Mediterranean Temporary Pools Volume 2 ... - Desde el Sekano

Mediterranean 

Temporary Pools 

Volume 2 

Species information sheets 

Grillas P., P. Gauthier, N. Yavercovski & C. Perennou

Production: Station biologique de la Tour du Valat 

Design: Tapages Publics 

Drawings: Antoine Catard, except pages 55, 83, 85, 88, 91 and 118 

Translated from French by Janet Clayton and John Phillips 

Cover: photos Jean Roché 

© 2004 Station biologique de la Tour du Valat 

Le Sambuc - 13200 Arles - France 

Readers are invited to reproduce texts and drawings featured in this publication 

provided credit is given to the texts and drawings authors and to the Station Biologique de la Tour du Valat. 

All photos rights reserved. No photographic part of this work may be reproduced or transmitted 

in any form or by any means, electronic or mechanical, including photocopying 

except as may be expressly permitted in writing from the publisher. 

ISBN : 2-9103-6850-5

Mediterranean 

Temporary Pools 

volume 2 

Species information sheets 

Editors: Grillas P., P. Gauthier, N. Yavercovski & C. Perennou 

Associate editors: Jakob C., H. Michaud, G. Paradis & L. Rhazi

Editors, associate editors, authors and collaborators 

Antonetti P. (CNBMC), Calvière T. (TDV), Catard A (CEEP Var), Cheylan M. 

(EPHE), Delaugerre (CELRL Corse), Fuselier J. (ADENA), Garnéro S. (CEN-LR), 

Garraud L. (CBNA), Gauthier P. (TDV), Gendre T. (CEN-LR), Grillas P. (TDV), 

Hébrard J. P. (Université d’Aix-Marseille III-IMEP), Houssard C. (CEN-LR), 

Hugonnot V. (ad. perso. 1), Jakob C. (EPHE and TDV), Klewscewski M. (CEN-LR), 

Lombardini K. (EPHE), Michaud H. (CBNMP), Molina J. (CBNM-LR), Papazian 

M. (SOF), Paradis G. (Université de Corse, ad. perso. 2), Perennou C. (TDV) Pozzo 

di Borgo M. L. (OEC), Rhazi L. (université Hassan II), Rhazi M.(TDV and IMEP 

Université d’Aix-Marseille III), Rombaut D. (CEEP Var), Ruchon F. (AGRN-RH), 

Samraoui B. (Université d’Annaba), Souheil H. (AGRN-RH), Soulié-Märsche I. 

(Université Montpellier II, CNRS), Thiéry A. (Université de Provence, Aix- 

Marseille I), Titolet D. (Lycée international Georges Duby), Yavercovski N. (TDV). 

ADENA (Fuselier J.) 

Association de Défense de la Nature des Pays d’Agde 

Domaine du grand Clavelet, F-34300 Agde 

Tél.: +33 (0)4 67 01 60 23, fax: +33 (0)4 67 01 60 29 

adena-bagnas@tiscali.fr 

AGRN.RH (Ruchon F., Souheil H.) 

Association de Gestion de la Réserve Naturelle de Roque-Haute, 

1, rue de la Tour, F-34420 Portiragnes 

Tél/fax: +33 (0)4 67 90 81 16 

roque.haute@espaces-naturels.fr 

CBNA (Garraud L.) 

Conservatoire Botanique National Alpin de Gap-Charance 

Domaine de Charance, F-05000 Gap 

Tél: +33 (0)4 92 53 56 82, fax: +33 (0)4 92 51 94 58 

cbn-gap@wanadoo.fr; l.garraud@cbn-alpin.org 

CBNMC (Antonetti P.) 

Conservatoire Botanique National du Massif central 

Le Bourg, F-43230 Chavaniac-Lafayette 

Tél: +33 (0)4 71 77 55 65, fax: +33 (0)4 71 77 55 74 

cbnmc@mail.es-conseil.fr 

CBNMP (Michaud H, Molina J.) 

1. siège Conservatoire botanique national de Porquerolles 

Castel Sainte Claire, F-83418 Hyères cedex 

Tél: +33 (0)4 94 12 82 30, fax: +33 (0)4 94 12 82 31 

cbn.siege@pnpc.com.fr 

2. Unité de conservation et collections (Michaud H.) 

Le Hameau, F-83400 Ile de Porquerolles 

Tél.: +33 (0)4 94 12 30 32 , fax: +33 (0)4 94 12 30 30 

Courriel: cbn.ile@pnpc.com.fr 

3. Antenne Languedoc-Roussillon (Molina J.) 

Institut de Botanique, Rue Auguste Broussonet, F-34090 Montpellier 

Tél.: +33 (0)4 99 23 22 11, fax: +33 (0)4 99 23 22 12 

cbn.lro@wanadoo.fr 

CEEP: Conservatoire Études des Écosystèmes de Provence Alpes du Sud-Var 

1, place de la Convention, F-83340 Le Luc 

Tél: +33 (0)4 94 50 38 39 / 06 16 97 82 03 

2. CEEP Var ((Catard A., Rombaut D.) 

1, Place de la Convention, F-83340 Le Luc 

Tél: +33 (0)4 94 50 38 39, fax: 04 94 73 36 86 

antoine.catard@libertysurf.fr 

dominique.rombaut@wanadoo.fr 

CEN-LR (Gendre T., Garnéro S., Houssard C., Klewscewski M.) 

Conservatoire des Espaces Naturels du Languedoc-Roussillon 

20 rue de la République, Espace République, F-34000 Montpellier 

Tél.: +33 (0)4 67 22 68 28, fax: +33 (0)4 67 22 68 27 

cen-lr@wanadoo.fr 

CELRL Corse (Delaugerre M.) 

Conservatoire des Espaces Littoraux et des Rivages Lacustres 

Délégation Corse 

3, rue Luce de Casabianca, F-20200 Bastia 

Tél: +33 (0)4 95 32 38 14, fax: +33 (0)4 95 32 13 98 

corse@conservatoire-du-littoral.fr 

m.delaugerre@conservatoire-du-littoral.fr 

EPHE (Cheylan M., Jakob C., Lombardini K.) 

Ecole Pratique des Hautes Etudes 

Laboratoire de Biogéographie et Ecologie des vertébrés, Case 94, 

Université de Montpellier II, Place E. Bataillon, 

F-34095 Montpellier cedex 5 

Tel: +33 (0)4 67 14 32 90, fax: +33 (0)4 67 63 33 27 

cheylan@univ-montp2.fr 

christiane.jakob@gmx.net 

katia89@hotmail.com 

Lycée international Georges Duby (Titolet D.) 

200, rue Georges Duby 

F-13080 Luynes 

titolet@yahoo.fr 

OEC (Pozzo di Borgo M. L.) 

Office de l'Environnement de la Corse 

Avenue Jean Nicoli, F-20250 Corte 

Tél.: +33 (0)4 95 45 04 00, fax: +33 (0)4 95 45 04 01 

SFO (Papazian M.) 

Société Française d’Odonatologie 

7, rue Lamartine, F-78390 Bois d’Arcy 

Fax: +33 (0)1 34 60 68 63, site internet: www.libellules.org 

TDV (Calvière T., Gauthier P., Grillas P, Jakob C., Perennou C., Rhazi M., 

Yavercovski N.) 

Station Biologique de la Tour du Valat, Le Sambuc, F-13200 Arles 

Tél: +33 (0)4 90 97 20 13, fax: +33 (0)4 90 97 20 19 

nom@tourduvalat.org, site internet: www.tourdu valat.org 

Université d’Aix-Marseille III – IMEP1 (Hébrard J.P.) 

Institut Méditerranéen d'Ecologie et de Paléoécologie - CNRS UMR 6116 

Université d'Aix-Marseille III, Faculté des Sciences et Techniques de Saint- 

Jérôme, Case 461, F-13397 Marseille cedex 20 

Tél: +33 (0)4 91 28 85 35, fax: +33 (0)4 91 28 80 51 

Université de Provence, Aix-Marseille I (Thiéry A) 

E.A. Biodiversité et environnement, Université de Provence, 3 place Victor 

Hugo, F-13331 Marseille cedex 3 

Tél: +33 (0)4 91 10 64 25, fax: +33 (0)4 91 10 63 03 

alain.thiery@univ-avignon.fr 

CNRS, Université de Montpellier II (Soulié-Märsche I.) 

Laboratoire de Paléobotanique - UMR 5554 du CNRS, Université Montpellier 

II, C.P. 062, Place E. Bataillon, F-34095 Montpellier cedex 5 

Tél: +33 (0)4 67 14 39 78, fax: +33 (0)4 67 14 30 31 

marsche@isem.univ-montp2.fr 

Université d’Annaba (Samraoui B.) 

Laboratoire de recherche des zones humides, Université d’Annaba, 

4, rue Hassi-Beïda, Annaba, Algeria 

bsamraoui@yahoo.fr 

Université Hassan II (Rhazi L.) 

Faculté des Sciences Aïn Chock, Laboratoire de Biologie et Physiologie 

Végétale, BP 5366, Maarif Casablanca, Morocco 

Tél.: (212) 037 86 33 10, fax: (212) 022 23 06 74 

lrhazi@hotmail.com 

ad. perso. 1 (Hugonnot V.) 

Le Bourg, F-43270 Varennes Saint Honorat 

Tél/Fax: +33 (0)4 71 00 23 07 

vincent.hugonnot@wanadoo.fr 

ad. perso. 2 (Paradis G.) 

7, cours Général Leclerc, F-20000 Ajaccio 

Tél: +33 (0)4 95 50 11 65 

guilhan.paradis@wanadoo.f 

Aknowledgements 

The Station biologique de la Tour du Valat would like to warmly thank all the 

editors, authors and everyone who collaborated on this volume as well as 

Mohand Achérar (CEN-LR), Joël Bourideys (DIREN PACA), Christine Bousquet 

(AME), Jean Boutin (CEEP), Maddy Cancemy (OEC), Marie-Luce Castelli (OEC), 

Paul Chemin (DIREN-LR), Claire Chevin (MEDD), Béatrice Coisman (CEEP), 

Natacha Cotinaut (Mairie du Cannet-des-Maures), Geneviève Coutrot (TDV), 

Daniel Crépin (DIREN LR), Florence Daubigney (TDV), Marie-Antoinette Diaz 

(TDV), Christian Desplats (CELRL PACA), Arnaud Dorgère (TDV), Aude Doumenge 

(AGRN-RH), Emilien Duborper (TDV), Renaud Dupuy de la Grandrive (ADENA), 

Roger Estève (CELRL PACA), Sabine Fabre (CEN-LR), Mauricette Figarella (DIREN 

Corse), Guy-François Frisoni (Réserve Naturelle des Bouches de Bonifacio), Jean 

Giudicelli (Maison régionale de l’eau, Barjols), Denis Gynouvès (ONF Var), Jean- 

Claude Heidet (CEEP), Bruno Julien (European Commission), Emilio Laguna 

(Generalitat de Valence, Espagne), Nicolas Leclainche (TDV), Olivier Limoges (Pôle 

relais Mares et Mouillères), Stéphanie Lieberherr (CEEP Var), Gilles Loliot (CELRL- 

LR), Isabelle Lourenço de Faria (Commission Européenne), Margarida Machado 

(Université d’Algarve, Portugal), Marc Maury (Ecosphère), Leopoldo Medina, 

Olivier Nalbone (ARPE), Georges Olivari (Maison régionale de l’eau, Barjols), Eric 

Parent (Agence de l’Eau RMC), Jean-Claude Pic (TDV), Marc Pichaud (TDV), 

Marlène Savelli (OEC), Pierre Quertier (ONF Var), Bertrand Sajaloli (Pôles relais 

Mares et Mouillères), Nathalie Saur (Agence de l’Eau RMC), Alain Sandoz (TDV), 

Laurine Tan Ham (TDV), Florence Verdier (CELRL LR) and Myriam Virevaire 

(CNBMP) for their contribution to the LIFE “Temporary Pools” project.

Summary 

Introductory notice. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

Plant species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

Apium crassipes (Koch ex Reichenb.) Reichenb. fil. . . . . 7 

Artemisia molinieri Quézel, Barbero & Loisel. . . . . . . . . 9 

Crypsis schoenoides (L.) Lam . . . . . . . . . . . . . . . . . . . . . . 12 

Damasonium polyspermum Coss. . . . . . . . . . . . . . . . . . . 15 

Elatine brochonii Clavaud . . . . . . . . . . . . . . . . . . . . . . . . 18 

Eryngium pusillum L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 

Illecebrum verticillatum L. . . . . . . . . . . . . . . . . . . . . . . . 23 

Isoetes duriei Bory (1) & Isoetes histrix Bory (2) . . . . . . 26 

Isoetes setacea Lam. (1) & Isoetes velata A. Braun (2) . 30 

Littorella uniflora (L.) Ascherson . . . . . . . . . . . . . . . . . . 34 

Lythrum borysthenicum (Schrank) Litv. (1) 

& Lythrum tribracteatum Salzm ex Sprengel (2). . . . . . 37 

Lythrum thymifolium L. (1) 

& Lythrum thesioides M. Bieb. (2). . . . . . . . . . . . . . . . . . 41 

Marsilea strigosa Willd. . . . . . . . . . . . . . . . . . . . . . . . . . 45 

Mentha cervina L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 

Myriophyllum alterniflorum DC. . . . . . . . . . . . . . . . . . . . 52 

Nitella opaca (Bruzelius) Agardh . . . . . . . . . . . . . . . . . . 55 

Ophioglossum azoricum C. Presl (1) 

& Ophioglossum lusitanicum L. (2) . . . . . . . . . . . . . . . . . 58 

Pilularia minuta Durieu ex A. Braun . . . . . . . . . . . . . . . . 62 

Ranunculus lateriflorus DC. (1) 

& Ranunculus nodiflorus L (2) . . . . . . . . . . . . . . . . . . . . 65 

Ranunculus revelieri Boreau . . . . . . . . . . . . . . . . . . . . . . 69 

Genus Riccia L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 

Riella helicophylla (Bory & Mont.) Mont.. . . . . . . . . . . . 76 

Teucrium aristatum Perez Lara . . . . . . . . . . . . . . . . . . . . 80 

Macrocrustaceans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 

Imnadia yeyetta Hertzog, 1935. . . . . . . . . . . . . . . . . . . . 83 

Linderiella massaliensis Thiéry & Champeau, 1988 . . . . 85 

Tanymastix stagnalis (Linnaeus, 1758) . . . . . . . . . . . . . . 88 

Triops cancriformis (Bosc, 1801) (1) 

& Lepidurus apus (Linné, 1758) (2). . . . . . . . . . . . . . . . . 91 

Odonata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 

Ischnura pumilio (Charpentier, 1825) (1) 

& Ischnura genei (Rambur, 1842) (2) . . . . . . . . . . . . . . 94 

Lestes barbarus (Fabricius, 1798) (1) 

& Lestes virens (Charpentier, 1825) (2) . . . . . . . . . . . . . 96 

Sympetrum fonscolombii (Sélys, 1840) (1) 

& Sympetrum meridionale (Sélys, 1841) (2) . . . . . . . . . . . . 99 

Amphibians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 

Bufo calamita, Laurenti 1768 . . . . . . . . . . . . . . . . . . . . . 102 

Discoglossus sardus Tschudi, 1837 . . . . . . . . . . . . . . . . . 105 

Pelobates cultripes (Cuvier, 1829). . . . . . . . . . . . . . . . . . 108 

Triturus cristatus, (Laurenti, 1768) . . . . . . . . . . . . . . . . . 111 

Triturus marmoratus, Latreille 1800 . . . . . . . . . . . . . . . . 114 

Glossary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 

Macrocrustacean inventory form . . . . . . . . . . . . . . . . . . . . . . 120 

Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 

5

Introductory notice 

This second volume of “Mediterranean Temporary Pools” presents, 

in the form of information sheets, a summary of the current 

knowledge on a selection of the remarkable species of 

Mediterranean temporary pools (29 plant species, 16 animal 

species). This knowledge is either bibliographical in origin or was 

acquired during the LIFE-Nature project “Conservation of 

Mediterranean Temporary Pools No. 99/72049”, presented on 

page one of volume 1. 

The species presented have been selected according to a number 

of criteria: 

• Their dependence on temporary aquatic habitats; 

• Their rarity (this criterion is, however, very variable from country 

to country); 

• The Mediterranean nature of their distribution; 

• Their presence at one or more of the seven sites of the LIFE 

project; 

• Their biological characteristics or their ecological requirements, 

illustrating various adaptive strategies. 

Plant species 

Amphibious vascular plants, flagship species of temporary pools, 

are the best represented with 26 species, i.e. almost a quarter of 

the 107 rare plants of the oligotrophic* temporary pools censused 

in 18 countries of the Mediterranean Basin (see volume 1, table 2). 

A discreet and often poorly known group, the liverworts are 

described on two information sheets. One is devoted to a very 

rare species of temporary pools (Riella helicophylla), the other to 

a genus particularly well adapted to temporary wetland areas, 

the genus Riccia. Finally, the choice of a charophyte* (algae) 

dependent on temporary Mediterranean pools, Nitella opaca, 

illustrates a poorly known but important group in temporary 

wetland areas. 

Macrocrustaceans 

Of the fifty or so species of the Mediterranean Basin, five have 

been selected from three different Orders. Two species are very 

rare in France (Linderiella massaliensis is endemic, Imnadia 

yeyetta is present on two sites). Water requirements differ 

among the species chosen (ephemeral habitats for Tanymastix 

stagnalis, semi-permanent habitats for Lepidurus apus and Triops 

cancriformis). Twelve Mediterranean countries support one or 

more of these species. 

Odonata 

In the Odonata, the 25 species with a Mediterranean distribution 

have life-history characteristics which are adapted to temporary 

flooding and can vary according to the latitude. The six species 

chosen illustrate restricted (Ischnura genei, endemic to the 

islands of the western Mediterranean) to wide (Sympetrum fonscolombii) 

Mediterranean distribution as well as several strategies 

of adaptation to summer drought, based on the mechanism 

and dates of egg laying, the existence of diapauses (egg or larva), 

the periods of emergence of the adults, the number of generations 

per year, wether or not migratory, the greater or lesser 

capacity to colonise new pools, etc. 

6 

Amphibians 

Among the sixty or so species found in the Mediterranean Basin, 

five which are adapted to temporary pools have been chosen in 

two different groups (anurans and urodeles), illustrating the various 

environmental requirements and biological characteristics: duration 

of larval cycle, size of clutches, etc.). 

Information sheet headings 

Latin name: current nomenclature 

Classification: except otherwise stated, the class then the family 

of the species (current nomenclature) are mentioned (in small 

upper-case letters) immediately under the title of each information 

sheet. 

Main synonyms: synonyms used in “classic” guides to flora and 

fauna. 

Common names: when they exist, the common French, Spanish, 

Portuguese, Italian, English, Moroccan and Algerian names are 

given. The names used at a national scale are given, with the 

exception of Morocco, where certain local names of oral tradition 

are known 265 , though it is not possible to give any precise 

region where these are used. For European countries, the French 

Internet site www.tela-botanica.org was consulted. In addition, 

the French names of the plant species were researched in the 

major flora of Bonnier 42 and in the Livre Rouge de la Flore 

Menacée de France (French Red Data Book of threatened flora) 274 , 

the Spanish and Portuguese names in Flora Iberica 64 , the Italian 

names in the Italian Red Data book of threatened flora 86 and in 

Flora d’Italia 292 . 

Subspecies: the subspecies currently recognised are mentioned. 

Description/identification criteria: a variety of bibliographic 

sources have been used, as well as the personal observations of 

the authors. 

Similar species: the species with which some confusion is possible 

are mentioned, as well as their distinctive characteristics. 

Distribution - Ecology 

Distribution/range 

The distribution of the species in France, then in other Mediterranean 

countries is given whenever possible, in particular in the 

countries of the western Mediterranean (Spain, Italy, Maghreb). 

Habitat 

This paragraph contains, for France, and if possible, for other 

Mediterranean countries: 

•A general description of the stations; 

• The mention of the habitats of “European Community Interest” 

or priority habitats for the species, included in annexe I of the 

Habitats Directive 12, 119 , accompanied by their Natura 2000* code; 

The mention of French habitats defined according to the CORINE 

Biotopes 39 typology; 

• For the flora: the list of the phytosociological communities* 

characterised by the species or to which the species belongs, and 

where possible the main companion species. 

Ecology 

This paragraph provides the main biological characteristics and 

environmental requirements useful for the conservation management 

of the species.

Biological characteristics 

In France, and if possible in other Mediterranean countries, information 

relating to the biology and life-history traits characteristics 

of the species is given. 

• For the flora: the biological type, methods of sexual reproduction 

(aerial, aquatic), description of seeds, and the biological 

cycle (germination, growth, flowering, fruiting, dispersal). 

• For the fauna: feeding, the site and the mechanisms of egg 

depositing, the biological cycle, (reproduction, egg laying, development), 

longevity, distinctive behaviour, interannual and spatial 

variations in the phenology. 

On each information sheet, a figure at the end of this paragraph 

illustrates the annual phenological cycle of a population and an 

average hydrological state of the pool. 

Example: the phenological cycle of Marsilea strigosa 

France 

Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec. 

Emergence of fronds and vegetative growth 

? 

Formation of sporocarps 

generally flooded phase (or of saturation of the soil for 

certain species) 

flooded phase some years 

dry phase 

Withering of fronds 

This schematic illustration cannot take into account all interannual 

variability in the Mediterranean climate (temperature, precipitation) 

or individual variability in the response of organisms, 

but it can provide the manager with an average state. Some 

authors express the variability of the response of a population to 

variations in the hydrological regime by dots extending the solid 

line of the different phases of a cycle. 

Environmental conditions 

In France, and if possible in other Mediterranean countries, the 

following are given (within the limits of the information available): 

• For the flora: the hydrological conditions required (depth of 

water, duration of flooding and drying out, germination of seeds, 

chemical characteristics, etc.); the nature of the substrate 

(acidic, calcareous, granulometry, etc.); sensitivity to interspecific 

competition (in relation to light in particular); the impact of perturbations 

such as cutting, grazing, trampling by livestock, etc. 

• For the fauna: the breeding requirements (duration and date 

of flooding, depth, type of pool, etc.); the territories, the wintering 

or summering habitat and the necessity for complementary 

habitats (terrestrial phase of amphibians, for example); the dispersal 

distance of the adults and the young after the aquatic 

phase; environmental perturbations and their impact (rain, 

drought, soil humidity, etc.); interspecific relations (competition, 

predation, etc.); the main factors in natural mortality, etc. 

Conservation - Management 

Assessment of populations 

In France, and in other Mediterranean countries, the number of 

sites is indicated (with, for the fauna, an estimation of the population 

numbers, if possible. 

Conservation status 

The status of the populations (stable, expanding or regressing) is 

provided, as well as the main sites of recent appearance of the 

species or of disappearance. 

Legal status of the species/level of protection 

A table provides for each species: 

• Inclusion in the IUCN Red Lists 176, 399, www.redlist.org . 

• Inclusion in the annexes of the Bern Convention 87 . 

• Inclusion in the annexes of the “Habitats” Directive 119 . 

• The status of protection at national or regional level in France 

and in other Mediterranean countries: 

- The protection status at national level for the flora was examined 

for Italy 86, 292 , Spain (Internet site: www.mma.es/conserv_nat/ 

acciones/esp_amenazadas/html/catalogo/flora.htm), France, Algeria 

(decree no. 93-117 of 23/11/1993 fixing the list of protected 

non-cultivated plant species). 

- The protection status at national level for the fauna was examined 

for France, Spain (Internet site: www.mma.es/conserv_nat/ 

acciones/esp_amenazadas/html), and Portugal (amphibians 59 ). 

- The protection status at regional level for the flora and fauna 

was examined for France, and in some regions of Italy for the 

fauna (Sardinia and Tuscany). 

• Inclusion in the Red Data Books and Lists in France and in 

other Mediterranean countries: 

- For vascular flora, the national Red Data Books and Lists of 

France 274 , Italy 86 , Spain 6 , Morocco 133 , Greece 290 , and Malta 344 were 

consulted as well as the regional Red Data Book for the Balearics 

331 . For the bryophytes, the Red Data Book of the bryophytes 

of Europe 128 , and the Red Data Book for France, which is currently 

being produced, served as references. 

- For the fauna, the National Red Data Book 242 was consulted for 

France, as well as the Red Data Lists for Spain (amphibians 293 ), 

Portugal (amphibians 59 ) and Italy (vertebrates 14 ). 

Conservation problems/threats 

The following factors are examined: 

• Anthropogenic factors: destruction, infilling, drainage, pollution, 

etc. 

• Natural factors: dynamics of the vegetation, etc. 

• Risks relating to populations: risks of extinction at a site. 

Management and conservation measures 

Introductory notice 

Current measures 

The main protected sites where the species are found are listed 

(Natural Parks, Nature Reserves, Natura 2000 sites, etc.) as well 

as the management measures in place. 

Recommendations 

These concern the management measures to put in place 

(hydrology, grazing, scrub clearing), modifications in the protection 

status of the species, protection of the habitat, etc. 

7

Mediterranean temporary pools 

Bibliography 

The authors of the references are listed in alphabetical order at 

the end of each information sheet, with a numbered cross-reference 

to the complete reference in the general bibliography at the 

end of the volume. 

Other information 

Acronyms and abbreviations used in the information sheets 

CBNBP: Conservatoire Botanique National du Bassin Parisien. 

CEEP: Conservatoire Etude des Ecosystèmes de Provence - Alpes 

du Sud. 

CEN-LR: Conservatoire des Espaces Naturels du Languedoc- 

Roussillon. 

MEDD: Ministère de l’Ecologie et du Développement Durable 

(France). 

ONF: Office National des Forêts. 

PACA (region): Région Provence-Alpes-Côte-d’Azur (France). 

IUCN: International Union for the Conservation of Nature 

8 

Journal officiel: Journal Officiel de la République Française (official 

bulletin for the public, giving details of laws). 

s.l.: sensu lato (in the broad sense) 

subsp.: subspecies 

sp.: Indeterminate species (from the Latin species). For example, 

Isoetes sp. signifies an indeterminate species of the genus Isoetes 

spp.: several species of the genus. For example, Isoetes spp. signifies 

“all or some species of the genus Isoetes”. 

em.: amended by (abbreviation of the Latin emendavit, from the 

Latin verb emendare: to modify, amend, improve). Used in phytosociology, 

for example: “the alliance Isoetion Br.-Bl. 1931 of 

the order Isoetalia Br.-Bl. 1931 em. Rivas-Goday, 1970”. 

µS.cm -1 : micro-siemens per cm (unit of measurement for conductivity 

of water) 

Glossary 

The technical and scientific terms not found in a current dictionary 

(such as the French dictionary Le Petit Robert), are defined 

in the Glossary. They are marked in the text with an asterisk* the 

first time they are encountered in each information sheet.

Apium crassipes (Koch ex Reichenb.) Reichenb. fil. 

ANGIOSPERMS 

APIACEAE 

Main synonyms 

Helosciadum crassipes Koch ex Reichenb. 

A. inundatum (L.) Reichenb. fil.var. crassipes Paol. 

A. inundatum (L.) Reichenb. fil subsp. crassipes Landi 

Sium crassipes (Reichenb.) Sprengel 

French name: Ache 

Italian name: Sedano di Sardegna 

Subspecies 

None 

50, 292 

Description/identification criteria 

• Herbaceous plant, hydrophytic* or amphibious, either 

hemicryptophytic* or annual. 

• Stem hollow and grooved, at first briefly prostrate and radicant, 

then upright, erect or floating, capable of growing up to 40 cm 

long in water. 

• Leaves alternate, of two types: basal leaves submerged, twoto 

four- pinnatisect* and completely divided into long, slender 

strips; emergent leaves simply pinnatisect with toothed segments. 

• Inflorescence: umbel composed of three to five partial umbels, 

no bracts, three to six bracteoles. 

• Flower with calyx invisible, corolla with five small white petals 

(0.5 mm) inferior ovary with two loculi, style slightly longer than 

the stylopodium* (apex of the ovary). 

• Fruit dry, consisting of two monospermic carpels (mericarps), 

indehiscent, 1.25 to 1.5 mm long, at the tips of short pedicels 

which are very swollen below. 

Similar species 

Apium inundatum is distinguished by its slightly larger size and 

bigger fruits, and especially by the non-swollen fruiting pedicels. 

The two species only coexist in Sicily (no risk of confusion in 

Corsica or Sardinia). 



Apium crassipes is considered to be a Tyrrheno-Sicilo-Numidian 

endemic species 50 . 

France 

The species only occurs in Corsica. 

Other Mediterranean countries 

Occurs in Italy (Sardinia, Pontins marshes and Sicily 292 ) and in 

North Africa (Tunisia and eastern Algeria). Casper & Krausch 63 

include the Nile Delta in the range. 

1 cm 

Habitat 

1. “Mediterranean temporary pool” habitat 

Plant species 

General description 

In Corsica and Sardinia, Apium crassipes is an important constituent 

in terms of the biomass* of the flooded phase of many 

temporary pools. In southern Corsica, A. crassipes, together with 

various floating species (Ranunculus peltatus, R. ophioglossifolius, 

Illecebrum verticillatum, Baldellia ranunculoides), is abundant 

in the temporary pools of Padulellu, Tre Padule de Suartone 

and Padulu219, 221, 284 . 

In Tunisia, A. crassipes is associated with Isoetes velata, Myosotis 

sicula, Illecebrum verticillatum, Eryngium barrelieri (E. pusillum), 

Ranunculus ophioglossifolius298 . 

Habitats Directive 

“Mediterranean Temporary Pools” (code 3170). 

CORINE Biotopes 

22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia). 

22.341 Short Mediterranean amphibious swards (Isoetion). 

22.3412 Aquatic Mediterranean Quillwort swards: communities 

containing Isoetes velata in water bodies of 

variable depth. 

Phytosociology* 

In its aquatic form, Apium crassipes belongs to formations of the 

submerged phase of the Isoetion, within communities of Isoetes 

velata. Some authors include the hydrophyte* formations in 

which A. crassipes is found within the communities of the Potamogetonetea 

pectinati class 284 . 

2. Other habitats 

Apium crassipes 


Apium crassipes is also found in ditches and in wet meadows. 

In these biotopes, which dry out rapidly at the end of winter, 

9


A. crassipes grows only to a small size and does not produce slender 

strap-like leaves. 


In Algeria, around the Mafragh, Tonga and Oubeira lakes, Géhu 

et al. 156 , for seasonally flooded meadows resulting from aggradation, 

created the association with Paspalum distichum and Apium 

crassipes which they named Helosciadi crassipes-Paspaletum distichi 

and which they included in the class Agrostietea stoloniferae, 

the order Plantaginetalia majoris and the alliance Paspalo-Agrostion. 

Ecology 


Life form 

Hydrophyte*, either hemicryptophytic* (perennial) or therophytic* 

(annual). 

Reproduction 

Aerial. 

Flowering 

In Corsica, from mid-March to late April. 

Adaptive strategy (sensu Grime 163 ) 

Stress-tolerant (S). 

Description of achenes 

Length: c. 1.25 to 1.5 mm. 

Phenological cycle 

Corsica 

� 

10 


Germination 

Vegetative growth 

Flowering 

Fruiting 


� 

Germination 

Hydrology 

Flooding in winter, even of short duration, appears to be essential 

for germination or the reactivation of buds and for the 

growth of young plants. 

Substrate 

Loose substrate, quite thin, (less than 10 cm deep), of varying 

particle size (sand, silt or clay) and acid. 

Interspecific competition 

A heliophilous* (light-demanding) plant, which does not thrive in 

the shade of tall species. 

Impact of perturbations 

•A low level of disturbance (by grazing, trampling), which 

causes a limited degree of substrate compaction, appears to be 

beneficial. 

• Heavy, frequent disturbance, for example due to all-terrain 

vehicles, that causes severe substrate compaction, are very detrimental 

(Chevanu pool, in Corsica). 



In Corsica, probably over 20. 


In Corsica 

The populations fluctuate from one year to another, but appear 

to be stable in the medium term at the Padulellu, Tre Padule and 

Padulu pools. On the other hand, as a result of the cessation of 

cattle grazing several decades ago, the Arasu, Mura dell’Unda 

and Muratellu pools now have small populations. The population 

of the Chevanu temporary pool, which is very bare and where the 

substrate is very compacted, is heading for extinction. 


IUCN 1997 

Bern Convention 


National and regional protection 

National Red Data Books and Red Lists 


Natural factors 

In Corsica, the likely cessation of extensive cattle grazing in the 

near future threatens to cause a severe fall in numbers at almost 

all sites, as a result of the spread of tall herbaceous plants 

(Schoenus nigricans, Scirpus holoschoenus, Oenanthe globulosa, 

O. lachenalii, etc.) and woody species (Phillyrea angustifolia, Erica 

scoparia, Myrtus communis, Pistacia lentiscus, etc.). 



In Corsica, currently no management measures, as a result of the 

absence of ownership control over land at the sites. 


In Corsica 

• The maintenance (and/or re-establishment) of extensive cattle 

grazing is recommended, as well as the maintenance of routes 

for walkers and hunters. 

• At least in the case of the Chevanu pool, a ban on using the 

site as a car park for several years in succession would probably 

enable the numbers of this species to increase again. 

Bibliography 

Anonymous, 1999 12 ; Bissardon & Guibal, 1997 39 ; Briquet & de 

Litardière, 1938 50 ; Casper & Krausch, 1981 63 ; Géhu et al., 1994 156 ; 

Grime, 1979 163 ; Lorenzoni & Paradis, 1997 219 , 2000 221 ; Paradis et 

al., 2002 284 ; Pignatti, 1982 292 ; Pottier-Alapetite, 1952 298 . 

Authors: Paradis G. & M. L. Pozzo di Borgo 

Collaborators: Grillas P. & N. Yavercovski 

- 

- 

- 

- 

-

Artemisia molinieri Quézel, Barbero & Loisel 

ANGIOSPERMS 

ASTERACEAE 

Main synonyms 

None 

French name: Armoise de Molinier 

Subspecies 

None 


• Sub-woody plant, very aromatic, with the same scent (like 

camphor) as Artemisia arborescens, 30 to 60 cm tall. 

• Stem branching above the neck. Leaves bluish-green, finely 

hispid, especially on the lower sides. Petiole equal in length to 

the leaf, which is triangular in outline, bipinnatifid* with threadlike 

lobes 1-1.5 mm wide. 

• Capitula fairly small, ovoid (2.5 x 2.5 mm), with 10-15 florets 

arranged in a dense spike. Inner involucral bracts lanceolatespatulate. 

The achenes (0.8-1 x 0.2-0.3 mm) are greyish with five 

ill-defined angles, smooth, slightly longitudinally grooved, and 

cut off somewhat obliquely at the apex*. 


The genus Artemisia is difficult. Historically, A. molinieri has been 

confused with A. alba Turra 31 , which belongs to another section. 

A. molinieri belongs to the Artemisia section which is characterised 

by its capitula having a glabrous receptacle and being 

allogamous* with the outer florets female and the disk florets 

hermaphrodite, all fertile. Within this section, A. molinieri is close 

to the species with green leaves and with the petiole lacking 

basal auricles*. Two of these species may be found in France: 

• Southernwood, (A. abrotanum L.), a cultivated wormwood whose 

origins are disputed, and which is distinguished from A. molinieri 

by its lemon scent, larger globose capitula (3-4 x 3-4 mm) with 

25-30 florets arranged in a leafy panicle, and its oval inner 

involucral bracts. 

• A. chamaemelifolia Vill. of dry grassland in the mountains of 

southern Europe and beyond, from Spain to Iran. This wormwood 

is distinguished from A. molinieri by its sessile paired cauline 

leaves with the lower lobes clasping the stem, and its slightly 

larger capitula: 2.5-3 mm. 

A. molinieri, A. chamaemelifolia and some populations of A. abrotanum 

have 2n = 18. 

All the wormwoods are found in dry habitats (rock, steppes, rubble 

etc.), apart from A. molinieri which is the only wetland wormwood. 

Notes: 

• According to Couteaux & Pons 93 , this taxon could be of hybridogenic* 

origin and its two stations could be artificial. 

•A small buprestid beetle, (Agrilus lacus), whose larva feeds solely 

on A. molinieri, has recently been described as endemic to the 

pools of the Centre Var 94 . 



Plant species 

Endemic* species of the Centre Var (France), occurring in the 

communes of Besse-sur-Issole and Flassans-sur-Issole. 

Habitat 

Artemisia molinieri 


Large natural depressions of karstic* origin, flooded irregularly 

depending on the year: flooding varying from more than six 

months to completely absent. 



1 cm 

CORINE Biotope 

22-342: Tall Mediterranean amphibious swards (Preslion cervinae 

Br.-Bl. 1931), with a type specific to this habitat: the 

Artemisia molinieri community 245, 307 . 


Class: Isoeto-Nanojuncetea 

Order: Isoetalia 

Alliance: Preslion cervinae 

Association*: Artemisia molinieri community 307 

Companion species 

At Gavoty and Redon: 

• Schoenoplectus lacustris, Scirpus maritimus, Phalaris arundinacea, 

Alisma lanceolatum, Eleocharis palustris, Oenanthe globulosa, 

Rorippa sylvestris, Ranunculus ophioglossifolius, Sisymbrella 

aspera, Veronica anagalloides. 

• Annual amphibious companion species of high natural-heritage 

value (for France), subject to competition from the wormwood 

11


and favoured by the opening-up of the wormwood mat (by trampling 

associated with grazing, or cultivation during dry years): 

Lythrum tribracteatum, Damasonium polyspermum, Heliotropium 

supinum, Schoenoplectus supinus, Chenopodium urbicum, Crypsis 

schoenoides, etc. 

Ecology 


Life form 

Artemisia molinieri is a competitive perennial species (chamaeophyte*), 

which forms dense species-poor stands. The few gaps, 

deriving from past disturbance, permit the development of a 

diverse annual flora. 

Biological cycle 

Vegetative growth begins in February with the appearance of the 

first green leaves. Its reproductive cycle takes place in summer 

(from mid-July to the end of August). The seeds begin to be 

mature from mid-September. The last green leaves wither in 

October/November. 

The plant produces runners, also perhaps layers, and it also readily 

propagates vegetatively. Couteaux & Pons 93 considered that vegetative 

reproduction was the plant’s only means of propagation. 

The plant is anemphilous 382 but the pollen is very poorly dispersed 

because the stamens remain contained within the corolla of the 

hermaphrodite florets* 93 . In addition, the pollen has a very low viability 

(10 to 30% of pollen viable depending on the population) 382 . 

However, the plant produces abundant achenes, which have no 

pappus* but are very light and are without doubt wind-distributed. 

Their production is irregular among individuals and stations 307 , and 

germination rates are extremely low 382 . This latter trait must be 

related to the irregularity of pollen production in a taxon whose 

origin probably derives from hybridisation* 93 , or to the demanding 

ecological conditions with which the wormwood is confronted 

(grazing, flooding) 382 . This is no doubt compensated for by the large 

number of achenes produced by a single plant 382 . At Lake Redon, 

many germinations were visible in 2001 after the water had fallen, 

in areas which had previously been ploughed. 

12 


Leafing and vegetative growth 


Flowering 

Fruiting 

Hydrology 

The habitat of this amphibious species is characterised by: 

•A flooding period which is often long, from autumn to spring 

(up to six or nine months), but in some years may be short or 

absent. 

•A maximum water depth of 20 to 70 cm, with the optimum 

apparently around 40 cm. 

Reproduction 

Flowering, which is late, is aerial. 

Substrate 

Colluvial silts of anthropogenic origin (erstwhile erosion of cultivated 

catchment areas) 93 . Meso-eutrophic* substrate. 


Artemisia molinieri is heliophilous*, and the establishment of 

woody plants (especially Fraxinus angustifolia) during dry years, 

or of helophytes* during wet years, is probably harmful to it (no 

data). 


This species does not appear to be grazed either by sheep (Redon) 

or by horses (Gavoty), probably because of its high terpene content 

during the vegetative stage* 241 . 


Assessment of populations and conservation status 

There are two well-known sites, Lake Gavoty and Lake Redon, 

where the plant is extremely abundant (several hundred thousand 

aerial shoots). 

The cultivation of part of Lake Redon, in 2001, destroyed a significant 

part of the station*. The Artemisia molinieri population has 

since proved to be healthy, many germinations having been seen 

during the subsequent years. 

There are several other depressions, which seem capable of having 

supported the wormwood in the past 307 , within a radius of a few 

kilometres from Gavoty and from Redon. Drainage operations 

and intensive cultivation could have wiped it out here. At three 

of these depressions, on fallow land, the recent discovery of small 

populations of A. molinieri (from one to under ten stalks) raises 

the question of whether these may be relict or alternatively pioneer 

populations. Moutte & Triat 261 found wormwood pollen in 

the sediment at a fossile lake northwest of Brignoles. They hypothesised 

that there was formerly a population of A. molinieri at 

this lake, which is several kilometres away from the currently 

known populations. However, Couteaux & Pons 93 questioned this 

hypothesis. 


IUCN 1997 




National Red Data Books 

and Red Lists 


“endangered” (E) 399 

- 

- 

France, PACA regional list: 

Decree of 09/05/1994/Journal 

Officiel 26/07/1994 

France, national Red Data 

Book: priority species274 Anthropogenic factors 

The deforestation and subdivision of catchment areas leads to 

the infilling of depressions and changes in water quality. The cultivation 

of pools in dry years probably constitutes a risk factor.


At Lake Gavoty, woody plants (ash, elm) are becoming established 

as a result of natural succession. This may eventually transform 

the depression into woodland, not susceptible to flooding and 

incompatible with the survival of a healthy Artemisia molinieri 

population. 

Risks relating to populations 

The risk of extinction at any site is very low in both of the populations 

known for a long time, which are very large (see vol. I, 

chapter 3, box 27). 



• Management plan in progress in the context of the production 

of the objectives document for Natura 2000* sites, including the 

two main pools with Artemisia molinieri. 

• Land acquisitions completed and in progress at Redon, in the 

framework of LIFE “Temporary Pools”. 

• Collection and ex-situ preservation of seeds by the Conservatoire 

Botanique National Méditerranéen de Porquerolles, and 

precautionary cultivation of the species on Conservatoire land 

since 1995. 


• Research into the role of disturbance in the dynamics and conservation 

of formations containing Artemisia molinieri and of 

other priority habitats (Lythrion tribracteati and Heleochloion) 

subject to competition from the wormwood. 

Plant species 

•Taxonomic study of A. molinieri, whose status as a species and 

as an endemic has been questioned. This type of study may only 

be considered at the scale of the Abrotanum section. 

• Retention of the band of low woody plants (Christ’s Thorn/ 

Jerusalem Thorn Paliurus spina-christi, Small-leaved Elm Ulmus 

minor and Blackthorn Prunus spinosa) around the pools, at the 

edge of the flooded zones. This plays a vital role as a natural 

water filter before it discharges into the depression and as a trap 

for fine materials which could cause the pools to silt up. 

Bibliography 

Aboucaya et al,. 2002 1 ; Berner, 1968 31 ; Camus, 1903 60 ; 

Couteaux & Pons, 1987 93 ; Curletti & Ponel, 1994 94 ; Guinochet, 

1982 167 ; Loisel, 1976 215 ; Masotti et al., 2003 241 ; Médail & 

Quézel, 1994 245 ; Moutte & Triat, 1968 261 ; Olivier et al., 1995 274 ; 

Quézel et al., 1966 307 ; Torrel et al., 1999 382 ; Tutin et al., 1964- 

1980 386 ; Walter & Gilett, 1998 399 . 

Authors: Michaud H 

Collaborators: Rombaut D. & N. Yavercovski 

13


Crypsis schoenoides (L.) Lam. 

ANGIOSPERMS 

POACEAE 

Main synonyms 

Heleochloa schoenoides (L.) Host ex Roem. 

French names: Crypside ovoïde, Crypside faux choin 

Italian name: Brignolo ovato 

English name: Swamp Pricklegrass 

Subspecies 

None 

Description/identification criteria 292 

• Annual herbaceous plant (therophyte* scapous*). 

• Stem branched from the base, 5 to 25 cm long, prostrate 

(rarely erect). 

• Leaves very typical of grasses (Poaceae) with glabrous, slightly 

swollen sheaths*, ligule divided into hairs, blade 1 to 3 cm long 

and 3 to 4 mm wide, at an angle to the stem. 

• Inflorescence a cylindrical spike, 5 to 8 mm in diameter and 1 

to 3 cm long, comprising many spikelets each consisting of one 

flower with three stamens. 


Young stages of Crypsis aculeata and Cynodon dactylon, but at 

the flowering stage confusion is not possible: 

• The inflorescence of C. dactylon is formed of digitate spikes, 

long and very slender. 

• The inflorescence of C. aculeata has the appearance of a tiny 

capitulum buried among the bracts*, with two stamens per flower. 



The taxon is considered to be Paleo-subtropical 292 . 

France 

This species is found at a number of ponds, watercourses and pools 

on the Mediterranean coast (Roussillon, Languedoc, Provence) 

and the coast of Corsica. It is also found on the Atlantic coast, 

from the Gironde to Finistère. 

Italy 

Several provinces in peninsular Italy (Trentino, Lombardy, 

Piedmont, Liguria, Emilia-Romagna, Tuscany, Lazio, Abruzzo), 

Sardinia and Sicily. 

Spain 

Scattered throughout the country; most frequent in the centre 177 . 

Morocco 

Tingitan peninsula, northwest Morocco, Middle Atlas. 

Algeria 

Tell region. 

14 

Habitat 

Crypsis schoenoides 


Crypsis schoenoides may be found in summer and early autumn, 

on the dried-out clay-silt edges of mesotrophic* ponds, on the 

banks of a few hydro-electric reservoirs (Corsica), and in temporary 

pools (in France, in Morocco in the dayas of the plains and 

low mountains etc.). It sometimes occurs on the silty edges of 

watercourses (Corsica, Algeria), in ditches, and also in poached 

pastures subject to flooding. 

Its optimal living conditions are clay-silt substrates which undergo 

an alternation of a lengthy flooded phase and a drying-out phase 

during summer (sometimes up to the beginning of autumn). 

The maximum water depth may vary between a few tens of centimetres 

(pools or ditches) to several metres (hydro-agricultural 

dams). This species will tolerate a slightly saline substrate. 



1 cm 


22.343 Mediterranean halo-nitrophilous swards (Heleochloion). 

“Post-aestival slightly halophilic and nitrophilous vegetation of 

temporarily flooded land, with Crypsis schoenoides, C. aculeata, 

C. alopecuroides and Centaurium spicatum”. 

54, 240, 250, 277, 281, 286, 318, 319 


Crypsis schoenoides (formerly named Heleochloa schoenoides) is 

characteristic of Heleochloion formations. 

In Corsica and Spain 


Order: Nanocyperetalia 

Alliance: Heleochloion 

Associations* and communities given in the literature: 

- Crypsio (aculeatae)-Heleochloetum schoenoidis 

Oberdorfer 1952 (southern mainland Europe, Corsica).

- Heleochloo schoenoidis-Fimbristyletum biumbellattae 

Br.-Bl. & Rivas Goday in Rivas Goday 1956 

corr. 54 (Spain). 

- Heliotropio supini-Heleochloetum schoenoidis Rivas 

Goday 1955 (Spain, Corsica). 

- Lythro flexuosi-Heleochloetum schoenoidis Rivas 

Martinez 1966 (Spain). 

- Crypsio schoenoidis-Cyperetum micheliani Martinez 

Parras et al., 1988 (Spain, Corsica). 

- Chenopodio chenopodioidis-Crypsidetum schoenoidis 

Paradis & Lorenzoni 1994 (Corsica). 

- Echinochloo cruris-galli-Crypsidetum schoenoidis 

Paradis & Lorenzoni 1994 (Corsica). 

- Community with Crypsis schoenoides and Corrigiola 

litoralis (in Paradis & Lorenzoni 277 ) (Corsica). 

- Community with Crypsis schoenoides and Cotula 

coronopifolia (in Paradis & Lorenzoni 277 ) (Corsica). 

In mainland France 

The Helochloion communities have not been studied in detail; a 

single community is indicated, that with Crypsis aculeata 215 , in 

which C. schoenoides and sometimes Cressa cretica also appear. 

In Morocco 

Crypsis schoenoides is most often found with Corrigiola littoralis, 

Heliotropium supinum and Hypericum tomentosum. 

Note 

whereas the Isoetion communities appear in spring and are oligotrophic*, 

the Helochloion communities appear in summer-autumn 

and are meso-eutrophic*. 

Ecology 


Life form 

Therophyte* (annual species). 

Reproduction 

Aerial. 

Flowering and fruiting 

• In Corsica, from mid-August to the end of September. 

• In Morocco, between June and August. 


Stress-tolerant ruderal (S-R). Disturbance is caused by flooding 

and by animal grazing during the summer and early autumn. 

Stress results from the drying-out of the soil during the summer 

and early autumn, which is variable depending on the topography 

and on edaphic* characteristics. 

Caryopses 

Length: 0.5 mm maximum. 


• At the end of spring or the beginning of summer: germination 

of caryopses followed by seedling development during the dry 

period (date variable, depending on climate and hydrology), on a 

substrate which is still damp or under a very shallow layer of 

water (less than 10 cm). 

• During summer: growth of plants on a dry substrate (competitive 

spring vegetation disappears completely or partly, giving way to 

the development of summer species), followed by flowering on 

the substrate which has now become very dry. 

• At the end of summer and beginning of autumn: fruiting (end 

of August to end of September), then death of the plants and 

break-up of the spikes. 

Corsica 


Morocco 


Hydrology 

Alternation between a long period of flooding in winter-spring 

and summer drying appears to be essential. 

Substrate 

Meso-eutrophic* substrate with fine particles (clay and silt), rich 

in organic debris and in nutrients, particularly nitrates. It is 

sometimes oligotrophic*, in the dayas* of Cork-Oak woods in 

Morocco. It may be slightly brackish at the surface (often more 

saline deeper down), which explains the presence of Tamarix spp. 

close to many coastal Crypsis schoenoides stations. There is usually 

a fairly significant degree of substrate compaction. 


As a light-demanding (heliophilous*) species, Crypsis schoenoides 

does not readily tolerate shading by tall species such as Phragmites 

australis, Typha latifolia, T. angustifolia, various Scirpus or 

Paspalum distichum, hence its restriction to bare areas. 


The potential for the establishment and survival of Heleochloion 

plant communities depends on denudation, generally due to human 

activity: 

• Deliberate cutting of helophytes*, to create shooting areas for 

example. 

• Grazing and trampling by cattle: trampling by cattle causes a 

fairly high degree of substrate compaction, which prevents tall 

helophytes from establishing themselves and does not hamper the 

germination and growth of the therophytes of the Heleochloion. 

• etc. 



Germination 

In Corsica 

This species occurs around ten sites: 

Growth 

Flowering 

Fruiting 


Germination 


Flowering and fruiting 

Dispersal of caryopses 

Plant species 

15


• Three on the west coast, in the lower Taravo valley (Etang de 

Tanchiccia, Etang de Canniccia 276, 277, 278, 279, 281 (Paradis unpublished 

observations, 2003). 

•Two near the Stabiaccio river at Porto-Vecchio 280 . 

• Three on the east coast, Etang de Gradugine 283 , Alzitone reservoir 

near Ghisonaccia, Teppe Rosse reservoir near Aléria 277, 278 . 


It occurs at more than twenty stations: 

• Pyrénées-Orientales: Villeneuve-la-Raho reservoir and alluvial 

plain of the Agly. 

• Aude and Hérault: Capestang, Poilhès, Ouveillan, Estagnol 

ponds and elsewhere in the lower valley of the Aude. Sesquiers 

marsh at Mèze, pools of Notre-Dame-de l’Agenouillade at Agde, 

pools and ditches at Roque-Haute, Saint-Martin-de-Londres and 

Saint-Nazaire de Pézan, etc. 

• Gard: marsh at Aigues-Mortes, ponds of Pujaut and Capelle, 

and alluvial plain of the Gardon at Dions. 

•Vaucluse: flood meadows at Monteux. 

• Bouches-du-Rhône: in the Camargue (ditches at the Tour-du- 

Valat, Verdier marsh). 

•Var: Lake Redon at Flassans, Badelune basins at Cannet-des- 

Maures, pools at Grimaud and Hyères, ditches at Roquebrunesur-Argens. 

No recent records from Fréjus. 

In Morocco 

More than twenty stations are known from the Tangier, Larache- 

Casablanca and Middle Atlas regions. 


In France 

The population at Tanchiccia (Corsica) appears to be decreasing 

in comparison with previous observations by Paradis 281 . On the 

other hand, all the other populations on the island are either stable 

or increasing. 

In mainland France, populations are large and sometimes in very 

high numbers in the region of Capestang, Ouveillan, Poilhès, etc. 

The species appears to have decreased on the coast in the Pyrénées- 

Orientales and has disappeared from the Alpes-Maritimes coast 332 . 

It is surviving well elsewhere: the number of new sites largely 

compensates for unconfirmed sites mentioned in earlier records. 

It seems to have appeared in the Camargue and in the floodplain 

of the Rhône between Arles and Tarascon as a result of the 

desalination which followed the increase of ricefields 256 . 

In Morocco 

The populations appear to be fairly stable despite the increasing 

pressures on the sites. 


IUCN 1997 





and Red Lists 

16 

- 

- 

- 




France, National Red Book: 

“to be observed” 274 


Anthropogenic factors 

The infilling of pools and ponds (in Morocco and at Tanchiccia in 

Corsica), drainage for agriculture or permanent flooding for 

shooting (mainland France), and overdeepening for the extraction 

of sediments or rock (Morocco) are the main threats. In Morocco, 

some sites located near large towns are also threatened with 

destruction in the mid-term by urban development. 


This plant is favoured by sheep or cattle grazing (France, Morocco) 

as well as by certain types of management for hunting, such as 

the creation of pools in reedbeds. 


In mainland France, the isolation of the Var populations and of 

the very small inland pools in Languedoc-Roussillon constitutes 

a potential threat. 



• In Corsica: at present there are no management measures in 

place, due to the absence of control over land ownership at the 

sites. 

• In mainland France: the objectives document for the Natura 

2000* site pools of the Centre Var takes this species into account, 

but no conservation measures are known for the other sites. The 

Conservatoire Botanique Méditerranéen de Porquerolles collects 

seeds and preserves them (ex-situ). 

• In Morocco: the species is not protected and does not benefit 

from any management measures. 


• Ensure that management by extensive cattle grazing is carried 

out and that an appropriate hydrological regime is maintained 

(mainland France, Corsica, Morocco). 

• In Morocco, organise site monitoring as well as control of urban 

development in the short term. 

Bibliography 

Anonymous, 1999 12 ; Bissardon & Guibal, 1997 39 ; Braun- 

Blanquet et al., 1952 49 ; Brullo & Minissale, 1998 54 ; Grime, 

1979 163 ; Loisel, 1976 215 ; Maire, 1952-1987 230 ; Martinez Parras 

et al., 1988 240 ; Molero & Romo, 1988 250 ; Molinier & Tallon, 

1974 256 ; Paradis & Lorenzoni, 1994 277, 278 ; Paradis & Orsini, 

1992 276 ; Paradis & Pozzo di Borgo, 2000 280 ; Paradis & Piazza, 

1995 279 ; Paradis et al., 2002 283 ; Paradis, 1992 281 ; Peinado Lorca 

et al., 1988 286 ; Pignatti, 1982 292 ; Rivas Goday, 1964 318 , 1970 319 ; 

Rivas-Martínez et al., 2001 320 ; Salanon et al., 1994 332 ; Valdés et 

al., 2002 391 . 

Authors: Paradis G. & M. L. Pozzo-di-Borgo 

Collaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski

Damasonium polyspermum Coss. 

ANGIOSPERMS 

ALISMATACEAE 

Main synonyms 

D. stellatum Thuill. var. polyspermum (Coss.) Loret & Barrandon 

D. stellatum Thuill. var. polyspermum (Coss.) P. Fourn. D. alisma 

Mill. proles polyspermum (Coss.) Rouy 

“D. alisma Mill. var. polyspermum Loret & Barrandon” in Guin. & 

R.Vilm. 

D. alisma Miller subsp. polyspermum (Coss.) Maire 

French name: Etoile d’eau à nombreuses graines 

Spanish names: Saeta de agua, Cola de Golondrina, Almea, 

Azumbar 

Italian name: Mestolaccia siciliana 

Moroccan name: Mizmar er raaï 

English names: Starfruit, Thrumwort 

Subspecies 

None 


• Small annual plant with the leaves all basal and arranged in a 

rosette. 

• First submerged leaves linear and not persisting, subsequently 

replaced by floating leaves with long petioles and narrow lanceolate 

blades, attenuated or more or less truncated at the base, 

with faint transverse venation. 

• Leaves produced during the non-flooded phase similar to the 

floating leaves but with a shorter petiole and a thicker blade. 

• Inflorescence most often with one, sometimes several superposed 

umbelliform whorls. Flowers hermaphrodite with three persistent 

green sepals and three deciduous coloured petals. Petals 

2 to 2.5 times as long as the sepals (5.4 ± 0.4 mm x 4.1 ± 0.6 mm), 

white, marked with yellow at the base and more or less tinged with 

pink at the tip. 

• Fruit consisting of 6-9 carpels arranged in a star shape (follicles), 

falling when ripe. Five to more than twenty cylindrical seeds with 

transverse ridges (dimensions: 0.9-1.2 mm x 0.5-0.7 mm), in each 

mature carpel. 


Damasonium alisma Mill. has only two large seeds (1.7-2.5 mm 

x 0.8-1.2 mm) per carpel when the fruit is ripe. According to some 

authors, additional distinguishing characters include the plant’s 

often larger size and more robust build, leaves rounded to cordate* 

at the base with oblique transverse veins visible against the 

light, smaller petals (4.2 ± 0.5 mm x 3.7 ± 0.5 mm) lacking pink 

colour at the tips, and shorter anthers (0.75 ± 0.006 mm v. 0.82 

± 0.12 mm). However, the relevance of these characters needs to 

be confirmed, especially in relation to the influence of the environment 

(depth of water, length of submersion, etc.). 

The two Damasonium are not always separated by botanists. According 

to Rich & Nicholls-Vuille 316 , they are also separable on the 

basis of their largely different geographical ranges and by their 

chromosome numbers (D. alisma 2n = 28, tetraploid; D. polyspermum 

2n = 14, diploid*). 

Plant species 

Baldellia ranunculoides (L.) Parl. is distinguished, at all stages of 

development, by the scent of coriander that is released when the 

leaves are crushed. During flowering or fruiting, the arrangement 

of the many obovate carpels (≈ 2 mm) in a globose head removes 

any doubt. 

Alisma spp.: The three Alisma species included in the French flora 

are usually far more robust than Damasonium polyspermum. At 

flowering or fruiting they are easily distinguished by their many 

carpels, which are blunt oval shaped, very tightly packed and 

whorled in a single row. Unlike Damasonium, these are all perennial* 

species. 



Western Mediterranean species. 

France 

Mediterranean France from Biterrois (Hérault) to the Centre Var 

(Var). 


North Africa: in freshwater dayas*, especially in mountains, in 

Cyrenaica, in the High Plateaux of northwestern Algeria and in 

the Middle Atlas in Morocco. 

Italy: southwest Sicily in the coastal zone 292 . 

Edges of ponds and pools in the Iberian Peninsula, in central and 

southern Spain as well as Portugal, Greece and Syria. 

Habitat 

Damasonium polyspermum 

1 cm 


Land temporarily flooded by fresh or slightly brackish water in 

the meso-Mediterranean zone: pools, edges of ponds and lagoons. 

17





22.32 Euro-Siberian dwarf annual amphibious swards 

(Cyperetalia fusci). 

22.34 Mediterraneo-Atlantic amphibious communities (Isoetalia). 

22.341 Short Mediterranean amphibious swards (Isoetion 

Br.-Bl. 1931). 


• Euro-Siberian dwarf annual amphibious swards (Cyperetalia 

fusci): 

- Lythrion tribracteati (e.g. the Lythrum tribracteatum-Teucrium 

aristatum association* 215, 252 . 

- Elatino triandrae-Eleocharition ovatae (e.g. Elatinetum 

macropodae). 

• Southern amphibious communities (Isoetalia): 

- Short Mediterranean amphibious swards (Isoetion). 

• Damasonium polyspermum sometimes overlaps into formations 

appearing later in the year and having a more distinctly eutrophic* 

or even halophilic* character: Heleochloion schoenoidis. 

These various vegetation types are frequently found in mosaics 

with perennial communities based on Eleocharis palustris, 

Bolboschoenus maritimus, Phragmites australis, Mentha cervina, 

Artemisia molinieri, Isoetes setacea, etc. (Preslion cervinae, 

Isoetion, Phragmition, etc.). 


The species most often associated with Damasonium polyspermum 

are: Lythrum tribracteatum at all the French localities, with 

in addition, depending on the site, L. hyssopifolium, Pulicaria vulgaris, 

P. sicula, Myosurus minimus s.l., Juncus bufonius s.l., 

J. tenageia and Herniaria glabra. 

Ecology 


Life form 

Annual species (therophyte*). 

Reproduction 

Aerial. 

Seeds (see above § Description) 

The lifespan of seeds in the sediment is not known, but observations 

of the dynamics of natural populations indicate that it is 

long. 


Seed germination and growth of leaves during the flooded phase 

(end of winter, beginning of spring). Flowering in spring (from 

April to June) up to the first dry periods, but may be prolonged 

or repeated in autumn if the substrate is sufficiently wet. Ripening 

of fruits and dispersal of seeds during the dry phase (summer) 

and before the autumn rains which mark the end of the 

cycle. 

18 

Southern France 


Germination and vegetative growth 

Flowering 


Formation and ripening of fruits 

Seed dispersal 

Hydrology 

The germination of seeds, which is very variable from one year to 

another, requires immersion in water (a few centimetres) or waterlogged 

soil in spring, and little vegetation cover (strong insolation). 

The plant can remain in the vegetative stage under very shallow 

water, with characteristic floating leaves, but must emerge quickly 

for flowering and fruiting. The size and density of individuals and 

the length of their life cycles exhibit spatial variability within the 

pool, in relation to hydrological gradients (duration of flooding, 

dates of flooding and drying out) that are themselves dependent 

on topographical gradients. 

Substrate 

Plant tolerant of a wide range of soils and of slightly saline conditions. 


A very light-demanding (heliophilous*) species, and therefore 

very sensitive to any plant cover at the germination stage. 


Disturbance which helps to control competing vegetation is beneficial, 

whereas disturbance resulting in the burial of seeds 

impede germination (trampling and digging by livestock or wild 

animals, ploughing, overdeepening or dredging of the pool, etc.) 

(see Vol. 1, box 38). 



France 

The species currently occurs at 10 sites: 

• Hérault: Vendres/Sauvian (Malhol de l’eau), Agde (Notre-Dame 

de l’Agenouillade), Montarnaud (pool on the Tamareau garrigues). 

• Gard: La Capelle-et-Masmolène (Etang de la Capelle). 

• Bouches-du-Rhône: Arles (Tour-du-Valat and Lanau), Saint- 

Antonin-sur-Bayon (Plateau du Cengle). 

•Var: Gonfaron (Lake Bonne Cougne), Flassans-sur-Issole (Lake 

Redon), Besse-sur-Issole (Lake Gavoty). 

• Absent from Corsica. 

Italy 

Only one site is known, on the coast of Sicily 86 . 

Spain (Medina, pers. com.) 

The species occurs in 13 provinces in the centre and north of the 

country (between 50 and 100 sites), and three provinces in the 

south of the country (about twenty sites).

Portugal 

A single site supports this species (Medina, pers. com.). 

Morocco 

The species is not considered to be rare 133 . 

Note 

As the two species of Damasonium have not always been separated 

by botanists, some data remain uncertain: for example old 

publications list D. alisma sensu lato from the closed depressions 

of the Roussillon plain (between Têt and Tech) 351 , in the coastal 

pools at Argelès-sur-Mer 155 and in the Montmajour marsh 

(Arles/Fontvieille) (Jacquemin 1848 in Molinier 257 ). In the French 

Mediterranean region, D. alisma is only known with certainty 

from the extreme south of Larzac 30 and appears therefore to be 

absent from the meso-Mediterranean zone. 


In France, eight populations have disappeared, or have not been 

seen again, in the communes of Sauvian (Vendres plateau), 

Portiragnes/Vias (Roque-Haute), Agde (Rigaud pools), Redessan, 

Jonquières, Manduel (pools of Jonquières, Campuget and 

Redessan), Bellegarde, La Barben (Estagnolet pool). 


IUCN 1997 





and Red Lists 


- 

- 

- 

- France, national list: Decree 

of 20/01/1982/Journal Officiel 

13/05/1982, amended by Decree 

of 31/08/1995/ 

Journal Officiel 17/10/1995 

- France, national Red Data Book: 

“to be observed” 274 

- Greece, national Red Data Book: 

“vulnerable” 290 

- Italy, national Red Data Book: 



Several populations have disappeared following the infilling or 

drainage of the depressions which supported them, in connection 

with agricultural developments (Vendres plateau, pools and ponds 

of the Costière Nîmoise) or with urban development (Rigaud 

pools at Agde). 

Plant species 


As the species is an annual, it does not compete well with colonial 

perennial species (Artemisia molinieri, Mentha cervina, 

Isoetes spp. Eleocharis palustris, etc.), this being partly compensated 

for by its abundant seedbank. In many depressions, the appearance 

of Damasonium is irregular from one year to another 37 . 


This risk is low in the absence of any major perturbations, due to 

the fecundity of the plant and the longevity of the seeds in the 

sediment. 



The Lanau pool (Bouches-du-Rhône) was acquired by the CEEP in 

1998. 

The only management measures known at present are: 

• Monitoring of the population of the Lanau pool in the Crau, in 

relation to grazing management. 

•A management plan in progress for the pools of the Centre Var, 

in the context of drawing up the Objectives Documents for 

Natura 2000* sites. 

• The ex-situ preservation of seeds by the Conservatoire 

Botanique National Méditerranéen de Porquerolles. 


• Promotion or maintenance of grazing, which is favourable to 

the growth of the species, as is any other means whereby perennial 

vegetation communities are opened up. 

• Promotion of the acquisition and management of Damasonium 

sites, which are in addition often rich in other rare species. 

Bibliography 

Bernard, 1997 30 ; Bigot, 1955 36, 37 ; Braun-Blanquet, 1935 48 ; Conti 

et al., 1992 86 ; Danton & Baffray, 1995 96 ; Fennane & Ibn Tattou, 

1998 133 ; Gautier, 1898 155 ; Guinochet & Vilmorin, 1978 166 ; Maire, 

1952-1987 230 ; Médail et al., 1998 246 ; Michaud & Molina, 1999 249 ; 

Molinier, 1981 257 ; Molinier & Tallon, 1947 252 ; Molinier & Tallon, 

1948 253 ; Olivier et al., 1995 274 ; Phitos et al., 1995 290 ; Pignatti, 

1982 292 ; Pouzolz de, 1862 100, 101 ; Rich & Nicholls-Vuille, 2001 316 ; 

Rivas-Goday, 1970 319 ; Simonneau, 1967 351 ; Vuille, 1987 398 . 

Author: Michaud H. 

Collaborator: Yavercovski N. 

19


Elatine brochonii Clavaud 

ANGIOSPERMS 

ELATINACEAE 

Main synonyms 

E. hydropiper L. var. pedunculata (Moris) Fiori 

E. hexandra (Lapierre) DC. subsp. brochonii (Clavaud) P. Fourn. 

French name: Elatine de Brochon 

Subspecies 

None 

76, 96, 138, 381 


• Herbaceous plant, amphibious, short (2-7 cm) with a relatively 

long and significant root system. 

• Leaves opposite, spatulate, more or less petiolate, with a 

smooth slender stalk, prostrate, much-branched and rooting at 

the nodes. 

• Flowers hermaphrodite, actinomorphic*, small (not more than 

5 mm), sessile, solitary in the leaf axils, with three free sepals 

persisting in fruit scarcely longer than the fruit, and three free 

petals, white with pink veins. Androecium of six stamens 

arranged on two verticils. 

• Fruit: spherical, slightly flattened capsule* with three valves. 

Seeds many, small, reticulate, straight or slightly arcuate, with 

longitudinal ribs, transversely grooved. 

• Plant brownish-coloured at the end of the cycle. 


• Elatine hexandra has pedicellate flowers (sessile in E. brochonii). 

• Lythrum borysthenicum has crimson petals (not whitish-pink), 

sepals longer than the capsules (scarcely longer in E. brochonii), 

and a rough stem which is not slender. 



A western Mediterranean species. 

France 

Occurs in the southwest (Gironde, Landes, Pyrénées-Atlantique) 

and in Corsica 81, 136, 219, 274, 346, 392 . 


Spain 76 , Morocco 303 , Algeria 292, 386 . 

Habitat 


In France, Elatine brochonii is found on the edges of temporary 

pools in southwest France on siliceous substrates with irregular 

flooding between years. In Corsica, this species occurs at a single 

site (Padulellu pool) in a depression that cuts into a granitic 

ridge (105 m altitude) covered with low matorral*. Flooding and 

drying out, which are irregular from one year to another, take 

place through rainwater and evaporation. The maximum water 

level recorded in 1996 (a very wet year) was 80 cm 219 . 

20 

On the Atlantic plains of Morocco, temporary pools with E. brochonii 

are found in Cork-Oak woods (Mamora and Benslimane) 

under sub-humid to semi-arid climatic conditions. Flooding usually 

takes place in December-January, solely by rainwater, and 

they dry out during April-May. E. brochonii also occurs in 

Morocco in mountain pools in siliceous terrain 133, 228, 229, 230 . 




22.34 Southern amphibious communities (Isoetalia). 



Elatine brochonii is found together with Juncus pygmaeus, J. bufonius, 

Lythrum hyssopifolia, Illecebrum verticillatum, Exaculum 

pusillum and Isoetes velata, which means that it is included in 

two alliances, the Isoetion velatae and the Cicendion 219 : 



Alliance: Isoetion velatae Br.-Bl. 1931 

Alliance: Cicendion filiformis Rivas Goday (1961, 1964) 

Br-Bl. 1967. 

Ecology 

0.25 cm 


Life form 

Therophyte* (annual). 

Adaptive strategy 

Stress-tolerant-ruderals (S-R) 163 . 

Reproduction 

Aerial. 

Elatine brochonii

Seeds 

Seeds are abundant (33 to 50 seeds per capsule), long-lived* in 

the sediment, small (length: 0.63 mm, width: 0.27 mm), and 

weigh less than 1.4 mg. 


Amphibious plant with a very short life cycle. It begins its biological 

cycle under water or just after emergence and ends it out 

of water. Its appearance, irregular from one year to another and 

on varying dates, has been observed in southwest France 346 . 

In Corsica germination takes place at the end of spring (May- 

June), only during wet years. Flowering occurs from May to July, 

fruiting in June and July, and the seeds are dispersed in July and 

August. 

In Morocco, Elatine brochonii germinates in February-March and in 

April forms flower buds which very quickly develop into flowers. 

Fruiting takes place from April to June with the dispersal of seeds 313 . 

Corsica 


Morocco 


Germination 

Flowering 


Germination 

Fruiting 

Flowering 

Fruiting 



Hydrology 

Seed germination requires the soil to be flooded (a few centimetres 

of water) or saturated during March, and intense insolation 

(bare soil). The plant may remain in the vegetative stage under 

very shallow water, but must emerge quickly for flowering and 

fruiting 314 . 

Substrate 

Acid, oligotrophic* at Moroccan stations, mesotrophic* in 

Corsica. 


A very light-demanding species (heliophilous*), Elatine brochonii 

is very sensitive to any plant cover at the germination stage. 


Perturbations which help to control competing vegetation are 

favourable to Elatine brochonii, while those resulting in burial of 

the seeds will impede germination (87% germination rate for 

seeds on the surface compared with only 13% and 6% for seeds 

buried in sand to depths of 2 mm and 5 mm respectively). This 

burial may be caused in the field by the passage or digging of 

wild and domestic animals, sedimentation (Padulellu) and the 

movements of vehicles. 



Plant species 

In mainland France, 13 stations were formerly known in the 

southwest, of which only one appears to have survived 274 ; there 

is one site in Corsica (Padulellu pool 219 ), and at least eight in 

Morocco 133, 346 . 


The interannual variability of the species causes difficulty in 

evaluating the status of the populations. However, it no longer 

appears to be present at most of its mainland French sites, and 

the Corsican population remains vulnerable. In Morocco the populations 

appear to be better preserved, but are subject to rapidly 

increasing human pressures, especially around Rabat. The high 

survival rate of the seed stocks* in the soil gives grounds for hoping 

that populations may still exist in the dormant state. 


IUCN 1997 





and Red Lists 


- 

- 

- 

- France, national list: Decree of 

20/01/1982/Journal Officiel 

13/05/1982, amended by the 



- France, national Red Data 

Book274 : priority species 

- Spain, national Red List: 


- Morocco: 

133, 181, 381 

“very rare” (RR) 


Digging out and removal of sediments, frequent vehicle movements 

creating tracks through the pool (Mamora pool in Morocco), 

or the formation of a sandy alluvial fan partly covering the Elatine 

station (Padulellu pool in Corsica). 


Competition from terrestrial vegetation (Cistus monspeliensis, 

Pistacia lentiscus, Myrtus communis, etc.) or from helophytes 

(Scirpus maritimus, Eleocharis palustris, etc.) only constitutes a 

temporary problem, inherent to Mediterranean temporary pools, 

if the hydrological regime is not altered. 


Despite its large seedbank, which enables populations to persist, 

risks of extinction may arise in the future, especially at the sites 

south of the Mediterranean where the species may be considered 

to be at the southern limit of its range. The high frequency of dry 

years in these countries (Morocco) could result in the long term 

in a decline in the populations of Elatine brochonii. The low number 

of sites increases the risk of extinction. 

21




None 


• In France and in Morocco systematic monitoring of the known 

populations is recommended, as well as an assessment of the 

environment, the condition of the populations (including seedbanks), 

and the possible opportunities for their restoration. 

• At the Padulellu pool (Corsica), it is a matter of urgency that 

the process of sedimentation be halted (sand originating from 

the erosion of the track running next to the pool) and, to this 

end, that the belt of Myrtle maquis be reinstated; the driving and 

parking of vehicles in the pool should also be prohibited. 

• In Morocco, the continuance of grazing appears to be beneficial 

in controlling the amount of competing vegetation. 

22 

Bibliography 

Anonymous, 1999 12 ; Anonymous, 2000 13 ; Bissardon & Guibal, 

1997 39 ; Cirujano & Velayos, 1993 76 ; Clavaud, 1883 81 ; Danton & 

Baffray, 1995 96 ; Fennane & Ibn Tattou, 1998 133 ; Fiton, 1916 136 ; 

Fournier, 1936 138 ; Grime, 1979 163 ; Jahandiez & Maire, 1931- 

1934 181 ; Lesouëf & Richard, 1995 210 ; Lorenzoni & Paradis, 1997 219 ; 

Maire, 1926 228 , 1932 229 , 1952-1987 230 ; Médail et al., 1996 243 ; Olivier 

et al., 1995 274 ; Pignatti, 1982 292 ; Quézel & Santa, 1962-1963 303 ; 

Rhazi et al,. 2001 313, 314 ; Rivas-Goday, 1970 319 ; Schotsman & Bosserdet, 

1966 346 ; Schotsman, 1985 347 ; Titolet & Rhazi, 1999 381 ; 

Tutin et al., 1964-1980 386 ; Vanden Berghen 392 , 1966 ; Vivant, 

1960 396 . 

Authors: Rhazi L. 

Collaborators: Grillas P., G. Paradis & D. Titolet

Eryngium pusillum L. 

ANGIOSPERMS 

APIACEAE 

Main synonyms 

E. barrelieri Boiss. 

French name: Panicaut nain de Barrelier 

Italian name: Calcatreppola di Barrelier 

Subspecies 

None 


• Short, thistle-like plant, in low-growing clumps. 

• The clump is composed of a more or less large number of units, 

depending on its age. A unit consists of a main stem, bearing a 

rosette of leaves at its base and ending in a capitulum (inflorescence). 

The stem may branch and give rise to new units. 

• The young rosette leaves are elongated and not prickly. As they 

age they become stiff, with thin spines along their edges. 

• There may be very many capitula on a single clump. Each carries 

several very prickly bracts. The flowers are very small and bluish. 

They produce tiny achenes (indehiscent dry fruits) 220, 274, 292 . 


None 



Southern Mediterranean species 

France 

Only in southern Corsica 


Sardinia, Sicily, southern Italy, Tunisia, Algeria, Syria, Morocco 274, 292 . 

Habitat 


The Musella depression is a poljé* (karstic* depression) which 

passes through very different ecophases* over the course of a 

year, whose two extremes are a winter flooded ecophase and a 

very dry summer ecophase. 

Flooding, which is due mainly to the rising water table in the 

underlying limestone, occurs in winter during wet years and may 

in exceptional cases last from November to April (e.g. in 1996 

and 2001). Drying out begins in March (in April in very wet 

years), by lowering of the water table. 

In summer, the clay-silt substrate becomes very cracked as it dries out. 




In winter flooding phase 

22.5 Oligo-mesotrophic* calcium-rich waters. 

In spring 

22.34 Mediterraneo-Atlantic amphibious communities. 

Plant species 


In summer and early autumn: Mentha pulegium and Eryngium 

pusillum meadows, classifiable within the Trifolio-Cynodontion 

alliance (order Plantaginetalia majoris 156 and class Molinio- 

Arrhenatheretea), which does not appear to correspond to any 

categories in CORINE Biotopes. 

Ecology 

1 cm 


Life form 

Perennial species, hemicryptophyte*. It is formed of sympodial* 

units. Rosettes of leaves develop at the base of the units, which 

are monocarpic (only flower once). The different units together 

form a low clump 220 . 

Germination 

Aquatic. 

Reproduction 

Aerial for flowering 220 . 

Eryngium pusillum 

Survival strategy of the clumps 

Stress-tolerant (S) according to the terminology of Grime 163 


Germination in November if the site is flooded, dispersal of 

young plants during the flooded period (hydrochory), vegetative 

growth from November to May, flowering end of May and June, 

fruiting in June and July, dispersal of achenes for a very short distance 

from the parent plant and dispersal of the seedlings by 

floatation (hydrochory) 220 . During and especially at the end of 

summer the leaves wilt and disappear, while new leaves appear 

with the first rain of September-October. 

23


� 


Hydrology 

The alternation of a flooded ecophase (from the end of autumn 

to the beginning of spring) and a dry ecophase is necessary. In 

the absence of winter flooding the seeds do not germinate in 

spring. An absence of germination for several years is, however, 

not a handicap for this species, as the large amount of germination 

during wet years, plus vegetative* reproduction, enable the 

population to rebuild itself. 

Substrate 

Mesotrophic*. 


A strictly heliophilous* species, negatively affected by the shade 

from taller vegetation. 


A prickly species once the site dries out, not (or very little) 

grazed, but highly stimulated by grazing cattle, which restrict 

competing species and create bare areas which are favourable 

for germination. 



A single site in France, in the Musella depression (Bonifacio limestone 

plateau, southern Corsica). 

In Sardinia (Italy) this species occurs at temporary pools of the 

Giara di Gesturi basaltic plateau 258 . 


In Corsica, the Musella population is increasing, slowly but surely 220 , 

but it has disappeared from the two Corsican stations at Vix and 

Vico 274 . 


24 



Flowering 

IUCN 1997 





and Red Lists 

Fruiting and dispersal 

of achenes 


Germination 

and dispersal 

of seedlings 

- 

- 

- 



13/05/1982, amended by the 




Book : priority species274 - Morocco: 

133, 181 


� 



At Musella, changes in the substrate have been observed as a 

result of ploughing and sowing. In addition, the cessation of cattle 

grazing by the site manager in 2002 and its replacement with 

grazing by a flock of sheep poses the risk, for the future, of altering 

the ecological conditions at the site. 

At Vix, development works destroyed the population and its 

habitat 274 . 


At Musella, the Eryngium pusillum population should not experience 

any problems in the short term. However, the development 

of woody plants (Blackthorn, Ulmus minor and Rubus ulmifolius 

could be detrimental in the medium term (shading). 



• Satisfactory at the Musella site up to 2001 as a result of the 

regular extensive cattle grazing. Moreover, the Musella pool is 

part of a Natura 2000* site. 

• Seed collection and ex-situ storage by the Conservatoire 

Botanique National Méditerranéen de Porquerolles, and precautionary 

cultivation of the species on Conservatoire land since 

1993. 


As shading has a detrimental effect on the species, surveillance 

and control of the spread of woody plants is advisable (scrub 

clearing, maintenance of grazing). Accordingly, keeping a flock of 

sheep at Musella for part of the year is strongly recommended. 

Bibliography 

Anonymous, 1999 12 ; Bissardon & Guibal, 1997 39 ; Fennane & Ibn 

Tattou, 1998 133 ; Géhu et al., 1994 156 ; Grime, 1979 163 ; Jahandiez 

& Maire, 1931-1934 181 ; Lorenzoni & Paradis, 1998 220 ; Mossa, 

1986 258 ; Olivier et al., 1995 274 ; Pignatti, 1982 292 ; Walter & 

Gillett, 1998 399 . 

Author: Paradis G.

Illecebrum verticillatum L. 

ANGIOSPERMS 

CARYOPHYLLACEAE 

Main synonyms 

Corrigiola verticillata (L.) Kuntze 

French name: Illécèbre verticillé 

Italian name: Corriggiola verticillata 

Portuguese name: Aranhoes 

Moroccan name: Souifa diel el ma 

English name: Coral-necklace 

Subspecies 

None 


• Annual herb with thread-like stem, often radicant, quadrangular, 

glabrous, pinkish. 

• Leaves opposite (can appear whorled due to the development 

of axillary clusters), entire, sub-sessile, obovate, 1.3 to 2.6 mm, 

slightly fleshy, with stipules small (1 mm) and scarious. 

• Inflorescence of pseudo-whorls*, each formed of two cymes* 

per node. Each cyme includes four to six flowers. 

• Flower: calyx with five sepals (1.5 to 2.5 mm), white, recurved, 

concave on the interior side, mucronate* at the apex. No corolla. 

Flower with five very small fertile stamens and five minute sterile 

stamens. Pistil with small ovoid ovary, with very short style 

and two stigmas. 

• Fruit: achene with membranous pericarp, retained within the 

persistent calyx. 

• Seed sub-elliptical, shining brown, about 1 mm long. 

The species exists in two very different forms, one terrestrial and 

one aquatic 63 : 

• The terrestrial form develops in spring when the pool has dried 

out. The stems are short (about 5 to 20 cm) and prostrate, with 

very short internodes. The flowers are very abundant. 

• The aquatic form develops in winter and spring. The stems are 

upright and unranked and may be up to 60 or 70 cm long, the 

internodes are very long and the submerged leaves are very fine. 

In March and April the upright stems develop floating branches 

at the surface of the water which bear a few inflorescences and 

resemble the terrestrial form, i.e. short internodes and more obovate 

leaves. The aquatic form produces very few flowers compared 

with the terrestrial form. 


The terrestrial form may be confused with Paronychia echinulata, 

but this lives on dry substrates. 

In the vegetative stage, confusion is also possible with Corrigiola 

littoralis, but this species has long basal leaves (up to 6 cm) in 

rosettes. 


63, 174, 292, 348 


Plant species 

This species is “widely distributed in Europe from southwest 

Spain to Sweden and Poland, in a band extending 200 to 400 km 

inland from the Atlantic seaboard” 348 . It is rarer in the 

Mediterranean region. 

France 

Western France and Corsica. 


Western half of Spain (absent from the Balearic Islands), 

Portugal, Italy (Lombardy, Piedmont, Tuscany, Marche, Lazio, 

Sardinia) and near the coasts of North Africa (Tunisia, Algeria and 

Morocco). 

Non-Mediterranean region 

Azores, Canaries. 

Habitat 

Illecebrum verticillatum 

1. “Mediterranean temporary pool” habitat 

1 cm 


In Corsica, Sardinia, North Africa and more rarely elsewhere, 

Illecebrum verticillatum is a component, significant in biomass* 

terms, of the flooded phase of several temporary pools (aquatic 

form). 

I. verticillatum is usually associated here with Apium crassipes, 

Ranunculus peltatus, R. ophioglossifolius, Myriophyllum alterniflorum, 

Baldellia ranunculoides, Eryngium barrelieri, Isoetes 

72, 219, 221, 298, 314 

velata, etc. 

I. verticillatum also occurs in its terrestrial form at the edges of 

pools, in small transitory rain pools within Mediterranean 

maquis, and in cupular pools* (for example in Corsica at Evisa). 

Here it forms communities in association with Anagallis parviflora, 

Cicendia filiformis, Exaculum pusillum, Kickxia cirrhosa, 

Radiola linoides, Silene laeta, Solenopsis laurentia etc. 54 

25







22.3412 Mediterranean aquatic communities with Isoetes: 

communities including Isoetes velata of water bodies of 

variable depth. 


In its aquatic form Illecebrum verticillatum is a member of the 

submerged phase of the Isoetion formations within the Isoetes 

velata communities. Some authors place the hydrophyte* formations 

to which it belongs in the communities of the Potamogetonetea 

pectinati class 221, 284 . 

The communities and associations* which include I. verticillatum 

and which are situated in places undergoing brief periods of 

flooding have received more attention from phytosociologists*. 

Thus, according Brullo & Minissale 54 , the phytosociological affinities 

of such communities and associations are as follows: 



- Alliance Cicendio-Solenopsion laurentiae 

Various associations, in which I. verticillatum is a 

species which ranks highly in terms of 

occurrence 

- Alliance Agrostion pourretii 

Association Illecebro-Agrostietum pourretii 

2. Other habitats 

Illecebrum verticillatum also occurs, in its terrestrial form, in 

other biotopes which flood temporarily and dry out rapidly at the 

end of the winter: ruts in tracks over a wide area of Europe, small 

transitory rain pools in clearings in the Atlantic heathlands, 

cupular pools (Massif Armoricain), or small bare depressions in 

overgrazed grasslands. 

Ecology 


Life form 

Therophyte (annual). 

Reproduction 

Aerial. 

Flowering 

From mid-March to the end of May (Corsica, Morocco). 


Stress-tolerant-ruderal (SR). 

Description of seeds 

Length approx. 1 mm. 


In the deep parts of temporary pools: 

• Germination during winter (from December). 

26 

• Growth during winter in the water body. 

• Flower branches on the water surface in April. 

• Flowering (limited) in April and May. 

• Fruiting during the drying-out of the pool (from May, often June). 

• Seeds in the dormant stage in summer and until the rains of 

autumn-winter. 

Morocco 

� 


Germination 

Corsica 



� 

Germination 


Flowering 

Flowering 


Fruiting 

Fruiting 

� 

Germination 

� 

Germination 

Hydrology 

Winter flooding or saturation of the sediment, even for a brief 

period, appears to be necessary for the germination of Illecebrum 

seeds. This germination, which takes place rapidly as soon as the 

first rain falls, enables the species to appear in the field almost 

every year 314 . 

Substrate 

Loose, very thin substrate (less than 3 cm deep) acid, siliceous, 

sandy or gritty and oligotrophic*. 


This strictly heliophilous* species does not tolerate cover from 

other species, hence its confinement to substrates that are very 

thin and poor in absorbable mineral salts. In very turbid pools 

(Mamora, Morocco) it only grows in abundance after the water 

has fallen. 


Disturbance resulting in the opening up of the vegetation is 

clearly favourable (e.g. trampling). 



In Corsica 

The species probably occurs at more than 20 sites. 

In Morocco 

There are over thirty stations distributed between the Rif, northern 

Atlantic Morocco, and mid-Atlantic Morocco. These sites are 

located on siliceous soils of the plains and low mountains subject 

to semi-arid, sub-humid and humid Mediterranean bioclimates* 

134 .

In Spain 

The species is present at 76 sites scattered through some twenty 

provinces 177 . 


In Corsica, Spain, Portugal and Morocco the species is not 

threatened, appearing to be stable despite more or less wide 

interannual fluctuations depending on the site. 


IUCN 1997 







In Corsica 

Up until now, there appear to be potential anthropogenic threats 

at very few sites. 

In Morocco 

Urban development and the filling-in of pools are the most 

important threats. Thus some sites in the Mamora forest have 

disappeared following the construction of the Rabat-Fez motorway. 

Some other sites close to large towns (Casablanca, Rabat, 

Benslimane, Bouznika, etc.) are also threatened by urban development 

in the medium term. 


In Corsica 

There is a risk that the likely discontinuation of extensive cattle 

grazing in the near future will result in a severe decline at several 

sites due to the spread of tall herbaceous plants (Schoenus 

- 

- 

- 

- 

- 

Plant species 

nigricans, Oenanthe globulosa etc.) and woody species (Phillyrea 

angustifolia, Erica scoparia, Myrtus communis, Pistacia lentiscus 

etc.). This is true of the temporary pools of southern Corsica, where 

the Illecebrum verticillatum population sizes are very small (Mura 

dell’Unda, Arasu, Muratellu pools), probably due to the cessation 

of cattle grazing several years ago which has led to the establishment 

of a heavy biomass of herbaceous plants. 

In Morocco 

Practically all the sites are grazed, allowing the vegetation to be 

kept open and preventing any possible spread by the surrounding 

woody plants. 



At present, in Corsica as well as in Morocco, there are no management 

measures, due to the lack of control over land ownership 

at the sites. 


• In Corsica, the maintenance (and/or re-establishment) of 

extensive cattle grazing is advisable, as well as the maintenance 

of routes for walkers and hunters. 

• In Morocco control of urban development in the short term is 

recommended. 

Bibliography 

Abbayes des, 1946 106 ; Anonymous, 1999 12 ; Bissardon & Guibal, 

1997 39 ; Brullo & Minissale, 1998 54 ; Casper & Krausch, 1981 63 ; 

Chevassut & Quézel, 1956 72 ; Fennane et al., 1999 134 ; Grime, 

1979 163 ; Herra 1990 174 ; Lorenzoni & Paradis, 1997 219 , 2000 221 ; 

Paradis et al., 2002 284 ; Pietsch, 1973 291 ; Pignatti, 1982 292 ; 

Pottier-Alapetite, 1952 298 ; Rhazi et al., 2001 314 ; Schumacker, 

1978 348 ; Sissingh, 1957 352 ; Valdés et al., 2002 391 . 


Collaborators: Grillas P., L. Rhazi & N. Yavercovski 

27


Isoetes duriei Bory (1) & 

Isoetes histrix Bory (2) 

LYCOPODIOPHYTES 300 

ISOETACEAE 

Main synonyms 

(1) I. duriaei Bory, I. durieui Bory 

(2) I. histrix Bory subsp. sicula (Tod.) P. Fourn. 

I. histrix Bory subsp. Delalandei (Lloyd) P. Fourn. 

French names: (1) Isoète de Durieu, (2) Isoète épineux 

Italian names: (1) Calamaria di Durieu, (2) Calamaria istrice 

English name: (2) Land Quillwort 

Subspecies 

None 

96, 300 


•Terrestrial perennial plants with a very short stem forming a 

kind of underground bulb bearing roots, and many fronds (“leaves”) 

arranged in rosettes at ground level. The fronds of Isoetes duriei 

are decurved towards the ground, while those of I. histrix are 

narrow and tough. 

• Bulb: usually firmly embedded, ringed with blackish spiny scales 

corresponding to the sclerified* remains of the bases of the preceding 

years’ fronds. 

• Fronds: dark green, 4 to 10 cm long, all fertile (sporophylls*), 

hollowed out at the base on the inner (upper) side to form a pit 

containing a sporangium. Outer fronds carrying macrosporangia, 

covered in a complete veil in I. duriei and a partial veil in I. histrix. 

Innermost fronds carrying microsporangia. 

• Macrosporangia containing macrospores (females) and 

microsporangia containing microspores (mâles). 


The two species are similar in appearance and in their ecology. 

One difference between them visible to the naked eye is the 

presence around the bulb of spiny scales, very obvious in Isoetes 

histrix, less conspicuous and briefly tridentate in I. duriei. 

Examination of the megaspores using a microscope or a good 

binocular lens allows the two species to be easily distinguished: 

Macrospores medium sized (0.4 to 0.6 mm) and with many 

tubercles in I. histrix. 

Macrospores larger (0.6 to 0.8 mm), with no tubercles and with 

a honeycombed surface in I. duriei. 



(1) Isoetes duriei 

Western Mediterranean species (distribution map in Quézel 306 ). 

France 

On the mainland, it is common at Les Maures and Estérel, fairly 

rare elsewhere (Biot Massif, between Béziers and Montpellier, 

southern catchment of the Caroux, southern foothills of the Montagne 

Noire, Albères, the Roussillon plain, valleys of the Gardon 

28 

1 cm 

1 cm 

Isoetes duriei 

frond base 

megaspore 

frond base 

0.35 mm 

megaspore 

0.25 cm 

Isoetes histrix 

0.25 cm 

0.35 mm 

rivers in the southern Cévennes); in Corsica, it is fairly common 

in the coastal zone and at low altitudes up to 300 m (but has 

been found exceptionally at 1,000 m). 


Portugal, Spain (west of the country and Catalonia), Balearics 

(Minorca), Tyrrhenian area of Italy (from Liguria to Calabria), 

Sardinia, Sicily, Greece, Cyprus, Algeria, Morocco and Turkey. 

(2) Isoetes histrix 

Mediterranean-Atlantic species (distribution map in Quézel 306 ). 

France 

It is very well represented along the Atlantic (Brittany, Poitou- 

Charentes, Pays de la Loire) and in Corsica, but local in the Mediterranean 

part of the mainland where it only occurs at Les Maures 

(Hyères region and the northern edge of the Plaine des Maures).


Central and western Iberian Peninsula (Spain, and all the Portuguese 

provinces), mainland of Italy, Sardinia, Sicily, Macedonia, 

Greece, Crete, Malta, Croatia, Morocco, Algeria, Tunisia, Middle 

East (Syria, Lebanon) and Turkey. 

Habitat 


The most suitable habitats are short Mediterranean swards, wet 

or waterlogged in winter and the beginning of spring and very 

dry in summer. The substrate is usually siliceous (silts and sands) 

and relatively rich in organic matter. At Les Maures and the 

Estérel, Isoetes duriei also grows equally well in soils with little 

organic matter where, however, it grows to a smaller size than in 

accumulation depressions. 

These kinds of habitat, which are not (or are only exceptionally) 

flooded, are located on the edges of temporary pools and streams, 

on various flat areas and in wet clearings in the maquis 48, 233, 296 . 

The communities described based on I. histrix are a little less wet 

than those defined for I. duriei, and form a stage transitional to 

the annual swards of drier biotopes (with Tuberaria guttata and 

Anthoxanthum ovatum). 


(1) I. duriei 

• In Corsica: Ophioglossum lusitanicum, various Serapias, Aira 

capillaris, Bellis annua, Cicendia filiformis, Radiola linoides, Linum 

bienne, Anagallis arvensis subsp. parviflora, Lythrum hyssopifolia, 

Juncus bufonius, J. tenageja, J. capitatus, Ranunculus revelieri and 

Lotus conimbricensis. 

• In mainland France 21, 22, 215 : the same species, with in addition 

Isolepis cernua (=Scirpus savii), Nasturtium asperum, Agrostis 

pourretii, Airopsis globosa, Veronica acinifolia, Juncus pygmaeus, 

J. fasciculatus and Mentha pulegium. 

(2) I. histrix 

• Species common to the various sites: Radiola linoides, Linum 

bienne, Anagallis arvensis subsp. parviflora, Serapias lingua, Juncus 

bufonius, Isolepis cernua (= Scirpus savii), Bellis annua. 

• At the former sites of the littoral Maures, where the species 

has not recently been seen: Allium chamaemoly, Romulea columnae 

and Isoetes duriei 22 . 

• In Morocco: Poa annua, Filago (= Logfia) gallica, Rumex bucephalophorus 

and Bromus mollis 275 . 


• At the edges of pools (1) and (2): 


• In the absence of disturbance, humid grassland vegetation 

often invades the short swards where Isoetes histrix occurs, and 

it is then included in the following habitat: 

“Mediterranean tall humid grasslands of Molinio-Holoschoenion” 

(code 6420). 


22.34 Amphibious southern communities (Isoetalia). 


22.3411 Terrestrial Quillwort communities: formations 

with Isoetes histrix, I. duriei, of ephemeral aquatic habitats. 

Plant species 


Owing to the difference in ecology between Isoetes velata (of 

periodically flooded habitats) on the one hand, and I. duriei and 

I. histrix (of non-flooded habitats) on the other, de Foucault 98 

split the Isoetion of Braun-Blanquet 48 and Rivas-Goday 306, 319 into 

two alliances: 

The Antinorio agrostideae-Isoetion velatae for topographically 

low-lying areas, periodically flooded. 

The Ophioglosso lusitanici-Isoetion histricis for topographically 

higher areas, not flooded. 

The Quillwort communities of non-flooded habitats are therefore 

included in the Ophioglosso lusitanici-Isoetion histricis alliance, 

order Isoetalia velatae and class Isoetea velatae 98, 194 . 

In the Isoetion s.l. (Isoetalia), the habitat of these two species 

corresponds to the following communities: 

(1) I. duriei 

• Isoetetum duriei association*, defined in Languedoc 48 and also 

described from Spain 319 . 

• Association with I. duriei and Juncus capitatus described from 

Corsica 233 . 

• Association with I. duriei and Nasturtium asperum described at 

Les Maures 21 . 


• In France, in the Serapion alliance, Aubert & Loisel 1971: subassociation 

with I. histrix of the Serapio-Oenanthetum, described 

by Barbéro 22 from Les Maures. 

• In Spain, in the Isoetion alliance 319 : 

- Association Isoetetum histricis s. l. 

- Association Wahlembergio-Isoetetum histricis. 

• In Corsica, in the Ophioglosso lusitanici-Isoetion histricis (Br.-Bl. 

1931) de Foucault 1988: community with I. duriei and I. histrix. 

• In North Africa, in the Isoetion s.l.: association with I. histrix 

and Radiola linoides, described by Chevassut & Quézel 72 . 

Ecology 


Life form 

Perennial species: bulb-forming geophyte* 292 . 

Adaptive strategy 163 

Stress tolerant-ruderal (S-R). 

Reproduction 

Aerial for maturation of spores. 

Description of spores (macrospores: see above, § “Similar species”) 

• (1) granular microspores. 

• (2) microspores covered with fine entangled fibrils. 


• In France: growth of fronds with the onset of the first autumn 

rains 21 and in winter when the substrate is very wet; ripening of 

spores in spring, followed by rapid withering of the fronds at the 

end of spring, when the habitat is drying out rapidly. 

29


•In Morocco: the cycle of Isoetes histrix begins at the end of 

winter after the first rain and ends at the beginning or towards 

the middle of spring with the formation of spores. 

Isoetes duriei 

France 

30 


� 

and vegetative growth 

Isoetes histrix 

France 

� 

Maturation of spores 


Germination 


vegetative growth 

Morocco 

� 








� 

� 

Germination and 

Hydrology 

A very wet substrate is necessary at the beginning of the cycle 

but these plants do not like to be submerged for long. 

Substrate 

(1) not (or very slightly) oligotrophic* in Corsica, oligotrophic on the 

mainland (Permian sandstones at the Plaine des Maures, rhyolites 

at the Colle du Rouet). 

(2) more or less oligotrophic (usually sand-silt). 


Low growing species, light-demanding (heliophilous*) and consequently 

not very tolerant of competition from taller plants. 


The main perturbations are the “ploughing” of the substrate by 

wild boars, and trampling (by cattle and people). They are beneficial 

as they limit the amount of cover and litter produced by taller 

plants. 

Temperature 

It appears to govern the distribution of Isoetes histrix at Les 

Maures, where it is confined to very warm coastal stations. It is 

usually replaced by I. duriei away from the coast, in the Maures 

plain and massif in particular 22 . 

� 



(1) I. duriei 

• France (mainland and Corsica): large number of sites (see 

above, § Distribution), but number of stations within the sites 

impossible to estimate accurately, given the very scattered nature 

of the species’ distribution. 

• Spain: not very widespread (provinces of Cadiz, Huelva, Gerona, 

Badajoz, and one station on Minorca 64, 79 ). 

•Portugal: locally present in six provinces. 

- Italy: fairly rare (Sardinia, Sicily, Capraia, Liguria and west coast 

of the peninsula) 137, 292 . 


• France: large number of sites in Corsica, but only three 

remaining populations in the Mediterranean part of the mainland 

(Hyères, La Londe-les-Maures, Le Cannet-des-Maures). 

• Spain: four stations on Minorca, one on Majorca 331 , and present 

in 15 provinces in the west of the peninsula and three provinces 

in the northeast. 

•Portugal: in all provinces 64, 79 . 

• Italy: rare species (Sicily, Sardinia, and the Tuscany, Lazio and 

Puglia regions) 292 . 


(1) I. duriei 

The populations appear to be stable. 


The populations appear to be stable in Corsica and Morocco as 

they are fairly resistant to anthropogenic and natural threats. 

They have declined in mainland France, in Roussillon (extinction 

due to the destruction of the Saint-Estève pool) as well as at Les 

Maures (not found recently in the Saint-Tropez region, or at the 

capes of Lardier, Benat, and Taillat, but recently rediscovered at 

Hyères, at La Londe-les-Maures and Le Cannet-des-Maures). In 

Malta, the species has not been seen again since its discovery at 

the end of the 19th century 344 . 

Legal status/level of protection 

IUCN 1997 





and Red Lists 

- 

- 

- 

France (1) and (2), national list: 


Officiel 13/05/1982, amended by 



- Balearics (2): “endangered” 

(EN) 331 

- Malta (2): “presumed extinct” 

(Ex ?) 344



France 

In Corsica, development projects (such as construction works) are 

a potential threat to some coastal sites. 

In Provence, in Les Maures, urbanisation and tourist developments 

(golf courses) have destroyed a large number of Isoetes 

duriei stations*. Similarly, development along the coastline has 

destroyed all the coastal I. histrix stations (region of Saint-Tropez, 

Le Lavandou, Les Bormettes), and constitutes a threat to the other 

stations. 

Morocco 

Along with urban development, the greatest threat to the I. histrix 

populations is the infilling of pools with debris. 


• In the absence of disturbance, the closing-up of the habitat due 

to the spread of taller plants (Schoenus nigricans, Scirpus holoschoenus, 

Dittrichia viscosa and especially various maquis species) 

could constitute a threat for several sites in Corsica. This is equally 

true at Les Maures where the cessation of grazing is favouring 

the growth of perennial herbaceous species (Paspalum dilatatum, 

Juncus conglomeratus, Scirpus holoschoenus), and even maquis 

species. This is not the case at the Moroccan sites, where grazing is 

commonly practised. 

• The frequency of dry years does not affect the Isoetes histrix 

populations, which sustain themselves by their spores as much as 

by their bulbiform rhizome. 


(1) I. duriei 

No extinctions currently known in Corsica or on the mainland. 


No extinctions currently known in Corsica. On the mainland, the 

inherent risks to populations are low, but possible, due to the 

limited numbers and the very marked isolation of each of the 

populations, which are not very vigorous. 

Management measures 


No management either in Corsica or on the mainland, or in 

Morocco, but grazing by free-ranging cattle, wild boar, and the 

movements of hunters and walkers are unintentional management 

activities which are beneficial to these Quillworts. 

Plant species 

At Les Maures: 

• The clearing of firebreaks (provided that it is sensitive, i.e. that 

it does not upset the microtopography), and especially their 

grazing, are beneficial. 

•A Natura 2000* area, in the process of being established in the 

Plaine des Maures, provides the opportunity to take the requirements 

of management for the conservation of I. duriei into 

account, as does a Réserve Naturelle project that includes the 

majority of the stations. 

• The LIFE “Temporary Pools” project has enabled experimental 

testing of the management of plant communities with I. duriei to 

be carried out, as well as the purchase by the CEEP of land where 

this species grows. 


France 

In Corsica, retention of extensive cattle grazing at the sites 

where it is still present, and its reintroduction at sites where it is 

no longer practised. 

In mainland France: 

• Retention of horse grazing (one Isoetes histrix station) and 

pedestrian traffic (two I. histrix stations); avoidance of destructive 

activities (mountain biking,* scrambling, fly-tipping). 

• Reintroduction of extensive grazing, beneficial for the conservation 

of I. duriei, at Les Maures. 

Morocco 

Avoidance of repeated cultivation of the catchment areas nearest 

to I. histrix sites, as the colonisation of the habitat by arable 

weeds may have a negative effect (competition). 

Bibliography 

Anonymous, 1999 12 ; Barbero, 1965 21 , 1967 22 ; Bissardon & 

Guibal, 1997 39 ; Braun-Blanquet, 1935 48 ; Castroviejo, 1986- 

2001 64 ; Chevassut & Quézel, 1956 72 ; Cirujano et al., 1992 79 ; 

Danton & Baffray 1995 96 ; Foggi & Grigioni, 1999 137 ; Foucault 

de, 1988 98 ; Grime, 1979 163 ; Julve, 1993 194 ; Loisel, 1976 215 ; 

Malcuit, 1962 233 ; Olivier et al., 1995 274 ; Ould Louleid, 1991 275 ; 

Pignatti, 1982 292 ; Poirion & Barbero, 1966 295 ; Prelli, 2001 300 ; 

Quézel et al., 1979 308 ; Quézel, 1998 306 ; Rivas Goday, 1970 319 ; 

Saez & Rossello, 2001 331 ; Schembri & Sultana, 1989 344 . 


Collaborators: Catard A., H. Michaud, L. Rhazi, D. Titolet & 

N. Yavercovski 

31


Isoetes setacea Lam. (1) & 

Isoetes velata A. Braun (2) 

LYCOPODIOPHYTES 300 

ISOETACEAE 

Main synonyms 

(1) I. delilei Rothm, (2) I. variabilis Le Grand 

French names: (1) Isoète sétacé, Isoète grêle, 

(2) Isoète à voile, Isoète voilé 

Italian names: (1) Calamaria setacea, (2) Calamaria velata 

Moroccan name: (2) Lehyet-el-rebb 

English name: (1) and (2) Quillwort (genus name) 

Subspecies: (2) Isoetes velata A. Braun subsp. velata 

I. velata A. Braun subsp. intermedia (Trabut) Maire & Weiller 

I. velata A. Braun subsp. tegulensis Batt. & Trabut 

I. velata A. Braun subsp. perralderiana 

I. velata A. Braun subsp. tenuissima (Boreau) O. Bolos & Vigo 

96, 299 


•Perennial amphibious plants: aquatic at least in winter and 

early spring, then terrestrial. 

• Habit is an upright tuft when submerged. Stem very short, 

forming a kind of bulb which bears roots and many slender, fragile 

fronds. 

• Fronds (“leaves”) pale green, long, 10 to 40 cm (1), 5 to 15 cm 

(2), linear and arranged in a dense rosette. These fronds are fertile 

(sporophylls*): each has a depression at its base on the interior 

(upper) side, occupied by a large sporangium. The sporangia 

of I. setacea are bare, those of I. velata are partly or completely 

covered by a well-developed veil. 

• Outer fronds bear macrosporangia (bare or covered with a veil 

according to species), containing macrospores. The inner fronds 

bear microsporangia (bare or covered depending on the species), 

containing microspores, which are very abundant. 

• Main difference between the two species: a very obvious veil 

over the sporangia of I. velata, while this veil is absent in I. setacea. 


• The more terrestrial Quillworts (Isoetes histrix and I. duriei) 

during the dry phase have their bulbs surrounded by old, persistent 

sclerified* frond bases; these scales are absent in I. setacea 

and I. velata whose bulbs, at least in winter and spring, have a 

whitish appearance 299 . 

• The subspecies I. velata subsp tenuissima is endemic to central 

France, so there is no risk of confusion with subspecies velata. 



(1) Isoetes setacea 

Western Mediterranean species (distribution map in Quézel 306 ). 

32 

1 cm 

1 cm 

Isoetes setacea 

Isoetes velata 

megaspore 

frond base 

megaspore 

frond base 

0.25 mm 

0.2 cm 

0.25 mm 

0.2 cm 

France 

Extremely rare plant, occurring in: 

• The Hérault: temporary pools in the Plateau de Roque-Haute 

and the Plaine de Béziers (Grand-Bois). 

• The Pyrénées-Orientales: Torremila pool and the Plateau de 

Rodès (Fenouillèdes). 

Note: Quézel’s map 306 wrongly includes Corsica for I. setacea. 


Spain (centre and west of the country and in Catalonia), Balearics 

(Minorca), Portugal (centre and south), and the Maghreb (Morocco, 

where it has recently been discovered). 

(2) Isoetes velata 

Mediterranean-Atlantic species (distribution map in Quézel 306 ). 

France 

Rare on the mainland (Plaine des Maures and Estérel in the Var, 

Plateau de Rodès in the Pyrénées-Orientales) and fairly well represented 

in Corsica.


• Spain (in more than half of the country, mainly in the west and 

on the island of Minorca), Portugal (in almost the whole of the 

country), mainland Italy, Sicily, Sardinia (subspecies tegulensis) 86 

and Greece. 

• Maghreb (Morocco, Algeria, Tunisia, Libya). The subspecies 

intermedia and tegulensis occur in Tunisia, Algeria and Morocco, 

subspecies perralderiana in Algeria. The subspecies velata occurs 

in all the countries of the species’ Mediterranean range. 

Habitat 


(1) I. setacea 

At Roque-Haute, I. setacea lives in pools hollowed out in the 

Quaternary basalt. 

(2) I. velata 

Habitats suitable for I. velata are flooded in winter and spring, 

and very thoroughly dried out in summer. Their substrate is thin, 

composed of compact rock (granite, schist, rhyolite) or of very 

fine sediment with a very low content of organic matter (sand, 

silt and clay). Most of the sites are on siliceous rock, but the temporary 

pool of Padulu (Bonifacio, Corsica), has a calcareous subsoil. 

The floodwater is oligotrophic* and its depth varies between 

sites and between years: approximately 10 to 60 cm. 



Braun-Blanquet 48 described an Isoetetum setacei for these pools, 

with the following companion species: Juncus pygmaeus, Myosotis 

sicula, Lythrum borysthenicum, Cicendia pusilla, Lotus angustissimus, 

Lythrum thymifolium and Marsilea strigosa. 


Species associated with Isoetes velata are varied, according to 

the depth of water (zonation) the time of year (phenophase) and 

also the substrate 21, 22, 219, 221, 284, 302 : 

• Aquatic plants: Tolypella glomerata (Charophytes), Callitriche 

truncata, Ranunculus peltatus s.l., R. ophiogossifolius, Myriophyllum 

alterniflorum, Illecebrum verticillatum, Pilularia minuta, 

Apium crassipes, Alopecurus bulbosus, Baldellia ranunculoides, 

Littorella uniflora, Glyceria fluitans, etc. 

• Other species, more or less associated with flooding: Crassula 

vaillantii, Lythrum borysthenicum, Juncus pygmaeus, Myosotis 

sicula, Laurentia michelii, Ranunculus sardous, Scirpus setaceus, 

Agrostis pourretii, Bellis annua, Elatine brochonii, Corrigiola littoralis, 

Hypericum tomentosum, etc. 







including Isoetes setacea or I. velata of water 

bodies of variable depth. 


Plant species 


• For France, Braun-Blanquet 48 described the Isoetetum setacei 

(see above). 

• For Spain, Rivas Goday 319 considered I. setacea to be a characteristic 

species of the Cicendion alliance (order Isoetalia). 


Various communities and associations* with I. velata have been 

distinguished around the Mediterranean, for example: 

• In North Africa, associations with I. velata and Myosotis sicula, 

and the Eryngium barrelieri and I. velata association 72, 298 . 

• In the Colle du Rouet Massif (northwest of the Estérel), the 

I. velata and Crassula vaillantii association 295 . 

• In Corsica, Lorenzoni & Paradis 219, 221 and Paradis et al. 284 

described some further I. velata communities. 

Note 

Owing to the wide ecological differences between, on the one 

hand, I. setacea and I. velata living in periodically flooded habitats, 

and on the other hand I. duriei and I. histrix living in nonflooded 

habitats, de Foucault 98 split the Isoetion of 

Braun-Blanquet 48 , Malcuit (for I. velata) 233 and Rivas-Goday 319 

into two alliances 306, 308 : 

• The Antinorio agrostideae-Isoetion velatae for low topographical 

levels, periodically flooded. 

• The Ophioglosso lusitanici-Isoetion histricis for higher topographical 

levels, not flooded. 

The I. setacea and I. velata communities are thus included in the 

Antinorio agrostideae-Isoetion velatae alliance, order Isoetalia 

velatae, class Isoetea velatae. 

Julve 194 accepted this classification. 

Ecology 


Life form 

•Perennial species. However, annual populations of Isoetes 

velata have been identified in the cupular pools of the Colle du 

Rouet Massif (Var); they have a very low level of reproductive 

success 295 . 

• (1) hydrophyte* 292 and geophyte* 48 . 

• (2) hydrophyte 292 , geophyte 48 and sometimes therophyte* 295 


Stress tolerant-ruderal (S-R). 

Reproduction 

Maturation of spores: (1) aquatic (2) aerial. 

Description of spores 

The megaspores of the two species are fairly similar. Those of 

I. setacea (0.4 to 0.9 mm) have a few inconspicuous tubercles 

and those of I. velata (0.4 to 0.5 mm) have more or less numerous 

tubercles on their various sides. The microspores are echinate 

when mature. 

33



Growth of fronds in the water during winter (from the first 

autumn rain on the mainland for Isoetes velata); maturation of 

spores in spring (1), or after the water has receded (2); rapid 

death of fronds at the beginning of summer, when the habitat is 

no longer flooded and dries out severely. 

Isoetes setacea 

� 

34 



Isoetes velata 

� 



Germination 






Germination 

Hydrology 

These species grow in wet soil and can tolerate water depths of 

up to 50 cm. The duration of the periods of flooding (six months 

or more) and drought (sometimes reduced to four or five months) 

varies depending on climatic conditions. Spores begin to germinate 

in autumn (November) in soil which is saturated with water 

or flooded. The hydrological conditions have a significant effect 

on the growth of Isoetes setacea. 

Substrate 

Usually non-calcareous, oligotrophic and thin. 


Light-demanding species (heliophilous) and therefore intolerant 

of competition. 


Favourable, by opening up of the habitat. 




France 

I. setacea occurs in about 100 pools in the Roque-Haute Nature 

Reserve, in one pool in the Béziers plain, one at Torremila, several 

pools and ditches at Rodès. 

Spain and Portugal 

It occurs in the centre and west, as well as in Catalonia and on 

the island of Minorca (13 provinces in Spain and six in Portugal) 64 . 

� 

� 


France 

• In Corsica, about twenty sites. 

• On the mainland, a single station in the Roussillon, with the 

remaining populations confined to the Var: 

- In the Plaine des Maures, three small recently discovered stations. 

- In the Plaine de Palayson, the Catchéou pool and a few temporary 

streams. 

- In the Estérel, about fifty small cupular pools* on the rhyolite 

escarpments of the Colle du Rouet (distributed between about 

fifteen stations), and the Barres de Roussiveau (a few stations). 

Morocco 

More than 50 stations for Isoetes velata. 


The species is scattered throughout the Iberian Peninsula (apart 

from the northern and western provinces), and on the island of 

Minorca 64 . 

Italy 

Very rare species (Sicily, Sardinia, Tuscany and Lazio regions). 


(1) In mainland France a more or less slow decline of the populations 

of Isoetes setacea and of the Isoetion plant associations 

is taking place. In particular, the populations of the pools at 

Saint-Estève (Pyrénées-Orientales), Grammont (Hérault), and 

Redessan (Gard) have disappeared. 

(2) In Corsica as well as in Morocco, at sites with loose, thin, even 

overgrazed substrates, the populations of I. velata are either stable 

or expanding. On the French mainland the populations in the 

Roussillon (Saint-Estève) and the Gard (Redessan, Pazac) have 

decreased, while they appear to be stable at Les Maures and 

Estérel. 


IUCN 1997 



National and regional 

protection 


and Red Lists 

- 

- 

- 

- France (1) and (2), national list: 



Decree of 31/08/1995/ 



priority species274 - Italy, national Red Data Book: (2) 

“vulnerable” (VU) 86 

- Balearics: (2) “vulnerable” (VU) 331 

- Morocco: (1) “very rare” (RR), 

133, 181, 380, 381 

(2) “rare?” (R?)




At Roque-Haute, the discontinuation of sheep grazing, accumulation 

of organic matter and significant shading are having an 

effect on the growth of I. setacea (reduction in weight of bulbs 

and in number and length of fronds) and on its reproduction 

(reduction in number of macrosporangia, weight of macrospores, 

number and weight of microsporangia 315 ) (vol. 1, box 46). 


• In France: the spread of agriculture has destroyed some 

I. velata stations in the Costière Nîmoise, as well as the principal 

station in the Pyrénées-Orientales (Saint-Estève pool). In Les 

Maures and Estérel, urbanisation and developments affecting the 

nature of the soil (plantations, golf, improvement of grazing in 

the maquis by overseeding and fertilisation) have caused a 

reduction in the number of pools capable of supporting this 

species; the Saint-Raphaël and Fréjus stations, among others, 

have disappeared. Some small artificial hill reservoirs are apparently 

becoming suitable for its colonisation. 

• In Morocco, the problems are mainly associated with infilling, 

extraction of sediment and drainage of pools. 


In mainland France and Corsica (but not in Morocco, where grazing 

still takes place) the closing up of the habitat and the accumulation 

of litter, resulting from the increase of tall plant species 

(Schoenus nigricans, Scirpus holoschoenus, Dittrichia viscosa and 

various maquis species), is deleterious to these species, as is 

infilling by eroded sediments (Plaine des Maures and Colle du 

Rouet). 


(1) No short-term risk of extinction. 

(2) In Corsica, several small stations are threatened with disappearance 

as a result of the closing up of the habitat (pools of 

Muratellu, Arasu and Mura dell’Unda). The recently rediscovered 

Roussillon population is very vulnerable due to its small size. 




The Roque-Haute site has legal “Réserve Naturelle” (Nature 

Reserve) status, which is not being implemented at present due 

to the impossibility of gaining access to the station (no control 

over usage of the site). The I. setacea sites have been the subject 

of a survey addressing water levels, the dynamics of woody vegetation 

and clonal* species 315 . 


In France and Morocco, the opening up of the tallest vegetation 

through grazing by cattle (Tre Padule Nature Reserve, Capandula 

pools in Corsica, Plaine de Palayson on the mainland) and by 

sheep (Padulu site in Corsica) is beneficial. 

In France, in the Var, further measures are in progress: 

Plant species 

• Diagnosis of management problems and implementation of a 

protocol for monitoring the cupular pools at the Colle du Rouet 

as part of the LIFE “Temporary Pools” project. 

• Classification in the “ecological series” by the ONF* of the 

Catchéou pools (Plaine de Palayson), as a prelude to enabling the 

natural-heritage issues associated with these pools to be integrated 

into forestry management. 

• Establishment of Natura 2000* areas at the Var sites, providing 

the opportunity to include the protection of Quillwort in the 

conservation objectives for these sites. 



• Ensuring that the pools are not kept permanently flooded. 

• Reintroduction of grazing in the pools with helophytes*, and in 

the pools with woody plants (after clearing). 

• Removal of litter from the pools to allow the spore stocks to 

germinate satisfactorily. 

• Evaluation of the dynamics of the vegetation following the 

introduction of grazing (and any modifications of management). 


•In France: 

- In Corsica and on the mainland, maintenance of cattle (and 

sheep) grazing at the sites where it still takes place and its reintroduction 

at sites where it is no longer practised, while avoiding 

intensive practices (overseeding, fertilisation), and following 

clearing operations at sites which have been heavily colonised by 

woody vegetation (Corsica). 

- At the Colle du Rouet, implementation of monitoring at the 

cupular pools in the context of contractual management which 

is being put in place by the ONF, the commune and the CEEP, and, 

in the case of pools which are silting up, planning for minor 

intervention (removal of litter). 

• In Morocco: 

Discouraging any modification of the hydrology of the pools 

(drainage, infilling) as well as extraction of sediment, which 

destroys the seedbank. 

Bibliography 


Guibal, 1997 39 ; Braun-Blanquet, 1935 48 ; Castroviejo, 1986- 

2001 64 ; Chevassut & Quézel, 1956 72 ; Conti et al., 1992 86 ; 

Danton & Baffray, 1995 96 ; Fennane & Ibn Tattou, 1998 133 ; 

Foucault de, 1988 98 ; Gaudillat & Haury, 2002 153 ; Grillas & Tan Ham, 

1998 162 ; Grime, 1979 163 ; Jahandiez & Maire, 1931-1934 181 ; Julve, 

1993 194 ; Lorenzoni & Paradis, 1997 219 , 2000 221 ; Malcuit, 

1962 233 ; Médail et al., 1998 246 ; Molina, 1998 251 ; Olivier et al., 

1995 274 ; Paradis et al., 2002 284 ; Pignatti, 1982 292 ; Poirion & 

Barbero, 1965 295 ; Pottier-Alapetite, 1952 298 ; Prelli, 2001 300 ; 

Quézel & Zevaco, 1964 302 ; Quézel et al., 1979 308 ; Quézel, 

1998 306 ; Rhazi et al., sous presse 315 ; Rivas Goday, 1970 319 ; Saez 

& Rossello, 2001 331 ; Titolet & Oualidi, 2000 380 ; Titolet & Rhazi, 

1999 381 . 


Collaborators: Catard A., H. Michaud, L. Rhazi, M. Rhazi & 

N. Yavercovski 

35


Littorella uniflora (L.) Ascherson 

ANGIOSPERMS 

PLANTAGINACEAE 

Main synonyms 

L. lacustris L. 

French name: Littorelle à une fleur 

Italian name: Littorella 

English name: One-Flowered Shoreweed 

Subspecies 

None 

63, 292 


Perennial herbaceous plant, hydrophytic* or amphibious, with 

two morphological types: a sterile submerged (aquatic) form and 

a terrestrial flowering form. 

Submerged form 

It has dense rosettes of leaves and stolons. It does not flower. 

Each rosette consists of: 

•A vertical rhizome, very short, very flattened (4-5 mm thick), 

covered by the remains of dead leaves. 

• Three to 14 leaves, light green, thick, cylindrical, stiff, glabrous, 

broadly elliptical in cross section, swollen at the base and finely 

pointed at the tip, with air vessels, and measuring 3 to 15 cm long 

and 2 to 4 mm thick. These submerged leaves have no stomata. 

• Many adventitious roots on the rhizome. 

From the leaf axils arise stolons, 3 to 60 cm long, bearing leaves 

that are reduced to scales. At their tips, the stolons form new 

rosettes of leaves and take root in the soil by forming adventitious 

roots. Hence the stolons enable the plant to propagate itself 

vegetatively to a considerable degree under the water. 

Terrestrial form 

This results form the transformation of the aquatic form after the 

water has dropped. It takes the form of dense rosettes of leaves 

but does not produce stolons. It produces flowers (the species is 

monoecious). 

Each rosette includes: 

•A very short vertical rhizome with adventitious roots. 

• Leaves many (5 to 25), small (2.5 to 4-10 cm long) and with 

many stomata 321 . The upper surface, especially of the internal 

leaves, has a longitudinal groove, while the lower surface is 

semi-cylindrical. 

The male flowers are borne at the tips of fairly long peduncles 

with the female flowers at their bases. The male flowers are small 

(sepals 4 mm long and stamens with filaments 10-20 mm), the 

scarcely visible female flowers are protected by foliar sheaths 

and surrounded by long woolly hairs. The flowering of each plant 

is usually rapid and concentrated over one to two weeks 321 . 

Flowering follows the emergence of the substrate from the 

water. 

Fruit dry, indehiscent, with one seed (achene), 2 mm in length. 

36 


Littorella is subject to confusion with: 

•Young stages of Baldellia ranunculoides. However, this species 

fairly quickly develops lanceolate leaves and has the scent of 

coriander when crushed. 

• The well-advanced stages of some Quillworts (such as Isoetes 

velata), but their lack of stolons enables them to be distinguished. 


63, 292 


Littorella uniflora is a west European and sub-Atlantic species, 

absent from the areas around the Mediterranean apart from Corsica, 

Sardinia, central and southern Iberian Peninsula (Medina, 

pers. com.) and Morocco. It extends from latitude 38°45’ in the 

south (at Lisbon) to 68°20’ in the north and, in longitude, from 

the Azores (31°W) to Karelia (35° E). Over this very wide range, 

its distribution is discontinuous. 

France 

A species of the Atlantic seaboard, but also occurs in Corsica. One 

locality, not recently confirmed, in the Crau (Etang du Luquier, 

Bouches-du-Rhône). 


Morocco in the Rif region, Sardinia (where it is extremely rare 

and reaches the southern limit of its European range), northern 

Italian provinces (in a non-Mediterranean climate), Spain and 

Portugal. 

Habitat 

1 cm 

Littorella uniflora 


1. Commonest habitats in Europe (non-Mediterranean) 

Littorella uniflora occurs on the edges of oligo-mesotrophic* water 

bodies: 

• Lakes and pools in peaty areas, fish ponds, sand and gravel 

pits, ditches. 

•Periodically flooded shallow pools on various substrates (silt, 

sand, gravel and even stones). The optimum living conditions are 

provided by water bodies that dry out in summer and have a 

maximum depth of 1.5 m in winter.

2. Mediterranean temporary pools 

In southern Corsica, Littorella uniflora currently occurs in six 

temporary pools: four in the Tre Padule de Suartone Nature Reserve 

and two pools, of artificial origin, on the Frasselli plateau. 

It is associated with the following species: Isoetes velata, Pilularia 

minuta, Baldellia ranunculoides and Ranunculus ophioglossifolius. 


In the Mediterranean region: 






containing Isoetes velata in water bodies of variable 

depth (where I. velata has a much earlier phenology than 

Littorella uniflora). 


Littorella uniflora is a companion species in the Isoetes velata communities 

(Corsica). Elsewhere it is characteristic of formations of 

the class Littorelletea in Spain 79 and in northwest Europe where 

it is also a component of submerged plant associations* of the 

class Potamogetonetea pectinati 47, 82,127 . 

Ecology 


Life form 

Perennial species: hemicryptophytic* hydrophyte*. 

Reproduction 

Aerial. 

Flowering 

In Corsica from mid-May to the end of June, and in April-May in 

Morocco. 


Stress-tolerant (S). 


At the beginning of spring, under water 10 to 30 cm deep: 

• The achenes germinate and give rise to new individuals which 

send out stolons that produce rosettes (submerged form). 

• The rhizomes break dormancy and produce short stolons giving 

rise to new rosettes (submerged form). 

At the end of spring, when the pool has dried out or in very shallow 

water (less than 10 cm): 

•Transformation of rosettes from the submerged form to the 

terrestrial form (beginning of May). 

• Flowering of plants that have emerged or are under very shallow 

water (end of May and June). 

In summer: 

• Fruiting (June and beginning of July). 

•Transition of achenes and short rhizomes into the dormant 

state. 

Corsica 


Plant species 


Morocco 

Germination and development of stolons 

Transformation aquatic � terrestrial form 



Flowering 

Flowering 

Fruiting 

Hydrology 

Winter flooding is essential for germination and for the activation 

of the buds on the short rhizomes to produce stolons. Emergence 

from the water is essential for flowering 117, 127 . 

Substrate 

Substrate acid, loose, sometimes pisolithic, fairly thin (less than 

10 cm deep), of variable particle size (silt, sand, gravel), without 

(or with little) organic debris and oligo-mesotrophic. 


A heliophilous* plant, intolerant of cover from tall species (Eleocharis 

palustris, Phragmites australis, various Scirpus). 


Low-intensity disturbance (grazing, trampling) that causes a 

small amount of substrate compaction is beneficial 127 . 

Drainage, by altering the hydrology of the pools, may have an effect 

on the germination of the achenes and sexual reproductive success. 

Light 

Light-demanding (heliophilous plant), which could explain its 

absence from turbid pools in Morocco. 



Fruiting 

• France: 

- In Corsica, only 6 sites (see above). 

- In non-Mediterranean France: the species has a scattered distribution 

in the western, central, eastern and extreme northern 

regions of the country 96 . 

• Italy: several scattered sites in the northern regions (Piedmont, 

Lombardy, Venetia, Emilia-Romagna); a single site in Sardinia 292 . 

• Spain and Portugal: about a hundred sites scattered through 

the northwestern half of the Iberian Peninsula (Medina, pers. 

com.). 

• Morocco: the species has been recorded only in the sub-humid 

to humid mountainous region of the Rif (Issaguene dayet). 

37



• In Mediterranean France: in Corsica the populations fluctuate 

from one year to another, but appear to be stable in the medium 

term at the Tre Padule pools. There is a lack of data from the 

Frasselli pools. The four plants recorded by Lorenzoni 225 at the 

Mura dell’Unda pool (north of Porto-Vecchio) have not been 

found again (looked for in 2001, 2002 and 2003). Gamisans 145 

reported its occurrence at the Lac de Ninu (1,800 m altitude), but 

the species could not be found there during a recent study of the 

lake. In the Crau, where the species was collected in 1954 at the 

Etang du Luquier 257 , it has not been seen again and could have 

disappeared (changes in water quality, invasion of the banks by 

Paspalum distichum). 

• In non-Mediterranean France, it is considered to be scattered 

and vulnerable 96 . 

• In Italy, the species is decreasing in the north of the country 86 . 

• In Morocco, the populations appear to be stable in the 

medium term, despite wide interannual fluctuations (linked with 

variable rainfall). 


IUCN 1997 




protection 


and Red Lists 



In Corsica, up until now, these kinds of threats appear to be 

absent. 

38 

- 

- 

- 

France, national list: 





133, 230 

- Morocco: “very rare” (RR) 

- Italy, national Red Data Book: “vul- 

86, 181 nerable” 

- Spain, regional lists (Medina, pers. 

com.) 

In Morocco and Spain, drainage is the most important threat, as 

well as overgrazing in Morocco. In northern Italy, increasing 

eutrophication* of the water is resulting in the decline of the 

species 86, 292 . 


In Corsica, the thorough drying out at the end of spring and in 

summer and the shallowness and nutrient-poor status of the 

loose substrate, as well as extensive cattle grazing at the six pools 

with L. uniflora, are sufficient to prevent the spread of more 

competitive species (such as Eleocharis palustris). 



Corsica: the Tre Padule site is protected by its legal “Réserve 

Naturelle” status. 

Morocco: no management measures. 


Corsica: continuation of extensive cattle grazing. 

Morocco: work towards limiting the numbers of livestock. 

Bibliography 

Anonymous, 1999 12 ; Bissardon & Guibal, 1997 39 ; Bournérias et al., 

2001 47 ; Casper & Krausch, 1981 63 ; Cirujano et al., 1992 79 ; Clément 

& Touffet, 1983 82 ; Conti et al., 1992 86 ; Danton & Baffray, 1995 96 ; 

Dierssen, 1983 117 ; Duvigneaud, 1971 127 ; Fennane & Ibn Tattou, 

1998 133 ; Gamisans, 1988 145 ; Grime, 1979 163 ; Jahandiez & Maire, 

1931-1934 181 ; Lorenzoni & Paradis, 2000 221 ; Lorenzoni, 1997 225 ; 

Maire, 1952-1987 230 ; Molinier, 1981 257 ; Pignatti, 1982 292 ; Robe & 

Griffiths, 1998 321 . 


Collaborators: Grillas P., H. Michaud, L. Rhazi & N. Yavercovski

Lythrum borysthenicum (Schrank) Litv. (1) & 

Lythrum tribracteatum Salzm ex Sprengel (2) 

ANGIOSPERMS 

LYTHRACEAE 

Main synonyms 

(1) Peplis erecta Moris, P. hispidula Durieu, P. nummulariaefolia 

Jord., L. biflorum (DC.) J. Gay. 

(2) L. Salzmannii Jord., L. dibracteatum Guss., L. hyssopifolium 

L. subsp. salzmanni (Jordan) Bonnier 

L. hyssopifolia L. var. pseudo-bibracteatum Tallon 

French names: (1) Péplis dressé; (2) Salicaire à trois bractées 

Spanish name: (2) Alheli silvestre 177 

Italian names: (1) Salcerella a foglie ovali; (2) Salcerella con due 

brattee 

English names: (1) Loosestrife (genus name); (2) Threebract 

Loosestrife 

Subspecies 

None 


(1) Lythrum borysthenicum 

• Leaves sessile or nearly so, ciliate when young, rounded oval, 

opposite, the upper leaves in a spiral. 

• Stem ciliate and fairly rough at the tip, erect, from 5 to 15 cm 

in height. 

• Flowers solitary, subsessile in the leaf axils. 

• Calyx expanded at the base, in a short tube, almost as wide as 

long, without prominent ribs and longer than the capsule, with 

five or six short teeth, blunt, equal or almost equal. 

•Petals crimson, five or six in number, or absent. 

• Stamens six, enclosed in each flower. 

(2) Lythrum tribracteatum 

• Green plant, glabrous, low (5 to 30 cm) more or less prostrate. 

• Leaves elliptical, small, entire, linear to oblong (0.3-1.5 cm long 

by 0.1 - 0.3 cm wide), sessile, alternate. 

• Flowers small, solitary in the axils of the branch leaves, subsessile, 

crimson; five or six petals, equalling approximately half 

the length of the calyx tube. 

• Calyx narrowed at the base, very elongated into a tube, much 

longer than wide (0.5 - 0.6 cm in length), with prominent ribs 

and 8 to 12 very short, triangular, unequal, blunt teeth, and two 

linear bracts, green, equal in length to the calyx and resembling 

leaves, or alternatively very short and scarious. 

• Stamens five to six. Style enclosed. 

• Fruit: cylindrical capsule* approximately equal to the calyx. 

Seeds yellow, slightly constricted. 


(1) Lythrum borysthenicum may be confused with L. (Peplis) portula, 

but this species has its leaves clearly petiolate and spatulate, 

a prostrate stem, petals liliaceous, and calyx shorter than 

the capsule. 

(2) L. tribracteatum may be confused with L. thymifolia and 

L. hyssopifolia, but in both these species, the colouration is glaucous, 

the habit is upright, the external calyx teeth are longer 

than the internal, the calyx is tubular and the stem unbranched 

or very little branched. 

calyx 

calyx 

1.5 mm 

1.25 mm 

Lythrum borysthenicum 

Lythrum tribacteatum 

Plant species 

1 cm 

1 cm 

39




(1) Lythrum borysthenicum 

Sub-Mediterranean species. 

France 

Provence-Alpes-Côte-d’Azur (Plaine des Maures, Colle du Rouet, 

Bois de Palayson, Hyères islands, Plateau d’Evenos), Languedoc- 

Roussillon (Roque-Haute, Béziers Quaternary plain, Plateau des 

Fenouillèdes , alluvial terraces of the Agly), and Corsica. 

Extension beyond the Mediterranean zone into west-central France 

as far as the Loire valley and the Sarthe. Previously recorded in 

the Dombes and surrounding area, but the data are however, 

highly dubious 266 (R. Dupré, CBNBP, pers. com.). 


Portugal: absent from the north of the country. 

Spain: western half, to Gerona and Valencia; absent from a large 

area of Galicia and Asturias 64 . 

Italy: Tuscany, Lazio, Sardinia, Sicily 292 . 

Morocco: Atlantic plains and mountains of low to medium altitude, 

in a sub-humid, semi-arid Mediterranean bioclimate*. 

Also present in Greece, Cyprus, Turkey, Algeria, Tunisia, Libya, Israel. 

(2) Lythrum tribracteatum 

Mediterranean species. 

France 

Atlantic region: départements of Loire-Atlantique, Vendée, 

Charente-Maritime, Aveyron. 

Mediterranean region: 

• Languedoc-Roussillon region: lower floodplain of the Aude, 

plateaux of Vendres, Roque-Haute and Caux-Fontès-Pézenas, Agde, 

Littoral ponds of Languedoc, Garrigue pools of the Montpellierais, 

Etang de Capelle and Costière nîmoise, Petite- 

Camargue. 

• Provence-Alpes-Côte-d’Azur Region: Lanau pool in the Crau, 

pools of Cerisières in the Camargue, Plateau des Quatre-Termes, 

Plateau du Cengle and Centre Var. 


Portugal, Spain, mainland Italy (in all provinces except Marche, 

Abruzzo and Basilicata), Sardinia, Sicily, Albania, Greece, Morocco, 

Algeria, Tunisia, Libya, Turkey, Syria, Lebanon and Egypt. 

Habitat 


(1) L. borysthenicum: temporary pools on a non-calcareous substrate 

(sandstone, granite, quartzite, schist, basalt, etc.), flooded 

in winter and very dry in summer, with oligotrophic water, but 

also edges of marshes and pools, or slow-flowing shallow rivulets 

(in Les Maures). 

(2) L. tribracteatum: temporary pools in plains, low and medium 

altitude mountains in sub-humid and humid Mediterranean bioclimates. 

In western France, L. tribracteatum also occurs in brackish 

grasslands near the coast in depressions with bare wet soil, and 

places that are trampled by livestock 96 . 

40 




22.34 Southern amphibious communities (Isoetalia) 

22.341 Short Mediterranean amphibious swards (Isoetion) 

(France): (1) and (2) 

22.343 Halo-nitrophilous Mediterranean amphibious swards 

(Heleochloion) (Spain): (2) 

22.32 Mediterranean annual amphibious swards (Nanocyperetalia) 

(France and Spain): (2) 


Morocco 

Both species belong to the Isoetion (velatae). Juncus pygmaeus, 

J. bufonius, J. capitatus, Lythrum thymifolia, L. hyssopifolia, 

Isoetes velata, etc. are associated with them. 

France continentale 

(1) In the Var (Plaine des Maures, Plaine de Palayson), L. borysthenicum 

is characteristic, along with Ranunculus revelieri, of an 

Isoetion association which is endemic* to this region. In Languedoc- 

Roussillon, it occurs in Isoetes setacea formations (Isoetetum 

setacei Br.-Bl.) 

(2) L. tribracteatum is often associated with Damasonium 

polyspermum as a characteristic species of the Lythrion tribracteati 

alliance (order Nanocyperetalia flavescentis) 153, 215 . It also 

occurs in rice fields in the Camargue, and fairly often together 

with Crypsis on the edges of ponds in Languedoc. It is also characteristic 

of the Elatinetum macropodae Br.-Bl. (1931), formerly 

described from Agde. 

On the central Atlantic coast, it is characteristic of a Junco 

hybridi-Lythretum tribracteati which occurs in sub-halophilic 362 

marshes. 

Corsica 

L. borysthenicum is associated with Isoetes velata and Juncus 

pygmaeus in the Tre Padule de Suartone pools 221 , with Elatine 

brochonii and Juncus pygmaeus in the Padulellu pool 219 , with 

L. hyssopifolia and Cotula coronopifolia in the coastal temporary 

pool at Tour d’Olmeto Point 282 . 

Spain 319 

L. tribracteatum is characteristic of the Lythrion tribracteati 

alliance, as well as the Lythro-Heleochloetum schoenoidis association 

Rivas Martinez 1966, within the Heleochloion alliance. 

Ecology 


Life form 

Therophyte (annual species); scapigerous*. 

Reproduction 

Aerial. 


In Morocco and Corsica, Lythrum borysthenicum and L. tribracteatum 

begin to germinate in February if the site is flooded; flowering

lasts from the end of April to July for L. borysthenicum and from the 

beginning of May to July for L. tribracteatum. Fruiting begins at the 

end of May for L. borysthenicum and mid-June for L. tribracteatum, 

accompanied by seed dispersal a short distance from the parent plant. 

In mainland France, flowering is slightly later, taking place in June- 

July for L. borysthenicum and in May-June for L. tribracteatum. 

In Corsica, L. borysthenicum has the following cycle: germination 

in February and March, growth in March and April, flowering and 

fruiting in May and the beginning of June, death of the plant 

when the habitat has dried out, in June. 

Lythrum borysthenicum 

Mainland France 


Morocco 


Flowering 

Fruiting and seed dispersal 



Lythrum tribracteatum 


Flowering 



Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec.. 

Morocco 


Flowering 





Flowering 


Hydrology 

These two amphibious pool species are often situated in more 

hydromorphic (low-lying) positions than L. thymifolium and 

L. thesioides. The germination of the seeds requires the soil to be 

saturated with water (flooding is not essential). At the vegetative 

stage the plant can tolerate shallow water (


in the Hautes-Alpes is vulnerable. In the Aveyron, this Lythrum 

has recently been observed as a adventive plant in the valley of 

the Tarn, but the other stations in this département have not 

recently been seen. 

• In the Atlantic area, the populations of L. tribracteatum, formerly 

fairly numerous, appear currently to be reduced to a small 

number of stations* 274, 362 . 


L. borysthenicum may be increasing due to its ability to colonise 

artificial new habitats (Medina, pers. com.). 


IUCN 1997 




protection 


and Red Lists 



The discontinuation of traditional stock rearing, conversion to 

arable land, urbanisation, infilling with debris, the use of pools 

that are dry or drying out (e.g. as car parks), and drainage, are the 

causes of the declines and the vulnerability of the populations of 

both these species. 


In Morocco, at sites with a semi-arid bioclimate, a high frequency 

of dry years, which prevent the renewal of the seed 

stocks, is a potential threat for both these taxa. In France, the 

main threat for both species comes mainly from the closing-up 

of the habitat due to the spread of woody plants. 


The fairly small sizes of the currently existing populations render 

them vulnerable. 



Morocco 

There are no management measures. 

42 

- 

- 

- 

France: 

(1) regional list for Languedoc- 

Roussillon: Decree of 29/10/1997/ 


(2) national list: Decree of 


13/05/1982, amended by Decree of 

31/081995/Journal Officiel 

17/10/1995 

- (2) France, national Red Data 

Book: priority species274 France 

Lythrum tribracteatum benefits from the legally protected status 

of some sites (Nature Reserves of Roque-Haute in the Hérault 

and Saint-Denis-du-Payré in western France), and L. borysthenicum 

from that of the Tre Padule de Suartone Nature Reserve 

(southern Corsica). 

L. borysthenicum also benefits at the Plaine des Maures (Var) from 

experimental management measures, in the context of the LIFE 

“Temporary Pools” Project. 

L. tribracteatum is the subject of annual monitoring at the Lanau 

pool (Bouches-du-Rhône), and its seeds are preserved ex-situ by 

the Conservatoire Botanique National Méditerranéen de Porquerolles. 

Finally, several sites (Roque-Haute, Les Maures plain and 

massif, Notre-Dame de l’Agenouillade at Agde, etc.) are included 

in Natura 2000* areas. 


• Promoting a more accurate census and a survey of the stations. 

• Avoiding any alterations to the hydrology of the pools 

(drainage and infilling). 

• Gaining control over urbanisation and site management. 

•Keeping the habitat open to a certain extent by grazing or by 

maintaining the plant cover via clearing. 

• Implementing legal protection measures for the species and 

the sites. 

• Raising the level of awareness among the managers involved. 

Bibliography 

Anonymous, 1999 12 ; Aubert & Loisel, 1971 18 ; Barbero, 1965 21 ; 

Castroviejo, 1986-2001 64 ; Chagneau, 2003 67 ; Danton & Baffray, 

1995 96 ; Fennane & Ibn Tattou, 1998 133 ; Fennane et al., 1999 134 ; 

Fournier, 1961 139 ; Gaudillat & Haury, 2002 153 ; Loisel, 1976 215 ; 

Lorenzoni & Paradis, 1997 219 , 2000 221 ; Lorenzoni et al., 1994 226 ; 

Maire, 1952-1987 230 ; Médail et al. 1998 246 ; Molina, 1998 251 ; Nétien, 

1993 266 ; Olivier et al., 1995 274 ; Paradis et al., 1999 282 , 2002 284 ; 

Pignatti, 1982 292 ; Quézel & Santa, 1962-1963 303 ; Rhazi et al., 

2001 314 ; Rivas-Goday, 1970 318 ; Terrisse, 1996 362 ; Valdés et al., 

2002 391 ; Walter & Gilett, 1998 399 . 

Authors: Michaud H. & L. Rhazi 

Collaborators: Paradis G. & M. L. Pozzo di Borgo

Lythrum thymifolium L. (1) & 

Lythrum thesioides M. Bieb. (2) 

ANGIOSPERMS 

LYTHRACEAE 

Main synonyms 

(1) L. hyssopifolia L. subsp. thymifolium (L.) Bonnier & Layens 

L. thymifolia L. 

(2) L. geminiflorum Bertol. 

L. thesioides M. Bieb. subsp. geminiflorum (Bertol.) Rouy & E.G. 

Camus 

L. purpurascens Châtenier 

French names: (1) Lythrum à feuilles de thym, Salicaire à feuilles 

de thym; (2) Lythrum faux-Thésium, Salicaire faux thésion 

Spanish names: (1) Salicaria-menor (catalan) 

Portuguese names: (1) Salicaria-menor 

Italian names: (1) Salcerella con foglie di timo; (2) Salcerella a 

fiori appaiati 

English names: (1) Thymeleaf Loosestrife; (2) Loosestrife (name 

of genus) 

Subspecies 

None 

96, 139, 292, 386, 394 


(1) Lythrum thymifolium 

• Annual plant, glaucous, more or less scabrid*, from 3-10 cm in 

height, with stem glabrous, erect, spindly, very leafy and not very 

much branched. 

• Leaves linear, sessileµ, alternate, small (0.5-0.9 cm long x 0.1- 

0.2 cm wide), narrow, close together, very finely toothed on the 

edges. 

• Flowers very small, solitary in the axils of the leaves on the 

branches, with short pedicels and with two linear bracts at the 

base of the calyx that are variable in size from one individual to 

another, sometimes within a single population 234, 386 . 

• Calyx elongated, tubular, short (0.20-0.25 cm long), bearing 

eight very unequal teeth at the tip: four long (0.5-1 mm) and 

papillose, alternating with four very short and membranous. 

• Corolla with four pink petals scarcely extending beyond the 

calyx teeth. 

•Two to four stamens per flower. 

• Fruit: capsule*, equal to or less than the length of the calyx 

tube. 

(2) Lythrum thesioides 

•Very similar, though a little taller (up to about forty centimetres). 

• Flowers very small, grouped in pairs or threes in the axils of all 

the leaves (solitary in L. thymifolium). 

• Calyx short, bell-shaped, with 8-12 teeth, and corolla rosepurple, 

with 4-6 petals with darker central veining (in L. thymifolium 

the calyx is cylindrical with eight teeth, and the corolla 

has four uniformly pink petals). 

calyx 

calyx 

1.5 mm 

1 mm 

Lythrum thymifolium 

Lythrum thesioides 

Plant species 

1 cm 

1 cm 


• Lythrum tribracteatum is usually more or less prostrate, with 

very branching stems, leaves often elliptical, flowers with a narrow 

tubular calyx with 10-12 not very obvious very short equal 

triangular teeth and a purple corolla with 5-6 petals. This is a 

rare species, more hygrophytic* than L. thymifolium, and is found 

in oligotrophic * to eutrophic* habitats. 

• L. hyssopifolium L. is a common species in all sorts of humid 

to flood-prone habitats. It is usually robust, not very glaucous, 

with lanceolate leaves tapering at the base, and flowers usually 

43


hexamerous* and pedicellate. The six petals clearly extend 

beyond the calyx, which has 12 unequal teeth at the tip (six 0.5- 

1 mm long, not very papillose, alternating with six short and 

membranous). The flower has six stamens. Ambiguous forms, 

intermediate between L. hyssopifolium and L. thymifolium, have 

(rarely) been found, and their determination is always difficult. 

• Frankenia pulverulenta is always a plant of saline habitats, on 

the coast or inland. It has a prostrate habit, a stem that is slightly 

woody at the base and very branching, petals purplish pink with 

serrated elongated limbs, and narrow spatulate leaves slightly 

curled at the edges. 



(1) L. thymifolium 


France 274 

Départements of Allier, Puy-de-Dôme, Haute-Loire, Ardèche, 

Drôme, Hautes-Alpes, Pyrénées-Orientales, Aude, Hérault, Gard, 

Bouches-du-Rhône, Var and Alpes-Maritimes. 

386, 394 


Portugal, Spain, Italy, Greece, Algeria and Morocco. 

(2) L. thesioides 

Sub-pontic species. 

France 

Lower Rhône valley: Pierrelatte (Drôme), Caderousse and Orange 

(Vaucluse), Tresques, La Capelle-et-Masmolène, Meynes, and 

Jonquières-Saint-Vincent (Gard). 


Two old records from Italy in the floodplain of the Pau. 

Outside the Mediterranean region 

Three old records from the Caucasus, lower valleys of the Volga 

and the Don. 

Habitat 


(1) Habitats suitable for L. thymifolium include pools that are 

temporarily flooded or simply saturated in winter, and very dry in 

summer, and also the edges of streams, marshes and ponds. The 

substrate is usually siliceous (sandstones, schists, basalts, etc.), 

the floodwater oligotrophic* and of variable depth between sites 

and years. 

(2) L. thesioides is also associated with habitats that are floodprone 

or very wet during the winter, such as the banks of natural 

freshwater ponds or the alluvial terraces of the Rhône or its 

tributaries. On the edges of ponds, L. thesioides occurs in the outer 

band of vegetation, leaving the centre of the marsh to the low, 

more hygrophytic formations of the Isoetion (formerly at Pazac, 

in the Gard) or the Heleochloion (Etang de la Capelle, and probably 

formerly at Jonquières, in the Gard). 

44 




(1) 22.34 Southern amphibious communities (Isoetalia) 

22.341 Short Mediterranean amphibious swards (Isoetion) 

(1) and (2): 22.32 Annual Mediterranean amphibious swards (Nanocyperetalia) 

Phytosociology* and companion species 

(1) Lythrum thymifolium 

It belongs to the class Isoeto-Nanojuncetea and to the Cicendion 

filiformis and Isoetion s.l. alliances 274 . 

• In Languedoc: Isoetetum duriaei Braun-Blanquet. 

• In the Provence-Alpes-Côte-d’Azur (PACA) Region: 

- Spirantho-Anagallidetum tenellae Aubert & Loisel. 

- Isoeto duriaei-Nasturtietum (=Sisymbrelletum) asperum 

Barbero. 

There are a large number of associated species: Juncus pygmaeus, 

J. capitatus (especially in Languedoc), J. bufonius, L. borysthenicum, 

L. hyssopifolia, Isoetes velata, I. duriei (ecologically 

more similar, in France, to L. thymifolium than to I. velata), Laurentia 

michellii. 

In the Crau (Lanau pool), the species is associated with species of 

the Lythrion tribracteati alliance. 

(2) Lythrum thesioides 

Very few facts are known about the habitat of this species, which 

has been observed as much in cultivated land in flood-prone areas 

as on the shores of temporary ponds. At Jonquières-Saint-Vincent 192 

as well as at La Capelle-et-Masmolène 249 , L. thesioides was abundantly 

accompanied by L. tribracteatum. At this station there also 

occurred Potentilla supina, formerly present at Jonquières. At Meyne, 

L. thesioides was accompanied by several Centaurium (C. pulchellum, 

C. spicatum and C. tenuiflorum), Blackstonia serotina, 

Deschampsia media, Brachypodium phoenicoides, Phleum pratense 

subsp. serotinum, L. hyssopifolium etc. 359 . The habitat of L. thesioides 

thus appears to be quite different from that of L. thymifolium 

and belongs to the Lythrion tribracteati alliance or even 

the Deschampsion mediae. 

Ecology 


Life form 

Therophytes (annual species). 

Reproduction 

Aerial. 


(1) Seeds glabrous, brownish, near ovoid in shape (length: 0.5 mm; 

width: 0.4 mm), very finely punctuate and light (weight


(1) In Morocco, germination at the beginning of February if the 

site is flooded, vegetative growth from February to the end of 

March, flowering in April-May, fruiting end of May and June, and 

seed dispersal a short distance from the parent plant. In France 

flowering is slightly later, taking place in May and June. 

(2) Cycle unknown. Based on examination of herbarium samples, 

flowering takes place mostly in August-October and rarely in 

June (one record). The phenology* of this species is therefore 

later than for L. thymifolium, and closer to L. tribracteatum for 

example. The stamens have anthers which extend to the same 

level as the stigma, which raises the a priori possibility of autogamy*. 

This mode of reproduction is all the more probable as the 

flowers often lack petals. The capsule, which barely extends outside 

the calyx, opens at the tip to form four very short valves. 

Seed dispersal is consequently probably limited. The seeds 

undoubtedly remain viable for a long time, as the plant may not 

appear every year and may disappear for many years between 

appearances at a given place. 

Lythrum thymifolium 

Morocco 


France 


Flowering 


Hydrology 

For seed germination it is only necessary for the soil to be saturated 

with water. Flooding is not essential. At the vegetative 

stage the plant can tolerate shallow water (< 5-7 cm). 

Substrate 

Sandy to sandy-silty, even silty for L. thesioides. Substrate above 

all non-calcareous (silica, basalt) for L. thymifolium. 


Deep shade from tall plants may be unfavourable for these heliophilous* 

species, especially at sites which are not grazed (France). 


Favourable in that the habitat is opened up. 



Fruiting 




Flowering 

? 

Fruiting 

? 



• In France, from west to east: Massif des Fenouillèdes and Plaine 

de la Têt in the Pyrénées-Orientales, plateaux of Roque-Haute, 

Plant species 

Caux-Fontès-Pézenas and Agde, Quaternary siliceous plains of 

Béziers and Montpellier in the Hérault, Costière Nîmoise in 

the Gard, Plateau du Coiron in the Ardèche, the Issoire area in the 

Puy-de-Dôme, Upaix in the Hautes-Alpes, the Lanau pool in 

the Bouches-du-Rhône, Plateau de l’Evenos , the Plaine and Massif 

des Maures, Colle du Rouet and Bois de Palayson, Estérel of 

Fréjus in the Var and the Biot Massif in the Alpes Maritimes 246, 274 . 

• In Spain and Portugal: on the Balearic Islands and distributed 

over a large part of the peninsula, with the exception of the 

Cordillera Cantabrica 64 . 

• In Italy: in the three regions of Puglia, Basilicata and Lazio, 

and in Sardinia 292 . 

• In Morocco: pools in the Atlantic plains (Benslimane) and pools 

in the Atlas Mountains. 


In France the plant has only been found 14 times at seven localities 

since it was discovered by A. Jordan in August 1841 at the 

Etang de Jonquières. The Etang de la Capelle is the only site at 

present (1998). 



Populations declining due to the transformation of their 

biotopes. This species is assumed to have disappeared from some 

former stations (Allier, Haute-Loire, Saint-André-d’Embrun, 

Aude, former Etang de Jonquières, etc.).. 


• In France, the following localities have been profoundly modified 

and the possibility of Lythrum surviving there these days 

appears very remote: the Etang de Jonquières, now drained and 

cultivated, where the species has not been seen since 1902 58 ; the 

Etang de Pazac, drained and converted to vineyards and pasture, 

where the species was only seen once, in 1951 360, 361 ; the areas 

around Orange (seen on one occasion in 1892), Caderousse 70 , 

Tresque (appeared en masse, but only once, in 1853) and 

Pierrelatte (1915, not seen since) 71, 148 . These days all these areas 

are used for agricultural activities that are not very propitious for 

the survival of annual Lythrum. The plant has not been seen for 

a long time elsewhere in Europe or in the former USSR. 

• In Italy, it is considered to be extinct, its two sites not having 

produced records since the 19th century 274 . 


IUCN 1997 




protection 


and Red Lists 

- 

(2): strictly protected (annexe I) 

- 

(1) and (2) France, national list: 





1) and (2) France, national Red 

Data Book: priority species274 133, 181, 381 

(1) Morocco: “rare” (R) 

(2) Italy, national Red Data Book: 

“extinct” 86 

45




Urban development, as well as drainage of pools and ponds, their 

filling in and their conversion to agriculture, constitute the greatest 

threats for both species. 


In Morocco, successions of dry years prevent the renewal of the 

L. thymifolium seedbank*, which could pose a problem for some 

populations in the long term. In France the closing-up of the 

habitat, resulting from the steady increase of woody plants, constitutes 

a threat for this species. 


•(1) In France, the L. thymifolium stations are vulnerable and 

subject to the risk of local extinction, especially as the species is 

at the northern limit of its range. 

• (2) The same is true of L. thesioides whose very insecure status 

in France is perhaps partly mitigated by the longevity of the 

seedbank in the soil. However, the very severe decline in the 

species throughout its entire range exposes the single currently 

confirmed population to particular risk. 



No direct management measures either in Morocco or in France. 

In France, only collection of the seeds of the two species (preserved 

ex-situ) has been carried out, by the Conservatoire Botanique 

National Méditerranéen de Porquerolles. 


• Avoidance of any alterations to the hydrology of the pools and 

ponds (drainage, infilling or, conversely, permanently flooding). 

46 

• Control over urbanisation. 

• Maintenance or promotion of a certain degree of opening of 

the habitat by grazing or by managing the vegetation cover 

(clearing). Scraping the surface of the soil appears to be beneficial 

for L. thesioides. 

• In France, gaining control over the ownership of plots of land 

at the Etang de la Capelle, in order to implement management 

and to remove the potential threats of destruction which are still 

faced by this last known site for L. thesioides. Looking for the 

species among the seed stocks at this site. Searching at the former 

localities where this species has been observed in the past, 

as well as in the few remaining wetlands in the lower Rhône valley, 

in an attempt to find new populations. 

Bibliography 

Anonymous, 1999 12 ; Aubert & Loisel, 1971 18 ; Cabanès, 1903 58 ; 

Castroviejo, 1986-2001 64 ; Charrel, 1913 70 ; Châtenier, 1922 71 ; 

Conti et al., 1992 86 ; Danton & Baffray, 1995 96 ; Fennane & Ibn 

Tattou, 1998 133 ; Fournier, 1961 139 ; Garraud, 2003 148 ; Jahandiez 

& Maire, 1931-1934 181 ; Jordan, 1847 192 ; Lesouëf, 1997 210 ; 

Mandin & Hugonnot, 2001 234 ; Medail et al., 1998 246 ; Michaud & 

Molina, 1999 249 ; Olivier et al., 1995 274 ; Pignatti, 1982292 ; Poirion 

& Barbero, 1966 296 ; Rhazi et al., 2001 314 ; Rouy & Camus, 1901 327 ; 

Tallon, 1923-1969 359 , 1953 360 , 1967 361 ; Titolet & Rhazi, 1999 381 ; 

Tutin et al., 1964-1980 385 ; Velayos, 1997 394 ; Webb, 1968 400 . 

Authors: Michaud H. & L. Rhazi 

Collaborators: Antonetti P., L. Garraud, J. Molina & G. Paradis

Marsilea strigosa Willd. 

POLYPODIOPHYTES 300 

MARSILEACEAE 

Main synonyms 

M. pubescens Ten. 

M. fabri Dunal 

French names: Fougère d’eau pubescente, Marsilée pubescente. 

Spanish names: Trébol de agua, Trébol de cuatro hojas, 

Marsilia180 32, 265 

Moroccan names: Ouarda el maa, Rjel-el-Ketta 

Algerian names: Arbas ourrak, Qoub303 English name: Clover Fern (a part of the genus) 

Subspecies 

None 

Description/identification criteria 2 

• Heterosporous* fern (with macrospores and microspores), 

aquatic, with long stolons and thick downy rhizomes with short, 

close internodes. 

• Fronds characteristic, formed from 4 leaflets arranged in a 

cross at the apex of the petiole (rachis), like a Four Leaves Clover 

leaf. 

•Two types of fronds: 

- The first with a floating blade, glabrous, borne on a long petiole, 

formed during the aquatic phase. 

- The second with a rigid, upright petiole and downy blade, 

appearing during the dry phase. 

• Fruiting: sporocarps* villous, axillary, subsessile, arranged in 

two rows along the rhizome. 


Water Shamrock (Marsilea quadrifolia L.; syn.: M. quadrifoliata 

(L.) L.) has always glabrous leaflets and sporocarps with short 

pedicels, solitary or clustered in twos or threes. In France, 

M. quadrifolia grows in the temperate region 2 , whereas M. 

strigosa is a strictly Mediterranean species, which reduces the 

risk of confusion. 

In Spain, M. strigosa can be confused with M. quadrifolia as well 

as with an endemic species of the Iberian Peninsula, 

M. batardae (Medina, pers. com.). The latter species has the rhizomes 

much longer and branched, and widely spaced sporocarps, 

distinctly pedicellate, each pedicel bearing a conical tooth 300 . 

In Morocco, M. minuta (= M. diffusa) has a slender rhizome and 

distinctly pedicellate sporocarps, grouped in twos or threes 


160, 386 


Marsilea strigosa is a western Mediterranean species; it also 

occurs in some localities to the north of the Caspian Sea. It is 

present in France, Italy, Sardinia, Spain, the Balearics, Algeria, 

Morocco, Egypt, Romania and Ukraine (Crimea). Its existence in 

Portugal has been disproved. 

Plant species 

France 

It is only found today in three localities: 

• Hérault: pools in the Roque-Haute plateau and a wet depression 

in the wine-growing plain of Vendres. 

• Pyrénées-Orientales: alluvial terraces of the Têt, in the uncultivated 

land of Torremila. 


Italy: south of the peninsula (several localities in the province of 

Puglia, region of Taranto) and Sardinia 86, 292 . 

Spain: between 0 and 400 m, in 14 provinces (Medina, pers. com.). 

Above all found in the regions of Catalonia, Levante, Almeria, 

Extramadura, Castilla y Leon and on the islands of Majorca and 

Minorca 64 . 

Algeria 303 : temporary pools of the Algerois and the Oranais 

(coastal sahels, coastal plains, Tellian Atlas). 

Morocco 134 : dayas* of the Atlas ranges, up to 900 m in altitude 

(Middle Atlas, High Atlas and Anti-Atlas) and the Atlantic plains 

(Benslimane, Sidi Bettache). 

Egypt: Nile Valley and Delta 46 . 

Habitat 

Marsilea strigosa 

1 cm 


Marsilea strigosa appears to occupy a relatively wide range of 

natural habitats and substrates, and to be part of several phytosociological 

units, some of which remain to be specified 2 . 

47


The species grows mainly in temporary pools with nutrient-poor 

and poorly mineralised water (Medina, pers. com.). It is found 

within amphibious vegetation which are submerged for a fairly 

long time (in France at Roque-Haute, or in Spain), dominated by 

Isoetes setacea or Mentha cervina, or in Crassula vaillantii- 

Lythrum borysthenicum formations 2, 251, 319 . 

In Morocco, in the Atlantic plains, it grows in Isoetes velata, 

Ranunculus peltatus and Myriophyllum alterniflorum formations 

312 . 

It also occurs in regularly flooded former vineyards, cleared of 

vines on siliceous gravel* (Vendres plateau and Plaine du Roussillon, 

France), often in furrows in which the water collects. 







France 

Marsilea strigosa is considered a characteristic species of the 

Isoetetum setaceae association* Br.-Bl. (1931) 1935 215 . On abandoned 

farmland, the formations have yet to be incorporated into 

phytosociological units 2 . These are most likely Isoetion formations 

heavily disturbed by viticulture. 

Spain 

The species is characteristic of the Sisymbrello-Preslietum association 

(Preslion cervinae alliance 319 ). 


France 

• In temporary pools (Roque-Haute and Torremila): Isoetes 

setacea, Pilularia minuta, Mentha cervina, Lythrum borysthenicum, 

L. thymifolium, Eleocharis palustris, Myosurus breviscapus, 

Juncus pygmaeus, Pulicaria vulgaris and Polygonum 

romanum subsp. gallicum 49 (Grillas & Tan Ham, pers. com. CEN-LR, 

pers. com. ). 

• On abandoned farmland, an impoverished form of this community 

can be found (disappearance of Isoetes, Mentha cervina, 

Pilularia minuta and Eleocharis palustris) augmented by ruderal 

species (Conyza spp., Dittrichia viscosa, etc.) 2, 246 . 

Morocco 

Isoetes velata, Ranunculus peltatus Lythrum borysthenicum, 

Myriophyllum alterniflorum, Eleocharis palustris and Baldellia 

ranunculoides 312 . 

Ecology 


Life form 

Species considered to be perennial* (radicant hydrophyte*, producing 

stolons, traditionally considered as a geophyte* with a rhizome), 

but which appears to behave as an annual (therophyte*) 395 . 

48 

Sporocarps 

Suborbicular, tightly packed, sessile* or with very short pedicels 

(2 to 2.5 mm) covered in articulated hairs 300 , with a diameter of 

3 to 5 mm. These are highly efficient survival organs due to their 

capacity to survive several decades, or even over a century of 

drought. In experimental conditions, a sporocarp aged 103 years, 

taken from a herbarium, was able to produce 15 embryos 83 . 

Size of microsporangia: 50-62 microns 180 . 

Size of macrosporangia: 450-500 microns 180 . 


The sporocarps*, often saturated with water from the autumn, 

open in late winter and early spring, releasing the microsporangia 

and macrosporangia. Fertilisation occurs over the following 

days. The first, filiform, fronds appear within a few days, followed 

by fronds with two leaflets, then with four leaflets. They appear 

under a thin layer of water, during the dry phase of the pool, in 

winter in Morocco (January-February) and in spring in France (in 

March-April at Roque-Haute). 

When the soil is waterlogged, vegetative growth is prolific. The 

rhizome continues to grow, emitting a large number of fronds 

(dense tufts) as well as stolons which take root, enabling new 

rhizomes to develop. The formation of sporocarps and spores has 

been observed in March-April in Italy 86 and from April to June in 

France. 

When the pool dries out completely, the plant acquires its terrestrial 

habit. The fronds wither and disappear towards the end 

of the summer in Morocco (Rhazi L., pers. com.) and up until the 

autumn in France. The plant then only survives in the form of 

sporocarps arranged in two tight rows on either side of the driedout 

rhizome. 

France 


Emergence of fronds and vegetative growth 

? 

Formation of sporocarps 



Hydrology 

The habitat of Marsilea strigosa is characterised by a fairly long 

inundation period, between five and eight months: from December 

to May-July in France (Rhazi M., pers. com.); from December 

to March in North Africa 312 , and by a maximum water depth 

of around 40 to 50 cm. In experimental conditions (not published), 

the opening of the sporocarps and fertilisation are observed 

under a thin layer of water, or even on saturated ground. 

Substrate 

Usually non-calcareous (basaltic plateau at Roque-Haute, siliceous 

gravels at Vendres and at Saint-Estève). 


This species is sensitive to competition by perennial grasses and 

to the shade of woody plants (heliophilous* species).


The rhizomes of Marsilea do not appear to be sought by Wild 

Boar, though they can be dug up by chance by these animals. 

They are resistant to the turning-over of the top layer of soil in 

vineyards. 



France 

Very rare species with only three sites known. At Roque-Haute, 

over an ensemble of 200 pools, Marsilea appears to be disseminate 

in small populations in 10 to 26 pools depending on interannual 

variations in rainfall 162, 246 . On Torremila, the species is 

present in small scattered populations in three humid areas of 

uncultivated land and one temporary pool. 

Italy 

Considered as very rare (a few localities). 

Spain 

Fifty or so localities in the regions of Catalonia, Levante, Almeria 

and Extramadura, including five which have now disappeared; 

very rare species in the Balearics 64 (a few localities in southern 

Majorca and northern Minorca (Médina, pers. com.). 

Morocco 

Many localities. The species, more common than M. minuta, is 

not considered to be rare or threatened 133 . 

Algeria 

Rare species 303 . 


France 

The populations have decreased: 

• The species has disappeared from the Clape d’Agde pool, 

where it was found in 1831, and from Aigues-Mortes where it 

was observed at the end of the 19 th century 2 . 

• The station at Vendres is very seriously threatened; the plants 

have not been observed since 1998 but the sporocarps are still 

present in the soil. 

• The populations of the Saint-Estève pool and the uncultivated 

lands of Torremila, discovered in 1968, have been destroyed. The 

species has disappeared since 2000 from two other sites of 

uncultivated land discovered in 1996; however, two new stations 

have since been found on this site. 

Morocco 

The species is considered as stable (Rhazi L, pers. com.). 

Spain 

The species is in decline (province of Valencia in particular); 10% 

of stations are expected to disappear over the next 20 years 

(Medina, pers. com.). 


IUCN 1997 




protection 


and Red Lists 



Plant species 

France 

These are the main threats. The habitat of the Vendres and 

Torremila stations has been degraded and partially destroyed by 

drainage and cultivation. It has returned to a wild state, where 

the species survives, though it risks being put under intensive 

cultivation again in the short term 66 . Only one intact pool survives 

among the vineyards of Torremila. At Roque-Haute, despite 

the “Réserve Naturelle (Nature Reserve)” status, the absence of 

control over land ownership and usage no longer permits management 

or monitoring of the site, and seriously compromises 

the conservation of the Marsilea populations. 

Spain 

The main threats are the drying-out of the pools (by drainage), 

overgrazing and climate change 179 (Medina, pers. com.). 


- 

Strictly protected species (Annexe I) 

Annexes II and IV 

- France, national list: 

Decree of 20/01/1982/ Journal Officiel 

13/05/1982, amended by Decree of 

31/08/1995/ Journal Officiel 

17/10/1995 


priority species. 



- Spain, regional lists (Medina, pers. com.) 

- Balearics: “vulnerable” 331 

Vegetation dynamics 

At Roque-Haute, following the abandonment of grazing several 

decades ago, the spread of woody plants and invasion by Scirpus 

are threatening the habitat of Isoetes setacea 315 , which is also 

that of Marsilea 251 . At Torremila, the species is subject to competition 

from perennial grasses (mainly Deschampsia media), 

encouraged by the natural dynamics of the uncultivated land. 

Variability of hydrological conditions 

The interannual fluctuations in flooding cause fluctuations in the 

abundance of Marsilea without jeopardising the survival of the 

populations. 


Its very reduced capacity for dispersal results in a poor aptitude 

for the colonisation of new sites, and significant isolation of populations, 

both geographically and genetically 395 . 

49




France 

The Roque-Haute site is protected by its legal “Réserve Naturelle” 

status. Marsilea was the subject of regular monitoring here from 

1992 to 2001. 

At Vendres, the site is included in a Natura 2000* area, and the 

Conservatoire des Espaces Naturels du Languedoc-Roussillon (CEN- 

LR) has acquired land to favour the restoration of the hydrological 

regime. 

At Torremila, the Conservatoire has drawn up the Objectives Document 

(DOCOB) of the Natura 2000 site (Natura 2000 FR 9102001 

“Torremila Humid Uncultivated Land”). 

The two sites of Vendres and Torremila have been selected for the 

setting-up of a LIFE project for the conservation of Alpine and 

Mediterranean flora of EEC interest at the European scale 

(France, Spain, Italy). 

Conservation ex-situ: sporocarps have been collected by the 

Conservatoire Botanique National Méditerranéen de Porquerolles. 

Cultivation trials successfully carried out in January 2004, particularly 

within the framework of a project for the restoration of 

the Vendres site, show the viability of harvested sporocarps. 

Spain 

Only a few localities are included in Natura 2000 areas. Some are 

the subject of particular programmes (Province of Valencia, 

Balearics) (Medina, pers. com) such as the creation of two microreserves 

in the lagoons of Sinarcas (Province of Valencia) and the 

production of sporocarps ex-situ from fragments of rhizomes 

with a view to the reintroduction of the species 179 . The species is 

classified as “threatened” on the regional lists of the regions 

where it is present (Medina, pers. com.). 

Italy 

The species is cultivated in the botanical gardens at Padua 86 . 

Recommendations (for France) 

Viticulture 

Ban intensive viticulture on all sites, as it is totally unfavourable 

to the survival of Marsilea populations through the impact of 

various agricultural practices (drainage, burying of sporocarps, 

herbicides). On the other hand, outside of the temporary-pool 

habitat, extensive viticulture (no herbicides, light cultivation of 

50 

the soil surface) is not incompatible with the conservation of 

Marsilea populations, provided it permits flooding from autumn 

to spring. 

Competition from perennials and woody plants (closing-up of the 

habitat) 

• In the Roque-Haute Nature Reserve: implement the management 

plan and more generally restore a concerted management 

with a natural-heritage objective: reintroduce grazing, preferably 

by sheep and goats, together with an appropriate monitoring 

programme. 

• At Torremila: promote management of perennial grasses by 

water-weed cutting and grazing. 

Isolation of populations 395 

Genetic isolation has implications for conservation strategies: to 

properly protect this species, and preserve its adaptive potential, 

it is essential to protect a large number of its populations. 

Absence of control over land ownership and uses 

Restore control over land ownership and/or usage, an important 

prerequisite for the implementation of conservation management 

of the populations, particularly at Roque-Haute. 

Bibliography 

Aboucaya et al., 2002 2 ; Aizpuru et al., 2000 6 ; Amigo, 1987 10 ; 

Anonymous, 1999 12 ; Baudière & Cauwet, 1968 23 ; Bertrand, 

1991 32 ; Bissardon & Guibal, 1997 39 ; Boulos, 1995 46 ; Braun- 

Blanquet et al., 1952 49 ; Castroviejo, 1986-2001 64 ; CEN-LR, 

2001 66 ; Colas et al., 1996 83 ; Conti et al., 1992 86 ; Bern 

Convention,1979 87 ; Danton & Baffray, 1995 96 ; Directive 

92/43/CEE, 1992 119 ; Fennane & Ibn Tattou, 1998 133 ; Fennane et 

al., 1999 134 ; Foucault de, 1988 98 ; Gaudillat & Haury, 2002 153 ; 

Greuter et al, 1984-1989 160 ; Grillas & Tan Ham, 1998 162 ; Ibars 

& Estrelles, 1997 179 ; Ibars et al., 1999 180 ; Lewin, 2000 213 ; Loisel, 

1976 215 ; Médail et al.,1998 246 ; Molina, 1998 251 ; Nègre, 1956 264 , 

1961-1962 265 ; Olivier et al., 1995 274 ; Pignatti, 1982 292 ; Prelli, 

2001 300 ; Quézel & Santa, 1963 303 ; Quézel, 1998 306 ; Rhazi, 

2001 312 ; Rhazi, 2004 315 ; Rivas Goday, 1970 319 ; Saez & Rossello, 

2001 331 ; Tutin et al., 1964-1980 386 ; Vitalis et al. 2002 395 ; Walter 

& Gilett, 1998 399 . 

Author: Yavercovski N. 

Collaborators: Garnéro S., C Houssard., M. Klesczewski, 

H. Michaud & F. Ruchon

Mentha cervina L. 

ANGIOSPERMS 

LAMIACEAE 

Main synonyms 

Preslia cervina (L.) Fresen. 

French name: Menthe des cerfs 

Spanish names: Menta de burro, Poleo de ciervo 

Moroccan name: Fliyou 

Subspecies 

None 

274, 292, 386 


•Perennial plant with rhizome very close to the surface42 , 

recognisable by its very musky mentholated scent, its upright 

habit, its very narrow (1 to 2.5 cm long and 1 to 4 mm wide) 

opposite leaves (in the axils of which grow fascicle of smaller 

leaves borne on very short branches), and above all, by the characteristic 

palmate shape of the bracteoles* surrounding the 

whorls of flowers, which evoke the antlers of a deer, and to which 

the species owes its name. 

• Upright stems, hollow and angled at the base (10 to 60 cm), 

square-shaped in section and slightly branched. 

• Glabrous leaves, sessile* but attenuate* at the base, entire or 

roughly toothed. 

• Flowers hermaphrodite, forming compact whorls, widely 

spaced on the upper half of the stem. Calyx tubular and villous 

at the throat, with four triangular teeth terminating in a fine 

point. Corolla twice as long as the calyx, with two equal lips, pink 

or white. Floral bracts identical to the leaves, but larger. 


During and after flowering, no confusion is possible: it is the only 

mint which has deeply lobed bracteoles and whose calyx has four 

teeth and not five, hence its former inclusion in the genus Preslia. 

In the vegetative state, the narrow, sessile leaves are characteristic 

and differentiate it from the Pennyroyal (Mentha pulegium), 

frequently present in the same habitats but whose leaves are 

rounded with short petioles. 

Before flowering, the young shoots do not yet have their scent of 

mint, and can be confused with young Hedge Hyssop (Gratiola 

officinalis) plants. 



Western Mediterranean species, found in six countries 160, 386 . 

France 

Present mainly in Languedoc-Roussillon 246 . In the PACA region 

(Vaucluse and Bouches-du-Rhône), its presence has not been 

confirmed recently 306 , but it was found in 2003 at the Etang des 

Aulnes (Willm, pers. com). 

Plant species 


Spain and Portugal: western half of the Iberian Peninsula, with 

some localities in the eastern part AND in the Balearics (Medina, 

pers. com.). 

Italy: once found in the wetlands and temporary pools of 

Abruzzo, but not seen recently 292 . 

Algeria: temporary pools of the high plateaux of the Algerois and 

the Oranais, Geryville region 303 . 

Morocco: dayas* and edges of peat bogs of the Rif regions (Rhazi 

L., pers. com.). 

Habitat 

Mentha cervina 


Species of the thermo and meso-Mediterranean zone. 

1 cm 

- In France, this mint occupies the long-flooded parts of temporary 

pools and lavognes*, and a number of temporary water 

courses of the Montpellierais garrigues (Molina, pers. com.). It 

once occurred in the ox-bows of the Rhône (breaches and 

launes* around Tarascon, Avignon, etc.), where hydraulic modifications 

caused its disappearance. It forms, with the often perennial 

species associated with it, areas of tall-herb grassland of 

several dm 2 to several m 2 . 

- In North Africa (Morocco, Algeria), the species grows at an altitude 

where the rainfall is sufficient (above 1,000 m in the 

51


Moroccan Rif). In the temporary pools of the plain, it gives way 

to the Pennyroyal, which is less water-demanding (Rhazi L, pers. 

com.). 





22.342 Tall Mediterranean amphibious swards (Preslion cervinae): 

vegetation of tall annuals of land covered by deep 

water for long periods, with Mentha cervina. 




Alliance: Preslion cervinae 

Associations*: 

•In France 49, 215 : 

- Preslietum cervinae: Mentha cervina-Eleocharis palustris association, 

described from western Languedoc. 

- Preslio-Trigonelletum ornithopodioides: association described 

from the Costière Nîmoise (Gard), from where it has disappeared. 

A related formation is perhaps present in the Crau 98 . 

• In Spain 319 : 

- Preslietum cervinae 

- Sisymbrello-Preslietum: Mentha cervina is accompanied by 

Sisymbrella aspera, Veronica anagalloides, Cyperus Badius, Marsilea 

strigosa, Eryngium galloides, Callitriche platycarpa, etc. 

- Preslio-Eryngietum corniculati: M. cervina is present here with 

Eryngium corniculatus, I. velata, I. setacea, etc. 


France 

At Roque-Haute, the mint grows with Marsilea strigosa and 

Isoetes setacea, or with Eleocharis palustris in the deepest parts 

of the pools 251 . 

In the Montpellierais and Uzégeois garrigues the companion 

species are Eleocharis palustris, Sisymbrella aspera, Juncus articulatus 

and Mentha pulegium. 

Morocco 

Eleocharis palustris, Callitriche truncata and Alisma plantagoaquatica 

are present at its stations. 

Ecology 


Life form 

Perennial species with vegetative growth (suckers, rhizomes). 

Although considered by several authors as hemicryptophyte*, in 

France this species is more of a rhizomatous geophyte*, as it is 

not visible before the beginning of the spring, and the young 

shoots appear towards the month of April. 


In France, its vegetative growth occurs in the spring, and its 

reproductive cycle in summer. At Roque-Haute, it flowers from 

late June to mid-August, fruits from late July to September and 

withers in October. 

52 

In Morocco, its development is earlier: the shoots appear in 

March, flowering commences in May and fruiting in early summer. 

France 


Morocco 

Emergence of shoots and vegetative growth 

Flowering 

Fruiting 




Flowering 

Fruiting 

Hydrology 

In France, the habitat of this species is characterised by a long 

period of flooding, from the autumn to spring 153 (six to eight 

months depending on the year), a depth of water of between 20 and 

70 cm (or 100 cm), with an optimum of around 40 cm (Rhazi M., 

pers. com.). Germination takes place under a thin layer of water 

(non-turbid) or waterlogged ground (Morocco). 

Substrate 

Various substrates, oligotrophic* 274 or meso-eutrophic* 98 , on 

basaltic (Roque-Haute), calcareous (garrigues), marl (Minervois), 

or granitic (Fenouillèdes) bedrock, or on alluvium (borders of 

ponds). 


This heliophilous* species is sensitive to shade. The water depth 

limits invasion by woody species, but on the other hand creates 

conditions favourable to colonisation by fiercely competitive 

helophytes* (reeds, Scirpus, Reedmace, sedges) 153 . 



• In France, two major sites in terms of the number of populations: 

the Montpellierais garrigues (Hérault), and the Uzégeois 

and Bagnolais garrigues (Gard). Then two other less important 

sites: Fenouillèdes (Pyrénées-Orientales) and Roque-Haute 

(Hérault). Finally, ten or so secondary sites in Languedoc and the 

Crau, where the plant forms populations which appear to be isolated, 

and sometimes very small (a few plants in the Etang de 

Pujaut, Gard). 

• In Italy, the species is considered to have disappeared 292 . 

• In Algeria, a very rare species 303 . 

• In Morocco, fewer than five localities 133 . Other stations could 

be discovered by searching in the mountainous regions of the Rif. 

• In Spain and Portugal, the species is not considered rare.


France 

The populations of this species seem to be undergoing significant 

declines over the whole of its range, except in the Hérault and 

the Gard. 

• Gard: the species remains abundant in the Uzégeois-Bagnolais 

(plateaux of Méjannes-le-Clap, Lussan, etc.), and is surviving in the 

region of Quissac (pools at Vibrac). It is still present in the marshes 

of Pujaut 157, 246 and further north in a lavogne* in the eastern end 

of Larzac, but it has disappeared from the Costière Nîmoise 

(Redessan marsh, Etang de Pazac and around Beauvoisin). 

• Hérault: the Grammont station has disappeared 208, 246 , but the 

other populations are stable (temporary pools and streams of the 

Montpellierais garrigues, the Bittérois plain, and the plateau of 

Roque-Haute, the Assignan pool in the Minervois). 

• Pyrénées-Orientales: the species has disappeared from the 

Saint-Estève pool (alluvial plain of the Têt) and the Catalan coast 246 

but survives on the Rodès plateau. 

• Bouches-du-Rhône and Vaucluse: With one exception (Etang 

des Aulnes), all the stations of the “Crau humide” (Marais du 

Vigueirat, Marais de l’Audience, Marais de Raphèle) and the 

“Crau sèche” (Etang du Luquier) appear to have disappeared 246 , 

as have the stations of the sandy banks of the Rhône 157, 158, 254, 255, 311 . 


• In Italy, the Abruzzo populations are considered to have disappeared 

292 . 

• In Spain and Portugal, the species could be declining due to the 

destruction of its habitat (Medina, pers. com.). 

• In Morocco, the populations appear to be stable. 


IUCN 1997 





and Red Lists 


- 

- 

- 





priority species274 181, 133 

- Morocco: “very rare” (RR) 


In France, habitat destruction is responsible for the disappearance 

of several important populations: 

• Reorganisation of land followed by drainage and cultivation 

(vineyards of the Costière Nîmoise). 

•Permanent flooding (Grammont and Saint-Estève pools). 

• Hydraulic modifications (banks of the Rhône). 

• Silting up, shading (cessation of grazing and lack of upkeep of 

the lavognes of the garrigues). 

• Game management of lavognes (sealing of the pool bed with 

a layer of concrete). 

• Extraction of materials: the Plateau de Rodès station is now 

threatened by the creation of a granite quarry 246 . 

Plant species 


The shade linked to invasion by woody species, as a result of the 

abandonment of grazing at some sites (France), is without doubt 

unfavourable for this species. 


In North Africa and in France, with the exception perhaps of the 

Montpellierais and Uzégeois garrigues and the Roque-Haute plateau, 

a risk of extinction of populations exists in the medium to 

long term, because of their small number, their distance from one 

another, and their reduced size. 


Current measures 274 

Several stations benefit from protected status: 

• The Site Classé of the Gardiole (commune of Gigean in the 

Hérault). 

• The Roque-Haute Nature Reserve. 

• The Etang des Aulnes, property of the département of 

Bouches-du-Rhône, included in the network of Espaces Naturels 

Sensibles*. 


• Maintain a favourable hydrological regime 

• Maintain open vegetation by cutting or grazing. 

• Protect the habitat at the remaining stations (Rodès plateau in 

the Pyrénées-Orientales; Etang de Pujaut, Quissac pools etc, in 

the Gard). 

• Raise awareness among the rural public of the interest and 

richness of the many lavognes of garrigue zones so they can be 

maintained and prevent their beds from being concreted over. 

• Intensify prospecting in the areas where only a few populations 

are known (Minervois, the piedmont plain of the Cévennes: 

Quissac, Alès etc.). 

• Carry out inventories and monitoring of these populations 

(dynamics), which are vulnerable due to the fragility of their 

habitat. 

Bibliography 

Aizpuru et al., 2000 6 ; Anonymous, 1999 12 ; Bissardon & Guibal, 

1997 39 ; Bonnier, 1990 42 ; Braun-Blanquet et al., 1952 49 ; Conti et 

al., 1992 86 ; Bern Convention, 1979 87 ; Le Dantec et al., 1998 208 ; 

Directive 92/43/CEE, 1992 119 ; Fennane & Ibn Tattou, 1998 133 ; 

Foucault de, 1988 98 ; Gaudillat & Haury, 2002 153 ; Girerd,1990 157 ; 

Goujard, 1997 158 ; Greuter et al., 1984-1989 160 ; Jahandiez & 

Maire, 1931-1934 181 ; Loisel, 1976 215 ; Médail et al., 1998 246 ; 

Molina, 1998 251 ; Molinier & Tallon, 1950 254 , 1950-1951 255 ; 

Molinier, 1981 257 ; Olivier et al., 1995 274 ; Pignatti, 1982 292 ; Quézel 

& Santa, 1962-1963 303 ; Quézel et al., 1979 308 ; Quézel, 1998 306 ; 

Reynier, 1883 311 ; Rivas Goday, 1970 319 ; Tutin et al., 1964-1993 386 ; 

Walter & Gillett, 1998 399 . 

Authors: Michaud H. & N. Yavercovski 

Collaborators: Rhazi L., M. Rhazi, F. Ruchon F. & H. Souheil 

53


Myriophyllum alterniflorum DC. 

DICOTYLÉDONES 

HALORAGACEAE 

Main synonyms 

M. verticillatum L. subsp. alterniflorum (D. C.) Bonnier & Layens 

French name: Myriophylle à fleurs alternes 

Spanish name: Ovas de rio 

Italian name: Millefoglio d’acqua gracile 

English names: Alternate-flowered Water Milfoil, Slender Water 

Milfoil 

Subspecies 

None 


• Aquatic plant, herbaceous, slender (10 to -50 cm), rooted, 

glabrous, branching. 

• Leaves green, 6 to 30 mm in length, sometimes reddish, pinnate 

hair-like. Submerged leaves whorled, aerial leaves situated 

on the fertile branches and almost all alternate. 

• Bracts of upper flowers entire or toothed, shorter than the 

flowers. 

• Flowers yellowish, forming a spike, generally with 4 petals. 

Calyx with a short tube with four deciduous lobes. Flowers few, 

single-sex, alternate, in a very slender, short, at first drooping 

spike, consisting of three to four female flowers in a whorl at the 

base and five to eight male flowers in a spike at the tip (with 

eight stamens). 

• Ovary inferior, with four carpels, each one containing four loculi 

with single ovules. Tetragon-shaped fruit, finely tuberculated. 


• Myriophyllum spicatum: has many pink flowers, all whorled, 

forming a flowering spike, robust and always upright 265 . 

• M. verticillatum: has the bracts of the upper flowers all pectinate 

and longer than the flowers. The spike is very elongated, 

terminating in leaves; the flowers are pink. 



Broad distribution: Europe, North Africa, North America. 

Mediterranean countries 64, 86, 386 : France, Spain and Portugal (dispersed 

over all the peninsula), Italy (Lombardy, Lazio, Sila, Sicily 

and Sardinia), Tunisia, Algeria and Morocco. 

France 

• On the mainland, the species is absent from the 

Mediterranean region, and elsewhere occurs in streams and 

ponds with a siliceous soil. 

• In Corsica, it is found in seven temporary pools in the south 

east of the island: four pools in the Tre Padule de Suartone 

Nature Reserve and three pools, of artificial origin, on the 

Frasselli plateau. 

Habitat 

54 

Myriophyllum alterniflorum 


In Morocco and in Corsica, this species is found only in temporary 

pools. In Morocco, it is present in the pools on the plains 

(region of Benslimane) and mountains in the Rif region (Outka and 

Issaguene), on siliceous soils (sandstone, quartzite) and in a semiarid 

bioclimate*, sub-humid or humid. These pools are generally 

filled with rainwater in December-January and dry out in April- 

May. 



0.5 cm 


This species appears in the aquatic phase of oligotrophic* temporary 

pools. 



In Morocco, the species is characteristic of Myriophyllum alterniflorum-Ranunculus 

peltatus communities. The species associated 

with this community are Callitriche brutia, Glyceria fluitans, 

Chara vulgaris, Nitella opaca and Illecebrum verticillatum. 

In the pools of Corsica (pools with Isoetes velata), the associated 

species are Ranunculus peltatus, Illecebrum verticillatum and 

Apium crassipes, as well as Potamogeton pectinatus on the 

Frasselli pools (which indicates slight eutrophication linked to 

the presence of livestock).

Ecology 


Life form 

Perennial species in permanent waters (hydrophyte*) but with an 

annual cycle (therophyte*) in temporary Mediterranean pools. 

Reproduction 

Aerial pollination. Flowering in March-April. 


Seeds: four per fruit, small (length: 1.5 mm; width: 1 mm to 1.25 

mm). 


This begins under the water and often ends out of the water. 

In Morocco and Corsica, germination takes place in February- 

March, when the pool is under water, and flowering in March- 

April. Fruiting, which begins during the aquatic phase, takes 

place in May-June, but the maturation of fruits and seeds, and 

their dispersal a few centimetres from the mother plant, occurs 

in June-July after the drying-out of the pool. In addition to this 

sexual reproduction, the plant reproduces vegetatively during the 

aquatic phase, by division and regrowth. 

Morocco and Corsica 


Germination 

Flowering 


Fruiting 


Hydrology 

The germination of seeds requires flooding or at least waterlogging 

of the soil. The plant is adapted to submerged conditions 

(divided leaves, etc.) and can withstand high water levels provided 

there is sufficient light: tolerance to water depth increases 

with the clarity of the water (for example 50 cm in the pools of 

Benslimane in Morocco or Padule Maggiore in Corsica). The plant 

forms its flowers under the water but they emerge above the surface 

of the water for opening and pollination (anemophily). The 

water is oligotrophic* to mesotrophic*. 

Substrate 

Acidic to neutral. 


M. alterniflorum is abundant in the centre of pools, where herbaceous 

species such as Ranunculus peltatus, Callitriche brutia, etc. 

grow, but it becomes less abundant or even absent in areas with 

helophytes* (Scirpus maritimus, Eleocharis palustris), due to competitive 

exclusion. 

Plant species 


Favourable when they facilitate the opening-up of the habitat. 

Light 

The species is heliophilous*, which explains its absence in the 

turbid pools of the Cork-Oak forest of Mamora in Morocco and 

in deep pools (with greater depths of water). 



• In Morocco, Myriophyllum alterniflorum has been found in 

14 pools 133, 230 in the Atlantic plains (Benslimane) and the mountainous 

region of the Rif. 

• In Algeria, the species is considered to be rare. There are few 

sites with M. alterniflorum and these are confined to just two 

regions, the Algiers region (coastal subregion) and the Kabyle 

region (Greater and Lesser Kabyle and Numidia) 303, 338 . 

• In Mediterranean France, there are seven pools, with M. alterniflorum, 

three of them artificial, and all of them in Corsica. 

• On the Iberian Peninsula and the Balearics, the species is widely 

distributed (40 provinces in Spain, five in Portugal). 


In Morocco, in Spain and in Corsica, the populations of 

M. alterniflorum appear to be stable. 

In Italy, the stations in Tuscany have disappeared 86 . 


IUCN 1997 





and Red Lists 


- 

- 

- 

- 

- Morocco: 

133, 181, 381 





Filling in, urbanisation, cultivation and the modification of the 

hydrology of pools (drainage) constitute the main threats for this 

taxon. In Morocco, there is a significant risk of extinction of populations 

at some stations because of urbanisation and the fillingin 

of pools. 


In Morocco, the frequency of dry years limits the appearance of 

this species 313 and reduces the chances of renewal of the seed 

bank *, which could pose problems in the long term. 

In Corsica, the closing-up of the habitat (invasion by woody 

species or by helophytes) is a potential threat for the species if 

grazing is abandoned. 

55




In Corsica, the populations of the Tre Padule de Suartone pools 

benefit from the legal status of the site (Nature Reserve). In Italy, 

the Sicilian stations are included within a Regional Park 86 . 


• Maintain the healthy hydrological functioning of the sites, 

avoiding drainage and silting up. 

• Control urbanisation. 

• Maintain an open habitat by controlling the dynamics of the 

vegetation (woody species and helophytes), and maintaining or 

introducing grazing. 

• Implement legal measures for conservation of non-protected 

sites. 

56 

Bibliography 

Anonymous, 1999 12 ; Castroviejo, 1986-2001 64 ; Conti et al., 

1992 86 ; Fennane & Ibn Tattou, 1998 133 ; Fournier, 1961 139 ; 

Jahandiez & Maire, 1931-1934 181 ; Lorenzoni, 1997 225 ; Lorenzoni 

& Paradis, 2000 221 ; Maire, 1952-1987 230 ; Marchetti, 1997 235 ; 

Nègre, 1961-1962 265 ; Pignatti, 1982 292 ; Quézel & Santa, 1962- 

1963 303 ; Rhazi et al., 2001 313 ; Rhazi, 2001 312 ; Samraoui & de 

Belair, 1998 338 ; Titolet & Rhazi, 1999 381 ; Tutin et al., 1964- 

1980 386 . 

Author: Rhazi L. 

Collaborators: Paradis G. & M. L. Pozzo di Borgo

Nitella opaca (Bruzelius) Agardh 

CHAROPHYCÉES 

CHARACEAE 

Main synonyms 

Chara opaca Bruzelius 1824 

N. flexilis var. flexilis f. opacoides R.D. Wood 1962 

French name: none 

English name: Dark stonewort 

Subspecies 

None 

90, 164, 200, 358 


• Dark green plants, small in size (10-30 cm), living entirely submerged. 

Thallus with whorled structure, entirely without cortication. 

• Whorls composed of 6-8 primary rays (branchlets), divided 

once to form 2-3 secondary rays (dactyls). Dactyls always unicellular 

with a blunt tip. Gametangia* situated in the forks of the 

primary rays. 

• Dioecious species, very fertile with a marked sexual dimorphism. 

• Male plants characterised by dense fertile heads with short 

dactyls; antheridia* solitary and of large size (diameter 650- 

775 µm). 

• Female plants with longer primary and secondary rays, giving 

them a more open appearance; fertile nodes carrying up to three, 

though usually two oogonia* 600-700 µm long and 500-600 µm 

wide. Oogonium bearing a coronule* composed of 2 x 5 tiny cells. 

Oospores laterally compressed, dark brown to black, c. 350-500 µm 

long and 350-400 µm wide with 6-7 spiral ridges. In the fresh 

state, the oospores have characteristic “winged” extensions of 

the spiral ridges. 

- Incrustations of calcite forming more or lesse regularly spaced 

rings around yhe axis and branchlets, often occur at stations with 

a low water level. This feature, known as “banded phenomenon of 

the Characeae” 310 , may be attributed to the effect of intense light356 . 


Two other dioecious species can ressemble Nitella opaca: 

• N. syncarpa (Thuillier) Chevalier, but this species grows in 

summer or autumn, not in spring. 

• N. capillaris (Krokeil) Groves and Bullock-Webster, which also 

has spring growth but is distinguished by gelatinous mucus 

around the male and female gametangia 90 , and by its oospores 

which are ornamented with papillae 131, 357 . 

N. opaca is, with N. tenuissima, the only representative of this 

genus so far recognised in Languedoc-Roussillon and southeast 

France. It is easily distinguished from N. tenuissima which has 

primary branches with two to four forks (multiple dichotomy). 



Plant species 

France 

Nitella opaca is mainly present in central and western France 

(Aquitaine andBritany) 90 . The numerous localities listed by Corillion 

all date from earlier than 1957 and even from the beginning of 

the 20 th century. A single earlier mention of N. opaca concerns 

southern France near Hyères in the Var (coll. Boulu 1851, determined 

and then cited by Hy 178 ). Today, there are only five localities 

known in the south of France: Lanau pool (Bouches-du- 

Rhône), the temporary lakes of Bonne Cougne and Redon, (Var) 358 , 

some pools in the natural reserve of Roque-Haute (Hérault) and 

one pool at the Tour du Valat in the Camargue (Grillas, pers. 

com.). It has disappeared from the Moulin du Rouet site (Hérault) 

where it was recorded by W. Krause. 


Rare in Spain 200 and Portugal 90 ; unknown in Italy and Greece. 

Algeria: three localities previous to 1950 132 . 

Morocco: five localities in 1974 165 and from 1985-1990 130 . 

Tunisia: the species has been identified in the Holocene sediments 

of a fresh water locality close to the Mediterranean coast 131 . 

Habitat 

male female 

Nitella opaca 

In the Mediterranean region, the species has been identified only 

in temporary freshwater habitats, characterised by flooding at 

the end of the autumn and drying out in May/June. This type of 

habitat seems to be consistent both in southern France and the 

localities in North Africa. 



1 cm 

57 

Modified from Groves & Bullock-Webster 164





22.342 Tall Mediterranean amphibious swards (Preslion cervinae). 


N. opaca is part of the aquatic phase of Preslion and Isoetion 

communities (Isoetalia). 

This species is also characteristic of the Nitelletum opacae association, 

often in monospecific* 90 populations or associated with 

Sphaerochara (personal observation). 


At the Lanau pool in the Crau (Bouches-du-Rhône), N. opaca is 

associated with Sphaerochara intricata 355 . These two species precede 

the other hydrophytes* and decompose during the emergence 

of communities of amphibious annuals with Damasonium 

polyspermum, Lythrum tribracteatum, Mentha pulegium, etc. 

At Lake Redon (Var), N. opaca is accompanied by Sphaerochara 

prolifera. The growth of this species also early and precedes that 

of Ranunculus ophioglossifolius, Baldellia ranunculoides and Butomus 

umbellatus. 

At Roque-Haute, it grows side by side with Isoetes setacea, 

Mentha cervina and Callitriche brutia in some deep pools. 

In the pools of Morocco, N. opaca is associated with Myriophyllum 

alterniflorum, Callitriche brutia, Glyceria fluitans, Chara 

vulgaris and Illecebrum verticillatum 312 . 

Ecology 


Life form 

Nitella opaca is an annual species. 


This is a vernal species, which grows while the water is cold. 

Thus, it appears very early in shallow habitats, while in western 

France (Britany) and in northern Europe, it is found up until the 

summer in permanent lakes and even down to a depth of 40 m 

in Sweden 90, 200 . In the Mediterranean region, it is present from 

February onwards and fructifies abundantly from the end of 

March to May. The plants then decompose entirely. 

58 


� 


Reproduction 

Decay 

� 


Figure established on the basis of monthly observations at the temporary lakes of 

Bonne Cougne and Redon in 2001 358 . 

90, 200 


Hydrology 

N. opaca is a fresh water species of habitats with a pH* close to 

neutral (6.5-7.5). N. opaca can be found in both temporary and 

permanent habitats and also adapts to weak currents. The minimum 

duration of submersion necessary to complete a full biological 

cycle is estimated at 5-6 months. 

Substrate 

Indifferent to substrate. 


Because of its early appearance, N. opaca is not subjected to 

much competition from other hydrophytes. 


Because it is abundantly fertile, the species is not very sensitive to 

“mechanical” disturbances, and even seems favoured by the trampling 

of livestock (Lanau pool) and wild boar (the longer lasting wet 

parts of Lake Bonne Cougne), which limit the terrestrial vegetation. 

Temperature 

Germination in cold water, fruiting between 12 and 18°C; 

N. opaca decays when the temperature of the water exceeds 

20°C (personal observation). 

Light 

Not very light-demanding; however, sexual reproduction is reduced 

in deep lakes 90 . 



France 

Distribution is concentrated in central and western France where 

it nonetheless appears to be in decline. An estimate of the number 

of sites in these regions is not currently possible. Five localities 

are known in the French Mediterranean region. 


France 

In overall decline: more of 32 localities listed by 91 for western 

France, mainly Britany, had only been collected at the beginning 

of the 20 th century. 

Other European countries 

Unevenly distributed in northern Europe; considered as fairly frequent, 

but nonetheless vulnerable, in Spain 79 , while Krause 200 mentions 

rare sites in central Spain. The species has been recorded 

recently in the Balkan countries 40 . 


IUCN 1997 





- 

- 

- 

- 

-

There is currently no legal protection for charophytes in France 

and the Mediterranean countries, nor are they included in any 

Red List of threatened species. 



In France: 

• The hydrological regime and the water quality of the Lake 

Bonne Cougne (Var), which is partly supplied by a karstic* aquifer, 

are seriously threatened by a golf-course project. 

• At Roque-Haute, the absence of control over land ownership 

and usage (ban on access to the site by owners) is preventing the 

monitoring and the management of populations of Nitella opaca. 



At Roque-Haute, the site benefits from the status of Nature 

Reserve. The pools underwent monitoring from 1997 to 2001. 

However, since 2002, the manager of the Nature Reserve has 

been prevented by the owners of the land from carrying out his 

monitoring and management activities. 

The Lanau pool has been acquired by the CEEP and undergoes 

regular monitoring. It is included in a Natura 2000 area, like the 

temporary lakes of Redon and Bonne Cougne. 

Plant species 


It is essential to maintain the temporary nature of the hydrological 

regime with flooding during the cold season. In particular, in 

the Mediterranean region, care must be taken to maintain the 

natural alternation of flooding/drying out, which favours sexual 

reproduction and, because of this, enables the plant to spread to 

new pools. 

Populations of N. opaca are vulnerable due to the fragility of 

their habitat and it is thus important that the dynamics of these 

populations are inventoried and monitored. 

Bibliography 

Blazencic & Blazencic, 2003 40 ; Cirujano et al., 1992 79 ; Corillion, 

1957 90 , 1975 91 ; El Khiati et al., 2002 131 ; El Khiati, 1995 130 ; 

Feldmann, 1946 132 ; Groves & Bullock-Webster, 1920 164 ; 

Guerlesquin, 1978 165 ; Hy, 1913 178 ; Krause, 1997 200 ; Raven et 

al.,1986 310 ; Rhazi, 2001 312 ; Soulié-Märsche & Thiéry, 1998 355 ; 

Soulié-Märsche & Vautier, 2004 356 ; Soulie-Märsche, 1989 357 ; 

Soulié-Märsche, 2003 358 ; Wood, 1962 403 . 

Author: Soulié-Märsche I. 

59


Ophioglossum azoricum C. Presl (1) & Ophioglossum lusitanicum L. (2) 


OPHIOGLOSSACEAE 

Main synonyms 

(1) O. ambiguum Cosson & Germ 

O. polyphyllum auct. non A. Braun 

(2) O. vulgatum L. subsp. lusitanicum (L.) Bonnier & Layens 

French names: (1) Ophioglosse des Açores, (2) Ophioglosse du 

Portugal 

Italian names: (1) Ophioglosso delle Azzorre, (2) Ophioglosso 

lusitanico 

Portuguese names: (1) Lingua de cobra, (2) Lingua-de-cobra, 

Lingua-de-cobra-menor 

English names: (1) Small adder’s tongue, (2) Least adder’s tongue 

96, 299 


•Perennial plants, delicate and unobtrusive from 5 to 10 cm (1), 

and 2 to 5 cm (2), present in the form of small, fairly localised 

populations 

•Two types of fronds (“leaves”) very distinct, as in all the 

Adder’s-Tongues: a sterile type, with an extended blade, oval or 

lanceolate, sometimes lying flat but usually upright and curled 

over to form a furrow; the other fertile, reduced to a fruiting 

spike with a peduncle and bearing sporangia stacked up on top 

of each other in two symmetrical rows (at the origin of the name 

“ophioglosse” from the Greek glossa “tongue” and ophis “snake”). 

• Fronds of the two types separated at ground level (the sterile 

fronds generally sheathing the fertile fronds), light green or yellowish-green 

in colour, and yellowing completely before wilting, 

after sporulation (production of spores). 

• Generally one single sterile frond per plant, but can grow in 

pairs in O. azoricum, in which the fronds are broader. 

• Spikes (or fertile fronds) do not invariably appear on every 

plant. 


Confusion is easy between the three species of Ophioglossaceae, 

Ophioglossum azoricum, O. lusitanicum and O. vulgatum. 

O. vulgatum is the largest (15 to 30 cm) and most widespread. 

Although present in the Plaine des Maures in proximity to near 

temporary pools, it is generally found in more eutrophic* habitats, 

undergrowth or damp grasslands. 

O. azoricum and O. lusitanicum are closer in size and share habitats 

that are at first sight similar, but are in fact different in 

terms of hydrology. The easiest distinction criterion is the 

autumn-winter growth period for O. lusitanicum, spring for 

O. azoricum. In early spring the two species can be present at the 

same time but their stages of development (colour, sporulation) 

are visibly out of step. 

Finally, note that the number of chromosomes differs between 

the three species: O. lusitanicum is diploid* (2n=240 chromosomes), 

O. vulgatum is tetraploid (2n=480) and O. azoricum 

hexaploid* (2n= 720), suggesting that this latter species is derived 

from a cross between the other two. 

60 



1 cm 

Ophioglossum azoricum & Ophioglossum lusitanicum 

(1) Ophioglossum azoricum 

Western European species present in France and around the 

Mediterranean in Spain, Portugal, Sardinia and Italy 160 . It also 

occurs locally in Slovakia and in the Czech Republic, the Canaries 

and Madeira (Azores) 300 . 

France 

•Atlantic coast. 

• Mediterranean region: O. azoricum is present in the Plaine des 

Maures and the Estérel (Var), in the garrigues of the Hérault, the 

Cévennes in the Gard and Lozère, and in Les Fenouillèdes (Pyrénées- 

Orientales). In Corsica, this species is only found in rares 

pozzines* on the Tenda Massif and the Plateau du Coscione 142 . 

(2) Ophioglossum lusitanicum 

Mediterranean and Sub-Atlantic species present all around the 

Mediterranean Basin (France, Portugal, Spain, Italy, Yugoslavia, 

Greece, Crete, Croatia, Algeria, Tunisia, Morocco, Turkey, Lebanon) 

as well as the Canaries, Madeira and England. 

France 

• Atlantic coast (Béarn, Massif Armoricain and its islands). 

• Mediterranean region: present in Les Maures and the Estérel, the 

Biot Massif and Les Albères. Formerly reported from the marshes 

of Coustière in the Crau. Fairly common in Corsica at low altitude 141 .


• Europe: Portugal, Spain, Italy, Yugoslavia, Greece, Crete and 

Croatia. 

• North Africa: Algeria (in the Constantinois Tell), Tunisia and 

Morocco (Middle Atlas, Rif mountains, north and mid-Atlantic 

plains). 

•Turkey, Lebanon. 

Habitat 


(1) Ophioglossum azoricum 

The habitats favourable to this species are very wet in winter and 

spring but rarely flooded for a long period. These are generally 

depressions, beds or edges of small streams and springtime temporary 

runnels on well-exposed hillsides on thin soils (Cévennes 

and Avants-Monts in Languedoc). 

Its habitat, in the Mediterranean region, is short swards, open or 

in a mosaic with the shrubby maquis vegetation, on thin, rather 

acidic, soil. The habitats found in the Atlantic region are coastal 

swards, dune slacks, sandstone shelves or other types of rock surfaces. 

On the mainland, the stations are not found above 700 m 

in altitude but in Corsica they are found up to 1,500 m in peaty 

basins (pozzines), which remain damp in the summer. 

The substrate is not very thick, sandy and often acidic. 

(2) Ophioglossum lusitanicum 

The habitats are comparable to those of the preceding species 

(temporarily wet short swards): rock shelves on coastal hillsides, 

edges of pools and temporary streams, damp depressions on the 

maquis and small basins in rocky outcrops. However, the species 

rarely mixes with the preceding species: it seems to seek stations 

with even thinner substrates, and seems less demanding of a 

water supply. It is clearly more thermophilic and is even found 

away from the low-altitude coastal zone. 

In Morocco, O. lusitanicum is mainly found in the sandy or stony 

pastureland of the plains and low hills under a Mediterranean 

mild semi-arid and sub-humid bioclimate. In Algeria, its presence 

is limited to sandy soils. 

(1) and (2): Ophioglossum lusitanicum profits from temporary runnels 

resulting from autumn rains, which are generally more significant 

than spring rains, whereas the cumulative effect of the 

supply of water in the depressions seems better adapted to the 

demands of O. azoricum. On a typical profile through a temporary 

pool on mainland France (Maures), O. lusitanicum will thus 

be situated higher than O. azoricum. In Corsica, the two species 

have very different distributions: O. lusitanicum is found at low 

altitude, while O. azoricum is only present in some higher-altitude 

pozzines. 

Directive Habitat 






Plant species 

Mediterranean regions 

• In a European list, Brullo & Minissale 54 include Ophioglossum 

lusitanicum among the characteristic species of the alliance 

Cicendio-Solenopsion laurentiae (within the order Isoetalia, class 

Isoeto-Nanojuncetea). In Spain, Rivas Goday 319 has described an 

Ophioglosso-Cicendietum filiformis association. 

• In crystalline Provence, Loisel 215 described O. lusitanicum as 

characteristic of the association Isoeto-Nasturtietum. 

• In Languedoc, in the Avants-Monts, O. azoricum is noted as a 

participant in a hygrophytic variant of a Scillo-Ranunculetum paludosii 

98 . In the Cévennes, O. azoricum forms part of Isoetion communities 

with Isoetes duriei which have not been described 44, 195 . 

• In Roussillon, O. azoricum is associated with various undescribed 

Isoetion communities 212 . O. lusitanicum is associated in temporary 

runnels with I. duriei 272 . 

Other regions 

• In western France, Ophioglossum azoricum is part of a north- 

Atlantic Ophioglosso azorici-Agrostietum caninae or a continental 

thermo-Atlantic Ophioglosso azorici-Isoetetum histricis whereas 

O. lusitanicum is one of the characteristic species of a coastal 

thermo-Atlantic Ophioglosso lusitanici-Isoetetum histricis 98 . In 

the Natura 2000* Habitats Register, this latter community is 

divided into two: Romuleo columnae-Isoetetum histricis and a Chamaemelo 

nobilis-Isoetetum histricis 153 . 

Ecology 


Life form 

Perennial species persisting in the summer through a rhizome 

(geophyte* with rhizome). 

Adaptive strategy 163 

Stress-tolerant (of summer drought). 

Reproduction 

Production of spores aerial, fusion of gametes in wet conditions. 


Growth of fronds with flooding (autumn or spring rains according 

to the species), maturation of spores in winter for Ophioglossum 

lusitanicum in late spring/early summer for O. azoricum; death of 

the leaves after sporulation, which coincides with the summer 

drying-out period for O. azoricum. The phenology* depends on 

the region. In the Mediterranean region, the cycles can differ 

from one year to the next depending on the weather conditions. 

In Morocco, the biological cycle of O. lusitanicum also begins in 

autumn (November). It terminates in the spring (May) with the 

withering of the fronds. 

61


O. azoricum 

France 


O. lusitanicum 

France 


Hydrology 

Germination of spores requires a very wet soil, supplied regularly 

during this phase with oligotrophic* water. These plants withstand 

occasional flooding. 

Substrate 

Acidic, fairly thin 


Heliophilous* species, which does not thrive in the shade of taller 

species. 


Generally favorable, but overgrazing constitutes a mid-term 

threat for Moroccan stations. 



France 

• In mainland France, the two species seem very local; the number 

of stations is not exactly known. 

• In Corsica, Ophioglossum lusitanicum is fairly frequent on the 

coast, O. azoricum is local. 

Morocco 

Ten or so stations* with O. lusitanicum have been censused, and 

this number could increase with a more systematic exploration 

of the more inaccessible regions of the Middle Atlas and the Rif. 

Spain 

O. lusitanicum is present in the western half of the country 

(19 provinces), and in the northeast in the province of Girona, 

and in the Balearics. O. azoricum is scattered in several provinces 

in the centre and northeast 64 . 

Italy 

O. lusitanicum is fairly widespread on the peninsula, but rare on 

the islands (Sicily, Sardinia) 292 . 

62 










(1) O. azoricum 

The number of stations is noticeably increasing in France, no 

doubt because of increased knowledge (the plant is either absent, 

or very poorly described in older works) and improved surveying, 

even though the habitats in which this species are found are a 

priori vulnerable. The species has not been found again at Roque- 

Haute (Hérault). 

(2) O. lusitanicum 

The number of stations is in noticeable decline in France, particularly 

in the Atlantic zone. In Morocco, the number of stations is 

fairly stable, although the populations are slightly increasing at 

the mountainous stations. In Algeria, the species is considered to 

be rare. 


IUCN 1997 





and Red Lists 


- 

- 

- 

- (1) France, national list: 


Officiel 13/05/1982, amended 

by Decree of 31/08/1995/ 


- (2) France, regional lists: 

• Languedoc Roussillon: Decree 


16/10/1998 

• PACA: Decree of 9/05/1994/ 


- (1) France, national Red Data 

Book: priority species 274 


The very specific requirements of the Adder’s-Tongues, in both 

trophic and hydrological terms, mean that these plants are 

species which are very sensitive to modifications in the balance 

of their environment. The localised nature of the stations accentuates 

their vulnerability. Modification in the hydrology of the 

sites is a frequent degradation factor. As the habitats occupied 

by these species are generally not very fertile for agriculture, the 

most immediate threat remains the development of sites, in particular 

for all the coastal stations. The Adder’s-Tongues are often 

established on thin soils which are sensitive to erosion. Excessive 

numbers of visitors can contribute to degradation at some stations. 


The closing-up of the habitat by vegetation can be a limiting factor 

for populations. However, the sites being generally not very 

productive (skeletal soils), the growth of the competitive vegetation 

is slow: in particular, populations of Adder’s-Tongues on rocky 

shelves are in a virtually climax* situation and not vulnerable to 

this factor. 

In addition, the competitive vegetation is often sensitive to perturbations 

(grazing, scrub clearing, etc.).



France 

• The Conservatoire Botanique National Méditerranéen de Porquerolles 

is conserving O. lusitanicum spores ex-situ. 

• In the Plaine des Maures: 

- Some areas supporting these species belong to the CEEP or the 

Conservatoire de l’Espace Littoral et des Rivages Lacustres. 

- Within the framework of the LIFE “Temporary Pools” project, 

experimental management measures are in place in several sites 

supporting these species (Péguière pool, Les Aurèdes site) and a 

site has been acquired in the Bois de Rouquan by the CEEP. 

- The grazing management and scrub clearance implemented for 

the Défense Forestière Contre les Incendies can be favourable for 

these species. 

- Several stations are included in the Natura 2000* area. 

Morocco 

There are no measures for the management of Adder’s-Tongue 

sites. 


• Management of the sites supporting Adder’s-Tongue populations, 

through statutory measures guaranteeing the maintenance 

of ecological conditions favourable to their survival, is the best 

way of conserving the stations. In particular, the implementation 

of decrees for biotope protection, notably for certain coastal 

populations, is recommended. 

Plant species 

•A more exact census of the stations of these inconspicuous 

plants is also recommended. In most favourable natural habitats, 

their presence is undoubtedly underestimated. Inclusion of the 

sites in various inventories (ZNIEFF*, Natura 2000, etc.) is essential. 

• From time to time, management measures aiming to maintain 

an open habitat or re-establish a satisfactory hydrological regime 

can be useful to their conservation, following evaluation. 

Bibliography 


Guibal, 1997 39 ; Boudrié, 1995 44 ; Braun-Blanquet, 1935 48 ; Brullo 

& Minissale, 1998 54 ; Castroviejo, 1986-2001 64 ; Chevassut & 

Quézel, 1956 72 ; Conti et al., 1992 86 ; Danton & Baffray, 1995 96 ; 

Fennane et al., 1999 134 ; Foucault de, 1988 98 ; Gamisans & Guyot, 

1991 142 ; Gamisans & Jeanmonod, 1993 141 ; Gaudillat & Haury, 

2002 153 ; Greuter et al., 1984-1989 160 ; Jahandiez & Maire, 

1931 181 ; Kessler, 2000 195 ; Lewin & Escoubeyrou, 1997 212 ; Loisel, 

1976 215 ; Nozeran & Roux, 1958 272 ; Olivier et al., 1995 274 ; 

Pignatti, 1982 292 ; Prelli, 2001 300 ; Prelli & Boudrié, 1992 299 ; 

Quézel, 1998 306 ; Rivas Goday, 1970 319 

Author: Catard A. 

Collaborators: Michaud H., G. Paradis, L. Rhazi & N. Yavercovski 

63


Pilularia minuta Durieu ex A. Braun 


MARSILEACEAE 

Main synonyms 

P. globulifera L. subsp. minuta (A. Braun) Bonnier & Layens 

French name: Pilulaire délicate 

Italian name: Pilularia minore 

English name: Pillwort (name of genus) 

Subspecies 

None 

64, 96, 292, 300 


• Amphibious plant, perennating by means of its filiform creeping 

rhizome, just slightly buried under the surface of the ground. 

• Fronds (“leaves”) green, filiform, cylindrical, upright from 1 to 

3 cm in length and arranged singly along the rhizome, the internodes 

being around 1 cm. 

• Sporocarps* (“fruits”) globular, from around 0.75 to 1 mm in 

diameter, densely downy, with short pedicels (the pedicel is 

recurved at the apex) and dark brown in maturity. 


Seedlings of rushes, grasses and Isoetes. The distinctive characters 

of Pilularia minuta are the filiform rhizome very slightly buried 

in the soil, the globular sporocarps and the filiform leaves. 




France 

Hérault, Alpes-Maritimes and Corsica. 


Portugal, Spain (Minorca and probably in Andalusia and 

Catalonia) 64 , Italy (Sardinia, Sicily, Rome region), North Africa 

(Morocco and Algeria), Greece, Cyprus and Turkey. 

Habitat 


Pools with Pilularia minuta are mainly (mainland France, Corsica, 

Morocco), confined to low-altitude forests or matorrals* dominated, 

depending on the site, by Cork Oak (Q. suber), Myrtle 

(M. communis), Broom Heather (E. scoparia), Mastic (P. lentiscus) 

and Phillyrea angustifolia. Other pools with P. minuta are situated 

in degraded vegetation dominated by Woody Fleabane (Dittrichia 

viscosa). The soil there is sandy to sandy-silty and the bedrock 

sandstone, quartzite, basalt or schist. In Morocco, floooding usually 

occurs in December-January, solely by rainwater, and drying 

out in April-May. At Roque-Haute (France) and in Corsica, flooding 

begins in autumn (November) and ends between April and July, 

depending on the pools. 

64 






22.3412 Aquatic Mediterranean Quillwort swards. 


The species associated with Pilularia minuta are numerous: 

Isoetes velata, I. setacea, Myosotis sicula, Elatine brochonii, 

Juncus pygmaeus, J. bufonius, Illecebrum verticillatum, Lythrum 

thymifolia, L. borysthenicum, L. hyssopifolia, Laurentia michelii, 

Exaculum pusillum, etc. 


• Traditional phytosociological system 319 : communities with 

Pilularia minuta belonging to the alliance Isoetion Br.-Bl. 1931 of 

the order Isoetalia Br.-Bl. 1931 em. Rivas-Goday 1970 and the 

class Isoeto-Nanojuncetea Br.-Bl. & R. Tx. 1943. 

Braun-Blanquet 48 considers P. minuta to be one of the characteristic 

species of his Isoetetum setaceae or Isoetes setacea- 

Lythrum borysthenicum association (= Peplis hispidula) of Roque- 

Haute (Hérault). 

• More recent phytosociological system 98 : communities with 

P. minuta belong to the Antinorio agrostideae-Isoetion velatae 

alliance (Br.-Bl. 1931) of de Foucault 1988, of the order Isoetalia 

velatae (Br.-Bl. 1931) of de Foucault 1988 and the class Isoetea 

velatae (Br.-Bl. & R. Tx. 1943) of de Foucault 1988. 

Ecology 


Pilularia minuta 

0.5 cm 

Life form 

Perennial species: rhizomatous geophyte* or radicant 

hydrophyte* 292 .

Reproduction 

Aerial for the maturation of the spores. 


Germination of spores in very damp or flooded conditions and 

growth of the fronds in the water in March-April, followed by the 

formation of sporocarps. In late spring, maturation of the sporocarps 

and the spores after complete drying out; withering and 

rapid death of the fronds. 

Morocco 


France 









Hydrology 

Germination of the spores requires the saturation or flooding of 

the substrate. The plant can remain in the vegetative stage under 

oligotrophic, very shallow water (



Risk of extinction of populations of Pilularia minuta in mainland 

France, due to their reduced numbers and their isolation. 



Morocco 

No current management measures. 

France 

• Mainland France: at Roque-Haute, on certain sites with Pilularia 

minuta, the clearing of woody species and the removal of 

litter to encourage Isoetes setacea were tested within the framework 

of the LIFE “Temporary Pools” project. However, the conservation 

of this population, situated on a Nature Reserve, is now 

uncertain (ban on access, and unilateral management by owners). 

• Corsica: the creation in December 2000 of the Tre Padule de 

Suartone Nature Reserve (Bonifacio) should allow management 

measures to be put in place which are favourable to this species 

in the four pools of the Reserve. 

• Collection and conservation ex-situ of the spores by the Conservatoire 

Botanique National Méditerranéen de Porquerolles. 


• Control urbanisation. 

• Limit the plant cover surrounding the stations of Pilularia minuta, 

to prevent it spreading and to enable the opening-up of the 

habitat to a certain extent. To attain this objective, the extensive 

grazing of cattle should be be maintained or encouraged. 

66 

• Promote legal conservation measures for stations which are 

not yet protected. 

• Implement the management plan at Roque-Haute, and more 

generally restore concerted management with natural-heritage 

objectives on this Nature Reserve. 

Bibliography 

Anonymous, 1999 12 ; Anonymous, 2000 13 ; Bissardon & Guibal, 

1997 39 ; Boudrié et al. 1998 45 ; Braun-Blanquet, 1935 48 ; Castroviejo, 

1986-2001 64 ; Conti et al., 1992 86 ; Danton & Baffray, 

1995 96 ; Daumas et al., 1952 97 ; Fennane & Ibn Tattou, 1998 133 ; 

Fennane et al., 1999 134 ; Foucault de, 1988 98 ; Grime, 1979 163 ; 

Hébrard, 1990 172 ; Jahandiez & Maire, 1931-1934 181 ; Lorenzoni 

& Paradis, 1997 219 , 2000 222 ; Lorenzoni, 1994 223 , 1996 224 ; Maire 

1952-1987 230 ; Médail et al., 1998 246 ; Olivier et al., 1995 274 ; 

Phitos et al., 1995 290 ; Pignatti, 1982 292 ; Poirion & Vivant, 

1969 294 ; Prelli, 2001 300 ; Quézel & Zevaco, 1964 302 ; Quézel, 

1998 306 ; Rhazi et al., 2001 314 ; Rita, 2000 317 ; Rivas Goday 1970 319 ; 

Saez & Rossello, 2001 331 ; Titolet & Rhazi 1999 381 ; Tutin et al. 

1964-1993 386 ; Walter & Gillett, 1998 399 . 

Author: Rhazi L. 

Collaborators: Michaud H., G. Paradis & M. Rhazi

Ranunculus lateriflorus DC. (1) & Ranunculus nodiflorus L (2) 

ANGIOSPERMS 

RANUNCULACEAE 

Main synonyms 

(1) R. nodiflorus L. subsp. lateriflorus (DC.) P. Fourn. 

(2) R. nodiflorus L. subsp. nodiflorus 

R. lateriflorus DC. var. charbonelii Rouy & Foucaud 

French names: (1) Renoncule à fleurs latérales 

(2) Renoncule à fleurs nodales 

Italian name: (1) Ranuncolo a fiori sessili 

Spanish name: (1) Ranunculo con flores laterales 

Subspecies 

These two taxa are sometimes considered to be two subspecies 

of Ranunculus nodiflorus. 


• Annual herbaceous plants (5-20 cm) pale green with slender, 

fasciculate roots. 

• Species with erect fistulous* cylindrical stems, sometimes radicant 

at the lower nodes, dichotomously branching. 

• Basal leaves petiolate with elliptical or lanceolate-oval blades, 

sometimes floating and in that case with long petioles; middle 

and upper leaves with progressively shorter petioles, entire (or 

slightly sinuate-toothed), lanceolate* to linear-lanceolate. 

Petiole extended at the base into a whitish membranous sheath. 

• Flowers very small, from 2.5 to 3 mm in diameter in Ranunculus 

lateriflorus, smaller (1.5 to 2.5 mm) in R. nodiflorus, solitary, 

sessile, (lower sometimes with short pedicels), and located 

in the forks of the stem or between two leaves on the unbranched 

parts of the main stem. Five pale yellow petals approximately 

equal to the sepals. 

• Achenes not deciduous, with surfaces covered in tubercles, 

very compressed, 1.5 to 2 mm excluding the beak, which approximately 

equals the main body of the achene in R. lateriflorus and 

is very short, not more than one-third of it, in R. nodiflorus. 

• R. lateriflorus is diploid (2n = 16), R. nodiflorus tetraploid 

(2n = 32). 

• Further characteristics for distinguishing the two species, given 

by some authors: in R. lateriflorus the petals are spoon-shaped, 

in R. nodiflorus they are flat (however, this criterion is difficult to 

apply objectively to very small petals which are quickly shed, 

especially as it is variable on a single flower); also, more over of 

equal size, R. lateriflorus may be more robust than R. nodiflorus. 


Other annual Ranunculus species with entire leaves and small to 

tiny yellow flowers may grow in identical habitats (R. revelieri, R. 

longipes, R. ophioglossifolius). However, in reasonably welldeveloped 

individuals there can be no confusion with the two 

above species, which are the only ones with sessile flowers in the 

leaf axils and in the forks of the stem branches. The other species 

have pedunculate flowers. 

Ranunculus lateriflorus 

Ranunculus nodiflorus 

Plant species 

1 cm 

1 cm 

67




(1) Ranunculus lateriflorus 

Paleo-tropical species with a wide but everywhere patchy distribution, 

around the Mediterranean and east to central Asia and 

Siberia: France, Italy, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, 

Algeria, Morocco and Croatia (?). 

In France: basaltic outcrops in the Roque-Haute area (Hérault), 

basaltic plateau of Caux-Fontès-Pézenas (Hérault), Plaine du Regard 

in the south of the basaltic plateau of the Coiron (Ardèche). 

In Italy, three currently known localities: in Sicily, in the Apennines, 

and in the Abruzzo 86 . 

(2) Ranunculus nodiflorus 

Franco-Iberian endemic. 

In France, the stations are scattered especially through the centre 

and west of France: Massif des Fenouillèdes (Pyrénées-Orientales), 

Massif Central in the southern Aveyron and on the basaltic plateaux 

(Chaux) of the Puy-de-Dôme, the Haute-Loire and the Cantal, 

the Paris region around Fontainebleau, the Loiret near Orléans, 

the Indre in the Brenne and the Massif Armoricain. 

In the Iberian Peninsula (Spain and Portugal) they occur in the 

centre and the north. 

Habitat 


In France: non-calcareous areas that are temporarily flooded in 

winter and dry out in spring (pools, ditches or nutrient-poor grassland). 


It grows in fairly deep pools and ditches, even in poor grasslands 

prone to flooding. At latitudes further south than France, it is 

found especially in mountains, sometimes on a peaty substrate 

(pozzines of the High Atlas, pools of the Taurus, Sicily, etc.). 

Outside France, R. lateriflorus sometimes grows on a non-siliceous 

substrate 218 . 


Outside the Mediterranean area, it shows a preference for flatbottomed 

basins, very shallow, covered in a clay-gravely silt of 

maximum depth 3 to 4 cm and scattered around on plateaux 

where there is much exposed bare rock 47, 106 . In the rest of its 

range, this species is found in similar habitats to R. lateriflorus. 

In Corsica, the plant is rare 141 and known only from the mountains 

in the south of the island, from about 900 to 1,600 m altitude: 

• In the pozzines* of the Massif de l’Incudine, in the Plateau du 

Coscione 99, 143, 144 . 

•A stream which dries up from June onwards in the Forêt de 

Marghèse in the Massif de l’Ospedale-Cagna 107 . 

Several of the pools where these Ranunculus grow have partly 

anthropogenic origins: granite quarries in the Pays Bigouden 80 , 

basalt quarries at Roque-Haute, stock watering ponds in the 

Ardèche 234 . 

68 

CORINE Biotope 


(22-11 to 22-13) x 22-32 Oligotrophic to mesotrophic annual 

communities, topographically low-lying, of plain levels, with 

Atlantic affinities, of the class Isoeto-Juncetea. 

Mediterranean region 





Community of EC interest “Oligotrophic to mesotrophic annual 

communities, acidophile, of medium plain to montane levels at 

altitudes ranging from plains to mountainous, of the Isoeto- 

Juncetea” (code 3130). 

Mediterranean region 



Class: Isoeto-Nanojuncetea. 

Order: Isoetalia. 

Alliances: Cicendion filiformis, Isoetion. 

Associations*: 

(1) Ranunculus lateriflorus: 

• France: Isoetetum setaceae 

• North Africa: Sedum nevadense-Juncus pygmaeus community 304 . 

• Eastern Iberian Peninsula: Ranunculus lateriflorus-Damasonium 

polyspermum community 98 . 

• Sicily: Ranunculo-Antinorietum insularis and Myosuro- 

Ranunculetum lateriflori 53, 98, 354 . 

•Taurus (Turkey): R. lateriflorus-Sedum annuum community 305 . 

• Eastern Europe: Ranunculo lateriflori-Limoselletum aquaticae 54 . 


• Outside the Mediterranean area: Bulliardio-Ranunculetum 

nodiflori 106 (= Crassulo vaillantii-Ranunculetum nodiflori Abbayes 

1946) and Ranunculus lateriflorus var. charbonellii (= R. nodiflorus)- 

Sedum villosum community 38 . 

• Mediterranean France: undescribed communities, notably with 

Isoetes setacea, Les Fenouillèdes. 

• Corsica: Gamisans 144 , based on three releves from the pozzines 

in the Plateau du Coscione, described the Ranunculeto-Juncetum 

bulbosi association, which should be renamed Ranunculo nodiflori- 

Juncetum bulbosi. 


In France, both species are accompanied by suites of species 

which depend mostly on the area where they are growing: 

• Languedoc-Roussillon: Isoetes setacea, Lythrum thymifolia, 

L. borysthenicum, Myosotis sicula, Juncus pygmaeus, Herniaria 

glabra, Polygonum aviculare subsp. rurivagum, etc. 

• Ardèche: Crassula vaillantii, Lythrum thymifolia, Polygonum 

aviculare subsp. rurivagum, Ranunculus hederaceus, etc. 

• Massif Central: Sedum villosum, Montia minor, Sagina procumbens, 

Poa annua 

• Region of Fontainebleau and Massif Armoricain: Crassula vaillantii, 

Illecebrum verticillatum, Lythrum portula, Polygonum aviculare, 

Spergularia rubra, Poa annua, Myosotis sicula, etc.

• High Atlas (Morocco): Sedum nevadense, Elatine macropoda, 

Ranunculus batrachioides, etc. 

• Iberian Peninsula: Sedum nevadense, Pulicaria paludosa, Illecebrum 

verticillatum, Exaculum pusillum, Lythrum borysthenicum, 

Juncus pygmaeus, Lythrum thymifolium, Myosotis sicula, etc. 

• Sicily: Antinoria insularis, Lythrum portula, Mentha pulegium, 

etc. 

• Taurus (Turkey): Sedum annuum, Lythrum thymifolia, 

Eleocharis palustris, etc. 

Ecology 


Life form 

Annuals (therophytes*). 


In France, both these Ranunculus germinate during the winter to 

produce floating leaves. Flowering takes place in spring when the 

water level is falling (April-June). The flowers quickly produce 

fruits: flowering and fruiting are almost simultaneous. The plants 

die and wither from the beginning of summer. 

R. nodiflorus is preferentially autogamous and its achenes are 

very buoyant 196 . This is probably also true of R. lateriflorus which 

has the same floral characteristics (very small petals, flowers 

without scent) and similar-sized achenes. The seed bank* of 

course plays a vital role in both species 196 . 

Ranunculus lateriflorus and R. nodiflorus 



Germination 

Growth 

Flowering 


Fruiting 

Hydrology 

Typically amphibious species that require a fairly long period of 

winter flooding in the spring, extending into March-April for 

Ranunculus nodiflorus in Brittany 106 and to April-May for R. lateriflorus 

in Mediterranean France. In a relatively wet climate 

(Massif Armoricain, Paris Basin) the basins occupied by R. nodiflorus 

are shallow, allowing rapid warming in spring followed by 

early drying out (des Abbayes, 1947; Bournérias et al., 2001). In a 

Mediterranean climate, the habitats occupied by R. nodiflorus or 

R. lateriflorus may be under several tens of centimetres of water 

and may dry out later. 

Substrate 

These are both species of oligotrophic* pools, which attain their 

optimum on acid mineral substrates: sandy-gravely soil (pH 5.2- 

6.5) on the Roque-Haute basalts 49 , gravely-clayey silts (pH 5.6- 

5.8), very thin (optimum 1-2 cm), lying directly over rock in the 

Massif Armoricain 106 , granitic sands at Rodès (Fenouillèdes). 

Plant species 


Very sensitive to competition from perennial species. In Brittany 

for example, simply root-stripping of the pool vegetation 

resulted in an increase from 47 R. nodiflorus plants in 1995 to 

3000 the following year at a single site 80 . 




In France, this species occurs in the mountains of southern Corsica 

and a large population is known from Rodès in Les Fenouillèdes. 

At Montalba-le-Château, a station near the above and known in 

the 19th century (Castanier in Herbier Oliver, Montpellier University), 

has not been seen since. There are old reports from the 

Pyrénées-Atlantique and the Hautes-Alpes. It has been mistakenly 

reported from Savoie, the Alpes-Maritimes and the Var. 

Overall, in France, the species is declining due to the disappearance 

of pools (urban development) and the growth of vegetation 

following the cessation of grazing 274 . 


There are three remaining localities in France: Roque-Haute, Caux- 

Fontès-Pézenas and Coiron. It disappeared from the Costière 

Nîmoise in the late 1960s or early 1970s following land ownership 

reorganisation which led to the filling-in of many of the small pools 

(laquets) which were scattered across this plateau (formerly at the 

Laquet de l’Oli at Beauvoisin, on Quaternary siliceous gravels). 

The populations of Aosta and Calabria (Italy) have not been seen 

recently 86 . 


IUCN 1997 





and Red Lists 


- 

- 

- 

- (1) and (2) France, national list: 

Decree of 20/01/1982/ 

Journal Officiel 13/05/1982, 

amended by Decree of 

31/08/1995/ Journal Officiel 

17/10/1995 

- (1) and (2) France, national Red 

Data Book: priority species274 133, 134, 181 

- (1) Morocco: “rare” (R) 

- (1) Italy, National Red Data Book: 



The depression with Ranunculus lateriflorus at Costière (France, 

Gard) was filled in during agricultural developments in the 

1960s-1970s. The population at the Plateau de Caux-Fontès- 

Pézenas survives only in a system of canals used to drain a natural 

depression that has been taken into cultivation. The R. nodiflorus 

population at Les Fenouillèdes is threatened by a quarrying proposal. 

69



The natural succession of perennial formations suppresses the 

development of these Ranunculus; however, they appear to be 

capable of surviving in clearings, even very small ones, within dense 

vegetation. In addition, the seed bank is of considerable size. 


Low risk of local extinction due to the size and longevity* of the 

seed bank. 



In France 

Only the populations of Ranunculus lateriflorus at Roque-Haute 

are located within a nature reserve which benefits from a management 

plan. However, the management plan cannot put into 

practice at this reserve owing to a dispute with the owners. 

The Conservatoire Botanique National Méditerranéen de Porquerolles 

has made collections of R. nodiflorus seeds and is preserving 

them ex situ. 

In Italy 

No measures 

70 


Work towards the acquisition of land at the richest sites (Fenouillèdes, 

etc.) as well as conservation management by a competent 

organisation. 

Bibliography 

Biche, 1881 34 ; Billy, 2002 38 ; Bournérias et al., 2001 47 ; Brullo & 

Grillo, 1978 53 ; Brullo & Minissale, 1998 54 ; Citoleux et al., 

1991 80 ; Conti et al., 1992 86 ; Des Abbayes, 1946 106 ; Deschâtres, 

1991 107 ; Fennane & Ibn Tattou, 1998 133 ; Fennane et al., 1999 134 ; 

Foucault de, 1988 98 ; Gamisans & Jeanmonod., 1993 141 ; Gamisans, 

1970 143 , 1976 144 ; Gaudillat & Haury, 2002 153 ; Jahandiez & 

Maire, 1931-1934 181 ; Kirchner et al., 2003 196 ; Litardière de, 

1955 99 ; López González 1986 218 ; Maire, 1952-1987 230 ; Mandin 

& Hugonnot, 2001 234 ; Médail et al., 1998 246 ; Olivier et al., 

1995 274 ; Quézel, 1957 304 , 1973 305 ; Rouy & Foucaud, 1893 328 ; 

Rouy, 1909 329 ; Sortino et al., 1977 354 ; Tallon, 1967 361 . 


Collaborators: Paradis G. & N. Yavercovski.

Ranunculus revelieri Boreau 

ANGIOSPERMS 

RANUNCULACEAE 

Main synonyms 

R. ophioglossifolius Vill. subsp. revelieri (Boreau) P. Fourn. 

French names: Renoncule de Revelière (Var subspecies: 

Renoncule de Rodié) 

Italian name: Ranuncolo di Reveillière 

English name: Crowfoot (name of part of genus) 

Subspecies 

Ranunculus revelieri Boreau subsp. rodiei (Litard.) Tutin 

(Renoncule de Rodié), and R. revelieri Boreau subsp. revelieri 

(Renoncule de Revelière). 

These two subspecies are, however, poorly differentiated 16, 85 and 

are not accepted by all botanists 141, 109 . 


• Small plant, 10 to 20-30 cm tall, with the stem upright, fistulous* 

(rarely in rodiei), branched and more or less hollow. 

• Basal leaves surrounding the stem, with long petioles and oval 

lanceolate blades. Stem leaves with shorter petioles and more 

narrowly lanceolate. 

• Flowers in a pale yellow perianth, with long pedicels. Petals 

glabrous, shorter than the sepals (very villous on the lower surface 

in revelieri and glabrous or slightly downy in rodiei). 

• Fruits: ovoid achenes, blunt, finely grainy, with a very short 

beak* 96, 274, 386 . 


Ranunculus fontanus C. Presl and R. ophioglossifolius (Vill.) 

Boreau have the petals longer than the sepals 96 . R. ophioglossifolius 

in addition has noticeably larger flowers than R. revelieri, 

and its first lowest two leaves are cordate, not lanceolate 16, 85 . 

R. ophioglossifolius, which is diploid (2n = 16) would have given 

rise to R. revelieri which is tetraploid (2n = 32). R. revelieri would 

thus be an apoendemic*, i.e. a polyploid* taxon no doubt derived, 

through evolution, from the diploid taxon R. ophioglossifolius 85 . 



Species endemic* to the Var (Maures and Estérel), Corsica and 

Sardinia, up to 1,000 m altitude. 

France 

Ranunculus revelieri rodiei in the Var (Maures, Estérel) and 

R. revelieri revelieri in Corsica. 


R. revelieri revelieri in Sardinia (Italy) 16, 85, 96, 292 . 

Habitat 

Plant species 


In Les Maures, Ranunculus r. rodiei is found in the shallow parts 

of pools, but also in ditches and on the edges of temporary 

rivulets. In the Plaine de Palayson, this species occurs in clay-silt 

pools situated in sandy depressions resulting from the erosion of 

Permian sandstones246 (R. r. rod.). 

In Corsica, the R. r. revelieri stations are mainly on the edges of 

16, 85, 108, 

temporary pools, as well as depressions and wet meadows 

109, 222, 285, 292 . 

Ranunculus revelieri 



1 cm 





• In the Var, Ranunculus revelieri occurs in an association* of the 

Isoetion histricis, the Lythrum borysthenicum and R. revelieri 

subsp. rodiei association 18 . This association may also be found 

under the old name “Peplis erecta (= Lythrum borysthenicum) 

and R. revelieri association” 18, 21, 215, 308 . 

71


• In Corsica, R. revelieri is frequently located among the following 

perennials: Schoenus nigricans, Plantago lanceolata var. timbali, 

Oenanthe globulosa, Cynodon dactylon and Carex serrulata. 

Associated annuals are: Juncus pygmaeus, J. tenageia, Solenopsis 

laurentia, Cicendia filiformis, Lythrum hyssopifolium, Illecebrum 

verticillatum (non-submerged form) and Agrostis pourretii. This 

type of community should not be included in the Isoetion histricis 

but in the Cicendion. 

Ecology 


Life form 

Therophyte* (annual species) with a scape* 96, 274, 292 . 

Reproduction 

Aerial. 

96, 386 

Seed characteristics 

Length: about 1.5 mm; width: about 1 mm. 


Seedlings may appear as soon as the first rain falls in autumn. 

They subsequently grow very quickly in April and flower in April- 

May. Flowering is advanced if rainfall comes early and delayed in 

the opposite case. The plant then spends the summer and part of 

the autumn in the form of seeds. The Lythrum borysthenicum and 

Ranunculus revelieri association is the most ephemeral among 

the Isoetion histricis and lasts for a maximum of two or three 

weeks: the community, submerged for the winter and a high proportion 

of the spring, sometimes disappears from the end of May, 

or more commonly in June with the first hot days 21, 96 . 

France 


72 

Germination Germination 


Flowering 

Fruiting 


Hydrology 

A hygrophytic* species found in shallow water (0-20 cm) but 

which can survive, in exceptional winter conditions, at depths of 

up to 40 cm. Germination takes place under water or on wet soil 

and the plant can tolerate prolonged immersion of the substrate 

for several months per year, from mid-autumn to the beginning 

of spring 153 . Climatic instability causes interannual fluctuations 

in the abundance of individuals. Ranunculus revelieri can also 

grow in slow-flowing streams and rivulets 18, 153 . 

Substrate 

It usually grows on siliceous soils. 


For this heliophilous* species, according to Barbero 21 , competition 

frequently causes population variations (instability). 

Experiments carried out within the context of the LIFE 

“Temporary Pools” project show that in the absence of grazing, 

competition with perennial herbaceous plants is very detrimental 

to this species, in particular on soils whose depth is greater 

than 20 cm. 


There are probably few direct effects on populations (uprooting 

of individuals), but a positive indirect effect of irregular disturbance 

of the substrate (stream dynamics, wild boar, etc) which 

restricts perennial species or improves the hydrological regime. 



In France 

• Several stations in the Var: Plaine des Maures (abundant, 

widely scattered), Massif des Maures, Plaine de Palayson (local), 

Colle du Rouet (very local). 

• Fifteen to twenty known sites in Corsica in 2001 285 : Agriate, 

Massif de Cagna (at 1,030 m altitude), communes of Pianottoli, 

Figari, Bonifacio and Porto-Vecchio 107, 108, 222, 225, 246, 274 and map in 

Paradis et al. 285 . 

In Italy 

Several stations in northern Sardinia, in the Limbara mountains, 

near Macomer, at Molara, and on San Pietro island west of 

Cagliari (map in Arrigoni 16 , Conti et al. 86 ). 


In Corsica, the populations appear to be maintaining themselves, 

but as with many therophytes, few individuals are visible in years 

when there is very little rainfall in winter and spring (as was the 

case in 2001-2002). 

In Provence, populations appear to be stable overall, despite the 

destruction of some stations. 


IUCN 1997 399 





and Red Lists 

- Ranunculus r. revelieri: 

“vulnerable” (V) for Corsica 

(France), and “endangered” (E) 

for Sardinia (Italy). 

- Ranunculus r. rodiei: 

“vulnerable” (V) for France 

- 

- 

- France, national list: Decree of 


13/05/1982, amended by Decree 


17/10/1995 




“threatened” 86



“Cleaning up”, drainage or infilling of the habitat, urban development 

96, 274 . In Provence, several stations have been destroyed 

directly or indirectly (disruption of hydrology) by tourism developments 

(golf), transport infrastructure works and urban development. 


Natural vegetational succession may lead to the habitat closing 

up and to processes of aggradation by siltation. For example, 

biotopes suitable for the Lythrum borysthenicum and Ranunculus 

revelieri association at the Péguière pool (Plaine des Maures) are 

progressively changing into drier habitats, which are prone to 

colonisation by other plant associations 153 , as a result of infilling 

with sediments from the catchment area. More generally in Les 

Maures, the disappearance of grazing favours the development 

of woody plants and perennial herbs (Juncus conglomeratus, 

Scirpus holoschoenus), to the detriment of R. revelieri. 

In Corsica, in the absence of disturbance (by cattle grazing, the 

activities of wild boar and of hares), there is a risk that the 

R. r. revelieri swards will in future be invaded by maquis species 

(Myrtus communis, Phillyrea angustifolia, Pistacia lentiscus). 


At Ranunculus r. rodiei stations, the populations are not visible 

very year as their development depends closely on the pattern of 

spring rainfall. Similarly, in Corsica, populations of R. r. revelieri 

are only visible in years when the winter-spring rainfall is sufficient 

to enable germination to take place (as was the case in 

2000-2001). However, their instability does not imperil the populations. 



Corsica 

The populations at the Tre-Padule de Suartone pools (Bonifacio) 

benefit from the site’s “Réserve Naturelle” (Nature Reserve) status 

(Ranunculus r. revelieri). 

Mainland France (Maures) 

•A Nature Reserve project at the centre of the plain should protect 

the majority of the stations on this site from anthropogenic 

Plant species 

threats (R. r. rodiei) 153, 274 . 

• The creation of a Natura 2000* area should enable the necessary 

requirements for the conservation of this species to be taken 

into account. Currently, the grazing which has been reintroduced 

along the firebreaks is favourable for it. 

• LIFE “Temporary Pools” has enabled experimental habitat management 

methods to be tested for this species, as well as the 

acquisition by the CEEP of two sites where it is present (Bois de 

Rouquan, Vallon de Sauronne). 

• Seeds have been collected and are preserved ex-situ by the 

Conservatoire Botanique National Méditerranéen de 

Porquerolles. 


In Corsica 

Managers should ensure that ground disturbance, due for example 

to extensive cattle grazing and the movements and “rooting” 

of wild boar (and hares), is continued. 


• Continuation of: 

- Station monitoring. 

- Raising the awareness of managers involved, in the context of 

taking the species into consideration during physical planning. 

- Control over management by acquisition or by contractual 

means. 

• Creation of protected areas wherever the species is present. 

• Re-establishment of extensive grazing. 

Bibliography 

Anonymous 1999 12 ; Arrigoni, 1983 16 ; Aubert & Loisel, 1971 18 ; 

Barbero, 1965 21 ; Bissardon & Guibal, 1997 39 ; Contandriopoulos, 

1962 85 ; Conti et al., 1992 86 ; Danton & Baffray, 1995 96 ; 

Deschâtres, 1993 108 ; Deschâtres et al., 1991 109 ; Gamisans & 

Jeanmonod, 1993 141 ; Gaudillat & Haury, 2002 153 ; Loisel, 

1976 215 ; Lorenzoni & Paradis, 2000 222 ; Lorenzoni, 1997 225 ; 

Médail et al., 1998 246 ; Olivier et al., 1995 274 ; Paradis et al., 

2002 285 ; Pignatti, 1982 292 ; Quézel et al., 1979 308 ; Tutin et al., 

1964-1993 386 ; Walter & Gillett, 1998 399 . 

Author: Calvière T. 

Collaborators: Catard A., P. Grillas, G. Paradis & N. Yavercovski 

73


Genus Riccia L. 

MARCHANTIOPSIDA 

RICCIACEAE 

Main synonyms 

None still in use nowadays. 

French name: Riccia 

English names: Crystalwort, Riccia 

Sous-genres 

Two sub-genera occur in the Mediterranean area: 

- Riccia with a single section, Riccia. 

- Ricciella (A. Braun) Rchb. with two sections: Ricciella and 

Spongodes Nees. 


• Small liverworts with gregarious thalli, forming either complete 

rosettes or more or less connected linear lobes, from a few millimetres 

to a few centimetres in diameter, with a tough, compact 

or spongy consistency. 

• May be annual or perennial* species. 

• Lobes usually having the dorsal surface longitudinally grooved 

and swollen at the edges into more or less prominent tumid 

ridges, varying widely in colour depending on the species and the 

growing conditions, from pure white to blackish-violet. Edges of 

the lobes sometimes with hairs, cilia or papillae and with scales 

which often have a crimson tint. 

• In species with a compact thallus (subgenus Riccia), dorsal tissue 

consisting of uniseriate columns of almost contiguous vertical 

cells; in species with lacunose thalli (subgenus Riciella) dorsal 

tissue very loose and with a network of air-filled gaps. 

• Dorsal epidermis pitted with more or less distinct pores. 

Ventral tissue more or less compact in all species in the genus. 

• Some species dioecious, others monoecious. 

• Male gametangia* (antheridia*) embedded in the dorsal tissue 

of the thallus, more or less projecting on the upper surface in the 

form of small upright protuberances. 

• Female gametangia (archegonia) also contained within the 

thallus and more or less protruding on the upper surface. Once 

fertilisation has taken place, capsule development within the 

thallus tissue; at maturity, the capsule may project from the 

upper or lower surface or not at all. 

• Spores released when the thallus decomposes at its base, 

although it may continue to grow at the tip. No elater* among 

the spores. Spores coloured at maturity, turning from brown to 

black and very often exhibiting polarity (the distal surface, which 

is roughly hemispherical, contrasting with the proximal surface 

which has three triangular facets). Distal surface (less often the 

proximal surface) usually provided with alveoli demarcated by 

walls which may have tubercles in their angles (this spore ornamentation 

is often very important for identifying species, hence 

it is always worthwhile to look for mature thalli in the field). 

- Vegetative* reproduction very rare. 

Introduction to a few species 

The genus Riccia includes over thirty species in the Mediterranean 

area, making it the most widely represented liverwort genus. The 

species are often fairly difficult to identify in the field, although 

some of them are easy to recognise with a little experience. They 

are not all strictly dependent on temporary pool biotopes but may 

74 

colonise any other habitats which are subject to a dry period/wet 

period alternation. 

Riccia sorocarpa, R. macrocarpa, R. glauca, R. sommieri, R. subbifurca, 

R. warnstorfii, R. nigrella, R. michelii, R. beyrichiana, 

R. crozalsii, R. crystallina and R. canaliculata for example are 

fairly typical of the surroundings of temporary pools. 

For species identification see among others the many works of 

Jovet-Ast 193 and of Schuster 349 . 



France 

The genus Riccia is particularly well represented in the 

Mediterranean Midi but many species have a wider distribution. 

Various species with Mediterranean affinities are found along 

the Atlantic seaboard (Riccia crozalsii, R. michelii, etc.) or in the 

Massif Central (R. trichocarpa). Many taxa are rare and only 

occur at a very limited number of stations. For example, the 

Massif des Maures, the Biot area and the Nature Reserve de Roque- 

Haute are sites that are extremely rich in liverworts of the genus 

Riccia. 


All the countries around the Mediterranean are rich in Riccia, 

although many of them have a very uneven distribution at this 

scale. For example, Riccia macrocarpa is much commoner in the 

west of the Mediterranean Basin than in the east; R. sommieri is 

only found in west of Mediterranean Basin; R. atromarginata is 

much less rarer in North Africa than in the north of the Mediterranean 

Basin, etc. 

Habitat 

Riccia sorocarpa 

1 mm 


Suitable habitats include, for example, short and pioneer grasslands 

with many gaps in the vegetation cover, thoroughly wetted 

in winter and spring and drying out completely in summer. The 

physical characteristics of the substrate are important. Riccia 

need a substrate which is firstly stable and relatively consolidated 

and secondly has a certain capacity to retain water. Sands, 

silts and clays (or mixed substrates) are the most suitable substrates 

in this respect. 

Biotopes with Riccia are often located close to temporary pools 

but not actually at the lowest topographical levels. Generally

speaking, it may be said that the Riccia of temporary pools and 

streams are linked with associations* of medium topographic 

levels. They are concentrated mainly among populations that are 

kept damp by capillary action or that emerge from the water 

early in the year, for example Isoetes duriei populations. 

Higher plant species that are usually associated are mostly small 

annuals (for example the dwarf Juncus, Radiola linoides, Laurentia 

michelii, Aira capillaris, Crassula vaillantii, Lythrum thymifolium, 

Lotus angustissimus, etc.). 


Communities rich in Riccia may on the whole be included in 

“Temporary Mediterranean Pools” (3170) or in “Very weakly mineralised 

oligotrophic waters of sandy plains in the Western Mediterranean 

with Isoetes “ (3120), in a very general way. It should 

be added that the phytosociological status of liverworts such as 

Riccia within vascular-plant communities is generally complex 

and cannot be simply expressed by means of these codes. 


These communities with Riccia may be placed in: 

22.34 “Southern amphibious communities” (Isoetalia), in particular, 



The relationships between liverworts and the vascular-plant 

communities are often complex. Liverworts and bryophytes* are 

in general very ephemeral and delicate plants whose populations 

(and thus the communities which they form) are subject to great 

variability in space and time. The communities are often patchy 

at the scale of a single station, where their appearance is frequently 

associated with openings in the herb layer. 

Some authors 21, 319 have considered the Riccia to be characteristic 

of the overall Isoetion alliance. Barbéro 21 and then Hébrard 171 

for their part emphasise the closeness of the sociological links 

which join the communities with liverworts to the vascular-plant 

communities Isoetion, but they also point out that it is very difficult 

to give an opinion on the true phytosociological affinities 

of the bryophytes. In addition, Hébrard 171 mentions cases of very 

clear extensions of bryological communities typical of temporarily 

wet biotopes into Cistus formations, when the soil humidity 

is sufficiently high. 

The Campylopetum introflexi Hébrard 171 is an association which 

unites the bryophytes of the rocky seepage surfaces of crystalline 

Provence, in which Riccia are considered to be companion species. 

The Riccietum crustatae is another circum-Mediterranean association, 

well represented in Spain, characteristic of wet, salty 

soils which dry out severely in summer 326 . However, most Riccia 

cannot tolerate the presence of salt in the substrate. 

It may be seen that the phytosociological characterisation of 

bryological communities involving Riccia is only in its early stages 

and that much descriptive work still needs to be carried out. 

Ecology 

Plant species 

Riccia are temporarily hygrophytic*, heliophilous*, thermophilic 

and terricolous species. However, the individual species’ ecological 

requirements show wide variations with respect to this general 

pattern. There are species for which the chemical 

characteristics of the substrate are immaterial while others are 

distinctly calcicolous or silicicolous. 


Life form 

Generally hepatico-therophytic. Annual or perennial species. The 

species considered to be annual may also revive after the dry 

season with the resumption of activity in the apical meristem or 

sometimes the whole thallus. It is possible to observe a resumption 

of activity of Riccia meristems that have been kept in a 

herbarium for over ten years, which is a good illustration of their 

powers of regeneration! 

Adaptive strategy (sensu During 126 ) 

•Type “a”, corresponding to “annual shuttle”: lifespan less than 

one year, spores large and produced in fairly small numbers; 

major reproductive investment. 

• More rarely type “s”, corresponding to “short lived shuttle 

species” (maximum of a few years): short-lived perennials with 

high investment in reproduction and few fairly large spores. 

Reproduction 

Fertilisation requires a film of water on the surface of the thalli 

or even the substrate; spore maturation takes place during the 

wet season and may continue into the beginning of the dry season. 

Spore production is therefore aerial. 

Spores 

They are usually fairly large (of the order 100 µm) and heavy, and 

are dispersed mostly through the movements of animals (birds, 

mammals, humans, etc.) but also by flowing water in some cases. 

Dispersal by anemochory* 349 appears to be less important and more 

unpredictable. In most cases the spores are ornamented and this 

could facilitate their attachment to vectors, although this hypothesis 

has not been experimentally confirmed. Fragments of thallus 

or propagules* (very rare) may also be carried by animals. 


Riccia are capable of carrying out the whole of their biological 

cycle very rapidly following the appearance of water in the substrate 

(rain, runoff) or merely on the thallus (condensation, nocturnal 

dewfall, etc.), enabling them to avoid the driest period. 

They are therefore able to tolerate the stresses of the driest 

months to a certain degree through the suppression of any 

physiological activity combined with an extreme longitudinal 

folding of the thallus, and by the production of spores that have 

very tough walls and that retain their germinative capacity for a 

very long time. 

The active phase begins with the germination of the spores, 

which takes place as soon as the first autumn rain starts to fall. 

The growth of the thalli continues throughout the whole of the 

wet season. Gamete production may take place very soon after 

germination (a few weeks for the fastest species), but may also 

continue until the beginning of the dry season of the following 

75


year. The spores are released by the decomposition of the old 

parts of the thalli as growth continues at the tips of the lobes. 

The complete desiccation of the substrate and the thalli causes 

the virtually complete disappearance of Riccia and the release of 

all the spores produced; the latent phase then begins, with the 

survival in the substrate of the spores, which may enter into a 

dormant state. 



Fertilisation and maturation of spores 

Release of spores 


Hydrology 

A very wet substrate is necessary at the beginning of the cycle 

but, in most species, flooding must not be too prolonged. 

Substrate 

More or less compacted substrates are definitely the most suitable, 

whether they are sandy, clayey or silty. Terra Rossa*, more or less 

muddy alluvia, etc., also provide potential sites. 


All the species are pioneers and struggle to survive in the presence 

of competition from other more colonist bryophytes and especially 

perennial vascular plants, which will eliminate them fairly 

readily. 


The main perturbations (“ploughing” of the substrate by wild 

boar, trampling by cattle and humans) may have a positive effect, 

due to the elimination of taller plants, the destruction of the litter 

from these plants and the opening of the herb layer through 

the creation of bare patches (“tonsures”) which are very suitable 

for Riccia. 

Temperature 

All the species are thermophiles. 

Light 

All the species are heliophiles, but very slight shading which 

moderates the intensity of the solar radiation may be a positive 

factor in the hottest regions of the Mediterranean Basin. 



Practically all temporary pools and streams in France are colonised 

by Riccia. Some rare species are, however, very local. 

The temporary pools and streams of the countries surrounding 

the Mediterranean are equally rich in Riccia. The number of sites 

is impossible to estimate but is probably very high. Spain, Algeria 

and Morocco are particularly rich in liverworts of this genus. 

76 

Latent spores and thalli 


The conservation status of Riccia populations is intimately linked 

with the general health of the temporarily wet biotopes. These 

liverworts have undergone a major decline following the disappearance 

of many temporary pool biotopes. 

In Malta, R. melitensis was known from one locality, where it has 

never been seen again since its discovery 344 . 


IUCN 1997 





and Red Lists 

- 

- 

- 

- 

- Europe: Red Data Book for 

bryophytes 128 : 

Riccia crustata: 

“endangered” (E) 

Riccia frostii: 

“rare” (R) 

R. huebeneriana: 


R. ligula: 


R. melitensis: 

“extinct” (K) 

R. perennis: 

“rare”(R) 

R. sommieri: 


R. trabutiana: 


- France, Red Data Book for 

bryoflora (proposed) 105 : 

Riccia crustata 

R. ligula 

Further species in this genus are locally rare to very rare, but are 

not included on the protected lists as they are little known. 



All the threats with which temporary pool and stream biotopes 

are faced in the area around the Mediterranean consequently 

also threaten populations of liverworts of the genus Riccia. 

Urbanisation, infilling with rubble, drainage and conversion to 

agriculture, etc. lead to the irreversible loss of “Ricciological” 

natural heritage. 


The absence of any factors that will disturb the vegetation, and 

the closing-up of the natural habitat, will eventually cause the 

disappearance of pioneer liverworts. The loss of the temporary 

character of pools, through infilling or through disruption of the 

hydrological network, is very deleterious to Riccia as this allows 

the spread of common and much more competitive species of 

hygrophytic bryophytes (usually pleurocarp* species). However, 

the presence of dormant spores in the substrate is a positive factor 

in the re-establishment of populations following a reopening 

of the habitat.

Extinction risks relating to populations 

Major interchanges probably take place between neighbouring or 

widely separated sites, but there are no experimental results to 

support this assumption. 



The sites at Roque-Haute and the Massif and Plaine des Maures 

are included within Natura 2000* sites (no. Fr 9101430 and 

no. Fr 9301622 respectively) which should allow particular 

attention to be paid to the conservation of the temporary pools 

and their associated species. 

The Roque-Haute site also has legally protected “Réserve Naturelle 

(Nature Reserve)” status, which is currently unenforced due 

to the ban on access imposed on the manager by the owners. 


• Promote a better knowledge of the distribution of the species 

(surveys and inventories) and raise the awareness of managers. 

• Maintain the hydrological network in an intact condition 

(hydrological regime and water quality). 

Plant species 

• Promote the perpetuation of extensive agricultural practices 

which create moderate disturbance, or management activities 

having the same rationale. Extensive grazing may be favourable 

at stations where the vascular vegetation has developed too 

much. Inputs of excess organic matter should, however, be monitored. 

Bibliography 

Anonymous, 1999 12 ; Barbero, 1965 21 ; Bissardon & Guibal, 1997 39 ; 

Deperiers-Robbe, 2000 105 ; Dierssen, 2001 118 ; During, 1979 126 ; 

ECCB, 1995 128 ; Hébrard, 1970 171 ; Jovet-Ast, 1986 193 ; Rivas-Goday, 

1970 319 ; Ros & Guerra, 1987 326 ; Schembri & Sultana, 1989 344 ; 

Schuster, 1992 349 . 

Author: Hugonnot V. 

Collaborator: Hébrard J. P. 

77


Riella helicophylla (Bory & Mont.) Mont. 

MARCHANTIOPSIDA 

RIELLACEAE 

Main synonyms 

Duriaea helicophylla Bory & Mont. (basionyme*) 

French name: Riella à thalle hélicoïde 

English names: Liverwort, Riella (genus names) 

Subspecies 

None 


• Aromatic aquatic liverwort (scent fairly like coriander, sometimes 

strong), measuring up to 3 cm high but often much smaller, 

annual, soft-tissued, formed from an upright thallus composed of 

a main branch with a more or less undulate single-layer membranous 

wing (a single layer of cells) on one side and small lanceolate 

scales on the other. Wing more or less helicoid, very fragile, 

and usually reduced to barely recognisable fragments due to various 

disruptions and to predation by phytophagous invertebrates. 

• Thallus fixed into the substrate by many rhizoids* growing 

from the base (plant appearing to be placed gently on the substrate). 

• Dioecious species whose male plants bear very small antheridia*, 

arranged in a row on the edge of the membranous wing of 

the thallus. 

• Female plants at maturity bearing bottle-shaped capsules* at 

the tip of the thallus, pedicellate but lacking hairs, indehiscent, 

protected by a papillose, membranous involucre without a wing 

(characteristic of the subgenus Riella). 

• Spores, released when the wall of the capsule decomposes, 

measuring 70 to 95 µm in diameter, with surface smooth to 

slightly granulous, bristling with many large “spines” (6 to 10 µm 

long) which are truncated and widen at the tip. No elaters* 

among the spores. 

The sporophyte (capsule), which is normally attached to the 

gametophyte*, may become quite independent of it (the thallus), 

when unfavourable conditions (drought, predation, etc.) result in 

its premature disappearance. The sporophyte then continues to 

develop in a completely independent way, which is very unusual, 

if not unique, among bryophytes*. 

The formation of single-layered propagules with indeterminate 

growth, arising from the apical meristem, has been described. 

These propagules, in contrast to spores, do not have a dormant 

stage and are therefore capable of growing immediately after 

their release. 


Confusion is possible in the field with all the other species in the 

genus. Identification can only be carried out by a thorough examination 

using appropriate optical equipment. Sterile material is 

practically impossible to identify to species. Mature sporophytes 

must therefore be looked for in the field amidst the population. 

In addition, mixtures of species are relatively common. 

78 

spore 

The species of the subgenus Trabutiella (Riella affinis M. Howe & 

Underw. and R. cossoniana Trab.) are fairly easily distinguished 

from R. helicophylla by their longitudinally winged involucre. 

R. notarisii (Mont.) Mont. is monoecious (often a difficult character 

to assess) and its spores have a rough surface and shorter spines 

(4 to 5 µm). 

R. parisii Gottsche has much smaller spores (about 60 µm), with 

the spines shorter (5 to 6 µm) and not widening at the tip. 

R. numidica Trab. also has small spores of about 60 to 70 µm. 

R. bialata Trab. has the wing divided into two. 

The species of the genus Riella are relatively poorly known from 

the taxonomic point of view. Great confusion reigns in the 

nomenclature regarding various taxa*. A complete and worldwide 

review of the genus appears to be essential to clarify the 

many points which remain uncertain. 



20 µm 

male 

Riella helicophylla 

0.4 cm 

female 

France 

Species endemic* to the Mediterranean Basin. 

Extremely rare species in France, only in the Hérault, known from 

two localities, close together: 

• The Notre-Dame de l’Agenouillade site (Agde), discovered 

recently in 2001 353 ; it is currently still present at this site.

• Station by the Etang de Thau (Marseillan), discovered in 

1966 124 , and very probably disappeared following the siting of a 

huge refuse tip on the exact site. 


Species occurs in the Iberian Peninsula (Spain and Portugal) and 

the Balearics, Malta 206 , quite common in the Maghreb (Algeria, 

Tunisia), and present in the Near East (Israel). 

Habitat 


The ecological characteristics necessary for the development of 

Riella helicophylla populations are: 

•A temporary water body (prolonged dry period). 

• Clear, relatively shallow water. 

• Soil that is bare or with sparse plant cover. 

The most favourable habitats corresponding to these criteria are 

therefore of two types: 

•Temporary pools, lakes, estuaries etc. with saline to supersaline 

or brackish water (France, Spain, North Africa). 

•Possibly, temporary freshwater pools (France). 

The muddy, sandy or clayey substrate is often base*-rich (pH* ranging 

from about 7 to 8.5). 

The depth of the water seems to be very important: often very 

shallow (a few centimetres) and a maximum of 1 metre. 

In Spain, Riella helicophylla has been collected from sites with a 

wide range of salinity and chemical composition. The salinity 

(total dissolved solids) varies from about two to almost 190 g/l, 

though the species grows best in water with values typically 

greater than 10g/l 77, 78, 189 . Continuous evaporation at some sites 

leads to the formation of saline crusts on the surface. R. helicophylla 

appears particularly to prefer waters containing sodium 

chloride although it also colonises those with magnesium sulphate. 

The waters in which it grows are subject to considerable 

variations in salinity over a single year and also from one year to 

another, depending on rainfall, runoff, evaporation, etc. 


The species most frequently associated with Riella helicophylla 

are other species of the genus Riella (in particular R. cossoniana 

and R. notarisii), aquatic vascular plants (Ruppia spp., Althenia 

spp., Potamogeton spp.,) Characeae (Chara galioides, C. canescens, 

etc., Tolypella spp., Lamprothamnium spp.), rarely bryophytes 

(Leptodictyum riparium, Drepanocladus aduncus, etc.), Green 

Algae (Ulothrix spp., Enteromorpha spp., etc.), Cyanophyceae 

(Microcoleus, Lyngbya, etc.) and sulphur bacteria. When the substrate 

becomes completely dried out, the vegetation then changes 

its appearance, with the frequent arrival of a suite of more or less 

xerophytic vascular and bryophyte species. Note that Riella only 

really disappears when the substrate is completely dry. 


The simplest approach is to include the Riella community under 

code 3170, with the overall title “Mediterranean Temporary 

Pools” for the freshwater communities. On the other hand, the 

subhalophilic and halophilic* communities could be included 

under code 1150, with the title “Lagoons”, or even code 1160, 

Plant species 

“Large shallow inlets and bays”. Insofar as these codes did not 

originally take the associated bryophyte communities into 

account, the affinities with one or other suggested code should 

be explained in detail, so as to avoid any ambiguity or confusion. 


The most appropriate CORINE codes appear to be: 

• For freshwater communities: 

22.34 Mediterranean amphibious communities. 

• For halophilic vegetation types: 

11.4 Vegetation beds of brackish waters. 


The Riella helicophylla communities are these days included 

within the class Rielletea helicophyllae, order Rielletalia helicophyllae, 

alliance Riellion helicophyllae, association* Rielletum 

helicophyllae 78 . These vegetation types represent a transitory 

cover of species that are highly specialised for the colonisation 

of a very restrictive temporary biotope, at the stage where other 

communities, dominated by various groups of species (Charetea, 

Potametea, Ruppietea, Isoeto-Nanojuncetea, etc.), are absent or 

at an embryonic stage of development. 

Ecology 

It should be borne in mind that Riella helicophylla is a basiphilous*, 

halotolerant* species, capable of withstanding hypersaline 

water; a heliophilous pioneer. 


Life form 

Hepatico-hydrotherophyte. Annual species (capable of living for 

more than one year in culture if it is permanently submerged). 

Adaptive strategy (sensu During 126 ) 

Type “a”, i.e. “annual shuttle” species (= lifespan less than one 

year, spores large and produced in fairly small quantities; major 

reproductive investment). 

Reproduction 

Maturation of spores aquatic then aerial; release aerial. Spores 

exhibiting dormancy and capable of retaining their germinative 

capacity for at least three months. 

Germination 

It is at its maximum with low salt concentrations, whereas the 

thallus grows better at higher salinities. 

Dispersal of spores 

Dispersal by water birds (waders, web-footed species), sometimes 

migratory, by external or internal ornithochory*. This mode of 

dispersal is assisted by the very large number of spines on the 

spores. anemochory* appears never to have been described. 


Germination of the spores takes place after the first autumn rain, 

followed by the slow growth of the gametophyte during winter 

and the beginning of spring. The spores are released in about 

May and constitute the resistive stage during the summer 

79


drought. When the ecological conditions are favourable, it is possible 

to see veritable population explosions of this species, which 

then excludes other species and attains coverage rates of close 

to 100%. 


� 


80 

Fertilisation and maturation of spores 


Release of spores 

Spores dormant 

� 


Hydrology 

• The optimal water depth ranges between a few centimetres 

and a few tens of centimetres. 

• The survival of the species is closely dependent on the annual 

flooding/drying regime. A disruption of the hydrological regime, 

a delay in the arrival of the rains or too low a level of annual 

rainfall may considerably restrict or even prevent the development 

of the species one or several years in succession (temporary 

disappearances). 

•Water quality is important, especially salinity. Riella helicophylla 

appears to be not at all obligatorily halotolerant* but, 

being a very uncompetitive species, it finds in salt water the conditions 

which allow it to exist without or almost without any 

competition. 

• The species can tolerate a certain degree of eutrophication* of 

the substrate and of the water. 


Perturbations due to the weather (rainstorms, frosts, etc.), anthropogenic 

disruption (trampling, ploughing, etc.), or zoogenic disturbances 

(grazing, disturbance of the substrate, etc.) may be 

favourable to the extent that they restore pioneer surfaces, provided 

they take place during the dry period (i.e. after the maturation 

of the spores). 


Natural succession inevitably leads to the disappearance of pioneer 

Riella helicophylla communities under increasing pressure 

from vascular plants and algae, whose increasing biomass* and 

ground cover result in greater competition for light and other 

essential resources. R. helicophylla does not survive in competition 

with other plants. 



A single site in France; many sites in Spain, distributed among 13 

provinces 79 ; several sites in North Africa. 

Populations appear to be stable or even increasing in Spain and 

the Maghreb, but are highly threatened in France due to the 

pressure of urban development in the Agde area. 


IUCN 1997 





and Red Lists 



Urban development around towns such as Agde, Tunis or Algiers 

constitutes a key threat. The filling in of pools with debris needs 

to be monitored, particularly at the Notre-Dame de l’Agenouillade 

site (Hérault, France). The illegal dumping of rubbish and largescale 

official tips are leading to the irreversible disappearance of 

the species. 


Natural vegetational succession leads inexorably to the spontaneous 

disappearance of pioneer taxa* such as Riella helicophylla, 

by increased competition within the vegetation mat. In addition, 

tall plants on the edges of pools (Tamarix sp., Phyllirea sp., etc.) 

are unfavourable for visiting birds (especially Anatidae) which 

are capable of carrying spores from one site to another. This 

reduction in the “accessibility” of some sites is clearly detrimental 

to the survival of viable metapopulations*. In addition, a high 

frequency of dry years may explain the species’ decrease or even 

extinction at some of these stations. A minimum of wetness suffices 

to trigger the germination of dormant spores but a sufficient 

amount of water is absolutely necessary to complete the 

cycle and to produce spores for a new generation. Several dry 

years may completely deplete a fairly small stock of spores. 


The risk of extinction is high, given the low numbers at the 

Notre-Dame de l’Agenouillade station (Hérault, France). The isolation 

of the French populations compared with those of North 

Africa and Spain, and their limited area, are a further risk factor. 


- 

Strictly protected species 

(Annexe I) 

Annexe II 

Malta: 

protected species since 1993 206 

- Europe, Red Data Book for 

Bryophytes 128 

“endangered” (E) 

- France, national Red Data Book 

(proposed) 105 


No specific management measures appear to be taking place at 

sites supporting Riella helicophylla. 

In France, the Notre-Dame de l’Agenouillade site (Hérault), is 

included: 

• In the European LIFE “Temporary Pools” project, which has 

enabled proceedings for obtaining control over land ownership to 

be undertaken, hydrological monitoring to be carried out and 

protocols for monitoring the pools’ vegetation to be drawn up. 

• In a Natura 2000* area (site Fr 9101416), which should enable 

the environmental conditions favourable for the conservation of 

this species to be maintained.


•A major survey exercise should be organised in order to locate 

any possible new population centres, especially in France. 

• Any management measures which have the aim of maintaining 

or restoring the pioneer character of the pools and their surroundings, 

such as controlled grazing or vegetation clearing, are 

to be encouraged, especially at sites where these activities have 

been discontinued. 

• Complete protection of the biotopes (control over land ownership, 

etc.) should be achieved in order to prevent any destructive 

urban development projects. 

• Dumping of rubbish, infilling, and illegal tourism-related 

activities should be strictly forbidden. 

Bibliography 

Plant species 

Aboucaya et al., 2002 2 ; Anonymous, 1999 12 ; Bissardon & Guibal, 

1997 39 ; Cirujano et al., 1988 77 ; Cirujano et al., 1992 79 ; Cirujano 

et al., 1993 78 ; Deperiers-Robbe, 2000 105 ; Dierssen, 2001 118 ; Dubois 

& Hébant, 1968 124 ; During, 1979 126 ; ECCB, 1995 128 ; Jelenc, 

1955 189 , 1957 190 ; Lanfranco & Lanfranco, 1999 206 ; Proctor, 1961 301 ; 

Skrzypczak, 2001 353 ; Trabut, 1891 383 ; Trabut, 1942 384 . 

Author: Hugonnot V. 

Collaborator: Hébrard J. P. 

81


Teucrium aristatum Perez Lara 

ANGIOSPERMS 

LAMIACEAE 

Main synonyms 

T. cravense Molinier & Tallon 

French names: Germandrée de Crau, Germandrée aristée 

English name: Germander (genus name) 

Subspecies 

None 


• Small annual plant 10 to 20 cm, stem slender taproot. 

• Stem square in cross-section, upright, usually branching 

almost from the base; branches erect-spreading. 

• Leaves pinnatilobed*, with lobes broad, short and blunt, shed 

from the time of flowering. 

• Flowers geminate, with short peduncles. Calyx lobes erect, 

spreading, terminated by a beard. Corolla villous externally, lavender 

blue with a dark-blue V-shaped mark on the inner base of the 

middle anterior lobe; posterior lobes streaked with crimson internally. 

• Fruits: each flower produces four achenes after fertilisation. 


Teucrium campanulatum L. is a species of wet clay soils and 

flood-prone depressions in North Africa, Spain and southern Italy: 

it is a perennial* plant with the stems often radicant (and not an 

annual with upright stems), with a shorter and more flared calyx 

than in T. aristatum, and with whitish, not lavender blue, flowers. 



Endemic species of Iberia and Provence. 

France 

Crau d’Arles in the Bouches-du-Rhône. 


Only in Spain, where it was formerly known (more than 100 years 

ago) in three provinces in the southwest (Huelva, Cadiz and 

Seville) 177 (Medina, pers. com.), and where it was rediscovered in 

2002 in the centre of the country (Morales, pers. com.). 

Habitat 


In France, this Germander occurs on the edges of a clayey basin 

created in the Crau pudding stone, whose water is slightly calcareous 

252 . 



82 


22.34 Southern amphibious communities (Isoetalia) 


In the Crau the vegetation of the Lanau pool is allied to a specific 

association* 215, 252 : 

Order Nanocyperetalia 

Alliance Lythrion tribracteati 

Association: Lythrum tribracteatum-Teucrium arista- 

215, 252 tum 

At this site, the species also forms part of the Deschampsion 

mediae alliance. 


Species associated with Teucrium aristatum in France are: 

Lythrum tribracteatum, Mentha pulegium, Achillea ageratum, 

Deschampsia media, Herniaria glabra, Polygonum aviculare subsp. 

depressum, Damasonium polyspermum, Lythrum thymifolium. 

Ecology 

1 cm 


Life form 

Therophyte* (annual species). 

Teucrium aristatum

Reproduction 

Aerial: the floral organs as well as the fruits develop out of water. 

Seeds 

Achenes oboval-obovate (2 mm x 1.1 mm), glandular, rounded, 

and with dense very short curved hairs at the tips. 


Germination of seeds after the water has receded, on very wet 

soil (in May in France). Flowering occurs at the end of spring 

(June) on a dry substrate, followed by the ripening of the fruits 

and the dispersal of seeds during summer. 

France 




Flowering 

Fruiting and ripening of fruits 

? ? 

Dispersal of seeds 

Hydrology 

This Germander is above all a species of the edges of pools, which 

needs a brief period of submersion to develop. It is probable that 

the saturation of the substrate with water at a given period (end 

of April or May) triggers germination. 

Substrate 

In France, silt-clay soils, poor in calcium and oligotrophic*. 


Very sensitive to competition with perennial herbaceous plants. 


Disturbance (grazing, trampling) is favourable to this heliophilous* 

species since it controls the cover from perennial grasses. At the 

higher parts of the Lanau pool, the Germander is most often seen 

on patches of bare soil: roosting places, cattle tracks, holes made 

by hoofs, and former ditches, where it forms small dense populations. 

Conservation-Management 


France 

A single site is known, Lanau pool in the Crau (Bouches-du- 

Rhône). Although it undergoes wide interannual fluctuations, 

this population has been seen regularly since its discovery in 

1946, and is not in danger at present. The temporary reduction 

of grazing at this site seems to have noticeably reduced the size 

of the visible population, which is in the process of re-establishing 

itself. 

Plant species 

Spain 

The species is very rare here, with a very limited range (in the 

provinces of Cadiz, Seville and Huelva), but a few new stations 

were discovered in 2002 in the centre of the country. 


IUCN 1997 





and Red Lists 



The acquisition by the CEEP of the site supporting the only 

French population has enabled the pool to be saved from possible 

threats of habitat destruction, and management favourable 

to this species to be introduced. 


This species is very sensitive to competition with perennial species 

that produce dense cover (Carex divisa, Agrostis stolonifera, etc.). 

Towards the end of the 1990s, it had disappeared from a part of 

the Lanau pool where the cessation of grazing had facilitated a 

massive growth of perennial grasses, whereas it always maintained 

itself in the grazed part of the pool. 


The unique nature of the population (very isolated) entails a high 

potential risk of extinction at the site. 



World: “rare” (R) 

- France: “endangered” (E) 

- Spain: “rare” (R) 

- 

- 



13/05/1982, amended by 

Decree of 31/08/1995/ 




France 

• The CEEP became owner and manager of the site in 1998, 

thanks to financial support from the “Réseau Ferré de France”, in 

the context of compensatory measures linked with the routing of 

the LGV Méditerranée (High Speed Rail Link). 

• The Conservatoire Botanique National Méditerranéen de 

Porquerolles carries out regular monitoring of the Germander 

populations, and of the other rare species at the site. 

• Seeds have been collected and preserved ex situ by the 

Conservatoire Botanique. 

•A grazing management experiment, carried out by the Station 

Biologique de la Tour du Valat, has been running since 2001. The 

initial results show a substantial spread of Teucrium aristatum in 

the grazed areas in 2002 and 2003 (see Vol. 1, box 49). 

83


Spain 

The species and the sites which support it are not subject to any 

specific measures for protection or management (Medina, pers. 

com.) 


France 

• Study the hydrological regime of the site and prevent any disruption 

of it. 

• Maintain grazing (formerly sheep, nowadays cattle) which is 

favourable to the development of annual species like Teucrium 

aristatum. 

• Study the population biology of T. aristatum, in particular the 

key spring recruitment phase (ecology of germination) and the 

effects of natural and anthropogenic perturbations. 

84 

Bibliography 

Devesa, 1987 110 ; Loisel, 1976 215 ; Maire et al., 1947 232, 233 ; 

Molinier & Tallon, 1947 252 ; Molinier & Tallon, 1950-1951 255 ; 

Olivier et al., 1995 274 ; Tutin & Wood, 1972 385 ; Vivant, 1980 397 ; 

Walter & Gilett, 1998 399 . 


Collaborators: Grillas P. & N. Yavercovski

Imnadia yeyetta Hertzog, 1935 

Class: BRANCHIOPODS 

Order: SPINICAUDATA 

Family: LIMNADIIDAE 

Main synonyms 

I. banatic Marincek & Petrov, 1982 

I. cristata Marincek, 1972 

I. panonica Marincek & Petrov, 1984 

I. voitestii Botnariuc & Orghidan, 1941 43 


English name: Clam Shrimp (general name for Spinicaudata) 

Subspecies/variability 

No valid subspecies. Intraspecific variability more or less marked 43 . 

In Yugoslavia, Marincek 239 and Marincek & Petrov 237 have included 

several species as being synonymous with I. yeyetta. 


The Spinicaudata have a bivalve carapace that is generally striated, 

laterally compressed and protecting the body. But unlike 

numerous Spinicaudata such as Cyzicus, Leptestheria, etc., the 

bivalve carapace of Imnadia yeyetta is not striated over its entire 

surface: the upper part of the valves is smooth, around the umbo, 

while their outer edges are finely striated. 

Sexual dimorphism is not very marked: the males have 17 pairs 

of legs whereas the females have only 14; the males have legs P 1 

and P 2 prehensile, armed with “claspers”, enabling them to clasp 

the female by the edge of the carapace during mating; the 

females have the pairs P 9 and P 10 modified into adhesive cylinder-like 

structures to agglutinate the eggs between their body 

and their carapace. The carapace, is yellowish to greenish white 

in colour, slightly transparent, and measures a maximum of 11 to 

12 mm. 

The eyes are sessile. 


The only other species of Spinicaudata possessing a carapace 

which is partially smooth around the umbo is a species of the 

Limnadiidae family, Limnadia lenticularis (L., 1761), found in the 

Ile-de-France and eastern France. This species is Palaearctic: its 

range extends over central and northern Europe 52 . L. lenticularis 

is also present in Italy. 



I. yeyetta is a species mainly present in central Europe (Austria, 

Slovakia, Hungary, Serbia and Romania). In the Mediterranean 

region it is found in northern Greece and in the former Yugoslavia 

52 ; in France, it is only found in the Bouches-du-Rhône. 

Habitat 



I. yeyetta colonises usually turbid temporary pools, in the 

Camargue Delta and the Crau. Its presence reflects the dispersal 

into biotopes of resting eggs carried by migratory birds flying 

between central and southern Europe, such as the Anatidae, 

which are particularly abundant in the region. 




Lanau pool (Crau) and temporary pools in the Camargue: 

22.32 Annual Mediterranean amphibious swards (Lythrion tribracteati). 

Ecology 


Imnadia yeyetta 

2 mm 

male 

egg (150-190 µm) 

Feeding 

Microphagous* species feeding by filtering water and sediments. 

Detrivorous* feeding regime. 

Behaviour 

Usually benthic* species, living on the sediment or partly buried 

in superficial sediments. 

Description of eggs 

Eggs ovoid, 150 to 190 µm in size, characterised by their spiralled 

surface. Confusion is possible with the gyrogonites* (résistant 

forms) of charophytes, which are distinguished by the presence 

of five spiral-shaped apical cells 355 . 

Site and mechanisms of egg laying 

The eggs, grouped together on the abdomen of the female, are 

discharged into the water during each moult. Their distribution 

on sediments depends on the position of the female at the time 

of the moult, the influence of vegetation and areas of open water 

as well as the wind, in particular the mistral (northerly wind), which 

creates movements in shallow water through convection cells. 

The distribution of the eggs can thus be very variable depending 

on the biotope. 

85 

Modified from Defaye et al. 102


Longevity 

Longevity is estimated at less than eight weeks when conditions 

are favourable (no premature drying out). However, as for all 

branchiopods, it is very variable depending on the temperature. 


I. yeyetta is most often present in the spring but can also be 

found in autumn (November-December). A complete cycle was 

studied in the Camargue by Nourisson & Aguesse 270 . The females 

can lay up to 400 eggs at a time (the size of the clutch depends 

on the age of the female). This occurs shortly after mating. The 

chitinous* carapace with two lobes appears rapidly, generally at 

the metanauplius* stage. This first, or “heilophore”, bivalve stage 

occurs three or four moults after the nauplius* stage. The morphology 

of the larval stages has been studied by Eder 129 . Unlike 

most crustaceans, and in particular the branchiopods, the carapace 

is not discarded totally during successive moults, which 

leads to the appearance of growth lines 374 . 


Interspecific relationships 

I. yeyetta can be found with the anostracans Tanymastix stagnalis, 

Branchipus schaefferi, Chirocephalus diaphanus and/or the 

notostracan Triops cancriformis. For more data on the ecological 

conditions of coexistence of several species of branchiopods in a 

temporary pool in a Mediterranean climate, see Thiéry 373 . 

Requirements for reproduction 

No data. 

Environmental perturbations 

Natural perturbations corresponding to the constraints linked to 

habitats with fluctuating water levels. 

Main natural mortality factors 

Not well known. However, as for most branchiopods, the individuals 

develop in a cohort from the onset of flooding 371 , which limits 

coexistence with predators (larvae of Coleoptera and notonectid 

Heteroptera, for example) which only colonise the habitats several 

weeks after inundation (see vol. 1, chapter 3e). Note, however, 

that there could be a possibility of predation, not yet quantified, 

by Rhabdocoela (Platyhelminthes), which are common in the temporary 

stillwater habitats of Provence (Thiéry, original data). The 

impact of birds is poorly known. 

86 


� 

Adult stages (clutches) 

Egg bank 

� 

Hatching and growth 

of larvae 



Number of sites 

The species is only present in France in the Camargue Delta and 

the Plaine de la Crau (Lanau pool). In central Europe, I. yeyetta 

has a fairly wide distribution 52 . 

Populations 

The densities of the populations are quite low, with an irregularity 

in the hatching periods and thus of appearance. 


IUCN 






No data are available relating to the damage thresholds linked to 

anthropogenic actions, nor are any toxicological data available. 

Recent observations (Thiéry, original data) are inconclusive. 


No data. 


In the Lanau pool, the population appears to be fairly stable, given 

the state of the cyst bank 355 ; no data on the state of the populations 

in the central Camargue. 



At two known sites, situated in Natura 2000* areas, the populations 

appear to be stable, and their biotopes safe from any 

human pressure. The Lanau pool was acquired by the CEEP in 

1998 and the temporary pools where this species lives in the 

Camargue benefit from the status of “Réserve Naturelle” (Nature 

Reserve) (Tour du Valat estate). 


Given the current state of knowledge, no conservation measure 

can be proposed. Only the maintenance of the biotopes in their 

current state is recommended. 

Bibliography 

AGRN-RH, 2000 3 ; Anonymous, 1999 12 ; Botnariuc, 1947 43 ; Brtek 

& Thiéry, 1995 52 ; Defaye et al., 1998 102 ; Eder, 2002 129 ; Hertzog, 

1935 175 ; Marincek & Petrov, 1984 237 ; Marincek, 1972 239 ; 

Nourisson & Thiéry 1988 269 ; Nourisson & Aguesse, 1961 270 ; 

Soulié-Märsche & Thiéry, 1998 355 ; Thiéry & Gasc, 1991 365 ; Thiéry 

& Pont, 1987 366 ; Thiéry, 1987 371 , 1991 373 , 1996 374 ; Thiéry et al., 

1995 375 . 

Authors: Thiéry A. & T. Calvière 

- 

- 

- 

- 

-

Linderiella massaliensis Thiéry & Champeau, 1988 364 


Order: ANOSTRACA 

Family: LINDERIELLIDAE 

Main synonyms 

None 

French name: None 


No subspecies. 

Very low morphological variability; low genetic variability (Thiéry, 

original data). 


Anostracans have an elongated body without a carapace and 

swim on their backs in open water. They possess two cercopods* 

(furca) at the end of the abdomen. 

Linderiella possesses 11 pairs of phyllopods* and two cercopods, 

orange in colour, borne on the last abdominal segment (telson). 

The males are distinguished from the females by the morphology 

of the antennae: in the males, they are much more developed, 

biarticulate, and their pincer shape enables them to clasp the 

female during mating. The males possess two spiny ventral hemipenises 

at the base of the abdomen (genital apparatus). The 

females, with shorter antennae, have, on maturity, a heartshaped 

brood pouch*, (bicoloured blue/brown) at the base of the 

abdomen, in which the eggs acquire their outer envelope. 

The individuals, creamy-greenish in colour and transparent, measure 

a maximum of between 10 and 13 mm. The females are 

slightly larger than the males (morphological dimorphism). 


For the non-specialist, all anostracans are very similarly morphologically: 

they can only be differentiated with an identification 

key 102, 269 . Linderiella massaliensis resembles Linderiella sp. of 

Spain as well as L. africana which inhabits the temporary pools 

of the Middle Atlas in Morocco. 



France 

Species endemic to southeastern France (Var). 

Biogeography 

Linderiella massaliensis is a vicariant* species of the Moroccan 

species L. africana Thiéry 1986 370 which inhabits several dayas 

(temporary pools) of the Middle Atlas on the Azrou Plateau. Three 

other vicariant species are so far known: two in California, L. occidentalis 

Dodds, 1923 and L. santarosae Thiéry & Fugate, 1994 367 , 

and one in Spain Linderiella sp. 7 . These species together constitute 

a case of adaptive radiation through fragmentation of the 

distribution range of a “mother” species at the time of the opening 

of the Atlantic and the Thetys. 

Habitat 



Linderiella massaliensis is found in temporary pools which are 

flooded in autumn and winter, such as Lake Redon. Generally 

speaking, L. massaliensis occupies pools of karstic origin (poljés*, 

dolines), of varying depths. The waters are of low turbidity, poorly 

mineralised (temperature between 5 and 20°C, pH between 7.2 

and 8.4, C 20 between 250 and 600 µS.cm -1 ) and rich in microcrustaceans 

(more than 25 species of cladocerans and copepods 

at Bonne Cougne). 




22.34 Southern amphibious communities (Preslion cervinae) 

22.32 Annual Mediterranean amphibious swards (Lythrion tribracteati) 

Ecology 

2 mm 

egg (240-260 µm) 


Linderiella massaliensis 

male 

Feeding 

Linderiella massaliensis, like all anostracans, is microphagous* 

and feeds by filtering suspended food particles from the water 

(microplankton, micro-organisms and organic material), using its 

legs which are covered in bristles. Anostracans can also disturb 

sediments in order to bring particles into suspension. 


The eggs are exclusively “resting eggs”, spherical and spiny. They 

have a diameter of 240 to 260 µm. 

Longevity 

One generation (from hatching to the disappearance of the adults) 

can last, depending on the conditions, for a maximum of three to 

four months. 


The eggs are expelled in small jets by the females and fall to the 

bottom in a random fashion. 

87 

Crustacean drawing modified from Thiéry & Champeau 364



Although the nauplius* stage only lasts a few hours, growth is 

slow as the adults are only present from February to March (breeding 

period) whereas the eggs hatch in autumn, from October to 

December. The species disappears from the pools at the end of 

the winter even if they remain flooded, generally until April-May. 

Little interannual variability in phenology. As for most anostracans, 

the population functions in cohorts (hatching once the pool 

has flooded, synchronous growth of the individuals, death). 

� 




Linderiella massaliensis can live alone (Petit Laucien, Var) or be 

found with Lepidurus apus, Chirocephalus diaphanus (Lake Redon) 

and Branchipus schaefferi 373 . 


The species requires an autumn and winter flooding regime which 

corresponds to its biological cycle (fresh water). 


Early drying-out of the pool, in March for example, does not negatively 

affect the species as the adults are present from February 

and have been able to begin breeding. The introduction of Pumpkinseed 

Sunfish (Lepomis gibbosus, a species introduced from the 

USA) into Lake Bonne Cougne in 2001-2002 caused a collapse in 

the population of L. massaliensis during that hydrological cycle. 

The situation returned to normal after the summer drying out 

(eradication of the fish) and the autumn flooding. 


Few natural factors affect the biological cycle of this anostracan. 

For example, the species is not sensitive to freezing temperatures 

(population present under the ice at Bonne Cougne in 

January 2001, or a minimum water temperature of 1.8°C at Petit 

Laucien, in February 2003, Thiéry, original data). However, the 

species does not appear to tolerate water which is too warm. 



Number of sites 

This endemic species has only been found in five temporary pools 

in the Var (around Saint-Maximin and Brignoles) including Lake 

Redon, Lake Gavoty, and the Petit Laucien (type locality). 

88 

Adult stages (egg laying) 

Egg bank 

� 

Hatching and larval 

Populations 

Rare species: the population densities are between 0.5 and 2 ind.L -1 

at Bonne Cougne and Petit Laucien. The distribution, which is 

often regular, can sometimes become aggregated (influence of 

beds of charophytes, Ranunculus, etc.). Two studies (McNutt & 

Thiéry and Meglecz & Thiéry) are ongoing, one to map the distribution 

of the cysts of L. massaliensis within Lake Bonne Cougne, 

the other to quantify the consequences, through the genetic bottleneck* 

effect, of the excavation of sediments reducing the pool’s 

egg bank (analysis of enzymatic polymorphism, levels of mutation 

and ‘genetic drift’*). 


IUCN 






The Lake Bonne Cougne (Centre Var) is seriously threatened by 

both local activities (overdeepening, dyke construction, filling in 

with spoil brought in from building work or hospital waste containing 

contaminants, etc.) and by a project for the creation of a 

golf course on the land above the depression. Although the planning 

application deposited at the Var préfecture in Toulon was 

initially refused, it has just been approved. Generally speaking, 

the modifications of the water quality due to pumping from the 

karstic water table, extraction of the run-off waters of the topographical 

catchment area and to water running into the Lake 

Bonne Cougne from the golf course above will jeopardise in the 

short term (estimated at less than 10 years), the populations of 

branchiopods as well as those of planktonic crustaceans, cladocerans, 

copepods, etc. 377 


In the Lake Gavoty (Var), the development of the phytocenosis* 

considered to be natural, is in fact more likely to be influenced 

by the nutritional inputs provided by the catchment area (homes 

without a collective treatment system, livestock raising - llamas, 

goats, horses, etc.). The increase in nitrogen and phosporus levels 

reduces the hatching rate of L. massaliensis, whose mechanism 

depends on osmotic processes 374 . In addition, the covering of the 

sediments by a very dense vegetation bed reduces the mobility of 

the larvae and increases natural mortality. 


Two populations are threatened (Bonne Cougne and Gavoty) out 

of the five known. If the possibilities for connectivity* are considered 

for the five populations which are in a fragmented habitat 

within this metapopulation*, these threats represent a potential 

loss of 40% of the genetic heritage entering into the gene flows 

necessary for the maintenance of the metapopulation. 

In 2002, an identical situation in California for the vicariant 

species L. santarosae 367 resulted in the implementation of conservation 

measures. In contrast, no actions, or conservation measures 

are planned in France. 

- 

- 

- 

- 

-



Most of the sites in the Centre Var are included in Natura 2000* 

areas. 


Imperative protection of the sites in their current condition 

where the species is present. Inclusion of the species on a Red 

List. 

Bibliography 


AGRN-RH, 2000 3 ; Alonso, 1996 7 ; Anonymous, 1999 12 ; Brtek & 

Thiéry, 1995 52 ; Defaye et al., 1998 102 ; Nourisson & Thiéry, 1988 269 ; 

Thiéry & Champeau, 1988 364 ; Thiéry & Gasc, 1991 365 ; Thiéry & 

Fugate, 1994 367 ; Thiéry, 1986 370 , 1991 373 , 1996 374 ; Thiéry et al., 

1995 375 , 2002 377 . 


89


Tanymastix stagnalis (Linnaeus, 1758) 


Order: ANOSTRACA 

Family: BRANCHIPODIDAE 

Main synonyms 

Cancer stagnalis Linnaeus, 1758 

Gammarus stagnalis (Linnaeus), Fabricius, 1775 

Astacus stagnalis (Linnaeus) Pennant, 1777 

Branchipus lacunae (Guérin-Méneville) Baird, 1852 

Chirocephalus lacunae Frauenfeld, 1873 

Tanymastix lacunae Daday, 1910 


Spanish, Italian names: species is present in these two countries 

but there is no common name. 

English name: Fairy Shrimp (general name for Anostraca) 


No subspecies. 

Variability of adults: Margalef 236 and Nourisson 271 report aberrant 

T. stagnalis with hypertrophied antennae. The lenticular cysts can 

present some differences in diameter depending on their geographical 

origin 375 . An aberrant morphology also exists, corresponding 

to a deformation of the lenticular shape 376 . 


Body without carapace, eyes pedunculate, 11 pairs of phyllopods*, 

abdomen without appendages, and two cercopods, vermilion 

in colour, forming the furca, at the end of the abdomen. 

The males are distinguished from the females by their prehensile 

antennae which enable mating to take place. A frontal complex 

(clypeal) unfurls to play a tactile role and for partner recognition. 

The genital apparatus of the male (2 retractable hemi-penises) is 

situated at the base of the abdomen. The female’s brood pouch*, 

also borne on the ventral side at the base of the abdomen, bears 

two widely spaced spines. The structure and development of the 

cyst envelopes have been studied by Garreau de Loubresse 149, 150, 151 . 

The males are a pale green colour and the females have a bluebrown 

to orange oviger*. 

Their maximum size varies from 6 to 20 mm. 


Within the genus, three other endemic species are morphologically 

very similar: T. stellae (Corsica, Sardinia), T. affinis (Morocco) 

and T. motasi (Romania). Only males can be distinguished with 

certainty (examination of antennae). Generally speaking, T. stagnalis 

does not coexist with any of these endemic species. 

90 



T. stagnalis is a Palaearctic species, present throughout the 

Mediterranean region (Europe and North Africa) but it is very 

unevenly distributed 52 . 

France 

It has also been found in the Paris region 309 , in the Camargue 270 , 

the Var 363 and the Rhône Valley (Thiéry, original data ). 


Spain 7 , Italy (including Sardinia ) 92 , Macedonia and North Africa 

(Algeria) 154, 339 . 

Habitat 

male 


Tanymastix stagnalis is present in habitats which dry out rapidly 

such as shallow pools (sometimes on rock shelves as in the Fontainebleau 

Forest), vegetated ditches or sansouires (Salicornia 

scrub) in the Camargue. 

France 

• Roque-Haute Nature Reserve in the Hérault 260 . 

• Cupular pools of the Estérel Massif and the Colle du Rouet (see 

vol. 1, box 7). These basins of several dm 2 of surface area, fill up 

from October to May, depending on the year. They are covered 

with an acidic sandy-silty layer (average pH 5.2) from 5 to 10 cm 

in depth 363 . 

• Oligo-brackish temporary pools on calcareous clayey-silty 

substrate in the Camargue (Tour du Valat estate). 

Italy 

Temporary clear water with sparse vegetation 92 . 


Tanymastix stagnalis 

Mediterranean pools 


2 mm 

egg (190-430 µm)

Salicornia scrub 

Mediterranean and Thermo-Atlantic halophilous scrubs (code 1420) 


T. stagnalis is present in Mediterranean temporary pools as well 

as in other types of seasonally flooded habitat. 

Temporary pools 


Salicornia scrub (seasonally flooded) 

15.61 Mediterranean salt-meadow scrub. 

Ecology 


Feeding 

Tanymastix stagnalis, like all the anostracans, is microphagous* 

and feeds by filtering the food particles in suspension in the 

water (microplankton, micro-organisms and organic material), 

using its legs which are covered with bristles. 


The eggs, whose lenticular shape is specific to the genus Tanymastix, 

are coppery brown in colour and measure between 190 

and 430 µm in diameter. 

Behaviour 

The adults react to variations in light intensity; behaviour involving 

fleeing towards the pool bed, or even burying in the mud, have 

been observed when shade is cast over them. 

Longevity 

This varies according to the temperature and the season when 

the species appears in the habitat (from 30 days in summer to 

over 60 days in winter). 


The eggs are laid in open water, where they usually float, and 

accumulate towards the edges of the pool 262 . 


Tanymastix lives for a period of around three months at all seasons 

of the year, both in winter and in summer . It grows rapidly 

and, depending on the temperature of the water, can reach sexual 

maturity after seven to 40 days. The nauplius* stage only lasts a 

few hours. 

The biological cycle of T. stagnalis is very variable between sites 

and from year to year. The cycle begins with the flooding of the 

pool and its duration depends on the temperature. 


Hatching � larval � adults (egg laying) Hatching� larvae � adults 

(egg laying) 

Egg bank 

One cycle per year in general, rarely two, in which case poor breeding success is 

observed during the second cycle (Lanau pool) (Thiéry, original data). 




Tanymastix stagnalis can be associated with the anostracans Chirocephalus 

diaphanus, Branchipus schaefferi, with the notostracan 

Triops cancriformis and with Imnadia yeyetta (Spinicaudata). 

On the other hand, it is often the only species present in cupular 

pools, the other species having longer biological cycles. 


Pioneer species, colonising clear, fresh and poorly mineralised water. 


Species present in pools with dry sediments in the summer, without 

a groundwater capillary link. Any modification in the environment 

which can cause an increase in the mineral content 

(conductivity


Legal status of the species/level of protection control over land ownership and usage) and the pools of the 

Colle du Rouet are included in a Natura 2000* site. 

IUCN 






In general, the infilling of pools and the use of pesticides in the 

rural environment threaten populations. 



Some Italian pools containing Tanymastix are situated in protected 

areas 263 . In France, the Roque-Haute site has a legal “Réserve 

Naturelle” status (currently ineffective because of the absence of 

92 

- 

- 

- 

- 

- 


It is vital that the small shallow biotopes be maintained in a 

healthy condition. 

Bibliography 

Alonso, 1996 7 ; Artom, 1927 17 ; Brtek & Thiéry, 1995 52 ; Cottarelli 

& Mura, 1983 92 ; Defaye et al., 1998 102 ; García & de Lomas, 

2001 146 ; Garreau de loubresse, 1965 149 , 1974 150 , 1982 151 ; 

Gauthier, 1928 154 ; Margalef, 1958 236 ; Médail et al., 1998 246 ; 

Moubayed, 1998 260 ; Mura, 1991 262 , 2001 263 ; Nourisson, 1960 271 ; 

Nourisson & Aguesse, 1961 270 ; Rabet, 1994 309 ; Samraoui & 

Dumont, 2002 339 ; Terzian, 1979 363 ; Thiéry & Gasc, 1991 365 ; 

Thiéry, 1991 373 , 1996 374 ; Thiéry et al., 1995 375 , en préparation 376 . 

Authors: Thiéry A. & T. Calvière


Triops cancriformis (Bosc, 1801) (1) & Lepidurus apus (Linné, 1758) (2) 


Order: NOTOSTRACA 

Family: TRIOPSIDAE 

Main synonyms 

(1) Limulus cancriformis Lamrck, 1801; Apus cancriformis Bosc, 

1802; Apus viridis Bosc, 1802; Triops simplex Ghigi, 1921 

(2) Monoculus apus L., 1758; Binoculus palustris Müller, 1776 

Apus productus Bosc, 1802; Apus apus (Linnaeus) Latreille, 1802 

Lepidurus productus (Bosc) Leach, 1819 

French names: (1) Triops cancriforme (sometimes inaccurately 

called apus – Apus is a bird genus), (2) Lépidure 

English name: Tadpole Shrimp (general name for Anostraca) 

Subspecies 

Triops c. cancriformis and Lepidurus a. apus are the only two French 

subspecies of Notostraca, but two other subspecies of 

T. cancriformis exist (T. cancriformis simplex and T. cancriformis 

mauretanicus) as well as subspecies of L. apus including L. apus 

lubbocki in the countries of the Mediterranean Basin (Spain, 

Morocco, etc.). 


These crustaceans, with a dorsoventrally flattened body, have an 

oval, streamlined dorsal carapace covering the head, the thorax 

and a part of the abdomen. They can reach a maximum size of 10 

to 11 cm, but the two cercopods* (furca), at the end of the 

abdomen, are almost as long as the body. 

The compound eyes are sessile* and dorsal. The phyllopods*, the 

first two pairs of which are locomotive, are more numerous in 

Triops (48 to 57 pairs) than in Lepidurus (35 to 48 pairs). 

The posterior abdominal segments are apodal (no legs): the last 

four to five segments in Lepidurus, the last four to seven segments 

in female Triops and the last five to nine segments in male 

Triops. 

Female Triops are also distinguished from the males by the structure 

of the 11 th pair of thoracic legs which are transformed into 

a brood pouch (oostegopod), where the eggs are stored. 

The first pair of legs have flagelliform endites (extensions) 

extending some way beyond the edge of the carapace in Triops 

but less so in Lepidurus. The latter is characterised also by the 

presence, on the final segment (the telson), of a caudal ramus 

with toothed edges. 

The dorsal shield is greenish-brown in Lepidurus and brownish in 

Triops, in which the thoracic legs can be reddish or greenish. 


These are the only two species of French Notostraca. Separation 

of the two species is easy: telson with or without caudal ramus. 

Identification of the various subspecies, on the other hand, is 

very difficult. Specialist advice is needed. 

1 cm 

Triops cancriformis 

Lepidurus apus 



1 cm 

egg (340-400 µm) 

egg (430-520 µm) 

France 

Triops is present over the whole country (including Corsica) and 

is fairly frequent in the east and in the coastal regions, particularly 

the Mediterranean regions. Lepidurus appears to be absent 

from eastern France and Corsica, but is more frequent in the north 

than Triops. Generally speaking, the two species are allopatric 

(separate ranges). The distribution of the two species seems clearly 

determined by precipitation, as is the case in southern France 372 

(P 600 mm for Lepidurus), in Israel 201 , 

Algeria 154 and Morocco 369, 371 . The genus Triops is found in drier 

regions than Lepidurus. 

These distributions may be erroneous, as the two species seem to 

have been often confused. Furthermore, their presence at certain 

stations has not been confirmed since 1950. For further details, 

consult Nourisson & Thiéry 269 , Brtek & Thiéry 52 and Defaye et al. 102 . 


Both species are present throughout Europe, as well as in the 

Mediterranean Basin for Lepidurus apus apus. 

• Spain: Lepidurus apus apus, Triops cancriformis simplex, 

T. c. mauretanicus 7 (which replaces T. c. cancriformis in southern 

Spain). 

•Portugal: T. cancriformis. 

• Italy: L. apus (mainland Italy, Sicily, Sardinia ), T. cancriformis, 

and T. numidicus (= T. granarius), a species of African origin present 

in Calabria. 

93 

Crustacean drawings modified from Alonso 7


• Malta: T. cancriformis 205 . 

• Croatia: L. apus. 

• Former Yugoslavia: L. apus, T. cancriformis 238, 289 . 

•Morocco: T. c. simplex, T. c. mauritanicus, L. a. lubbocki 369, 371 . 

• Israel: L. a. lubbocki, T. c. cancriformis (Thiéry, original data ). 

Habitat 


The two species are found in temporary pools. 

Whereas Triops can be found in sometimes quite turbid pools, 

Lepidurus is usually found in fresh, clear water, and sometimes in 

temporary watercourses with a weak current. 

Triops is also present in artificial habitats flooded in the summer, 

such as the ricefields in the Camargue 297 . 





(1) 22.32 Annual Mediterranean amphibious swards (Nanocyperetalia). 

(2) 22.34 Southern amphibious communities (Isoetalia). 

Ricefields 

(1) 82.41 Ricefields. 

Ecology 


Feeding 

They take a wide range of food and they can be considered as 

predators, herbivores or detritivores. In turbid waters, they are 

scavengers: they feed on non-living particles (dead organisms, 

organic detritus, mineral particles) in suspension in the water, 

and on the bacteria that that build up on these particles. They dig 

up the mud with the front part of their carapace. 

Feeding can vary during the growth of the animal, which can 

pass from a strictly phytoplanktonophagous* feeding regime, 

when it measures less than 5 mm, to a herbivorous regime (in the 

broadest senses); when it is larger than 1 cm, it also feeds on 

microcrustaceans. As an adult, it is capable of devouring weakened 

anostracans (usually Branchipus schaefferi), chironomid larvae, 

Oligochaete worms and tadpoles (Pelobates cultripes). 

Behaviour 

They usually swim close to the bottom, but can come up to the 

surface, ventral side uppermost, if there is a lack of oxygen. 

Reproduction 

Mating is ventral-ventral in Lepidurus, whereas it appears to be 

dorso-ventral in Triops. Parthenogenetic reproduction is frequent 

in Triops, but when both sexes are present, sexual reproduction is 

the rule. The sex ratio* varies according to latitude with an 

absence of males beyond 50°N, a low percentage between 45° 

94 

and 50°N (northern and central Europe) and an equal sex ratio 

below 45°N (southern, western Europe and North Africa), with 

exceptions (mainland Italy, Sicily, etc.). 

The parthenogenetic females possess a hermaphroditic gland but 

the spermatozoa formed degenerate and play no part in reproduction. 


The eggs acquire their external envelope in the brood pouch*. 

They are smooth. Their diameter is greater in Lepidurus (430-520 µm) 

than in Triops (340-400 µm). 


The eggs (60 to 70 in Lepidurus) are deposited just before the 

moult, in clumps, on small pebbles where they remain during the 

dry period. In Lepidurus, the female can lay eggs on the stems of 

aquatic vegetation or bury her eggs in the mud. 


After hatching, the nauplius* larvae evolve rapidly into metanauplius* 

larvae, with the dorsal shield already present. 

Lepidurus, which is present from January to June (most often 

from February to April), has a life expectancy of 4 to 6 months. 

Triops, which can be found in all seasons of the year in the 

flooded phase, from March to January (most often from May to 

October), has fairly rapid growth and can produce resting eggs 

from the 15 th day, when it is still small in size. 

Triops cancriformis 

Temporary pool 


� 

stages 

Lepidurus apus 

adult stages (egg laying) 


Egg bank 

� 



� 

stages 

adult stages (egg laying) 

Egg bank 



Their cycle being relatively long, these notostracans require a 

submersion period of three to five months. 

Triops, which can be found from March to January, is more thermophilic 

than Lepidurus, which is present from January to June. 

Triops cannot withstand temperatures higher than 35°C and thus 

from June to August is practically absent from habitats flooded 

in summer, such as the ricefields in the Camargue. 

�


Triops is usually found with Branchipus schaefferi and more 

rarely with Chirocephalus diaphanus. In the Camargue, it can also 

live alongside Tanymastix stagnalis, Imnadia yeyetta and Eoleptestheria 

ticinensis. 

In ricefields, when its populations explode (densities reaching 

300 individuals per m 2 have been measured in the Camargue 297 ), 

it can become an important part of the diet of the Little Egret 

Egretta garzetta 169 . 

Lepidurus usually lives with Chirocephalus diaphanus but can, in 

the Var, live alongside Linderiella massaliensis. 

Territories 

In notostracans, there is a differential micro-distribution between 

the sexes in the pool. 


Early drying out of the pool (case of the Lanau pool, for example) 

can cut short the otherwise significant life span of these species. 

Predation, notably by Ardeidae, can play an important role in certain 

populations (Camargue) 169, 297 . 



In France, they are present at many stations: Triops is found notably 

at the sites of Roque-Haute (Languedoc-Roussillon), Padulu (Corsica), 

in the Camargue and the Crau (Lanau pool), while Lepidurus 

has been found in the pools of the Var, Lake Redon, Lake Gavoty 

and Saint Maximin (PACA). A description of the distribution of 

the two species in Languedoc-Roussillon is given by Thiéry 372 . 

In Italy, these species are fairly widespread, in particular in the 

ricefields. 


In France, these two species are common but Triops seems to 

have been often confused with Lepidurus and their presence at 

some stations has not been confirmed since 1950. 


IUCN 





- 

- 

- 

- 

Malta: rare species (R) 344 




As Lepidurus can be found in the overflow pools of rivers (Loire, 

for example), physical planning of watercourses and their regulation 

in order to limit flooding can represent an indirect threat 

to their populations. 

Triops is sometimes considered to be a nuisance in ricefields 

where it can be extremely abundant (it uproots young plants, 

eats their growing tips, and reduces photosynthesis activity by 

bringing sediments into suspension). Others estimate that these 

perturbations are minor and that it acts as an extremely efficient 

biological control agent to combat invading plants. It is, however, 

eliminated by insecticides or copper sulfate. 

More generally, the infilling of pools and the use of pesticides 

destroys populations. 



In France the pools of Roque-Haute and the Camargue are situated 

in Nature Reserves (at Roque-Haute, however, conflicts with 

the owners do not allow management of the reserve to take place). 

The lakes of the Centre Var, the Padulu pool in Corsica, and the 

Lanau pool, acquired by the CEEP in 1998, are included within 

Natura 2000* areas. 


Ensure the conservation of the habitat (hydrological functioning 

and water quality). 

Bibliography 

Alonso, 1996 7 ; Anonymous,1999 12 ; Boix et al., 2002 41 ; Brtek & 

Thiéry, 1995 52 ; Defaye et al., 1998 102 ; García & de Lomas, 

2001 146 ; Gauthier, 1928 154 ; Hafner, 1977 169 ; Knoepffler, 1978- 

1979 199 ; Kuller & Gasith, 1996 201 ; Lafranco, 1990 205 ; Longhurst, 

1955 217 ; Marincek & Petrov, 1992 238 ; Moubayed, 1996 259 ; Nourisson 

& Thiéry, 1988 269 ; OEC, 2001 273 ; Petrov & Petrov, 1997 289 ; 

Pont & Vaquer, 1986 297 ; Scanabissi & Mondini, 2002 342, 343 ; 

Schembri & Sultana, 1989 344 ; Thiéry & Gasc, 1991 365 ; Thiéry, 

1985 368 , 1986 369 , 1987 371 , 1988 372 , 1991 373 , 1996 374 ; Thiéry et al., 

1995 375 ; Zaffagnini & Trentini, 1980 404 


95


Ischnura pumilio (Charpentier, 1825) (1) & Ischnura genei (Rambur, 1842) (2) 

Order: ODONATA 

Suborder: ZYGOPTERA 

Family: COENAGRIONIDAE 

Main synonyms 

(1) Agrion rubellum Curtis, 1839; Agrion cognata Sélys, 1841 

(2) Agrion genei Rambur, 1842 

French names: (1) l’Agrion nain, (2) l’Agrion de Gené 

English name: (1) Scarce blue-tailed damselfly 

95, 401 


Like all the Zygoptera, these species have the eyes well separated 

and the four wings of exactly the same shape. The family characteristics 

include the absence of any metallic reflections and the 

entirely hyaline (transparent) wings, which are petiolate (stalked at 

the base). Both species have a mostly dark or black abdomen with 

a bronze tint and, in males, the eighth segment at least partly 

blue (also the ninth in Ischnura pumilio). The pterostigma (dark 

mark on the wing) is scarcely longer than wide. In I. genei, the 

pterostigma is slightly larger on the forewings than on the hindwings, 

while in I. pumilio it is almost twice as large on the 

forewings as on the hindwings. Only I. genei has a median tubercule 

on the prothorax* and this is clearly indented. 

Abdomen: 18 to 27 mm. 

Wings: 12 to 19 mm. 

Subspecies 

None 


Ischnura elegans (similar to I. genei in particular). 



France 

Ischnura pumilio occurs only on the mainland (especially the 

Rhône Valley, Pre-Alps, Champagne-Ardenne, Centre-Ouest, and 

Var), and I. genei is only found in Corsica 122, 159 . 


Ischnura genei occurs only on the islands of the western 

Mediterranean (Sardinia, Sicily, Capri, Malta, Elba, etc.). 

I. pumilio is more widely distributed, along the whole of the coast 

of Mediterranean Europe and part of the North African and 

Middle Eastern coasts 95, 401 . 

Habitat 

Both species are pioneer colonisers and are therefore attracted to 

recently created habitats, temporary or otherwise, usually with 

still water. They may be found in brackish water, and sometimes 

in still areas of water associated with watercourses (oxbow lakes, 

etc.). They are usually seen below 800 m altitude, however some 

sites are known at up to 2,100 m (Ischnura pumilio) 55, 95, 173 . 

96 

I. pumilio does not like acid biotopes (Samraoui, unpublished 

data). It is also known to disappear from some sites after a few 

years, probably due to habitat succession 68, 207 . 

Ecology 


male 

Ischnura pumilio 

0.5 cm 

(1) Ischnura pumilio: in southern France, the adults, or imagos, 

can be seen from the beginning of April to the end of September 95 , 

sometimes March to October (Papazian, pers. com.), with 

ephemeral populations 323 , while in Spain and northeast Algeria 

the species appears in two waves, one in August, the other in 

September/ October (Samraoui, unpublished data). 

The lifespan of the imagos is fairly short (of the order 40 days) 

and their sexual maturation is also very rapid. Females are capable 

of breeding six to ten days after emergence 69 and the males 

possibly even sooner 207 . This maturation is all the more rapid if 

the individual emerges late in the season, thereby maximising the 

chances of breeding 207 . After mating, the female goes alone to lay 

the eggs on the stems of plants emerging from shallow water, 

choosing species with stems which are fairly soft (often pithy), 

such as certain rushes 68 . The female stations herself just above 

the surface of the water and plunges her abdomen in so that she 

can lay her eggs in the submerged part of the stem. She prefers 

to lay at the level of the leaf sheaths (rushes), where the growth 

of the plant tissues will completely cover the eggs; this appears 

to be advantageous for their survival during dry conditions 68 . The 

eggs can survive for at least two weeks in an emergent plant if 

it has its base in the water. The eggs hatch quickly (after 15 days 

at an ambient temperature of 25°C 68 , most often between 20 and 

30 days (Papazian, pers. com.). Rapid larval development enables 

the adults of the second generation to emerge before the pool 

dries out completely. In temporary habitats, this second generation 

disperses and awaits the return of the water before laying 

(August to October, depending on the date of re-filling with water), 

usually in another pool. The eggs then hatch fairly quickly and 

the young larvae spend the winter in the diapause* stage.

In the Mediterranean region, the short hatching time, rapid larval 

development, precocious emergence spread over a long 

period of time (advantageous in view of the interannual variability 

of the hydrological conditions), and the rapid maturation of 

newly emerged individuals enable two generations of the species 

to be produced per year. The rapid dispersal of individuals following 

emergence (migratory instinct), together with the rapid 

turnover of generations, enable I. pumilio easily to colonise temporary 

habitats, forming new colonies 69 . 

(2) I. genei: few facts are known regarding this species. In France, 

its phenology is very similar to I. pumilio (Papazian, pers. com.). 

In Sardinia, adults are present from the beginning of May to the 

end of September 55 . 

Regarding its ecology, one may refer partly to I. elegans, which is 

very similar. Like I. pumilio and I. elegans, I. genei has two generations 

per year, with rapid development of the pre-imaginal* stages 5, 95 . 

Simplified phenological cycle of Ischnura pumilio 

Temporary pool in southern France 


First generation (pool A) 

� 

followed by larval growth 

Emergence of adults, egg laying and migration 

Second generation 


Larval development, emergence 

and dispersal to other pools 

Hydrology 

Both species lay their eggs in submerged stems. The eggs die if 

the plant is out of the water for more than about fifteen days (no 

summer diapause at the egg stage), as do the larvae if the pool 

dries up too early. 

Vegetation 

The sites used are rather sunny and the emergent aquatic vegetation 

need not necessarily be very well developed 173, 204 . 

The plants used by Ischnura pumilio for egg laying have a grasslike 

form 330 such as rushes (Juncus bulbosus, J. tenuis, J. subnodulosus, 

J. inflexus, J. articulatus) or spike-rushes (Eleocharis 

palustris) 68,173 . I. pumilio prefers to colonise water bodies with 

clayey beds and fairly shallow average depth (less than one metre). 

Competition 

Ischnura pumilio has a low level of tolerance of competition from 

other dragonfly species. It is a requirement therefore that there are 

few of these at the sites or at least that some areas are free from 

competitors (notably I. elegans), or that the vegetation is well-developed 

enough to allow I. pumilio to find refuge 173 . Accordingly, the adults 

often only occupy a limited territory within a suitable biotope 330 . 


Egg laying (pool B) 


� 

Winter larval diapause, 

Hatching of eggs 

Larval diapause… 

(1) Ischnura pumilio is less common than I. elegans. In the French 

Mediterranean region, it is known at the pools in the Plaine des 

Maures 244 , the Centre Var 377 , the Colle du Rouet, Porquerolles 

� 

Odonata 

(oxidation ponds), the marshes and rice fields of the Camargue, 

on the Durance (small temporary pools at Puy-Sainte-Réparade), 

in the Crau (Vergière canal) and on the Touloubre at Saint-Chamas 

(Papazian, pers. com.). 

(2) I. genei occurs at a number of sites in Sardinia 55 and at 16 out 

of 50 stations visited as part of the dragonfly survey in Corsica 

159, 323 , notably in the temporary pools at Tre Padule de Suartone. 


IUCN 







Interference with habitats that dry out in summer (drainage, filling 

in), and lack of maintenance of pools. 


Overgrowing of vegetation at water bodies, siltation and overgrowing 

of wetlands; feeding competition with other dragonflies 

(such as Ischnura elegans), predation by spiders and natural mortality 

during heavy rain (very frail species). 


Early and prolonged drought may result in the complete destruction 

of all the larvae at a site. Only recolonisation will then 

enable these Ischnura to return. 



In Corsica, Tre Padule benefits from its legally protected “Réserve 

Naturelle (Nature Reserve)” status. The sites at the Plaine des 

Maures, the Centre Var and the Colle du Rouet are included in 

Natura 2000* areas. 


The aquatic vegetation should be maintained in an undeveloped 

condition if the site is not much used by dragonflies, and even 

more so if there is strong competition. Vegetation management 

should favour small rushes or club-rushes, with sufficiently low 

water levels. In addition management should aim for the pool to 

be flooded until spring (from November to April-May) to enable 

the larvae to develop. 

Bibliography 

- 

- 

- 

Italy, regional protection 

(Tuscany): regional law n° 56 

of 06/04/2000 

- 

Aguesse, 1968 5 ; Anonymous, 1997 11 ; Bucciarelli et al., 1983 55 ; 

Cham, 1992 68 , 1993 69 ; D’Aguilar & Dommanget, 1998 95 ; 

Dommanget, 1994 122 ; Grand & Papazian, 2000 159 ; Heidemann & 

Seidenbusch, 2002 173 ; Landmann, 1985 204 ; Langenbach, 1993 207 ; 

Médail et al., 1993 244 ; Roche, 1990 323 ; Rudolph, 1979 330 ; Thiéry et 

al., 2002 377 ; Wendler & Nüss, 1994 401 . 

Author: Gendre T. 

Collaborators: Jakob C., M. Papazian, B. Samraoui & N. Yavercovski 

97


Lestes barbarus (Fabricius, 1798) (1) & Lestes virens (Charpentier, 1825) (2) 


Suborder: ZYGOPTERA 

Family: LESTIDAE 

Main synonyms 

(1) Agrion nympha Hansemann, 1823 

Agrion barbara Fabricius, 1798 

Lestes barbara Selys, 1840 

(2) Lestes vestalis Rambur, 1842 

Agrion virens Charpentier, 1825 

French names: (1) Leste sauvage, (2) Leste verdoyant 

English names: (1) Shy Emerald Damselfly, Southern Emerald 

Damselfly, (2) Small Emerald Damselfly 

Subspecies 

L virens virens (in southern France) 

L. virens vestalis (in Italy) 


(1) Lestes barbarus has a clearly bicoloured pterostigma (dark 

mark in the wing) when mature: brown in its basal half and yellow-white 

in its apical half. The overall body colour is clear 

metallic green at first, becoming more or less coppery, even dull 

brown in old individuals. The lower hind part of the head, the 

coxa, the metathorax, and the thoracic and abdominal lines are 

pale yellow. The male has short cerci*. The female has a rounded 

vulvar scale, untoothed, and ochre-coloured legs in most cases. 

Measurements of male: abdomen 26 to 35 mm; hind wings 19 to 

25 mm. 

Measurements of female: abdomen 26 to 33 mm; hind wings 21 

to 25 mm. 

(2) Lestes virens has a uniformly pale brown pterostigma, edged 

with white veins (apart from in juveniles). The general body 

colour is brilliant bronze-green, the sides of the thorax and the 

tip of the abdomen are pale blue in mature males. The lower hind 

part of the head is pale yellow, as in L. barbarus. The male has 

very short cerci. The female has a vulvar scale with a short, only 

slightly pointed tooth. L. virens is smaller than L. barbarus. The 

subspecies from southern France, L virens virens, shows a thin 

pale yellow line along the anterior part of the metathorax*. 

Measurements of male: abdomen 24 to 34 mm, hindwings 16 to 

24 mm. 

Measurements of female: abdomen 24 to 30 mm, hindwings 17 

to 23 mm 95, 173, 322, 402 . 


Lestes barbarus is very similar to L. virens (especially at the juvenile 

stage) and to its subspecies L. v. vestalis. 

98 



France 

(1) West and southeast France (but mainly a southern species: 

coast and Corsica). 

(2) Southern France (coast and Corsica). 


(1) Southern and temperate Europe, North Africa, Middle East. 

(2) Western Mediterranean Basin except mainland Italy. 

Habitat 

(1) Lestes barbarus is found in still, sunlit water bodies, clear, 

shallow, often but not exclusively oligotrophic*, up to 250 m altitude. 

As a pioneer species, it appears more particularly to prefer 

habitats which dry out in summer (temporary pools) and brackish 

coastal marshes, where there are often large populations. 

(2) L. virens occurs in still bodies of fresh water, including temporary 

pools, colonised by vegetation, up to 1,000 m altitude. 

Ecology 

males 


Lestes barbarus 

Lestes virens 

1 cm 

(1) In southern France, the adults of Lestes barbarus (known as 

imagos) are fairly common from the end of May (immature at 

that time, they become mature in June) to the end of September 

(at which time they colonise new pools). In spring, after emergence, 

the immature imago leaves the water, to return when it is 

sexually mature, usually in June-July. The period of maturation 

between flying away and returning to the pool is estimated to be 

15 days on average for Zygoptera 89 . In other regions maturation

may take place following a period of aestivation, as is the case in 

particular of the populations of southern Italy and northern 

Algeria 338, 340, 390 . 

The duration of the maturation period influences the egg-laying 

period: in southern France laying may take place from the end of 

June to September (Papazian, pers. com.), while in southeast 

Algeria (in Numidia) the females become mature in September 

and no breeding behaviour is seen before October (Samraoui, 

unpublished data). 

Mating (forming a heart shape) and the tandem flight which follows, 

may last for between 30 and 160 minutes (for both species) 

and usually takes place during late morning 387 . After mating, the 

partners continue their tandem flight and the females begin to 

lay eggs using their ovipositor (egg-laying organ). In both species, 

eggs are laid into the aerial parts of plant stems 4 : the female’s 

ovipositor pierces the plant to insert the eggs, which are often 

laid in groups in the same stem, with gaps of a few millimetres 

between them. The distances covered between stems range from 

a few centimetres to a few metres 387 . Often, the male breaks 

away from the tandem formation after a certain time, and the 

female then continues to lay by herself. The total duration of egg 

laying may last for up to six hours, depending on the weather 

conditions and diurnal rhythm. 

After a period of diapause* inside the plant, which varies according 

to the date of laying, the eggs hatch in the spring of the following 

year, and give birth to elongated larvae which develop fairly 

quickly, taking about two months and moulting about ten times. 

These larvae are carnivorous, like all dragonfly larvae, and eat 

twice their own weight in food every day. 

Thanks to the diapause eggs (a primitive protective characteristic 

of the species) and to the short larval development period (six 

to eight weeks) very synchronised emergence may take place in 

spring, with practically all the imagos emerging over a short 

time. 

The sex ratio* (calculated at a Camargue site) is 50% females for 

larvae and 54.1% females for exuviae 4 . The adult dies later in the 

autumn. The maximum adult lifespan of a damselfly such as 

L. barbarus or L. virens in southern France is one and a half months, 

while in populations further to the south it extends to more than 

two and a half months 4 . 

(2) The mature adults of Lestes virens may be seen from June to 

September in southern France, and from May (occasionally end 

of April) to November in northeast Algeria (Samraoui, unpublished 

data). The female, who may or may not be accompanied by 

the male, lays her eggs in pairs, in spring and summer, into plant 

stems. After a winter egg diapause (often preceded by a summer 

diapause), larval development takes place in the following spring 

as with L. barbarus, and lasts for eight to ten weeks 402 . 

In the North African part of the Mediterranean Basin, a proportion 

of Lestes also lays in summer, before the site becomes flooded; 

the diapause eggs then withstand the drought, then overwinter 

and do not hatch until the following spring 340 . In Numidia, this is 

true of L. virens at lower altitudes (laying in June-July), while at 

higher altitudes breeding activity in this species does not begin 

until August (Samraoui, unpublished data). 

Both species (L. barbarus, L. virens) only have one generation per 

year. 

Phenological cycle of Lestes barbarus 


� 


Odonata 


Larval development 

Emergence of adults 

Maturation of adults 

Winter temperatures 

Lestes barbarus and L. virens are adapted to the Mediterranean 

climate due to their rapid life cycle and their temperature 

requirements 95 . However, in southern France, L. barbarus requires 

a period of cold, of at least fifteen weeks below 10°C, to trigger 

the hatching of all of the eggs 4 . 

Hydrology 

As far as is known for these two species, the eggs are laid most 

often in a dry environment; the pool need not be flooded at the 

time of laying for the eggs to survive 4, 340 , since they are capable 

of entering a long period of diapause inside the plant stems. 

On the other hand, the emergence of adults, which is delayed, 

must take place before the pool dries out completely in spring or 

summer, and this is usually the case thanks to the rapidity of larval 

development. 

At the time of the emergence of the prolarva*, it sometimes happens 

that the place where the eggs were laid is dry; the prolarva, 

which on hatching allows itself to drop off the stem, is then 

capable of crawling to the water over a distance of a metre or 

more (Papazian, pers. com.). 

Vegetation 

According to Utzeri et al. 387 , the plants preferred by Lestes virens 

vestalis for egg laying at a temporary pool in Italy (Rome), are 

Carex sp. and Juncus effusus (green or dry stems). L. barbarus 

shows a preference for the same species, as well as Juncus articulatus, 

Polygonum hydropiper, Alisma plantago and Mentha sp. 

Samraoui (unpublished data) reports that in Numidia, L. barbarus 

prefers to lay in the stems of Juncus effusus. 



Egg laying 

Death of adults 

� 

Eggs in diapause 


Both species are known from the Alpes Maritimes at the Pic des 

Courmettes (Papazian, pers. com.), as well as in the Var: Plaine 

des Maures 244 , Colle du Rouet 325 , Joyeuse valley at Callas, lakes of 

the Centre Var 377 . Lestes barbarus has been observed at several 

other localities in the Var (Palayson wood, Giens Peninsula) 363 (Papazian, 

pers com.), as well as in the Bouches-du-Rhône in the Crau 

and in the Camargue 363 , and in the Hérault at Roque-Haute 260 . 

L. virens occurs in the Vaucluse along the Durance (Papazian, 

pers. com.) 

99


Corsica 

Lestes barbarus, a migratory and pioneer species, appears to be 

commoner than L. virens, with six stations out of the 50 visited 

during the survey of dragonflies in Corsica, compared with only 

two stations for L. virens. Both species are known in particular 

from Tre Padule and Padule Maggiore 159, 323 . 


No information. 


Lestes barbarus and L. virens have no legal protection at the present 

time 227 . 

IUCN 







Drainage and infilling of the sites used. 


Siltation and overgrowing of wetlands; feeding competition with 

other dragonflies (e.g. Sympetrum fonscolombii), predation by 

water beetles, parasites, interannual variability of hydrological 

conditions (early drying out may result in the death of all larvae). 



In France: 

• The Tre Padule de Suartone and Roque-Haute sites are legally 

protected due to their “Réserve Naturelle (Nature Reserve)” status 

(at Roque-Haute, the management of the site is compromised 

as there is no control over land ownership or the activities 

for which the site is used). 

100 

- 

- 

- 

- 

- 

• The lakes of the Centre Var, the Plaine des Maures, the Palayson 

wood and the Colle du Rouet are included within Natura 2000* 

areas, as well as the Lanau pool, which was acquired in 1998 by 

the CEEP. 


• Attempt to ensure, at temporary pools and fresh or brackish 

still waters, that there are flooded conditions until spring (April- 

May), suitable for the development of the larvae. 

• Maintain and promote aquatic vegetation and bands of vegetation 

around the edge of the pool, suitable for egg laying 

(rushes, sedges, water plantains, mints, bistorts). 

Bibliography 

Aguesse, 1961 4 ; Anonymous, 1997 11 ; Carchini & Nicolai, 

1984 62 ; Corbet, 1999 89 ; D’Aguilar & Dommanget, 1998 95 ; 

Donath, 1981 123 ; Grand & Papazian, 2000 159 ; Heidemann & 

Seidenbusch, 2002 173 ; MNHN, 1993 227 ; Médail et al. 1993 244 ; 

Moubayed, 1998 260 ; Robert, 1958 322 ; Roche, 1990 323 ; Rombaut, 

1994 325 ; Samraoui & Corbet, 2000 340 ; Samraoui et al., 1998 341 ; 

Shiemenz, 1954 350 ; Terzian, 1979 363 ; Thiéry et al., 2002 377 ; 

Utzeri et al., 1976 388 , 1984 390 , 1987 387 , 1988 389 ; Wendler & Nüss, 

1997 402 . 

Authors: Jakob C. & J. Fuselier 

Collaborators: Papazian M., B. Samraoui & N. Yavercovski

Sympetrum fonscolombii (Sélys, 1840) (1) & 

Sympetrum meridionale (Sélys, 1841) (2) 


Suborder: ANISOPTERA 

Family: LIBELLULIDAE 

Main synonyms 

(1) Libellula erythroneura Schneider, 1845 

Libellula insignis Brittinger, 1850 

(2) Libellula hybrida Rambur, 1842 

Libellula nudicollis Hagen, 1850 

Libellula meridionalis Sélys, 1841 

French names: (1) Sympetrum de Fonscolombe, 

(2) Sympetrum méridional 

English names: (1) Red-veined Darter, (2) Southern Darter 


Sympetrum have six to eight veins in the forewings. The bases of 

the hindwings have a yellow mark which is fairly well-developed 

in S. fonscolombii (but often absent in S. meridionale). 

(1) In S. fonscolombii, the legs are black with a yellow stripe on 

the outer edge. 

Measurements of male: abdomen 22 to 29 mm, hindwings 26 to 

30 mm. 

Measurements of female: abdomen 22 to 28 mm, hindwings 

26 to 31 mm. 

(2) S. meridionale has yellowish or brown thorax with practically 

no black, and yellowish legs with scarcely any black markings. 

The pterostigma is pale yellow to reddish, edged with black veins. 

The abdomen is red in the male and brown-yellow in the female. 

Measurements of male and female: abdomen 22 to 28 mm, hindwings 

20 to 30 mm 95, 322, 401 . 


No confusion possible with Sympetrum fonscolombii, whereas 

S. meridionale may be confused with S. vulgatum ibericum and 

S. sinaiticum. 



France 

(1) S. fonscolombii is found especially in the Mediterranean area, 

including Corsica, and in the Rhône Valley; it is much rarer in the 

rest of the country. 

(2) S. meridionale has the same range but appears to be commoner 

in the rest of the country 122, 401 . 


The whole of the Mediterranean region 95 . 

Habitat 

Odonata 

(1) Sympetrum fonscolombii prefers still, occasionally brackish 

water (pools, ponds, gravel pits, marshes, boggy edges of lakes), 

from sea level up to 2,000 m (sometimes much higher for wandering 

individuals). It prefers wetlands that are very little vegetated 

(or vegetated with helophytes* and hydrophytes*), well 

exposed to the sun, and of very variable water depth. 

(2) S. meridionale also prefers still water, more or less colonised 

by vegetation, up to 1,800 m altitude 95, 173 . 

Ecology 


1 cm 

Sympetrum fonscolombii 

Sympetrum meridionale 

1 cm 

males 

The eggs of Sympetrum often enter a diapause* phase after being 

laid, and do not hatch until they have spent the summer or winter 

in egg form 89 . However, in S. striolatum in Corsica it is also 

possible to observe larvae hatching out before the winter and 

these spend the winter in the water of the pool (Papazian, 

unpublished data.) In both cases, this type of annual cycle, with 

the eggs laid in autumn, corresponds to the flooding cycle of 

Mediterranean temporary pools 340 . S. meridionale and S. fonscolombii 

lay directly into the water (a highly evolved characteristic) 

but have partly retained the diapause at the egg stage. 

101


(1) Adult S. fonscolombii show well-marked exploratory behaviour 

and undertake movements over great distances (migratory 

species) 95, 122 , which explains their sudden appearance in more or 

less large numbers 322 . The imagos are usually seen from March 

(an early species) to December, mainly from March to June and 

from August to December in temporary habitats. The mass movements 

of various Sympetrum that are observed in the countries 

around the Mediterranean are explained as being flights between 

pools which dry out in summer and aestivation sites 338 . 

In this species, there are at least two and even three generations 

per year in the Mediterranean region: the first emerges in March 

and, following a period of rapid maturation, lays eggs which 

hatch after very short incubation period; the larvae grow quickly 

before the water body dries up completely, and the emergence of 

the second generation takes place 340 (Papazian, pers. com.). This 

second generation lays at the end of summer or in autumn, as 

soon as the water returns. In southern France, either the eggs (or 

a proportion of the eggs) of the second generation hatch quickly 

and the larvae spend the winter in diapause, or they do not hatch 

until the following spring, depending on biological and ecological 

factors (Papazian, pers. com.). In northeast Algeria 

(Numidia) 340 , embryonic or larval diapause does not take place; 

the second generation eggs hatch and the larvae undergo an 

autumn and winter growing phase which is prolonged due to the 

low temperatures; in addition, if there is water available in summer 

the second egg laying takes place earlier, and the species 

manages to lay for a third time late in the season; this is also the 

case in southern Europe in ricefields 340 . 

Laying usually takes place in tandem (the male still clasping the 

female). In full flight, the female, who like all Libellulidae has no 

ovipositor, taps the water surface with her abdomen: the eggs 

thus released into the water remain caught up on a support (an 

aquatic plant in most cases), before falling to the bottom. In 

France the larval stage of S. fonscolombii (first generation) is 

among the shortest observed among Odonata in France (along 

with Hemianax ephippiger). In the Mediterranean region, possibly 

after a diapause due to the drying out of the pool or to wintry 

temperatures in the water, the eggs hatch in a short time 

(rapid incubation). Emergence is usually nocturnal, extending 

from March in the Mediterranean region 95 , to November. 

(2) In southern regions, adult S. meridionale may be seen from 

the beginning of June (a late species) to mid-October 95 . At pools 

in northeast Algeria, Samraoui et al. 341 have noted the presence 

of adult S. meridionale around the pools in May-June (immature 

imagos) and in September-October (mature). 

In temporary habitats, the adults have just enough time to 

emerge before the summer drought. They mature over the summer. 

At this time in northeast Algeria 340 , they disperse to refuges 

(woodland) at high altitudes where they slowly develop sexual 

maturity (imaginal diapause); adults may travel up to 35 km to 

aestivate at altitudes greater than 1,000 m. 

After the summer phase of maturation, the adults return, at the 

end of summer or in autumn, to the egg laying sites, which by 

now are usually flooded, to breed. In southern France (Papazian, 

pers. com.), the eggs may either hatch rapidly, the larvae then 

spending the winter in diapause, or delay hatching until the following 

spring. On the other hand in northeast Algeria larval 

102 

development continues until the emergence of adults in the spring 

of the following year; there is no embryonic diapause if water is 

available, but if it happens that the species lays in habitats that 

are still dry, the eggs can then remain resistant to desiccation 

until the first rains 340 . Contrary to S. fonscolombii therefore, 

S. meridionale only has one generation per year in temporary habitats 

(southern France, North Africa) 341 (Papazian, pers. com.). The 

eggs (up to 550) are laid in places with shallow water, among 

reeds and Carex, even in puddles. 

S. meridionale is a migratory species like S. fonscolombii. The 

lifespan of the adults which survive to aestivation is among the 

longest known among Anisoptera. 

Simplified phenological cycle of Sympetrum fonscolombii 



Hydrology 

• Larval development: 

Aguesse 4 showed that laying does not always take place at times 

and places that are favourable for the development of the larvae. 

Among Libellulidae such as Sympetrum, females are frequently 

seen laying eggs on the dry outer edge of a pool, which will not 

be submerged at a suitable time, or on the water surface at a 

pool which dries out a few weeks later. While larval development 

is jeopardized by the failure of the laying site to refill with water 

or by early drying out of the pool, these laying strategies remain 

nonetheless effective overall thanks to egg diapause (facultative), 

which takes place if the site is dry and which enables the 

eggs to survive the dry period. It should be noted that this group 

of Odonata has the widest geographical range 4 . 

• Adult diapause: 

For North Africa, Samraoui et al. 341 described a prolonged, diapaused 

adult phase in S. meridionale (3-4 months during the 

summer dry period), when there is no reproductive activity. Von 

Hagen (in Samraoui et al. 341 ) described the same phenomenon in 

S. striolatum in southern Spain. Papazian (pers. com.) has 

observed it in Corsica and in southern France. 

` 


First generation 

� 

or larvae, followed by larval development 

Second generation 

Emergence of adults, sexual maturation, 

followed by egg laying and migration 

Larval development, 

emergence of adults, migration 

Aestivation and maturation of adults 

Eggs laying and migration 

� 

Diapause of eggs




Both species are fairly common near the Mediterranean, especially 

in the PACA region (more local in the north): 

•Var: Plaine des Maures 244 and lakes of the Centre Var 377 ; 

Sympetrum fonscolombii has in addition been recorded from the 

Colle du Rouet 325 , the Plaine de Palayson 363 , the oxidation ponds 

at Porquerolles and the Quinson reservoir on the Verdon, and 

S. meridionale on the Giens peninsula (Papazian, pers. com.). 

• Bouches-du-Rhône: S. fonscolombii is recorded from the Crau 363 

and the Camargue (Papazian, pers. com.) 

•Hérault: S. fonscolombii is known from the Roque-Haute 

plateau 3 . 

Corsica 

Sympetrum fonscolombii occurs at six out of the 50 stations visited 

during the survey of the dragonflies of the island, and S. 

meridionale occurs at ten of these 50 stations 323 . Notably, both 

are present at Padule Maggiore and Tre Padule. 


IUCN 







(1) The presence of people at water bodies is a constraint on 

Sympetrum fonscolombii, which is very timid 322 . 


(1) Shading of water bodies and the growth of very dense aquatic 

vegetation is probably harmful to Sympetrum fonscolombii. 

(2) Male S. meridionale are often parasitized by mites (Acarus 

libellulae De Geer) 322 . 

- 

- 

- 

- 

- 


Odonata 


In France, some sites are wholly or partly legally protected by 

their Réserve Naturelle status (Roque-Haute, Camargue, Les Tre 

Padule de Suartone); others are included in Natura 2000* areas 

(Colle du Rouet, Plaine des Maures, lakes of the Centre Var, Lanau 

pool), and among these some have been the subject of operations 

of land ownership control in the context of the LIFE “Temporary 

Pools” project (some parts of the Plaine des Maures and the 

Lanau pool). 


While Sympetrum fonscolombii is a common species in the French 

Mediterranean region, S. meridionale is much more confined to 

the coastal fringe, and the survival of these populations therefore 

depends above all on the conservation of coastal pools. 

Bibliography 

AGRN-RH, 2000 3 ; Aguesse, 1961 4 ; Aguesse, 1968 5 ; Anonymous, 

1997 11 ; Corbet, 1999 89 ; D’Aguilar & Dommanget, 1998 95 ; 

Dommanget, 1994 122 ; Heidemann & Seidenbusch, 2002 173 ; Médail 

et al., 1993 244 ; Robert, 1958 322 ; Roche, 1990 323 ; Rombaut, 

1994 325 ; Samraoui & Corbet, 2000 340 ; Samraoui et al., 1998 341 ; 

Terzian, 1979 363 ; Thiéry et al., 2002 377 ; Wendler & Nüss, 1994 401 . 

Authors: Jakob C., T. Gendre & J. Fuselier 

Collaborators: Papazian M., B. Samraoui & N. Yavercovski 

103


Bufo calamita Laurenti 1768 

AMPHIBIA 

BUFONIDAE 

Main synonyms 

None 

French names: Crapaud calamite, Crapaud des joncs 

Spanish name: Sapo corredor 

English name: Natterjack Toad 

Subspecies 

None 


The length varies between 40 and 80 mm (sometimes up to 90 mm 

in the female). The build is sturdy with prominent paratoid 

glands* arranged in parallel, a horizontal pupil and pale greenishyellow 

iris, and a greenish to brownish dorsal surface generally 

crossed by a pale yellow mid-dorsal line 125, 333 . The breeding male 

develops brown nuptial pads on the toes. The species has a characteristic 

mode of locomotion: unlike most amphibians the 

Natterjack does not jump, but walks rapidly or runs. It is easy to 

spot thanks to its far-carrying song, which may be heard from up 

to a kilometre away. 


The Green Toad (Bufo viridis) has no yellow dorsal line; it has 

handsome green markings contrasting with its ash-grey back. 

The Common Toad (B. bufo) has a golden-red iris. 



France 

The Natterjack is found in all parts of France except for the high 

mountains (maximum 1,500 m in the Pyrenees) and Corsica. This 

species extends all the way into built-up areas in some towns. It is 

abundant in the south but its distribution becomes more discontinuous 

towards the north. 


The Natterjack occurs in Spain (except for part of the northwest 

and the Pyrenees) and in Portugal 168 , but is absent from Italy. 

Habitat 

The Natterjack is typically an amphibian of open habitats, with 

short vegetation and few trees. It prefers loose substrates, either 

sandy (sandy back dunes) or stony (Plaine de la Crau, Plaine des 

Maures), degraded garrigue and maquis and also cool heathlands 

(Plateau de l’Aubrac). It adapts to man-made sites: parks and 

wasteland on the edges of built-up areas. Its aquatic habitat is 

fairly varied: live stock-watering ponds, flooded areas among 

sand dunes, puddles, Mediterranean wadis, quarry lakes, etc. 

104 

1 cm 

These habitats are typified by often shallow water (a few tens of 

centimetres at most), an absence of predators, and sparse or even 

completely non-existent aquatic vegetation. 

Ecology 


Bufo calamita 

This is a pioneer species, which is able rapidly to colonise recently 

created habitats such as temporary pools. It prefers loose, sandy 

soils. In search of suitable habitats, Natterjacks may easily cover 

distances of more than 2.5 km 268 . The males may live for seven 

years while females can sometimes live for 17 years 125 . Like almost 

all European amphibians, the Natterjack has terrestrial and aquatic 

phases. Breeding usually starts in March in the Southern France 

with the young toads leaving the water at the start of summer 

(June), extending into July further north. In the south, a second 

breeding season is sometimes observed at the end of summer, 

depending on the storms of August and September. Young frogs 

from this breeding season emerge at the end of September or 

beginning of October. Males form choirs and sing in shallow 

water to attract the females. After mating, the adults quickly 

leave the pool, and the eggs, which are laid in strings, (2500- 

4000 at a time) hatch after a few days. Aquatic larval development 

is relatively brief (four weeks to three months). Their rapid 

development does not always protect the young Natterjacks from 

falling victim to early drying out 184, 185 . The juveniles are very small 

at metamorphosis compared with other species, but grow quickly. 

Sexual maturity is generally attained when they are three years 

old. The tadpoles, which are completely black, may feed on algae 

or other plants as well as detritus. They are sometimes cannibalistic 

268 . Both adults and juveniles are predators of terrestrial 

insects.

Annual phenological cycle 



Normal cycle 

� 

hibernation 

Exceptional cycle 

hibernation 

Arrival 

Spawning 


then metamorphosis 

Departure 


Hydrology 

Flooding must take place at the latest at the beginning of May, 

the limiting date for breeding in the Mediterranean region. 

However, a brief period of flooding (two to five months) does not 

disrupt the process of larval development and the year’s breeding 

success. 

Substrate for spawning 

This is one of the rare species capable of breeding in the complete 

absence of any aquatic vegetation, in concrete-lined ponds, 

rocky wadis or bare sandy areas. 


This species avoids temporary pools used by other species, especially 

the Green Frog (Rana perezi) or the Common Toad; the 

digestive tract of some species contain an alga which inhibits 

the larval growth of Natterjack Toad tadpoles 191 . 


This species is highly adapted to unstable habitats and as a result 

is very tolerant of perturbations in the aquatic environment. Its 

short larval development period, tolerance of high temperatures, 

ability to cover great distances to breed and production of a large 

number of eggs make it a species well-adapted to the risks of 

drying out and for the colonisation of new sites. On the other 

hand, it needs open habitats for feeding and moving around, and 

declines when the habitat becomes wooded. 

Shade 

The Natterjack prefers pools with strong insolation and rather 

short terrestrial vegetation, the larvae being capable of withstanding 

high temperatures, up to 40°C for brief periods. 


� 

Aestivation by adults then terrestrial phase and 

Spawning 


Departure, terrestrial phase and 


The Natterjack Toad is fairly abundant in the Mediterranean 

region, rarer and more sparsely distributed in northern Europe 125 . 

It occurs at about 29% of Réserves Naturelles de France land and 

10% of Réserves Naturelles Volontaires. However, if only those 

Réserves Naturelles located within its French range are included, 

it is present at about 48% of them 135 . 


IUCN 





and Red Lists 


Amphibians 

- 

Annexe II 

Annexe IV 



09/09/1993 

- Spain, national list293 - Portugal, national list59 - Europe: “to watch” 26 

- France, national Red Book: 

“to watch” 242 

- Spain, national Red Book: 

“low concern” (LC) 293 

- Portugal, national Red Book: 

“not threatened” (NT) 59 

The Natterjack Toad is a relatively little-threatened species in the 

Mediterranean region, aside from the impact of increasing amounts 

of woodland. 


The transformation of a shallow temporary pool into a deep or 

even permanent pool poses problems for this species. Such a pool 

can support greater numbers of invertebrate predators than a 

shallow pool, and also enables other amphibian species to breed, 

thereby impeding or preventing spawning among Natterjacks. 

However, it is able to breed in some places alongside the Western 

Spadefoot (Pelobates cultripes), Parsley Frog (Pelodytes punctatus), 

Midwife Toad (Alytes obstetricans), Marbled Newt (Triturus 

marmoratus), Great Crested Newt (T. cristatus), Palmate Newt 

(T. helveticus), Painted Frog (Discoglossus pictus) or Stripeless Tree 

Frog (Hyla meridionalis). This is the case for example at the 

Valliguières pool and the Opoul pool, where temporal segregation 

from the other species allows it to breed successfully. 

The introduction of fish causes the decline and disappearance of 

this species. 

Finally, the degradation, destruction or modification of the terrestrial 

habitat has harmful effects, by destroying the aestivation 

and hibernation sites which are essential for the survival of amphibians 

in the Mediterranean (stumps of trees or bushes, rocks, 

vegetation near the water bodies). 


If the aquatic and terrestrial habitats become overgrown through 

natural succession, this poses problems for pioneer species of open 

habitats like the Natterjack. The increase of terrestrial plant 

cover, in particular, has been proved to be a factor in its decline: 

on a very small scale, plant cover provides shelter, but on a larger 

scale it may form an obstacle to moving around, and a bare habitat 

is greatly preferred (see the Plaine de la Crau, where the species 

reaches extremely high densities!). 


The disappearance of shallow temporary water bodies and the 

destruction of breeding sites increase the risk of local extinction 

as a result of the increasing isolation of habitats. 

105




Recent scrub clearing around shallow pools such as those at 

Roque-Haute (terrestrial vegetation) is improving the quality of 

the habitat and migratory routes. The overdeepening of pools, 

frequently undertaken in the case of pools which have become 

very shallow through aggradation*, has a negative impact on the 

population 56 , as does the creation of permanent pools next to 

shallow temporary pools 35 : both these measures favour other, competing 

species of amphibians. 


In most cases, the reinforcement of populations or the substitution 

of recreated habitats for destroyed habitats are effective 

106 

methods and are often more prudent than deepening the pool. 

Population reinforcement may be achieved by creating a series of 

temporary pools with a range of varying depths around a site 27, 35 . 

Bibliography 

Beebee, 1977 24 , 1983 25 , 1985 26 , 1996 27 , 1997 28 ; Beebee et al., 

1993 29 ; Biggs et al. 2001 35 ; Buckley, 2001 56 ; Cabral et al., 

1999 59 ; Duguet & Melki, 2003 125 ; Fiers, 1998 135 ; Günther, 1996 168 ; 

Jakob et al., 1998 185 , 2003 184 ; Joly, 1994 191 ; Maurin, 1994 242 ; 

Nöllert & Nöllert, 1996 268 ; Plueguezuelos, 2002 293 ; Salvador & 

Garcia-Paris, 2001 333 

Author: Jakob C.

Discoglossus sardus Tschudi, 1837 

AMPHIBIA 

DISCOGLOSSIDAE 

Main synonyms 

None 

French name: Discoglosse sarde 

Italian name: Rana Tirrenica 

English name: Tyrrhenian painted Frog 

Subspecies 

None 


This Painted Frog is a small amphibian of slender appearance with 

a strong resemblance to a frog of the genus Rana. The males, 

which are slightly larger than the females, may reach 5 cm in 

length from snout to cloaca. The colour of the upperside is very 

variable, most often brownish but sometimes reddish, greyish or 

even greenish, more or less heavily marked with irregular dark 

areas all over the body. A pale band runs between the eyes across 

the top of the head. This pattern gives the animal the appearance 

of a dead leaf, which makes it very cryptic on the ground and in 

the water. The hind legs have pale and dark bands. The underside 

is a rather uniform pearly cream colour. The pupil is shaped like 

a water droplet and the eardrum is not visible. 

Eggs are laid in tens at a time, separately or in the form of a 

sheet at the bottom of the water. They are greyish or bicoloured, 

with a diameter of 1 to 1.5 mm, and are contained within a 3 to 

4 mm gelatinous capsule. 

The tadpoles are small and dark-coloured, fairly similar to Bufo 

tadpoles. Like all the Discoglossidae, they have a ventral spiracle*. 


The Tyrrhenian Painted Frog forms a species pair with the Corsican 

Painted Frog, with which it was confused until 1984. Compared 

with this species the hind leg is shorter: if extended along 

the body, the heel usually reaches as far as between the eye and 

the tip of the snout, but no further. The snout is more pointed 

and the part between the eye and the nostril is slightly sloping. 



This Painted Frog is restricted to the French and Italian Tyrrhenian 

islands: Sardinia and some of its islets, Corsica and the Lavezzi 

Islands, the Tuscan Archipelago (Giglio and Montecristo Islands), 

the island of Monte Argentario, and the eastern Hyères Islands 

(Port-Cros and Levant). This isolation is undoubtedly very longstanding, 

such that each of these populations has evolved distinctive 

and specific characteristics. In Corsica, it ranges over a good 

part of the island, from sea level up to at least 1300 m. At Port- 

Cros it is found in all the streams on the island, notably the Vallon 

de la Solitude and the Vallon Notre-Dame. It seems to be rare on 

the Île du Levant where surveys are still required to ascertain its 

precise status. 

Amphibians 

Habitat 

This species lives in a wide range of habitats, from low maquis to 

mature oak or pine woodland. It is undemanding in its choice of 

breeding sites: springs, coastal marshes, canals, brooks and streams, 

depressions cut off from the main channels of rivers, reservoirs 

and fountains. In particular it uses impermanent bodies of water 

which in coastal regions are fed by unpredictable events (isolated 

puddles, temporary runnels, temporary pools, etc.). It is not uncommon 

for the eggs to fail to hatch, or to be dried out by evaporation 

or carried away by floods. This species can breed in relatively 

brackish water and, in contrast with the Corsican Painted Frog, 

can adapt to disturbed or modified habitats (polluted streams on 

the edges of villages, artificial reservoirs). Outside the breeding 

season, the species may be observed under piles of leaves or under 

trunks of fallen trees, several hundred metres from the nearest 

aquatic habitat. 

Ecology 

Discoglossus sardus 


1 cm 

This species is active for a good part of the year, by day as well 

as by night, but comes out mainly during wet weather in autumn 

(October-November) and at the end of winter (February-April). 

Breeding may be observed from October to April, usually following 

heavy rain (two to three nights after the rain starts). The males’ 

song is quiet, not very far-carrying and restricted to the breeding 

seasons. Males arrive at the breeding sites first and remain there 

longer than the females. Mating is brief and the eggs are laid in 

bundles of 20 to 50. The small tadpoles hatch out after about 

three days. These tadpoles grow rapidly and metamorphose after 

four to six weeks. Some, originating from eggs laid in autumn, 

grow much more slowly (20-25 weeks) during the winter. In 

Corsica as well as at Port-Cros, the eggs are laid over a very long 

period of time. Several generations of tadpoles coexist in a single 

place and it is common to see eggs, tadpoles of all sizes and 

juvenile frogs all present simultaneously. 

At metamorphosis individuals are very small (about 8 mm) and 

resemble adults in every respect. They reach 16 mm a year later, 

30 mm after two years and 45 mm at the end of the third year. 

The lifespan may certainly extend to nine or ten years. 

The diet consists of small invertebrates caught on land: woodlice, 

spiders and various insects. 

107





Usual cycle 

Occasional cycle 

� 


Hydrology 

This species is perfectly adapted to the unpredictability of the Mediterranean 

hydrological regime (very extended breeding period, 

rapid development of larvae) reducing its vulnerability to climatic 

hazards. 

Aquatic habitat 

This Painted Frog uses almost all types of aquatic habitat for 

breeding, with the exception of flowing water and very large water 

bodies. This species ranges widely over the countryside, without 

specialising in any one type of habitat. Pools are therefore not 

essential for the survival of its populations, in contrast with other 

amphibians. 

A slight degree of salinity is tolerated, as well as a high degree 

of turbidity (muddy puddles). Organic pollution does not seem to 

have any affect on breeding. In general, this species uses small 

temporary water bodies, preferably exposed to sunlight, although 

a fairly large amount of plant cover may be tolerated. 

Terrestrial habitat 

Almost all kinds of habitat may be used: dense woodland, maquis, 

or partly built-up areas. 



Most of the populations are large, except on some islets which 

have practically no fresh water such as the Lavezzi islets. Corsica 

undoubtedly supports the largest numbers. In Sardinia this species 

seems to be less common and it is only known there from a small 

number of sites. There are no quantitative data available for the 

Tuscan archipelago, but according to Knoepffler (1962), the islands 

of Giglio and Montecristo have very large populations due to the 

presence of abundant fresh water. 

On Port-Cros, Knoepffler estimated the population size at around 

5000 adults at the end of the 1950s. Recent observations indicate 

that numbers may be lower at present. The introduction 

some years ago of fish into the artificial reservoir where a high 

proportion of the island’s breeding adults were concentrated had 

reduced the breeding capacity of the population. The eradication 

of the fish in 2001 made the site once again available for the 

Painted Frogs, which bred there in numbers in 2002-2003. 

(Joyeux, pers. com.). On the Levant Islet, Knoepffler 198 considered 

108 

Extended egg laying 

Larval growth and metamorphosis 

Larval growth and metamorphosis 

Departure and 

terrestrial phase 

� 

Egg laying (sometimes) then 

the species to be common during the 1950s-60s, but threatened 

by the deforestation of the island, the destruction of watering 

places and the impact of the Viperine Snake on the larvae and 

the adults. No surveys have been carried out since then to assess 

the current status of this population. Most of the occupied sites 

are situated on steep islands that are not very suitable for agriculture 

and are well preserved overall. 


IUCN 





and Red Lists 



Owing to its lack of specialisation, the threats to this species are 

not very serious. Dams interrupt the distribution of populations 

and reduce the number of breeding sites; water pollution undoubtedly 

affects the eggs and larvae. One threat which cannot be 

ignored is that of forest fires, which as well as destroying adults 

have the effect of profoundly altering the hydrological regime of 

the watercourses (more torrencial regime) and the nature of the 

breeding sites (input of ash and sediment). Coastal populations 

are no doubt affected by anti-mosquito treatments (Corsica) and 

by the spread of the Italian Pool Frog (Rana bergeri), introduced 

probably recently into Corsica. The introduction of fish into closed 

water bodies (small reservoirs, ponds) is recognised as constituting 

a threat (for example the case of the Parc de Port-Cros). It is 

acknowledged that the introduction of alien competitive or predatory 

species in particular should be controlled (American crayfish, 

fish, frogs), as these will undoubtedly have catastrophic effects 

on this species. 


Natural threats are few in number apart from the problem of fire, 

which may in some cases constitute a serious threat (small wooded 

islands such as Port-Cros). Reafforestation is known to constitute 

a threat to some populations by its effect on the breeding sites 

(shading of open water, and reduction of open water due to the 

accumulation of litter). 


It is necessary to maintain sufficient numbers to counter the risk 

of chance extinctions, which is particularly high in an island situation. 


- 

Annexe II 




09/09/1993 

- Italy, regional lists 

(Sardinia, Tuscany): 

regional law of 29/07/1998 


Book: “rare” 242 

- Italy, national Red Data Book14 Current measures 

The Tyrrhenian Painted Frog occurs at many sites in France and 

Italy which benefit from official protection either specifically for

wildlife (Parc National de Port-Cros, Nature Reserves of Etang de 

Biguglia, Scandola and Bouches de Bonifacio in Corsica, Parc 

Naturel Régional de Corse, Tuscan Islands National Park) or as a 

result of laws relating to land use (in France, sites belonging to 

the Conservatoire de l’Espace Littoral et des Rivages Lacustres 

and Forêts domaniales de Corse). 

The inclusion in the Natura 2000* Network of most of these areas, 

where the conservation of the species is usually taken into account 

in the management plan, is recommended. The species is the subject 

of research in the Parc National de Port-Cros and in Corsica. 

An inventory of the populations of the Ile du Levant is planned. 


• It would be useful to clarify the status of some populations 

which are still poorly known: the islets of the Tuscan archipelago 

(Italy) and the Ile du Levant (France). 

• On Port-Cros, it would be worthwhile to contain the growth of 

forest cover around the main breeding sites by selective streamside 

felling. In addition, it would be worth reassessing the dry 

Amphibians 

stone dams along the watercourses, which were constructed in 

the early years of the Park to hold back sediment: the piles of 

leaves accumulated over the years have had the effect of blocking 

the beds of the island’s streams, tending to cause the water to 

run underground, with consequent losses of the amount of open 

water available for use by the aquatic fauna. The number of 

breeding sites has decreased significantly as a result. 

Bibliography 

Anonymous, 2004 14 ; Capula & Corti, 1993 61 ; Cheylan & 

Massemin, 1999 73 ; Cheylan, 1983 75 ; Delaugerre & Cheylan 

1992 103 ; Delaugerre 1999 104 ; Gasc et al., 1997 152 ; Knoepffler, 

1962 197 , 1973 198 ; Maurin, 1994 242 ; Pesme, 2001 287 ; Salvidio et 

al., 1997 335 , 1998 337 , 1999 336 ; Vandenbrouck, 1996 393 

Authors: Cheylan M. & M. Delaugerre 

109


Pelobates cultripes (Cuvier, 1829) 

AMPHIBIA 

PELOBATIDAE 

Main synonyms 

None 

French names: Pélobate cultripède, Crapaud à couteaux 

Spanish name: Sapo de espuelas 

Portuguese name: Sapo-de-unha-negra 

English names: Western Spadefoot, Iberian Spadefoot Toad 

Subspecies 

None 


The Western Spadefoot is a fairly large amphibian (10-11 cm) 

with a squat appearance. The head is short and surmounted by 

large protuberant eyes. The eardrum is not visible. The skin is 

smooth, and variable in colour from greenish-yellow to brown, 

strongly marked with more or less contiguous brown blotches, 

giving each individual its own unique pattern. In the field it can 

be immediately recognised by its upright stance. There is little 

sexual dimorphism, apart from the presence in the male of a yellow 

lens-shaped protuberance on the forelegs. On the heel there 

is a down-turned black spade used for digging in the soil. The 

tadpoles, which are very large when fully-grown (up to 12 cm) 

are distinguished by a bulky oval body and a lead-grey to greenish 

colouration, with fine golden markings. The caudal crest* is lanceolate 

shaped, pointed at the tip and practically unmarked. When 

they leave the water the young measure on average 25 mm and 

weigh 1.5 to 2 g. The spawn takes the form of a long string, 12 

to 20 mm in diameter and 1.1 metres long on average. Spawn 

from different females is often mixed up, tangled together at the 

edge of a water body, usually caught up in vegetation 0 to 20 cm 

deep. It may be distinguished from the spawn of “true” toads by 

the irregular distribution of the eggs within the string and by the 

larger number of eggs. 


Impossible to confuse with other species, except in the case of 

the tadpoles. 



This Spadefoot is found in the Mediterranean regions of southwest 

Europe: Spain, south and southwest France, as well as in Portugal. 

In France the populations form two distinct nuclei: a Mediterranean 

nucleus which extends eastwards to the Fréjus area in 

the Var département and to the Ardèche in the Rhône Valley, and 

an Atlantic nucleus extending from the mouth of the Gironde to 

the Guérande peninsula in the département of Loire-Atlantique. 

Habitat 

The Spadefoot especially prefers open habitats: dry plateaux, 

sandy areas, degraded garrigue and maquis, grazed meadows and 

110 

1 cm 

Mediterranean arable land, up to 800 metres altitude in southern 

France and 1,700 m in the Iberian Peninsula. In Spain it is common 

in the dehesa* country typical of southwest Iberia and around 

the mouths of rivers (Marismas del Guadalquivir). 

The breeding sites are fairly varied: permanent or temporary pools, 

dune slacks, marshland ditches, concreted lavognes*, gravel pits, 

hill reservoirs. 

Ecology 


Pelobates cultripes 

Like most Mediterranean amphibians, the Western Spadefoot is 

active especially in winter and spring, from November to May in 

southern Spain, from March to June and then from the end of 

August to November in France. Strictly nocturnal, they are especially 

active during rain, whenever the temperature reaches at 

least 8°C. In the daytime the animal remains buried in the soil at 

a depth of 6 to 20 cm, up to 40 cm or deeper in winter. 

The breeding season varies depending on the local climatic conditions. 

It extends from October to February in the south of the 

Iberian Peninsula and from the end of February to the beginning 

of May in central Spain. In southern France the breeding cycle is 

markedly bimodal with one more or less consistent breeding period 

in autumn (September to November) and another in spring, usually 

between the third week in February and the end of April, 

sometimes to the beginning of May. The onset of this event is 

triggered by continual rain over several days (20 mm), with an air 

temperature of 9 to 16°C. The breeding adults arrive at the 

breeding sites at dusk and remain under a few centimetres of 

water immediately next to the water’s edge. The largest males 

arrive first, sometimes before the females, and remain for one to 

eight nights. The females usually stay for one to three nights at 

the breeding place. During amplexus*, which lasts an average of 

72 hours, the male and the female both give a call which resembles 

the clucking of a chicken. 

In Spain, the average number of eggs laid is 2318, while at a site 

at the Ile d’Oléron it is 960 eggs. The strings of eggs are wound 

around the vegetation and are laid in shallow water, usually less 

than 20 cm below the surface. The tadpoles feed mostly on plant 

material, both algae and higher plants, but also detritus. When 

food becomes scarce they may eat the tadpoles of other species 

or of their own species. Larval development usually takes three

and a half months for eggs laid in spring, up to eight months for 

autumn eggs. Metamorphosis takes place mostly in June and July. 

Some tadpoles do not metamorphose in summer and are still in 

the water in autumn. In this case, they probably do not undergo 

metamorphosis until they are one year old. 

The diet of the adults consists entirely of invertebrates: spiders, 

harvestmen, beetles, bugs, hymenopterans, butterflies and moths, 

grasshoppers, diptera, ants and insect larvae. 

Some individuals may live for 10-15 years; sexual maturity 

appears to develop at two years old. They have many aquatic 

predators: dragonfly larvae, water beetles, water snakes, herons. 




Usual cycle 

Occasional cycle 

� 

metamorphosis 

Arrival 

Spawning 

Larval development and metamorphosis 


Departure, aestivation and terrestrial phase 

Arrival 

Spawning 

Hydrology 

The length of the larval cycle constitutes a major constraint for 

this species, particularly in view of the unpredictability of the 

Mediterranean climate. Many breeding attempts are therefore 

doomed to failure, whether due to the breeding site drying out 

earlier than usual or to delayed flooding. Ideally, the site should 

start to fill with water at the end of October and remain flooded 

until the end of July. 


It is usually characterised by fairly deep water, most often deeper 

than 70 cm. The spawn is laid preferably on submerged vegetation, 

very close to the water’s edge, in water containing not more 

than 5 g of salt per litre (the tadpoles will tolerate slightly higher 

salinities, up to 10 g.l -1 ). The breeding sites are typically poor in 

aquatic vegetation: the highest densities of tadpoles are found in 

sites with hydrophyte biovolumes of less than 10 kg/m 3 . 


The presence of open areas appears to be a major factor. Otherwise 

the habitat may be very varied: coastal dunes, dry grasslands, 

open heathlands, garrigue, oak or pine woodland, dry plateaux, 

arable land, on sandy, stony or even rock substrates. In sandy 

areas, the presence of rabbits appears to be favourable for the 

Western Spadefoot: it uses their burrows for shelter in the daytime. 

Grazing, especially by sheep, is also beneficial. 

� 

Larval development and 



Amphibians 

The Spadefoot is currently declining over the whole of its range. 

This decline is due to the destruction of its breeding sites or terrestrial 

habitats, the widespread use of pesticides, and the 

increasing fragmentation of its populations. 

In France, the total number of known sites is not greater than 150. 

This species is particularly threatened at the edges of its range, 

especially in the Var département where only two out of the four 

historically known sites still remain, but also in the Bouches-du- 

Rhône, the Vaucluse and all the Atlantic coastal départements. 

Most of the breeding sites hold fewer than 100 individuals, a few 

tens being the norm. Six Conservatoire de l’Espace Littoral et des 

Rivages lacustres sites and four Nature Reserves support the 

species. 

In Spain, the situation is a cause for concern in most provinces. 

The decline seems to have been particularly steep over the course 

of recent decades. In many cases, the survival of the species 

depends on the preservation of a single breeding site. 


IUCN 





and Red Lists 


- 

Annexe II 

Annexe IV 



09/09/1993 

- Spain, national list293 - France, national Red Data Book: 


- Spain, national Red Book: 

“almost endangered” (NT) 293 

- Portugal, national Red Book: 

“not threatened” (NT) 59 


Impacts of an anthropogenic nature include the destruction of 

both terrestrial habitats (conversion to arable, urban development) 

and aquatic habitats (infilling of pools), the introduction of competitive 

or predatory alien species (fish, American crayfish, Marsh 

Frogs), the use of toxic chemicals (pesticides) and the destruction 

of the animals (road traffic). Coastal populations are especially 

affected by tourism developments and by anti-mosquito treatments. 

Further inland, it is the abandonment of farmland and the spread 

of woodland which are the two principal causes of decline. 


The main threats are due to weather conditions, either when the 

breeding site dries out before the tadpoles have metamorphosed, 

or when a sudden dry period lowers the water level to below the 

strings of spawn, which are usually laid at water level at the water’s 

edge. Some predators (notably Polecats) can have a major impact 

on the adults during the breeding season. 

111



Essentially, it is necessary to maintain high enough numbers to 

overcome stochastic risks, as well as a dense enough network of 

connections between populations to ensure interchanges of individuals 

and hence gene flow. Distances of less than a kilometre 

between sites would appear to be desirable. 



In France, if some of this species’habitats are situated in protected 

areas, the species is not the subject of specific conservation 

measures, apart from the monitoring protocols set up at some 

sites in Charente-Maritime, in the Camargue (Tour du Valat estate), 

at the Valliguières site in the Gard département, in the Roque- 

Haute Nature Reserve in the Hérault, in the Parc du Lubéron in 

the Vaucluse and at the Suze-la-Rousse site in the Drôme. 


It would be helpful, in the first place, to maintain the spatial 

cohesion of the breeding sites, if necessary by recreating suitable 

sites in recently abandoned areas. This could be attempted experimentally 

at a few sites: the Giens peninsula and the Plaine de 

Palayson in the Var, the Vallée du Calavon in the Vaucluse, the 

Plateau d’Aumelas in the Hérault. 

In addition, it would be useful to carry out some activities at certain 

breeding sites: providing information for owners and communes, 

112 

setting up management agreements, etc. As far as management is 

concerned, the most important actions are the following: to control 

the introduction of undesirable species (Louisiana Crayfish, 

fish, terrapins etc.), to ensure that the hydroperiod* is compatible 

with the species’ breeding cycle, to control plant cover over an 

area of radius 100 to 200 m by grazing and/or scrub clearing, and 

to place shelters by the edges of the pools to promote the survival 

of the newly metamorphosed young. 

Bibliography 

Alvarez et al., 1990 8 ; Busack & Zug, 1976 57 ; Cabral et al., 1999 59 ; 

Cei & Crespo, 1971 65 ; Cheylan & Poitevin, 1999 74 ; Conservatoire 

de l’Espace Littoral et des Rivages Lacustres, 1998 84 ; Diaz-Paniagua 

& Arrizabalaga, 1987 111 ; Diaz-Paniagua, 1983 112 , 1985 113 , 1986 114 , 

1988 115 , 1990 116 ; Dohogne, 1999 120 ; Doménech, 1994 121 ; Duguet & 

Melki, 2003 125 ; Fiers, 1998 135 ; Garcia-Paris, 1990 147 ; Gasc et al., 

1997 152 ; Lescure, 1984 209 ; Lizana et al., 1994 214 ; Maurin, 1994 242 ; 

Nöllert & Nöllert, 1995 267 ; Petit & Lomont, 1958 288 ; Pleguezuelos 

& Lizana, 2002 293 ; Rodriguez, 1988 324 ; Salvador & Garcia-Paris, 

2001 333 ; Salvidio & Quero, 1987 334 ; Thirion, 1996 378 , 2001 379 . 

Author: Cheylan M.

Triturus cristatus (Laurenti, 1768) 

AMPHIBIA 

SALAMANDRIDAE 

Main synonyms 

None 

French name: Triton crêté 

Italian name: Tritone crestato 

English names: Great crested Newt, Warty Newt 

Subspecies 

None 


Large newt (total length of adults: 19 cm), dark brown or almost 

black in colour, finely spotted with white on the flanks. The underside 

is yellow/orange, marked with black. The skin is very characteristically 

warty. During the breeding season, the male develops 

a high, spiky dorsal and caudal crest. This has a silvery streak 

along its side. Females and juveniles have an orange dorsal line. 

The throat is greyish, spotted with white. 


• Blasius’ Newt (originally described as a separate species, Triturus 

blasii) is a hybrid between the Great Crested and Marbled 

Newts. It is distinguished from the Great Crested by its ventral 

surface, which is greenish black above and yellowish with black 

markings and fine white spots below. 

• The range of the Italian Crested Newt (T. carnifex) is almost 

completely separated from that of the Great Crested Newt. Its feet 

are longer, its body is more elongated and its upperside is paler, 

showing many black marks; the male’s crest is lower and less 

deeply indented. 



France 

This species is found throughout the northern half of France, 

north of a line from Charentes to Savoie. In the southern half of 

France it is currently known only from a few sites: one at Arles, 

Bouches-du-Rhône, in the Gard 51 , two in the Drôme and one in 

the southern Ardèche. 

Other countries 

The range extends from Great Britain in the west to the Ural Mountains 

in Russia and, north-south, from southern Scandinavia to 

Austria and northern Romania. This species is absent from Spain 

and Portugal. 

In the southern Mediterranean countries, it is replaced by two 

vicariant species formerly treated as subspecies of Triturus cristatus: 

• T. carnifex (Laurenti, 1768) in Italy and the Balkans; 

• T. karelinii (Strauch, 1870) by the northwest coasts of the 

Aegean Sea (Turkey, Greece). 

1 cm 

Habitat 

Amphibians 


The Great Crested Newt is a species of lowlands which lives mainly 

in fish-free permanent pools and/or temporary pools which dry 

out in summer. 

The three sites in the Mediterranean region take the form of: 

•a marlpit in a calcareous area covered with pine woods, garrigue 

with Holm Oak, heathland with juniper and vineyards 

(Valliguières in the Gard, see vol. 1, box 26), 

•a small quarry pool fringed with Yellow Flag Iris and reeds 

(Peyreguil, in the Gard), 

•a marshy area with willows and reeds, surrounded by built-up 

areas and degraded (Trinquetaille, at Arles). 

Ecology 


Triturus cristatus 

male 

In the Mediterranean region, the only information concerning 

the species’ life cycle comes from the Etang de Valliguières. At 

this site the aquatic phase is variable in length between years, 

depending on the flooding period. In “normal” conditions, the 

newts arrive at the pool in November/December and leave again 

around May. The length of time spent in the water is about 80- 

90 days 33 . 

The adults feed mostly on aquatic and terrestrial insects 19 . The 

larvae feed on aquatic prey (especially microcrustaceans). 

Breeding begins in April-May. The female lays 200 to 500 eggs 

per year 248 , placed one by one underneath the leaves of submerged 

plants which she then folds up with her hind feet (mainly 

species with broad leaves such as Ranunculus gr. peltatus, Glyceria 

fluitans, etc.). Embryonic development takes from two to 

three weeks depending on the water temperature. It is followed 

by a two- or three-month larval stage which ends with metamorphosis, 

usually at the end of the summer (mid-July to August, 

exceptionally September if the pool has not completely dried up). 

The young newts then leave the water and seek out a damp 

refuge in the surrounding area, where they hide and feed. After 

two years, in optimal conditions, they attain sexual maturity 

(males usually one year earlier than females 140 , and return to the 

water to breed. Immature T. cristatus tend to appear in the pool 

later than the adults. Adult survival is fairly high 247 : between 60 

and 75% depending on the year. The maximum lifespan in the 

wild is from 14 to 18 years. 

113


The Great Crested Newt is known for its fairly strong fidelity to 

breeding sites, local variations in breeding success being compensated 

for by its long life expectancy. The newts do not appear 

to colonise new breeding sites more than 400 m away from their 

natal pool 20 . Juveniles may disperse for up to a kilometre 15, 202 . 

During the terrestrial phase, the individuals hide underneath 

stones, under pieces of wood or in crevices in the ground, a few 

hundred metres, or further, from the breeding sites. Fidelity to 

terrestrial sites appears to be linked to the different “scent complexes” 

which distinguish them, orientation being carried out by 

means of olfaction 170 . 




� 

overwintering (aquatic phase) 

? 


Hydrology 

Successful breeding depends on the presence of water up to the 

end of July, which is essential for the metamorphosis of the juveniles. 


The site must be relatively large and deep (>1 m) open to the sun 

and lacking fish. The presence of macrophytes* is essential for 

egg laying and for the development of an abundant invertebrate 

fauna; however, vegetation should not cover more than 20% of 

the water surface (maximum tolerated by breeding individuals) 88 . 


The presence of wooded areas immediately adjacent to the site 

appears to be necessary for the survival of adults (and newly 

emerged young) during the terrestrial phase 188 . The presence of 

natural refuges (logpiles, piles of stones, stone walls) is also a 

favourable factor. Ditches and hedges provide biological corridors 

which facilitate dispersal. 



Due to their isolation, the populations of Mediterranean southern 

France are highly endangered in the short term. The site at 

Trinquetaille (Bouches-du-Rhône) is situated within very degraded 

semi-urban surroundings. The population at Valliguières (Gard) has 

114 

Eggs laying 

Larval growth 

Larval metamorphosis 

Departure, terrestrial phase and aestivation 

Arrival and 

(juveniles, adults) 

� 

been subject to conservation measures for three years but its isolation, 

low numbers (about 200 individuals) and irregular breeding 

mean that its chances of survival in the medium to long term 

are questionable. The population at Peyreguil (Gard) is still poorly 

known. The presence of fish here constitutes a serious threat. 


IUCN 





and Red Lists 



The destruction of breeding sites (filling-in of pools), changes to 

the farmed environment resulting from new management practices, 

the isolation of different populations through habitat 

fragmentation 203 , and the introduction of competitive alien species 

(Marsh Frogs, fish, American crayfish) are the main causes of 

decline. 


The repeated drying-out of breeding sites may eventually 

threaten the survival of the population 161 ; the effect of wild boar 

during the emergence of the young newts also represents a 

threat. 


- 

Annexe II 




09/09/1993 


Book: “vulnerable” 242 


The population of the Etang de Valliguières has been monitored 

since 2000 216 as part of the LIFE “Temporary Pools” project. 

Among the planned work (see vol. I, Box 50) are included: 

• Digging out the pool in order to extend the period of flooding 

and thereby to encourage more regular breeding. 

• The creation of an artificial pool close to the existing site, in 

order firstly to encourage more regular breeding and secondly to 

create a second breeding site with the aim of improving the 

probability of survival of the population. 

• The control of invasive species (fish, crayfish, Marsh Frog). 


• Produce a complete inventory of the sites used by the species 

in the Mediterranean region. 

• Establish a network of potential breeding places in close proximity 

to one another (a few tens or hundreds of metres). 

• Control introduced species. 

• Maintain “biological corridors” around the pools. 

• Educate the public so that they do not fill in or bank up pools. 

• Control the impact of wild boar. For this, various solutions may 

be considered: installing electric fences around all or part of the 

site during the period when the young are emerging, digging out 

a substitute pool, planting bramble bushes around the pool to 

increase the number of refuges for the young newts.

• Urgently introduce conservation measures at the Trinquetaille/ 

Arles site in the Bouches du Rhône (translocation of individuals 

to a more suitable site could be considered). 

Bibliography 

Arntzen & Teunis, 1993 15 ; Avery, 1968 19 ; Baker & Halliday, 

1999 20 ; Besnard, 2002 33 ; Brogard et al., 1996 51 ; Cooke et al., 

Amphibians 

1994 88 ; Francillon-Vieillot et al., 1990 140 ; Griffiths & William, 

2000 161 ; Hayward et al., 2000 170 ; Jehle, 2000 188 ; Kupfer & Kneitz, 

2000 202 ; Laan & Verboom, 1990 203 ; Lombardini et al., 2002 216 ; 

Maurin, 1994 242 ; Miaud, 1990 247 , 1991 248 . 

Authors: Lombardini K. & M. Cheylan 

115


Triturus marmoratus Latreille 1800 

AMPHIBIA 

SALAMANDRIDAE 

Main synonyms 

None 

French name: Triton marbré 

Spanish name: Triton jaspeado 

Portuguese names: Tritão-marmorado, Tritão-verde 

English name: Marbled Newt 

Subspecies 

T. m. pygmaeus in the Iberian Peninsula, recently considered to be 

a species in its own right. 

333, 405 


Large newt (total adult length: 16 cm) black to brown in colour 

with green marbling on the back. Females and juveniles have an 

orange-red line down the spine. In the breeding season, the male 

develops a smooth crest with vertical black and yellowish-white 

bars. The underside is black, dotted with small white spots 

(orange-yellow with large black markings in the Great Crested 

Newt). 


Pygmy Marbled Newt (Triturus pygmaeus), Blasius’ Newt (T. blasii), 

Great Crested Newt (T. cristatus). 

The Great Crested Newt differs in its orange underside marked 

with black. Blasius’ Newt is a hybrid between T. marmoratus and 

T. cristatus. It may show morphological characters of both species, 

but it is completely absent from the Mediterranean region. The 

Pygmy Marbled Newt, an endemic species of southern Spain, has 

a black underside like the Marbled Newt, but is smaller. 



France 

This species occurs in the western half of the country, as far 

north as the Seine. The eastern limit of its range lies in the Sommières 

area, in the Gard département. 


Iberian Peninsula, north of the river Tagus, from the coast up to 

2,100 m in the central mountains. 

Habitat 

The Marbled Newt, during its aquatic phase, is found in small 

bodies of still water, lacking fish: Mediterranean temporary pools 

(code 3170), shaded woodland pools, sometimes fed by springs 333 . 

The terrestrial habitat is varied: arable land or pasture, open grassy 

areas (Causses) and woodland. 

116 

Ecology 


Like almost all the European amphibians the Marbled Newt has 

a bimodal cycle, with a terrestrial phase and an aquatic phase. 

At all stages of development, they are carnivorous. The larvae 

feed mostly on aquatic invertebrates 405 , the adults and juveniles 

are predators of tadpoles and of aquatic and terrestrial insects. 

The lifespan is long, up to 14 years in the wild 345 . 

In Mediterranean pools, the aquatic phase occurs mainly in winter 

owing to the unfavourable condition of the terrestrial habitat 

in summer. In southern France the period for which they are in 

the water extends from mid-November to mid-June, with considerable 

variation between sites and years. 

Breeding takes place in February-March in the south of France, 

October to May in the Iberian Peninsula. This phenology is slightly 

variable depending on interannual variations in the flooding 

period. Larval development takes a relatively long time (two to 

three months 140 ). A single female may take several weeks to lay 

all her eggs. The eggs are laid one by one, in a fold in a leaf of an 

aquatic plant, with a preference for species with broad, flexible 

leaves such as Mentha pulegium (Pennyroyal), Ranunculus spp. or 

Callitriche sp. 9 . 

The terrestrial habitat in evidently very important for the aestivation 

of amphibians in the Mediterranean as the animals run 

the risk of desiccation. Juveniles and adults hide under stones or 

among the roots of bushes. They do not move far, travelling 

partly underground during the summer drought. 



� 

Hibernation 

1 cm 


male 

Egg laying 


Triturus marmoratus 

Larval metamorphosis 

Departure, then terrestrial phase and aestivation 

(juveniles, adults) 

� 

Arrival and feeding 

(juveniles, adults)

183, 184, 405 


Hydrology 

Breeding sites must be holding water by February at the latest to 

allow the eggs to be laid. A flooding period of five months is then 

necessary for completion of larval development and successful 

breeding. Breeding failure in a given year does not constitute a 

major problem for the population, unless it recurs too frequently. 


The water should be of a reasonable depth (about 0.8 m) and 

provide sufficient growth of aquatic vegetation to support the 

development of sufficient invertebrates for the feeding requirements 

of the larvae and adults. 


It may be very varied in character (open heathland, garrigue, oak 

or pine woodland, arable or even built-up areas). Hedges and copses 

around the edge of breeding sites provide important terrestrial 

habitats and migration routes 135 . 



In Spain there are large populations which are relatively stable, 

apart from in the Ebro Valley and in north Catalonia where it 

appears to be more threatened. In France, its status gives no cause 

for concern although some local disappearances have been 

recorded, due mainly to the filling-in of pools or to land consolidation 

(destruction of hedges). 

It occurs at about 13% of Réserves Naturelles and 3% of Réserves 

Naturelles Volontaires. However, if only those Réserves Naturelles 

located within the newt’s French range are included, it occurs 

in 42% 182 . 


IUCN 





and Red Lists 

- 

Annexe III 

Annexe IV 


of 22/07/1993 Journal Officiel 

09/09/1993 

- Spain, national list293 - Portugal, national list59 - France, national Red Data 

Book: “vulnerable” 242 

- Portugal, national Red Data 

Book: “not threatened” (NT) 59 

- Spain, national Red Data 

Book: “low concern” (LC) 293 


Amphibians 


Impacts of an anthropogenic nature include the destruction of 

habitats, both terrestrial (grubbing out of hedges and woods, 

destruction of old stone walls) and aquatic (filling-in of pools), 

the introduction of competitive or predatory alien species (fish, 

American crayfish, Marsh Frog (Rana ridibunda), and destruction 

of the animals (road traffic). 


Too short a period of flooding (i.e. through silting up) may prevent 

the completion of larval development, which will jeopardise 

the presence of the newt at a site. 


The destruction of breeding sites and terrestrial habitats brings 

the risk of extinction by reducing the interconnections between 

populations. 



No specific measures are currently being applied at the Roque- 

Haute Nature Reserve (Hérault, France). The effect of scrub clearing 

around the pools (terrestrial vegetation) on the survival of adults 

and their migration routes remains to be assessed. 


• Maintain the cohesion of populations at the local and regional 

scales. 

• Control the introduction of fish and American crayfish into 

breeding sites. 

• Maintain well-developed aquatic vegetation. 

• Do not burn the area around pools or the sites used as refuges 

in the terrestrial phase. Dry years, when breeding does not take 

place, do not present real problems for this long-lived species, as 

long as the aestivation refuges enable good numbers of adults to 

survive. The maintenance of vegetation corridors around the 

pools is strongly recommended. 

Bibliography 

Alvarez et al., 1989 9 ; Cabral et al., 1999 59 ; Fiers, 1998 135 ; 

Francillon-Vieillot et al., 1990 140 ; Gasc et al., 1997 152 ; Hilton-Taylor, 

2000 176 ; Jakob, 2001 182 ; Jakob et al., 2003 183, 184 , 1999 186 ; Jehle & 

Arntzen, 2000 187 ; Maurin, 1994 242 ; Pleguezuelos et al., 2002 293 ; 

Salvador & Garcia, 2001 333 ; Schoorl & Zuiderwijk,1981 345 ; Zuiderwijk, 

1997 405 . 

Author: Jakob C. 

Collaborator: Cheylan M. 

117

Glossary 

Actinomorphic: (= regular) describes a flower, or a group of 

identical organs within a flower, having radial symmetry. 

Active limestone: very fine particles of limestone in the soil, 

which are easily dissolved by the action of carbon dioxide. The limestone 

ions thus released are capable of being absorbed by plants. 

Aggradation: Situation resulting from gradual infilling by accumulated 

material (soil, silt, sand, gravel, etc.). 

Allogamous: having a mode of sexual reproduction involving 

cross-fertilization among plants. 

Amplexus: among amphibians, mating without internal fertilisation. 

The eggs and sperm are emitted simultaneously and fertilisation 

is external. Depending on the species, amplexus may be 

lumbar (the male clasps the female around the lower part of the 

abdomen) or axillary (the male clasps the female under the 

armpits). 

Anemochory: mechanism of dispersal of seeds, spores, eggs, etc. 

of certain animal and plant species by the wind. 

Anemochory: mechanism of dispersal of seeds, spores, eggs, etc. 

of certain animal and plant species by the wind. 

Antheridium: organ in which the male gametes are produced, in 

algae, mosses, liverworts and ferns. 

Apoendemic: describes a new plant taxon which has been produced 

in a given region by polyploidisation (by becoming polyploid*) 

from diploid taxa* which are more or less widely 

distributed in the neighbouring regions. 

Association (phytosociological): see volume 1, chapter 2, box 4. 

Attenuate: width gradually decreasing (tapering). 

Auriculate: having auricles (in botany, small lobes situated at the 

base of a leaf, petiole, ligule, etc.). 

Bases: all the exchangeable positive ions in the soil which raise 

the pH*. 

Basionym: original name under which a species was first 

described. 

Basiphile: inhabiting an environment (soil, water) rich in bases*. 

Beak: in botany, a stiff pointed projection on the end of a dry fruit. 

Benthic: inhabiting the bed of a wetland or an ocean. 

Bioclimate: climatic conditions which characterise the various 

types of continental environment, either at the scale of natural 

regions of limited area or at the scale of stations occupying small 

areas. 

Biomass: total mass of living material, animal and vegetable, in 

a defined biotope at a given time. 

Bipinnatifid: used of a pinnatifid* leaf whose divisions are themselves 

pinnatifid. 

Bract: more or less modified leaf situated in the axil of a flower 

or an inflorescence. 

Bracteole: small bract situated in the axil of the ray of a 

branched inflorescence or in the axil of each of its constituent 

flowers. 

Brood pouch: see oviger 

Bryophytes: group comprising both mosses and liverworts. 

Capsule: in mosses and liverworts, designates a hollow dry-walled 

organ of the sporophyte, in which the spores develop. Among 

higher plants, designates a dry dehiscent fruit (i.e. opening spontaneously 

when ripe) containing more than one seed. 

Caudal crest: membrane on both sides of the tail in tadpoles (in 

the Anura) and larvae (in Urodela) used for propulsion. In some 

species, this crest extends up the back to the back of the head 

(Tree Frog tadpoles for example). 

118 

Cercopods: (= cerci) in arthropods and crustaceans, appendages 

located on the last abdominal segment (usually two in number, 

one on each side). 

Chamaeophyte: (= chameophyte) low woody plant whose buds 

are at least 25 cm above the soil (see Figure) 

Charophytes: specialised algal group consisting of one family, 

the Characeae, characterised by the whorled structure of the 

thallus and by the highly complex structure of the reproductive 

organs (antheridia and oogonia). 

Chitinous: formed of chitin, a molecule involved in the structure 

of the arthropod cuticle. 

Climax: used of a plant community which has developed to a 

sustainable state of equilibrium with the climatic and edaphic 

conditions of the environment, in the absence of human intervention. 

Clonal: used of an organism (animal or plant) derived from a single 

cell or a single individual, by asexual reproduction (known as 

vegetative propagation* in plants) 

Cohort: set of individuals which have experienced the same 

event at the same time (individuals born at the same time or 

breeding at the same time in pools, for example). 

Connectivity: facilitation of the movement of individuals of a 

species between local sub-populations to form a single functional 

demographic unit. 

Cordate: describes a heart-shaped flat organ. 

Coronule: short apical cells which develop a crown shape at the 

tip of the oogonium* in the Characeae. 

Cupular pool: small temporary pool located in bowl created in 

rock by natural erosion (see volume 1, chapter 2a). 

Cyme: branched inflorescence in which each flower is located at 

the end of a branch. 

Daya: a temporary pool in Morocco. 

Dehesa: Spanish name used to describe a landscape specific to 

certain regions in the centre and south of Spain, consisting of 

pasture or cereal cultivation scattered with evergreen oaks or 

Cork Oaks, which are usually cut. 

Detrivore (or Détritivorous): feeding on dead organic material. 

Diapause: period during which metabolic activity and the development 

of an insect is suspended at a particular stage (egg, larva, 

nymph or adult), as a result of the action of internal or external 

factors. 

Diploid: describes an organism whose cell nuclei have a double 

set of chromosomes (2n). 

Echinulate: covered in small spines. 

Ecophase: during its life cycle, a species passes through different 

stages (egg, larva, juvenile, etc.). An ecophase corresponds to 

one of these stages having a different ecology from the other 

parts of the cycle. 

Edaphic: refers to ecological factors linked with the soil as well 

as to their relationships with plants. 

Elater: elongated cell attached to a spore, and capable of flexible 

movements. 

Endemic: used of a species exclusively confined to a given biogeographical 

area, often of limited extent. 

Espace naturel sensible (Sensitive Natural Areas): areas bought 

by each département in France since 1985 “to preserve the quality 

of sites, landscapes and natural habitats” in their territories, 

using a tax levied on building. 

Eutrophic: used of water rich in nutrients (nitrates, phosphates) 

and with little oxygen at depth. Opposite of “oligotrophic*”. 

Eutrophication: process of enrichment of a water body with 

mineral nutrients (phosphates, nitrates) often accompanied by a

proliferation of algae and/or aquatic higher plants, leading to 

oxygen depletion of the deeper water as well as a build-up of 

organic matter. 

Fistulous: hollow and cylinder-shaped. 

Gametangium: organ of the gametophyte* in which the 

gametes are formed. 

Gametophyte: in the life cycle of algae, mosses, liverworts and 

ferns, designates the haploid* phase of the organism, which produces 

gametes; in higher plants refers to the haploid organ 

which produces the gametes. 

Genetic bottleneck: sudden decrease in the size of a population 

associated with a decrease in the total genetic variability. 

Genetic bottleneck: sudden decrease in the size of a population 

associated with a decrease in the total genetic variability. 

Genetic drift: within small populations, chance fluctuations in 

gene frequency with each generation, enabling rare genes (which 

would be eliminated by selective pressure within large populations) 

to be continuously expressed. 

Geophyte: plant species which withstand the unfavourable season 

thanks to the presence of bulbs, rhizomes or any other type 

of underground reserve organ (see Figure). 

Gravel bed/Gravels: loose sedimentary formation consisting of 

rock fragments. 

Gyrogonites: calcified female fructifications produced by charophytes, 

corresponding to fossil or living forms after dispersal. 

They are invariably made up of five cells in the form of lefthanded 

spirals, joined at the tips. 

Halotolerant: tolerant of a certain amount of salt in the environment 

(water, soil, etc.). 

Haploid: organism or organ whose cell nuclei each contain a 

set of n chromosomes, comprising a single copy of each chromosome. 

Heliophilous: used of a plant which grows in conditions of strong 

sunlight. 

Helophyte: marsh plant whose budding parts, which enable it to 

survive during the bad season, are laid down in the sediment, 

while in the good season they develop an aerial structure which 

extends above the water surface (Reed, for example). 

Hemicryptophyte: perennial herbaceous plant whose budding 

parts, which enable it to survive during the bad season, remain 

on the surface of the soil, at the very base of the stems (or of the 

tuft for caespitose Graminae) (see Figure). 

Heterosporous: producing spores of two types (in algae, mosses, 

ferns and liverworts): microspores and macrospores. 

Hexamerous (6-merous): describes a plant organ the number of 

whose constituent equivalent parts (petals, sepals, stamens, etc.) 

is six or a multiple of six. 

Hexaploid: said of an organism whose cell nuclei contain three 

pairs of each homologous chromosome. 

Hybridogenic: taxon derived from a cross between two or several 

different taxa usually different species or sub-species. 

Hydroperiod: period during the year when the pool contains 

water. 

Hydrophyte: plant that lives in an aquatic environment (Water 

Milfoils, Water Lilies). 

Hydrophytic: refers to a hydrophyte* plant. 

Hygrophilous: organisms dependent on biotopes characterised 

by high soil water content. 

Karstic: relating to karst (the whole of the surface and subterranean 

structure of a limestone massif, which results from the 

dissolution of rock and is characterised by a wide-ranging system 

of underground water flow). 

Glossary 

Laune (or lône): a local term meaning a former branch of a 

watercourse which can reconnect with the present watercourse 

when water levels are medium or high. 

Lavogne: local term meaning depressions dug out for watering 

livestock, in the Causses of the Cevennes and the Uzégeois and 

Montpellierais garrigues (France). 

Life form: five life forms (see Figure) were defined by Raunkiaer 

in his ecological classification of plants based on their strategy 

for surviving through the unfavourable season. Each life form is 

defined by the position of the buds in the architecture of the 

plant, as well as by their degree of protection during this season. 

Longevitous: living for a long time 

Macrophytes: generic term including all plants that are visible 

to the naked eye. 

Matorral: word of Spanish origin, in areas with a Mediterranean 

climate usually applied to a bushy vegetation community, adapted 

to drought and often dominated by evergreen species, often with 

small leaves. 

Meso-eutrophic: see Mesotrophic*. 

Mesotrophic: intermediate between eutrophic* and oligotrophic*. 

Metanauplius: larval stage following the nauplius* stage in 

crustaceans, characterised by the beginnings of body segmentation. 

Several moults, over a few weeks or a few months, are then 

necessary to attain the adult stage. 

Metapopulation: set of populations that are interconnected via 

migration events (gene flow) and are subject to extinction and 

recolonisation. This concept may be extended to include any set 

of populations developing in a more or less independent way 

which are, however, interconnected through rare instances of 

migration. 

Metathorax: posterior part of the thorax in insects, to which are 

attached the last pair of legs (and sometimes the second pair of 

wings). 

Microphagous: feeding on small prey. 

Mucronate: terminating abruptly in a sharp point. 

Natura 2000: ongoing European policy, with the aim (through 

the implementation of the Habitats Directive 119 and of the 

Council Directive 79/409/EEC of 2 April 1979 on the conservation 

of wild birds) of reconciling human activities and natural habitats 

within a network of sites having great ecological interest, 

some designated for their diversity of birds (Special Protection 

Areas) others for the rarity of their fauna, their flora and their 

natural habitats (Special Conservation Areas). 

Nauplius: first larval stage of crustaceans, followed by the 

metanauplius* stage. 

Oligotrophic: describes nutrient-poor water (low in nitrate, 

phosphate, sulphate): its opposite is “eutrophic”. 

Oogonium: reproductive cell of lower plants (algae, mushrooms) 

which produces the female gametes. 

Ornithochory: mechanism of seed* or spore dispersal by birds. It 

may be external (carried on the body) and/or internal (absorption 

and passage through the digestive tract). 

Pappus: plume of hairs crowning an achene. 

Parotoid glands: glands located at the back of the head among 

some species of amphibians (true toads, salamanders), producing 

a poisonous secretion as a deterrent against predators. 

Perennial: in botany, describes a plant species which lives for 

more than one year (as opposed to annual). 

Phenology: description of the various phases in the life cycle of 

a species. 

Phytocenosis: collection of plant species having a homogenous 

appearance and colonising the same habitat (syn.: plant community). 

119


Phytoplanktonophagous: feeding on phytoplankton. 

Phytosociology: see volume 1, chapter 2, box n° 4. 

Pinnatilobed: used of a divided leaf when the divisions cut into 

the lamina for less than half its width. 

Pinnatipartite: used of a pinnate leaf when the divisions cut into 

the lamina for more than half its width. 

Pinnatisect: used of a pinnate leaf when the divisions cut completely 

into the lamina as far as the midrib. 

Pleurocarp: describes a bryophyte* whose female gametangia* 

(and hence sporophytes) appear in a lateral position and not at 

the ends of the gametophyte* stems. 

Polje: karstic* plain or subsidence valley, resulting from the coalescence 

of several karstic erosion systems (“ouvalas”) and covered 

in decalcified clays. A polje is subject to more or less prolonged 

seasonal flooding which sometimes occurs suddenly, depending 

on the underground karstic network. 

Polyploid: condition of an organism whose cells contain more 

than two sets of homologous chromosomes, i.e. more than 2n 

chromosomes. 

Pozzines: high altitude swards dominated by Carex, growing on 

a peaty substrate and mostly dotted with pools (Corsica, Pyrénées, 

Sierra Nevada, etc.). 

Pre-imaginal stage: final larval stage before the adult stage, in 

insects. 

Figure. The main biological types, according to Raunkiaer** 

120 

Pistacia 

lentiscus 

Artemisia 

molinieri 

Eryngium 

pusillum 

Phanerophytes Chamaeophytes Hemicryptophyte Cryptophytes 

Geophytes with bulb 

Prolarva: temporary larval stage (of very short duration) immediately 

after hatching. 

Propagule: any part of an organism, produced by asexual or sexual 

reproduction, capable of producing a new individual. 

Prothorax: in Odonata, the anterior part of the thorax, very 

short, bearing the head and the first pair of legs. 

Pudding stone: consolidated sedimentary rock of the conglomerates 

group, of continental or marine origin, composed of pebbles 

bound together with a limestone or sandstone cement. 

Recruitment: the adding of new individuals to a population. Recruitment 

takes place through reproduction, immigration and restocking. 

Rhizoid: hair-like rooting structure, particularly among mosses. 

Scabrid: rough (covered in protrusions). 

Scape: leafless peduncle of a flower (or inflorescence). 

Sclerified: impregnated with lignin (component of the wall of 

certain cells, in particular in bushes, trees and shrubs, making it 

rigid and impermeable). 

Seed stock (of the soil): all the viable seeds and spores in the soil. 

Seedbank: all the viable seeds in the soil. 

Sessile: Botanical: an organ (leaf, flower) having no petiole or 

peduncle. Zoological: microorganism attached to a support 

(stem, rock, etc.). 

Sex-ratio: ratio of the numbers of male and female individuals 

in a defined population. 

Ophioglossum 

lusitanicum 

Isoetes 

histrix 

In perennial plants, position of the buds 

surviving the bad season 

Cryptophytes 

Géophytes 

with rhizome 

Ranunculus 

revelieri 

Therophytes 

(surviving by seeds) 

Annual plants Perenial 

plants 

**: Raunkiaer, C, 1934. The life form of plants. Oxford, Clarendon Press. 632 pages 

seeds 

N. Beck

Sheath: a leaf base expanded to form a more or less cylindrical 

envelope, split longitudinally or not and clasping the stem. 

Sorus: group of sporangia*, arranged in a distinctive pattern on 

the lower surface of the fronds of some ferns. 

Spiracle: in tadpoles, opening which enables water to be ejected 

during respiration. In tadpoles of Painted Frogs, Midwife Toads 

and Yellow-bellied Toads, it is located in a median position on 

the underbelly. In other European amphibians it is situated on the 

sides of the abdomen. 

Sporocarp: in some ferns (Marsilea, Pilularia, etc.), a particular 

type of sorus*, closed and with tough walls, enclosing the spores. 

Sporophyll: in ferns, specialised leaf or frond, bearing one or 

more sporangia. 

Station (of plants): homogenous area of land where a plant 

species (or a particular plant community) is found. 

Stylopodium: small fleshy nectariferous disc which surmounts 

the fruit in Apiaceae (= Umbellifers), bearing two separate styles. 

Sympodic: describes the parts of a plant resulting from sympodial 

growth, i.e. lengthening of the stem in a succession of segments 

produced by lateral budding of the stems which have 

flowered in the previous year. 

Terra Rossa: type of soil found in Mediterranean regions, derived 

from the breakdown of limestones and characterised by an accumulation 

of ferric oxides. 

Glossary 

Therophyte: synonym for an annual plant, a herbaceous plant 

with a very short reproductive cycle, lasting a few months or in 

certain cases a few weeks, which survives the bad season in the 

form of seeds. (see Figure). 

Therophytic: describes a therophyte*. 

Trophic: everything relating to nutrition among plants and animals. 

Vegetative reproduction: mode of reproduction of a plant 

species using vegetative organs (stolons, rhizomes, tubers, etc.). 

Vegetative stage: part of the life cycle of plants relating to germination, 

growth and propagation (vegetative reproduction), 

excluding flowering and fruiting. 

Verticil (whorl): all the homologous organs arising from a stem 

at the same point and radiating out from the stem. 

Vicariant: used of animal or plant species that are taxonomically 

closely related and which inhabit environments with similar ecological 

characteristics in different geographical regions. 

ZNIEFF: (Zone naturelle d’intérêt écologique floristique et faunistique) 

one of a list (begun in 1982 by the MEDD) of French 

natural areas, both terrestrial and marine, whose interest lies 

either in the stability and the richness of the ecosystem or in the 

presence of rare and threatened animals and plants. 

121

Macrocrustacean inventory form 

Please send to: Danielle DEFAYE, Muséum National d’Histoire Naturel, Laboratoire de Zoologie - Arthropodes (Crustacés), 61, rue Buffon, F-75005 Paris 

1. Essential data 

A) Source 

Observer or code: 

Surname, first name: 

Adress: 

Name of form compiler if different from observer: 

B) Taxon 

Number of reference 

list: 

C) Place 

122 

or 

Coordinates 

– in grades (Paris meridian) 

Longitude: � E, �W 

Latitude: 

or coordinates 

– in degrees (Greenwich meridian) 

Longitude: � E, �W 

° ‘ ‘’ 

Latitude 

° ‘ ‘’ 

2. Additional data 

Type of contact: 

� seen 

� photo 

� captured/released 

� found dead 

� preserved 

� exuviae 

� .............................. 

or or 

Biblio. ref. or code: 

Author(s): 

Date: 

Title: 

Periodical: 

Volume, number, page: 

Genus Species Subspecies or variety 

Commune + locality: 

INSEE code: 

IGN map number: 

Coll. ref. or code: 

Where deposited: 

Inventory number: 

Name of form compiler if different from observer: 

Taxon 

or and and 

D) Date 

Abundance and stage: 

1 2-10 +10 

� � � male 

� � � female 

� � � oviger 

� � � sexe = ? 

� � � juvenile 

� � � larvae 

� � � cyst 

Altitude: 

At: 

hr 1 hr 2 day month year 

3. Additional information (in note form to aid validation) 

Depth of biotope: 

Duration of flooding: 

Colour on the living creature: 

Systematic notes, associated fauna, biometry, etc.: 

Biotopes: 

� clay 

� sand 

� rock surface 

� clear water 

� turbid water 

� grassy ground 

� bare soil 

� algae 

� vegetation beds 

� .............................. 

metres 

� salt marsh 

� salt water 

� fresh water 

� Isoetes 

� Ranunculus 

� .................... 

determined by: 

verified by: 

Name of pool or water body: 

Method of collection: 

� by hand 

� shrimp net 

� drag net 

� plankton net 

� bird stomach 

� amphibian stomach 

� fish stomach 

� .............................. 

v.c. 

Use reverse side of the form if necessary

Bibliography 

1. Aboucaya, A., H. Michaud & M. Delorme, 2002. Etude de la flore et de 

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