flash area is notable for the magnificent show of
Marsh Fragrant Orchids. More than 10,000 spikes
have been recorded on the site in some years.
Marsh Helleborine also occurs.
24 Ufton Fields
25 Draycote Meadow
Leicestershire
26 Lea Meadows
27 Cloud Wood
28 Muston Meadows
29 Cribbs Meadow
30 Great Merrible Wood
Rutland
31 Priors Coppice
Herefordshire
32 Davies Meadows
33 The Doward Reserves
35
36
37
38
39
40
Monkwood
Wyre Forest
Tiddesley Wood
Windmill Hill
Trench Wood
Eades Meadow
northamptonshire
41 Stoke Wood End Quarter
42 Glapthorn Cow Pastures
43 Short Wood
44 Collyweston Quarries
45 Bedford Purlieus
46 Barnack Hills and Holes
Worcestershire
34 The Knapp and Papermill
Northeast of Northwich on the far side of the Trent &

Mersey Canal. Access is from the minor road west of the
Flash. Park just before the village of Marston on the B5075.
STaFFORDSHIRE
2. Rod Wood
SJ 997531
With some fine meadows, woodland and marsh, the
interesting flora here includes Common Twayblade
and Early Purple Orchid.
29/1/09 12:49:29
CENTRAL ENGLAND
Near Leek. Accessed from the A523 Leek to Ashbourne

road; go past the RSPB Coombes Valley reserve (Site 3
below) to the top of the hill. Park past the cattle grid.
(Staffordshire WT)
3. Coombes Valley
SK 005525
A steep valley among old woodland with the
Coombes Brook running the length of the reserve.
There is some good flower-rich grassland here, with
Common Spotted and Greater Butterfly Orchids.
Southeast of Leek; take the A523 Leek to Ashbourne
road and go along the minor road signposted to
Apesford. (RSPB, Staffordshire WT)
DERBYSHIRE
4. Hilton nature Reserve
SK 249315
With lakes, pools, woodland and grassland, there are
stunning displays of Southern Marsh Orchids here.
Also several thousand Common Twayblades.
West of Derby, off the A516 to Willowpit Lane, north of
Hilton. (Derbyshire WT)
5. Rose End Meadows

SK 293567
A series of traditionally managed meadows, wetland
and woodland with a great variety of wildflowers. Species include Pyramidal, Common Fragrant and Bee Orchids, and there are many Common Spotted Orchids.
Above Cromford between the A5012 and the B5036
Cromford to Wirksworth road. Park in Cromford village.
(Derbyshire WT)
6. Derbyshire Dales
Five valleys make up the Derbyshire Dales NNR.
These are two of the best for orchids.
Lathkill Dale (SK 190658) is one of the finest,
with stunning scenery and woodland, grassland
and evidence of the lead-mining industry. Common
Twayblade, Common Spotted Orchid and Common Fragrant Orchid can all be found among a very
rich flora, with thousands of Early Purple Orchids on
the grassy slopes.
Southwest of Bakewell, with numerous footpaths
across the dale. There is parking at Monyash on the
B5055 and at Over Haddon on minor roads from
Bakewell.
Cressbrook Dale (SK 170731) is a beautiful place,
with Early Purple, Common Spotted and Common
Fragrant Orchids, and a few Dark-red Helleborines
and Common Twayblades.
Northwest of Bakewell between Cressbrook and Litton.
Park at Monsal Dale off the B6465 or at Tideswell
Dale off the B6049.
7. Deep Dale
SK 165698
Fantastic upland limestone grassland, with stunning
displays of Early Purple Orchids in spring. Other
orchids to be found include Common Twayblade.
443
Turn off the A6 towards Sheldon between Taddington

and Ashford-in-the-water. Access is along public footpaths. (Plantlife)
8. Millers Dale Quarry

SK 140731
One of the Wye Valley reserves, which also include
Chee Dale (SK 120727), and Priestcliffe Lees (SK
147730). They have some excellent habitats, and a
range of orchids can be found. These include Early
Purple, Common Fragrant and Common Spotted
Orchids, and Common Twayblade.
East of Buxton, with parking at Millers Dale off the
B6049 or at Monsal Dale Station. (Derbyshire WT)
nOTTInGHaMSHIRE
9. Bentinck Banks
SK 493550
A triangle of disused railway lines; the embankments
have some good limestone grassland, with Common
Fragrant Orchid, Frog Orchid, Common Twayblade
and Common Spotted Orchid.
Just south of Mansfield, off the B6018 between Kirkby
in Ashfield and the minor road to Nuncargate. Park on
the verge. (Nottinghamshire WT)
10. Wilwell Farm Cutting
SK 568352
One of the best wildflower sites in Nottinghamshire,
with grassland, acid fen and scrub on an old industrial site. There is a good display of Green-winged
Orchids, as well as Bee, Common Spotted, and
Southern Marsh Orchids and Common Twayblade.
Vast hybrid swarms of orchids also occur.
Just off the B680 between Ruddington and Wilford.
Parking is available beside the reserve. (Nottinghamshire WT)
11. Eaton Wood
SK 727772
Ancient pasture woodland with a visible ridge and
furrow system that holds several species, including
Greater Butterfly, Early Purple and Common Spotted
Orchids, and perhaps Broad-leaved Helleborine.
Off the minor road from Upton to Eaton at East Retford,
opposite Gamston Wood. (Nottinghamshire WT)
LInCOLnSHIRE
12. Messingham Sand Quarry
SE 908032
Old flooded lagoons with fringing vegetation and
remnant heath, with woodland, grassland and marsh.
Several orchid species occur, including Pyramidal,
Common Spotted, Early Purple and Bee Orchids.
South of Scunthorpe at Messingham, on the A159. Access is off the B1400 opposite Scallow Grove Farm. A
track leads to a small car park. (Lincolnshire WT)
13. Kingerby Beck Meadows
TF 051941
A series of herb-rich meadows with mixed boundary hedges. Orchids present include a colony of the
diminutive Frog Orchid.
29/1/09 12:49:29
444
SITE GUIDE
Northwest of Lincoln near Market Rasen.The meadows

lie between Kingerby Beck and North Owersby; access
from North Owersby on the minor road to Glentham.
Park near the entrance. (Lincolnshire WT)
14. Donna Nook and SaltfleetbyTheddlethorpe Dunes

TF 467917
A large area of coastal saltmarsh and sand dunes, with

Bee and Pyramidal Orchids and Marsh Helleborine
in the dune slacks. The area of freshwater marsh can
have fantastic displays of Early and Southern Marsh
Orchids and their hybrids.The Saltfleetby reserve occupies the coast between Mablethorpe North End in
the south and Saltfleet Haven in the north.
Access to Donna Nook is from North Somercotes,
which is on the A1031 coast road; park at Stonebridge.
Do not enter the danger area when the ranges are
in use. Saltfleetby is easily accessed from several car
parks off the A1031. (NNR, NE, Lincolnshire WT)
15. Candlesby Hill Quarry
TF 460682
An old chalk and lime pit on the eastern edge of the
Lincolnshire Wolds. There is wood and scrub, and
newly colonised chalk grassland with Bee and Common Spotted Orchids, among others.
Northwest of Skegness off the A1028 Skegness to
Louth road, just north of Gunby Corner roundabout.
Park on the verge. (Lincolnshire WT)
16. Heaths Meadows
TF 484640
Old meadows divided into small fields surrounded by
large hedgerows.The meadows are traditionally managed and support a good grassland flora, including
Green-winged and Common Spotted Orchids.
West of Skegness on the A158, just southwest of
Burgh-le-Marsh at Bratoft Ings. Access is from Ings
Lane. Park on the roadside. (Lincolnshire WT)
17. ancaster Valley

SK 984434
A narrow, steep-sided valley with limestone grassland,
scrub and woodland. Bee and Common Fragrant
Orchids can be found here. Early Purple Orchid occur in the beech woodland.
On the south side of the A153 Sleaford to Grantham
road near Ancaster. Access is via the bridleway signposted east of the Ancaster crossroads. Park in the village. (Lincolnshire WT)
18. Moulton Marsh
SK 344336
The bank on the south side of the River Welland has
a fantastic display of Pyramidal Orchids, with over
4,000 spikes.
Off the A17 south of Fosdyke Bank, on a minor road
that leads on to the river. There is a car park. (Lincolnshire WT)
SHROPSHIRE
19. Wem Moss
SJ 472342
One of Shropshires lowland mires, with open water,
carr, woodland and a fen community that includes

Early Marsh and Lesser Butterfly Orchids.
Between Wem and Ellesmere, near Northwood on the
B5063. Park on the minor road to Dobsons Bridge just
west of Northwood. (Shropshire WT, NNR)
20. Granville Country Park

SJ 719124
Based around reclaimed mine workings near Telford, this site contains woodland and old grassy pit
mounds; the meadows have vast numbers of Southern Marsh Orchids and hybrids.
Just outside Telford, on Granville Road, Oakengates,
Wellington. (Shropshire WT, Telford & The Wrekin
Council)
21. Llynclys Common
SJ 273238
A mixture of habitats: woods, meadows, old quarries and scrub, where up to 12 orchid species have
been recorded. These include Common Spotted,
Early Purple, Pyramidal, Greater Butterfly and Bee
South of Oswestry on the A495. Park in the lay-by west
of Llynclys crossroads opposite Dolgoch. (Shropshire
WT).
22. Wenlock Edge
SO 595988
A magnificent limestone escarpment running from
Much Wenlock to Craven Arms, with habitats ranging from scrub woodland and grassland to marsh.
The pockets of limestone grassland have good displays of Pyramidal and Greater Butterfly Orchids,
along with other limestone species.
Harton Hollow Woods (SO 481878), on minor
roads off the B4368, is an excellent place to start a
walk along the escarpment. The National Trust has a
car park here. (NT, Shropshire WT)
WaRWICKSHIRE
There are several interesting Roadside Nature Reserves in the county, notably on the A429 Ettington
bypass, which has a spectacular display of Bee and
Pyramidal Orchids.
23. Snitterfield Bushes

SP 200603
One of the best ancient woodlands in the county,
the ash and birch woods have a good shrub layer
and ground flora, with Early Purple, Greater Butterfly and Common Spotted Orchids, Common Twayblade, and Broad-leaved Helleborine.
Near Stratford-on-Avon, on the Bearley Road out of
Snitterfield; the reserve is on both sides of the road,
with a car park. (Warwickshire WT)
24. ufton Fields
SP 378615
Old limestone workings with scrub, grassland and
ponds and a good range of orchid species, with Common Twayblade, Greater Butterfly, Common Spotted
and Bee Orchids.
29/1/09 12:49:29
CENTRAL ENGLAND
Between Royal Leamington Spa and Southam, off the

A425 at Ufton. There is a car park available. (Warwickshire WT)
25. Draycote Meadow

SP 448706
Warwickshires best meadow, with thousands of
Green-winged Orchids among other specialities.
Southwest from Rugby, with access off the B4453 at
Draycote village. A public bridleway gives access to the
meadow. (Warwickshire WT)
LEICESTERSHIRE
26. Lea Meadows
SK 506115
Unimproved meadows beside a clear stream. Signs
of ridge and furrow agriculture betray medieval usage, but the fields are now traditionally managed.
They contain a rich flora that includes great displays
of Common Spotted and Heath Spotted Orchids.
Northwest of Leicester; take the minor road to Newton
Linford from the A46. The site is on Ulverscroft Lane.
(LRWT)
27. Cloud Wood
SK 899188
A traditionally coppiced ancient woodland with good
rides and glades. Several orchids can be found including Bee, Birds-nest, Common Spotted and Greater
Butterfly Orchids and Broad-leaved Helleborine.
Northeast of Ashby de la Zouch on the minor road
linking Griffydam on the B5324 and Tonge on the
A453. Park in the lay-by. (LRWT)
28. Muston Meadows
SK 824367
One of the best lowland meadows in England with
unimproved ridge and furrow grassland. Among its
rich flora is a colony of more than 10,000 Greenwinged Orchids.
West of Grantham off the A52 at Muston. Park off the
minor road south to Woolsthorpe. (LRWT, NNR)
29. Cribbs Meadow
SK 899188
Ancient grassland, ponds and a disused railway line.The
fields lie on boulder clay and support a rich flora, including Green-winged and Common Spotted Orchids.
Between Wymondham and South Witham on the minor road between Sewstern and Thistleton. Park on the
road verge. (LRWT, NNR)
30. Great Merrible Wood
SP 834962
A mixed woodland with a very ancient history. With
ash, oak, and a varied shrub layer, the ground flora
includes Broad-leaved and Violet Helleborines.
Southwest of Uppingham off the Horninghold to Great
Easton road, southeast of the crossroads with the
B664.There is roadside parking. (LRWT)
RuTLanD
31. Priors Coppice
SK 834052
An ancient woodland on a steep slope. Under the
445
ash-maple and ash-wych elm coppice is a rich flora,

with a large colony of Early Purple Orchids and
Broad-leaved and Violet Helleborines; the often-wet
rides contain Common Spotted Orchids.
Southwest of Oakham, on minor roads connecting
Braunston to Leighfield. Park on the road to Leighfield
Lodge. (LRWT).
HEREFORDSHIRE
32. Davies Meadows
SO 375485
With three unimproved meadows, an ancient orchard, marshland and old hedgerows, this is an
enchanting place to visit. The meadows contain
Green-winged and Common Spotted Orchids.
On the A480 from Hereford to Lyonshall. Entrance
is just past the Three Horseshoes pub at Eccles
Green; park on the road verge. (Herefordshire WT,
Plantlife)
33. The Doward Reserves
SO 549161
Several ancient oak and beech woodland reserves
with areas of unimproved limestone grassland. They
skirt the Doward, a rock outcrop in a bend of the
River Wye. At Woodside (SO 555147), the limestone grassland has a good flora with Early Purple
and Greater Butterfly Orchids. Leeping Stocks (SO
548162) has White and Broad-leaved Helleborines
and Greater Butterfly Orchid, while White Rocks
(SO 550158) has Bee Orchid.
South of Ross-on-Wye on the A40 to Whitchurch, from
where the reserves are signposted. (Herefordshire WT)
WORCESTERSHIRE
34. The Knapp and Papermill
SO 751522
One of the finest nature reserves in the Midlands, this
is a mixture of old meadows, woods and orchards beside the Leigh Brook. Species include Green-winged,
Early Purple, Greater Butterfly and Common Spotted
Southwest of Worcester on the A4103. Follow minor
roads from Bransford along the BransfordSmith End
GreenAlfrick Pound road. The reserve entrance is on
the left after about 3 miles, where the road crosses the
Leigh Brook. There is limited parking near the bridge.
(Worcestershire WT)
35. Monkwood
SO 804606
A coppiced woodland important for butterflies
and moths, but also supporting many Early Purple
Orchids, some Common Spotted and Greater Butterfly Orchids, and a few, hard-to-find Broad-leaved
and Violet Helleborines.
Northwest of Worcester; near Grimley, on the minor
road to Monkwood Green from the A443. (Worcestershire WT, Butterfly Conservation)
29/1/09 12:49:30
446
SITE GUIDE
36. Wyre Forest
SO 743740
The Wyre forest covers over 6,000 acres and contains some of the best stands of ancient woodland
in Britain (it is the largest continuous area of ancient
woodland in England). Although predominantly Sessile Oak, just over half of the forest consists of conifer
plantations. The Dowles Brook runs through one of
the richest and most diverse areas of the forest, including meadows and abandoned orchards. Eleven species
of orchid have been recorded, including Sword-leaved
Helleborine, although there are just a few scattered
populations of the latter that are hard to find. Common Spotted Orchid is abundant in some rides, and
other species include Common Fragrant Orchid and
Green-winged Orchid in some of the meadows (and
perhaps also Heath Fragrant Orchid).
West of Bewdley on the A456 Ludlow road, with parking and a Visitor Centre at the FCs Callow Hill, which
gives access to the North Worcestershire path; the
New Parks Beech Wood area is a good place to start.
Access is only possible on open days contact the

Worcestershire WT. (Worcestershire WT)
37. Tiddesley Wood
43. Short Wood
SO 929462
An ancient mixed woodland near Pershore with a
good shrub layer. Common Spotted Orchid is widespread; there are also Early Purple and Greater Butterfly Orchids and Common Twayblade, with Bee
Orchids in the open grassland; Birds-nest Orchid
and Violet Helleborine are also present but can be
hard to find.
Next to the minor road signposted to Besford and
Croome from Pershore on the A44. Park beside the
wood. (Worcestershire WT)
38. Windmill Hill

SP 072477
Windmill Hill has a small area of limestone grassland
and scrub. There is a spectacular display of Pyramidal Orchids as well as Common Spotted Orchid,
Common Twayblade and small numbers of Bee and
Greater Butterfly Orchids.
East of Evesham on the south side of the B4510, close
to the Fish and Anchor Inn and just before the brow
of the hill; park on the roadside opposite the entrance.
(Worcestershire WT)
39. Trench Wood
SO 930589
An ancient woodland, with the wood managed
partly as high forest and partly as scrub or coppice.
Orchids present include Common Spotted, with
smaller numbers of Pyramidal and Greater Butterfly
Orchids.
Southeast of Droitwich on the Shernal Green to Sale Green
minor road, just west of the M5. (Worcestershire WT)
40. Eades Meadow
A traditionally managed meadow on heavy clay soils,

Eades Meadow has a spectacular display of Greenwinged Orchids in the spring. Common Spotted Orchid,
Common Twayblade, Greater Butterfly Orchid and a
few Common Fragrant Orchids are also present.
nORTHaMPTOnSHIRE
41. Stoke Wood End Quarter
SP 800861
This ancient oak and ash woodland has an excellent
flora on the mainly calcareous clay soils. Orchids
include Early Purple Orchid, Broad-leaved Helleborine, Common Twayblade and Greater Butterfly
Orchid.
North of Desborough on the B669 to Stoke Albany.
Park in the lay-by. (BCNPWT, WT)
42. Glapthorn Cow Pastures
TL 005903
A scrubbed-over pasture, the resulting thickets are a
haven for wildlife. The area has a rich flora, including
Early Purple and Common Spotted Orchids.
Access from the minor road halfway between Glapthorn and Benefield, north of Oundle. Park on the road
verge. (BCNPWT)
TL 015913
Rich deciduous woodland and coppice on boulderclay (said to be the finest Bluebell wood in the county), with Early Purple, Birds-nest, Greater Butterfly
and Common Spotted Orchids, Common Twayblade,
and Broad-leaved and Violet Helleborines.
Northwest from Oundle to Southwick on a minor road.
Access is along a signposted footpath on this road.
(BCNPWT)
44. Collyweston Quarries
TF 004038
A series of old grassed-over quarries, the open-cast
pits and mines have created a hills and holes appearance, and a rich flora has developed. Common Twayblade, Man, Bee, and Common Fragrant Orchids are
present, while Pyramidal Orchid is abundant.
South of Stamford, on the A43 between Easton-on-thehill and Collyweston. (BCNPWT)
45. Bedford Purlieus
TL 034997
A large, remarkably diverse woodland reserve,
which many botanists consider to be the richest of
all British woods more than 400 species of vascular plants have been recorded on the site. Fly and
Greater Butterfly Orchids can be found here, as can
large numbers of Common Spotted Orchids.
North of Oundle on the A47; park on the farm track between Wansford and the Kings Cliffe road. (FC, NNR)
46. Barnack Hills and Holes
TF 075045
A superb area of calcareous grassland on longabandoned medieval limestone workings. There is
an important Man Orchid colony (more than 1,000
spikes) as well as Early Purple, Pyramidal, Bee, Common Fragrant and Common Spotted Orchids.
East of Stamford; the reserve is off the B1443 on minor roads at the western end of Barnack Village, with
several car parks around the perimeter. (NNR)
29/1/09 12:49:30
NORTHERN ENGLAND
447
northern England and Isle of Man

1
4
3
32
5
6
33
34
12
35
36
11 9
10 14
13
30
31
29
28
27
26
17
25
18
24
19 20
23
15
northumberland
1 Holy Island
2 Beltingham River Gravels
3 Williamston Reserve
Cumbria
4 Geltsdale
5 Augill Pasture
6 Little Asby Inrakes and Outrakes
7 Smardale Gill
8 Waitby Greenriggs
9 Hutton Roof Crags
10 Arnside Knott
11 Latterbarrow
12 Whitbarrow
13 Sandscale Hawes
Lancashire
14 Gait Barrows
15 Ainsdale Sand Dunes
16 Nob End
Yorkshire
17 Southerscales
16
21
22
18 Brae Pasture
19 Malham Tarn Estate
20 Grass Wood
21 Sherburn Willows
22 Brockadale
23 Skipwith Common
24 Wharram Quarry
25 Flamborough Cliffs
County Durham
26 Coatham Marsh
27 Castle Eden Dene
28 Hawthorn Dene
29 Wingate Quarry
30 Raisby Hill Grassland
31 Bishop Middleham Quarry
32 Upper Teesdale
Isle of Man
33 Ayres Visitor Centre
34 Moaney and Crawyns Meadows
35 Close Sartfield
36 Dalby Mountain
29/1/09 12:49:30
448
SITE GUIDE
nORTHuMBERLanD
1. Holy Island
nu 100432
The island is around three miles long, and sand
dunes extend along much of the north shore. The
dunes and dune slacks form an excellent orchid
habitat, especially at the Snook at the western end
of the island. The great speciality is Lindisfarne Helleborine, formerly treated as a subspecies of Dune
Helleborine (which it greatly resembles). Following
recent genetic research, the Lindisfarne Helleborine
is now treated as a distinct species endemic to the
island. The population is small (150-300 plants) and
scattered, and it is advisable to contact the warden
for advice; tel. 01289 381470. Other notable species
include Coralroot Orchid in very small numbers in
the dune slacks, Marsh Helleborine, Northern and
Early Marsh Orchids and Common Spotted Orchid.
The road to the island is signposted off the A1 at West
Mains Inn. It is impossible to cross the causeway for
at least 2 hours either side of high water, and the
strength of the wind and the height of the tide must
also be taken into account.Tide tables are displayed on
the causeway, which is a tarmac road.
2. Beltingham River Gravels
nY 785640
Grassland polluted with heavy metals, woodland
and scrub where Tyne Helleborine, the inland form
of Dune Helleborine, can be found.
Off the A69 south of Bardon Mill, on the minor road
from Beltingham to Willimontswick. There is roadside
parking and a walk along the river to the reserve.
(Northumberland WT)
3. Williamston Reserve
nY 681521
This small reserve on the eastern bank of the River
South Tyne comprises birch and willow woodland;
this has grown on gravel and silt contaminated by
tailings from lead and zinc mines. The reserve is notable for its large colony of Tyne Helleborine, the
inland form of Dune Helleborine. Common Spotted Orchid is also present.
Off the A689 Alston to Brampton road, south of Slaggyford village, signposted to Barhough Hall. Park by the
road; the entrance to the reserve is via a stile immediately over the river bridge. (Northumberland WT)
CuMBRIa
4. Geltsdale
nY 561557
Twelve-thousand acres of upland heather moorland,
working farms, old coal and limestone workings and
woodland, along the River Gelt. There are some
very rich habitats where several species of orchid
can be found, including Greater Butterfly Orchid
and Common Fragrant Orchid.
East of Carlisle, south of the A69 off the B6413 at

Castle Carrock. Park in the RSPB car park. (RSPB)
5. augill Pasture
nY 817147
Species-rich grassland and some woodland with
Common Fragrant and Greater Butterfly Orchids,
among others.
Just east of Brough off the A66 to Augill House Farm.
Park at the end of the road. (Cumbria WT)
6. Little asby Inrakes and Outrakes
nY 699101
A small area of rough grazing notable for a colony of
Small White Orchids, as well as Northern and Early
Marsh Orchids, Common Spotted Orchid, Early
Purple Orchid and Heath Fragrant Orchid.
West of Kirkby Stephen, which lies on the A685. The
area lies north of the minor road immediately north
of Little Asby.
7. Smardale Gill
nY 727070
A disused railway line with a rich orchid flora; Fly and
Frog Orchids can be found in small numbers, along
with Greater Butterfly, Common Spotted, Northern
Marsh and Common Fragrant Orchids, Common
Twayblade, and Marsh Helleborine. A few Lesser
Butterfly Orchids are also here but hard to find.
West of Kirkby Stephen, off the A685. Access is from
the south side of the minor road running from Kirkby
Stephen to Crosby Garrett. (Cumbria WT, NNR)
8. Waitby Greenriggs
nY 760085
An old railway line running down a beautiful valley,
with woodland and herb-rich meadows. The rich
flora includes Marsh Helleborine, all three fragrant
orchids, Fly, Frog, Northern Marsh and Common
Spotted Orchids, and Common Twayblade.
East of Ravenstonedale at Waitby, off the A685. Park
close to the railway bridge. (Cumbria WT)
9. Hutton Roof Crags
SD 543783
A large area of woodland, scrub, heath and fantastic limestone pavement, with great views. Dark-red
Helleborine can be found amongst the grikes, and
Fly Orchids grow in the grassland.
Off the A6070, between Burton and Hutton Roof. Park
on the roadside or walk from Hutton Roof, Burton or
Holme villages. (Cumbria WT)
10. arnside Knott
SD 456775
Woodland, scrub and limestone pavement sloping
down to saltmarsh, with great views. Several species of orchid are present, including Early Purple, Fly
and Lesser Butterfly Orchids and Broad-leaved and
Dark-red Helleborines.
On the south side of the Kent estuary. Park in the
National Trust car park signposted from Arnside village,
which is at the end of the B5282. (NT)
29/1/09 12:49:31
NORTHERN ENGLAND
11. Latterbarrow
SD 441827
Excellent limestone grassland, scrub and woodland
with Early Purple and Greater Butterfly Orchids.
Near Grange-over-Sands just off the A590 at Longhowe End. Park beside the road just past the Derby
Arms. (Cumbria WT)
12. Whitbarrow
SD 436859
Largely limestone, Whitbarrow Scar is just over 650
feet high. The reserve sits on top; it consists of limestone pavement, grassland, juniper scrub and woodland. It is an interesting place to visit, and Dark-red
Helleborine and Fly Orchid can be found here.
Southwest of Kendal off the A5074. There is a small
car park at the Kennels at Witherslack. Access is via
steep and rocky paths. (Cumbria WT, FC, NNR)
13. Sandscale Hawes
SD 200756
This extensive dune system holds 10 species of orchid, with specialities including the largest population of Coralroot Orchid in England (3,000 were
counted across five slacks in 1991), Dune and
Green-flowered Helleborines (often more than
1,000 spikes of each), as well as Marsh Helleborine,
Northern Marsh Orchid and Bee and Pyramidal
Orchids.
North off the A590 to the northwest of Dalton-inFurness, on the minor road to Roanhead. Park at the
beach car park and walk west along the beach and
then into the dunes (the first blow-out after 550 yards
holds a few Coralroot Orchids, but the farther slacks
are better). (NT, NNR)
LanCaSHIRE
14. Gait Barrows
SD 483775
Some of the finest limestone pavement and Yew
woodland in the country. Dark-red and Broadleaved Helleborines, and Common Fragrant, Common Spotted and Northern Marsh Orchids are
important parts of a fascinating flora.
Gait Barrows is just in Lancashire, east of Silverdale and
close to Arnside Knott. Off the minor road linking Carnforth to Arnside, with roadside parking. (NNR)
15. ainsdale Sand Dunes
SD 397129
Part of the huge Sefton Coast wildlife area, the speciality here is Dune Helleborine, which is common,
as well as Green-flowered Helleborine (both these
helleborines can also be found in the pine plantations to the rear of the dunes). Other orchids include
Marsh Helleborine, Bee and Pyramidal Orchids and
Early Marsh Orchid (both pink-flowered plants and
the red-flowered subspecies coccinea). Northern
and Southern Marsh Orchids also occur here.
There is open access to the dunes, which stretch from
Southport to Formby, with plenty of car parks. Ainsdale
beach car park is a useful start. (NNR)
449
16. nob End

SD 743070
An unusual site made up entirely of alkaline industrial waste, generated through soda crystal manufacture in the 19th century. Its weathered surface
now has a unique flora, with eight species of orchids.
Common Spotted Orchid and Early Marsh Orchid
(subspecies coccinea) are common. Northern and
Southern Marsh Orchids occur, and Common
Fragrant Orchid and Common Twayblade are also
common. Harder to find are Marsh Helleborine and
Bee Orchid. The marsh orchid hybridisation at this
site is stunning.
Just off the A6053 west of Little Lever, near Bolton.
Access is on foot, with parking near Rock Hall Visitor
Centre in Moses Gate Country Park.
YORKSHIRE
17. Southerscales
SD 742769
Part of the much larger Ingleborough NNR, this reserve has good limestone pavement with its clints
and grikes, and acidic and limestone grassland. Early
Purple Orchid, Dark-red Helleborine and Common
Twayblade are present.
Off the B6255 Ingleton to Hawes road in North Yorkshire. Park at the Old Hill Inn or on the roadside at
Chapel-le-Dale. (Yorkshire WT, NNR)
18. Brae Pasture
SD 790741
A small area of limestone pasture and damaged
limestone pavement that has a very rich flora; Early
Purple and Frog Orchids can be found here.
On the B6479 between Horton-in-Ribblesdale and
Selside. Park on this road. A Public Footpath sign points
to the reserve.
19. Malham Tarn Estate
SD 890660
Upland hay meadows and limestone grassland, limestone pavements and the NNR at Malham Tarn make
up this wonderful landscape. The variety of habitats
have Dark-red Helleborine and, in the wetter areas,
Early, Northern and Pugsleys Marsh Orchids.
Northwest of Skipton on minor roads from the A65;
the estate is north from Malham village, where there is
a Visitor Centre, parking and footpaths. (NT, NNR)
20. Grass Wood
SD 983652
A wooded limestone hillside that slopes down to
the River Wharfe. The wood has formed on a series
of limestone terraces, and has rocky outcrops and
open areas. Early Purple and Birds-nest Orchids can
be found, among others.
In Wharfedale, north of Grassington alongside the
Grassington to Coniston minor road. There is a small
car park. (Yorkshire WT)
29/1/09 12:49:31
450
SITE GUIDE
21. Sherburn Willows

SE 487326
An interesting mix of habitats, with magnesian limestone grassland, scrub, and fen to the side of a small
stream. Common Spotted and Bee Orchids and
Common Twayblade can be seen.
West of Selby; access along Mill Dyke, off the B1222
between Sherburn-in-Elmet and South Milford. Park on
New Lane or walk from South Milford Station. (Yorkshire WT)
22. Brockadale and Thompson Meadow
SE 513174
Brockadale is a wooded valley with some old quarry
workings and traditionally managed meadows. One
of these, Thompson Meadow, is on a sloping hillside
beside the River Went. The rich limestone flora includes Common Spotted, Common Fragrant and
Early Purple Orchids.
Between Wentbridge and Kirk Smeaton, off the B6474
south of Pontefract, just east of the A1.The reserve car
park is on an unmarked track out of Little Smeaton.
(Yorkshire WT, Plantlife)
23. Skipwith Common

SE 669378
A large area of heath, marsh and woodland with
Common Twayblade, Common Spotted Orchid and
Broad-leaved Helleborine.
Northeast of Selby, Skipwith is north off the A163 on
minor roads after North Duffield. Park and walk from
the village.
24. Wharram Quarry
SE 858653
A disused chalk quarry, with steep chalk slopes and
grassland. A typical chalk flora has developed, and
Pyramidal Orchid, Common Spotted Orchid and
abundant Bee Orchids can be seen.
Southeast of Malton close to Wharram-le-Street, on the
B1248 Malton to Wetwang road. Park on the roadside
close to the old railway line. (Yorkshire WT)
25. Flamborough Cliffs
Ta 240722
The spectacular chalk cliffs attract thousands of seabirds but also support a varied flora on the cliff-tops.
Pyramidal and Common Spotted Orchids occur
here and Northern Marsh Orchids can be found
in wet flushes.
East of Bridlington on the B1255 to Flamborough.
There is a car park at North Landing and access to
the clifftop footpaths. (Yorkshire WT)
COunTY DuRHaM
26. Coatham Marsh
nZ 586248
A wetland reserve of lakes, marsh and wet meadows.
The wide variety of wildflowers include Common
Fragrant Orchid, and good displays of Northern
Marsh and Bee Orchids.
Off the A1085 Redcar to Middlesbrough road. Park off

Tod Point Road or on the sea front on Manjuba Road.
(Tees Valley WT)
27. Castle Eden Dene

nZ 435397
This spectacular wooded gorge is on magnesian
limestone and boulder clay. Fly and Birds-nest Orchids can be found in the woods, and Common
Spotted and Common Fragrant Orchids occur in
the open grassland.
On the south side of Peterlee, east of the A19. Castle
Eden Dene is signposted from the A19 and from Peterlee town centre. Car parking is at the main entrance,
and a small car park is available at Oakerside Dene
Lodge. (NNR)
28. Hawthorn Dene and Meadow nZ 433457
and Beacon Hill
nZ 440455
A coastal site with extensive, steep-sided mixed
woodland on limestone and some excellent
grasslands with a great flora. Early Purple, Lesser
Butterfly and Birds-nest Orchids can be found
among the ground flora in the woods, with Bee,
Common Fragrant, Common Spotted and Northern Marsh Orchids in the pastures.
At Hawthorn off the B1432, between Easington and
Seaham on the Durham coast. The site is at the end
of a minor road signposted Quarry Traffic. Park in the
lay-by opposite the cottage. Access to Beacon Hill is
along the Coastal Footpath or through the southern
end of Hawthorn Dene. (Durham WT, NT).
29. Wingate Quarry

nZ 373375
An important example of magnesian limestone
grassland one of Britains rarest habitats. There are
large numbers of Common Fragrant Orchids here.
Northwest of Hartlepool, off the A181 west of Wingate.There is a car park. (Durham WT)
30. Raisby Hill Grassland
nZ 337375
Magnesian limestone grassland with fen and pools.
There is a small disused quarry which has an important population of Dark-red Helleborines.
Off the A177 from Middlesbrough at Coxhoe Quarry.
(Durham WT)
31. Bishop Middleham Quarry
nZ 330324
This large disused quarry on the Durham limestone
is superb. It holds the largest colony of Dark-red
Helleborines in Britain, with around 2,000 spikes
(perhaps more than all the other populations put
together). Other species include Pyramidal and Bee
Orchids, Common Spotted Orchid, Marsh Fragrant
Orchid and Common Twayblade.
Northwest of Middlesbrough, off the A177 west
through Bishop Middleham. The reserve is on a minor
road north of the village. Park in the lay-by opposite the
entrance. (Durham WT)
29/1/09 12:49:32
NORTHERN ENGLAND
451
32. upper Teesdale

nY 907282
UpperTeesdale is a botanists paradise, with the famous
reserves of Cronkley and Widdybank Fell above Cow
Green Reservoir on the sugar limestone. At lower
altitudes, wonderful flower meadows still exist along
the River Tees from Langdon Beck to Middleton-inTeesdale.There are many orchids in the meadows and
around the paths. Heath Spotted, Common Spotted
and Early Purple Orchids and Common Twayblade
can be found. As well as Early Marsh, Northern Marsh
and Lesser and Greater Butterfly Orchids, there can
be spectacular displays of hybrids.
North of Middleton-in-Teesdale on the B6277 there is
an excellent Durham WT Visitor Centre at Bowlees. Information on good walks in the area is available from
there. (NNR)
ISLE OF Man
33. ayres Visitor Centre
nx 435038
The shingle near the northern tip of the island supports an unusual lichen-heath community. Denseflowered Orchid grew here from 1966-1986 at its
only British site. The lichen heath lies behind the
dunes, and Pyramidal Orchid can be found here. In
the dune slacks there are Early Marsh and Northern
Marsh Orchids.
On the A10 Ballaghennie Road, west of Bride. There is
a Visitor Centre and car park. (Manx WT)
34. Moaney and Crawyns Meadows
SC 375957
Traditional hay meadows with a rich flora.The meadows are on the edge of the Ballaugh Curragh wetland. Heath Spotted Orchids occur in abundance.
Off the A14 from Sulby to Ballaugh Curragh. Park in
the lay-by. (Manx WT, Plantlife)
35. Close Sartfield

SC 358956
Part of the Ballaugh Curragh, this is the largest and
most important wetland on the Isle of Man. It is a
mixed area with bog, willow and birch woodland,
and rich hay meadows. There is an amazing display
of many thousands of orchids flowering from late
May, including Heath Spotted, Common Spotted,
Early Marsh and Northern Marsh Orchids, and
Common Twayblade.
On minor roads off the B9 between Ballaugh village
and Sulby Glen.The entrance and car park are located
about 25 yards along a track. (Manx WT).
36. Dalby Mountain
SC 233769
A traditional moorland with good displays of Heath
Spotted Orchids.
The A27 Dalby to Round Table road passes through
the reserve; there is parking on this road close to the
track to Eary Cushlin. (Manx WT)
29/1/09 12:49:32
452
SITE GUIDE
Wales
3
2
9
4
10
5
6
7
8
12
14
13
15
17
18
28
23
25
24
26
27
anGLESEY
1. newborough Warren
SH 415635
Lying at the southern tip of Anglesey, this is one of
the largest sand dune systems in Britain, covering
more than 3,000 acres. Although two-thirds of this
area has been planted with Corsican Pine, there
are still some fantastic dunes and dune slacks in the
treeless south-eastern section of the Warren. Dune
Helleborine (once again a British endemic) is a speciality, as are Northern and Early Marsh Orchids,
Common Twayblade and Marsh Helleborine.
Access is from the Menai Bridge along the A4080 to
Aberffraw. There are several car parks along this road
around Newborough, and footpaths crossing the dunes
to the foreshore. (NNR, CCW)
2. South Stack Cliffs

SH 207823, SH 218805
Situated three miles west of Holyhead on Holy
Island at the western extremity of Anglesey, this
area has some of the finest coastal cliff scenery in
Wales. Inland, there are areas of maritime heathland rising to 853 feet at Holyhead Mountain, with
a separate area of heath to the south at Penrhos
Glamorganshire
23 South Gower Cliffs
24 Oxwich Bay
25 Crymlyn Bog
26 Kenfig
27 Merthyr Mawr
Cardiganshire
11 Ynyslas
21
22
Gwynedd
5 Caeau Tan y Bwlch
6 Morfa Harlech
7 Morfa Dyffryn
8 Roman Steps
Wrexham
10 Marford Quarry
16
19
Pembrokeshire
19 Dowrog Common
20 St Davids Head
Conwy
9 Great Orme
11
20
anglesey
1 Newborough
Warren
2 South Stack Cliffs
3 Cemlyn
4 Cors Goch
29
30
31
Carmarthenshire
21 Allt Rhyd y Groes
22 Pembrey Burrows
Monmouthshire
28 Cwm Clydach
29 Pentwyn Farm
30 Springdale Farm
31 Newport Wetlands
Powys
12 Lake Vyrnwy
13 Dyfnant Meadows
14 Llanymynech
Rocks
15 Elan Estate
16 Nant Irfon
17 Burfa Bog
18 Pwll y Wrach
Feilw Common. Heath Spotted Orchid is common

on Holyhead Mountain and four species of orchid,
including Northern Marsh, can be found at Penrhos
Feilw Common.
West of Holyhead on minor roads via Llaingoch and
Twr to South Stack. For Penrhos Feilw Common (The
Range) follow minor roads south from South Stack, or
the B4545 from Dyffryn to Trearddur, and then take
the minor coast road through Penrhosfeilw, parking at
Gors-goch. (RSPB)
3. Cemlyn
SH 331932
On the north coast of Anglesey, a shingle storm
beach has sealed off the entrance to Cemlyn Bay
to form a saline lagoon. There are Autumn Ladystresses on the more stable parts of the beach.
Off the A5025 between Dyffryn and Cemaes, on the
minor road from Tregele. There are car parks at both
the eastern (Traeth Cemlyn) and western (near Bryn
Aber) ends of the shingle ridge. (NT, North Wales
WT)
4. Cors Goch
SH 504817
A large and diverse reserve with areas of fen, heath
and grassland. The orchids here include Early Purple,
29/1/09 12:49:32
WALES
Green-winged, Early Marsh and Northern Marsh

Orchids, Common Fragrant and Lesser Butterfly
Orchids, and Marsh Helleborine. There are several
splendid fens on Anglesey, but this is the only one
with open access and with footpaths and a boardwalk. For information and permits for the others
contact the CCW.
Cors Goch is off the A5025 turning to Llanbedrgoch.
Parking is available in the lay-by. (North Wales WT,
CCW)
GWYnEDD
5. Caeau Tan y Bwlch
SH 431488
A sloping calcareous grassland site with wet flushes;
more than 2,000 Greater Butterfly Orchids can be
seen in some years. Heath Spotted and Common
Spotted Orchids also occur.
Off the A499 Caernarfon to Pwllheli road, near Clynnog-fawr on the Lleyn Peninsula. (North Wales WT,
Plantlife)
6. Morfa Harlech
SH 555350
One of two major sand dune systems near Harlech,
Morfa Harlech has a rich flora. There are various
orchids present, including Early Marsh Orchid of the
red-flowered subspecies coccinea, and also Marsh
Helleborine.
Morfa Harlech can be reached by turning off the A496
to the public car park at Harlech and walking along
the beach. (NNR, CCW)
7. Morfa Dyffryn
SH 560250
The second of the dune systems between Harlech
and Barmouth, Morfa Dyffryn has extensive areas
of dune slack with Early Marsh Orchid of the subspecies coccinea, Marsh Helleborine and Greenflowered Helleborine.
Morfa Dyffyn can be reached from public car parks
at each end of the reserve, south of Llanbedr on the
A496. (NNR, CCW)
8. Roman Steps
SH 500775
This area of hill pasture and moorland is well known
for its population of Lesser Twayblades. Look for
these orchids in places where the footpath passes
through stands of heather.
From the B4573 at Harlech follow the minor roads to
Cwm Bychan (SH 647314). Then follow the Roman
Steps footpath for around 1 mile.
COnWY
9. Great Orme
SH 780832
Lying immediately north of Llandudno, this massive
limestone headland covers around two square miles
and rises to 675 feet, forming the eastern boundary
of Conwy Bay. Dark-red Helleborine grows on the
453
limestone pavement, with Autumn Ladys-tresses on

the short turf capping.
From Llandudno, Marine Drive encircles the head and
a minor road bisects the plateau, while a tramway also
climbs to the top.The whole area is a Country Park.
WRExHaM
10. Marford Quarry
SJ 357560
An old sand and gravel quarry, now grassed over
and developing an interesting flora, which includes
Common Spotted, Pyramidal and Bee Orchids.
North of Wrexham on the B5445, between Rossett
and Gresford west of the village of Marford. The reserve is on Springfield Lane; park beside the railway
bridge. (North Wales WT)
CaRDIGanSHIRE
11.Ynyslas
Sn 610941
Lying on the southern side of the Dyfi estuary, the
dunes here are excellent, with Marsh Helleborine,
Early Marsh Orchid (including the red-flowered
subspecies coccinea), Northern Marsh, Southern
Marsh, Pyramidal and Bee Orchids.
Leave the A487 from Machynlleth onto the B4353,
and turn north at Ynyslas onto the minor road to the
dunes. There is a car park on the sand beside the Information Centre. (NNR, CCW)
POWYS
12. Lake Vyrnwy
SJ 015191
The largest artificial lake in Wales, fringed by scrub
woodland and meadows with heather and grass
moorland on the slopes. There are large areas of
conifer plantation and deciduous woodland too.
Heath Spotted Orchid is locally common and Lesser Twayblade occurs, although it is hard to find; look
on damp north-facing slopes under heather.
Northwest of Welshpool, off the B4393 from Llanfyllin
to Llanwddyn, from where the road continues on to
circumnavigate the lake. The Information Centre is on
the minor road 100 yards south of the west end of the
dam. (RSPB)
13. Dyfnant Meadows
SH 998155
A typical traditionally managed upland pasture, comprising eight fields, with acidic bogs and flushes containing large numbers of Heath Spotted Orchids.
Near Dyfnant (in Dyfnant Forest), off the B4395 at
Hendre, then on minor roads and forest tracks. Park by
the reserve gates. (Montgomeryshire WT)
14. Llanymynech Rocks
SJ 267218
An outcrop of limestone, part of an old quarry with
some woodland and a rich grassland flora; several
29/1/09 12:49:33
454
SITE GUIDE
orchids occur including Greater Butterfly, Early Purple, Bee and Pyramidal Orchids.
Near Llanymynech, between Oswestry and Welshpool,
off the A483, with a footpath from the small village
of Pant. Park at the end of the cul-de-sac called Pant
Underhill Lane. The reserve is on the border between
England and Wales. (Shropshire WT)
15. Elan Estate

Sn 930652
A vast mosaic of moorland, blanket bog, Sessile
Oak woodland, conifer plantations, rivers and reservoirs around the Rivers Elan and Claerwen. In the
river valleys the meadows have Common Fragrant,
Heath Spotted and Greater Butterfly Orchids; very
much harder to locate is Bog Orchid in the upland
plateau bogs.
West of Rhayader on the B4518; the reservoirs are
well signposted. There is a Visitor Centre with full details of the 80 miles of leafleted walks and nature
trails, and a Countryside Ranger service.
16. nant Irfon
Sn 840550
A mixture of upland habitats in a spectacular rocky
valley with some beautiful Sessile Oak woodland.
There are also extensive areas of pasture, meadows,
flushes and upland streams with Early Purple and
Common Fragrant Orchids, among others.
On the minor road from Abergwesyn to Terrain. Nant
Irfon is part of a large protected area, which includes
the National Trust reserve of Abergwesyn Common
and another NNR at Claerwen.There is general access
along paths from either end of the valley. (NNR)
17. Burfa Bog
SO 275613
The wet woodland, grasslands and mire have large
colonies of Heath Spotted Orchids that produce a
marvellous display in summer.
Off the B4362 Walton to Ditchyeld Bridge road, near
Presteigne. Park on the road verge. (Radnorshire
WT)
18. Pwll y Wrach
SO 165326
Deciduous woodland beside the River Enig with
Early Purple and Birds-nest Orchids among the interesting flora.
Pwll y Wrach is in the Brecon Beacons National Park
off the A479 Talgarth to Abergavenny road, southeast
of Talgarth. Park at the reserve. (Brecknock WT)
PEMBROKESHIRE
19. Dowrog Common
SM 775273
Some fantastic heathland and bog, grassland, willow
carr and pools. With the neighbouring commons of
Tretio and Waun Fawr, this is a large and botanically
rich area. There are good colonies of Lesser Butterfly and Heath Spotted Orchids, and Southern Marsh
Orchids in the wetter parts.
Just outside St Davids on the A487 to Fishguard, with

footpaths leading from the road. There is a small car
park on the western boundary at SM 772275. (NT)
20. St Davids Head

SM 734272
The walks along the Pembrokeshire Coast Path
close to St Davids encompass cliff grassland and
coastal heathland, with good displays of Common
Spotted, Heath Spotted and Pyramidal Orchids.
Common Twayblade and Lesser Butterfly Orchid
can also be found here.
Access is from St Davids on the B4583 to Whitesand
Bay.
CaRMaRTHEnSHIRE
21. allt Rhyd y Groes
Sn 760480
Ancient woodland on the slopes of a rocky river
valley; Greater Butterfly Orchids can be found in the
adjacent meadows.
The site is in the Cambrian Mountains west of Llanwrtyd Wells, on minor roads from Rhandirmwyn (off the
A483) to the A482 Lampeter road. There are various
paths from the minor roads. (NNR, CCW)
22. Pembrey Burrows
SS 415007
Dunes and a forest of Corsican Pines on former
dunes; the wide, grassy and sheltered rides are an
excellent habitat for Bee Orchid and Marsh Helleborine.
Northwest of Swansea, off the B4311 at Burry Port.
Car parking is at Pembrey Country Park.
GLaMORGanSHIRE
23. South Gower Cliffs
SS 470844
Large areas of the Gower Peninsula are protected
as nature reserves or are in National Trust ownership. There are a wide variety of habitats; the South
Gower Cliffs Reserve contains interesting cliff-top
and old dune grassland, with Green-winged and
Early Purple Orchids.
Access to the Gower is via Swansea or via Junction 47
of the M4. There is car parking at the beach car park
at Port Eynon. (WT)
24. Oxwich Bay
SS 506870
Oxwich Bay, on the south shore of the Gower Peninsula, contains large areas of dunes and dune slacks,
woodland on limestone and freshwater marsh.There
are a good number of orchid species with Pyramidal
and Southern Marsh Orchids, the coccinea subspecies of Early Marsh Orchid, Marsh Helleborine and,
more rarely, Bee Orchid and Green-flowered Helleborine in the dune slacks, and Broad-leaved Helleborine in Nicholaston and Oxwich Woods.
Oxwich village is on a minor road south off the A4118,
29/1/09 12:49:33
WALES
with open access to the dunes and beach and marked

paths in Nicholaston and Oxwich Woods. (NNR,
CCW)
25. Crymlyn Bog

SS 694947
Just north of Swansea docks, this is the largest area
of lowland fen in Wales and is of international importance for its valley mire communities. The vegetation includes rich fen, reedbed and old pasture.
There are Southern Marsh, Common Spotted and
Heath Spotted Orchids and hybrids.
Access is via minor roads leading off the A4217 and
A483 around Kelvey Hill. There is a Visitor Centre on
the west side. (NNR, CCW)
26. Kenfig
SS 780820
North of Porthcawl, this extensive area of calcareous dunes in various stages of development is one
of the richest orchid sites in Wales. The speciality is
Fen Orchid (of the broad-leaved variety ovata). This
grows in the younger dune slacks and has been in
decline in recent years due to a lack of suitable new
slacks. It is hard to find; the best way is to contact
the Information Centre in advance to check flowering dates, which can vary a lot (any time from early
June onwards), and to make sure someone will be
on hand to give precise directions. Other notable
species include a few Green-flowered Helleborines,
and also Broad-leaved Helleborines of the unusual
subspecies neerlandica (known as Dutch Helleborine), although this is erratic in its appearances and
hard to find. Other orchids include Marsh Helleborine, Early Marsh Orchid (including the subspecies
coccinea), Southern Marsh, Pyramidal and Greenwinged Orchids, and Common Twayblade.
Kenfig can be reached from Junction 37 of the M4
and is signposted from North Cornelly, Pyle and Porthcawl. (Bridgend County Borough Council, Tel: 01656
743386; NNR)
455
limit) on a steep slope beside a river gorge. Little

grows on the ground, but Birds-nest Orchid can be
found in some numbers.
West of Abergavenny on the A465, accessed via footpaths from minor roads which run from Clydach to
Llanelly Hill. (NNR, CCW)
29. Pentwyn Farm

ST 523095
A small hill farm reserve of unimproved hay meadows with great views towards the Forest of Dean.
Several orchid species can be found, including
Common Twayblade, Early Purple, Greater Butterfly, Green-winged and Common Spotted Orchids
among the meadows and lanes.
Located at Penallt near Monmouth, the reserve is adjacent to the Bush Inn with parking. (Gwent WT)
30. Springdale Farm
ST 401992
Woodland and grassland on limestone. The woods
contain Early Purple Orchid, Common Twayblade
and Broad-leaved Helleborine, and there are also
many Common Spotted Orchids.
Park beside the almshouse on the minor road south
between Usk and Llantrisant. (Gwent WT)
31. newport Wetlands
ST 334834
A new reserve on the edge of Newport with reedbeds and wet grassland. Heath Spotted and Southern Marsh Orchids can be seen from the many
paths.
The car park is on West Nash Road between Nash
village and Uskmouth Power Station. (NNR, CCW)
27. Merthyr Mawr

SS 870771
The newest NNR in Wales is adjacent to Kenfig. It
has sand, perched sand dunes on top of limestone,
and limestone outcrops, pools and scrub a vast
and interesting area. The dune slacks hold the usual
complement of orchids, including large numbers of
Marsh Helleborines, and Autumn Ladys-tresses.
The reserve has open access, with parking at Candleston Castle at the end of a minor road through Merthyr Mawr village, located off the B4265 southeast of
Bridgend. It can also be accessed from the beach car
park at Newton, Porthcawl.
MOnMOuTHSHIRE
28. Cwm Clydach Woodlands
SO 207123
Important beechwoods (close to their northern
29/1/09 12:49:33
456
SITE GUIDE
Scotland
1
9
8
2
10
7
13
11
12
6
14
15
20
21
18 19
5
24
16
17
25
4
28
27
22
23
Shetland
1 Lumbister
Orkney
2 Birsay Moors and
Cottascarth
Outer Hebrides
3 Balranald
Inner Hebrides
4 Killinallan Dunes, Islay
5 Mull
6 Tokavaig Wood, Skye
Highland
7 Beinn Eighe and Torridon
8 Inchnadamph
9 Invernaver
10 Talich
11 Glen Affric
12 Abernethy Forest and
Loch Garten
Moray
13 Culbin Forest
aberdeenshire
14 Glen Tanar Estate
26
15 St Cyrus
argyll and Bute
16 Ballachuan Hazelwood
17 Knapdale
Perth and Kinross
18 Black Wood of Rannoch
19 Keltneyburn
20 Killiecrankie
angus
21 Glen Cova
ayrshire
22 Auchalton Meadow
23 Feoch Meadows
Fife
24 Tentsmuir Point
East Lothian
25 Aberlady Bay
Dumfries and Galloway
26 Grey Mares Tail
Borders
27 Gordon Moss
28 St Abbs Head
29/1/09 12:49:34
SCOTLAND
SHETLanD ISLES
1. Lumbister
Hu 485967
A series of lochs among heather moorland and sea
cliffs. This reserve covers over 4,000 acres; Heath
Spotted Orchids are easy to spot, but Lesser Twayblade is very inconspicuous here.
Located west of the A968 and northeast of Whale
Firth; access to the site is from the lay-by just northwest of Mid Yell on the Island of Yell, north of Mainland.
The Daal of Lumbister, a steep narrow gorge, makes a
good walk. (RSPB)
ORKnEY ISLES
2. Birsay Moors and Cottascarth HY 368197
Heather moorland with dry and wet heath. This
large area includes the raised mire of Glims Moss
and a calcareous valley mire known as the Dee of
Durkadale. Orchids found here include Lesser Twayblade, and Heath Spotted, Northern Marsh and
Early Marsh Orchids.
On Mainland, on both sides of the B9057.The entrance
to Cottascarth is north of Finstown off the A966, signposted RSPB. Park in the farmyard. (RSPB)
OuTER HEBRIDES
The well-protected machair of the north coast of
North Uist is home to the endemic Hebridean Marsh
Orchid. Other species typical of the machair include
Hebridean Spotted Orchid (Common Spotted Orchid
of the hebridensis subspecies), Northern Marsh Orchid, Early Marsh Orchid of both the pink-flowered
incarnata and red-flowered coccinea subspecies,
Lesser Butterfly Orchid, Frog Orchid and Common
Twayblade.
3. Balranald
nF 714699
An area of machair and dunes interspersed with
lochs and marshes, on the west coast of North
Uist. Predominantly grazing land, the large stretches
of machair are herb-rich, with Frog Orchid; the
marshes have Northern Marsh and Early Marsh
Orchids.
West of the A865 between Tigharry and Paiblesgarry,
northwest of Bayhead; follow signposts for Hougharry.
The cottage at Goular has information and maps.
(RSPB)
InnER HEBRIDES
4. Killinallan Dunes, Islay
nR 3071
These low dunes, which stretch for a few hundred
yards behind the beach, are good for Pyramidal and
Frog Orchids, with small numbers of Early Purple,
457
Early Marsh and Northern Marsh Orchids, Common Spotted and Common Fragrant Orchids and
Common Twayblade. The heaths on either side of
the lane contain Lesser Butterfly Orchid and Heath
Spotted Orchid. A small colony of Marsh Helleborines is a speciality here.
On the northern coast of Islay beside Loch Gruinart.
There is limited parking beside the kissing gate.
5. Mull
Mull is a diverse island, with mountains, sea cliffs
and dune habitats and great wildlife. Thirteen
species of orchid have been recorded here. Broadleaved Helleborine and Early Purple Orchid can be
found in woodland near Tobermory Lighthouse,
while pulchella Early Marsh Orchids occur around
Loch Spelve in the south. Other species to look
for include Small White, Greater Butterfly, Lesser
Butterfly, Common Fragrant, Common Spotted and
Northern Marsh Orchids.
Mull can be accessed by ferry from the mainland at
Oban.
6. Tokavaig Wood, Skye

nG 620120
This wood is on the southern shore of Loch
Eishort on a wide variety of rock types and support a varied flora. Lesser Twayblade grows in
the sandstone and quartzite areas; limestone areas hold Frog Orchid, Dark-red Helleborine and
Common Twayblade.
The Isle of Skye can be reached from the A87, crossing
at the Kyle of Lochalsh. The woods are off the A851
towards Ord and Tokavaig. (NNR)
HIGHLanD
7. Beinn Eighe and Torridon
nG 950621
Some of the finest mountain scenery in Scotland,
with mountain habitats, grassland, moorland, cliffs
and woodland. There is a rich flora in places with
Creeping Ladys-tresses in the Caledonian pinewoods.
Between Loch Torridon and Loch Maree; enclosed by
the A896 and A832, northwest of Kinlochewe.There is
a Visitor Centre with information on walks. (NNR)
8. Inchnadamph
nC 250220
This is Scotlands largest area of limestone, with
limestone pavements and outcrops at the head of
Loch Assynt. Species of interest include Dark-red
Helleborine, Common Fragrant, Frog and Small
White Orchids.
To the east of the A837 at the eastern end of Loch
Assynt, east of Lochinver and north of Ullapool.
(NNR)
29/1/09 12:49:34
458
SITE GUIDE
9. Invernaver
nC 681612
Seashore, dunes, machair and moorland by the
mouth of the River Naver, this area has some of the
most interesting plant communities in the whole of
Scotland. Small White, fragrant (probably Common
Fragrant) and Greater Butterfly Orchids and Darkred Helleborine can be found.
On the north coast of Sutherland around the village of
Bettyhill. Access from Invernaver village, off the A836.
Slightly to the west at Durness, the limestone cliffs
have several orchid species too.
10. Talich
nH 850786
An area of woodland, meadow and marsh on the
bed of a former loch. The wet pastures contain
Heath Fragrant, Heath Spotted, Early Purple and
Lesser Butterfly Orchids.
North of Invergordon on the A9, at Rhynie off the
B9165. (Scottish WT)
11. Glen affric
nH 235240
One of the most attractive valleys in Scotland, there
are remnants of Caledonian pine forest, grassland
and lochs, with Northern Marsh and Lesser Butterfly Orchids.
Southwest of Inverness, on a minor road into the Glen
from Cannich on the A831.There are several car parks
and footpath information. (NNR).
12. abernethy Forest and Loch Garten
nH 980188
Pine forest, lochs and wet moors part of a remnant of Caledonian pine forest. A large area with
an interesting flora, there are several orchid species,
including Creeping Ladys-tresses and Lesser Twayblade.
Off a minor road between Boat of Garten and Nethybridge. Park at the RSPB centre. (RSPB)
MORaY
13. Culbin Forest
nH 997614
This vast area stretches from Nairn in the west to
Findhorn in the east. There are sand dunes which
have been extensively planted with Corsican and
Scots Pines, which hold Creeping Ladys-tresses,
Lesser Twayblade and Coralroot Orchid.
Off minor roads north of the A96.There are several car
parks in the area.
aBERDEEnSHIRE
14. Glen Tanar Estate
nO 480964
The Dees wooded valley has areas of relict Scots
Pine and birch woodland. The surrounding slopes
contain bog and moorland, and extend to the peak
of Beinn aBhuird. In the valley bottom are Lochs
Davan and Kinord. Among several species of orchid

are Lesser Twayblade, Creeping Ladys-tresses and
Heath Fragrant Orchid.
Off the B976 just south of Aboyne. There is a Visitor Centre across the Water of Tanar from Braeloine.
(NNR)
15. St Cyrus
nO 743635
A wonderful part of Montrose Bay, there are dunes
and slacks, and coastal grasslands and cliffs with a
rich flora, including Northern Marsh, Heath Fragrant,
Heath Spotted and Lesser Butterfly Orchids.
North of Montrose off the A92.There is a car park and
Visitor Centre just south of St Cyrus. (NNR)
aRGYLL anD BuTE

16. Ballachuan Hazelwood
nM 763146
An abandoned hazel coppice with a good ground
flora. Heath Fragrant and Northern Marsh Orchids,
and Sword-leaved Helleborine can be found.
At the southern tip of Seil Island, west of the A816. Access from Ballachuan Farm on the B8003 near Cuan.
(Scottish WT)
17. Knapdale
nR 720835
An area of ancient Atlantic oak woodland and coniferous plantation beside Loch Sween, Knapdale is a
good site for Sword-leaved Helleborine.
At the head of Loch Sween, at the north end of the Kintyre Peninsula. Off the B8025 to Tayvallich. (Scottish
WT)
PERTH anD KInROSS

18. Black Wood of Rannoch
nn 570536
The most extensive patch of original Caledonian
pine forest in Perthshire, with Lesser Twayblade and
Coralroot Orchid present.
West of Pitlochry on the B8019 and B846, to the
south side of Loch Rannoch; there are two car parks
with information boards.
19. Keltneyburn
nn 771496
Keltneyburn is a steep wooded gorge with an adjacent herb-rich meadow, Balchroich Meadow. There
are eight species to see here, including Birds-nest,
Greater Butterfly, Heath Fragrant and Small White
Orchids.
Turn west off the B846 c. 5 miles west of Aberfeldy
towards Fortingall, then north off this minor road, just
west of Keltneyburn bridge, on a rough track to the
reserve. (Scottish WT)
20. Killiecrankie
nn 910620
This site includes oak woodland, farmland, birchclad crags and moorland with wet lime-rich areas.
The boggy areas have Northern Marsh Orchid,
29/1/09 12:49:35
SCOTLAND
and Lesser Twayblade can be found amongst the

heather. Birds-nest Orchid occurs in the woodland.
Overlooking the famous Pass of Killiecrankie, west
of the A9 to the north of Pitlochry. The reserve is
signposted from the centre of Killiecrankie village.
(RSPB)
anGuS
21. Glen Clova
nO 330730
One of the five Glens of Angus, Glen Clova is
popular with walkers. Within it grow Heath Fragrant, Small White, Frog and Common Spotted
Orchids.
Access from the B955, to the north of the village of
Kirriemuir. Parking is available at Clova.
aYRSHIRE
22. auchalton Meadow
nS 335036
These meadows on the site of old limestone workings contain a good orchid flora, with Frog and
Northern Marsh Orchids, Lesser and Greater Butterfly Orchids, and Common Twayblade.
Park off the B741 near Broomland Cottage, between
Girvan and Maybole. (Scottish WT)
23. Feoch Meadows
nx 263822
Traditionally managed herb-rich meadows which
have retained a good flora, including Greater and
Lesser Butterfly, Heath Fragrant, Frog and Small
White Orchids.
Off the A714 Newton Stewart to Barrhill Road at
Killantringan. (Scottish WT)
459
A large bay to explore, with many footpaths.

Northeast of Edinburgh. Access via the A198 with a
car park just outside the village of Aberlady. (NTS)
DuMFRIES anD GaLLOWaY

26. Grey Mares Tail nature Reserve
nT 185145
Famous for its waterfall, the botanical richness of this
mountainous landscape should be explored. There
are upland heaths, blanket bog, rocky slopes, a loch
and wonderful scenery. It is a good place to search
for Lesser Twayblade under the heather.
In the Moffat Valley, northeast of Moffat on the A708
Moffat to Selkirk road. There is a Visitor Centre and
car park. (NTS)
BORDERS
27. Gordon Moss
nT 635425
The largest area of semi-natural woodland in the
Borders, this site consists of dense birchwoods on
peat moss. Rich in plants, there are Coralroot Orchids, Lesser Butterfly Orchids and several other
species to be found.
Just outside Gordon, south of the A6105 Gordon to
Earlston road. Park at the reserve entrance on this
road. (Scottish WT)
28. St abbs Head
nT 914692
As well as great coastal scenery and seabird colonies, the grasslands along the top of the cliffs have
many Early Purple Orchids.
From the A1107 take the B6438. This leads to the
car park at Northfield Farm. (Scottish WT, NTS,
NNR)
FIFE
24. Tentsmuir Point
nO 500270
Tentsmuir Forest lies on the southern bank of the
Tay Estuary across the river from Dundee. There is
an extensive dune system, one of the fastest growing in Britain, with a rich flora that includes a very
few Coralroot Orchids in the dune slacks.
Access from Tayport near to the B945.Walk from here
along the beach, or park at the Forestry Commission
car park at Kinshaldy Beach. (FC, NNR)
EaST LOTHIan
25. aberlady Bay
nT 465801
The heathland and sand dunes around Aberlady
Bay hold a rich flora with several orchid species,
including Pyramidal, Common Spotted, Frog and
Heath Spotted Orchids, and Common Twayblade.
29/1/09 12:49:35
460
SITE GUIDE
Ireland (Republic of Ireland and northern Ireland)

28
26
27
25
24
20
18
30
31
22
21
29
32
33
34
23
36
19
35
17
16
14
12
15
2
13
11 10
9
Co. Kerry
1 Killarney National Park
Co. Clare
2 The Burren National Park
Co. Waterford
3 Tramore Dunes
Co. Wexford
4 Bannow Bay
5 Cahore Polders and Dunes
Co. Wicklow
6 Wicklow Mountains National
Park
7 Bray Head
Co. Laois
8 Ballyprior Grassland
9 Clonaslee Eskers and Derry Bog
Co. Galway
10 River Shannon Callows
11 Glenloughaun Esker
12 Rahasane Turlough
13
14
15
16
Coole/Kilcolgan/Ardrahan
Lough Corrib
Inishmore Island
Slyne Head Peninsula and
Aillebrack
17 Connemara National Park
Co. Mayo
18 Lough Carra
19 Urlaur Lakes
20 Lough Conn and Lough Cullin
21 Blacksod Bay
Co. Sligo
22 Mullaghmore
27 Lough Beg
Co. antrim
28 The Giants Causeway
29 Carrick-a-rede and Whitepark Bay
30 Fairhead and Murlough Bay
31 Slieveanorra
32 Shanes Castle
33 Lough Neagh
Co. Down
34 Belfast Harbour Reserve
35 Killard Point
36 Murlough NR
Co.Monaghan
23 Kilroosky Lough Cluster
Co. Fermanagh
24 Crom Estate
25 Castle Coole
Co. Londonderry
26 Umbra
29/1/09 12:49:35
IRELAND
CO. KERRY
1. Killarney national Park
R 970860
The mountainous park to the south and west of
Killarney around Lough Leane has good bog and
moorland vegetation, and there are extensive
woodlands around the famous Lakes of Killarney.
There are many interesting orchids and Irish Ladystresses has been recorded here.
The National Park Visitor Centre is at Muckross
House, at Muckross on the N71, where information
on various walks is available.
CO. CLaRE
2. The Burren
R 324954
One of Europes finest areas of limestone pavement, and of outstanding botanical interest. There
are turloughs (seasonal loughs), fens and some
small stands of woodland, with dunes covering a
large area on the west coast of Co. Clare around
Lisdoonvarna and Kilfenora. The Burren is bisected
by the N67, and many other minor roads allow
easy exploration. Of the 30 species of orchid found
in Ireland, 23 have been recorded in The Burren.
Dense-flowered Orchid is the real speciality. This
favours short, closely grazed turf; it is widespread
but hard to spot. Another speciality is Early Marsh
Orchid of the spotted-leaved subspecies cruenta
(Flecked Marsh Orchid), which is found around
some of the loughs in the east of the region. Early
Purple Orchids are abundant and spectacular, and
Heath and Common Spotted Orchids are widespread (with white-flowered Common Spotted
Orchids of the variety okellyi common here). Irish
Marsh Orchid is scattered and sometimes found in
dry as well as wet grassland habitats. Fly Orchids
occur in open habitats, on rocky hillsides and in fens
around the turloughs; Common Twayblade and, later in the season, Dark-red Helleborine and Marsh
Fragrant Orchid can also be found.
The Burren National Park covers a fairly small area
in the east; the green road at R 304945 gives good
access. Other productive areas include Loch Gealain,
with numerous Flecked Marsh Orchids, Lough Bunny, and the pastures near the coast at Poullsallagh,
which have some Dense-flowered Orchid colonies.
Keelhilla at Slieve Carron on the northeastern edge
of The Burren is also a good area; Fly and Denseflowered Orchids occur there (roadside parking for
a few cars). The grid reference above is for Mullaghmore (National Trust for Ireland) in the centre
of the region, but the whole of the Burren is worth
exploring.
461
CO. WaTERFORD
3. Tramore Dunes
S 580005
A series of fixed dunes and dune slacks with Bee
Orchids present.
South of Waterford on the R675. Park in the town of
Tramore.
CO. WExFORD
4. Bannow Bay
S 804064
A mosaic of sand dune habitats on both sides of the
estuary leading into Bannow Bay. There are damp
slacks where both Bee and Pyramidal Orchids grow.
On the south coast between Wexford and Waterford;
the R734 leads to Fethard on the west side, and the
R736 leads to Newtown on the east side.
5. Cahore Polders and Dunes
T 220462
An important wildlife site on the southeast coast.
A 2/2 mile-long sand dune system extends south
from Cahore Point; behind the dunes there are
areas of grassland, wetland and drainage channels.
Pyramidal Orchid occurs on the fixed dunes at the
northern end. The newer dunes to the south also
have a rich flora, and Marsh Helleborine has been
recorded.
Between Wexford and Arklow, south of Courtown.Take
minor roads off the R742 to reach the site.
CO. WICKLOW
6. Wicklow Mountains national Park
T 099980
This National Park covers much of upland Wicklow,
extending to Blessington Reservoir in the west, Vartry Reservoir in the east, Cruagh in the north and
Lybagh in the south; the Park also includes Glendalough Wood Nature Reserve. There is heathland,
blanket bog and upland grassland, small rivers and
scattered lakes, with a rich flora; the Liffey Head
blanket bog is one of the best of its kind in eastern
Ireland. Small White Orchid and Bog Orchid occur
here but are difficult to find.
The Visitor Centre is in Glendalough Valley off the R756
Green Road close to the Upper Lough (where there is
a car park). It is popular walking country with many
footpaths taking in the major habitat types.
7. Bray Head
O 284174
This coastal site is a plateau of high ground with
rocky knolls and sea cliffs. Heath is the principal
habitat but in some areas there is calcareous grassland with several species of orchids, including Bee,
Fragrant, Pyramidal and Common Spotted Orchids
and Common Twayblade.
29/1/09 12:49:35
462
SITE GUIDE
Off the R761 out of Bray; walk from Bray or Greystones.

The site is signposted from the road.
CO. LaOIS
8. Ballyprior Grassland
S 572924
Located at the north end of the Castlecomer Plateau
and largely underlain by limestone, this site contains
much orchid-rich calcareous grassland, with Early
Purple, Fragrant and Common Spotted Orchids.
Green-winged Orchid has also been recorded here.
The site is 2 1/2 miles south of the village of Stradbally,
which is on the N80 between Portlaoise and Carlow.
9. Clonaslee Eskers and Derry Bog S 265110
A series of glacial moraines and eskers, including
Derry Bog. The wide ranging habitats support an
interesting flora including Fragrant and Fly Orchids.
Three miles west of Clonaslee, northwest of Portlaoise
on the R422.
CO. GaLWaY
10. River Shannon Callows
n 014304
Callows are seasonally flooded wet grasslands.
These form the largest area of lowland semi-natural
grassland in Ireland. They have a good range of characteristic species, varying according to their tolerance to flooding. There are some areas of limestone
pavement and calcareous grassland where Greenwinged Orchid occurs, among other species.
Along the River Shannon between the towns of Athlone
and Portumna;The NationalTrust for Ireland has a reserve
at Mongan Bog (N 032318) and an information point
with parking at Clonmacnoise National Monument.
11. Glenloughaun Esker
M 8226
A fine example of dry, mostly unimproved, orchidrich grassland on an esker. The flora includes large
populations of Green-winged and Early Purple
Orchids.
Three miles southwest of Ballinasloe, off the N6, along
the minor roads to Kilnahown.
12. Rahasane Turlough
M 472238
On the course of the River Dunkellin, this turlough
is surrounded by damp grassland, limestone outcrops and scrubby woodland. A flora similar to that
of The Burren occurs here.
Just west of Craughwell, southeast of Galway City on
the N6. Access from the N6, north on a minor road
towards Athenry.
13. Coole Lough

M 430040
An area of low-lying limestone karst with deciduous
woodland, pasture and limestone heath. Especially interesting are the turloughs, with limestone
pavement extending to their edges in some places.
Pyramidal, Heath Spotted, Common Spotted, Greater Butterfly, Fragrant and Fly Orchids occur here.
Similar habitats occur just slightly north around Caranavoodaun turlough south-east of Kilcolgan. The
dry, calcareous grassland among the limestone pavement here holds Early Marsh, Lesser Butterfly, Fragrant and Dense-flowered Orchids, Broad-leaved
Helleborine and Autumn Ladys-tresses. On the
Ardrahan limestones, the Ballinderreen turlough has
similar species with the addition of Fly Orchid.
Coole Lough (and Visitor Centre) is off the N18, northwest of Gort. (M 430040). Caranavoodaun turlough is
off the N18 SE of Kilcolgan (M 4317). Ballinderreen
turlough is S of Ballinderreen on the N67 (M 3914).
14. Lough Corrib

M 121429
The second largest lake in Ireland, this is surrounded by some excellent habitats, including spring-fed
fen, raised bog, limestone pavement, woodland and
orchid-rich grassland. Pyramidal, Common Spotted,
Early Purple, Frog, Fragrant and Greater Butterfly
Orchids and Marsh Helleborine can all be found.
There are good populations of Irish Ladys-tresses
around the lough and on the Doorus Peninsula.
To the north of Galway City, with information points at
Oughterard on the N59 and Cong on the R345; there
are various picnic spots around the Lough.
15. Inishmore Island
L 830090
The largest of the three Aran Islands, Inishmore is
geologically an extension of The Burren in Co. Clare.
There are some fantastic cliffs, sand dunes, machair,
orchid-rich grassland and limestone pavement, as
well as ancient meadows surrounded by stone walls.
The rich flora includes Common Twayblade, Early
Purple, Common Spotted and Heath Spotted Orchids. Dense-flowered Orchid is a speciality, and Bee
Orchid occurs in the coastal machair.
The Aran Islands can be reached by ferry from Rossaveal or Doolin in Co. Clare and explored by foot.
16. Slyne Head Peninsula and aillebrack
L 620445
In the far west of Galway, this low-lying peninsula
has some extensive areas of machair (particularly
good at Mannin Bay and Aillebrack) and sand dunes
backed by a mosaic of tiny fields, grassland, heath,
lakes, marshes and fens. The specialities are Denseflowered Orchid and, very locally in spring-fed fens,
Pugsleys Marsh Orchid. The dry grassland supports
Early Purple and Green-winged Orchids, and both
Greater and Lesser Butterfly Orchids.
The area lies west of Ballyconneely on minor roads off
the R341.
17. Connemara national Park
L 670520
Connemara has mountains, bogs, heaths, grasslands and woodlands. The predominant habitats are
29/1/09 12:49:36
IRELAND
blanket bog and heathland. Lesser Twayblade is present but, as usual, is hard to find.
Entrance to the park is on the Clifden side of Letterfrack village on the N59.There is a car park and Visitor
Centre, numerous trails and helpful staff. (NPWS)
CO. MaYO
18. Lough Carra
M 190740
A large, shallow, marl lake surrounded by limestone,
with fens, woodland, grassland and limestone pavement. Early Purple and Dense-flowered Orchid
occur. Nearby Lough Mask has Irish Ladys-tresses
along its eastern shore.
Alongside the N84, between Castlebar and Ballinrobe.
Access from minor roads around the lough, and park
on the east side south of Carrownacon.There are many
picnic areas on the east side of Lough Mask on the
scenic road.
19. urlaur Lakes
M 520899
Three small calcareous lakes, Lough Nanoge, Lough
Roe and Urlaur Lough, lie in the upper catchment
of the River Lung, a major tributary of the Boyle.
Swamps and reedbeds fringe the lakes, and in some
areas there is flushed, species-rich heath with Lesser
Butterfly, Pyramidal and Heath Spotted Orchids.
Common Twayblade can also be found.
North of Ballyhaunis, on either side of the N83 near
Urlaur. Access via minor roads.
20. Lough Conn and Lough Cullin
G 216048
There are fine oak woodlands around the southern shores of both these lakes, each with a mixed
shrub layer and good ground flora. Sword-leaved
Helleborine has been recorded here. Lesser Twayblade occurs further up the slopes on the blanket
bog, while there are important populations of Irish
Ladys-tresses around the lake margins.
There is information at the nearby town of Ballina. Picnic sites are on Lough Conn at Crossmolina on the
N59, and on Lough Cullin beside the R318.
21. Blacksod Bay
F 700330
The Mullet Peninsula and Blacksod Bay form an
important wildlife habitat. There are fixed dunes,
machair and dune heath, as well as alkaline fens
and coastal habitats. The dunes and machair can be
found in the north of the peninsula to the west of
Termoncarragh Lough, Tonamace and Cross Lough;
such habitat also occurs on the eastern shore of
the bay, around Doolough, Srah and Dooyork. Orchids to be found include Lesser Butterfly, Pyramidal,
Common Spotted and Early Marsh, and Pugsleys
Marsh Orchid has also been recorded here.
Situated in northwest Mayo, Belmullet is on the R313,
from where the peninsula can be explored.
463
CO. SLIGO
22. Mullaghmore
G 710580
This tiny site measures just 100 x 200 yards, but it is
extraordinarily orchid-rich; Pyramidal, Fragrant, Frog,
Lesser Butterfly and Northern Marsh Orchids all
occur, as do Common Twayblade and Marsh Helleborine. Common Spotted Orchid, the red-flowered
coccinea subspecies of Early Marsh Orchid, and
Heath Spotted Orchid are found in their hundreds.
On a headland off the N15 between Sligo and Bundoran, on the R279 to Mullaghmore.
CO. MOnaGHan
23. Kilroosky Lough Cluster
H 490276
Lough Kilroosky, Lough Burdautien, Lough Summerhill and Dummys Lough are a series of marl lakes,
low in nutrients and surrounded by fen and speciesrich freshwater marsh. Nearby lakes are surrounded
by reed swamp and wet woodland. Many species
of orchid can be found here, including Marsh Helleborine.
Northwest of Clones, straddling the border with Northern Ireland.
CO. FERManaGH
24. Crom Estate
H 455655 - 361245
On the shores of Upper Lough Erne, this is one
of Irelands most important conservation areas. Orchids include Common and Heath Spotted, Early
Purple, Fragrant, Birds-nest, and Greater and Lesser
Butterfly Orchids, plus Broad-leaved Helleborine
and Common Twayblade.
The Visitor Centre and car park lie west of Newtownbutler on the minor road to Crom, with walks through
the wetlands and woodlands. (NT)
25. Castle Coole
H 378788
The landscaped grounds of Castle Coole have walks
in woodland to Lough Coole, where Common
Twayblade, Birds-nest Orchid and Broad-leaved
Helleborine can be found.
Just east of Enniskillen on the A4. (NT)
CO. LOnDOnDERRY
26. umbra
C 726358
A dune system on the north coast, with woodland
and scrub behind the beach of Magilligan Strand.The
rich flora in the dune slacks includes Early Purple,
Common Spotted, Northern Marsh, Early Marsh
(two subspecies, incarnata and coccinea), Pyramidal
and Bee Orchids, Common Twayblade and Marsh
Helleborine.
29/1/09 12:49:36
464
SITE GUIDE
Access from the A2 Coleraine to Limavady road; the

site is adjacent to the Umbra level crossing on the A2,
and has roadside parking. The whole Magilligan dune
system is worthy of exploration. (Ulster WT)
27 Lough Beg
H 9895
Just north of Lough Neagh, this area holds Irish Ladys-tresses which can be found in damp grassland
on the western shore. Early Marsh Orchid (subspecies pulchella) also occurs.
A good area to look is between the Ballydermott Road
and Church Island (H 966953).
CO. anTRIM
28. The Giants Causeway
C 954453
These famous polygonal columns of layered basalt
are a World Heritage Site. Several species of orchid
can be found along and near to the North Antrim
Coastal Path and beside paths to the Causeway,
including Frog, Common Spotted, Heath Spotted,
Northern Marsh and Early Purple Orchids, and Irish
Ladys-tresses has recently been found here.
On the B146 2 miles from Bushmills. There are car
parks and a Visitor Centre. (NT)
29. Carrick-a-rede and Whitepark Bay
D 062450
On the North Antrim Coast east of the Giants
Causeway, this rocky outcrop is accessible only
via a rope bridge. The grasslands in this area have
Common Twayblade, Early Purple, Greater Butterfly,
Fragrant, Common Spotted, Northern Marsh and
Pyramidal Orchids. At nearby Whitepark Bay, the
dunes, grassland and scrub, with limestone cliffs to
the east, support similar species.
On the North Antrim coast road; there are picnic areas,
footpaths and a nature trail to Whitepark Bay. (NT)
30. Fairhead and Murlough Bay
D185430
On the north coast 4 miles east of Ballycastle, this
area of rugged cliffs has limestone outcrops alongside a wooded area behind Murlough Bay. Common
Twayblade, Birds-nest, Bee, Pyramidal, Northern
Marsh and Common and Heath Spotted Orchids
all occur.
Access from the A2, parking at Murlough Bay. There
are several paths and viewpoints. (NT)
31. Slieveanorra Forest

D 132265
Mixed woodland and moorland; there are forest
tracks to the summit of Slieveanorra where there
are interesting plots of peat bog at various stages
of formation. The bogs contain a range of species,
including Lesser Twayblade and Heath Spotted Orchid.
Slieveanorra is on the Altarichard road off the Ballymoney to Cushendun Scenic Route.
32. Shanes Castle

J 111881
Lying on the north shore of Lough Neagh, this
area includes woods, parkland and farmland. Castle
Meadow holds carpets of Common Spotted Orchids; there are Broad-leaved Helleborines in the
woods, and Irish Ladys-tresses have been recorded
along the shores of the lough.
Enter via the park gate 1 mile west of Antrim on the
A6 Randalstown road. (RSPB)
33. Lough neagh
J 050620
Irelands largest freshwater lough, bounded by
meadows, reedbeds and woodland. There are many
nature reserves including the Montiaghs NNR and
Lough Neagh NNR. Irish Ladys-tresses (albeit much
reduced), Lesser Butterfly Orchid and Early Marsh
Orchid (subspecies pulchella) can be found.
Situated to the west of Belfast, many minor roads lead
to the shore with picnic areas.The grid reference is for
the Discovery Centre on Oxford Island where information can be obtained regarding access and flowering
times from the staff and the Environment & Heritage
Service wardens. (Craigavon Borough Council)
CO. DOWn
34. Belfast Harbour Reserve
J 398795
The reserve and surrounding area hold Bee Orchid,
Common Spotted and Northern Marsh Orchids
and good displays of Dactylorhiza hybrids. Pyramidal
Orchid and Common Twayblade may also be found.
3 miles north of Belfast city centre, off the A2 on the
south side of Belfast Lough. Speak to the warden for
access arrangements. (RSPB)
35. Killard Point
J 610433
Killard Point marks the southern limit of Strangford
Lough, and has dunes, lime-rich boulder clay grassland, scrub, heath and low cliffs. One of the best
orchid sites in Northern Ireland: Common Twayblade, and Common Spotted, Heath Spotted, Early
Marsh, Early Purple and Green-winged Orchids occur in good numbers and there are also Bee, Frog,
Pyramidal and Northern Marsh Orchids in smaller
numbers.
Approximately 7 miles east of Downpatrick. Take the
minor road between Kilclief and Ballyhornan from the
A2. Park in one of the lay-bys. (NNR)
36. Murlough nature Reserve
J 414351
This large area of dunes forms a peninsula that projects into Dundrum Bay.The rich flora includes Common Spotted, Heath Spotted, Pyramidal, Bee and
Northern Marsh Orchids, and Common Twayblade.
On the coast, about 3 miles north of Newcastle. Access
from the A2; park at Dundrum and walk over Downshire Bridge to the reserve. (NT, NNR)
29/1/09 12:49:37
GLOSSARY
465
Glossary
achlorophyllose lacking the green pigment chlorophyll and therefore unable to photosynthsise.
actinomorphic radially symmetrical; with more than one plane of symmetry.
adventitious buds and roots that appear in abnormal places on the stem.
ancient woodland woodland that has maintained a more or less continuous cover of trees, probably
for thousands of years.
annular ring-shaped.
anther the pollen-bearing, male reproductive organ. In most orchids, the pollen is grouped into two
pollinia.
anther cap in some orchids, such as the helleborines Epipactis, the anther lies on top of the column
and is hinged or stalked. It may be contrastingly coloured.
anthocyanins group of pigments that produce purple or reddish colours.
apomixis reproduction by seed, originating from unfertilised egg cells.
asymbiotic when a symbiotic fungus is absent.
auricles ear-like structures.
autogamy self-pollination with pollen from the same flower (see self-pollinate).
back-cross cross between a hybrid and one of its parent species.
base-rich soil with a high concentration of calcium or magnesium and a pH above 7.0.
bog plant community on wet, acidic peat.
bosses irregular swellings.
bract structure at the base of a flower stalk, varying in size and shape, but often leaf-like.
bulbils tiny, round growths, e.g. along the rim of the leaf of Bog Orchid, which can separate and are
capable of developing into a new plant.
bursicle the pouch-like structure on the column of some orchids that contains and protects the
viscidium (q.v.).
calcareous rich in calcium carbonate, e.g. chalk, limestone or sea shells.
Caledonian woodland ancient pine woodland, a relict of the Forest of Caledon that supposedly
once covered Scotland.
capsule the dry seed pod of an orchid.
carapace hardened shell.
caudicle the stalk present in some orchids that attaches the pollinium to the viscidium (qq.v.).
cilia minute, thickened or fleshy hair-like structures.
ciliate with cilia projecting from the margin.
chlorophyll a green pigment, important in photosynthesis, found in discrete organelles (chloroplasts)
in the cells of plants, usually in the leaves.
cleistogamy self-pollination in bud; after which the bud may remain closed or may open.
clinandrium depression on the top of the column, below the anther and behind the stigmatic zone,
in which the pollinia lie.
clone individual of identical genetic make-up to its parent that results from asexual, vegetative reproduction.
column specialist structure characteristic of orchid flowers in which the stamens and stigmas are
fused together.
crenate with scalloped margins.
cross-pollinate pollination in which pollen from one flower fertilises another; usually taken to mean
a flower on a different plant.
29/1/09 12:49:37
466
GLOSSARY
decurved curved downwards.

deflexed bent sharply downwards.
deltoid shaped like the Greek letter delta, i.e. triangular.
diploid having two matching sets of chromosomes. This is the normal state for plant cells.
ectomycorrhiza association of a fungus with the roots of a plant where the fungus forms a layer on
the outside of the roots.
endomycorrhizal association of a fungus with the roots of a plant in which the fungus penetrates
the tissue of the root.
epichile outer portion of the lip, often heart-shaped, in those orchid genera where the lip is divided
into two (e.g. Epipactis, Cephalanthera, Serapias).
epidermis skin or surface layers.
epiphyte plant growing on the surface of another plant without receiving nutrition from it.
esker glacial debris, often sands and gravels.
fen plant community on alkaline, neutral or very slightly acidic soil.
filiform thread-like.
flexuous wavy.
eutrophication enriched with plant nutrients, often leading to a luxuriant and eventually stifling
growth of vegetation.
geitonogamy fertilised by pollen from another flower on the same plant.
glandular hair short hair tipped with a small spherical gland containing oil or resin.
hanger a wood on a steep hillside.
herbarium (plural: herbaria) a collection of dried and pressed plant material.
hooded formed into a concave shape resembling a monks cowl.
hybrid plant originating from the fertilisation of one species by another.
hybrid swarm population in which the barriers between two species have largely or completely
broken down. Hybridisation is commonplace and at random, producing a population that forms
a continuous range of intermediates between the two parent species.
hybrid vigour when the first generation of hybrids between two species are exceptionally large and
robust.
hyperchromic intensely coloured, with an excessive amount of pigmentation.
hyperresupinate when the ovary and/or pedicel twist through 360 to position the lip at the top of
the flower, e.g. Bog Orchid.
hypha (plural: hyphae) fine, thread-like structures that make up the body of a fungus.
hypochile inner portion of the lip, often cup-shaped, in those orchid genera where the lip is divided
into two (e.g. Epipactis, Cephalanthera, Serapias).
intergeneric hybrid a hybrid whose parents are in two different genera.
internode section of stem between two nodes.
lanceolate narrowly oval, tapering to a more or less pointed tip.
lax loose, not dense.
lignify become hardened and woody.
Leblanc chemical process used to produce washing soda.
lip highly modified third petal of an orchid; also known as the labellum.
lough Irish term for loch.
machair sandy, lime-rich soil with a species-rich sward of short grasses and herbs. The machair is
confined to the coasts of western Ireland and western Scotland.
29/1/09 12:49:37
GLOSSARY
467
meadow grassy field from which stock are excluded for at least part of the year so that it can be cut
for hay.
monocarpic flowering once and then dying.
mutualism an intimate relationship between two or more organisms from which all derive benefits.
mycorhizome early stages in the development of the underground rhizome in which the seedling is
nourished entirely by fungi.
mycorrhiza association of a fungus with the roots of a plant in which the fungus may form a layer on
the outside of the roots (ectomycorrhizal, q.v.) or penetrate the tissue of the root (endomycorrhizal,
q.v.).
mucro very short, bristle-like tip.
mycelium the mass of branching filaments that make up the body of a fungus.
mycotrophic acquiring nutrition from fungi.
native growing in an area where it was not introduced, either accidentally or deliberately, by
humans.
node point on a stem from which leaves, flowers or lateral stems grow.
non-sheathing leaf a leaf with its base clasping the stem but not completely encircling it.
ovary female reproductive organ that contains the ovules.
ovule organ inside the ovary that contains the embryo sac, which in turn contains the egg.
pH measure of acidity.
parasitic organism that lives on or at the expense of other organisms.
pasture grassland that is grazed for some or all of the year but not cut.
petals inner row of perianth segments, one of which is modified to form the lip.
patent projecting more or less at right-angles.
papilla (plural: papillae) small, nipple-like projection.
pedicel stalk of the flower; very short in many orchids, with the cylindrical and sometimes slender
ovary forming the apparent stalk.
pendant hanging downwards.
peloton a coil-like structure formed by fungi inside the cells of an orchid.
pheromone chemical secreted by an animal, especially an insect, that influences the behaviour or
development of others of the same species.
photosynthesis production of food by green plants. In the presence of chlorophyll and light energy
from the sun, carbon dioxide and water are converted into carbohydrates and oxygen.
phototrophic acquiring nutrition through the process of photosynthesis.
pollen single-celled spores containing the male gametes.
pollinium (plural: pollinia) regularly-shaped mass of individual pollen grains which is transported as
a single unit during pollination; the pollinia are often divided into two.
propagules various vegetative portions of a plant such as a bud or other offshoots that aid in dispersal
and from which a new individual may develop.
protocorm initial stage of development for every orchid formed by a cluster of cells.
pseudobulb swollen or thickened portion of stem, covered in the leaf bases. It fulfils the same storage
function as a bulb or tuber; found in Fen and Bog Orchids and common in tropical species.
pseudocopulation attempts by an insect to copulate with an insect-mimicking flower.
pseudopollen structures in a flower that imitate pollen in order to attract insects, e.g. in the
Cephalanhera helleborines.
reflexed bent back or down.
resupinate when the ovary and/or pedicel twist through 180 to position the lip at the bottom of
the flower.
29/1/09 12:49:37
468
GLOSSARY
reticulation marked with a network of veins.

rhizoid hair-like structure on the surface of a protocorm or mycorhizome that facilitates the entrance
of fungi (i.e. a root hair).
rhizome underground stem that lasts for more than one year from which roots and growth buds
emerge; also a horizontal stem, either growing along the surface or underground.
rostellum a projection from the column that often functions to separate the pollinia from the stigma
and thus prevents self-pollination (the rostellum is actually a modified sterile third stigma). The
rostellum may exude a viscidium (q.v.).
runner horizontal stem that grows along or just below the surface of the soil.
saprophytic plants, fungi, etc. that feed on dead organic matter.
scale leaf a leaf that is reduced to a small scale.
secondary woodland woodland in which the continuity of tree cover has been broken for a substantial
period of time.
secund all facing in the same direction.
self-pollinate pollination of a flower by pollen taken from the same plant; usually used in the context
of autogamy, where pollination is by pollen from the same flower, but sometimes also geitonogamy,
where the pollen comes from another flower on the same plant.
sepal outer row of perianth segments that form the protective covering of the bud. In orchids they
are either green or brightly coloured and form a conspicuous part of the flower.
sessile stalkless.
symbiosis an intimate relationship between two or more organisms; formerly used for a
relationship where all participants derive benefits (q.v. mutualism) but now used in a broader
sense to include parasitism
sheathing leaf main leaf on an orchid with a base that completely encircles the stem.
sinus indentation between two lobes on the lip, used especially when describing the flowers of marsh
orchids and spotted orchids in the genus Dactylorhiza.
speculum pattern on the lip, often with a metallic lustre or shine, in the bee and spider orchids, genus
Ophrys.
spur sack-like extension of the base of the lip which contains nectar in some species of orchid.
stamens male reproductive organs of a flowering plant.
staminode sterile stamen that forms a prominent shield-shaped structure within the flower of
Ladys-slipper.
stigma receptive surface of the female reproductive organs to which the pollen grains adhere.
subsecund almost second, i.e. almost all facing to one side.
sympodial pattern of growth in which the tip of the stem or rhizome either terminates in a flower
spike or dies each year. Growth continues from buds formed at the base of the old stem.
synonym former scientific name.
synsepal structure formed when the two lateral sepals are joined for almost their entire length, found
in Ladys-slipper.
tetrads group of four pollen grains originating from a single mother cell.
tetraploid having four sets of chromosomes.
tubers swollen underground roots or stems, functioning as storage organs.
turbary an area where peat or turf is cut for fuel.
turlough Irish term, used for a seasonal lake on limestone in which the water level may fall dramatically
in summer.
viscidium (plural viscidia) detachable sticky exudation from the rostellum that attaches the pollinia
(sometimes via a short stalk, the caudicle) to a visiting insect (qq.v.).
zygomorphic having only one plane of symmetry.
29/1/09 12:49:38
BIBLIOGRAPHY
469
Sources of Information
and Bibliography
In researching this book we have consulted many books and articles, and a complete list of consulted
works appears below. We would, however, like to acknowledge several works in particular that have
been especially valuable. A great debt is owed to the Wild Orchids of Britain (Summerhayes 1968),
which beautifully summarises knowledge on the orchid flora up to that point. More recent advances,
and especially the matter of subspecies, varieties and hybrids, were expertly presented by D.M.
Turner Ettlinger in Notes on British and Irish Orchids (1997) and Illustrations of British and Irish
Orchids (1998). Several regional orchid floras have also been a goldmine of information, for Box
Hill (Sankey 2000), Dorset ( Jenkinson 1991), Hampshire and the Isle of Wight ( Jenkinson 1995),
Scotland (Allan & Woods 1993), Suffolk (Sanford 1991) and Sussex (Lang 2001). Two other works
are worthy of particular mention. Rasmussen (1995) is a comprehensive summary of development
from seed and later growth, while Van Der Cingel (1995) summarises information on pollination.
Note that, in the following, BSBI = Botanical Society of the British Isles.
Ackerman, J.D. & Mesler, M.R. 1979. Pollination biology of Listera cordata (Orchidaceae). American J. Bot. 66:
820-824.
Adcock, E.M., Gorton, E. & Morries, G.P. 1983. A study of some Dactylorhiza populations in Greater
Manchester. Watsonia 14: 377-389.
Allan, B. & Woods, P. 1993. Wild Orchids of Scotland. Edinburgh: HMSO.
Akeroyd, J. 1993 Wildlife reports. Flowering plants. Brit. Wildlife 4: 191-192.
Alexander, C. & Alexander, I.J. 1985. Seasonal changes in populations of the orchid Goodyera repens Br. and its
mycorrhizal development. Trans. Bot. Soc. Edin. 44: 219-227.
Allen, D.E. 1968. Neotinea intacta (Link) Reichb. in the Isle of Man. Proc. Bot. Soc. Brit. Is. 7: 165-168.
Allen, D.E. 1971. Dactylorhiza fuchsii subsp. okellyi (Druce) So. - Behaviour and characters in the Isle of Man.
Watsonia 8: 401-402.
Anon. 1956. The Military Orchid in Suffolk. Proc. Bot. Soc. Brit. Is. 2: 4-5.
Anon. 1958. Spiranthes romanzoffiana in South Devon. Proc. Bot. Soc. Brit. Is. 3: 37-38.
Arditti, J. & Ghani, A.K. 2000. Tansley Review No. 110 Numerical and physical properties of orchid seeds
and their biological implications. New Phytol. 145: 367-421.
Ayasse, M., Schiestl, F.P., Paulus, H.F., Ibarra, F. & Francke, W. 2002. Pollinator attraction in a sexually deceptive
orchid by means of unconventional chemicals. Proc. R. Soc. Lond. B 270: 517-522.
Bateman, R.M. 1985. Peloria and pseudopeloria in British orchids. Watsonia 15: 357-359.
Bateman, R.M. 2001. Evolution and classification of European orchids: insights from molecular and
morphological characters. J. Eur. Orch. 33: 33-119.
Bateman, R.M. 2004. Burnt Tips and Bumbling Bees: How many orchid species currently occur in the British
Isles? J. Hardy Orch. Soc. 1: 10-18.
Bateman, R.M. 2006. How many orchid species are currently native to the British Isles? In: Bailey, J. & Ellis, R.
G., eds, Contributions to taxonomic research on the British and European flora: 89110. London: BSBI.
Bateman, R.M. 2009. Whats in a Name? 1. The heavy responsibility of using a previously described name. J.
Hardy Orchid Soc. 6 (2).
Bateman, R.M. & Denholm, I. 1983a. A reappraisal of the British and Irish dactylorchids, 1. The tetraploid
marsh-orchids. Watsonia 14: 347-376.
Bateman, R.M. & Denholm, I. 1983b. Dactylorhiza incarnata (L.) Soo subsp. ochroleuca (Boll): F. Hunt &
Summerhayes. Watsonia 14: 410-411.
Bateman, R.M. & Denholm, I. 1983c. Diploid Marsh Orchids in the British Isles. Watsonia 14: 448.
29/1/09 12:49:38
470
BIBLIOGRAPHY
Bateman, R.M. & Denholm, I. 1985. A reappraisal of the British and Irish dactylorchids, 2. The diploid marshorchids. Watsonia 15: 321-355.
Bateman, R.M. & Denholm, I. 1989. A reappraisal of the British and Irish dactylorchids, 3. The Spottedorchids. Watsonia 17: 319-349.
Bateman, R.M. & Denholm, I. 1989. On measuring Marsh-orchids. Morphometric procedure, taxonomic
objectivity and Marsh-orchid systematics. Watsonia 17: 449-462.
Bateman, R.M. & Denholm, I. 1995. The Hebridean Marsh-Orchid: nomenclatural and conceptual clarification
of a biological enigma. Edinburgh J. Bot. 52: 55-63.
Bateman, R.M. & Farrington, O.S. 1987. A morphometric study of x Orchiaceras bergonii (Nanteuil) Camus
and its parents (Aceras anthropophorum (L.) Aiton f. and Orchis simia Lamarck) in Kent. Watsonia 16: 397407.
Bateman, R.M. & Farrington, O.S. 1989. Morphometric comparison of populations of Orchis simia Lam.
(Orchidaceae) from Oxfordshire and Kent. Bot. J. Linn. Soc. 100: 205-218.
Bateman, R.M. & Farrington, O.S. 1999. A new infrageneric orchid hybrid for Britain. BSBI News 80: 19.
Bateman, R. & Sexton, R. 2008. Ongoing HOS Platanthera Spur-Length Survey, a great success. J. Hardy
Orchid Soc. 5 (1): 20-23.
Bateman, R. & Sexton, R. 2008. Is spur length of Platanthera species in the British Isles adaptively optimized
or an evolutionary red herring? Watsonia 27: 1-21.
Bateman, R.M., Hollingsworth, M., Preston, J., Yi-Bo, L., Pridgeon, A.M. & Chase, M.W. 2003. Molecular
phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidaceae) Bot. J. Linn. Soc. 142:
1-40.
Bateman, R.M., Pridgeon, A.M. & Chase, M.W. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae,
Orchidaceae) based on nuclear ITS sequences. 2 Infrageneric relationships and reclassification to achieve
monophyly of Orchis sensu stricto. Lindleyana 12: 131-141.
Bidartondo, M.I., Burghardt, B., Gebauer, G., Bruns, T.D. & Read, D.J. 2004. Changing partners in the dark:
isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids and trees. Proc. Royal.
Soc. Lond. B 271: 1799-1806.
Blackmore, S. 1985. Bee Orchids. Princes Risborough: Shire Publications.
Bowler, J. 2003. Irish Ladys-tresses (Spiranthes romanzoffiana) discovered on the Isle of Tiree, Argyll. BSBI News
92: 25-26.
Braithwaite, M.E., Ellis, R.W. & Preston, C.D. 2006. Change in the British Flora 19872004. London: BSBI.
Brewis, A., Bowman, P. & Rose, F. 1996. The Flora of Hampshire. Colchester: Harley.
Briggs, M. 1995. Orchid seed dispersal. BSBI News 70: 19.
Brown, P.M. 2003. The Wild Orchids of North America North of Mexico. Gainesville: University Press of
Florida.
Burgess, J. 1991. The scent of Orchis morio, 2. BSBI News 58: 27.
Buttler, K.P. 1991. Field Guide to the Orchids of Britan and Europe. Swindon: Crowood Press.
Byfield, A. 1993. The Status and Ecology of Cephalanthera longifolia in Britain with Conservation Recommendations.
Plantlife/Hampshire Wildlife Trust.
Cameron D.D., Leake J.R. & Read D.J. 2006. Mutualistic mycorrhiza in orchids: evidence from plant-fungus
carbon and nitrogen transfers in the green-leaved terrestrial orchid Goodyera repens. New Phytol. 171: 405416.
Cameron D.D., Johnson I., Leake J.R. & Read D.J. 2006. Mycorrhizal acquisition of inorganic phosphorus by
the green-leaved terrestrial orchid Goodyera repens. Ann. Bot. 99: 831-834.
Carey, D. 1998. Modelling the spread of Himantoglossum hircinum (L.) Spreng. at a site in the south of England.
Bot. J. Linn. Soc. 126: 159-172.
Carey, D. 1999. Changes in the distribution and abundance of Himantoglossum hircinum (L.) Sprengel
(Orchidaceae) over the last 100 years. Watsonia 22: 353-364.
Carey, D. & Farrell, L. 2002. Biological Flora of the British Isles No. 221. Himantoglossum hircinum (L.) Sprengel.
J. Ecol. 90: 206-218.
Catling, M. 1980. Rain-assisted autogamy in Liparis loeselii (L.) L. C. Rich. (Orchidaceae). Bull. Torrey Bot. Club
107: 525-529.
29/1/09 12:49:38
BIBLIOGRAPHY
471
Cheffings, C.M. & Farrell, L. (eds) 2005. The vascular plant Red Data List for Great Britain. Species Status 7:
1-116. JNCC, Peterborough.
Clarke, D. & Iveson, D. 2003. A new Cumbrian site for the Dune Helleborine Epipactis dunensis (T. & T. A.
Stephensos) Godfery. BSBI News 93: 17-18.
Clarke, W.A. 1900. First Records of British Flowering Plants. London: West, Newman & Co.
Cobbing, P. 1989. Serapias parviflora Parl. BSBI News 52: 11-12.
Cole, S. 2008. Naming orchid taxa: what are the rules? J. Hardy Orchid Soc. 5(3): 86-90.
Coleman, R. A. 1995. The Wild Orchids of California. Ithaca: Comstock.
Coleman, R. A. 2002. The Wild Orchids of Arizona and New Mexico. Ithaca: Comstock.
Cowie, N.R. & Sydes, C. 1995. Status, distribution, ecology and management of Lapland marsh orchid
Dactylorhiza lapponica. Scottish Natural Heritage Review no. 42. Edinburgh: Scottish Natural Heritage.
Curtis, T.G.F. & McGough, H.N. 1988. The Irish Red Data Book. 1. Vascular Plants. Dublin: Stationary Office.
Dafni, A. & Woodell, S.R.J. 1986. Stigmatic exudate and the pollination of Dactylorhiza fuchsii (Duce) Soo.,
Flora, Morphol., Geobot., Oekophysiol. 178: 343-350.
Darwin, C. 1892. The Various Contrivances by which British and Foreign Orchids are Fertilised by Insects, and on the
Good Effects of Intercrossing. London: John Murray.
Davies, P., Davies, J. & Huxley, A. 1983. Wild Orchids of Britain and Europe. London: Hogarth Press.
Davies, P.H. 1989. Identification. British Bee Orchids. Brit. Wild. 1: 37-40.
Davies, P.H. 1990. Identification. Epipactis Helleborines. Brit. Wild. 2: 106-110.
Davies, K. L. 1996. Welsh notes: the rediscovery of Epipactis leptochila (Godfery) Godfery var. leptochila in south
Wales. Orch. Rev. 104: 299-300.
DAyala, R. & Snell, N. 2002. Listera cordata (Lesser Twayblade) in Bucks (V.C. 24). BSBI News 91: 33.
Delforge, P. 1995. Orchids of Britain & Europe. London: HarperCollins.
Delforge, P. 1995. Epipactis dunensis (T. & T.A. Stephenson) Godfery et Epipactis muelleri Godfery dans les les
Britanniques. Natural. Belges (Orchid. 8) 76: 103-123.
Delforge, P. 2001. Guides des Orchides dEurope. Lausanne: Delachaux et Niestl.
Delforge, P. 2006. Orchids of Europe, North Africa and the Middle East. A&C Black, London.
Delforge, P. & Gvaudan, A. 2002. Contribution taxonomique et nomenclaturale au groupe dEpipactis leptochila.
Natural.Belges 83: 19-35.
Duffy, K.J., Scopece, G., Cozzolino, S., Fay, M.F., Smith, R.J. & Stout, J.C. 2008. Ecology and genetic diversity
of the dense-flowered orchid, Neotinea maculata, at the centre and edge of its range. Annals of Botany.
Edmondson, T. 1979. Ophrys apifera Huds. in artificial habitats. Watsonia 12: 337-338.
Ehlers, B.K., Olesen, J.M. 1997. The fruit-wasp route to toxic nectar in Epipactis orchids? Flora, Morphol.,
Geobot., Oekophysiol. 192: 223-229.
Ennis, T. 2007. The occurrence of Dactylorhiza lapponica in Co. Antrim: a new Irish record. BSBI News 105:
13-15.
Ettlinger, D.M.T. 1976. British & Irish orchids. London: Macmillan.
Ettlinger, D.M.T. 1979. x Pseudorhiza bruniana (Brgger): F. Hunt in Orkney. Watsonia 12: 259.
Ettlinger, D.M.T. 1991. Two new varieties of British Dactylorhiza. Watsonia 18: 307-309.
Ettlinger, D.M.T. 1990. The 1976 Ophrys bertolonii in Dorset flora. BSBI News 54: 15-16.
Ettlinger, D.M.T. 1997. Notes on British and Irish Orchids. Dorking: Privately published.
Ettlinger, D.M.T. 1998. Illustrations of British and Irish Orchids. Dorking: Privately published.
Ettlinger, D.M.T. 1998. A new variety of Ophrys apifera Hudson (Orchidaceae). Watsonia 22: 105-107.
Ettlinger, D.T. 1992. More on white orchids. BSBI News 60: 7-8.
Ettlinger, D.T. 1997. Yellow-edged fly orchid flowers - finale. BSBI News 77: 33.
Evans, P.A., Evans, I.M. & Rothero, G.P. 2002. Flora of Assynt. Privately published.
Ewen, A.H. & Prime, C.T. (trans. & eds) 1975. Rays Flora of Cambridgeshire (Catalogus Plantarum circa
Cantabrigiam Nascentium). Hitchin: Wheldon & Wesley.
Farrell, L. 1985. Biological Flora of the British Isles. No. 160. Orchis militaris L. (O. galatea Poir, O. rivini Gouan,
O. tephrosanthes Willd. & Sw.). J. Ecol. 73: 1041-1053.
Farrell, L. 1991. Population changes and management of Orchis militaris at two sites in England. In T.C.E Wells
& J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague: SPB Academic Publishing.
29/1/09 12:49:39
472
BIBLIOGRAPHY
Fisher, J. 1991. A Colour Guide to Rare Wild Flowers. London: Constable.

Fitter, R. 1994. The second Oxfordshire Orchis simia site. BSBI News 66: 17.
Fitter, R.S.R. 1957. Cephalanthera rubra (L.) Richard in Buckinghamshire. Proc. BSBI 2: 234-35.
Foley, M.J.Y. 1987. The current distribution and abundance of Orchis ustulata L. in northern England. Watsonia
16: 409-415.
Foley, M.J.Y. 1989. Dactylorhiza traunsteineri (Sauter) Soo: variants in north-east Yorkshire. Watsonia 17: 355356.
Foley, M.J.Y. 1990. The current distribution and abundance of Orchis ustulata L. in southern England. Watsonia
18: 37-48.
Foley, M.J.Y. 1992. The current distribution and abundance of Orchis ustulata L. (Orchidaceae) in the British
Isles an updated summary. Watsonia 19: 121-126.
Foley, M.J.Y. 2000. Dactylorhiza incarnata (L.) Soo subsp. ochroleuca (Wstnei ex Boll): F. Hunt and Summerh.
(Orchidaceae): A comparison of British and European plants. Watsonia 23: 299-303.
Foley, M.J.Y. 2004. A summary of the past and present status of Spiranthes aestivalis (Poir.) Rich. (Orchidaceae)
(Summer Ladys-Tresses) in north-west Europe. Watsonia 25: 193-201.
Foley, M.J.Y. & Clarke, S. (2005) Orchids of the British Isles. Cheltenham, Gloucs: Griffin Press.
Forrest, A.D., Hollingsworth, M.L., Hollingsworth, P.M., Sydes, C. & Bateman, R.M. 2004. Population genetic
structure in European populations of Spiranthes romanzoffiana set in the context of other genetic studies on
orchids. Heredity. 92: 218-27.
French, C.N., Murphy, R.J. & Atkinson, M.G.C. 1999. Flora of Cornwall. Camborne: Wheal Seaton Press.
Garrard, I. & Streeter, D. 1983. The Wild Flowers of the British Isles. London: Midsummer Books.
Gay, P. & Philip, E. 1999. Early Spider Orchids at Samphire Hoe, Dover. Brit. Wildlife 10: 165.
Gebauer, G. & Meyer, M. 2003. 15N and 13C natural abundance of autotrophic and myco-heterotrophic orchids
provides insight into nitrogen and carbon gain from fungal association. New Phytol. 160: 209-223.
Gendle, A. 2006. Unusual Epipactis on the Cumbrian Limestone. J. Hardy Orchid Soc. 3 (1): 20-21.
Gerard, J. 1633. The Herbal or Generall Historie of Plantes. Facsimile edition (1975) revised by T. Johnson. New
York: Dover.
Gledhill, D. 2002. The Names of Plants. Cambridge: Cambridge University Press.
Goodfellow, G. 1991. The scent of Orchis morio. BSBI News 58: 27.
Graham, R.A. 1953. Epipogium aphyllum Sw. in Buckinghamshire. Watsonia 3: 33.
Green, P.R., Green, I.P. & Crouch, G.A. 1997. The Atlas Flora of Somerset. Somerset: Privately published.
Greenwood, E.F. & Gemmell, R.P. 1978. Derelict industrial land as a habitat for rare plants in S. Lancs. (v.c. 59)
and W. Lancs. (v.c. 60). Watsonia 12: 33-40.
Gulliver, R.L. 1996. The status of Spiranthes romanzoffiana Cham. (Orchidaceae) Irish Ladys-tresses, on
Colonsay, (v.c. 102) in 1995, with special reference to associated plant communities. Watsonia 21: 202-204.
Gulliver, R.L. 1997. Irish Ladys-tresses (Spiranthes romanzoffiana) on Colonsay (v.c. 102). Glasgow Natur. 23:
55-56.
Gulliver, R.L. 2005. Notes on seeds and capsules of Spiranthes romanzoffiana with special reference to the
drawing of a seed in Butchers A New Illustrated British Flora, (1961). BSBI News 99: 35-37.
Gulliver, R., Kiernen, M. Gulliver, M. & Sydes, C. 2000. Observations on Irish Ladys-tresses orchid (Spiranthes
romanzoffiana) on Colonsay (VC 102). Glasgow Natur. 23: 9-12.
Habron, N. & Habron, L. 2007. Cambrensis in Cumbria. J. Hardy Orchid Soc. 4 (2): 65-67.
Hagger, J. 2003. The Early Marsh Orchid (Dactylorhiza incarnata) in Northern Europe. I Introduction. J.
Hardy Orch. Soc. 27: 4-9.
Hagger, J. 2003. The Early Marsh Orchid (Dactylorhiza incarnata) in Northern Europe. II The purple-flowered
early marsh orchids. J. Hardy Orch. Soc. 29: 45-51.
Hagger, J. 2004. The Early Marsh Orchid in Northern Europe. III The British and Irish fen, marsh and bog
forms. J. Hardy Orch. Soc. 31: 18-23.
Hagger, J. 2004. The Early Marsh Orchid (Dactylorhiza incarnata) in Northern Europe. IV Northern forms,
blotched leaves and polymorphism. J. Hardy Orch. Soc. 32: 45-51.
Hagger, J. 2005. The Early Marsh-orchid (Dactylorhiza incarnata) in northern Europe. V Red flowers and
dune forms. J. Hardy Orchid Soc. 2 (2): 51-59.
29/1/09 12:49:39
BIBLIOGRAPHY
473
Hagger, J. 2005. The Early Marsh-orchid (Dactylorhiza incarnata) in northern Europe. VI The significance of
yellow flowers. J. Hardy Orchid Soc. 2 (4): 116-124.
Hagger, J. 2007. The Early Marsh-orchid (Dactylorhiza incarnata) in northern Europe. VII The nominate
variety. J. Hardy Orchid Soc. 4 (2): 52-58.
Hall, P.C. 1980. Sussex Plant Atlas. Brighton: Booth Museum of Natural History.
Halliday, G. 1997. A Flora of Cumbria. Lancaster: University of Lancaster.
Harding, K. & Perring, F. 1959. Spiranthes romanzoffiana Cham. Proc. BSBI 3: 288-289.
Harris, S.A. & Abbott, R.J. 1997. Isozyme analysis of the origin of a new endemic orchid species, Epipactis
youngiana, in the British Isles. Heredity 79: 402-407.
Hazeldon, E., Naisbitt, T. & Richards, A.J. 1991. Differential pollination efficiency within a hybrid swarm
between Dactylorhiza purpurella (T. & T.A. Stephenson) Soo and D. fuchsii (Druce) Soo. Watsonia 18:
391-393.
Hedley, R. No date. The flowering and fruiting performance of Cephalanthera longifolia and the implications for
conservation management. Hampshire Wildlife Trust (unpublished).
Hedley, R. 2003. Biodiversity Action Plan for Cephalanthera longifolia (the Sword-leaved Helleborine).
Hampshire Wildlife Trust.
Henderson, S.A. 2001. The vegetation associated with Spiranthes romanzoffiana Cham. (Orchidaceae), Irish
Ladys-tresses, on the Isle of Coll, Inner Hebrides. Watsonia 23: 493-503.
Heslop-Harrison, J. 1950. Orchis cruenta Mll. in the British Islands. Watsonia 1: 366-375.
Heslop-Harrison, J. 1953. Studies in Orchis L. II. Orchis traunsteineri Saut. in the British Isles. Watsonia 2:
371-391.
Hoare, A.G. 1997. An aberrant form of fly orchid in Surrey. BSBI News 75: 26-27.
Hollingsworth, M. 2003. Taxonomic complexity, population genetics and plant conservation in Scotland. Bot.
J. Scot. 55: 55-63.
Hollingsworth, P.M., Squirrell, J., Hollingsworth, M.L., Richards, A.J. & Bateman, R.M. 2006. Taxonomic
complexity, conservation and recurrent origins of self-pollination in Epipactis (Orchidaceae). Pp. 2744 in
J.P. Bailey & R.G. Ellis (eds.), Current taxonomic research on the British and European flora. London: BSBI.
Horn, C. 1991. Variation in the scent of Orchis morio. BSBI News 57: 42-43.
Horn, C. 1995. Some colour variants of Orchis morio in Bedfordshire. BSBI News 70: 11-12.
Horsman, F. 1989. The history of the recording of Spiranthes romanzoffiana in Britain. BSBI News 53: 18-20.
Horsman, F. 1990a. Peloria in Dactylorhiza. BSBI News 55: 16-18.
Horsman, F. 1990b. Spiranthes romanzoffiana and John Raven. BSBI News 56: 5-7.
Horsman, F. 1990c. On some curious dactylorchids. BSBI News 58: 29-31.
Horsman, F. 1999. The Irish Ladys Tresses Orchid. Coll Magazine 17: 9-11.
Horsman, F. 2005. Some observations on Spiranthes romanzoffiana in the British Isles. BSBI News 99: 37-40.
Hoy, J. 2002. New England Plant Conservation Program. Listera cordata (L.) R. Br. Heart-leaved Twayblade.
Conservation and Research Plan for U.S. Forest Service Region 9.
Hughes, T. 2004. British Ophrys Oddities. J. Hardy Orchid Soc. 1 (4): 111-113.
Hultn, E. & Fries, M. 1986. Atlas of North European vascular plants north of the Tropic of Cancer. Konigstein:
Koeltz Scientific Books.
Hutchings, M.J. 1987a. The population ecology of the Early Spider Orchid Ophrys sphegodes Mill. 1. A
demographic study for 1975 to 1984. J. Ecol. 75: 711-727.
Hutchings, M.J. 1987b. The population biology of the Early Spider Orchid, Ophrys sphegodes Mill. II. Temporal
patterns in behaviour. J. Ecol. 75: 729-742.
Hutchings, M. J., Mendoza, A. & Havers, W. 1998. Demographic properties of an outlier population of Orchis
militaris L. Orchidaceae in England. Bot. J. Linn. Soc. 126: 95-107.
Jenkinson, M.N. 1991. Wild Orchids of Dorset. Gillingham (Dorset): Orchid Sundries.
Jenkinson, M.N. 1995a. Wild Orchids of Hampshire and the Isle of Wight. Gillingham (Dorset): Orchid Sundries.
Jenkinson, M.N. 1995b. A new variety of Narrow-leaved Marsh-orchid in South Hampshire (v.c. 11). Watsonia
20: 263-273.
Jenkinson, M.N. 1996. A new variety of Narrow-leaved Marsh-Orchid in S. Hampshire (v.c.11). BSBI News
72: 64.
29/1/09 12:49:39
474
BIBLIOGRAPHY
Jenkinson, M.N. 1997. Marsh-orchid controversy a small or large argument. BSBI News 77: 29-31.
Jenkinson, M.N. & Hobson, A.G. 1989. Epipactis purpurata Sm. reappears in Dorset. Watsonia 17: 441-442.
Jermyn, S.T. 1974. Flora of Essex. Colchester: Essex Naturalists Trust.
Jones, S. 1998. Aspects of the population biology of Liparis loeselii (L.) Rich. var. ovata Ridd. Ex Godfrey
(Orchidaceae) in the dune slacks of South Wales, UK. Bot. J. Linn. Soc. 126: 123-139.
Julou, T., Burghardt, B., Gebauer, G., Berveiller, D., Damesin, C. & Selosse, M-A. 2005. A case for mixotrophy
in orchids: insights from a comparative study of green and achlorophyllous Caphalanthera damasonium. In
press.
Kemp, R.J. 1987. Reappearance of Orchis purpurea Hudson in Oxfordshire. Watsonia 16: 435-36.
Kendrick, F.M. 1983. Botany. Trans. Woolhope Naturalists Field Club 44: 124-125.
Kenneth, A.G. & Tennant, D.J. 1984. Dactylorhiza incarnata (L.) Soo subsp. cruenta (O. F. Mueller): D. Sell in
Scotland. Watsonia 15: 11-14.
Kenneth, A.G. & Tennant, D.J. 1987. Further notes on Dactylorhiza incarnata subsp. cruenta in Scotland.
Watsonia 16: 332-334.
Kenneth, A.G., Lowe, M.R. & Tennant, D.J. 1988. Dactylorhiza lapponica (Laest. ex Hartman) Soo in Scotland.
Watsonia 17: 37-41.
Kent, D.H. 1954. Plant notes. 675/1 Cypripedium calceolus L. Proc. BSBI 1: 39-40.
Killick, J., Perry, R. & Woodell, S. 1998. The Flora of Oxfordshire. Newbury: Pisces.
Kreutz, C.A.J. 2007. Epipactis dunensis (T. & T.A. Stephenson) Godfrey subsp. tynensis Kreutz, eine neue
Epipactis-Sippe aus Nordengland und Sdschottland. J. Eur. Orch. 39 (1): 123-134.
Kreutz, C.A.J. 2008. Update on British Orchids. J. Hardy Orchid Soc. 5 (1): 11-16.
Kull, T. 1999. Biological Flora of the British Isles. No. 208. Cypripedium calceolus L. J. Ecol. 87: 913-924.
Kull, T. & Kull, K. 1991. Preliminary results from a study of populations of Cypripedium calceolus in Estonia.
In T.C.E. Wells & J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague: SPB Academic
Publishing.
Lacey, W.S. 1955. Orchis traunsteineri Saut. in Wales. Proc. BSBI 1: 297-300.
Land, R. No date. Management guidelines for Fen Orchid. Norfolk Wildlife Trust (unpublished).
Laney, B. & Stanley, P. 2004. Ophrys apifera (Bee Orchid) in Ayrshire (VC75). J. Hardy Orch. Soc. 32: 43.
Lang, D. 1980. Orchids of Britain. Oxford: Oxford University Press.
Lang, D.C. 1991. A new variant of Ophrys apifera Hudson in Britain. Watsonia 18: 408-410.
Lang, D.C. 2001. Wild Orchids of Sussex. Lewes: Pomegranate Press.
Lang, D.C. 2004. Britains Orchids. Old Basing (Hampshire): WildGuides.
Laurence, R.J. 1986. Ophrys apifera Hudson subsp. jurana Ruppert found in Britain. Watsonia 16: 177-178.
Laurence, R. & Chalk, M. 2004. Hyperchromic form of Dactylorhiza maculata found in the New Forest. BSBI
News 96: 28-29.
Lewis, L. 2005. Unusual forms of the Fly Orchid, Ophrys insectifera. J. Hardy Orchid Soc. 2 (3): 81-88.
Lewis, L. 2008. Naming new orchid taxa: a reply but not an answer. J. Hardy Orchid Soc. 5 (4): 138-143.
Lewis, L. & Spencer, E.J. 2005. Epipactis phyllanthes var. cambrensis (C.A. Thomas) P.D. Sell and other unusual
Epipactis at Kenfig National Nature Reserve. Watsonia 25: 290-295.
Light, M.H.S. & MacConaill, M. 1991. Patterns of appearance in Epipactis helleborine (L.) Crantz. In T.C.E. Wells
& J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague: SPB Academic Publishing.
Lindop, M. 1992. Cypripedium conservation 1991 report. BSBI News 60: 45.
Lindop, M. 1994. Cypripedium conservation report 1993. BSBI News 66: 30.
Lord, R.M., Richards, A.J. 1977. A hybrid swarm between the diploid Dactylorhiza fuchsii (Druce) Soo and the
tetraploid D. purpurella (T. & T.A. Steph.) Soo in Durham. Watsonia 11: 205-210.
Lousley, J.E. 1969. Wild Flowers of Chalk and Limestone. London: Collins.
Lousley, J.E. 1976. Flora of Surrey. Newton Abbot: David & Charles.
Lowe, M.R., Tennant, D.J. & Kenneth, A.G. 1986. The status of Orchis Francis-Drucei Wilmott. Watsonia 16:
178-180.
Mabey, R. 1996. Flora Britannica. London: Sinclair-Stevenson.
29/1/09 12:49:40
BIBLIOGRAPHY
475
McClintock, D. 1975. The Wildflowers of Guernsey. London: Collins.

McKean, D.R. 1996. A chlorotic Epipactis helleborine in an Edinburgh housing estate. BSBI News 72: 35.
McKendrick, S.L., Leake, J.R. & Read, D.J. 2000. Symbiotic germination and development of myco-heterotrophic
plants in nature: Transfer of carbon from ectomycorrhizal Salix repends and Betula pendula to the orchid
Corallorhiza trifida through shared hyphal connections. New Phytol. 145: 539-548.
McKendrick, S.L., Leake, J.R., Taylor, D.L. & Read, D.J. 2000. Symbiotic germination and development of
myco-heterotrophic plants in nature: Ontogeny of Corallorhiza trifida and characterization of its mycorrhizal
fungi. New Phytol. 145: 523-537.
McKendrick, S.L., Leake, J.R., Taylor, D.L. & Read, D.J. 2002. Symbiotic germination and development of the
myco-heterotrophic orchid Neottia nidus-avis in nature and its requirement for locally distributed Sebacina
spp. New Phytol. 154: 233-247.
Madge, S. 1994. The status of Serapias parviflora Parl. in Britain. Botanical Cornwall, 6: 51-52.
Madge, S. 1995. A new orchid for Britain. Carradon Wildlife 11: 10.
Margetts, L.J. & David, R.W. 1981. A Review of the Cornish Flora 1980. Redruth: Institute of Cornish Studies.
Margetts, L.J. & Spurgin, K.L. 1991. The Cornish Flora Supplement 1981-1990. Zennor, St Ives: Trendine
Press.
Marren, 1999. Britains Rare Flowers. London: T & AD Poyser.
Merryweather, J. 2001. Comment. Meet the Glomales - the ecology of mycorrhiza. Brit. Wild. 13: 86-93.
Moore, S. 1997. Fly orchid aberration? Another record. BSBI News 76: 21-22.
Mrkvicka, A.C. 1992. Keimung, Entwicklung und Wachstumezyklen von Liparis loeselii (L.) L.C. Rich. am
natrlichen Wuchsort. Orchidee (Hamburg) 43: 35-36.
Murphy, R.J. 1994. Progress report. Botanical Cornwall, 6: 1-7.
Murphy, R.J. 1995. Hammarbya paludosa (Bog Orchid) refound in Cornwall. BSBI News 69: 74-75.
Neiland, M.R.M. 1994. Reproductive ecology of Dactylorhiza lapponica in Scotland. Edinburgh: Scottish Natural
Heritage.
Neiland, M.R.M. and Wilcock, C.C. 1994. Survey of Dactylorhiza lapponica. Edinburgh: Scottish Natural
Heritage.
Neiland, M.R.M. & Wilcock, C.C. 1995. Reproductive Biology and Morphological Variability of Two Rare Scottish
Orchids, Dactylorhiza lapponica and Dactylorhiza traunsteineri. Edinburgh: Scottish Natural Heritage.
Pankhurst, R.J. & Mullin, J.M. 1991. Flora of the Outer Hebrides. London: Natural History Museum.
Paul, V.N. 1964. Epipogium aphyllum. Reading Naturalist 16: 29-30.
Payne, R. 1991. Orchis morio in old Cambridgeshire. Nat. Camb. 33: 23-25.
Pearman, D.A. & Preston, C.D. 2000. A Flora of Tiree, Gunna and Coll. Dorchester: Privately published.
Perring, F.H. 1956. Spiranthes spiralis (L.) Chevall. in Britain, 1955. Proc. BSBI 2: 6-9.
Philip, E.G. 1982. Atlas of the Kent Flora. Kent: Kent Field Club.
Pillon, Y., Qamaruz-Zaman, F., Fay, M.F., Hendoux, F. & Piquot, Y 2006. Genetic diversity and ecological
differentiation in the endangered fen orchid (Liparis loeselii). Conservation Genetics 8: 177-184.
Preston, C.D. 2000. Engulfed by suburbia or destroyed by the plough; the ecology of extinction in Middlesex
and Cambridgeshire. Watsonia 23: 59-81.
Preston, C.D., Pearman, D.A. & Dines, T.D. 2002. New Atlas of the British and Irish Flora. Oxford: Oxford
University Press.
Pridgeon, A.M., Bateman, R.M., Cox, A.V., Hapeman, J.R. & Chase, M.W. 1997. Phylogenetics of subtribe
Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 1. Intergeneric relationships and
polyphyly of Orchis sensu lato. Lindleyana 12: 89-109.
Qamaruz-Zaman, F., Fay, M.F., Parker, J.S. & Chase, M.W. 1998. The use of AFLP fingerprinting in conservation
genetics: a case study of Orchis simia (Orchidaceae). Lindleyana 13: 125-133.
Ramsay, M.M. and Stewart, J. 1998. Re-establishment of the Ladys Slipper Orchid (Cypripedium calceolus L.)
in Britain. Bot. J. Linn. Soc. 126: 173-181.
Rasmussen H.N. 1995. Terrestrial Orchids from Seed to Mycotrophic plant. Cambridge: Cambridge University
Press.
Reinhammar, L.G. 1998. Systematics of Pseudorchis albida sl (Orchidaceae) in Europe and North America. Bot.
J. Linn. Soc. 126: 363-382.
29/1/09 12:49:40
476
BIBLIOGRAPHY
Rich, T. 1994. Narrow-leaved Helleborine (Cephalanthera longifolia) in Surrey. PlantLife Back from the Brink
Project Report, 38.
Rich, T.C.G. & Jermy, A.C. 1998. Plant Crib 1998. London: BSBI.
Richards, A.J. 1986. Cross-pollination by wasps in Epipactis leptochila (Godf.) Godf. S.L. Watsonia 16: 180182.
Richards, A.J. 1989. Some problematic Epipactis in northern Britain. BSBI News 51: 51.
Richards, A.J. & Porter, A.F. 1982. On the identity of a Northumberland Epipactis. Watsonia 14: 121-128.
Richards, A.J. & Swan, G.A. 1976. Epipactis leptochila (Godfery) Godfery and E. phyllanthus G.E. Sm occurring
in South Northumberland on lead and zinc soils. Watsonia 11: 1-5.
Riddelsdell, H.J., Hedley, G.W. & Price, W.R. 1948. The Flora of Gloucestershire. Bristol: Chatford House.
Roberts, R.H. 1959. Notes on the fen habitat of Ophrys insectifera in Anglesey. Proc. BSBI 3: 274-278.
Roberts, R.H. 1961. Studies on Welsh orchids. I. The variation of Dactylorchis purpurella (T. & T. A. Steph.)
Vermeul. in North Wales. Watsonia 5: 23-36.
Roberts, R.H. 1961. Studies on Welsh orchids. II. The occurence of Dactylorchis majalis (Reichb.) Vermeul. in
Wales. Watsonia 5: 37-42.
Roberts, R.H. 1966. Studies on Welsh orchids. III. The coexistence of some of the tetraploid species of marsh
orchid. Watsonia 6: 260-267.
Roberts, R.H. 1988. The occurrence of Dactylorhiza traunsteineri (Sauter) Soo in Britain and Ireland. Watsonia
17: 43-47.
Roberts, R.H. 1989. Errors and misconceptions in the study of Marsh-orchids. Watsonia 17: 455-462.
Roberts, R.H. 1997. A new variety of Fly Orchid in Angelsey. BSBI News 74: 24.
Roberts, R.H. & Foley, M.J.Y. 1997. The taxonomic status of Dactylorhiza majalis (Rchb. fil.): F. Hunt &
Summerh. subsp. traunsteineri (Saut. & Rchb. fil.) H. Sund. var. bowmanii M.N. Jenk. Watsonia 21: 374-376.
Roberts, R.H. & Gilbert, O.L. 1963. The status of Orchis latifolia var. eborensis Godfery in Yorkshire. Watsonia
5: 287-293.
Rose, F. 1949. Orchis purpurea. J. Ecol. 36: 366-77.
Rose, F. 1960. The rediscovery of Orchis simia Lam. in Kent, 1. Foreword. Trans. Kent Field Club 1: 50.
Rose, F. 1981. The Wild Flower Key. London: Frederick Warne.
Rose, F. 1991. A new subspecies of Gymnadenia conopsea (L.) R. Br. Watsonia 18: 319-320.
Rose, F. 1998a. Gymnadenia conopsea (L.) R. Br. In T.C.G Rich & A.C. Jermy (eds), Plant Crib 1998. London:
BSBI.
Rose, F. 1998b. A new orchid hybrid for Britain x Orchiaceras melsheimeri (Aceras anthropophora x Orchis
purpurea). BSBI News 79: 19.
Rose, F. & Brewis, A. 1988. Cephalanthera rubra (L.) Rich. in Hampshire. Watsonia 17: 176-177.
Rose, F. & Davey, S.R. 1996. The three forms of Fragrant Orchid. BSBI News 72: 71.
Sanford, M. 1991. The Orchids of Suffolk. Ipswich: Suffolk Naturalists Society.
Sanger, N. & Waite, S. 1998. The phenology of Ophrys sphegodes (the early spider orchid): what annual censuses
can miss. Bot. J. Linn. Soc. 126: 75-81.
Sankey, A. 2000. The Box Hill Book of Orchids. Dorking: Friends of Box Hill.
Scobie, A.R. 2007. Evidence of pollination and seed set in Scottish populations of Spiranthes romanzoffiana.
BSBI News 106: 9-12.
Scott, W., Harvey, P., Riddington, R. & Fisher, M. 2002. Rare Plants of Shetland. Lerwick: Shetland Amenity
Trust.
Scott, W. & Palmer, R. 1987. The Flowering Plants and Ferns of the Shetland Islands. Lerwick: Shetland Times.
Sell, D. & Murrell, G. 1996. Flora of Great Britain, Ireland, Isle of Man and the Channel Islands. Vol. 5: Butomaceae
Orchidaceae. Cambridge: Cambridge University Press.
Selosse, M-A., Faccio, A., Scappaticci, G. & Bonfante, P. 2004. Chlorophyllous and achlorophyllous specimens of
Epipactis microphylla (Neottieae, Orchidaceae) are associated with ectomycorrhizal septomycetes, including
truffles. Microb. Ecol. 47: 416-426.
Selosse M-A., Weiss, M., Jany, J-L. & Tillier A. 2002. Communities and populations of sebacinoid basidiomycetes
associated with the achlorophyllous orchid Neottia nidus-avis (L.) L.C.M. Rich. and neighbouring tree
ectomycorrhizae. Mol. Ecol. 11: 1831-1844.
29/1/09 12:49:40
BIBLIOGRAPHY
477
Sex, S. & Sayers, B. 2004. Irelands Wild Orchids. Portmarnock: Susan Sex Wild Orchids.
Shefferson R.P. & Tali K. 2007. Dormancy is associated with decreased adult survival in the burnt orchid,
Neotinea ustulata. J. Ecology 95: 217225.
Silvertown, J., Wells, D.A., Gillman, M., Dodd, M.E., Robertson, H., Kakhani, K.H. 1994. Short-term effects
and long-term effects of fertilizer application on a flowering population of Green-Winged Orchid, Orchis
morio. Biol. Cons. 69: 191-197.
Sinker, C.A., Packham, J.R., Trueman, I.C., Oswald, P.H., Perring, F.H. & Prestwood, W.V. 1991. Ecological
Flora of the Shropshire Region. Shrewsbury: Shropshire Wildlife Trust.
Socit Franaise dOrchidophilie 1998. Les Orchides de France, Belgique et Luxembourg. Paris: Collection
Parthnope.
Squirrell, J., Hollingsworth, M., Bateman, R.M., Tebbitt, M.C. and Hollingsworth, M.L. 2002. Taxonomic
complexity and breeding system transitions: conservation genetics of the Epipactis leptochila complex
(Orchidaceae). Mol. Ecol. 11: 1957-1964.
Stace, C.A. 1991. New Flora of the British Isles. Cambridge: Cambridge University Press.
Stace, C.A. 2004. Interactive Flora of the British Isles (CD-ROM). Amsterdam: ETI Bioinformatics.
Stace, C.A., Ellis, R.G., Kent, D.H. & McCosh, D.J. 2003. Vice-county Census Catalogue of the Vascular Plants of
Great Britain. London: BSBI.
Steel, D.T. & Creed, C. 1982. Wild Orchids of Berkshire, Buckinghamshire and Oxfordshire. Oxford: Pisces.
Stewart, A., Pearman, D.A. & Preston, C.D. 1994. Scarce Plants in Britain. Peterborough: Joint Nature
Conservation Commitee.
Stone, D.A. & Russell, R.V. 2000. Population biology of late spider orchid Ophrys fucifera A study at Wye
National Nature Reserve 1987-1998. English Nature Research Report 389.
Summerhayes, V.S. 1968. Wild Orchids of Britain. London: Collins.
Sumpter, J.P., DAyala, R., Parfitt, A.J., Pratt, P. & Raper, C. 2004. The current status of Military (Orchis militaris)
and Monkey (Orchis simia) Orchids in the Chilterns. Watsonia 25: 175-183.
Synge, H. (ed.) 1981. The Biological Aspects of Rare Plants Conservation. London: John Wiley.
Tali, K., Foley, M.J.Y. & Kull, T. 2004. Biological Flora of the British Isles. 232. Orchis ustulata L. J. Ecol. 92:
174-184.
Tali, K., Fay, M.F., & Bateman, R.M. 2006. Little genetic differentiation across Europe between early-flowering
and late-flowering populations of the rapidly declining orchid Neotinea ustulata. Biol. J. Linn. Soc. 87: 1325.
Tam, C.O. 1991. Behaviour of some orchid populations in a changing environment: Observations on permanent
plots, 1943-1990. In T.C.E. Wells & J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The
Hague: SPB Academic Publishing.
Taylor, D.L. & Bruns, T. 1997. Independent, speciliazed invasions of ectomycorrhizal mutualism by two
nonphotosynthetic orchids. Proc. Natl. Acad. Sci. 94: 4510-4515.
Tennant, D.J. & Kenneth, A.G. 1983. The Scottish records of Dactylorhiza traunsteineri (Sauter) Soo. Watsonia
14: 415-417.
Thomas, C. 1950. The Kenfig Epipactis. Watsonia 1: 283-288.
Thomas, S. 1997. Orchids from the 18th-century herbarium of Joseph Andrews (1688-1764). BSBI News 76:
17-21.
Tyteca, D. & Klein, E. 2008. Genes, morphology and biology the systematics of Orchidinae revisited. J.
Europischer Orchideen 40: 501544.
UK Biodiversity Group 1999. Tranche 2 Action Plans, Vol. III: Plants and Fungi. Peterborough: English
Nature.
Van Der Cingel, N.A. 1995. An Atlas of Orchid Pollination: European Orchids. Rotterdam: Balkema.
Vanhecke, L.E.M. 1991. Population dynamics of Dactylorhiza praetermissa in relation to topography and
inundation. In T.C.E. Wells & J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague:
SPB Academic Publishing.
Waite, S. and Farrell, L. 1998. Population biology of the rare military orchid (Orchis militaris L.) at an established
site in Suffolk. Bot. J. Linn. Soc. 126: 109-121.
Waite, S. & Hutchings, M.J. 1991. The effects of different management regimes on the population dynamics
29/1/09 12:49:41
478
BIBLIOGRAPHY
of Ophrys sphegodes. Analysis and description using matrix models. In T.C.E. Wells & J.H. Willems (eds),
Population Ecology of Terrestrial Orchids. The Hague: SPB Academic Publishing.
Wells, T.C.E. 1967. Changes in a population of Spiranthes spiralis (L.) Chevall. at Knocking Hoe National
Nature Reserve, Bedfordshire, 1962-65. J. Ecol. 55: 83-99.
Wells, T.C.E. 1981. Population ecology of terrestrial orchids. In Synge, H. (ed.) Biological Aspects of Rare Plant
Conservation. London: John Wiley.
Wells, T.C.E & Cox, R. 1991. Demographic and biological studies on Ophrys apifera: some results from a ten
year study. In T.C.E. Wells & J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague: SPB
Academic Publishing.
Wells, T.C.E. & Kretz, R. 1986. Spiranthes spiralis (L.) Cheval. from seed to flowering plant in five years.
Watsonia 16: 235.
Wells, T.C.E., Rothery, P., Cox, R. & Bamford, S. 1998. Flowering dynamics of Orchis morio L. and Herminium
monorchis (L.) R. Br. at two sites in Eastern England. Bot. J. Linn. Soc. 126: 39-48.
Wells, T.C.E. & Willems, J.H. (eds) 1991. Population Ecology of Terrestrial Orchids. The Hague: SPB Academic
Publishing.
Weston, R.P. 1983. A Lincolnshire Epipactis. Watsonia 14: 457-458.
Wheeler, B. 1998. Cephalanthera longifolia in England, 1997. Plantlife Report 89.
Wheeler, B.D., Lambley, W. & Geeson, J. 1998. Liparis loeselii (L.) Rich. in eastern England: Constraints on
distribution and population development. Bot. J. Linn. Soc. 126: 141-158.
Wigginton, M.J. (ed.) 1999. British Red Data Books 1: Vascular Plants. Peterborough: Joint Nature Conservation
Committee.
Wilks, H.M. 1960. The rediscovery of Orchis simia Lam. in Kent 2: Orchis simia in East Kent. Trans. Kent Field
Club 1: 50-55.
Willems, J.H. & Bik, L. 1991. Long-term dynamics in a population of Orchis simia in the Netherlands. In
T.C.E. Wells & J.H. Willems (eds), Population Ecology of Terrestrial Orchids. The Hague: SPB Academic
Publishing.
Willems, J.H., Melser, C. 1998. Population dynamics and life-history of Coeloglossum viride (L.) Hartm.: an
endangered orchid species in The Netherlands. Bot. J. Linn. Soc. 126: 83-93.
Williams, J.G., Williams, A.E. & Arlott, N. 1978. A Field Guide to the Orchids of Britain and Europe with North
Africa and the Middle East. London: Collins.
Willis, A.J. 1967. A new locality for Liparis loeselii. Proc. BSBI 6: 352-353.
Willis, A.J. 1980. Ophrys apifera Huds. x O. insectifera L., a natural hybrid in Britain. Watsonia 13: 97-102.
Willis, A.J., Martin, M.H. & Taylor, K.B. 1991. Orchis purpurea Hudson in the Avon Gorge, Bristol. Watsonia
18: 387-390.
Wilson, D.E. & Ingram, G.L. 1994. Distribution of Lady Orchid (Orchis purpurea Hudson) in an East Kent
wood. Watsonia 20: 154-156.
Wood, J. & Ramsey, M. 2004. Anacamptis laxiflora Orchidaceae. Curtiss Bot. Mag. 21: 26-33.
Young, D.P. 1949a. Studies in the British Epipactis. I. Epipactis dunensis and E. pendula. Watsonia 1: 102-108.
Young, D.P. 1949b. Studies in the British Epipactis. II. The differentiation of E. pendula from E. vectensis.
Young, D.P. 1952a. Studies in the British Epipactis. III. Epipactis phyllanthes G. E. Sm., an overlooked species.
Young, D.P. 1952b. Studies in the British Epipactis. IV. A revision of the phyllanthes-vectensispendula group.
Young, D.P. 1962a. Studies in the British Epipactis. V. Epipactis leptochila; with some notes on E. dunensis and E.
mulleri. Watsonia 5: 127-135.
Young, D.P. 1962b. Studies in the British Epipactis. VI. Some further notes on E. phyllanthes. Watsonia 5: 136139.
29/1/09 12:49:41
INDEX
479
Index of Species
aestivalis, Spiranthes 188
albida, Gymnadenia 236
albida, Leucorchis 236
albida, Pseudorchis 236
alpina, Chamorchis 416
AnACAmPtiS 361
anthropophora, Orchis 202
anthropophorum, Aceras 202
aphyllum, Epipogium 136
apifera, Ophrys 398
atrorubens, Epipactis 85
Autumn Ladys-tresses 181
Bertolonis Mirror Orchid 416
bertolonii, Ophrys 416
Bee Orchid 398
bifolia, Platanthera 243
Birds-nest Orchid 67
Bog Adders-mouth 152
Bog Orchid 152
borealis, Gymnadenia 263
Bracted Green Orchis 288
Broad-leaved Helleborine 92
Burnt-tip Orchid 345
Burnt Orchid 345
calceolus, Cypripedium 25
cambrensis, Dactylorhiza purpurella 332
CEPHALAntHERA 32
Chalk Fragrant Orchid 256
chlorantha, Platanthera 249
coccinea, Dactylorhiza incarnata 276
Common Fragrant Orchid 256
Common Spotted Orchid 294
Common Twayblade 62
conopsea, Gymnadenia 256
CORALLORHiZA 159
Coralroot Orchid 160
cordata, Listera 56
cordata, neottia 56
cordigera, Serapias 416
Creeping Ladys-tresses 166
cruenta, Dactylorhiza incarnata 276
CYPRiPEDiUm 23
Dutch Helleborine 97
DACtYLORHiZA 272
damasonium, Cephalanthera 48
Dark-red Helleborine 85
Dense-flowered Orchid 341
densiflora, Gymnadenia 267
densiflora, Gymnadenia borealis 263
densiflora, Gymnadenia conopsea 267
dunensis, Epipactis 113

Dune Helleborine 113
Dwarf Alpine Orchid 416
Early Coralroot 160
Early Marsh Orchid 276
Early Purple Orchid 228
Early Spider Orchid 411
ebudensis, Dactylorhiza 325
EPiPACtiS 73
EPiPOGiUm 135
ericetorum, Dactylorchis 303
ericetorum, Orchis 303
False Musk Orchid 416
Fen Orchid 142
Fly Orchid 392
Frivalds Fragrant Orchid 416
frivaldii, Gymnadenia 416
Frog Orchid 288
fuchsii, Dactylorhiza 294
fuciflora, Ophrys 405
fuschii, Dactylorchis 294
fuschii, Orchis 294
Ghost Orchid 136
GOODYERA 165
Greater Butterfly Orchid 249
Greater Tongue Orchid 384
Green-flowered Helleborine 126
Green-veined Orchid 372
Green-winged Orchid 372
GYmnADEniA 254
HAmmARBYA 151
Heart-flowered Tongue Orchid 416
Heart-leaved Twayblade 56
Heath Fragrant Orchid 263
Heath Spotted Orchid 303
Hebridean Marsh Orchid 325
helleborine, Epipactis 92
HERminiUm 192
HimAntOGLOSSUm 353
hircinum, Himantoglossum 354
Hooded Ladies-tresses 174
incarnata, Dactylorhiza 276
incarnata, Dactylorhiza incarnata 276
insectifera, Ophrys 392
Irish Ladys-tresses 174
Irish Marsh Orchid 334
Jersey Orchid 363
Lapland Marsh Orchid 321
Ladys-slipper 25
Lady Orchid 222
29/1/09 12:49:42
480
INDEX
lapponica, Dactylorhiza traunsteineri 321

Late Spider Orchid 405
latifolia, Dactylorchis 276
latifolia, Orchis 276
laxiflora, Anacamptis 363
laxiflora, Orchis 363
leptochila, Epipactis 107
Lesser Butterfly Orchid 243
Lesser Rattlesnake Orchid 166
Lesser Rattlesnake Plantain 166
Lesser Twayblade 56
Lindisfarne Helleborine 122
lingua, Serapias 384
LiPARiS 141
Lizard Orchid 354
Loesels Twayblade 142
loeselii, Liparis 142
longifolia, Cephalanthera 41
Loose-flowered Orchid 363
maculata, Dactylorhiza 303
maculata, neotinea 341
majaliformis, Dactylorhiza purpurella 332
Man Orchid 202
Marsh Fragrant Orchid 267
Marsh Helleborine 78
mascula, Orchis 228
militaris, Orchis 214
Military Orchid 214
Monkey Orchid 207
monorchis, Herminium 193
morio, Anacamptis 372
morio, Orchis 372
muelleri, Epipactis 107, 113
muscifera, Ophrys 392
Musk Orchid 193
Narrow-leaved Helleborine 41
Narrow-leaved Marsh Orchid 316
Narrow-lipped Helleborine 107
neerlandica, Epipactis helleborine 97
neglecta, Serapias 416
nEOtinEA 340
nEOttiA 54
Newfoundland Orchid 236
nidus-avis, neottia 67
Northern Marsh Orchid 328
occidentalis, Dactylorchis 334
occidentalis, Dactylorhiza 334
occidentalis, Dactylorhiza comosa 334
occidentalis, Dactylorhiza majalis 325, 334
occidentalis, Orchis 334
ochroleuca, Dactylorhiza incarnata 276
odoratissima, Gymnadenia 416
OPHRYS 387
ORCHiS 199
ovata, Listera 62
ovata, neottia 62
Pale-flowered Orchid 416
pallens, Orchis 416
paludosa, Hammarbya 152
paludosa, malaxis 152
palustris, Epipactis 78
parviflora, Serapias 381
pendula, Epipactis 126
Pendulous-flowered Helleborine 126
phyllanthes, Epipactis 126
PLAtAntHERA 241
praetermissa, Dactylorchis 309
praetermissa, Dactylorhiza 309
praetermissa, Orchis 309
PSEUDORCHiS 235
Pugsleys Marsh Orchid 316
pulchella, Dactylorhiza incarnata 276
purpurata, Epipactis 101
purpurea, Orchis 222
purpurella, Dactylorchis 328
purpurella, Dactylorhiza 328
purpurella, Orchis 328
pyramidalis, Anacamptis 366
Pyramidal Orchid 366
Red Helleborine 34
repens, Goodyera 166
romanzoffiana, Spiranthes 174
rubra, Cephalanthera 34
sancta, Epipactis 122
Scarce Tongue Orchid 416
SERAPiAS 378
Short-spurred Fragrant Orchid 416
simia, Orchis 207
Small-flowered Tongue Orchid 381
Small White Orchid 236
Southern Marsh Orchid 309
sphegodes, Ophrys 411
spiralis, Spiranthes 181
SPiRAntHES 172
straminea, Pseudorchis 236
Summer Ladys-tresses 188
Sword-leaved Helleborine 41
Tyne Helleborine 118
traunsteineri, Dactylorhiza 316
traunsteinerioides, Dactylorhiza 316
trifida, Corallorhiza 160
ustulata, neotinea 345
ustulata, Orchis 345
vectensis, Epipactis 126
Violet Helleborine 101
viride, Coeloglossum 288
viridis, Dactylorhiza 288
White Helleborine 48
Youngs Helleborine 97
youngiana, Epipactis 97
29/1/09 12:49:42

Orchids of Britain and Ireland

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Orchids of Britain and Ireland

Uploaded by

Copyright:

Available Formats

Orchids

of Britain and Ireland

000 prelims intro.indd 1

000 prelims intro.indd 2

Anne and Simon Harrap

000 prelims intro.indd 3

Published in 2005 by A&C Black Publishers Ltd,

000 prelims intro.indd 5

SOURCES OF INFORmATION AND BIBLIOGRApHY

000 prelims intro.indd 6

M A typical orchid flower.

000 prelims intro.indd 7

ORCHIdS OF BRITAIn And IRELAnd

000 prelims intro.indd 8

Fungi form several kinds of mycorrhiza:

000 prelims intro.indd 9

ORCHIdS OF BRITAIn And IRELAnd

fungi and rare orchids

germinAtion And underground growth

000 prelims intro.indd 10

cross-pollination versus self-pollination

000 prelims intro.indd 11

ORCHIdS OF BRITAIn And IRELAnd

000 prelims intro.indd 12

000 prelims intro.indd 13

ORCHIdS OF BRITAIn And IRELAnd

000 prelims intro.indd 14

marshes and fens

000 prelims intro.indd 15

ORCHIdS OF BRITAIn And IRELAnd

heaths and moors

Pits, quarries and railway lines

000 prelims intro.indd 16

road verges, green lanes, churchyards and lawns

dunes and dune slacks

000 prelims intro.indd 17

ORCHIdS OF BRITAIn And IRELAnd

000 prelims intro.indd 18

what you can do

orchids and the law

000 prelims intro.indd 19

ORCHIdS OF BRITAIn And IRELAnd

introductions to the genera

000 prelims intro.indd 20

000 prelims intro.indd 21

ORCHIdS OF BRITAIn And IRELAnd

Variation and varieties

Name and classification

history and conservation

000 prelims intro.indd 22

produces two buds at its tip: the larger will

O 31 May, Khabarovsk, eastern Sibera. Ladys-slipper

Critically Endangered, BAP

north-facing slope. They nestle in a sheltered

M 30 May, Lancashire (Sean Cole). This plant, probably

Confined to a single site in Yorkshire,

northern Mongolia, northeast China, Korea,

fleshy stigma, which lies on the lower part of

Variation and varieties

picking up a load of pollen in the process. The

Development and growth

seedlings at the base of the mother plant.

Name and classification

Past and present occurrence of Ladys-slipper in Britain