S T R E L I T Z I A 35

Transcription

1

2

3 S T R E L I T Z I A 35 Systematics and biology of Lapeirousia, Codonorhiza, Psilosiphon and Schizorhiza in southern Africa Peter Goldblatt and John C. Manning Pretoria 2015

4 S T R E L I T Z I A This series has replaced Memoirs of the Botanical Survey of South Africa and Annals of the Kirstenbosch Botanic Gardens, which the South African National Biodiversity Institute (SANBI) inherited from its predecessor organisations. The plant genus Strelitzia occurs naturally in the eastern parts of southern Africa. It comprises three arborescent species, known as wild bananas, and two acaulescent species, known as crane flowers or bird-of-paradise flowers. The logo of SANBI is partly based on the striking inflorescence of Strelitzia reginae, a native of the Eastern Cape and KwaZulu-Natal that has become a garden favourite worldwide. It symbolises the commitment of SANBI to champion the exploration, conservation, sustainable use, appreciation and enjoyment of South Africa s exceptionally rich biodiversity for all people. Peter Goldblatt B.A. Krukoff Curator of African Botany, Missouri Botanical Garden, St. Louis; Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa John C. Manning Compton Herbarium, South African National Biodiversity Institute, Cape Town; Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa EDITOR: PROOFREADER: COVER DESIGN & LAYOUT: COVER PHOTOGRAPHS: Yolande Steenkamp Alicia Grobler Elizma Fouché John C. Manning Citing this publication GOLDBLATT, P. & MANNING, J Systematics and biology of Lapeirousia, Codonorhiza, Psilosiphon and Schizorhiza in southern Africa. Strelitzia 35. South African National Biodiversity Institute, Pretoria. ISBN: Obtainable from: SANBI Bookshop, Private Bag X101, Pretoria, 0001 South Africa Tel.: bookshop@sanbi.org.za Website: Printed by: Seriti Printing, Tel.: , Website: Address: Unit 6, 49 Eland Street, Koedoespoort, Pretoria, 0001 South Africa Copyright 2015 by South African National Biodiversity Institute (SANBI) All rights reserved. No part of this book may be reproduced in any form without written permission of the copyright owners. The views and opinions expressed do not necessarily reflect those of SANBI. The author and publisher have made their best efforts to prepare this book, and make no representation or warranties of any kind with regard to the completeness or accuracy of the contents herein. All images in this book have been reproduced with the knowledge and prior consent of the artists concerned and no responsibility is accepted by the publisher or printer for any infringement of copyright or otherwise arising from the contents of this publication. Every effort has been made to ensure that the credits accurately comply with the information supplied by the author.

5 S T R E L I T Z I A 35 (2015) iii Contents Abstract Acknowledgements... Introduction... 1 Taxonomic history Morphology Chromosome cytology Floral biology Geography Ethnobotany Phylogeny and subgeneric classification Systematics Key to genera of tribe Watsonieae Systematics of Lapeirousia Key to sections Key to species A. Sect. Chasmatocallis (Spp. 1 8) B. Sect. Sophronia (Spp. 9 13) C. Sect. Lapeirousia (Spp ) Systematics of Codonorhiza Key to species Spp Systematics of Schizorhiza Systematics of Psilosiphon Key to species in southern Africa Spp New combinations for Psilosiphon species not represented in southern Africa References Index iv v

6 iv S T R E L I T Z I A 35 (2015) Abstract Plastid and nuclear DNA sequence analyses show that the sub-saharan African genus Lapeirousia Pourr. is paraphyletic as presently circumscribed. Species of Lapeirousia are retrieved as a clade in which are nested the tropical African genera Cyanixia Goldblatt & J.C.Manning and Savannosiphon Goldblatt & Marais as sister to Lapeirousia sect. Paniculatae Goldblatt plus the taxonomically isolated L. neglecta Goldblatt. To preserve taxonomic monophyly we dismember Lapeirousia, recognising the new genera Codonorhiza Goldblatt & J.C.Manning for L. sect. Fastigiatae Goldblatt (7 spp.) and Schizorhiza Goldblatt & J.C.Manning (1 sp.) for L. neglecta, both from the Cape Floristic Region, as well as Psilosiphon Welw. ex Goldblatt & J.C.Manning (15 spp.) for L. sect. Paniculatae, all from tropical and eastern southern Africa. This leaves Lapeirousia (27 species) centred in western southern Africa, but also represented in south tropical Africa. All four genera have derived corms with a flat base, each with distinctive and different corm tunics and sometimes corm shapes that are correlated with fundamental differences in leaf morphology and anatomy, floral bracts, and seed shape and surface sculpturing. Codonorhiza and Schizorhiza also have apomorphic, 1-banded pollen apertures. We revise the infrageneric taxonomy of Lapeirousia in light of the molecular phylogeny, recognising sect. Chasmatocallis (R.C.Foster) Goldblatt & J.C.Manning, sect. Lapeirousia and sect. Sophronia (Licht. ex Roem. & Schult.) Goldblatt & J.C.Manning. We provide dichotomous keys to the four genera in southern Africa and full revisions of all four genera. We recognise the following six new species: C. elandsmontana Goldblatt & J.C.Manning and C. pillansii Goldblatt & J.C.Manning from Western Cape, South Africa; L. purpurea Goldblatt & J.C.Manning from Western Cape; L. kalahariensis Goldblatt & J.C.Manning from southern Namibia and Northern Cape; L. kamiesmontana Goldblatt & J.C.Manning, a local endemic of the Kamiesberg of Northern Cape; and P. erongoensis Goldblatt & J.C.Manning from central Namibia. We also raise L. dolomitica subsp. lewisiana (B.Nord.) Goldblatt and L. littoralis subsp. caudata (Schinz) Goldblatt to species rank as L. lewisiana B.Nord. and L. caudata Schinz respectively, and recognise L. angustifolia Schltr., currently included in L. pyramidalis (Lam.) Goldblatt. We describe two new subspecies in the South African L. fabricii (D.Delaroche) Ker Gawl.; subsp. compressa Goldblatt & J.C.Manning and subsp. purpurascens Goldblatt & J.C.Manning; the new subsp. foliosa Goldblatt & J.C.Manning of L. plicata (Jacq.) Diels; and the new P. sandersonii subsp. limpopoensis Goldblatt & J.C.Manning.

7 S T R E L I T Z I A 35 (2015) v ACKNOWLEDGEMENTS We are grateful to Elizabeth Parker and Lendon Porter for assistance and companionship in the field and to Ingrid Nänni, SANBI, who helped considerably with travel arrangements and with field work. We thank Clare Archer, SANBI, Pretoria, for providing answers to our several questions about holdings at the National Herbarium in Pretoria; Mary Stiffler, Missouri Botanical Garden, who cheerfully helped solve bibliographic questions and provided copies of numerous articles not readily available to us; and Roy Gereau, Missouri Botanical Garden, for advice on nomenclatural questions. Michelle Smith prepared the distribution maps. This study was supported in part by grant from the US National Geographic Society. Collecting permits were provided by the conservation authorities of Northern Cape and Western Cape.

8

9 S T R E L I T Z I A 35 (2015) 1 INTRODUCTION Phylogenetic analysis of plastid and nuclear DNA sequences (Forest et al. 2014) shows that the sub-saharan African genus Lapeirousia Pourr. (± 45 species) is paraphyletic as presently circumscribed (e.g. Goldblatt 1990a; Goldblatt & Manning 2008). Three tropical African genera of tribe Watsonieae Klatt (subfamily Crocoideae Burnett) (Goldblatt et al. 2006), Cyanixia Goldblatt & J.C.Manning (1 sp.), Savannosiphon Goldblatt & Marais (1 sp.) (and by inference Zygotritonia Mildbr., 6 spp.) nest within the genus, where they are retrieved as sister to the largely tropical and eastern southern African L. sect. Paniculatae Goldblatt, excluding L. neglecta Goldblatt. This clade is in turn sister to the largely southern but also south tropical African subg. Lapeirousia, with this combined clade sister to L. neglecta and then with L. sect. Fastigiatae Goldblatt from the Cape Floristic Region (Figure 1). The characters used to unite Lapeirousia in the past were a flat-based, ± bell-shaped corm (unique in Watsonieae but also present in several other genera of Crocoideae), and often a strongly asymmetric karyotype (Goldblatt & Takei 1993). Most species also have deeply divided style branches, a character ancestral in Watsonieae but lacking in Cyanixia, Zygotritonia and the Cape endemic Pillansia L.Bolus as well as in some species of Lapeirousia sect. Paniculatae. Options for circumscribing genera in Watsonieae as monophyletic based on the molecular phylogeny are either to unite Cyanixia, Savannosiphon and Zygotritonia in an expanded and, in practice, morphologically indefinable Lapeirousia, or to treat L. sect. Paniculatae, L. neglecta and sect. Fastigiatae as separate genera. We select the latter option, transferring the tropical and eastern southern African species of sect. Paniculatae to the new genus Psilosiphon (naked or smooth tube) and those of sect. Fastigiatae to a second new genus Codonorhiza (bell-shaped root). We treat the phylogenetically isolated and morphologically intermediate L. neglecta as the new genus Schizorhiza. In this revised generic taxonomy all genera in the Lapeirousia clade are evidently monophyletic and readily distinguished by macro- and micro-morphological features, including leaf anatomy, pollen and seed morphology, and chromosome cytology. The inference to be drawn from the phylogenetic analysis is that flat-based, ± bell-shaped corms are ancestral to the greater Lapeirousia clade and were either lost by reversal in Cyanixia, Savannosiphon and Zygotritonia or were acquired independently in Psilosiphon and Codonorhiza plus Lapeirousia and Schizorhiza. Woody corm tunics of brown colour characterise Lapeirousia, and tunics of blackish, compressed fibres are unique to Codonorhiza (sect. Fastigiatae). Psilosiphon has pale or dark brown, fibrous tunics, usually becoming loosely fibrous with age, and Schizorhiza has pale, corky tunics and ± ellipsoid rather than bell-shaped corms with a flat base; the diameter of the base smaller than the middle of the corm. These differences make the latter hypothesis seem reasonable. Associated with the corm differences are smooth, round seeds in Psilosiphon, Schizorhiza and most species of Lapeirousia (in which L. caudata Schinz, L. littoralis and all members of sect. Sophronia have a sculptured seed surface). In contrast, Codonorhiza has unusually small, ellipsoid seeds with reticulate sculpturing and a prominent funicular collar. Differences in the floral bracts among the four genera correlate closely with the corm tunic, seed and leaf features and are consistent with separate generic status for the three clades. The molecular phylogeny renders the available taxonomic accounts of the Lapeirousia complex out of date and in need of substantial revision. This account presents a full revision of Codonorhiza, Lapeirousia, Schizorhiza and the

10 2 S T R E L I T Z I A 35 (2015) FIGURE 1. Phylogenetic relationships in Lapeirousia and allied genera based on ten plastid regions (matk, rpl16, trnqrps16, trnl, trnl-trnf, ndhj, ycf5, ndhf-rpl32, rpl32-trnl, ACCD-psa1) and one nuclear marker (RPB2) and obtained using the Bayesian inference implemented in MrBayes. Posterior probabilities are indicated on nodes (modified from Forest et al. 2014).

11 S T R E L I T Z I A 35 (2015) 3 southern African species of Psilosiphon, thus bringing together the results of a series of papers in systematics, chromosome cytology, floral biology and phylogeny and speciation published after a revision of Lapeirousia in the southern African winter-rainfall zone (Goldblatt 1972) and later for tropical Africa (Goldblatt 1990a). In this treatment, we recognise seven species in Codonorhiza, 27 species in three sections in Lapeirousia, 15 species in Psilosiphon and one in Schizorhiza. Four species of Psilosiphon occur exclusively in tropical Africa and we do not deal with them in detail here, providing only the new combinations in Psilosiphon. Six new species are included in the account, C. elandsmontana, C. pillansii, L. kalahariensis, L. kamiesmontana, L. purpurea and P. erongoensis, as well as new subspecies in L. fabricii and L. plicata. In addition, we recognise L. lewisiana B.Nord., which Goldblatt (1972) treated as subsp. lewisiana of L. dolomitica; L. caudata, currently treated as a subspecies of L. littoralis; and L. angustifolia Schltr., included by Goldblatt (1972) in L. pyramidalis.

12 4 S T R E L I T Z I A 35 (2015) TAXONOMIC HISTORY The genus Lapeirousia was described in 1788 by French naturalist, Pierre André Pourret for the single species L. compressa, which he believed to have been collected in Isle de France, now Mauritius. Pourret was unaware that several species corresponding to his genus had already been described but assigned to Gladiolus L. (species with zygomorphic flowers) or to Ixia L. and even Galaxia Thunb. (both genera for species with radially symmetric flowers). John Ker Gawler (1804) adopted Lapeirousia in his then novel treatment of southern African Iridaceae in which he described the new genera Anomatheca Ker Gawl., Babiana Ker Gawl., Geissorhiza Ker Gawl., Hesperantha Ker Gawl. and Tritonia Ker Gawl., while also recognising Gladiolus and Ixia. Ker Gawler s Lapeirousia included just seven species, three of them now referred to Codonorhiza. Shortly thereafter K.P. Sprengel (1817) erected the genus Ovieda to replace Lapeirousia without providing any reason for his action and the name was adopted for a short time despite the fact that it was not only a later synonym, thus superfluous and illegitimate, but also a later homonym for Ovieda L. (Verbenaceae). In the same year Roemer & Schultes (1817) described Sophronia for the acaulescent species, S. caespitosa, which had already been illustrated by Jacquin (1795) under the name Galaxia plicata (now L. plicata (Jacq.) Diels). In the Dictionaire des Sciences Naturelles, Poiret (1826) used the name Peyrousia for species of Lapeirousia, a name perhaps intended as a grammatical correction rather than a new genus since the circumscription of Peyrousia corresponded exactly to that of Lapeirousia. Then, for no evident reason, A. Dietrich (1833) transferred all the known species of Lapeirousia to a new genus Meristostigma A.Dietr., now regarded as an illegitimate, superfluous name. L. falcata (L.f.) Ker Gawl., L. fastigiata (Lam.) Ker Gawl. and L. corymbosa (L.) Ker Gawl., all described before 1800 in Gladiolus or Ixia, were included without question in Lapeirousia, then distinguished by the combination of bell-shaped corms and divided style branches. These three species form the core of the new genus Codonorhiza. The first species of Psilosiphon was discovered in Ethiopia by Henry Salt on his expedition to Abyssinia (Salt 1814). His collections were referred by Robert Brown in the report of the expedition to Geissorhiza as G. abyssinica, without description. Over 25 years later in the 1840s, the French explorers Quartin-Dillon and Petit again collected P. abyssinica; their collections being described as Montbretia abyssinica by A. Richard (1850). The species was later transferred to Lapeirousia by J.G. Baker in Quartin-Dillon and Petit also collected P. schimperi (Aschers. & Klatt) Goldblatt & J.C.Manning, as did the Austrian, Georg W. Schimper, who spent the years in Ethiopia. The species was first referred to Tritonia (Klatt 1866b), then to Acidanthera Hochst. because the white flower and elongate perianth tube corresponded to the circumscription of that genus. Only in 1934 was T. schimperi Aschers. & Klatt transferred to Lapeirousia. Then in the 1860s another Austrian, Friedrich Welwitsch, living in Angola, collected different populations of what we now regard as the same species, to which he gave the manuscript names P. cyanescens and P. fragrans. Welwitsch also discovered L. odoratissima Welw. ex Baker, naming his collections Psilosiphon odoratissima. J.G. Baker (1878) referred all Welwitsch s provisional Psilosiphon species to Lapeirousia. At the time Psilosiphon remained without formal description. Species of both Codonorhiza and Psilosiphon were included in Lapeirousia together with Ker Gawler s new genus Anomatheca (now included in Freesia Klatt) by Baker in his accounts of the Iridaceae for Flora capensis (Baker 1896) and Flora of tropical Africa (Baker 1898), evident-

13 S T R E L I T Z I A 35 (2015) 5 ly on the basis of their long-tubed flowers with divided style branches. Foster (1939) described the genus Chasmatocallis for the florally distinctive L. divaricata Baker, an action that was largely ignored by the botanical community. Baker s circumscription of Lapeirousia was revised by Goldblatt (1972), who recognised Anomatheca as a separate genus more closely allied to Freesia Klatt (1866a) than Lapeirousia. Goldblatt s (1972, 1990a) accounts of Lapeirousia for the southern African winter rainfall region and for tropical and summer rainfall southern Africa collectively admitted 34 species to the genus. Additional species were added in the years that followed (Goldblatt & Manning 1992, 1994) with the result that in 2010 Lapeirousia included over 40 species. A subgeneric classification based on corm and leaf morphology, proposed by Goldblatt & Manning (1990), recognised two subgenera, subg. Lapeirousia and subg. Paniculatae [as Paniculata], the latter with sect. Paniculatae (now Psilosiphon) and sect. Fastigiatae [as Fastigiata] (now Codonorhiza). Apomorphic features for subg. Lapeirousia are the prominently ridged leaves without a central vein, large outer floral bracts, leathery or succulent in texture, an inner bract much smaller than the outer, and particularly hard, woody corm tunics. For subg. Paniculatae the derived features are primarily the hard but ± fibrous corm tunics, and an inflorescence that is (usually) a false panicle of few-flowered spikes. The floral bracts are ± equal in length in both sections of subg. Paniculatae but in sect. Fastigiatae the bracts are leathery, green and obtuse to ± truncate vs. of softer texture, acute and becoming dry apically in sect. Paniculatae. Although first collected in 1943 by R.H. Compton, Schizorhiza neglecta was only described in 1972 as subsp. alta Goldblatt of Lapeirousia corymbosa and later treated as the separate species L. neglecta (Goldblatt & Manning 1992) and referred to sect. Paniculatae. Molecular phylogenetic studies in the Iridaceae (Goldblatt et al. 2006; Goldblatt & Manning 2008) showed that Lapeirousia in the broad sense falls in a clade that retrieves Watsonia Mill., Micranthus (Pers.) Eckl., Pillansia L.Bolus and Thereianthus G.J.Lewis as sister to Lapeirousia plus Savannosiphon and Cyanixia. The tropical African Zygotritonia was inferred to belong here as well, but was not included in these studies. This assemblage of genera currently constitutes the tribe Watsonieae of subfam. Crocoideae.

14 6 S T R E L I T Z I A 35 (2015) MORPHOLOGY Corm: although the corms of all three genera are bell-shaped to broadly obconic (triangular in outline), the corm tunics differ markedly among them (Figure 2). Tunics of Codonorhiza are dark grey and composed of concentric layers of compacted fibres with a broad basal lip. Those of Lapeirousia are dark brown to reddish brown and woody with a crenate to finely toothed or spiny basal margin. In Psilosiphon the tunics are usually light brown and of a hard, fibrous to leathery texture and lack marginal decoration (Table 1). Somewhat unusual for the entire Lapeirousia clade, Schizorhiza produces multiple cormlets at the subterranean and lower aerial nodes of the stem. The genus is also unusual in that several new corms replace the parent at the end of the growing season instead of the usual single new main corm. Stem: usually aerial and branched, stems are rounded to somewhat compressed in cross section. Branching is pronounced in Codonorhiza, with the branches arranged in corymbose fashion forming a compound, flat-topped inflorescence. Stems of most species of Psilosiphon have a similar branching pattern, especially prominently branched in P. coeruleus, P. erythranthus and P. sandersonii, whereas branching in P. gracilis and P. schimperi is limited, resulting in a rather different appearance. Aerial stems of Lapeirousia are more strongly flattened and branches bear multiple flowers in spikes. In species of L. sect. Sophronia the stem is subterranean and strongly contracted so that plants form low tufts at ground level with flowers crowded together. Leaves: leaf anatomy, especially the leaf margins, is typically consistent in genera of Crocoideae (Rudall 1995; Goldblatt et al. 2006) and is, as far as known, consistent in all genera of Watsonieae. Leaf margins in the Lapeirousia group of genera are of two types: simple and without an associated marginal vein or subepidermal strand of sclerenchyma; or with a welldeveloped marginal vein present with ± prominent sclerenchyma cap (Figure 3). Codonorhiza (C. corymbosa examined: Rudall 1995) and Schizorhiza have simple margins of the first type, lacking marginal veins or a sclerenchyma strand. In contrast, Lapeirousia (L. anceps, L. divaricata and L. odoratissima examined) and Psilosiphon (P. avasmontanus, P. bainesii, P. coeruleus, P. erythranthus examined) (Table 1) have margins A B C D E FIGURE 2. Corm morphology in Lapeirousia and allied genera. A, Lapeirousia plicata; B, L. dolomitica; C, Psilosiphon bainesii; D, Psilosiphon coeruleus; E, Codonorhiza micrantha. Artist: John Manning.

15 S T R E L I T Z I A 35 (2015) 7 TABLE 1. Comparison of taxonomically significant vegetative characters in the Lapeirousia clade Taxon Corm shape Corm tunics Codonorhiza bell, base flat compacted fibres, greyblack Lapeirousia Psilosiphon Schizorhiza bell to obconic, base flat ± bell, base flat ± ellipsoid with flat base woody, brown leathery or compacted fibres, brown cork-like, light brown Cyanixia globose fibrous, redbrown Savannosiphon globose fibrous, redbrown Zygotritonia globose fibrous, redbrown Foliage leaves 1 or 2, lower largest few, lower largest 2 several in basal fan few in basal fan 2 or 3 in basal fan several in basal fan 1 or 2, lower largest Leaf marginal vein absent present present absent present present Vascular bundles opposite; sheath complete alternate; sheath incomplete ± opposite; sheath incomplete alternate; sheath complete alternate; sheath incomplete opposite; sheath complete Leaf surface plane corrugately ribbed plane slightly pleated pleated pleated present n/a pleated of the second type, with a prominent marginal vein with sclerenchyma cap (Rudall & Goldblatt 1991; Goldblatt & Manning 1990). Codonorhiza and Psilosiphon have plane leaves with a visible and sometimes prominent main vein ± centrally placed to form a pseudomidrib, and with the primary and secondary vascular bundles ± opposite. In Lapeirousia at least the basal leaf is prominently closely plicate (corrugate) with the angles of the folds thickened and ribbed, without a pseudo-midrib, and with the major vascular bundles alternate (Goldblatt & Manning 1990). Schizorhiza has a distinctive leaf anatomy in which the main vascular bundles near the centre are ± alternate and surrounded by a complete sclerenchyma bundle sheath, also characteristic of Codonorhiza. The latter is unusual in having falcate, ± obtuse leaves with the margins often undulate to crisped. In both Codonorhiza and Lapeirousia the basal leaf is largest and is sometimes the only foliage leaf, whereas Psilosiphon and Schizorhiza have a fan of a few to several basal leaves. The differences in leaf anatomy and morphology among the four genera (Table 1) are congruent with the molecular phylogeny. Inflorescence: one of the reasons in the past for maintaining a broad circumscription of Lapeirousia has been the trend for inflorescences to form flat- or round-topped false panicles sometimes called corymbs (Table 2). Usually some flowers in ultimate branches are sessile as in a spike, the ancestral inflorescence type in the tribe and subfamily. Such false panicles are universal in Psilosiphon and Schizorhiza and almost so in Codonorhiza (the exception, C. falcata has a spike). All Lapeirousia species with aerial stems have spicate inflorescences. Because C. falcata is retrieved as sister to the remaining species of that genus, a spike may be inferred as ancestral in Codonorhiza as well as for Lapeirousia. It follows that the false panicles in Codonorhiza, Psilosiphon and Schizorhiza may be independently derived. That interpretation is again congruent with the molecular phylogeny.

16 8 S T R E L I T Z I A 35 (2015) A B C FIGURE 3. Leaf anatomy in Lapeirousia and allied genera. A, Lapeirousia odoratissima; B, Psilosiphon bainesii; C, Schizorhiza neglecta. Scale bar: 200 µm. Artist: John Manning. Floral bracts likewise, are consistent with the phylogeny in texture and morphology. In Lapeirousia the large, fleshy to succulent outer bract is usually twice as long as the membranous, 2-keeled inner bract. This contrasts with the leathery, green, subequal outer and inner bracts of Codonorhiza and Schizorhiza. In Codonorhiza the inner bracts are ± obtuse and not 2-keeled or forked apically although sometimes emarginate. Bracts of Psilosiphon represent a third type, relatively soft-textured, often becoming dry from the tips at anthesis, the outer only slightly longer than the inner or sometimes slightly shorter. Flowers: remarkably diverse in form, size and colour, flowers of Lapeirousia and Psilosiphon reflect their adaptation to a range of pollinators. All species have a well-developed perianth tube, shortest in white-flowered L. divaricata and L. spinosa (7 12 mm long) and in some species of Psilosiphon. In species with tubes exceeding 30 mm, the tube is narrow almost throughout. Long-tubed species have one of two predominant colours. One group, exemplified by L. silenoides and L. lewisiana, has flowers intensely pigmented in shades of purple, red or violet, often with white markings on the lower tepals or, in species with radially symmetric flowers, on all tepals. The second group, exemplified by L. anceps and L. fabricii, has a white to beige perianth with red markings on the lower tepals and often pink or red outside, which gives otherwise white flowers a pink cast. The flowers in both of these long-tubed groups are unscented. A third group of long-tubed species, including L. pyramidalis subsp. pyramidalis and L. odoratissima, has white or pale pink to dull yellow (or sometimes dull maroon) flowers usually without markings on the tepals but often intensely scented. The three groups represent the main pollination groups in the genus: long-proboscid flies of the genus Prosoeca; long-proboscid flies in the genera Moegistorhynchus and Philoliche; and moths. In Psilosiphon, P. bainesii and P. schimperi have flowers

17 S T R E L I T Z I A 35 (2015) 9 TABLE 2. Summary of inflorescence, floral and pollen grain differences among genera of the Lapeirousia clade Taxon Inflorescence Bracts Style branches Pollen grains Codonorhiza Lapeirousia Psilosiphon Schizorhiza spike or flat-topped false panicle spike (or basal rosette) flat- or roundtopped false panicle flat- or roundtopped false panicle leathery, subequal, inner entire fleshy, inner smaller, forked at tip soft (leathery), subequal, inner forked at tip firm to leathery, subequal, inner entire Cyanixia spike soft, green, inner smaller, forked at tip Savannosiphon spike leathery, subequal, inner entire Zygotritonia spike firm to leathery, short, becoming dry, inner forked at tip or not deeply divided deeply divided (entire) deeply divided or entire deeply divided entire deeply divided style undivided or branches short, undivided monosulcate, operculum band 1 monosulcate, operculum bands 2 monosulcate, operculum bands 2 monosulcate, operculum band 1 trisulcate polyaperturate trisulcate of the third group, pollinated by moths. Shortertubed species of Psilosiphon and Schizorhiza with darkly pigmented (or white) flowers, including P. erythranthus and P. sandersonii, exhibit no such specialised pollination systems and evidently have a generalist pollination strategy. Flowers are typically bilaterally symmetric (zygomorphic), most likely the ancestral condition, with unilateral stamens held erect or arched beneath the dorsal tepal and the style parallel to or arching over the stamens. All species of the acaulescent sect. Sophronia have radially symmetric (actinomorphic) flowers, sometimes with either relatively short or exceptionally long tubes (Lapeirousia oreogena and L. odoratissima, up to 140 mm long in the latter). Two species of Psilosiphon, P. avasmontanus and P. coeruleus, have radially symmetric flowers but their different vegetative morphology and cytology indicate that they are not immediately related. In Codonorhiza, C. coeruleus and C. fastigiata have radially symmetric flowers, again the derived condition, and possibly shared in the two (the molecular data are ambiguous). Radial floral symmetry has thus evolved at least four times in the four genera and also in the related genus Cyanixia (Goldblatt et al. 2004a). The lability of floral symmetry, perianth pigmentation and perianth tube length is typical of many African genera of Crocoideae (Goldblatt & Manning 2006). We discuss floral longevity under the chapter on Floral Biology (pg. 16). Apart from their orientation, the stamens offer few useful taxonomic characters. Anther and pollen colour is however, variable. Yellow anthers and pollen are most common, but in long-tubed species with white or beige flowers the anthers are often blue or purple, particularly noticeable in Lapeirousia fabricii. The unusual pollen colour may serve as orientation guides for visiting pollinators or as camouflage to deter pollen foraging by bees. In the gynoecium, the level of division of the style is often important, as is the nature of the style branches, either deeply divided or entire (Table 2). Parsimony suggests that deeply divided style branches are ancestral in the Lapeirousia clade and in Watsonieae. Thus ± entire style branches in L. plicata subsp. effurcata and several species of Psilosiphon are reversals of ancestral states. The style itself may divide below, opposite, or beyond the anther apices. The level of division thus may allow for direct contact of the pollen-bearing anthers and permits selfing in self-compatible species when pollination by outside agents fails. Most species with style branches

18 10 S T R E L I T Z I A 35 (2015) in contact with pollen have been experimentally shown to be facultatively autogamous (see discussion of Floral Biology, pg. 16). Capsules and seeds: capsules of the Lapeirousia clade offer few characters of taxonomic significance: they are ± globose to broadly ovoid, somewhat 3-lobed and contain relatively few seeds, the outlines of which are often evident as slight bulging of the capsule walls. Several species of Lapeirousia sect. Chasmatocallis, however, have unusual capsules with prominent locular ridges on the lateral walls, and in L. silenoides, L. verecunda and a few other species, the ridges continue distally as ear-like lobes. Rarely collected, seeds have virtually no practical value for species identification. Nevertheless, seed shape, size and surface features provide valuable information about generic and species relationships (Table 2, 3). Seeds are typically ± globose in Psilosiphon, Schizorhiza and most species of Lapeirousia, with a smooth (rarely slightly wrinkled) surface of domed (colliculate) epidermal cells (exceptionally tuberculate in L. pyramidalis). In Lapeirousia, all members of sect. Sophronia and two of sect. Lapeirousia (L. caudata and L. littoralis) have seeds with a reticulate or rugulose surface and plane outer epidermal cells walls. In contrast, seeds of Codonorhiza are smaller and ellipsoid with rounded ends and a prominent funicular collar at the micropylar end (Goldblatt & Manning 1992). Seeds are mostly mm long in Codonorhiza but up to 1.6 mm long in C. micrantha. In Lapeirousia and Psilosiphon, seeds are typically larger than 1.3 mm diam. and up to 2 mm diam. in L. odoratissima and the L. dolomitica group (Table 3). Most species of sect. Sophronia have unusually large seeds, mm diam., but L. plicata has smaller seeds than are typical of the section, mm diam. Among the largest in the clade, seeds of Schizorhiza are mm diam. Most species of the Lapeirousia group of genera show no adaptations for dispersal in their TABLE 3. Comparison of seed characteristics for genera and sections of the Lapeirousia clade. Seed length is measured from micropyle to chalaza Taxon Seed shape Length (mm) Sculpturing Surface cell outline and orientation Codonorhiza ovoid, collar prominent mostly rugulo-reticulate or reticulate flat, not organised Lapeirousia Sect. Chasmatocallis globose mostly smooth usually ± domed (rarely tuberculate to areolate), scalariform Series Sophronia globose mostly 1 2 rugose to reticulate flat, not organised Sect. Lapeirousia globose smooth (reticulate in 2 spp.) Psilosiphon globose (± angled by pressure) domed, scalariform mostly rugulose to smooth flat or domed, not organised Schizorhiza ± globose rugulose flat, not organised Savannosiphon ± globose rugulose ± domed Cyanixia ± globose ± rugulose ± domed, not organised Zygotritonia globose-ellipsoid ± 2 smooth flat foveate

19 S T R E L I T Z I A 35 (2015) 11 seeds or capsules, but dispersal by wind of the entire plant when dry is important in species of arid areas. In Psilosiphon bainesii, dry plants bearing mature, intact capsules may be blown significant distances across open veld. Similarly, in acaulescent species like L. exilis, L. montana, L. oreogena, L. plicata and even L. kalahariensis and L. oreogena, the dry plants detach from the corm and blow across the veld for some distance, shedding their seeds as the capsules break open on rough ground. Pollen grains: species of Lapeirousia and Psilosiphon (Table 2) have the ancestral type pollen grains for subfamily Crocoideae, ellipsoid in shape with a single broad sulcus in which lies a pair of longitudinal exine bands, the operculum (Goldblatt et al. 1991). In contrast, species of Codonorhiza have pollen grains with a single opercular band, as do both the diploid and tetraploid populations of Schizorhiza neglecta. The one-banded condition is an occasional character state in subfamily Crocoideae, consistent only in Ixia and Xenoscapa (Goldblatt) Goldblatt & J.C.Manning and found in one or more species of Freesia, Romulea Maratti and Thereianthus (Goldblatt & Manning 2009). The closely related genera Cyanixia and Zygotritonia have trisulculate grains (Goldblatt et al. 2004a). Other pollen apertures also occur in the family and are reviewed by Goldblatt & Manning (2009). Pollen grains are always shed singly, and the exine sculpturing is uniformly perforatescabrate, the ancestral and most common condition for Crocoideae (Schulze 1971).

20 12 S T R E L I T Z I A 35 (2015) CHROMOSOME CYTOLOGY Cytology has played an important role in our understanding of the systematics and evolution of Lapeirousia and the segregates, Codonorhiza and Psilosiphon. Early counts (Goldblatt 1971, 1972) revealed little of the variation now evident because the highly asymmetric and bimodal karyotypes include B chromosomes that obscured the base numbers in some species counted. Later work (Goldblatt 1990b) and especially by Goldblatt & Takei (1993) resolved much of the confusion and allowed a reinterpretation of earlier counts, here assembled in Table 4. The patterns we now perceive still need further clarification but correlate reasonably well with some morphological characters and are consistent with our taxonomy. Codonorhiza consistently has 2n = 20 (not including B chromosomes) and a karyotype of one long and nine much shorter chromosome pairs, thus x = 10. Basic chromosome number in Schizorhiza is x = 6 and that in Lapeirousia x = 9 and 8, and the karyotypes are also asymmetric with one long chromosome pair. This pattern characterises sect. Chasmatocallis except for L. divaricata, the two counts for which are 2n = 20, possibly representing B. Sect. Lapeirousia and sect. Sophronia have derived karyotypes with 2n = 16 (again excluding B chromosomes) (Table 4). The chromosome numbers are interpreted here as a descending series because related genera of the tribe have a basic chromosome number of x = 10 (Micranthus, Pillansia, Thereianthus) or 9 (Watsonia) and we infer x = 10 for the entire tribe. Ancestral base number for Lapeirousia is evidently x = 9 and the base of x = 8 is secondary, possibly derived independently at least twice, once in sect. Sophronia and once in sect. Lapeirousia. Psilosiphon has a wide range of diploid chromosome numbers, the highest recorded, 2n = 16 in P. avasmontanus, and the lowest 2n = 6 in some populations of P. bainesii (others 2n = 10). Karyotypes of most species are also strongly asymmetric with one long and the remaining much smaller chromosome pairs. Chromosome rearrangement is evident in P. erythranthus and its immediate allies (Table 4), 2n = 8, which have a relatively symmetric karyotype without gross size differences (Goldblatt & Takei 1993). Neopolyploidy is present in P. erythranthus, some populations of which have 2n = 14 or 12 (hypopolyploid on the base of 2n = 8), and hypopolyploid karyotypes characterise P. bainesii, P. otaviensis and P. schimperi (2n = 10); P. bainesii also has derived populations in central Namibia with 2n = 6. Ancestral karyotypes in Psilosiphon and Schizorhiza are distinctive in consisting of one long chromosome pair, mm long. In most species the remaining smaller pairs are 1 2 mm long. Because of the constant presence of the large chromosome pair and a constancy in chromosome length (which is correlated with quantity of DNA), species of all four genera with high base numbers of 10, 9, 8 or 6 must be regarded as palaeodiploid, as are some lower base numbers. Codonorhiza and Lapeirousia are also palaeodiploid. The northern population of S. neglecta is exceptional in its diploid number of 2n = 24 (tetraploid); one of the two southern populations examined for chromosome number has 2n = 12. A descending dysploid series is a striking feature of Psilosiphon: P. avasmontanus (2n = 16), P. gracilis (2n = 12) and P. sandersonii (2n = 10) maintain the putatively ancestral pattern of one large and the remainder small chromosome pairs, as is the total quantity of DNA per cell (Table 4). P. abyssinicus, P. coeruleus, P. setifolius and the southern populations of P. erythranthus have 2n = 8, but P. erythranthus in Zambia and Malawi, the northern part of its range, has 2n = 14 or 12, with the latter cytotypes having twice as

21 S T R E L I T Z I A 35 (2015) 13 TABLE 4. Chromosome numbers, mean total length of the chromosome complement and number of opercular bands in the pollen grain aperture in Codonorhiza, Lapeirousia, Schizorhiza and Psilosiphon (from Goldblatt 1971; Goldblatt 1990b; Goldblatt & Takei 1993). Karyotype measurements were made from photographs or camera lucida drawings of karyotypes of estimated comparable degree of contraction; all at the same magnification. The asterisk (*) indicates readings corrected for measurements made using a different cytological method by Goldblatt & Manning (1992). New counts are indicated in bold type and voucher data are as follows: C. azurea 2n = 20 +2B (Goldblatt 8016, Malmesbury commonage, NBG); Schizorhiza neglecta 2n = 24 (Goldblatt & Manning 10122, Bains Kloof, NBG). Genome sizes where known are from Goldblatt & Takei 1993) Taxon Diploid number 2n Total chromosome length (inferred ploidy level, x) Genome size (µm), pg Codonorhiza (7 spp.; C. elandsmontana, C. pillansii not counted) C. azurea 20, B 1 C. corymbosa (2x) 1 C. falcata 20 1 C. fastigiata 20 1 C. micrantha * (2x) 1 Psilosiphon (16 spp.; P. erongomontanus, P. masukuensis, L. teretifolius, P. zambesiacus not counted) P. abyssinicus * (2x) 2 P. avasmontanus * (2x) 2 P. bainesii * (4x) 2 P. coeruleus (2x) 2 P. erythranthus (2x) (4x) 2 12, B 54.1 (4x) P. gracilis (2x) P. otaviensis (4x) P. rivularis (4x) 2 P. sandersonii (2x) 2 P. schimperi (4x) P. setifolius (2x) 2 Schizorhiza (1 sp.) S. neglecta (2x) 1 24 (4x) 1 Lapeirousia (27 spp.) Sect. Chasmatocallis (8 spp.) L. anceps 18, B 30.1* (2x) 2 L. angustifolia 18 2 L. divaricata 20 (?18 + 2B) 2 Number of opercular bands in aperture

22 14 S T R E L I T Z I A 35 (2015) TABLE 4 (cont.). Chromosome numbers, mean total length of the chromosome complement and number of opercular bands in the pollen grain aperture in Codonorhiza, Lapeirousia, Schizorhiza and Psilosiphon (from Goldblatt 1971; Goldblatt 1990b; Goldblatt & Takei 1993). Karyotype measurements were made from photographs or camera lucida drawings of karyotypes of estimated comparable degree of contraction; all at the same magnification. The asterisk (*) indicates readings corrected for measurements made using a different cytological method by Goldblatt & Manning (1992). New counts are indicated in bold type and voucher data are as follows: C. azurea 2n = 20 +2B (Goldblatt 8016, Malmesbury commonage, NBG); Schizorhiza neglecta 2n = 24 (Goldblatt & Manning 10122, Bains Kloof, NBG). Genome sizes where known are from Goldblatt & Takei 1993) Taxon Diploid number 2n Total chromosome length (inferred ploidy level, x) Genome size (µm), pg Number of opercular bands in aperture L. jacquinii 18, B 25.7 (2x) 2 L. exilis B 29.8 (2x) 2 L. pyramidalis subsp. pyramidalis (2x) 2 subsp. regalis 18 2 L. silenoides 18, B 27.1 (2x) 2 L. verecunda 18 2 Sect. Sophronia (5 spp.; L. kalahariensis, L. montana not counted) L. odoratissima (2x) 2 L. oreogena 16, B 28.8 (2x) 2 L. plicata (2x) 2 Sect. Lapeirousia (14 spp.; L. kamiesmontana, L. macrospatha, L. purpurea, L. simulans, L. tenuis not counted) L. arenicola (2x) 2 L. barklyi 16 2 L. caudata (2x) 2 L. dolomitica 16, B 2 L. fabricii B 2 L. lewisiana (2x) 2 L. littoralis * (2x) L. spinosa 16 2 L. violacea 16 2 much DNA per cell as those with 2n = 8 (Goldblatt 1990b). They must, therefore, be regarded as hypopolyploid (perhaps derived from a hypothetical 2n = 16, so far not found in the wild). P. rivularis (one Zambian population counted) also has 2n = 12, but P. otaviensis and P. schimperi have 2n = 10 and P. bainesii has cytotypes with 2n = 10 and 6 (the latter recorded for two Namibian populations and the lowest chromosome number recorded in the Iridaceae). All four species have genomes with total chromosome length and, where known, DNA quantity per cell that marks them as polyploid, but with radically reorganised karyotypes. They are in fact polyploid but with lower chromosome numbers than their ancestral stock. The most parsimonious explanation is that the latter four species share a common polyploid ancestor. Our molecular phylogeny is not informative on this issue as too few species of Psilosiphon have been sequenced.

23 S T R E L I T Z I A 35 (2015) 15 Among the genera of Watsonieae believed to be most closely related to the Lapeirousia group of genera, Cyanixia has x = 10 and Zygotritonia x = 8, both with strongly bimodal karyotypes consisting of one large pair and the remaining pairs much smaller, thus consistent with the ancestral pattern for the entire Lapeirousia clade (Goldblatt 1989, Goldblatt et al. 2004a). Savannosiphon has x = 7 but a very different karyotype of large, medium and small chromosomes (Goldblatt & Marais 1979), without strong bimodality. The pattern of chromosome evolution, viewed in the light of the DNA-based phylogeny, is consistent with the scenario outlined above in which x = 10 and a bimodal karyotype are ancestral. Additional counts will help clarify the patterns already evident. The variability in number, even within some species of Psilosiphon, indicates that the genus is not adequately sampled. It would be particularly useful to determine the chromosome number of the Namibian populations of P. rivularis in light of the counts for the Zambian populations, which have 2n = 12. Counts are also needed for additional populations of P. sandersonii and P. gracilis while P. erongoensis remains uncounted.

24 16 S T R E L I T Z I A 35 (2015) FLORAL BIOLOGY Reproductive compatibility: compatibility relations in Lapeirousia, although inadequately known, are informative: at least one or more populations of L. anceps, L. arenicola, L. fabricii, L. jacquinii, L. oreogena, L. simulans and L. violacea have been determined to be self-compatible and facultatively autogamous using pollinator exclusion experiments (Goldblatt et al. 1995; Manning & Goldblatt 1997; Pauw et al. 2008). Notably, all of these species are either known or inferred to be specialised for long-proboscid fly pollination. Similar studies of bee pollinated L. barklyi and L. spinosa, long-proboscid fly pollinated L. lewisiana, L. pyramidalis subsp. regalis and L. silenoides, and moth pollinated L. pyramidalis subsp. pyramidalis show these species to be self-incompatible and unable to set seed even when self-pollen is transferred by hand to receptive stigmas in those species in which the stigmatic surfaces are not normally in contact with anthers and pollen. The association of selfcompatibility and often autogamy (facultative when species have regular pollinator visits) with specialist pollination strategies is not uncommon in the southern African Iridaceae (Goldblatt & Manning 2006) and is documented, for example in Babiana (Anderson et al. 2005; Goldblatt & Manning 2007; De Waal et al. 2011) and Sparaxis (Goldblatt et al. 2000; Goldblatt & Manning 2013). The phenomenon evidently represents a failsafe mechanism when pollinators are absent or visits are rare or unreliable. We suspect that L. arenicola and L. simulans may be truly autogamous. No pollinators have ever been recorded on flowers of either of these long-tubed species and both have flowers with a shorter tube and smaller tepals than typical fly pollinated taxa. Moreover, they flower earlier in the season than the long-proboscid flies, which are inferred to be their pollinators, are on the wing. Floral longevity: like most Crocoideae, flowers of the Lapeirousia group of genera are longlived, lasting at least three and up to five days (Goldblatt et al. 1995). Known exceptions are L. odoratissima and P. bainesii, both of which have scented, pale-coloured flowers and are known or inferred to be pollinated by sphinx moths. The tepals of both species unfold in the late afternoon, at which stage the flowers produce a strong fragrance, and remain open all night but begin to wilt early to mid-morning. Flowers lasting at most 24 hours are unusual in Crocoideae and we know of only two comparable situations in the subfamily in the Namibian Gladiolus diluvialis Goldblatt & J.C.Manning and Savannosiphon euryphyllus, both also evidently sphinx moth-pollinated (Manning et al. 2011; R.E. Gereau, pers. comm. 2013). Flowers of other sphinx moth-pollinated Crocoideae, notably Lapeirousia pyramidalis subsp. pyramidalis and Gladiolus tristis L., last at least three days. Flowers of those species examined for the character are protandrous. The anthers dehisce first, two to four hours after tepals unfold, but pollen remains in place on the anther surface until removed by insect activity, either passive or active, or is washed away by rain. The style branches, at first appressed to one another, only curve outward on the following days, exposing glistening, ciliate surfaces. Pollination: among the genera of the Lapeirousia clade, only the pollination systems of Lapeirousia have been extensively studied. As suggested by the remarkable floral variation in the genus, it exhibits a diversity of pollination strategies, ranging from generalist systems that rely on a mix of different bee families, butterflies, and even settling moths and bee-flies, to specialist systems using only pollinators of a single class, viz. sphinx moths or long-proboscid flies, or even a single species of pollinator (Goldblatt et al. 1995; Manning & Goldblatt 1996). The reward for most pollinators is nectar, retained within the perianth tube and secreted from septal nectaries, the pores of which open at the apex of the ovary

25 S T R E L I T Z I A 35 (2015) 17 at the base of the style. Nectar is sucrose rich or sucrose dominant (Goldblatt et al. 1995), irrespective of pollination system, and nectar quantity correlates closely with perianth tube length. Most striking in Lapeirousia are the longtubed species with purple, violet or cerise to magenta tepals with white markings on a dark background. These species, including L. jacquinii, L. dolomitica, L. oreogena, L. pyramidalis subsp. regalis, L. silenoides and L. violacea, are pollinated by two species of Prosoeca (Nemestrinidae), P. peringueyi in Namaqualand and northwestern Western Cape, and a second, unnamed Prosoeca species in the Bokkeveld Mtns and Plateau (Goldblatt et al. 1995). These flies, with probosces (15 )25 45 mm long, feed on nectar held within perianth tubes mm long. Pollen is deposited on the frons or dorsum. We infer that L. fabricii subsp. violacea, L. lewisiana and L. purpurea also belong to this pollination guild in light of their early flowering, floral morphology and pigmentation. A second group of long-tubed species in Lapeirousia with white to pale yellow flowers marked with pink or red near the tepal bases is pollinated by a second series of long-proboscid flies, including Moegistorhynchus longirostris (Nemestrinidae) and Philoliche gulosa (Tabanidae). Species with this pollination system include L. anceps, L. fabricii and by inference because of similarly shaped and coloured flowers, L. kamiesmontana, L. macrospatha and L. verecunda. Flies of both these pollination groups sometimes hover while feeding but more often grasp the tepals, especially the lower three, as they forage. In Psilosiphon, the northern Namibian P. otaviensis is a likely candidate for long-proboscid fly pollination but this pollination strategy has not been documented within its range to date. Fly mouthparts may be up to 95 mm in M. longirostris and up to 35 mm in P. gulosa and P. rostrata (Goldblatt & Manning 2000). Moth pollination is likely for long-tubed species with white flowers that are open and strongly scented at night, the most striking examples of which are Lapeirousia odoratissima (sect. Sophronia) and Psilosiphon schimperi, which have perianth tubes mm long. The night blooming cycle in L. odoratissima adds support to the inference of moth pollination. Psilosiphon bainesii most likely has a similar pollination system and we have seen but not captured sphinx moths visiting its flowers in Namibia. In Lapeirousia moth pollination also occurs in L. pyramidalis subsp. pyramidalis and is likely for L. caudata, L. kalahariensis and L. littoralis. Hawk moths are the likely pollinators. Codonorhiza micrantha, which has small, dull-coloured flowers, but a strong, clove-like scent at night, is almost certainly also pollinated by moths, specifically settling moths (e.g. Noctuidae) because of the small size of the flowers and relatively short, narrow perianth tube that are consistent with these pollinators (Goldblatt et al. 2004b). The remaining species appear to have less specialised pollination strategies. The shorttubed, open flowers of species such as Psilosiphon erythranthus and its allies, P. sandersonii and the Codonorhiza corymbosa group have a range of visitors including butterflies, female bees of several families, worker honey bees and even wasps, but the recorded pollinators of C. azurea are anthophorine bees (Anthophora) and hopliine beetles. L. divaricata and L. spinosa, which have short-tubed, gullet flowers, are known to be pollinated by large-bodied anthophorine bees and honey bees and in the late afternoon by noctuid moths (Goldblatt et al. 1995), possibly a bimodal system sensu Manning & Goldblatt (2005). In our study population of P. coeruleus, which has secondarily radially symmetric flowers, the only pollinator captured was a small bee (family unknown), which nestles in the flowers (which lack nectar) and presumably forages for pollen. While much research remains to be done to determine the pollination systems of several species, especially of Psilosiphon in summer rainfall southern and tropical Africa, it is clear that pollination systems in the Lapeirousia clade are even more labile than they are in most other genera of African Iridaceae (Goldblatt & Manning 2006). Matching the pollination system to the phylogeny (Figure 1) shows that the Prosoeca peringueyi pollination system evolved five or possibly six times (if ancestral in the genus), and the Moegistorhynchus Philoliche pollination system three times. Hawk moth pollination evidently evolved independently in L. odoratissima, L. pyramidalis and L. caudata L. littoralis (and possibly also in L. kalahariensis) in Lapeirousia as well as in Psilosiphon bainesii and P. schimperi (or their com-

26 18 S T R E L I T Z I A 35 (2015) mon ancestor). Long-proboscid fly pollination is not recorded in Psilosiphon but seems likely in P. masukuensis (perianth tube length mm). This lability of pollination system in Lapeirousia was first noted by Goldblatt & Manning (1996) and has been calculated as one shift in strategy for every 1.5 species by Forest et al. (2014), a level of lability unprecedented in Iridaceae and among all flowering plants. In Gladiolus and Babiana (Iridaceae), one pollination shift has been inferred for every five to six species (Goldblatt & Manning 2006), an estimate later confirmed for Gladiolus by Valente et al. (2012). Shifts in pollination system in Lapeirousia are closely associated with speciation events, but whether they are the direct cause of these events or are reproductive reinforcement following geographic isolation or edaphic shifts remains uncertain.

27 S T R E L I T Z I A 35 (2015) 19 GEOGRAPHY The Lapeirousia clade is entirely restricted to mainland sub-saharan Africa and is centred in southern Africa. The range of Codonorhiza is exclusively in the Cape Floristic Region (CFR), with C. micrantha the most widespread species in the genus, extending through the Cape Fold Mountains from the Gifberg in the north to the Langeberg in the southeast. The remaining species are restricted to the southwest of the CFR, mostly on the coastal lowlands and lower slopes of the mountains although C. falcata is essentially montane. Only C. corymbosa and C. pillansii occur on the Cape Peninsula. The monospecific Schizorhiza is also restricted to the CFR, known from just three populations in the extreme southwest of Western Cape. Lapeirousia, largely a genus of the southern African winter rainfall zone, has a far wider range. Centred in central Namaqualand in Northern Cape, South Africa, several species extend south through the Cederberg to the Western Cape coastal plain, but only L. anceps reaches the Cape Peninsula, extending along the southern coast to near Mossel Bay. Several species extend to the north of Namaqualand into the winter rainfall belt of southwestern Namibia, and L. plicata ranges from interior Western Cape through the Great and Upper Karoo to southern Namibia. L. plicata subsp. effurcata is surprising in occurring in the southern Cape limestone country south of Riversdale as well as in the western Little Karoo. The remaining L. caudata, L. kalahariensis and L. odoratissima are species of the southern African summer rainfall area, the latter species extending from central Namibia to Angola, Zambia, Malawi and Tanzania. The range of L. caudata broadly corresponds to that of L. odoratissima but reaches further south into Gordonia, the margins of Bushmanland and Limpopo in South Africa, with isolated populations in southern Mozambique, an exceptional disjunction for sub-saharan Iridaceae. Psilosiphon is exclusively a genus of summer rainfall sub-saharan Africa, its range virtually complementing those of Codonorhiza, Lapeirousia and Schizorhiza. Species extend from central Namibia and the northern provinces of South Africa across south tropical Africa. The most widespread species of Psilosiphon, the variable P. erythranthus, crosses the equator into northern Nigeria. Almost as widespread, P. schimperi extends in suitable, seasonally moist habitats from central Namibia and Botswana to Ethiopia, Somalia and possibly Yemen. P. abyssinicus is restricted to the Ethiopian highlands, P. setifolius to Malawi and southwestern Tanzania, P. teretifolius to Congo, and P. zambeziacus to western Zambia and adjacent Angola. The clade sister to Psilosiphon comprises the Socotran endemic, Cyanixia socotrana, and the south tropical African Savannosiphon euryphyllus. Zygotritonia, which has yet to be sequenced, is almost certainly immediately allied to these two genera: it shares with the tribe axillary corm development, with Cyanixia and Savannosiphon corms rounded at base and shaggy, brown fibrous tunics, plicate leaves, and with Cyanixia trisulcate pollen grains, unique in Crocoideae. The six species of Zygotritonia extend from West Africa to Ethiopia and south to Tanzania, Malawi and Zambia (Goldblatt 1989; Demissew et al. 2003; Goldblatt et al. 2015). Genera immediately related to the Lapeirousia clade (tribe Watsonieae) are all southern African, either restricted to the Cape Floristic Region or with their centre here. Codonorhiza, sister to the remaining members of the Lapeirousia clade is also a Cape genus. The origins of the Lapeirousia clade thus lie to the south, a pattern repeated in the few genera of Crocoideae that extend outside the Cape area.

28 20 S T R E L I T Z I A 35 (2015) ETHNOBOTANY As early as the 17th century, European explorers in southern Africa noted the use of corms of Lapeirousia and its allies as a food, for example in the manuscript Codex Witsenii (Wijnands et al. 1996), where the Khoisan vernacular names cabong and chabi were used for the relatively common L. jacquinii and L. fabricii. It was only in the early 20th century that the first published documentation appeared when, in 1912, Kurt Dinter reported that corms of Lapeirousia and Psilosiphon species were used as a food for humans. Dinter mentioned specifically that corms of L. caudata, L. odoratissima (as L. juttae), P. coeruleus (as L. coeruleus) and P. schimperi (as L. uliginosa) were valued foods eaten after roasting in hot ashes. Dinter provided vernacular names onduvi (ozonduvi pl.) (Herero) and garib (Khoi) for these species. Dinter s observation has been confirmed repeatedly by plant collectors and ethnobotanists, notably R. Story for the!kung Bushmen (P. coeruleus Story 6121 and L. caudata Story 6162). Story also noted that corms of P. coeruleus were a staple, eaten after roasting in hot ash. In addition, a collection of P. gracilis (Seydel 3419) documents the edibility of this species. The!Kung Bushmen are also reported to eat the corms of L. odoratissima for their water content (Marshall 1976), which seems unlikely, and to eat corms of P. bainesii after baking, or to pound the cooked corms into a meal, which is then eaten as a gruel with water (Fox & Young 1982). The ethnobotanist R. Rodin (1985) reported that corms of Psilosiphon coeruleus, P. bainesii (as Lapeirousia vaupeliana) and P. schimperi (as L. cyanescens) were eaten both raw and roasted by the Kwanyama Ovambo in northern Namibia. Leffers (2003) confirms the use of three species by the Jul hoansi subgroup of the San in northeastern Namibia, P. bainesii, P. coeruleus and P. schimperi, with the vernacular names g xàrú, g àrì and g!úíh respectively. Corms are shucked and then roasted in hot ash and either consumed directly or pounded and stewed with water or more recently, milk, usually without the addition of other ingredients. Lapeirousia and Psilosiphon appear to have little value to human communities outside Namibia and presumably adjacent Angola and Botswana. There are, however, isolated reports that corms of P. erythranthus are eaten in the Shire Highlands of southern Malawi in time of great famine (Buchanan 426). Collection notes (Simpathu 60) from Victoria Falls, Zimbabwe, also indicate that corms of P. erythranthus are eaten raw. In the light of the amply documented use of the corms of several Namibian species of Lapeirousia and Psilosiphon as food, the report cited, without source, by Watt & Breyer-Brandwijk (1962) that P. coeruleus is poisonous, must be regarded as doubtful at best, particularly as tests performed on a frog for cardiac glycoside action proved negative.

29 S T R E L I T Z I A 35 (2015) 21 PHYLOGENY AND SUBGENERIC CLASSIFICATION Analyses of DNA sequences of 10 plastid regions and one nuclear gene (Forest et al. 2014) have provided surprising and unanticipated insights into phylogenetic relationships among the Lapeirousia clade, prompting us to radically dismember Lapeirousia sensu Goldblatt (1972) in order to maintain monophyly of the genera. Results (e.g. Figure 1; Forest et al. 2014) show that two genera Cyanixia and Savannosiphon are nested in Lapeirousia s. lat. where they are retrieved together as sister to Lapeirousia sect. Paniculatae (one of two sections of subg. Paniculatae) [following the classification of Gold blatt & Manning 1990]. Sect. Fastigiatae of subg. Paniculatae is retrieved as sister to the remainder of the entire Lapeirousia clade, with subg. Lapeirousia placed as sister to sect. Paniculatae plus Cyanixia and Savannosiphon. Our preferred solution to restoring monophyly is the recognition at generic rank of both sect. Paniculatae and sect. Fastigiatae. As already outlined in the foregoing text, sect. Paniculatae is here recognised as the new genus Psilosiphon, sect. Fastigiatae as Codonorhiza, and the phylogenetically isolated L. neglecta as the monospecific Schizorhiza. This solution is absolutely congruent with morphology. Although the corms of these three new genera plus Lapeirousia s. str. have a flat base (the primary justification for their initial treatment as a single genus), corm tunic structure differs in all four genera, as do the floral bracts. Seeds of Codonorhiza are unique in Watsonieae and the pollen grains of Codonorhiza and Schizorhiza, which have a single banded operculum, differ from the 2-banded operculum in Lapeirousia and Psilosiphon. Chromosome cytology likewise differs in all three genera. Species relationships in Codonorhiza are well resolved and supported (Figure 1). Codonorhiza falcata is consistently retrieved as sister to the remaining species, a relationship that accords with its spicate inflorescence, the plesiomorphic condition in Watsonieae the remaining species have derived, round- or flat-topped, false panicles composed of multiple branches each bearing just one or two flowers. The two species C. corymbosa and C. fastigiata have (evidently independently) derived radially symmetric flowers, and C. micrantha is nested within the otherwise blue-flowered C. corymbosa group of species and sister to C. corymbosa. Only C. pillansii was unavailable for inclusion in our molecular analyses. Lapeirousia in its restricted sense is strongly supported as monophyletic (1.0 PP), and comprises three main clades or species groups. The first, the L. silenoides clade with eight species, includes L. angustifolia and the two subspecies of L. pyramidalis in a strongly supported subclade that is characterised morphologically by unique, retuse floral bracts. A close association of the long-tubed L. jacquinii with short-tubed L. divaricata (1.0 PP) is the only surprise here. Flowers of the latter closely resemble those of L. spinosa evidently convergent as their corms are very different. Corms with margins bearing minute teeth and unusual transparent windows in the floral bracts are the only evident morphological features linking L. jacquinii and L. divaricata. A second major clade (1.0 PP) in Lapeirousia comprises the four acaulescent species included in the study: L. oreogena, L. plicata, L. montana and L. odoratissima (one additional species, L. kalahariensis, was not available). Within this clade, the tropical African and hyper long-tubed L. odoratissima is retrieved as sister to the shorter-tubed southern African species. The acaulescent species all have a radially symmetric flower and seeds with a sculptured surface, both derived characters. The third major clade includes the remaining species. Surprisingly, L. caudata, L. littoralis

30 22 S T R E L I T Z I A 35 (2015) and L. barklyi are retrieved as sister to the remaining species of Lapeirousia in the combined analysis, a result that receives no support from morphology or chromosome cytology and defies explanation. In contrast, the association of white-flowered and short-tubed L. spinosa and L. tenuis (1.0 PP) is congruent with corm and floral morphology, as is that of L. arenicola and L. macrospatha (1.0 PP). We are somewhat surprised that the molecular analyses do not show a close relationship between L. dolomitica and L. lewisiana, the latter until now treated as a subspecies of the former, nor with L. kamiesmontana, until now regarded as merely a white-flowered race of subsp. lewisiana. We accept the results of the molecular analyses as support for the recognition of the three sets of populations as separate species. In the taxonomic account, however, we group the species in sequence for they are linked by specialised corm tunics and a long perianth tube, and L. lewisiana and L. kamiesmontana are further linked by unique, sparsely hairy leaves. The weak association of L. lewisiana and L. simulans in the molecular analysis (0.31 PP) is not supported by morphology the corms of L. simulans lack the specialised spines present in L. lewisiana. An association between L. fabricii and L. kamiesmontana is likewise weakly supported (0.55 PP), but in this case the two species share similar corms, suggesting a reflection of true relationship. Lastly, the species of Lapeirousia here segregated as Psilosiphon comprise a clade (1.0 PP) sister to Cyanixia plus Savannosiphon (0.85 PP). Relatively few species of Psilosiphon were included in our study (8 out of 15) but notably the hypopolyploid L. otaviensis and L. schimperi (both 2n = 10) are retrieved as sisters (1.0 PP) in all analyses. The two species with radially symmetric flowers, P. avasmontanus and P. coeruleus, are retrieved as only distantly related within the genus, a result congruent with their very different corm tunics and branching patterns. Undoubtedly one of the most surprising results of the molecular analysis is the position of Lapeirousia neglecta (now the monospecific genus Schizorhiza). This taxon, from the southern African winter-rainfall region and until now included in the otherwise tropical and summer rainfall southern Africa sect. Paniculatae, is retrieved with moderate support (0.84 PP) as sister to Lapeirousia + Psilosiphon + Cyanixia + Savannosiphon. Re-examination of its morphology, leaf anatomy, and pollen grains provides grounds for accepting this result. The corm shape, ellipsoid with a small flat base, and corky tunics are unique and are discordant with Psilosiphon, as are the simple leaf margins and the single-banded pollen operculum. Its chromosome base number, x = 6, which is shared in Psilosiphon with P. gracilis, must be seen as convergent. We propose the following infrageneric classification for Lapeirousia, based on the high level of congruence among the molecular phylogeny, morphology and cytology. The L. silenoides clade (1.0 PP) merits sectional rank, as sect. Chasmatocallis, and we likewise treat the clade (1.0 PP) of acaulescent species with radially symmetric flowers as sect. Sophronia. We treat the remaining species as sect. Lapeirousia despite their being retrieved as two separate clades, the L. littoralis clade (1.9 PP) and the L. fabricii clade (1.0 PP). We see no morphological or anatomical reason for their separation taxonomically and both have the derived basic chromosome number x = 8. Additional sampling of these species may resolve the current incongruence of morphology and molecular data. Codonorhiza is too small a genus for an infrageneric classification to be useful, and until more species of Psilosiphon are obtained for sequencing, establishing an infrageneric classification of the genus is premature. We continue to recognise the remaining genera of the Lapeirousia clade of Watsonieae, Cyanixia, Savannosiphon and Zygotritonia.

31 S T R E L I T Z I A 35 (2015) 23 SYSTEMATICS Key to genera of tribe Watsonieae 1a Corm bell-shaped, ellipsoid or obconic (triangular in outline), with flat base and roots emerging from margin of base: 2a Corm ± ellipsoid, diameter of flat base narrower than midline; lower axils of stem bearing multiple cormlets; inflorescence a many-branched false panicle Schizorhiza 2b Corm bell-shaped or obconic, widest at base; lower axils of stem with a single cormlet or without cormlets; inflorescence a spike or a false panicle: 3a Corm tunic layers woody, base sometimes serrated to spiny; leaves corrugate (plicately ridged); inflorescence a spike or a crowded rosette when plants lack aerial stem; floral bracts large, fleshy, the inner significantly smaller than outer; seeds globose... Lapeirousia 3b Corm tunic layers not woody, composed of compressed fibres, the outer layers often becoming loosely fibrous with age, base never serrated to spiny; leaves plane or terete; inflorescence a spike or a many-branched false panicle, the individual flowers always sessile (subtended by a pair of opposed bracts at base of ovary); floral bracts subequal, the inner slightly shorter or slightly longer than outer; seeds ovoid or globose: 4a Floral bracts green, firm and ± leathery, apices ± rounded or emarginate; seeds ovoid with surface reticulate to rugose; pollen with 1-banded operculum... Codonorhiza 4b Floral bracts green to dry, soft-textured, often becoming membranous, apices acute; seeds globose with smooth surface; pollen with 2-banded operculum... Psilosiphon 1b Corms globose to obconic with rounded base and roots emerging from lower half of the corm: 5a Style branches deeply divided: 6a Leaves ± clustered in a basal fan, with thickened, hyaline margins; flowers medium to large, usually at least 20 mm long, never blue; bracts usually green, sometimes becoming dry.watsonia 6b Leaves superposed, only lowermost basal, without thickened, hyaline margins; flowers small to medium, mostly less than 30 mm long, blue or mauve to white; bracts ± dry: 7a Spikes distichous; flowers small, < 12 mm long, crowded in dense spikes; bracts solid below with broad membranous margins... Micranthus 7b Spikes spiral; flowers > 15 mm long, with a ± straight perianth tube; bracts without broad membranous margins thereianthus 5b Style branches undivided or at most notched apically: 8a Inflorescence a panicle, individual flowers always pedunculate; flowers radially symmetric; perianth bright orange; leaves firm, leathery, plane; plants of the southwestern Cape.. Pillansia 8b Inflorescence a simple or branched spike and individual flowers sessile, rarely inflorescence reduced to a single flower; flowers actinomorphic or zygomorphic; perianth variously coloured; leaves shallowly pleated and relatively soft-textured: 9a Flowers radially symmetric; perianth pale blue; plants of Socotra... Cyanixia 9b Flowers zygomorphic; perianth shades of red to purple or white: 10a Style undivided or with short, entire branches; flowers relatively small and perianth tube shorter than dorsal tepal; perianth shades or red to purple or partly white; plants of tropical Africa... Zygotritonia 10b Style 3-branches, each branch deeply divided; flowers large and perianth tube much longer than dorsal tepal; perianth white... Savannosiphon

32 24 S T R E L I T Z I A 35 (2015) Systematics of Lapeirousia Lapeirousia Pourr. in Histoire et Mémoires de l Académie Royale des Sciences, Toulouse 3: 79 (1788). Ovieda Spreng.: 258 (1817), illegitimate homonym and superfluous name, not Ovieda L. (1753). Meristostigma A.Dietr.: 593 (1833), superfluous name. Peyrousia Poir.: 363 (1826), possible orthographic variant of Lapeirousia (also spelled Lapeyrousia), or superfluous name. Type: L. compressa Pourr. (= L. fabricii (D.Delaroche) Ker Gawl.). Sophronia Licht. ex Roem. & Schult.: 482 (1817). Type: S. caespitosa Licht. ex Roem. & Schult. (= Lapeirousia plicata (Jacq.) Diels). Chasmatocallis R.C.Foster: 40 (1939). Type: C. macowanii R.C.Foster (= Lapeirousia divaricata Baker). Named in honour of the Compte de la Peyrouse, 18th century patron of French science and contemporary of the Abbe Pourret, who described the genus. Deciduous geophytes. Corms bell-shaped with flat base, rooting from lower edges; axillary in origin; tunics of hard, woody layers, sometimes spiny at base. Foliage leaves few, lowermost linear to sword-shaped, inserted on stem near ground level, closely corrugate, upper leaves progressively smaller. Stem aerial or sometimes subterranean, compressed, often angled, angles sometimes winged, simple or branched. Inflorescence a spike or when acaulescent, a dense rosette; bracts paired and opposed, outer green and firm to ± succulent, sometimes ridged, keeled, crisped or toothed; inner much shorter, usually hidden by outer, 2-keeled and often partly membranous, forked at apex. Flowers zygomorphic or actinomorphic in acaulescent species, often salver-shaped, blue, purple, red, pink or white, lower or all tepals usually with contrasting darker or lighter markings, sometimes sweetly scented; perianth tube cylindric or obliquely funnel-shaped, short to extremely long; tepals subequal or unequal, with dorsal largest. Stamens unilateral and arcuate or symmetrically disposed; filaments slender, free; anthers linear, splitting longitudinally. Ovary ± ovoid; style filiform, 3-branched, style branches usually forked for ± 1 / 2 their length, occasionally not divided. Capsules ovoid to oblong, cartilaginous, often showing outline of seeds. Seeds globose, flattened at chalazal end, either smooth with surface cells domed (rarely tuberculate) or rugulo-reticulate or reticulate with surface cell walls flat. Basic chromosome number x = 9; diploid numbers 2n = 18, 16. Species 27; mainly of winter-rainfall, western South Africa and southwestern Namibia, also central South Africa and south tropical Africa, mainly in sandy soils or rock outcrops. Key to sections Notes: All measurements are from living plants; floral parts may shrink up to 20% depending on care taken with pressing specimens. Measure corm diameter at corm base, bract length in middle of spike if there is considerable variation along a spike. Seeds may decrease in size with age through desiccation; dimensions are for capsule-matured seeds no more than a year old. Measure perianth tube from base to point of separation of tepals, thus including narrow and wide parts of tube; tepal length is from point of separation from tube to tepal tip. 1a Corm bell-shaped, basal margin bluntly ridged to crenate or minutely spiny with fine teeth directed downward (not with laterally directed spines or teeth); flowers radially symmetric or zygomorphic: 2a Plants with aerial stems (unless depauperate); flowers zygomorphic sect. Chasmatocallis 2b Plants acaulescent; flowers radially symmetric sect. Sophronia 1b Corm bell-shaped to obconic, thus ± triangular in profile, basal margin serrated to spiny and bearing short to long teeth; flowers zygomorphic sect. Lapeirousia

33 S T R E L I T Z I A 35 (2015) 25 Key to species 1a Plants tufted, ± acaulescent with axis contracted above ground level and inflorescence congested; flowers radially symmetric: 2a. Floral bracts mm long; perianth tube mm long L. odoratissima 2b. Floral bracts mm long; perianth tube mm long: 3a. Flowers violet, lower 1 / 3 of tepals dark violet to black with wedge-shaped white mark in centre of dark zone; perianth tube mm long L. oreogena 3b. Flowers whitish or pale blue, sometimes with dark blue to purple mark near base of tepals; perianth tube mm long: 4a. Perianth tube mm long; tepals 7 9 mm long; anthers mm long, blue or whitish L. plicata 4b. Perianth tube (33 )35 60 mm long; tepals mm long; anthers 3 4 mm long, white or pale yellow: 5a. Flowers ± white; leafy upper parts of floral bracts ± straight, channelled to apex, not ribbed L. kalahariensis 5b. Flowers white or pale blue; leafy upper parts of floral bracts falcate, unifacial above and strongly ribbed L. montana 1b Plants with aerial stems unless depauperate (e.g. L. exilis, L. arenicola); flowers zygomorphic with stamens unilateral: 6a Flowers predominantly white to creamy white, often flushed pink outside; perianth tube 7 15 mm long: 7a Corm bell-shaped, basal rim minutely serrated or entire and lobed: 8a Perianth tube 7 10 mm long, obliquely funnel-shaped; corm margin with short fine teeth directed downward; plants of the Cederberg, Gifberg and Bokkeveld Mtns...1. L. divaricata 8b Perianth tube mm long, ± slender throughout; corm margins entire; plants of the Knersvlakte L. angustifolia 7b Corm obconic (triangular in profile), basal rim bearing prominent, spreading spines: 9a Dorsal tepal mm L. spinosa 9b Dorsal tepal mm L. tenuis 6b Flowers variously coloured; perianth tube mm long: 10a Corm bell-shaped, basal rim bluntly ridged to crenate or minutely serrate with teeth directed downward: 11a Outer floral bracts obtuse to retuse when opened flat L. pyramidalis 11b Outer floral bracts tapering to an acute apex: 12a Flowers deep violet or cherry-red to magenta, lower tepals with white to yellow markings near base: 13a Floral bracts with median keel, without white patches; flowers cherry-red to magenta; perianth tube mm long L. silenoides 13b Most or all floral bracts 2-keeled, fenestrate with white patches; flowers violet to purple, lower tepals streaked with white; perianth tube mm long L. jacquinii 12b Flowers not as above: 14a Outer floral bracts mostly 6 10 mm long; lower tepals ± linear to narrowly lanceolate with a short, filiform tooth near base L. anceps 14b Outer floral bracts (10 )12 25 mm long; lower tepals broadly lanceolate to ovate, with or without teeth: 15a Perianth white to pink, pale blue or maroon-brown; not marked with red on lower tepals:

34 26 S T R E L I T Z I A 35 (2015) 16a Perianth tube slender below, ± abruptly expanded to wider upper part; tepals ascending below to form part of floral cup, spreading distally; tepals predominantly pink to lilac with yellow nectar guides on lower tepals; style dividing opposite or beyond anther tips L. barklyi 16b Perianth tube slender ± throughout, slightly wider distally; tepals spreading from base; tepals predominantly white to ivory or pale blue; style dividing between base and middle of anthers: 17a Plants slender, less than 60 mm high, often unbranched; flowers pale blue (rarely ± white) with violet streaks on lower tepals L. exilis 17b Plants usually > 100 mm high, usually repeatedly branched from base; flowers white, pale yellow, yellow-brown or dull maroon, usually without contrasting markings on lower tepals: 18a Perianth tube mm long; tepals mm; bracts (12 )15 20( 25) mm long L. littoralis 18b Perianth tube (25 )30 45( 70) mm long; tepals mm; bracts 10 18( 23) mm long L. caudata 15b Perianth white to pink or creamy beige, lower tepals always marked with red to brown near base: 19a Perianth tube mm long; corm with basal margins entire to crenately lobed L. verecunda 19b Perianth tube (10 )12 35 mm long; corm with basal margins minutely serrated: 20a Lower tepals ± 13 5 mm L. macrospatha 20b Lower tepals mm: 21a Corm relatively shallowly buried, and underground part of stem not bearing cormlets at nodes; perianth tube mostly mm long L. arenicola 21b Corm relatively deeply seated, with underground part of stem bearing cormlets at nodes; perianth tube mostly mm long L. simulans 10b Corm obconic (triangular in profile) with margins bearing prominent, spreading spines: 22a Flowers white to beige or blue-mauve (rarely pink), usually with red markings on lower tepals: 23a Perianth tube slender below, expanded in upper 5 10 mm into a wider gullet; style dividing opposite base to upper 1 / 3 of anthers; basal leaf smooth L. fabricii 23b Perianth tube slender, ± uniformly wide throughout; tepals spreading from base; lower tepals not bearing a tooth at base; style dividing opposite or above anther tepals; basal leaf sparsely hairy L. kamiesmontana 22b Flowers predominantly blue, violet, purple or magenta, variously marked on lower tepals: 24a All or only median lower tepal with tooth-like projection near base: 25a Only lower median tepal with tooth near base; upper lateral tepals strongly recurved; lower tepals with white V-shaped mark near base on darker violet background L. purpurea 25b All three lower tepals with tooth near base; upper lateral tepals not obviously recurved; lower tepals white near base with large red to purple blotch in centre L. violacea 24b All lower tepals without tooth-like projections: 26a Basal leaf smooth or minutely papillate along edges of ribs; perianth tube (25 )35 45 mm long; perianth pale to deep blue to violet, lower tepals white( pale yellow) at base L. dolomitica 26b Basal leaf sparsely hairy; perianth tube mm long; perianth deep magenta to purple marked with white lines on lower margins and darker magenta in lower midline L. lewisiana

35 S T R E L I T Z I A 35 (2015) 27 In the descriptions that follow, we have tried to be as concise as possible. Stem characters are included in the opening sentence beginning Plants. Cataphylls offer no taxonomic insight and are mentioned only in the generic description. The ovary provides minimal information, always ± ovoid-oblong and its size changes rapidly after fertilisation, even before flowers fade so that dimensions are of little utility and we do not include information about the ovary. We do provide descriptions of capsules, which are variable, even though plants with well-developed capsules are rarely collected. A. Sect. Chasmatocallis (Spp. 1 8) A. Sect. Chasmatocallis (R.C.Foster) Goldblatt & J.C.Manning, comb. et stat. nov. Chasmatocallis R.C.Foster: 40 (1939). Type: C. macowanii R.C.Foster (= Lapeirousia divaricata Baker). Corm bell-shaped; tunics with basal rim bluntly lobed or minutely denticulate with minute teeth directed downward. Stem mostly aerial (sometimes ± subterranean in L. exilis and L. silenoides) and often branched, stems compressed with angles often crisped. Inflorescence a spike. Flowers bilaterally symmetric, lower tepals usually with nectar guides of contrasting colour; variously coloured and perianth tube short to elongate, cylindric or funnel-shaped. Capsules often with auriculate lobes and locular ridges. Seeds smooth (scarcely wrinkled) with surface cells domed or tuberculate (L. pyramidalis). Basic chromosome number x = 9 (or 10). 1. Lapeirousia divaricata Baker in Journal of Botany (London) 14: 337 (1876). Goldblatt: 64 (1972), including L. spinosa and L. tenuis as varieties. Type: South Africa, [Northern Cape], without precise locality or date, Thom s.n. (K, holo!). Gladiolus setifolius L.f.: 96 (1782 [as 1781]). Lapeirousia setifolia (L.f.) N.E.Br.: 30 (1928), hom. illegit. non Harms (1901). Type: South Africa, without precise locality, probably in 1774, Thunberg s.n. UPS-THUNB1067 (UPS-THUNB, holo.!). Chasmatocallis macowanii R.C.Foster: 40 (1939). Type: South Africa, [Western Cape], Pakhuis Pass, Oct. 1897, MacOwan 1975 (GH, holo.!; G!, SAM!, iso.). Plants mm high, with axillary cormlets below ground, stem simple or more often branched and then mostly well above ground, terete. Corm bell-shaped, ± 10 mm diam.; tunics dark brown, often glossy, basal margin finely serrated, teeth directed downward. Leaves 3 6 (or more), lowermost usually exceeding others, linear, suberect or falcate, mm wide, strongly ribbed, generally longer than stem, up to 350 mm long, upper cauline leaves shorter, sometimes resembling bracts and not ribbed, usually subtending branches. Inflorescences 5 12-flowered spikes, usually crowded; outer bracts green or flushed reddish, often white to semi-transparent below, firm, mm long, obtuse or acute, usually folded in midline or channelled, margins hyaline, inner bracts 1 / 2 to 2 / 3 as long. Flowers zygomorphic, white or sometimes flushed with pink to lilac (aging pink), lower tepals often with pink to purple streak in midline and bearing tooth-like ridge in lower midline; perianth tube short, 7 10 mm long, narrow below, curved and widening toward apex; tepals unequal, lanceolate, acute, dorsal mm long, erect, separating from tube below lower tepals, margins undulate, remaining tepals joined for ± 4 mm, upper lateral tepals sharply reflexed shortly above base, lower 3 tepals united for additional 2 3 mm, ± 12 mm long, bent at right angles shortly above base, directed downward distally. Stamens unilateral and arcuate; filaments ± 6 mm long, exserted 4 mm from tube; anthers ± 4 mm long, white; pollen white. Style dividing opposite anther tips; branches forked for ± 2 / 3 their length, recurved. Capsules obovoid, mm long, apices of locules forming auriculate lobes decurrent on low locular ridges. Seeds ± globose, flattened at chalazal end, mm diam., light brown. Chromosome number 2n = 20 (?18 + 2B). Flowering time: mainly late August to late September, but October at higher elevations. Distribution and biology: Lapeirousia divaricata has a restricted distribution from Citrusdal in the Olifants River Valley north through the valleys of the Cederberg to the Gifberg and Bokkeveld Mtns (Figure 4). It occurs exclusively in sandy ground in soils derived from Cape sandstones, in valley bottoms, seeps or poorly drained flats where the ground remains moist for some time. The flowers of Lapeirousia divaricata remain open at night, unlike most species of the

36 28 S T R E L I T Z I A 35 (2015) consistent with several other species of sect. Chasmatocallis. FIGURE 4. Distribution of Lapeirousia divaricata, ; L. silenoides,. genus, and are visited by a range of large bees including four species of Anthophora (A. braunsii, A. diversipes, A. krugeri and A. schulzei), Amegilla grisella, and a species of Megachilidae from midmorning until sunset, and in the later afternoon and early evening by settling moths Heliothis (Noctuidae) (Goldblatt et al. 1995). Both bees and moths forage for nectar, present in small quantities, and are probably effective pollinators of L. divaricata. Sometimes present in large numbers, the moths make a striking evening sight, fluttering from plant to plant. We regard L. divaricata as having a bimodal pollination system. The flowers are self-compatible and readily selfpollinate as the style branches remain in contact with the anthers during anthesis. Diagnosis and relationships: Lapeirousia divaricata, named for its uneven and wide-angled (divaricate) branching pattern, is readily recognised by its short-tubed white to pale pink flower, a stem normally branched well above the ground, and short floral bracts that are white to translucent on the sides. The lower tepals bear a conspicuous tooth-like ridge in the lower midline, a feature shared by only a few other species of Lapeirousia. The bell-shaped corm has finely toothed margins, the teeth directed downward. The relatively small capsules of L. divaricata are ridged with apically auriculate locules, a feature Lapeirousia spinosa, treated by Goldblatt (1972) as subsp. spinosa of L. divaricata, has similar, short tubed white flowers but a very different corm, triangular in profile with the basal rim bearing prominent spreading spines. A second subspecies of L. divaricata, subsp. tenuis, also now a separate species, has corms of the same type as L. spinosa and both taxa are now believed to be only distantly related to L. divaricata. The DNA-based molecular phylogeny (Figure 1) places the two in sect. Lapeirousia. The floral similarity in the three species is evidently convergent. The sister relationship of L. divaricata with L. jacquinii retrieved in the molecular phylogeny is unexpected as the flowers of the two are very different. They do, however, share virtually identical corms and distinctive fenestrate floral bracts with white or semi-transparent patches. History: Lapeirousia divaricata was first recorded by C.P. Thunberg, whose collection, most likely made in 1774, formed the basis for Gladiolus setifolius, described by the younger Linnaeus in The species was not transferred to Lapeirousia when this genus was understood to be distinct from Gladiolus and continued to be known as G. setifolius throughout the 19th century. After examining the Iridaceae in the Thunberg Herbarium in Uppsala, Sweden, N.E. Brown transferred the species to Lapeirousia in Unfortunately, by this time a tropical African species had been named L. setifolia (Harms 1901) (now Psilosiphon setifolius), and unknown to Brown, his combination was an illegitimate homonym. A second collection of the species was described in 1876 as L. divaricata by J.G. Baker. The American botanist, R.C. Foster named a third collection of the species Chasmatocallis macowanii in 1939, the epithet honouring Peter MacOwan, who made the type collection. Foster seems to have been unaware that his new genus Chasmatocallis differed in no significant way from either G. setifolius or L. divaricata. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Calvinia): sandy ground near Grasberg, west of Nieuwoudtville, ( AC), 18 Sept. 1980, Goldblatt 5839 (MO, PRE); Farm Driefontein, 18 km south of Nieuwoudtville, ( AC), 9 Oct. 1991, Manning 1043 (MO, NBG); Lokenburg, ± 700 m, ( CA), 26 Sept. 1953, Acocks (PRE).

37 S T R E L I T Z I A 35 (2015) 29 WESTERN CAPE (Vanrhynsdorp): Gifberg plateau, ( DC), 21 Sept. 1948, Acocks (PRE), Sept. 1911, Phillips 7617 (NBG, PRE), 7508 (NBG); Nardouw, ( DC), Sept. 1941, Stokoe s.n. (NBG) (Calvinia): Doornbos, ( CC), 4 Sept. 1948, Compton (NBG) (Clanwilliam): deep sand at old campsite, Clanwilliam, ( BB), 24 Sept. 1974, Goldblatt 2754 (MO, PRE) (Wuppertal): Biedouw Valley, ( AA), 14 Sept. 1976, Thompson 2922 (MO, PRE, STE), 7 Sept. 1953, Compton (NBG), 21 Sept. 1952, Lewis s.n. (PRE, SAM); Brandewyn River, ( AA), 20 Sept. 1933, Barker 254 (NBG), 2 Oct. 1940, Bond 615 (NBG), 8 Sept. 1974, Goldblatt 2538 (MO, PRE); Pakhuis Mtns [as Pakenberg], ( AA), Oct. 1897, MacOwan 3300 (G, PRE); Diamond Drift, Bidouw River, ( AA), Aug. 1939, Leipoldt s.n. (NBG 61341, PRE); Hoek, 27 Aug. 1896, ( AB), Schlechter 8712 (E, G, MO, PH, PRE); Koudeberg near Wuppertal, ± 900 m, ( AC), 4 Oct H. Bolus 9100 (NBG, PRE). 2. Lapeirousia jacquinii N.E.Br. in Journal of the Linnean Society, Botany 48: 20 (1928); Goldblatt: 56 (1972). Type: South Africa, Cape, without precise locality, illustration in Jacquin: t. 269 (1794) (specimens believed to have been destroyed). Plants ( 200) mm high, stem often ± prostrate below for a short distance, erect above, often branched from close to ground, compressed and triangular in cross section, one side narrower than other two, angles strongly winged, wings undulate and usually slightly denticulate. Corm bell-shaped, 7 9 mm diam.; tunics light to dark brown, basal rim minutely serrate, teeth directed downward. Leaves 3 5, ± crowded at ground level, lowermost longest, ascending or arcuate, often quite short, lanceolate to falcate, mm, strongly ribbed, remaining leaves smaller, weakly ribbed, progressively bract-like above. Inflorescences 2 5( 10)-flowered spikes; outer bracts green-glaucous, often ± translucent on sides, (10 )15 20 mm long, margins narrowly hyaline, prominently 2-keeled or lowermost sometimes 1-keeled, keels undulate to crisped, minutely denticulate, inner bracts ± 2 / 3 as long, membranous and translucent with two prominent green keels, forked apically for 2 3 mm. Flowers zygomorphic, violet (rarely white), lower 3 tepals darker violet or reddish in lower midline with margins white to pale yellow in proximal half, scentless; perianth tube cylindric, mm long, widening and weakly curved near apex; tepals unequal, dorsal largest, mm, oblanceolate, initially erect but usually reflexed when fully open, then held in same plane as lower; upper lateral tepals spreading at right angles to tube, weakly spooned, apices curved upward, lower 3 tepals directed forward with upcurved tips, 7 8 ± 2 mm. Stamens unilateral, ± erect; filaments ± 7 mm long, exserted 4 5 mm; anthers parallel, ± 3 mm long, violet; pollen usually blue-violet. Style dividing near base of anthers, style branches deeply forked and recurved. Capsules oblong, 7 9 mm long, apices of locules forming auriculate lobes decurrent on weakly developed locular ridges; seeds globose, mm diam., dark brown. Chromosome number 2n = 18. Flowering time: (late July) August to mid-september. Distribution and biology: one of the most common species of the genus, Lapeirousia jacquinii extends from Cape Town along the West Coast as far north as Garies in southern Namaqualand, extending inland through the Breede River valley east of Worcester as far as Overhex and Aan de Doorns, and into the western valleys of the Cederberg and Bokkeveld Mtns (Figure 5). It is especially common in the Olifants River valley in sandy, often disturbed, places. Plants flower fairly early in the season and are some- FIGURE 5. Distribution of Lapeirousia jacquinii, ; L. verecunda,.

38 30 S T R E L I T Z I A 35 (2015) times so common that they form sheets carpeting the ground under sparse low shrubs, notably in the Bokkeveld Mtns. Records indicate that L. jacquinii once occurred on the Cape Peninsula near Salt River and on the slopes of Lion s Head. Urbanisation around Salt River has long since eliminated native plants but L. jacquinii probably persists on Lion s Head (Lewis 1950). Lapeirousia jacquinii is self-compatible and facultatively autogamous, and reproduces very readily from seed, thus rapidly colonising new habitats. It is also often cross-pollinated in the parts of its range where long-proboscid flies are active. In the Bokkeveld Mtns plants are regularly visited by two species of Prosoeca, P. peringueyi and P. sp. (Nemestrinidae), and elsewhere only by P. peringueyi. Both insects are effective pollinators while foraging for nectar (Goldblatt et al. 1995). East of Pakhuis Pass and at the southern end of the Bokkeveld Mtns and in Biedouw Valley L. jacquinii frequently co-occurs with the florally similar L. violacea, also pollinated by Prosoeca peringueyi. Although flies visit flowers of both species, no hybrids between them are known. Diagnosis and relationships: Lapeirousia jacquinii is most easily recognised by the prominently 2-keeled outer floral bracts (occasionally 1-keeled, especially the lowermost) and even in fruit the species can be recognised by this feature. The stems are conspicuously 3-angled and the angles winged and undulate and sometimes minutely toothed. The long-tubed flowers are deep violet (rarely white) with darker violet or red makings outlined in white on the lower tepals, a colour combination very similar to that in L. violacea, with which it is sometimes sympatric. This last species has quite different corms, triangular in profile and bearing long spines radiating from the base, unlike the bell-shaped corm of L. jacquinii, the basal rim of which bears minute, downward-directed teeth. The outer bracts of the two species also differ, those in L. violacea being folded or keeled and the keels minutely crisped. The remarkable similarity in shape, tube length and colouration of the flowers in the two species is evidently due to convergence for the same long-proboscid fly pollinators. The immediate relationships of Lapeirousia jacquinii are uncertain, but the basic chromosome number, x = 9, is consistent with sect. Chasmatocallis and most species have similar, if not identical, bell-shaped corms either with crenate or minutely serrate margins. The plastid DNA sequence-based phylogeny consistently shows L. jacquinii to be most closely related to L. divaricata, which is very different in its shorttubed, white to palest pink flowers, and short, 1-keeled floral bracts. The corms of the two are, however, identical and they also share unique, fenestrate outer bracts with pale, almost translucent sides. History: although common, Lapeirousia jacquinii is poorly represented in older herbarium collections. It was first recorded in the Codex Witsenii, a collection of illustrations of plants encountered on Simon van der Stel s expedition to the Copper Mountains of Namaqualand. The painting, a very poor likeness, was copied in Jan Commelin s manuscript account of the Cape Flora, written between 1687 and 1692 (Wijnands et al. 1996), and called therein branching Sisyrinchium aethyopicum with helmeted purple flower. The name appears never to have been adopted in the botanical literature: the first post-linnaean record is evidently the plant figured by N.J. Jacquin in 1794, presumably collected north of Cape Town a few years earlier. Jacquin confused the species with Gladiolus (Lapeirousia) anceps of Linnaeus fil. and in doing so initiated the confusion that long surrounded these two very distinct, but long-tubed species. Ker Gawler (1804) perpetuated the error, and L. jacquinii continued to be considered conspecific with L. anceps (e.g. Sprengel 1824; Poiret 1826; Dietrich 1833) and was so treated by J.G. Baker in 1896 in Flora capensis. The situation was only clarified in 1928 by N.E. Brown, who then formally described L. jacquinii, naming it in honour of Jacquin. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Hondeklipbaai): ± 71 km west of Garies to Hondeklip Bay, dune ridges, ( AD), 18 Sept. (fr.), Goldblatt & Manning 9998 (MO); hills above Garies on road to Hondeklip Bay, ( DB), 20 Aug. 2001, Goldblatt & Porter (MO), 27 Aug. 1991, Goldblatt & Manning 9135 (MO, NBG) (Calvinia): Bokkeveld Mtns, Farm Avontuur, ( AA), 28 Aug. 2008, Goldblatt & Manning 13052A (MO); top of Vanrhyns Pass, ( AC), 6 Sept. 1963, Nordenstam 2990 (MO, S); Farm Driefontein, southwest of Calvinia, ( CA), 8 Sept. 1992, Goldblatt & Manning 9412 (MO); top of Botterkloof Pass

39 S T R E L I T Z I A 35 (2015) 31 (growing with L. violacea), ( DD), 3 Sept. 1994, Goldblatt & Manning 9966 (MO). WESTERN CAPE (Vanrhynsdorp): Biedouw River, Vanrhynsdorp, ( DA), 21 Aug. 1954, Barker 8599 (NBG), Lewis 1407 (SAM); hills north of Klawer, ( DC), 5 Aug. 1974, Goldblatt 2266 (MO); Windhoek, ( DC), 30 July 1896, Schlechter 8340 (MO) (Vredenburg): Stompneus, ( DB), 23 Aug. 1953, Hall 735 (NBG); Olifantskop, Langebaan, ( DD), 15 Aug. 1976, Goldblatt 2328 (MO) (Clanwilliam): east of Graafwater, on pass, ( BA), Booysen 48 (NBG); north of Piketberg, flats at foot of Zebrakop, ( DB), 1 Sept. 1981, Goldblatt 6128 (MO); Piekenierskloof, Schlechter 4916 (BOL) (Wuppertal): Pakhuis Pass, ( AA), Barker 1309 (NBG), Lewis 1854 (SAM); Cederberg, 2 km from Algeria to Clanwilliam, ( AC), 13 Aug. 1986, Van Wyk 2564 (MO, PRE); Citrusdal, ( CA), 24 Sept. 1928, Grant s.n. (MO) (Cape Town): near Hopefield, ( AB), H. Bolus (BOL); Darling Flower Reserve, ( AD), 11 Aug. 1960, Barker 9182 (NBG); west slope of Lions Head, ( CD), MacOwan (BM, BOL, K, SAM); 13 miles [18.5 km] from Cape Town to Kalbas kraal, ( DC), Hutchinson 174 (BOL, K, PRE) (Worcester): below Tulbagh Falls, deep sand, ( AC), 29 Aug. 1978, Goldblatt 4759 (MO); rocky flats between Worcester and Overhex, ( CA), 1 Sept. 1992, Goldblatt & Manning 9371 (MO); Aan de Doorns, Farm Reiersrus, ( CB), Walters 2163 (NBG). 3. Lapeirousia silenoides (Jacq.) Ker Gawl. in König & Sims Annals of Botany 1: 238 (1804); Goldblatt: 36 (1972). Gladiolus silenoides Jacq.: 168 (1792). Ovieda silenoides (Jacq.) Spreng.: 147 (1824); Peyrousia silenoides (Jacq.) Poir.: 365 (1826). Meristostigma silenoides (Jacq.) A.Dietr.: 596 (1833). Type: South Africa without precise locality or collector, illustration in Jacq.: t. 270 (1794) (herbarium specimens believed destroyed). Lapeirousia speciosa Schltr.: 105 (1900). Type: South Africa, [Western Cape], Kareebergen, 17 July 1896, Schlechter 8187 (B, holo.!; BM!, BOL!, COI!, E!, G!, GRA!, K!, PH!, PRE!, iso.). Plants ( 200) mm, taller in sheltered or shaded sites, ± acaulescent in exposed sites or in seasons of low rainfall, stem aerial often branched close to base, compressed, 2- angled, often 2-winged below first flower, wings often minutely serrate. Corm bell-shaped, 8 10 mm diam.; tunics light brown, basal rim ridged (minutely serrate in plants from Spektakel Mtns). Leaves few, lowermost longer than others, lanceolate to linear, ribbed or corrugate, ± as long as or longer than stem, mm wide; remaining leaves shorter and broader, progressively bract-like above. Inflorescence usually a crowded to lax spike, or a head-like cluster at ground level, up to 12-flowered, initially 2-ranked, spiral with age; outer bracts linear to lanceolate, sometimes very broad at base, occasionally ± succulent, outer mm long, unifacial for distal 2 4 mm, acute and curving upward, folded or lightly keeled below, keels usually minutely serrate, margins membranous; inner bracts ± 1 / 2 as long, with broad transparent margins. Flowers zygomorphic, bright cerise to dark magenta (rarely white), lower 3 tepals each with creamy white markings near base, limbs with large central blotch of dark red or purple, scentless; perianth tube cylindric, slender, ± erect, mm long, curved outward near apex; tepals subequal, ± broadly obovate, obtuse, limbs concave, ± 5 mm, dorsal erect or sometimes slightly inclined toward stamens, upper lateral tepals reflexed, lower 3 united basally for ± 1 mm, inclined at ± 45 to horizontal. Stamens unilateral, ± erect; filaments 7 8 mm long, exserted ± 4 mm from tube; anthers 3 4 mm long, blue violet; pollen white or pale blue to mauve. Style erect, dividing close to or just beyond anther tips, ultimately exceeding them; branches mm long, forked for ± 1 / 2 their length, recurved. Capsules obovoid to oblong, trigonous, mm, usually with well-developed wing-like locular ridges in distal half. Seeds globose, light brown, variable in size even on same plant, mm diam. Chromosome number 2n = 18. Flowering time: (July) August to mid-september. Distribution and biology: widespread in Namaqualand, Lapeirousia silenoides extends from near Nuwerus in the south to the Anenous Mtns west of Steinkopf in the north (Figure 4). It is restricted to granitic soils, and may grow either in deep, coarse granitic sand where plants tend to be robust in years of adequate rainfall or in granite outcrops where the corms are usually wedged in rock cracks, in little or no soil. On rocks, plants tend to be fairly dwarfed but in years of ample rainfall the main stem branches repeatedly at the base, producing a cushionlike form. In years of poor rainfall, plants can be virtually acaulescent with the flowers produced close to ground level. Similarly, at higher elevations in the Kamiesberg, the aerial stem is often

40 32 S T R E L I T Z I A 35 (2015) hardly or not at all produced above the ground, and plants assume a low, tufted growth habit. Self-incompatible, Lapeirousia silenoides often has relatively low seed set in the wild, and in cultivation does not produce capsules unless actively cross pollinated even though the recurved style branches do come into contact with pollen. Pollination studies show that the species is regularly visited by the long-proboscid nemestrinid fly, Prosoeca peringueyi, which appears to be its only legitimate pollinator (Goldblatt et al. 1995). Plants produce up to 3.6 ml of nectar of moderate sugar concentration (mean 26.5% sucrose equivalents). Plants reproduce very successfully by seed and L. silenoides can be seen flowering along roadsides in masses, the seed having fallen from parent plants growing on banks and slopes above the road. Diagnosis and relationships: the bright magenta to cerise flower with darker colour in the centres of the lower tepals and cream streaks or blotches on the lower tepal bases instantly identify Lapeirousia silenoides in Namaqualand. To the south, in the Vanrhynsdorp and Clanwilliam districts, L. pyramidalis may be similarly coloured, though more often purple to violet, but the two species can be separated by their different bracts; narrow to fairly broad and acute with an upturned apex in L. silenoides, broad and often retuse with the apex curving downward in L. pyramidalis. The two are undoubtedly related as is evidenced not only by their similarly marked and sometimes almost identically coloured flowers but by the similar, red-brown corms and capsules with wing-like locular ridges, a character shared by relatively few other species of Lapeirousia. When dry, L. silenoides can easily be confused with red-flowered L. pyramidalis subsp. regalis unless the bracts are carefully compared. Bract shape is fairly variable in Lapeirousia silenoides. In the centre of its range, in the Kamieskroon area, the bracts are narrow and often quite long. In this area too, the plants are often tallest and are occasionally slender. This form resembles the type illustration of Jacquin (1794). Both south and north of this area the plants are smaller, and generally less robust, with the bracts shorter and broader. Plants with particularly broad-based bracts occur in the south of its range, notably in the vicinity of Bitterfontein. This variant, named L. speciosa by Schlechter (1900), has flowers identical with those of plants from further north and although the difference in the shape of the bracts in Schlechter s type material is striking, the presence of populations intermediate for bract shape and the consequent difficulty in placing these makes an infrageneric classification seem unnecessary. History: as far as we have been able to determine, Lapeirousia silenoides was first collected by the French explorer, François le Vaillant, during his expedition to Namaqualand in 1783 (Gunn & Codd 1981). Although Le Vaillant collected few plant specimens, one attributed to him in the Lamarck Herbarium in Paris attests to his discovery of the species. L. silenoides was, however, described (as Gladiolus) by N.J. Jacquin whose painting in Icones plantarum rariorum (1794) of a plant cultivated and flowered in Vienna is regarded as the type of the species. The source of Jacquin s plant must have been Franz Boos and Georg Scholl who collected bulbs, corms and seeds for Jacquin in the period 1786 to 1788, just a few years after Le Vaillant visited Namaqualand. The species was not recorded by the Simon van der Stel expedition to the copper mountains of Namaqualand, probably because they only reached Namaqualand proper in late September (Wijnands et al. 1996) when L. silenoides would no longer have been in bloom. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): between Steinkopf and Klipfontein, ( BA), 17 Sept. 1929, Herre s.n. (BOL30710, PRE); Klipfontein, ( BA), Aug. 1883, H. Bolus 696 (BM, BOL, G, GRA, SAM); Anenous Pass, ( BA), 23 Aug. 1983, Van Wyk 6168 (PRE); top of Anenous Pass, ( BA), 27 Aug. 2008, Goldblatt & Manning (MO, NBG); ± 11 km northeast of Nigramoep, ( BC), Acocks (PRE); 12 km north of Komaggas, ( CD), 20 July 1970, Wisura 1692 (NBG); top of Spektakel Pass, west of Springbok, ( DA), 17 Aug. 1974, Goldblatt 2365 (MO); hill west of Springbok, ( DB), 24 Aug. 1992, Goldblatt & Manning 9322 (MO, NBG); Droëdap, ( DD), 27 Aug. 1941, Barker 1306 (NBG) (Hondeklipbaai): Grootvlei, ( BB), 18 July 1961, Middlemost 1636 (NBG), Lewis 1637 (SAM); hills between Garies and Kamieskroon near microwave tower, ( BD), 21 Aug. 1992, Goldblatt & Manning 9278 (MO), 16 Sept (fr.), Goldblatt & Manning 9467 (MO); hills at Brakdam, ( BD), 4 Sept. 1945, Compton (BOL); 24 July 1941, Esterhuysen 5442 (BOL), Leighton 1140 (BOL), 8 Sept. 1897, Schlechter L. montana Schltr. (BM, COI, E, GRA, K, MO, PH, SAM);

41 S T R E L I T Z I A 35 (2015) 33 Darter s Grave, ( BD), 7 Sept. 1950, Barker 6611 (NBG), 18 July 1961, Middlemost 2156 (NBG); Karkams, Kamiesberg, ( BD), 11 Sept. 1911, Pearson 6661 (BOL, SAM); 6 km north of Garies, ( DB), Schlechter 1400 (BM, BOL); 13.7 km from Kotzesrus to Garies, along seasonal stream, ( DD), 3 Sept. 2001, Goldblatt & Porter (MO) (Kamiesberg): top of Kamiesberg Pass at Farm Bo-Tuin, ( AA), 9 Sept. 1980, Goldblatt 5762A (MO); Twee Rivieren, Kamiesberg, ( AC), 13 Sept. 1911, Pearson 6478 (BOL, SAM); Kamiesberg, Damsland Kloof, northern approach to Rooiberg, ( AC), 19 Sept. 1991, Goldblatt 9259 (MO); ± 16 km south of Garies, ( CA), Barker 7330 (NBG); 10 km south of Garies, ( CA), 29 Aug. 1991, Goldblatt & Manning 9202 (MO); Kamiesberg, road between Karas and Farm Naartjiesdam, ( CA), 1 Sept. 2002, Goldblatt (MO). WESTERN CAPE (Vanrhynsdorp): 6.5 km west-northwest of Bitterfontein, ( AA), 26 Aug. 2008, Goldblatt & Manning (MO, NBG); Mierenkasteel, ( AA), Drège 2642 (MO, SAM); Bitterfontein ( AB), 23 July 1941, Bond 1091 (NBG); 23 July 1941, Esterhuysen 5407 (BOL); 10 km northeast of Bitterfontein toward Kliprand, ( AB), 22 Aug. 1974, Nordenstam & Lundgren 1456 (MO, NBG, S); 8 km north of Nuwerus, ( AB), 18 July 1948, Compton (NBG). 4. Lapeirousia verecunda Goldblatt in Contributions from the Bolus Herbarium 4: 38 (1972). Type: South Africa, [Northern Cape], Namaqualand, upper east slopes of Spektakel Pass, 24 Aug. 1970, Goldblatt 573 (BOL, holo.!; K!, MO!, NBG!, PRE!, iso.). Plants mm high, stem simple or with 1 3 branches from base, slightly compressed and 2-angled, one or both angles winged, wings decurrent on leaves or bracts. Corm bell-shaped, mm diam.; tunics redbrown, slightly rugose, basal rim slightly lobed. Leaves 5 7 (more when stem branched), lowermost longest, linear, ascending to falcate, closely ribbed, mm, seldom exceeding stem; upper leaves < 1 / 2 as long as lowermost, progressively shorter and bract-like distally. Inflorescences 9 12-flowered, 2-ranked, spikes; outer bracts green, firm, narrowly lanceolate, acute, often recurved at tips, mm long, margins membranous, channelled or folded in midline; inner bracts ± 1 / 2 as long, notched at tips. Flowers zygomorphic, white, tube and reverse of tepals copper-pink, lowermost or all 3 lower tepals with red, heart-shaped mark in lower midline; perianth tube cylindric, slender, 40 50( 55) mm long, slightly curved forward distally; tepals subequal, ovate-elliptic, slightly narrowed at base, sub-acute or obtuse, mm, dorsal erect, lower 3 tepals ± horizontal. Stamens unilateral, erect; filaments ± 6 mm long, exserted ± 3 mm; anthers parallel, contiguous, ± 3 mm long, purple; pollen purple. Style dividing opposite anther tips, branches ± 2 mm long, divided for 1 / 2 their length, recurved. Capsules obovoid-trigonous, ± mm, with prominent wing-like locular ridges in distal half. Seeds ± globose, smooth, dark brown, mm diam. Chromosome number 2n = 18. Flowering time: late August to the end of September, rarely early October. Distribution and biology: a narrow endemic of the Spektakel Mtns west of Springbok, Lapeirousia verecunda was thought, when first described in 1972, to consist of a single extended population on the eastern slopes of Spektakel Pass, at elevations of m. Now also recorded near Nigramoep, some 20 km to the north in the same mountain complex (Figure 5), the species is largely restricted to stony shale slopes (Nama Shale Formation), which has a scattered distribution in Namaqualand. Toward the eastern summit of Spektakel Pass where the shale merges with the surrounding granite, L. verecunda grows together with cerise-flowered L. silenoides. The latter, which is widespread in Namaqualand, always grows in granite derived soils or in crevices in granite outcrops. We have found hybrids between the two species on the farm Naries on Spektakel Pass where the ranges of the two species converge (Goldblatt & Manning 9969). The flower colour, white with red markings, and the long perianth tube of L. verecunda are stereotypical adaptations for the Moegistorhynchus longirostris long-proboscid fly pollination system, but to date no pollinators have been seen visiting L. verecunda. Hybrids are likely the result of the pollinator of L. silenoides, Prosoeca peringueyi visiting both species. Diagnosis and relationships: this attractive plant is easily recognised by its odourless, longtubed white flowers with red, round to heartshaped marks on one or more of the three lower tepals, and the red colour on the reverse of the tepals and tube. Aside from colour, the floral tube, usually mm long, plane tepals, and fairly long and comparatively narrow bracts with deflexed apices are distinctive. L. verecunda belongs to the group of species in sect. Chasmatocallis with

42 34 S T R E L I T Z I A 35 (2015) bell-shaped corms with tunics that are ridged or regularly lobed on the basal rim rather than being serrated. In its long perianth tube and fairly broad, ovate-elliptic tepals the flowers of L. verecunda most closely resembles those of L. silenoides, but because of the colour differences of the flowers the two are unlikely to be confused. The cerise flowers of L. silenoides with spoon-shaped lower tepals are rather different from the white perianth with flat lower tepals of L. verecunda. The species is named for its rarity (Latin verecunda = shy) and for the pale flowers with a slight blush from the pink colour on the reverse of the tepals. History: when described in 1972, Lapeirousia verecunda was thought to have been discovered only in August 1970, a year of ample rainfall in Namaqualand that encouraged collecting in the region. Later, we learned that there were at least three earlier records of the species. The first of these is was found in the personal herbarium of the American botanist, A.L. Grant, whose South African plant collection was bequeathed to the University of California, Los Angeles, and was later incorporated into the Missouri Botanical Garden Herbarium, St. Louis, in The specimens were collected by Harry Bolus, founder of the Bolus Herbarium at the University of Cape Town. Bolus collected in the Spektakel Mtns in September 1883, on his single expedition to Namaqualand. The collection, undated but accompanied by labels, some bearing the numbers 38 and 42, was provisionally identified as L. anceps (a name then misapplied to L. fabricii). Bolus s field register (University of Cape Town Archives) records L. anceps, number 36, collected on the Spektakelberg (not Kommagas Kloof as one of the labels indicates). It seems clear that Bolus did indeed make the first gathering of L. verecunda, but through a quirk of circumstance, the entire collection was acquired by the Missouri Botanical Garden, evidently because it was thought to be yet another collection of a common Namaqualand species and considered not worth retaining in the Bolus Herbarium collection. Later collections of L. verecunda were made by B.E. Martin in 1950 and J.P.H. Acocks in Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): west-facing slope south of Nigramoep, 800 m, ( BC), 4 Sept. 1983, Oliver 8033 (NBG, PRE); ± 23 km southsouthwest of Springbok, 25 Aug. 1957, ( DA), Acocks (PRE); Eselsfontein (Spektakelberg), ( DA), 8 Sept. 1950, Martin 515 (NBG); upper eastern slopes of Spektakel Pass, ( DA), 7 Sept. 1980, Goldblatt 5710 (MO, NBG), 23 Sept. 1977, Perry 383 (NBG), Sept. 1970, Thompson 1021 (NBG), 28 Aug. 1991, Goldblatt & Manning 9144 (MO); Naries, Spektakel Mtns., ( DA), undated [probably Sept. 1883], H. Bolus 38 & 42 (BOL, MO); Spektakel Mtns, at top of Pass, ( DA), 26 Sept 1974, Goldblatt 2789 (MO, NBG), 28 Aug. 9144, Goldblatt & Manning 9144 (MO); Kommagas Kloof, ( DA), but perhaps Spektakelberg intended, without date [1883?], H. Bolus s.n. (MO). 5. Lapeirousia anceps (L.f.) Ker Gawl. in König & Sims Annals of Botany 1: 238 (1804); Goldblatt: 33 (1972). Gladiolus anceps L.f.: 94 (1782). Ovieda anceps (L.f.) Spreng.: 147 (1824). Peyrousia anceps (L.f.) Poir.: 363 (1826). Meristostigma anceps (L.f.) A.Dietr.: 596 (1833). Type: South Africa, without precise locality, Sparrman 192 (59/19 LINN, holo!.). Lapeirousia pentheri Baker: 27 (1906). Type: South Africa, [Western Cape], Breede River, Penther 728 (BOL!, BM!, K!, syn.). [anceps two-edged referring to the compressed, usually two-angled stem.] Plants mm high; stem usually repeatedly divaricately branched, compressed and elliptic in cross section, 2- or 3-angled, angles slightly winged, wings weakly denticulate. Corm bell-shaped, red-brown, 8 10 mm diam.; tunics lightly ridged, rim bluntly lobed. Leaves 5 7, lowermost usually somewhat longer and wider than others, (40 ) mm, ascending to falcate, lanceolate, ribbed, remaining leaves shorter, mostly crowded near base, becoming increasing bract-like above. Inflorescences (2 )4 8-flowered spikes, at first 2-ranked, ultimately spiral; outer bracts grey-green, (5 )6 10 mm long, apices and margins often tinged red, channelled, folded along upper midline, acute in profile; inner 3 5 mm long, transparent with two green keels. Flowers zygomorphic, white to beige or faintly flushed pink, reverse of tepals and tube pink, lower 3 tepals each with red mark in lower midline, and usually with small, claw-like cusp near base; perianth tube narrow and cylindric, weakly arched distally, (20 ) mm long (3 6 times as long as tepals); tepals narrowly lanceolate to ± linear, acute, dorsal largest, mm, erect or recurved when fully open, upper lateral ± 3 mm, widely

43 S T R E L I T Z I A 35 (2015) 35 separated from dorsal, lower tepals united basally for 2 3 mm, free parts mm long, extended horizontally. Stamens unilateral, ± erect; filaments ± 6 mm long, exserted ± 4 mm from tube; anthers parallel, ± 2.5 mm long, pollen grey-blue or white. Style arching adaxial to anthers, dividing between base and middle of anthers; branches 2 3 mm long, divided for 1 / 2 their length and recurved. Capsules ± barrel-shaped, 9 10 ± 7 mm. Seeds red-brown, globose, smooth, somewhat flattened at chalazal end, ± 1.5 mm diam. Chromosome number 2n = 18. Flowering time: (late September )October, to late November at higher elevations and in the south. Figure 6. Distribution and biology: with one of the widest distribution ranges of the southern African species of Lapeirousia, L. anceps extends from the sandy coastal plain of central Namaqualand near Hondeklip Bay in the north through the mountains and coastal plain of Western Cape to near Mossel Bay in the east (Figure 7). Over this wide range, plants almost always occur in deep sandy soils, sometimes growing among tufts of Restionaceae. Like many geophytes of sandstone soils, L. anceps shows a pattern of profuse flowering following fires the previous season. Along the Cape west coast this is not, however, the rule but even here flowering is promoted by disturbance of the native, largely shrubby vegetation. Plants are typically scattered, but sometimes grow in masses, when the slender flowers then make a notable display. Pollination studies have shown that Lapeirousia anceps is pollinated by long-proboscid flies, either the nemestrinid Moegistorhynchus longirostris or the horseflies Philoliche gulosa and P. rostrata (Goldblatt et al. 1995; Manning & Goldblatt 1997; Pauw et al. 2008). The remarkable range of perianth tube length in different populations is closely correlated with variation in proboscis length in the pollinators, shorter-tubed populations being pollinated by flies with correspondingly shorter probosces (Pauw et al. 2008). Over much of its range L. anceps has a tube mm long, corresponding to the proboscis of P. gulosa, mm. In the southern Cape the tube tends to be shorter, sometimes only 20 mm long (e.g. Woodvine 91 from Hermanus and Bohnen 7863 from Riversdale). In the west, along the coastal plain and at sites in the Olifants River Mtns, some populations have tubes as long as mm. Both this very long-tubed form and the southern populations are pollinated largely by Moegistorhynchus, some morphs of which have the longest probosces of any fly in southern Africa. Diagnosis and relationships: Lapeirousia anceps is readily recognised by the combination of short floral bracts and white to ivory flowers, pink on the outside of the tepals and tube, and usually relatively long perianth tube, mostly mm but occasionally only mm, and narrow tepals 3 4 mm wide (sometimes the dorsal up to 5 mm). The lower tepals usually have a claw-like projection just above the base. The bell-shaped corms have either bluntly lobed or minutely serrated basal rims. The species has no close allies within the section and the short floral bracts of L. anceps, 5 10 mm long, are particularly surprising in view of its long perianth tube. History: the background to the protologue of Lapeirousia anceps was outlined in some detail by Goldblatt (1972) and we briefly summarise its history here. The draft description of the species was penned by Linnaeus senior, who called it Gladiolus decurrens, based on a collection of Anders Sparrman. The younger Linnaeus deleted that name in the amanuensis draft of Linnaeus s account and replaced the species epithet with the name anceps, which he copied from a draft description by C.P. Thunberg of a second species, now L. fabricii, which at the time was thought to be the same species. Thunberg (1800) and later authors, among them John Ker Gawler, included L. anceps, L. fabricii as well as a third species (now L. jacquinii) under one name, L. fabricii, described by D. Delaroche in Baker (1896) also confused the three species and in Flora capensis he treated them under the name L. anceps, distinguishing what we now know as L. fabricii as L. anceps var. aculeata. N.E. Brown (1928) was the first to establish that L. anceps, L. fabricii and L. jacquinii are in fact separate species. We now believe that L. anceps with its bell-shaped corm and L. fabricii with an obconic corm with prominent teeth are not closely related and that their floral similarities are due to convergence for long-proboscid fly pollination. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Hondeklipbaai): sandy flats west of Wallekraal, ( AD), Sept. (in bud), Goldblatt & Manning s.n. (sight record) (Calvinia): Lokenburg, sandstone hills ( CA), Story 4330 (GRA).

44 36 S T R E L I T Z I A 35 (2015) FIGURE 6. Lapeirousia anceps, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

45 S T R E L I T Z I A 35 (2015) 37 SAM) (Bredasdorp): De Hoop, Melkkamer, ( AD), 17 Oct. 1984, Fellingham 830 (NBG, PRE); near Struisbaai, ( CA), Leipoldt 2595 (BOL) (Riversdale): Farm Klein Jongensfontein, Riversdale, ( AD), 1 Oct. 1980, Bohnen 7863 (PRE); sandy dunes, Riversdale, ( AB), Ferguson s.n. (BOL); Albertinia commonage, ( BA), Muir 1139 (BOL) (Mossel Bay): between the Great and Little Brak rivers, deep sand on north-facing slope, ( AB), 28 Oct. 1988, Vlok 2035 (MO, NBG). 6. Lapeirousia exilis Goldblatt in Contributions from the Bolus Herbarium 4: 44 (1972). Type: South Africa, [Northern Cape], Namaqualand, Ratelpoort, ± 25 km north of Springbok, 6 July 1970, Tölken 3238 (BOL, holo.!; K!, MO!, NBG!, iso.). FIGURE 7. Distribution of Lapeirousia anceps, ; L. exilis,. WESTERN CAPE (Vanrhynsdorp): 3 km north of Vredendal, ( DA), 30 Sept. 1972, Bremer 234 (MO, S); near Klawer turnoff on road to Clanwilliam, ( DC), 4 Sept. 1980, Goldblatt 5664 (MO) (Vredenburg): Witklip Farm, 1 mile [1.5 km] south of Vredenburg, ( DD), 16 Oct. 1975, Thompson 2657 (PRE) (Clanwilliam): Nortier Experimental Farm, Lambert s Bay, ( AB), 6 Nov. 1974, Boucher 2579 (PRE); 11 km west of Clanwilliam toward Graafwater, ( BB), 13 Oct. 1976, Hugo 660 (PRE); 10 km from Clanwilliam on road to Graafwater, Farm Ysterfontein, ( BB), 10 Oct. 1995, Goldblatt & Manning (MO, NBG);Verloren Vlei, Piketberg District, ( DA), 29 Sept. 1943, Barker 2617 (BOL, MO, NBG) (Wuppertal): Vleikraal, southeast of Klawer ( AB), Sept. 1979, Walters 50 (PRE); Cederberg, 3 km west of Algeria on Clanwilliam road, burnt sandy slope, 16 Nov. 1979, Goldblatt 5147 (MO); 1 mile [1.5 km] south of The Baths, Farm Latjeskloof ( CC), 24 Oct. 1977, Emdon 103 (MO, PRE); Keerom, Olifants River Valley, ( CC), Esterhuysen (BOL) (Cape Town): Bloubergstrand turnoff from Malmesbury road, ( DC), 18 Nov. 1981, Snijman 578 (NBG, PRE); Jonkershoek, Farm Waterval, ( DD), 5 Nov. 1975, Oliver 6200 (PRE) (Worcester): Saron, 800 ft [244 m], ( AA), Oct. 1896, Schlechter (GRA, K, MO); near Tulbagh Falls, ( AC), MacOwan 2157 (E); Nuwe Kloof near Tulbagh, Drège 8507a (BM, K, MO); Botha Station, ( CB), Compton (NBG); Wemmershoek Mtns, ( CC), Esterhuysen s.n. (BOL30327) (Simonstown): Kommetjie, stony hillside near Ocean View Village, ( AB), 7 Nov. 1982, Goldblatt 6720 (MO); Kommetjie, ( AB), Galpin 4692 (GRA, K); Buffels Bay, Cape Point Reserve, ( AD), Salter 1843 (BM, BOL) (Caledon): Hermanus, Fernkloof Nature Reserve, ( AD), 11 Nov. 1981, Woodvine 91 (MO); Riviersonderend, ( BA), Schlechter 5635 (BM, GRA, K, Plants small, usually mm high, rarely taller, usually unbranched, stem subterranean or produced shortly above ground. Corm ± 8 mm diam., light brown, bell-shaped, ridged at base, margin slightly lobed (not serrated). Leaves usually 2, basal longest, falcate or ± trailing, mm, ribbed, second leaf much shorter, often only slightly longer than floral bracts, 2 3 mm wide, channelled in lower half. Inflorescence a congested spike, sometimes ± a basal rosette, 6 13-flowered, 2-ranked in bud, becoming spiral as flowers open; outer bract mm long, lowermost longest, with unifacial apices, upper leaves channelled, apices rounded without unifacial tips, not keeled, translucent over veins, margins pale, membranous; inner bract ± 1 / 3 as long as outer, transparent with two green keels. Flowers zygomorphic, white to pale blue with proximal 1 2 mm fading to ± white, lower 3 tepals with a dark blue median streak in lower midline (fading in dried material), strongly and sweetly scented; perianth tube slender, erect, curved and widening at apex, mm long; tepals subequal, ± 8 mm long, spreading, dorsal suberect at anthesis, later spreading, lower 3 tepals slightly smaller broadly to narrowly ovate, obtuse, narrowed at base and ± clawed. Stamens unilateral, ± erect; filaments ± 4 mm long, inserted close to top of tube, ± fully exserted; anthers ± 3 mm long, pale blue; pollen ± white. Style arching adaxial to stamens, dividing opposite anther bases, style branches forked for 1 / 2 their length and recurved. Capsules broadly ovoid, trigonous, 8 11 ± 6.5 mm. Seeds globose, smooth, flattened at chalazal end, ± 1.3 mm diam., light brown. Chromosome number 2n =

46 38 S T R E L I T Z I A 35 (2015) 18. Flowering time: mid-june to mid-august. Figure 8. Distribution and biology: Lapeirousia exilis has a scattered range across Namaqualand from Steinkopf north of Springbok southward to the rugged northern edge of the Bokkeveld Escarpment northeast of Nieuwoudtville (Figure 7). Plants grow in stony ground in granitic gravel, sand, or occasionally clay. Pollination studies show the species to be pollinated by large bodied bees and bombyliid flies (Goldblatt et al. 1995). The fairly long perianth tube appears to indicate a pollinator with long mouthparts, but we have found that nectar is held fairly high in the tube and within reach of long-tongued bees and bombyliids. Diagnosis and relationships: although known since at least 1898 when it was collected by Maximilian Schlechter, Lapeirousia exilis was included in L. plicata (sometimes as L. caespitosa), often only tentatively, until 1972 when it was recognised as a separate species (Goldblatt 1972). When pressed, the plant and its flowers can appear similar to L. plicata and may be difficult to distinguish from that species. The similarity to L. plicata, which is acaulescent and has radially symmetric flowers, is superficial. In L. exilis the stem in well-grown plants is short but produced above the ground and the flower is zygomorphic. The stamens are unilateral and the lower tepals bear markings only on the horizontal lower tepals. The branches, when produced, are short, but the internodes are not as severely contracted as in L. plicata except in the most depauperate plants with only a few flowers. L. exilis has floral bracts typical of Lapeirousia and not like those found in L. plicata, which have elongated leaf-like tips above the sheathing bases. The seeds (Table 3) accord with sect. Chasmatocallis in their smooth surface and domed surface cells, thus quite unlike the sculptured, ± reticulate to rugose surface and flat surface cells of L. plicata and its acaulescent allies. Molecular data support the fundamental differences in flower, bract and seed morphology in allying L. exilis with sect. Chasmatocallis rather than to L. plicata (sect. Sophronia). Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): 2 km west of Steinkopf, shale hillside, ( BA), 16 July 1981, Goldblatt 6004 (MO); Sabies [Zabies], ( BC), 16 FIGURE 8. Lapeirousia exilis, Goldblatt Scale bar: 10 mm. Artist: John Manning.

47 S T R E L I T Z I A 35 (2015) 39 June 1898, M. Schlechter 99 (BOL, GRA); 11 km north of Concordia, ( DB), 30 June 1935, Salter 5540 (BM, BOL); north of Okiep, ( DB), Barker 9170 (NBG), Esterhuysen (BOL) (Gamoep): Goegap (Hester Malan) Nature Reserve, gravel pit along Pofadder road, ( CA), 1 July 1975, Rosch & Le Roux 1160 (PRE); ± 28 km east of Springbok toward Pofadder, sandy plains, ( CA), 23 Aug (fr.), Goldblatt & Manning 9650 (MO, NBG) (Kamiesberg): Farm Klein Lieslap, ± 14 km southwest of Kliprand, exposed granite sheets, ( DA), 17 May 1993, Snijman 1313 (NBG) (Loeriesfontein): Taaiboskloof, north slopes of Langberg, ( CA), 4 July 1970, Oliver 9501 (NBG) (Calvinia): 17.6 km along Rondekop road off R27, Soetlandsfontein River, ( AD), 6 June 2005, Manning 2948 (NBG); 10 July 2008, Manning s.n. (NBG). WESTERN CAPE (Vanrhynsdorp): Knersvlakte, Farm Kalkgat, ( BB), 16 June 1987, Boucher 5137 (NBG); 2 July 1998, Orthen s.n. (NBG). 7. Lapeirousia angustifolia Schltr. in Botanische Jahrbucher für Systematik 27: 105 (1900). Type: South Africa, [Western Cape], Zoutrivier, 15 July 1896, Schlechter 8144 (B, holo.!; BOL!, BM!, COI!, F, G!, GRA!, K!, PH!, PRE!, iso.). [Schlechter s handwritten collection number has also been misread as 8149 or 8188] Plants ( 180) mm high, stem simple or sometimes 1 or 2 branched from base, slightly compressed. Corm bell-shaped, 7 10 mm diam.; tunics brown, surface lightly ridged, basal rim smooth or bluntly lobed. Leaves few, lowermost longest, linear, ascending, usually exceeding stem, to 100( 180) mm long, ribbed, cauline leaves broader and shorter than basal, ± bractlike. Inflorescence a 6 15-flowered spike, spiral in flower; outer bracts green, firm, 8 13 mm long, oblong to broadly ovate, channelled, obtuse (retuse when opened), apex deflexed in profile, margins hyaline; inner bracts ± 1 / 2 as long, transparent with two green keels. Flowers zygomorphic, white to pale lilac or mauve, lower 3 tepals pale yellow at base and with violet median streak, tube pink on reverse, lower tepals each with small claw-like tooth in lower midline, with light sweet scent; perianth tube mm long, slender below, ± abruptly curved and expanded in upper 5 6 mm; tepals unequal, dorsal erect or slightly arched over anthers, mm, oblanceolate, upper lateral tepals outspread, lower 3 united basally for ± 2 mm, 7 9 mm long, directed downward Stamens unilateral, suberect; filaments 7 8 mm long, exserted mm; anthers parallel, mm long, lilac; pollen ± white. Style dividing opposite middle 1 / 3 of anthers, style branches ± 2.5 mm long, forked for slightly > 1 / 2 their length. Capsules obovoidtrigonous, 5 8 mm long, without locular ridges. Seeds globose, somewhat flattened to irregularly lobed at chalazal end, mm diam., brown, smooth with outer epidermal walls lightly colliculate. Chromosome number unknown. Flowering time: early July to late August. Distribution and biology: restricted to the Knersvlakte of southern Namaqualand (Figure 9), Lapeirousia angustifolia grows in sandy or light clay soils, sometimes in areas with a surface of quartz pebbles. Its appearance, size and exact flowering is very unpredictable from season to season in this area of low and unreliable rainfall. Pollination of the pale, faintly scented flowers is unknown but is likely to be by long-tongued bees. Records from central and northern Namaqualand (e.g. Marloth 6730 PRE, Aug. 1925, Garies-Okiep), are evidently incorrect given the absence of any recent collections from this relatively well-botanised area. Diagnosis and relationships: evidently first collected by the German botanist Rudolf FIGURE 9. Distribution of Lapeirousia angustifolia, ; L. pyramidalis subsp. pyramidalis, ; L. pyramidalis subsp. regalis,.

48 40 S T R E L I T Z I A 35 (2015) Schlechter in 1896 and described by him in 1900, Lapeirousia angustifolia has been inextricably confused with the widespread and fairly common L. pyramidalis. These two species are alike in general appearance and share derived, obtuse floral bracts, and were treated as synonymous by Goldblatt (1972). Recent field work, including pollination biology, suggests that Schlechter was correct in regarding L. angustifolia as a separate species. The flowers, like those of L. pyramidalis subsp. pyramidalis, are white to pale lilac in colour and also pleasantly, if not as strongly sweet scented, but are readily distinguished by the shorter perianth tube, mm long vs mm, the claw-like cusps on the lower tepals, and the small capsules lacking the wing-like locular ridges characteristic of L. pyramidalis. The small seeds, mm diam., and slightly wrinkled with colliculate epidermal cells also differ from the slightly larger seeds of L. pyramidalis subsp. pyramidalis, mm diam., which have derived, tuberculate epidermal cells (hedgehog-like under the microscope). The molecular phylogeny (Figure 1) confirms the very close relationship of the two taxa. Until more is known about Lapeirousia angustifolia, it is more consistent to treat it as separate from L. pyramidalis, not merely a shorttubed local variant. Differences in capsule and seed morphology are significant in determining relationships in Lapeirousia and in the case of L. angustifolia and L. pyramidalis suggest that the two should be regarded as separate, closely allied species. This conclusion is supported by the co-occurrence of both taxa near Hol River north of Vredendal, without evidence of hybridisation or introgression between them. Representative specimens SOUTH AFRICA. WESTERN CAPE (Vanrhynsdorp): Knersvlakte, Quaggaskop reserve, ( BC), 27 Aug. 1991, Goldblatt & Manning 9127 (MO, NBG); Knersvlakte, road to Quaggaskop from N7, ( BC), 25 Aug. 2008, Goldblatt & Manning (MO, NBG); 7 miles [11.2 km] north of Vredendal, Farm Liebendal, ( CB), 22 July 1970, Hall 3663 (NBG); 4 miles [6.4 km] north of Hol River Station, near Moedverloor, ( CB), 10 Aug, 1962, Nordenstam 900 (NBG, S); 6 miles [9.6 km] north of Klawer, ( DA), 22 Aug. 1950, Lewis 2347 (NBG, SAM); Farm Varschrivier, ( DA), 11 July 2001, Boucher 6724 (NBG); north of Vredendal, deep red soil, ( DA), 3 Aug. 2007, Van der Niet 451 (NBG); between Vanrhynsdorp and Vredendal, ( DA), 24 Aug. 2007, Goldblatt, Manning & Porter (MO, NBG); near Olifants River, without date, Drège 8508 (SAM); Farm Troe-troe, 3 km from Vanrhynsdorp on Nieuwoudtville road, ( DB), 3 Aug. 1977, Le Roux 2010 (NBG); Vanrhynsdorp, Mauwerskop, ( DB), 2 July 1990, Oliver 9450 (NBG). 8. Lapeirousia pyramidalis (Lam.) Goldblatt in Contributions from the Bolus Herbarium 4: 41 (1972). Ixia pyramidalis Lam.: 334 (1789). Witsenia pyramidalis (Lam.) Pers.: 42 (1805). Type: said to be from Isle de France [Mauritius], but certainly from western South Africa, said to be collected by Commerson but probably by P. Sonnerat s.n. (P-Herb. Jussieu, holo.!). [See additional synonymy under subsp. pyramidalis.] Plants (50 ) mm high, stem simple or 1 few-branched from near base, slightly compressed. Corm bell-shaped, ± 10 mm diam.; tunics light red-brown, lightly ridged, basal margin entire or bluntly lobed. Leaves 4 or 5, lowermost much longer than others, up to 160 mm long, linear to lanceolate, 2 6 mm wide, strongly ribbed, upper leaves shorter and broader, becoming progressively bract-like above. Inflorescences 6 20-flowered spikes, initially 2-ranked, later becoming spiral; outer bracts green, firm, ± succulent, mm long, channelled, ovate when flattened, obtuse in side view (deltoid and retuse when unfolded), apex deflexed in profile, margins hyaline; inner bracts 1 / 3 to 1 / 2 as long, transparent with 2 green keels. Flowers zygomorphic, either white to pale mauve, pale blue or pale pink, lower tepals each with a small triangular to diamond-shaped median mark in lower half (subsp. pyramidalis) or violet, purple or dark carmine, lower tepals each with white or cream markings near base and a central zone of darker pigment (subsp. regalis), either intensely sweet scented day and night (subsp. pyramidalis) or evidently scentless (subsp. regalis); perianth tube mm long, slender, straight or widening slightly in upper 5 mm; tepals subequal, narrowly to broadly ovate, narrowed into claw-like base ± 1.5 mm long, lower tepals each occasionally with a small claw-like cusp near base, dorsal tepal mm, erect, held apart from others, upper lateral tepals slightly reflexed, lower 3 tepals usually held closely together and at right angles to tube, 8 5 mm. Stamens unilateral, ±

49 S T R E L I T Z I A 35 (2015) 41 erect to ascending; filaments 5 6 mm long, exserted ± 3.5 mm; anthers ± 3 mm long, usually pale yellow to white; pollen ± white. Style dividing between base and apex of anthers or slightly beyond; branches ± 2.5 mm long, forked for 1 / 2 their length, often becoming tangled in anthers. Capsules obovoid, 8 10 mm long, with wing-like locular ridges in distal half. Seeds ± globose to ovoid, usually somewhat flattened at chalazal end, mm, epidermal cells tuberculate. Chromosome number 2n = 18. Flowering time: July to September. Distribution and biology: Lapeirousia pyramidalis, one of the more widespread species of Lapeirousia, extends from the southern edge of Namaqualand though dry areas of the northwestern Cape to the Little Karoo and southern Cape (Goldblatt 1972) (Figure 9). Over much of its range it has a fairly consistent appearance. Plants are compact with broadly obtuse floral bracts, and the flowers are white to pale blue or pale pink to lilac with a perianth tube mm long, slightly curved and wider in the upper 5 7 mm, and always strongly scented, both during the day and in the evening. In the west of its range, in the Olifants and Biedouw River valleys, and at Karoo Poort to the south, plants segregated as subsp. regalis are distinguished by bright red, purple, or dark violet flowers with a long, nearly straight perianth tube, mm long, and have no discernible odour. We considered the two sets of populations too similar to be treated as separate species (Goldblatt & Manning 1994). As far as we know, the two forms never grow sympatrically and their ranges are more or less complementary. They also favour different habitats: subsp. pyramidalis normally grows on shale and clay soils, but subsp. regalis has only been recorded on stony sandstone ground. The floral differences between the two subspecies reflect a shift in pollination strategy (Goldblatt et al. 1995). Strongly scented subsp. pyramidalis is thought to be pollinated primarily by sphinx moths (Hippotion celerio is the only insect so far identified as a pollinator), which are most active at sunset and at night, whereas subsp. regalis is a member of the Prosoeca peringueyi pollination guild. This guild also includes L. silenoides (Goldblatt et al. 1995), which has flowers very like those of subsp. regalis, with a long, nearly straight tube, erect dorsal tepal, and darkly pigmented perianth with contrastingly pale nectar guides. Volume and quality of nectar produced by the two subspecies appears to reflect their different pollination systems: subsp. pyramidalis produces up to 3 ml of nectar of 30 34% sucrose equivalents whereas samples of subsp. regalis show that it produces up to 4.8 ml of nectar of approximately 28% sucrose equivalents. Diagnosis and relationships: Lapeirousia pyramidalis can always be immediately recognised by the broadly ovate to deltoid bracts with obtuse to retuse apices. In profile the bracts do not taper to a point but curve outward and have blunt tips. The flowers have shortly clawed, subequal tepals with broadly oval to rounded limbs resembling the Namaqualand species, L. silenoides and we consider the two to be closely allied. Both also have unusual capsules with prominent ridges on the locules, a feature shared with the local Namaqualand endemic, L. verecunda, and also similar red-brown corms with a ridged or lobed basal margin. The tuberculate seeds of L. pyramidalis are unusual in Lapeirousia, in which most species of sect. Chasmatocallis, including those of L. silenoides and L. verecunda, have epidermal cells with evenly domed (colliculate) outer walls. The difference is often striking, making it possible to distinguish the seeds of L. pyramidalis immediately from other species of the genus. There is a marked size difference between the seeds of the two subspecies. Over much of its range subsp. pyramidalis has seeds mm diam., but subsp. regalis has seeds ± 1 / 2 this size, mm diam. Outgroup comparison with related species suggests that the darkly pigmented flowers of L. pyramidalis subsp. regalis are plesiomorphic. A shift to pollination by sphinx moths is presumably a specialisation in the species, and associated with the dilution and loss of perianth pigmentation and the development of intense, clove-dominated floral scent. This hypothetical scenario is supported by the molecular phylogeny (Figure 1). History: Lapeirousia pyramidalis has a somewhat confused taxonomic history. The species was described by the French biologist, J.P.B. Lamarck (1789), based on a collection attributed to the botanist, Philibert Commerson ( ), who is not known to have ever visited the Cape (Gunn & Codd 1981). More likely

50 42 S T R E L I T Z I A 35 (2015) his colleague, Pierre Sonnerat, was responsible for the type collection. Sonnerat briefly visited the Cape on his journey to Mauritius, where he worked with Commerson, who had settled there. Plants cultivated in Vienna were described as Gladiolus fissifolius by N.J. Jacquin in 1792, and the fine illustration published shortly thereafter (Jacquin 1794) shows this clearly to be the same as Lamarck s species. Ker Gawler (1804) transferred G. fissifolius to Lapeirousia and the name remained in use for the species until 1972 when it became clear that Ixia pyramidalis was an earlier synonym, and that, in addition, Gladiolus fissifolius was nomenclaturally illegitimate, being superfluous for G. anceps, now L. anceps. The species was collected by C.P. Thunberg in the 1770s and later described by him as G. bracteatus (Thunberg 1800). Lapeirousia fissifolia was renamed L. homoidea F.W. Klatt in 1894, but the reason for his action is not clear. Key to subspecies 1a Flowers whitish to pale blue (occasionally pale pink to mauve), strongly scented; perianth tube mm long, cylindric below but expanded and curved outward in distal 5 mm.... 8a. subsp. pyramidalis 1b Flowers dark purple, violet-blue or red, not scented; perianth tube (35 )40 55 mm long, ± cylindric throughout... 8b. subsp. regalis 8a. subsp. pyramidalis Gladiolus fissifolius Jacq. in Collecteanea 4: 164 (1792) nom. illegit. superfl. pro G. anceps L.f. (= Lapeirousia anceps (L.f.) Ker Gawl.). Lapeirousia fissifolia Ker Gawl.: 238 (1804), as nom. nov. pro G. fissifolius Jacq. nom. illegit.; Baker: 92 (1896). Ovieda fissifolia (Ker Gawl.) Spreng.: 147 (1824). Peyrousia fissifolia (Ker Gawl.) Poir.: 364 (1826). Meristostigma fissifolium (Ker Gawl.) A.Dietr.: 595 (1833). L. homoidea Klatt: 191 (1895) as nom. nov. pro Gladiolus fissifolius Jacq., nom. illegit. Type: South Africa, without precise locality or collector, illustration in Jacq.: t. 268 (1794). Gladiolus bracteatus Thunb.: 186 (1800). Lapeirousia bracteata (Thunb.) Ker Gawl.: sub. t (1810). Ovieda bracteata (Thunb.) Spreng.: 147 (1824). Peyrousia bracteata (Thunb.) Sweet: 499 (1830a). Meristostigma bracteatum (Thunb.) A.Dietr.: 597 (1833). Type: South Africa, without precise locality, Thunberg s.n. (UPS-THUNB 1008, holo.!). Flowers whitish to pale bluish, or flushed with pale pink, lower tepals cream at base with small basal median streak of purple and larger median purple mark, intensely sweetly scented night and day; perianth tube mm long, narrow and cylindric below, broadened and curving outward in upper 4 5 mm; tepals subequal, dorsal ± erect, mm, upper laterals spreading to reflexed, lower 3 directed downward, with short claw ± mm, limbs abruptly expanded, broadly oval, mm. Stamens unilateral, ± erect; filaments 5 6 mm long, exserted ± 3.5 mm; anthers ± 3 mm long, cream or light purple; pollen yellowish to white. Style usually dividing shortly above anther bases, style branches becoming tangled in anthers. Seeds mm diam. Flowering time: late July to September. Distribution and biology: fairly widespread across the southern African winter rainfall zone, subsp. pyramidalis extends from Worcester and Grabouw in the southwest through the Little Karoo and the Overberg to Riversdale in the east and northward into the arid Tanqua Karoo, and dry sites along the interior Cederberg and Bokkeveld Mtns as far north as Loeriesfontein and Nuwerus (Figure 9). Populations in the Tanqua Karoo vary considerably in colour, from white to deep pink but even the darker forms are fragrant, with the perianth tube curved outwards at the end and thus represent the typical subspecies and not subsp. regalis. Plants are usually found on heavier soils and are mostly encountered on clays and shales of the Bokkeveld and Malmesbury Systems. Plants also occur in stony alluvial sand but rarely on the poor sandy soils of the Cape Sandstone formation. The closely related L. angustifolia, which has smaller flowers with a shorter perianth tube, mm long, and faintly scented flowers, is restricted to dry flats of the Knersvlakte west and north of Vanrhynsdorp. Both species have been recorded near Hol River, north of Vredendal. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Wuppertal): Stompiesfontein, Calvinia district, ( BA), 26 July 1941, Compton (NBG), Esterhuysen 5361 (BOL); Tankwa National Park, Volmoersfontein, ( BD), 20 July 2006, Rösch 384 (NBG); WESTERN CAPE (Lepelfontein): Koekenaap, coast between Olifants River and Brand-se-Baai, ( BD), Aug. 2002, Desmet 3595 (NBG) (Vanrhynsdorp):

51 S T R E L I T Z I A 35 (2015) km north of Koekenaap on road to Kotzesrus above Sout River, ( AA), 16 Sept. 2001, Goldblatt & Porter (MO, NBG); Nieuwerust [Nuwerus], ( AB), 7 Sept. 1945, Barker 3731 (NBG); 7 km on Kliprand Road east of Nuwerus, ( AB), 9 Sept [mainly fruiting], Thompson 2840 (NBG); 2 miles [3 km] south of Nuwerus, ( AB), Lewis 1385 (SAM); Hol River, ( AD), 18 Aug. 1970, Hall 3731 (NBG); Kareeberg, southeast of Bitterfontein, ( BA), 15 Aug. 1997, Desmet & Ellis 1226 (NBG); Olifants River, 20.8 km from National Road at Rooiberg turn-off, ( BC), 5 August 1977, Le Roux 2118 (NBG); between Vanrhynsdorp and Pass, ( BD), Barker 6439 (NBG) (Wuppertal): 4 km along road to Middelpos from Ceres/Calvinia road, Blaauwboschkolk, ( DA), 31 Aug. 1982, Snijman 609 (NBG) (Worcester): Kluitjieskraal, Ceres road ( AC), Aug. 1888, Tyson 1541 (G, K, SAM) (Worcester): Karoopoort, ( BA), 30 July 1950, Hall 282 (NBG), 26 Aug. 1935, Compton 5409 (BOL, NBG), 6 Aug. 2002, Manning 2762A (NBG);between Little and Great Doorn Rivers, ( BB), Burchell 1213 (K); between Osplaats and Tunnel siding, Hex River Valley, ( BC), without date, Rogers (G, K); Riverside, Worcester, 5 Sept ( CA), Walters 187, 590 (NBG); Brandvlei, ( CB), Hutchinson 134 (K, PRE); Karoo Veldreservaat [Veld Reserve], 29 Aug. 1962, Olivier 102 (NBG, PRE); Hex River Kloof, ( AD), Drège s.n. (K) (Montagu): near Ashton, ( CC), Compton (NBG); 10 miles [15 km] from Montagu to Barrydale, ( CD), Hurling & Neil s.n. (BOL 30715); Warmwaterberg, ( DD), 11 July 1948, Compton (NBG) (Ladismith): between Bosluiskloof and Prince Albert ( BC), Lewis 4435 (SAM) (Oudtshoorn): near Prince Albert, ( AA), Krige s.n. (BOL 10631); Great Doorn River ( CA), Thorne s.n. SAM51700 (SAM); Moeras River bridge, Robinson Pass road, ( CC), Van Niekerk 463 (BOL) (Caledon): roadside near Grabouw, ( AA), Garside 4496 (K); 8 miles [± 12 km] north of Bot River, Goldblatt 298 (BOL); Caledon Villiersdorp road intersection, ( AB), Barker 26 (BOL), between Houw Hoek and Caledon, ( AB), Oct. 1887, MacOwan 805 (BOL, G, GRA, K); Appelkraal, Riviersonderend, ( BB), Zeyher 4025 (G, K, SAM); Fairfield ( BD), Acocks (PRE); Napier, Barker 2510 (BOL, NBG); Bond 477 (NBG) (Bredasdorp): Bontebok Park, ( AB), Acocks (PRE); Bredasdorp ( CA), Metelerkamp s.n. (NBG6182, 6183) (Riversdale): Riversdale, ( AA), Muir 181 (BOL). 8b. subsp. regalis Goldblatt & J.C.Manning in Novon 4: 342 (1994). Type: South Africa, [Western Cape], Middelplaas, ± 5 km from Farm Biedouw on road to Wuppertal, 8 July 1941, Leipoldt 3866 (BOL, holo.!; BOL!, NBG!, iso.). Flowers red, purple, violet or navy blue, lower tepals with cream markings on claws and darker blue or red markings in centre of limbs, unscented; perianth tube 40 50( 55) mm long, cylindric and ± straight; tepals subequal, mm long, dorsal held apart from others when fully open. Stamens and style as in subsp. pyramidalis. Capsules oblongtrigonous, locules with prominent ridges, 7 9 mm long. Seeds mm diam. Flowering time: late July to early September. Distribution and biology: subspecies regalis is restricted to the northwestern corner of Western Cape and occurs in the middle Olifants River Valley between Citrusdal and Klawer, extending inland into the interior valleys of the Cederberg and Pakhuis Mtns (Figure 9). Plants typically grow on light sandy soils, in stony habitats. The flowers are adapted for pollination by the longproboscid fly, Prosoeca peringueyi (Goldblatt et al. 1995; Manning & Goldblatt 1996). Typical of species with this pollination system, the deeply pigmented red to violet flowers have pale markings on the tepals and lack scent. Representative specimens SOUTH AFRICA. WESTERN CAPE (Vanrhynsdorp): Olifants River Barrage, ( DD), 22 July 1941, Esterhuysen 5377 (NBG) (Clanwilliam): 9 miles [± 14 km] north of Clanwilliam, ( BB), 22 Aug. 1950, Barker 6420 (NBG); between Nardouwskloof and Bulshoek, 12 Aug. 1976, Goldblatt 3837 (MO); near Clanwilliam, Aug. 1945, L. Bolus s.n. BOL23183 (BOL); Olifants River valley south of Clanwilliam, ( BD), Martin s.n. BOL (BOL) (Wuppertal): top of Biedouw valley between Blinkvlei and the Doorn River, ( AB), 23 July 1961, Lewis 5824 (NBG); 3 miles [4.8 km] from Biedouw on road to Wuppertal, ( AB), 8 July 1941, Leipoldt 3866 (NBG); 15 km north of Citrusdal, ( CA), Lewis 2350 (SAM). B. Sect. Sophronia (Spp. 9 13) B. Sect. Sophronia (Licht. ex Roem. & Schult.) Goldblatt & J.C.Manning in Annals of the Missouri Botanical Garden 77: 373 (1990). Sophronia Licht. ex Roem. & Schult.: 482 (1817). Lapeirousia subg. Sophronia (Licht. ex Roem. & Schult.) Baker: 482 (1892). Type: S. caespitosa Licht. ex Roem. & Schult. (= Lapeirousia plicata (Jacq.) Diels). Corm bell-shaped; tunics with basal rim not produced into teeth or spines. Stem subterranean, internodes at ground level crowded. Inflorescence a head-like, congested spike. Flowers radially symmetric, with perianth tube ± cylin-

52 44 S T R E L I T Z I A 35 (2015) dric, short to elongate; stamens symmetrically arranged. Basic chromosome number x = Lapeirousia plicata (Jacq.) Diels in Engler & Prantl, Die naturlichen Pflantzenfamilien ed. 2, 15a: 488 (1930); Goldblatt: 46 (1972). Galaxia plicata Jacq.: t. 292 (1795). Ixia heterophylla Willd.: 199 (1798), as a new name for G. plicata in Ixia (not I. plicata L.). Meristostigma heterophyllum (Willd.) A.Dietr.: 199 (1833). L. heterophylla (Willd.) R.C.Foster: 4 (1939). L. fasciculata Ker Gawl.: 238 (1804), superfluous name for Galaxia plicata Jacq. Ovieda fasciculata (Ker Gawl.) Spreng.: 147 (1824). Peyrousia fasciculata (Ker Gawl.) Sweet: 499 (1830a). Type: South Africa, without precise locality, illustration in Jacq.: t. 292 (1795) (original specimens, if any existed, believed destroyed). Epitype: South Africa, Western Cape, Matjiesfontein, gravel flats near village, flowers white, 18 July 1974, Goldblatt 2115 (MO!, NBG!). [Additional synonymy under the subspecies.] Plants small, acaulescent (stem entirely below ground), forming diffuse or compact cushions, seldom > 50 mm high. Corm bell-shaped, red-brown, rough-surfaced, basal rim slightly crenate. Foliage leaves up to 3, lowermost longest, usually exceeding inflorescence, up to 300 mm long, blades linear, suberect to strongly inclined, closely ribbed, margins of sheaths undulate or often crisped, remaining leaves broader than basal. Inflorescence congested, head-like, few- to several -flowered; bracts ovate at base, outer larger than inner, with apex often produced as ribbed leaf-like tip, margins usually crisped especially in basal half. Flowers actinomorphic, pale blue, ± white or pale yellow, with blue to violet spot at base of each tepal, scented of cloves or with light, sweet odour; perianth tube mm long, ± cylindric, straight; tepals subequal, ± lanceolate, 7 9 mm long, acute, spreading or often slightly deflexed when fully open. Stamens symmetrically arranged around style; filaments exserted mm, arching outward later in anthesis; anthers mostly mm long, usually blue (or ± white), initially erect, later ± horizontal. Style dividing opposite base to middle of anthers, rarely below anther bases, style branches forked for 1 / 3 to 1 / 2 their length or undivided (subsp. effurcata). Capsules ovoid-oblong, 8 10 mm long. Seeds 6 9 per locule, ± globose, with reticulate to rugulose surface, dark brown, mm diam. Chromosome number 2n = 16 (subsp. plicata). Flowering time: autumn, winter or spring, mostly June to mid-august, occasionally in May. Figure 10A. Distribution and biology: widespread in southern Africa, Lapeirousia plicata extends from Vanrhynsdorp and Robertson in the southwest, through the Little and Great Karoo to Northern Cape, North West Province, Free State and into southern Namibia (Figure 11). Plants grow in fairly heavy clay soils, less often in light clay or sand, in fairly flat places, as well as on limestone or calcrete, usually where drainage is poor. Flowering is earliest in the north and east of its range, mainly in June but sometimes in May or July; in the west and south flowering usually begins in July lasting into August. Diagnosis and relationships: smallest of the several acaulescent species of sect. Sophronia, the tufts of Lapeirousia plicata seldom reach 50 mm including the flowers. The perianth tube ranges from 12 to 20 mm long in southern and western populations to as much as 30 mm in the northeastern populations. A distinctive feature of L. plicata is orientation of the stamens, the filaments of which are initially erect but arch outward during anthesis so that the dehisced anthers come to lie in a ± horizontal position. The style typically divides opposite the base to middle of the anthers, but sometimes below the anther bases and rarely slightly beyond the anther tips. Typification of Lapeirousia plicata has never been in doubt: the painting in Jacquin s Icones plantarum rariorum (Jacquin 1795), entitled Galaxia plicata represents the form of the species with white flowers and a style described as trifid. The type has until now been associated with the form of the species occurring in the western Karoo, as at Calvinia and nearby, but these often have blue flowers and almost always style branches divided for a third to half their length. The name can, however, be applied to whiteflowered plants from the Matjiesfontein area of Western Cape, where the species is common and plants have style branches notched apically, the style thus seeming trifid. We designate an

53 S T R E L I T Z I A 35 (2015) 45 A B FIGURE 10. A, Lapeirousia plicata subsp. plicata, Goldblatt 6066A; B, L. montana, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

54 46 S T R E L I T Z I A 35 (2015) FIGURE 11. Distribution of Lapeirousia plicata subsp. effurcata, ; subsp. foliosa, ; subsp. plicata,. epitype for L. plicata stabilising the application of the name, which otherwise might be associated with southern Cape plants, described by Lewis (1952) as L. effurcata, which always has white flowers but undivided style branches. Three subspecies of Lapeirousia plicata were recognised by Goldblatt (1972), based on morphological differences and by a moderate degree of geographical discontinuity. One of these, subsp. longifolia, is a separate species here described as L. kalahariensis. We continue, however, to recognise the other two subspecies, subsp. effurcata, which has unusually broad, widely channelled outer floral bracts with undulate margins and consistently white flowers and undivided style branches, whereas subsp. plicata has blue or white flowers, with apically notched to deeply divided style branches and somewhat narrower floral bracts, sometimes with short, leafy, ribbed apices. A third series of populations from Bushmanland, the Great and Upper Karoo extending into Free State mostly have white or cream-coloured flowers (sometimes described as pale yellow) but rarely blue and the outer floral bracts have leaf-like, ribbed tips up to 30 mm long, sometimes unifacial towards the tips. These populations are also usually early blooming, often in flower in May and June. We suggest separation at subspecies rank as subsp. foliosa for this morph, noting that there are populations intermediate for these features. This subspecies has also been described at species rank as L. galaxioides by J.G. Baker (1892), a name that he applied to plants from the Kimberley area, now in Northern Cape. Key to subspecies 1a Outer floral bracts channelled to apices or with short, ribbed, leafy tips < 10 mm long; perianth tube mostly mm long; flowers blue or white with blue to violet mark in lower midline of tepals: 2a Style branches usually forked for 1 / 3 1 / 2 their length, rarely undivided; flowers blue or white a. subsp. plicata 2a Style branches always undivided; flowers white, flushed blue outside. 9b. subsp. effurcata 1b Outer floral bracts with ribbed, leafy tips up to 30 mm long; perianth tube mostly mm long; flowers white to cream (pale yellow), occasionally with a blue to purple spot near tepal bases...9c. subsp. foliosa 9a. subsp. plicata Leaves usually 2 or 3, only lowermost with unifacial blade, ± linear, to mm, suberect, ribbed, usually ciliate or papillate on rib edges, margins plane or undulate, ciliolate. Bracts

55 S T R E L I T Z I A 35 (2015) 47 green, outer usually falcate, channelled throughout or with short, ribbed leafy apex up to 10 mm long, margins undulate to slightly crisped, usually ciliate. Flowers either blue with white zone at tepal bases and violet or purple triangular mark at distal edge of white zone or white with violet mark near tepal bases; perianth tube mostly mm long; tepals 7 8 mm long, spreading or slightly reflexed. Stamens: filaments exserted ± 2 mm, diverging; anthers mm long, blue to purple. Style dividing below to opposite base (rarely opposite middle) of anthers, style branches usually forked for 1 / 2 to 1 / 3 their length or shortly notched at apex, mm long, recurving. Flowering time: mainly July and early August. Figure 10A. Distribution: restricted to the southern African winter rainfall zone, subsp. plicata occurs in the Western Karoo from Calvinia and the Bokkeveld Plateau south through the Tanqua Basin to Touws River, Matjiesfontein and Laingsburg where it meets the range of subsp. foliosa in the western Little Karoo (Figure 11). Flowers have been reported as pollinated by honey bees (Apis mellifera) while foraging for nectar (Goldblatt et al. 1995) in the Calvinia area. Sophronia (= Lapeirousia) caespitosa is generally regarded as a synonym of L. plicata (Goldblatt 1972), but its typification is problematic. The type, evidently at one time at the Berlin Herbarium, is no longer extant, and we follow the interpretation of the species of past authors, treating it as a synonym of the Jacquin plant (although sometimes under the illegitimate, superfluous names L. heterophylla or L. fasciculata). Literal reading of the protologue is, however, a source of confusion. Flower colour is described as yellow (flava) and the divided style branches (an important consideration) are not mentioned. Nevertheless, the number of stamens (three) leaves no doubt that this is a species of Iridaceae and the radially symmetric flowers and acaulescent (caespitose) habit are consistent with L. plicata. Plants of L. plicata from near the type locality, Sakrivierspoort, which is southeast of Fraserburg, and near Fraserburg itself, have cream-coloured flowers and otherwise match L. plicata vegetatively, thus are reasonably consistent with the current interpretation of the name. We have chosen a neotype for the taxon, one from close to the type locality. It is now subsp. foliosa of L. plicata. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Calvinia): Calvinia, Akkerdam, ( BA), Lewis 5714 (NBG), 24 July 1981, Goldblatt 6066A (MO); Calvinia to Nieuwoudtville, ( BC), Goldblatt 3892 (MO, NBG); Volstruisvlakte, ( DC), 26 July, Bond 1169 (NBG) (Sutherland): Komsberg Nature Reserve, m, ( DB), 4 Aug. 2008, Rosch 175 (NBG). WESTERN CAPE (Montagu): foot of the Voetpadsberg, 21.5 km east of Touws River, ( AC), Goldblatt (MO); Matjiesfontein, gravel flats, ( BA), 26 July 2001, Burgoyne 8215B (PRE). 9b. subsp. effurcata (G.J.Lewis) Goldblatt in Annals of the Bolus Herbarium 4: 51 (1972). Lapeirousia effurcata G.J.Lewis: 7 (1952). Type: South Africa, [Western Cape], near Riversdale, Elands Drift, limestone koppies, Aug. 1946, Ferguson s.n. (BOL 24923!, SAM!, syn.). Plants with foliage leaves usually 2 or 3, only lowermost unifacial, ± linear, to mm, suberect to spreading, margins plane. Bracts green, outer much exceeding flowers, channelled throughout, margins undulate to slightly crisped. Flowers usually white (rarely pale blue) with blue mark in lower midline of tepals; perianth tube mostly mm long; tepals ± mm, ± spreading. Stamens: filaments exserted ± 3 mm, arching outward; anthers ± 2 mm long, blue. Style dividing slightly below to opposite base of anthers, style branches entire, recurved, ± 1 mm long. Flowering time: mainly July and August. Distribution: subsp. effurcata occurs on stony flats in the Worcester Robertson Karoo, the western Little Karoo and locally in the limestone belt of the southern Cape near Riversdale (Figure 11). The Riversdale populations, including the type collection, are from limestone outcrops, a habitat unusual for any species of Lapeirousia, but other collections are from stony, sandy or clay ground. A curious feature of subsp. effurcata is that the flowers are reported to be heavily scented toward evening. Flowers of subsp. plicata have a light, sweet scent during the day, but we have not noted an increase in scent toward evening.

56 48 S T R E L I T Z I A 35 (2015) Representative specimens SOUTH AFRICA. WESTERN CAPE (Worcester): 10 miles [15 km] west of Worcester to Robertson, ( DA), Barker 8256 (NBG) (Montagu): Anysberg Nature Reserve, near Vrede homestead, sandy flats, ( BC), 12 Aug. 1993, Germishuizen 6732 (PRE); 6 miles [9 km] east of Montagu, ( CC), Barnard s.n. (BOL); Montagu Baths (heavily scented toward evening), ( CC) Aug & 1920, Page s.n. (BOL); Kalkoensnes, Montagu, ( CC), 9 July, Schonken 263 (NBG) (Ladismith): Seweweekspoort, southern end, ( AD), Levyns 7442a (BOL); Seweweekspoort, middle of pass, ( AD), Levyns 7442b (BOL) (Riversdale): 18 km south of Riversdale, limestone flats at turnoff to Watergat, ( AA), 3 Aug. 1976, Goldblatt 3744 (MO). 9c. subsp. foliosa Goldblatt & Manning, subsp. nov. TYPE. South Africa, Northern Cape, 3019 (Calvinia): Loeriesfontein Dist., between Graafwater and Donkiedam, ( CC), 4 July 1990, Oliver 9484 (NBG, holo.). Sophronia caespitosa Licht. ex Roem. & Schult.: 482 (1817). Lapeirousia caespitosa (Licht. ex Roem. & Schult.) Baker: 174 (1892). Type: South Africa, [Northern Cape], Sakrivierspoort, Lichtenstein s.n. (probably originally at B, no longer extant). Neotype: South Africa, Northern Cape, 10 km northeast of Fraserburg on Loxton Road, 5 July 2000, Manning et al (NBG, holo). Lapeirousia galaxioides Baker: 174 (1892). Type: South Africa, [Northern Cape], diamond fields at Klipdrift, without date, Bowker s.n. (K, holo!). Plants often larger in vegetative features than subsp. plicata, with foliage leaves 2 or 3, lowermost longest, mm, suberect, ribbed, smooth or ciliate to papillate on rib edges, margins usually plane, often ciliate. Outer bracts upright below, tapering distally to leaf-like, channelled or unifacial tip mostly mm long, linear, straight or falcate. Flowers white to cream-coloured (pale yellow), occasionally pale blue, tepals often with purple spot in lower midline, often (?always) scented of cloves; perianth tube mostly mm long, well exserted from sheathing part of bracts; tepals 7 9 mm long. Stamens: filaments exserted ± 2 mm; anthers 2 3 mm long, blue (often fading white in herbarium specimens). Style dividing opposite base to apex of anthers (rarely exceeding anthers); style branches mm long, usually deeply forked. Flowering time: mainly May and June. Distribution: widespread across arid parts of southern Africa, subsp. foliosa extends from the Knersvlakte of southern Namaqualand in Western Cape, along the eastern margins of Namaqualand north to southern Namibia and east across Bushmanland to Gordonia, the Kimberley District, western Free State, the Upper Karoo as far east as Colesberg and Murraysburg and with records in Eastern Cape near Cradock (Figure 11). Diagnosis: subsp. foliosa is recognised by the floral bracts with leaf-like, ribbed, unifacial tips at least 12 and up to 40 mm long, versus floral bracts usually channelled to the tips or with short, unifacial tips up to 10 mm long. The distinction is not absolute, hence our rank of subspecies for the plant. Flowers are usually white to creamcoloured, occasionally pale yellow and are strongly clove scented according to notes with several collections. The flowers are sometimes larger than those of subsp. effurcata and subsp. plicata, with a perianth tube mm long. We include plants from the Roggeveld escarpment in subsp. foliosa, evidently the only blue-flowered populations of the subspecies, on account of the floral bracts with leafy tips up to 20 mm long. Representative specimens NAMIBIA (Warmbad): Warmbad, Farm Vrede, quartzite hill white, blue anthers fading to white, ( BD), 16 June 2005, Bartsch et al (K); Farm Graswater, Warmbad, ( BD), 17 May 1963, Giess, Volk & Bleissner 7041 (WIND); Farm Sperlingsspütz, granietgrobsandfläche, ( CA), 26 May 1972, Giess & Müller 12275A (WIND); Farm Eendoorn, in river sand, ( DB), 26 May 1972, Giess & Müller 12275B (WIND). SOUTH AFRICA. EASTERN CAPE (Somerset East): Cradock District, ( BB), May 1904, Rogers s.n. (BOL10551, MO). FREE STATE (Kroonstad): Kroonstad, ( CA), Pont 260 (PRE) (Boshof): Smitskraal, Boshof, fragrant, ( CA), June 1911, Burtt Davy (BOL) (Jagersfontein): Fauresmith, ( CB), Henrici 2654 (PRE) (Bloemfontein): near Bloemfontein, ( AA), Eyre s.n. (BOL, MO), Potts s.n. (BOL30761). NORTHERN CAPE (Vioolsdrif): Richtersveld, base of Tatasberg, ( AC), Williams 3543 (NBG) (Kakamas): ± 20 km west of Augrabies, road to Onseep-

57 S T R E L I T Z I A 35 (2015) 49 kans via Nous Wes, scented of cloves, ( CA), 23 June 1999, Meyer 2117 (PRE) (Upington): near Upington, ( AC), Glover (BOL) (Glen Lyon): Grie kwastad, Farm Elim, banks of spruit above waterfall, ( AA), May 1942, Cooke 6582A (KMG); Hay District, Farm Highlands, gravel and sandy ridges, ( AA), May 1942, Cooke 6582B (KMG Hay District, Bingap, Van Sittart Mtns, ( CD), 18 June 1938, Acocks s.n. (BOL 30737, K) (Grie kwastad): Griquatown, m, ( CC), June 1895, Marloth 2103 (NBG, PRE); dolomite slope, Brakpan near Campbell, ( DB), 5 June 1936, Acocks 414 (PRE) (Kimberley): Warrenton, ( BB), Adams 151 (GRA); Kimberley, ( DB), Wilman s.n. (NBG) (Gamoep): Haramoep near Pofadder, ( BA), Mauve 4143 (PRE); ± 27 km east of Springbok, sandy plains, ( CA), 2 Aug. 1994, Goldblatt & Manning 9896 (MO, NBG) (Pofadder): 30 miles [± 47 km] west of Pofadder, ( AC), 25 May 1961, Schlieben 8989 (B, BOL, MO, PRE) (Prieska): Prieska, ( DA), Bryant 313 (BOL, PRE) (Hopetown): Modder River, Kimberley, ( BA), Galpin s.n. (BOL30724) (Kamiesberg): ± 10 km north of Kliprand, red sand flats, ( DA), 13 Sept (fr.), Goldblatt & Manning 9730 (MO, NBG) (Colesberg): Colesberg, ( CA), Shaw s.n. (K) (Calvinia): Roggeveld Escarpment, Top of Perdekloof west of Middelpos, ( DD). 3 Aug. 1998, Goldblatt & Manning (MO) (Merweville): between Great Reed River and Stinkfontein, ( AC), Burchell 1394 (G, K, P) (Loxton): 36.5 km northeast of Loxton, ( BC), 5 July 2000, Manning 2266 (NBG); 3124 (Richmond): Vlakplaats [Vlakplaas], Richmond Dist. ( CB), H. Bolus 3827 (BOL). WESTERN CAPE (Vanrhynsdorp): Knersvlakte, Douse-the-Glim, ( BB), Oliver 9497 (NBG); Vars River [Knersvlakte], ( BC), 12 July 1896, Schlechter 8149 (BOL, GRA, K, MO) (Victoria West): between Middelkop and Murraysburg, ( DC), June 1879, Tyson 276 (BOL, GRA, MO, SAM) (Oudtshoorn): Swartberg near Klaarstroom, ( BC), Drège 2188 (K, MO, SAM). 10. Lapeirousia oreogena Schltr. ex Goldblatt in Contributions from the Bolus Herbarium 4: 53 (1972); t (1973). Type: South Africa, [Northern Cape], 6 km east of Nieuwoudtville, 14 Sept. 1970, Goldblatt 541 (BOL, holo.!; K!, MO!, PRE!, iso.). Plants forming dense tufts at ground level, mm high including flowers, stem subterranean, upper internodes contracted. Corm bell-shaped, ± 10 mm diam.; tunics light brown, surface lightly rugose. Foliage leaves usually solitary, linear, straight, ribbed, mm long. Inflorescence several- to many-flowered, congested in head-like tuft; outer bracts mm long, bifacial and channelled throughout, usually ± falcate, often twisted, margins undulate and crisped, inner up to 1 / 2 as long, margins often plane. Flowers actinomorphic, salverform, dark blue-violet, almost black in centre with white triangular to arrowhead markings close to tepal bases; perianth tube erect, cylindric, mm long; tepals subequal, mm long, lanceolate, acute, spreading but becoming slightly deflexed with age. Stamens symmetrically disposed around style, erect; filaments exserted ± 3 mm; anthers erect, contiguous around style, ± 3.5 mm long. Style erect, usually dividing opposite middle 1 / 3 of anthers, style branches ± 2.5 mm long, deeply forked and recurved. Capsules ovoid to oblong, 7 10 mm long, without locular ridges. Seeds globose, ± 2 mm diam., red-brown, reticulately sculptured. Chromosome number 2n = 16. Flowering time: August to mid-september, rarely late July. Distribution and biology: restricted to the Calvinia District of Northern Cape, Lapeirousia oreogena is common and well known from the rocky, dolerite hills just to the north of Nieuwoudtville, but it extends south at least as far as the Farm Matjesfontein, ± 12 km from Nieuwoudtville, and has also been recorded southeast of Calvinia at Driefontein-se-Berg and northeast of the town in the Agterhantam (Figure 12). Plants typically grow in heavy red, doleritederived clay soil, either in open ground or among rocks in shallow soil, but we have also seen plants on stony tillite ground. L. oreogena flowers fairly early in the season before the plants are overshadowed by annuals and taller geophytes that reach maturity some weeks later. Although plants are invariably autogamous when grown in the greenhouse, they produce ample nectar and are regularly pollinated by long-proboscid flies of the genus Prosoeca. Presumably L. oreogena is outcrossed when pollinators are available, but in seasons when pollinators are scarce (or emerge late), self-pollination ensures seed production. This pattern of facultative outcrossing occurs in other nemestrinid-pollinated species of Lapeirousia, including L. jacquinii and L. violacea. Although Lapeirousia oreogena has a very restricted distribution, and on a world scale must be considered a rare species, its survival seems assured by its rocky habitat, unlikely ever to be placed under the plough, and also because parts of its range falls within the Nieuwoudtville Nature Reserve, where it is locally common.

58 50 S T R E L I T Z I A 35 (2015) Selected specimens SOUTH AFRICA. NORTHERN CAPE (Calvinia): Oorlogskloof, hills north of village, ( AC), 27 July 1962, Nordenstam 742 (MO, S); hill slopes, Nieuwoudtville, ( AC), Sept. 1899, Leipoldt 827 (BOL, K, SAM); Klipkoppies, Nieuwoudtville, ( AC), Lewis 5858 (SAM); Wildflower Reserve, Nieuwoudtville, ( AC), Oct (fr.), Snijman s.n. (MO, NBG); Farm Oorlogskloof south of Nieuwoudtville, ( AC), 10 Aug. 1983, Van Wyk 1404 (PRE); Matjesfontein hills, ( AD), 20 Aug. 1897, Schlechter (B, BM, BOL, COI, E, G, GRA, K, PH, PRE); Agterhantam, 31.5 km from Calvinia to Klipwerf, dolerite clay flats, ( BB), 1 Sept. 2006, Goldblatt & Porter (MO); Farm Kareeboomfontein, Calvinia, ( DA), 5 September 1974, Hanekom 2392 (PRE); southwestern slopes of Driefontein-se-Berg, red clay among dolerite rocks, ( DA), 25 Sept (fr.), Goldblatt, Manning & Porter (MO, NBG). 11. Lapeirousia montana Klatt in Abhandlungen der naturforschenden Geselschaft zu Halle 15: 359 (Ergänzungen: 25) (1882). Baker: 93 (1896). Goldblatt: 52 (1972). Type: South Africa, [Northern Cape], without precise locality, as Hantams Gebirge, 1869, Meyer 10 (B, holo.!). FIGURE 12. Distribution of Lapeirousia montana, ; L. oreogena,. Diagnosis and relationships: the acaulescent Lapeirousia oreogena is distinguished from its allies of similar habit by its large dark blue to violet flowers with slightly reflexed tepals and a perianth tube mm long. The tepals are almost black at the base with striking white, triangular to arrowhead-shaped marks. The centrally placed, purple anthers form a ring around the style and have dark purple pollen. Like some other acaulescent members of sect. Sophronia the leafy outer bracts extend well above the flower buds and have wavy to crisped margins. The earliest known collection of Lapeirousia oreogena was made by Rudolf Schlechter on a visit to the Bokkeveld Plateau in August Despite its distinctive appearance the species was not formally described until 1972 when Goldblatt revised Lapeirousia for the winter rainfall area of southern Africa. Schlechter s collection was widely distributed under the manuscript name L. oreogena, the name eventually adopted for the species. Plants acaulescent, branched at base, forming compact tufts at ground level, mm high including flowers, stem underground. Corm bell-shaped, 8 10 mm diam.; tunics light brown, basal rim crenate. Leaves 1 or 2, lowermost always longest, linear, mm 2 3 mm, straight or falcate, remaining foliage actually floral bracts, but often resembling leaves and identifiable as bracts only by position. Inflorescence of several to many flowers in congested head-like structure borne at ground level, internodes very short but evident on dissection or in fruit; bracts leaf-like, wide and pale below for mm, tapering to a green unifacial upper half, resembling leaf blade and mm long. Flowers actinomorphic, white to pale blue, then with lower third of tepals white, usually with dark blue triangular or diamond-shaped median mark near base, slightly scented of narcissus, lily or carnation; perianth tube cylindric, (25 )32 45( 55) mm long, with filaments decurrent as intrusive ridges dividing tube into 3 compartments and ± blocking tube in lower half; tepals subequal, lanceolate, (10 ) mm, spreading horizontally or deflexed up to 30 below horizontal. Stamens symmetrically disposed around style; filaments exserted 2 3 mm from tube; anthers erect, 3 4 mm long, ± white, pollen white to ivory. Ovary oblong, 4 5 mm long; style straight and erect, usually dividing opposite middle to just below anther tips, style branches divided for 1 / 2 their length, mm long. Capsules ± ovoid, 8 10 mm long, showing outline of seeds.

59 S T R E L I T Z I A 35 (2015) 51 Seeds globose, dark brown, ± 2 mm diam., with reticulate sculpturing, Chromosome number unknown. Flowering time: late August to late September. Figure 10B. Distribution and biology: a relatively narrow endemic of the western Karoo, Lapeirousia montana is centred in the southern Roggeveld of Northern Cape, where it is known from the Roggeveld Escarpment south and west of Sutherland, at sites in the Klein Roggeveld and locally close to the escarpment west of Middelpos (Figure 12). The type is said to be from the Hantam Mtns ( Hantams Gebirge ) where the collector, Heinrich Meyer, is known to have collected (Gunn & Codd 1981), but the species has not since been found there. While it would be useful to confirm the type locality we have no doubt Meyer s specimen represents the Roggeveld species. Plants grow in seasonally wet ground in relatively light clay or sandy soil in valley bottoms, around seasonal pools or in shallow soil over rocky pavement. Observations on its pollination indicate that L. montana is a generalist despite its long perianth tube. Nectar ascends well up into the tube and is accessible to settling moths, butterflies, bee flies (Bombyliidae) and bees (only Apis), all of which have been captured visiting the flowers and found to carry host pollen (Goldblatt et al. 1995). Floral scent is usually strong but unusually variable, sometimes reminiscent of narcissus, lily, carnation or even an unpleasant acrid odour. Diagnosis and relationships: the acaulescent Lapeirousia montana is allied to the widespread L. plicata, which it replaces in the southern Roggeveld and is distinguished from that species by the larger flower with longer perianth tube, mostly mm long, and tepals mm long, always with ± straight margins. The anthers are ± white and 3 4 mm long. L. plicata which has smaller flowers with a perianth tube mm long, tepals 7 9 mm with undulate margins and anthers usually blue anthers and mostly mm long. The upper leaves and floral bracts are typically crisped in L. plicata versus the straight margins of the upper leaves and bracts in L. montana. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Calvinia) 56 km south of Calvinia on the Blomfontein road to Middelpos, rock sheets, ( DD), Goldblatt 5811 (MO) (Williston): Hartebeesfontein, west of Middelpos, ( CC), 5 Sept. 1986, Oliver 8930 (NBG); entrance to Farm Onderplaas, north of Middelpos, ( CC), 16 Sept. 1999, Steiner 3379 (NBG) (Sutherland): top of Gannaga Pass on track to Watervlei, ( AA), 16 Aug. 2007, Helme 4695 (NBG); Quaggasfontein, Roggeveld, rock outcrop, ( AB), 23 Sept. 1981, Goldblatt 6352 (MO, NBG, PRE); Ouberg Pass, ( AD), Helme 4695 (NBG); Roggeveld Escarpment at Farm Blesfontein, ( AD), 24 Sept. 2011, Goldblatt & Manning (MO, NBG); Oliviersberg, Farm Tonteldoosfontein, ( DA), 23 Sept. 2007, Clark & Kelly 182 (NBG); Roggeveld, north of top of Komsberg Pass, 5,269 ft [1 606 m], ( DB), Goldblatt & Porter (MO, NBG); 12.5 miles [± 20 km] south-southwest of foot of Komsberg Pass, ( DC), Acocks (BOL, PRE); 57 km south of Sutherland, below Komsberg Pass, valley bottom, ( DC), 31 Aug. 1993, Goldblatt & Manning 9672 (MO, NBG); 1 Oct (fr.), Goldblatt & Manning 9672C (MO). 12. Lapeirousia kalahariensis Goldblatt & J.C.Manning, new rank and name for L. plicata subsp. longifolia Goldblatt in Contributions from the Bolus Herbarium 4: 50 (1972). Type: South Africa, [Northern Cape], Daniels Kuil, in sand, 22 Mar. 1939, Esterhuysen 1157 (BOL, holo.!). Plants ± acaulescent or shortly caulescent, mm high excluding basal leaves, stem usually reaching mm above ground, several-branched, branches crowded to form a compact, stalked rosette. Corm bell-shaped, brown, mm diam., rough-surfaced, basal rim lightly crenate. Foliage leaves ± 5, usually lowermost longest, up to 200 mm long, linear, lightly ribbed, margins of sheathing portion ± straight, remaining leaves shorter than basal, becoming ± bract-like. Inflorescence congested, head-like, many-flowered; outer bracts ovate at base, tapering to apex, channelled throughout, often hooked at tips, mm long, leaf-like distally, reaching or slightly exceeding flowers, margins undulate to slightly crisped; inner ± 1 / 2 as long, transparent with 2 green keels. Flowers actinomorphic, evidently uniformly white to cream,?probably sweetly scented; perianth tube (30 )33 40 mm long, slender, straight; tepals subequal, ± lanceolate, ± 3 mm, acute, held ± at right angles to tube. Stamens symmetrically disposed in a ring around style; filaments exserted ± 2 mm, arching outward after anthers split; anthers (3 )4 mm long, white. Style dividing opposite middle 1 / 3 of anthers, style branches ± 2.3 mm long, recurved, forked

60 52 S T R E L I T Z I A 35 (2015) for ± 1 / 2 their length (undivided in McLea s.n.). Capsules obovoid, ± 5 mm long, concealed in leaf bases, papillate in upper 1 / 3. Seeds probably globose (only immature seen), surface reticulate. Chromosome number unknown. Flowering time: March, probably also April. Distribution and biology: with a relatively wide range across the southern edge of the Kalahari, Lapeirousia kalahariensis extends from Danielskuil and Kuruman west of Kimberley in Northern Cape, South Africa, to south-central Namibia (Figure 13). Most of the available collections do not describe the habitat except in the broadest terms, but one collection indicates that the species was rare on flats of compacted calcareous sand. Nothing is known about its biology but the relatively long-tubed and white flowers are consistent with pollination by moths. Diagnosis and relationships: a member of the Lapeirousia plicata alliance (sect. Sophronia), L. kalahariensis was treated by Goldblatt (1972) as subsp. longifolia of L. plicata. We have examined more material and with a more thorough understanding of the genus, we are now convinced that this is a separate species, which we rename at species rank L. kalahariensis. Most significantly, the bracts are channelled throughout and not unifacial distally and, not noted previously, the stem is produced up to 40 mm above the ground. The flowers are recorded as being uniformly white, or in one collection yellowish, always lacking the blue or purple markings of L. plicata. The tepals, mm long, and tube, mm long, are somewhat longer than in L. plicata, which has tepals 7 10 mm long and a tube typically mm long, rarely up to 35 mm. Significantly, L. plicata occurs within the range of L. kalahariensis but typically blooms there later in the year, in May and June, sometimes in July. L. kalahariensis also shows a resemblance to L. odoratissima, which occurs across southern tropical Africa, but that species has much larger flowers, the perianth tube mm long, and a well-developed, strongly ribbed basal leaf. Lapeirousia kalahariensis was first collected in 1875 by J.H. McLea at the vague locality Transvaal, presumably north of the Vaal River near Kimberley. Little appears to be known about his travels (Gunn & Codd 1981), but we assume the collection was made while en route from Kimberley to Graaff Reinet where he lived in his later years. When first described, the species was included in L. plicata as subsp. longifolia. Representative specimens NAMIBIA (Nauchas): Rehoboth, Farm Namibgrens, ( CA), 14 Mar. 1953, Walter 1818 (B, WIND) FIGURE 13. Distribution of Lapeirousia barklyi, ; L. kalahariensis, ; L. odoratissima, in southern Africa.

61 S T R E L I T Z I A 35 (2015) 53 (Aus): Luderitz District, Farm Gamochas, on sand, flowers yellowish, ( BA), 2 Apr. 1950, Kinges 2500 (PRE); between Ausis and Khuias, sandy valley, ( BB), Mar. 1885, Schenck 1914 (M). SOUTH AFRICA. NORTH WEST (Vryburg): Vryburg, Armadillo Creek, ( DC), 27 Apr. 1912, Burtt-Davy s.n. (PRE 13714) (Kuruman): Batlharos, open veld, ( AC), Mar. 1921, E. Silk 248 (K, KMG). NORTHERN CAPE (Glenlyon): Hay District, Farm Bermolli, ( BD), Feb. 1922, Wilman s.n. (KMG 2636); Hay District, Black Ridge, ( DC), Mar. 1934, Wilman s.n. (GRA); 8 miles [12 km] west-northwest of west entrance to Farm Padkloof, ( CB), 5 Apr. 1960, Leistner 1748 (KMG) (Griekwastad): Danielskuil, ( BA), 25 Mar. 1920, Pole-Evans 77 (K); Asbestos Hills, Danielskuil, ( BA), 21 March 1939, Lewis 389 (SAM 53479). Without precise locality: Transvaal, 1875, McLea s.n. (BOL 5970, K). 13. Lapeirousia odoratissima Welw. ex Baker in Transactions of the Linnean Society of London, Botany ser. 2, 1: 273 (1878); Sölch: 10 (1969); Geerinck et al.: 344 (1972); Goldblatt: 480 (1990a). Type: Angola, [Huila], sandy woods near Lopollo, Welwitsch 1551 (BM, lecto.!, so annotated in unknown hand; B!, BM!, C!, G!, K!, iso.). L. stenoloba Vaupel: 548 (1912). Type: Namibia, Omaheke, m, in brown sand, 17 March 1911, Seiner 329 (B, lecto.!, designated by Sölch on sheet). L. congesta Rendle: 435 (1895); Baker: 354 (1897). Type: Tanzania, between Zanzibar and Uyui [near Tabora], in 1866, Taylor s.n. (BM, holo.!). L. juttae Dinter: 13 (1912). Type: not cited, unknown. Plants ( 250) mm high, with condensed aerial axis and then rosette-like, rarely stem partly aerial and up to 100 mm long, with expanded internodes, extending up to 80 mm below ground, rarely 5 10 mm long, simple or with several branches, each subtended by leaves or by leafy bracts. Corm bell-shaped, mm diam.; tunics woody, brown, outer layers fragmenting irregularly, rarely becoming fibrous by decay. Leaves few, often only 2 (hardly distinguishable from the bracts except by position), linear, strongly ridged, lowermost inserted below ground, exceeding bracts and up to twice as long, to mm. Inflorescence comprising 1 or more congested spikes, ± umbellate in appearance, flowers 3 6 per branch; bracts leafy, outer mm long, lanceolate, lightly ridged, inner ± 1 / 3 shorter than outer, ± membranous. Flowers actinomorphic, salver-shaped, white to ivory, usually strongly scented especially in evenings; perianth tube cylindric, mm long; tepals narrowly lanceolate to ± ovate, often attenuate, (25 )5 40( 50) 3 11 mm, extended horizontally or sometimes slightly drooping. Stamens symmetrically arranged in a ring around style; filaments ± 4 mm long, exserted for mm; anthers 6 8 mm long, linear, white; pollen yellow. Style usually ultimately reaching to ± apex of anthers, sometimes shorter, style branches 3 4 mm long, forked for ± 1 / 2 their length, diverging and recurved. Capsules obovoid-oblong, mm long, concealed in leaf and bract bases. Seeds ± globose, mm diam., with reticulo-rugose sculpturing. Chromosome number 2n = 16, B. Flowering time: December to March, occasionally in April. Figure 14. Distribution and biology: distributed widely across south tropical Africa, Lapeirousia odoratissima is apparently common only in Namibia where it occurs most often in sandy flats in the central and northern half of the country. Collections from a number of scattered sites indicate a fairly wide range across south tropical Africa except along the east coast and near interior. It extends from southwestern Angola across Zambia and adjacent Shaba Province of Congo to central and northern Malawi, Zimbabwe, and in locally dry sites in western and central Tanzania (Figure 13). It probably also occurs in Botswana, although to date there are no records from there. Largely a plant of semiarid habitats, L. odoratissima also occurs in Brachystegia woodland and even in exposed places in montane sites such as the Nyika Plateau in Malawi and the Inyanga Highlands in Zimbabwe, as well as open grassland and Acacia savanna. Little is known about its biology but the large, long-tubed, and usually intensely fragrant flowers are clearly adapted for pollination by hawk moths. Our observations on plants in Namibia indicate that flowers open towards sunset, then with creamy white tepals. By morning the perianth has turned ivory to buff and the tepals droop slightly below the horizontal and wilt by the end of the day. The flowers are most intensely fragrant when they first open and maintain their scent through the following day. Ample nectar is produced in the perianth tube: we measured up

62 54 S T R E L I T Z I A 35 (2015) FIGURE 14. Lapeirousia odoratissima, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

63 S T R E L I T Z I A 35 (2015) 55 to 25 µl in one flower and nectar sugar concentration ranged from 16 25% sucrose equivalents in our study population. Observations made by Jean Pawek in Malawi confirm our phenological studies: she recorded that buds open between 17:30 and 18:00 but have wilted at least by 10:00 the following day (earlier than we recorded in Namibia). Flowers lasting at most 24 hours are unusual in Crocoideae in which flowers usually last at least three days, even when pollinated. We only know of two comparable situations in the subfamily in the Namibian Gladiolus diluvialis Goldblatt & J.C.Manning, also evidently moth-pollinated (Manning et al. 2011), and in Savannosiphon euryphyllus (R.E. Gereau pers. comm. 2013). The corms of Lapeirousia odoratissima are reported to be edible and to comprise part of the diet of the!kung Bushmen of Namibia. Diagnosis and relationships: the tufted or rosette-like growth habit combined with the large white flowers with an exceedingly long perianth tube, mm long, set Lapeirousia odoratissima well apart in the genus. The other tufted to rosette-forming species of sect. Sophronia are all southern African and have smaller flowers, coloured pale blue to violet, or white to pale yellow. The chromosome number, 2n = 16 (x =8) in L. odoratissima and its reticulo-rugose seeds correspond with the southern African species of sect. Sophronia. L. odoratissima and L. caudata are the only tropical African species of the genus. The interior southern African L. kalahariensis, which has white flowers and a perianth tube mm long, appears to provide a link between L. odoratissima and the smaller southern African members of sect. Sophronia. These seems no such link between the caulescent L. caudata, L. littoralis and L. odoratissima, although both L. caudata and L. littoralis have the same basic chromosome number of x = 8 and reticulate seed sculpturing, unlike other species of the genus, excepting of course sect. Sophronia. History: Lapeirousia odoratissima was discovered by Friedrich Welwitsch in southern Angola in His collection was distributed widely under the manuscript name Psilosiphon odoratissima and formed the basis for J.G. Baker s (1878) L. odoratissima. A second early collection made by the Rev. W.E. Taylor in central Tanzania in 1886, was described as L. congesta by A.B. Rendle, who distinguished it from L. odoratissima on the basis of a more congested habit. Rendle described the plant as having a stem forming a dense sessile head above the first leaf. This is not strictly true although the internodes of the stem and inflorescence axis are shorter than in the type specimens of L. odoratissima, which have among the longest internodes found in the species. From the ample material now available it seems clear that the variation between the extremes represented by the two collections is continuous and there is no reason to consider them separate species. Later collections from the then German South West Africa, made by Kurt Dinter and Franz Seiner, were described as Lapeirousia stenoloba by Vaupel (1912). He considered that these Namibian populations constituted a species related to but distinct from L. odoratissima, differing by a robust habit and particularly narrow and attenuate tepal apices. Many but not all Namibian specimens of L. odoratissima differ from those from tropical Africa in their branched stems and numerous flowers but intra-populational variation is considerable in this species and there are often few- or even unbranched plants in populations of more robust ones with several branches, thus the difference does not seem taxonomically significant. Representative specimens NAMIBIA. Kaokoveld (Ohopoho): Okakura, ( DA), 28 Feb. 1913, Dinter 3322 (SAM) (Kamanjab): north of Otjovasandu in red sand, ( BA), 11 Mar. 1976, Geiss & Loutit (WIND); Outjo. Etosha National Park, Kaross, ( B), 9 Apr (fr.), Volk & Le Roux 807 (WIND); between Kaross and Kamanyab, ( BC), Thorne s.n. (SAM ) (Okaukuejo) 10 km west of Farm Uitzig, ca. 60 km east-northeast of Otjiwarongo, grassveld on red sandy loam, ( CB), 9 Feb. 1983, Lavranos & Pehlemann (MO, WIND). Ovamboland (Ondangua): Ondonga, ( DD), Barnard 196 (SAM) (Okahakana): Onolongo Onambeke, ( BB), Apr. 1923, Barnard 136 (SAM). Grootfontein (Gobaub): Farm Norabis 387, thornveld, ( DD), 18 Mar (fr.), Goldblatt & Manning 8824 (MO) (Tsumeb): Auros (Otavi), ( DA), 10 Feb. 1925, Dinter 5599 (B, G, GH, PRE, SAM, Z) (Grootfontein): 30 miles [± 42 km] north of Gautscha Pan, ( CA), 10 Feb. 1958, Story 6460 (PRE); Farm Oliewenhof, sandy flats, ( CB), 8 Mar (fr.), Merxmüller & Giess (M, WIND). Otjiwarongo (Waterberg): Otjiwarongo, Farm Okosongomingo, ( CA), 4 Mar. 1974, Merxmüller & Giess (M, PRE, WAG, WIND). Okahandja. 2116

64 56 S T R E L I T Z I A 35 (2015) (Okahandja): Omatako View, red sand, ( BA), 4 Mar. 1974, Woortman 256 (WIND) (Otjosondu): Okahandja District, Farm Hochveld, brown sandy loam, ( BD), 30 Apr (fr.), Giess et al (WIND); Okahandja, ( DD), 22 Feb. 1928, Bradfield 385 (PRE). Windhoek (Otjosondu): 35 km from Steinhausen to Windhoek on Kapps Farm road, ( DD), 15 Mar. 1988, Goldblatt & Manning 8807 (MO) (Windhoek): Farm Bodenhausen, near river, ( BC), 7 Mar. 1959, Seydel 1771 (M, WIND). Gobabis (Tsumkwe) Grootfontein distr., ± 3 miles ([4.6 km] south of Nama Pan, ( DC), Story 6275 (M, PRE, WIND) (Kaukauveld): Kaukauveld, 56 miles north of Eiseb Omuramba towards Kano Vlei, ( AD), 13 Apr (fr.), Giess 9815 (WIND) (Steinhausen): 15 km from Steinhausen to Windhoek on Kapps Farm road, ( CC), 15 Mar (fr.), Goldblatt & Manning 8803 (MO). [Exsiccatae for Lapeirousia odoratissima from tropical Africa are cited in Goldblatt (1990a)]. C. Sect. Lapeirousia (Spp ) Corm obconic, ± triangular in outline; tunics with basal rim produced into teeth or long, usually spreading. Stem mostly aerial (sometimes ± subterranean in L. arenicola) and often branched, stems compressed with angles often crisped. Inflorescence a spike. Flowers bilaterally symmetric, lower tepals with nectar guides of contrasting colour; perianth tube ± cylindric or funnel-shaped, short to elongate; stamens unilateral. Capsules without auriculate lobes or locular ridges. Seeds smooth (scarcely wrinkled) with surface cells domed. Basic chromosome number x = Lapeirousia barklyi Baker, Handbook of the Irideae 171 (1892); Goldblatt: 68 (1972) [including L. macrospatha Baker]. Type: South Africa, Namaqualand, without precise locality, Sept. 1876, Barkly s.n. (K, lecto.!, designated here [not holo. as indicated by Goldblatt, 1972]). Plants mm high, usually branched, sometimes repeatedly, mainly from base; stem compressed and 2-angled to 2-winged, wings often lightly serrated, with cormlet at each of 2 underground nodes. Corm bell-shaped, usually mm below ground, ± 8 mm diam.; tunics brown, fairly smooth, basal rim lightly serrated, teeth directed downward. Leaves 4 6 (or more), lowermost 1 or 2 longest and slightly to about twice as long as stem, linear, ascending to ± falcate, mm wide, corrugately ribbed, upper leaves shorter than stem. Inflorescences 2 5-flowered spikes; outer bracts green, veins translucent, lanceolate and acute or acuminate, (10 )20 25 mm long, folded and angular along upper midline, often keeled near apex, margins broadly hyaline; inner bracts ± 1 / 2 as long as outer, transparent with 2 green keels, forked and turning brown at apex. Flowers zygomorphic, light purple-pink to lilac, lower tepals with basal ridge or claw-like tooth near base, marked with median yellow streak edged distally in dark red to purple, odourless; perianth tube obliquely funnel-shaped, (15 )20 25 mm long (often shorter in dry years, or apparently shorter in herbarium material), narrowly cylindric below, abruptly expanded in upper 4 5 mm into wide cylindrical throat; tepals subequal, lanceolate, acute, upper 3 slightly larger, ± 6 7 mm, lower 3 united for 2 3 mm and thus slightly shorter, mm, all weakly differentiated below into claws ± 8 mm long, claws forming wide cup, limbs spreading at right-angles to claws, when fully open held ± in one plane. Stamens unilateral, arcuate; filaments ± 10 mm long, exserted ± 5 mm from tube but included in cup formed by tepal claws; anthers 5 6 mm long, included in floral cup, shortly apiculate, usually ± white; pollen white. Style arching over stamens, dividing close to or 1 2 mm beyond anther tips; branches ± 5 mm long, divided for 2 / 3 their length, spreading beyond anthers. Capsules oblong to ovoid, 8 10 mm long. Seeds mm diam., round, smooth. Chromosome number 2n = 16. Flowering time: usually mid-september to early October, occasionally in late August. Distribution and biology: a species of the coastal Namib, Lapeirousia barklyi extends along the Atlantic coastal plain and adjacent coastal mountains from near Lüderitz Bay in the north to Lekkersing and Port Nolloth in the south (Figure 13). Where recorded, the soil type is sand. Although first collected in 1876, relatively early for a northern Namaqualand species, and described in 1892 by J.G. Baker, Lapeirousia barklyi remained poorly known until the later 20th century. The species was treated as including L. macrospatha by Goldblatt (1972), an error only realised later when the development of better roads

65 S T R E L I T Z I A 35 (2015) 57 made travel in the Richtersveld easier and living plants could be compared side-by-side. Lapeirousia barklyi was first recorded from Namibia in 1922 by the German botanist and collector, Kurt Dinter, but the widespread occurrence of the species across the southwestern part of the country only became evident after a series of collections were made after the mid-1970s. It is now clear that the larger part of the distribution of L. barklyi is in southwestern Namibia and that in South Africa it only occurs in the western Richtersveld. Diagnosis and relationships: the purple to violet perianth and relatively short-tubed flower combined with a broadly conical corm with well-developed basal teeth serve to distinguish Lapeirousia barklyi. The tube mm long is normally shorter to about as long as the rest of the flower. This contrasts with L. macrospatha, included in L. barklyi by Goldblatt (1972), which has a cream to beige perianth flushed pink on the reverse, and a tube mm long and half as long again as the rest of the flower. Goldblatt (1972) suggested that Lapeirousia barklyi was probably most closely related to white- to pink- (or pale yellow-) flowered L. fabricii, a species of Western Cape and southern to central Namaqualand. The latter has a similar corm with marginal teeth but differs notably in its longer-tubed flower. The two can also easily be distinguished by their bracts, which have an almost plane, smooth margin and keel in L. barklyi, but are strongly crisped and serrate in L. fabricii. Representative specimens NAMIBIA (Lüderitz): Lüderitz District, southern Namib, ( BC), 22 Sept. 1999, Mannheimer & Mannheimer 269 (WIND) (Bogenfels): Pomona, ( AB), 1922, Dinter s.n. (Z); Sept. 1925, Schaefer s.n. sub Marloth 6596 (PRE); Klinghardt Mtns, northern slope of quartzite hill ( BC/BD), 18 Aug. 1977, Merxmüller & Giess (BR, M, PRE, WIND); Gemsbokstal, Middagskupper, grassy flats south of Bankenberge, ( BD), 17 Aug (BD), Van Berkel 570 (MO, NBG, PRE) (Witputs): 14 km from Rosh Pinah to Obib Mtn near Spitskop road junction, ( DC), 1 September 1989, Van Wyk 8909 (PRE) (Oranjemund): 16 km southwest of Obib-Wasser on road to Orange River (Daberas), ( BA), 17 Sept. 1973, Giess (K, M, NBG, PRE, WAG, WIND); red sand flats, road to Obib, Sperrgebiet I, ( BA), 1 Sept. 1963, Merxmüller & Giess 3413 (BR, M, PRE, WIND); Obib, ( BB), Merxmüller & Giess 3413 (BR, M, PRE, WIND). SOUTH AFRICA. NORTHERN CAPE (Oranjemund): gravel flats near Arrisdrif [Arriesdrif], 70 m, ( BC), 21 Aug. 1925, Marloth (BOL, PRE); Daberas, sandveld, ( BD), 4 Sept. 1962, Nordenstam 1264 (MO, NBG, S); Beauvallon, gravel, ( DA), 2 Sept. 1980, Venter 8201 (PRE); ± 5 km east of Alexander Bay, ( DA), 17 Aug. 1986, Williamson 3554 (MO, NBG); Grootderm, ( DA), 12 Sept. 1973, Lavranos (MO, PRE); 37 km north of Lekkersing, ( DB), 9 October 1991, Dreyer 182 (PRE) (Vioolsdrif): sandy flats between Jasper s Werf and Doornpoort, 250 m, ( AC), 26 Aug. 1925, Marloth (PRE, STE) (Springbok): Lekkersing, ( AA), 24 Aug. 1925, Marloth (PRE, STE); Anenous plains ± 8 km west of Farm Grasvlakte, in deep red sand, ( AB), 15 Sept. 1992, Goldblatt & Manning 9460 (MO, NBG, PRE, WAG); near Abbevlakte, ( AD), Aug. 1883, H. Bolus 6576 (BOL). 15. Lapeirousia littoralis Baker in Transactions of the Linnean Society, London, Botany ser. 2, 1: 272 (1878). Type: Angola, [Namibe], sandy coastal hills at Praia da Amelia near Villa de Mossamedes, July 1859, Welwitsch 1546 (BM, holo.! see discussion of the type specimen). Lapeirousia burchellii Baker: 171 (1892). Lapeirousia caudata subsp. burchellii (Baker) Marais & Goldblatt: 30 (1972). Type: South Africa, [?North West], Morokweng, Chooi Desert, ( AC), Oct. 1812, Burchell 2350 (K, lecto!. designated by Goldblatt (1972: 30); G!, P!, iso.). Lapeirousia ramosissima Dinter: 255 (1931). Type: Namibia, Lüderitz: dunes near Gründorn, Dinter 5043 (B, holo.!; BOL!, BR!, G!, K!, PRE!, SAM!, Z!, iso.). Lapeirousia streyii Suess.: 88 (1951). Type: Namibia, Lüderitz: dunes south of the Kuiseb, Strey 2587 (M, holo.!; PRE, iso.! [not holo. as cited by Goldblatt, 1972]). Plants ( 200) mm high, several- to many-branched, branches short, clustered near base, stem ± rounded to weakly angled below nodes. Corm bell-shaped, mm diam.; tunics woody, brown, outer layers breaking irregularly, rarely becoming ± fibrous, basal rim shortly denticulate. Leaves few to several, linear, 2 4 mm wide, slightly corrugately ribbed, lowermost longest, inserted at or just below ground, often as long as inflorescence, rarely slightly longer, ascending to falcate, upper leaves progressively shorter. Inflorescence 1 or more short, 4 8-flowered spikes; outer bracts green, succulent, weakly keeled, (12 ) 15 20( 25) mm long; inner 2 / 3 to 1 / 2 as long, ± membranous with 2 green keels, forked at apex. Flowers zygomorphic, white to creamy white,

66 58 S T R E L I T Z I A 35 (2015) scented or not; perianth tube slender below, curving outwards and expanded above, mm long, slightly wider in upper ± 4 mm; tepals subequal, narrowly lanceolate, mm, acute, spreading. Stamens unilateral, arcuate; filaments 7 8 mm long, exserted 2 3 mm; anthers parallel, 4 5 mm long, ± white; pollen white or yellow. Style arching adaxially to stamens, branching between base and middle of anthers, style branches divided in distal 1 / 3 2 / 3, recurved above and usually tangled in anthers. Capsules globose, 8 12 mm long. Seeds globose, mm diam., dark brown, with reticulate sculpturing. Chromosome number 2n = 16. Flowering time: mainly July to October in southern Africa but also at other times, March and April in northern Namibia. Distribution and biology: relatively widespread, Lapeirousia littoralis extends from southwestern Angola and western, largely coastal Namibia to Bushmanland and Gordonia in Northern Cape, South Africa, as far east as Kimberley. In Namibia it extends from near the Brandberg in the central west to the Lüderitz District in the south. There is also a record from southern Botswana adjacent to Northern Cape (Figure 15). L. littoralis blooms early in the wet season, usually not long after the first soaking rains have fallen. Thus the Northern Cape and southern Namibian populations usually flower in the spring months (July to October) following the winter rainfall of the southwestern African coast. Elsewhere populations generally bloom in late summer. The species favours sandy, well-drained soils, where we have seen it on red sandy ground. The style branches are tangled with the anthers, suggesting that self-pollination may occur in the absence of pollinator visits. Diagnosis and relationships: characterised by white to creamy white, uniformly coloured flowers with relatively short, straight tepals, mm long, and a long perianth tube, mm long, expanded and curving outward distally, Lapeirousia littoralis can usually be recognised by its flowers alone. It is sometimes confused with L. caudata but this species, from northern Namibia, Zambia, Zimbabwe and Mozambique, has long, relatively lax spikes usually mm high, with short bracts, 10 18( 23) mm long. The flowers are creamy-yellow to dull maroon, colours not recorded in L. littoralis, and have long, narrow, attenuate, drooping tepals (18 )25 30 mm long and ± mm wide. As the flowers fade and dry the tepals appear virtually filiform. Plants from arid southwestern Angola, western and southern Namibia, and the springflowering populations from Northern Cape in South Africa tend to be shorter, often with the FIGURE 15. Distribution of Lapeirousia caudata, ; L. littoralis, in southern Africa.

67 S T R E L I T Z I A 35 (2015) 59 branches crowded at the base. In these plants the spikes are shorter, usually mm high, and sometimes congested, and the bracts are usually 15 20( 25) mm long. The flowers in these populations have lanceolate, acute tepals ± 20 mm long and mm wide. With a diploid chromosome number of 2n = 16, Lapeirousia littoralis has the dimorphic karyotype of one very long and seven short chromosome pairs typical of Lapeirousia: the base number and karyotype corresponds to that in sects. Lapeirousia and Sophronia. Molecular phylogenetic studies link the species with the western southern African L. barklyi and then to L. caudata. Vegetatively the two species are very alike, but the flowers differ sharply, L. barklyi having large pink flowers with yellow nectar guides, indicating a different pollination system. The seeds of L. littoralis and L. caudata are somewhat unusual, having a sculptured, ± reticulate surface, otherwise known only in sect. Sophronia. Seeds of L. barklyi have a smooth surface characteristic of other species of the genus. The presence of sculptured seeds in L. littoralis and L. caudata suggests a possible link with sect. Sophronia but neither molecular data nor any other morphological features point to a connection with that section. History: the type collection of Lapeirousia littoralis was made by Friedrich Welwitsch in fruit in July 1859, and was described by J.G. Baker in 1878, based on this collection alone. The type locality is near Mossamedes on gravelly hills around Praia da Amelia, in southwestern Angola, now the province of Namibe. The type material is in poor condition and consists of depauperate plants with short branches produced from near the ground, bracts ± 12 mm long, capsules ± 9 mm long, and linear, lightly corrugate leaves. The specimen matches a second collection from southwestern Angola, Torre 8824, which was first assigned to L. caudata, but the flowers correspond to the South African and southern Namibian L. burchellii, which mostly flowers in the spring months. L. burchellii was first recorded in October 1812 when the English botanist and explorer William Burchell reached the Kuruman area of Northern Cape and North West. Burchell s collection was only formally described by J.G. Baker in Goldblatt (1972) considered the taxon to be subsp. burchellii of L. caudata, the name used until now for L. littoralis. L. ramosissima, collected near Grundorn in southern Namibia by Dinter and described by him, matches the type collections of L. burchellii and L. littoralis. A third species, L. streyii, from west-central Namibia is close to being intermediate between L. littoralis and L. caudata from tropical Africa, but has the shorter, slightly broader tepals characteristic of L. littoralis. Selected specimens ANGOLA. Near Mossamedes close to Vila Arriaga, 7 Feb. 1956, Torre 8824 (LISC). BOTSWANA (Mafekeng): Moropedi ranch, ( CA), 15 Oct.1978, Hansen 3455 (C, K). NAMIBIA (Uis): Brandberg, B. W. Mine Uis Mine road, sand in rocky ground, ( AA), 14 Mar. 1963, Kers 942 (WIND); Messumberge, foot of mountains, ( AC), 19 Mar. 1967, Giess 9689 (M, PRE, WIND); Nabab Hill, Brandberg area, ( AB), 4 Mar. 1978, Craven 715a, 715b (WIND) (Trekkopje): 1 mile [1.5 km] northwest of Farm Bloemhof (AD), Giess et al (M, WIND) (Rostock): Namib Park, Tumasberg at campsite, ( BA), Giess (M, PRE, WIND) (Helmeringhausen): Farm Duwisib, ( BC), 17 May 1956, Volk (M, MO) (Aus): dunes 15 miles [ 23 km] west of Aus, ( CB), Giess & Van Vuuren 828 (WIND); 13 miles [± 20 km] west of Aus, coarse sand, Giess & Van Vuuren 824 (M, WIND). SOUTH AFRICA. NORTH WEST (Morokweng): Chooi Desert, Giraffe Station, Oct. 1812, ( AD), Burchell 2341 (K). NORTHERN CAPE (Mata-Mata): Kalahari Gemsbok Park, between the Nossob and Auob dunes, ( BC), 25 Apr. 1960, Barnard 807 (PRE); Kalahari Gemsbok Park, 6 miles [± 9 km], southeast of Kafferpan, loose red sand on dune top, ( CC), 22 Apr. 1960, Leistner 1874 (M, MO, PRE) (Vioolsdrif): Richtersveld National Park, Ghaapkop near Sandberg, ( AA), 7 July 1993, Van Jaarsveld 13246J (PRE); Namaqualand, 15 km south of Vioolsdrif, Koubank River, ( DA), 7 July 1976, Giess (M, WIND) (Warmbad): Goodhouse, ( CD), 27 July 1950, Barker 6263 (NBG), Lewis 2272 (SAM) (Kimberley): 18 miles south-southwest of Schmidtsdrift, red sand, ( CA), 24 Sept. 1961, Leistner 2866 (B, G, PRE) (Springbok): 25 miles [± 37 km] north of Okiep on Goodhouse road (old road through Leospoort), ( BD), 25 Aug. 1959, Lewis 5517 (NBG); Tuinvlak, Goegab (Hester Malan) Nature Reserve, ( DB), 9 Oct. 1974, Rosch & le Roux 953 (PRE) (Gamoep): road to Goodhouse ( ), sandy soil ( AA), 19 Sept. 1980, Van Berkel 255 (MO), NBG); Little Bushmanland, Keuzabies, ( AB), 18 June 1973, M. Schlechter s.n. or 103 (B, BOL, COI, E, G, MO, P, PRE); Aggenys, Brabees, red sand dunes and plains, ( BB), 15 Aug. 1990, Dean 1002 (PRE); ± 100 km west of Pofadder to Springbok, ( BC), 13 Sept. 1973, Coetzee & Werger 1752 (PRE);

68 60 S T R E L I T Z I A 35 (2015) Aggenys, 10 miles [± 15 km] west of farmhouse, ( BD, 4 Sept. 1971, Wisura 2222 (NBG); near Ratelkraal, ( CA), 7 Sept. 1950, Barker 6755 (BOL, NBG). 16. Lapeirousia caudata Schinz in Verhandlungen des botanische Vereins der Provinz Brandenburg 31: 213 (1890). L. littoralis subsp. caudata (Schinz) Goldblatt: 478 (1990a). Type: Namibia, Amboland, Olukonda, Rautanen 2 (Z, lecto.!, designated by Sölch on sheet; Z, iso.!). Lapeirousia delagoensis Baker: 171 (1892); Baker: 94 (1896). Type: Mozambique, Delagoa Bay, Lourenço Marques [Maputo], sandy places, H. Bolus 7618 (K, lecto.!, designated by Goldblatt (1990a: 478); BOL!, G!, SAM!, iso.). Lapeirousia lacinulata Vaupel: 546 (1912). Type: Zambia, Kantanina Hills, Kässner 2170 (B, holo.!; BM!, BR!, E!, HBG!, K!, P!, Z!, iso.). Plants mm high, usually with several (rarely up to 12) long branches, produced from near base, stem ± rounded to weakly angled below nodes. Corm bell-shaped, mm diam.; tunics woody, brown, outer layers breaking irregularly, rarely becoming ± fibrous, basal rim shortly denticulate. Leaves few to several, linear, mm wide, lightly corrugately ribbed, lowermost longest, inserted ± at ground level, usually slightly longer than stems, ascending to falcate or trailing, upper leaves progressively shorter. Inflorescence comprising (1 )several, relatively lax, 8 12-flowered spikes; outer bract green, succulent, weakly keeled, 10 18( 23) mm long; inner 2 / 3 to 1 / 2 as long, ± membranous with 2 green keels, forked at apex. Flowers zygomorphic, greenish yellow, light yellow-brown or dull maroon, with strong sweet scent; perianth tube slender below, curving outwards and expanded above, (8 )30 45( 70) mm long, wider upper part ± 6 mm long; tepals subequal, narrowly linear-filiform, (18 ) mm, attenuate, spreading at right-angles to tube, dorsal ± erect, upper laterals arching backward, lower 3 arching toward ground. Stamens unilateral, arcuate; filaments 7 8 mm long, exserted 2 3 mm; anthers parallel, 4 5 mm long, ± white; pollen white or yellow. Style arching adaxial to stamens, branching between base and middle of anthers, style branches divided in distal 1 / 3 2 / 3, recurved above and usually tangled in anthers. Capsules globose, 6 8( 11) mm long. Seeds globose, mm diam., dark brown, with reticulate sculpturing. Chromosome number 2n = 16. Flowering time: mainly January to April (almost any month in Mozambique). Distribution and biology: occurring in a wide swath across south tropical Africa and northern southern Africa, Lapeirousia caudata extends from northern Namibia and Zambia, across Botswana to Zimbabwe and in South Africa to western Limpopo and Northern Cape, and with isolated and surprising populations in southern Mozambique (Figure 15). It blooms late in the wet season, usually after January and sometimes as late as April. In southern Mozambique L. littoralis appears to bloom at almost any month, reflecting the non-seasonal rainfall pattern of the southeast coast. The species favours sandy, well-drained soils, sometimes growing on sand dunes. The likely pollinators are moths, in light of the pale coloured, long-tubed flowers and often clove-like fragrance. The style branches are tangled with the anthers, suggesting that selfpollination may occur in the absence of insectmediated pollen transfer. Diagnosis and relationships: characterised by uniformly coloured, pale dull yellow, yellowbrown or maroon flowers, usually with a long perianth tube expanded and curving outward distally, Lapeirousia caudata can usually be recognised by its flowers alone. The relatively long tepals, mm long, are up to 2 mm wide, and as the flowers fade and dry the tepals shrink, then appearing filiform and less than 1 mm wide. The habit is also distinctive: well grown plants have several (up to 12) long branches mm high, produced from near the base, and relatively lax spikes with short bracts, 10 18( 23) mm long. In plants we have seen alive the tepals spread outward from the base in somewhat lax fashion with the dorsal tepal ± erect and the lower three arching toward the ground. Plants from southern Mozambique, mostly from immediately around Maputo, stand out in having particularly long-tubed flowers and in often being very robust. The perianth tube is sometimes up to 70 mm long, compared with the mm usual in central African plants. Floral variation is notable in plants from western Zambia, and a collection from Mongu (Robinson 6749) has flowers with a perianth tube 8 13 mm

69 S T R E L I T Z I A 35 (2015) 61 long and tepals ± 18 mm long. Such occasional variation, particularly in tube length, also occurs in other species of Lapeirousia (see L. anceps) and appears to have no taxonomic significance. Lapeirousia caudata was treated by Goldblatt (1990a) as subsp. caudata of L. littoralis as collections then available suggested the existence of plants intermediate between the two species, especially for bract and perianth tube length. We have now seen populations of both taxa alive and were impressed by differences between them in overall habit and tepal dimensions and orientation. It seems more useful to recognise two separate, but no doubt closely allied, species. The overlap in perianth tube and bract length in the two species most likely varies independently in each and does not indicate the existence of truly intermediate plants. We include both species in sect. Lapeirousia, with which corm morphology and chromosome number accord. L. caudata and L. littoralis are, however, discordant in the section in having seeds with reticulate to rugose surface sculpturing, a feature otherwise found in Lapeirousia only in sect. Sophronia. History: first gathered in December 1885 at the Finnish Mission Station at Olukonda in northern Namibia by Hans Schinz, Lapeirousia caudata was described in 1890 by Schinz based on his and the Finnish missionary, Martti Rautanen s ample, but confusingly labelled, collections at the Zurich Herbarium. A sheet collected by Rautanen in 1887 was designated the lectotype by Sölch in 1959 and this choice appears to be as suitable as any other. Corresponding closely with L. caudata, although from northern Zambia, plants collected by T. Kässner in 1906 were described by Vaupel in 1912 as L. lacinulata on the basis of its slender habit, low stature, and shorter perianth tube, only 25 mm long (actually mm), in contrast to the type of L. caudata in which the perianth tube is ( 70) mm long. The distinction now seems minor and insufficient reason for their separation at any taxonomic rank. Several other collections of L. caudata have a short perianth tube similar to that in L. lacinulata, but not the slender habit. The first of the series of isolated populations near Maputo, Mozambique, were discovered by the Cape Town botanist Harry Bolus in 1886 and his collection was made the type of his new species L. delagoensis by J.G. Baker (1892). Selected specimens BOTSWANA (Siambiso): sandy floodplain of Kwando River, ( AB), Smith 2222 (K, PRE, SRGH) (Ukwi): 120 km west-northwest of Hukuntsi on track to Ncojane, open sandy savanna, ( DC), 13 Mar. 1979, Skarpe S-333 (K, PRE, SRGH, UCBG) (Kang): Kang, 320 km west of Gaborone, 3500 ft [1 067 m], ( DD), 20 Sept. 1975, Mott 772 (SRGH, UCBG). NAMIBIA (Ondangua): Ondangua, open sandy flats, ( DD), 5 Feb. 1959, De Winter & Giess 6877 (M, WIND) (Enana): Oniipa, eroded field, ( CC), 7 Mar. 1967, Soini 429 (WIND); Olukonda, ( CC), Dec. 1885, Schinz 15 or s.n. (B, Z), 30 Jan. 1887, Rautanen 2 (Z), Feb. 1894, Rautanen 170 (B, G, Z) (Runtu): 10 km east of Runtu, ( DD), Jan. 1956, Kruger 1 (PRE) (Katima Mulilo): Mpilila Island, locally common near marsh, 12 Jan. 1959, Killick & Leistner 3325 (PRE, SRGH, WIND) (Okahakana): Onambeke Onolongo, (?BB), Apr. 1923, Barnard 141 (SAM), Barnard 573 (SAM) (Tsumkwe): 15 km east of Tsumkwe on road to Botswana, ( DA), 10 Jan 1971, Geiss et al (WIND); Tsumkwe, hard black sand, Oberflachenkalk, ( DA), 13 Jan. 1971, Giess et al (WIND); Simkue, 157 miles [± 250 km] east of Grootfontein, Acacia-Combretum parkland, ( DB), 17 Jan. 1958, Story 6162 (M, PRE, SRGH, WIND) (Rehoboth): Farm Arovley, red sand, ( AA), 14 Feb. 1965, Giess 8401B (M, WIND) (Gibeon): sandbodem, Sudkalahari bis Gibeon, ( BA, BB), May 1912, Range 1455 (SAM). SOUTH AFRICA. LIMPOPO (Nylstroom): Waterberg, Zandrivierspoort near Alma, Farm Witpoort, ( AC), 5 Mar. 1979, Van Wyk 2680 (M, PRE). NORTHERN CAPE (Unions End): Kalahari Park, Khaapan, ( BD), Jan. 1978, Van der Walt (PRE) (Mata-Mata): Kalahari Gemsbok Park, 2 km north of Kameelsleep, ( DC), 26 Feb. 1962, Leistner & Werger 3498 (PRE) (Tsabong): Kuruman District, Vanzylsrust, Farm Floradale, red sand, dune woodland, ( CD), 30 Mar. 1988, Venter (PRE) (Kuruman): Farm Uitkoms, ( CA), Gubb 146/38 (PRE) (Upington): ± 40 km north of Upington at southern edge of Kalahari, ( AB), 1 Apr. 1980, Snijman 235 (NBG) (Glen Lyon): sandveld west of Padkloof, on dune running parallel to Langebergen, ( DA), 15 Mar. 1937, Acocks 2057 (PRE); 90 km west of Griquatown, ( CD), 6 Mar. 2010, Goldblatt & Porter (MO, NBG). 17. Lapeirousia macrospatha Baker, Flora capensis 6: 94 (1896); Goldblatt & Manning: 345 (1994). Type: South Africa, [Northern Cape], between Port Nolloth and Eleven Mile Station, 22 Aug. 1883, H. Bolus 697 (K, holo.!; B!, BOL!, G!, SAM!, iso.). [macrospatha Latin, large spathes but alluding to the prominent floral bracts]

70 62 S T R E L I T Z I A 35 (2015) Plants mm high including flowers, usually branched, then from base, stem compressed and lightly 2-angled, angles smooth or lightly serrated. Corm ± 7 8 mm diam., ± bellshaped, usually mm below ground; tunics dark brown, fairly smooth, basal margin lightly serrated with teeth directed downward. Leaves 3 6 (or more), lowermost usually conspicuously longer than others, ± twice as long as stem, linear, ascending to falcate, mm wide, strongly ribbed, upper cauline leaves shorter, sometimes resembling bracts and not ribbed, usually subtending branches. Inflorescences 2 4-flowered spikes; outer bracts green, firm, mm long, lanceolate, acute, slightly keeled, margins hyaline, those of lower flowers unifacial and ribbed distally; inner bracts forked at tips, transparent in centre, ± 1 / 2 to 2 / 3 as long as outer. Flowers zygomorphic, cream to ivory, fading to dull pink, pink on outside of tepals and tube, lower tepals each marked with a small yellowish basal mark surrounded by a dark purple zone, usually with a small claw-like cusp at base of lowermost or all 3 lower tepals; perianth tube mm long, widening and lightly curving outward near apex; tepals unequal, lanceolate and acute, dorsal largest, ± 20 7 mm, initially erect or arching forward over stamens later reflexed, upper lateral tepals slightly reflexed, lower 3 tepals united for ± 2 mm, ± 13 5 mm, held at right angles to tube, tilted at ± 30 below horizontal. Stamens unilateral, arcuate; filaments ± 9 mm long, reaching mouth of tube or exserted up to 2 mm; anthers contiguous, 3 5 mm long, mauve, pollen pale violet. Style dividing opposite middle of anthers, style branches ± 3 mm long, divided for ± 2 / 3 their length. Capsules oblong, ± 7 mm, apices of locules forming auriculate lobes decurrent on weakly developed locular ridges. Seeds globose, ± flattened or lobed at chalazal end, ± 1.6 mm diam., red-brown. Chromosome number unknown. Flowering time: August to late September. Figure 16 Distribution and biology: restricted to northern Namaqualand, Northern Cape, Lapeirousia macrospatha has a fairly localised distribution in the southern Richtersveld. It has been collected repeatedly on the Anenous plains between Port Nolloth and Anenous Pass, but occurs to the north near Eksteenfontein, near Steinkopf, and in the sandy country east and north of Port Nolloth. It also extends southward at least as far as Grootmis (Figure 17). L. macrospatha appears to occur mainly in deep, fine-grained red sand. Sometimes, possibly due to heavy overgrazing, it may be extremely common locally and even dominant, but more often it grows in association with sandveld annuals such as Cotula (Asteraceae), Manulea, Sutera and Zaluzianskya (Scrophulariaceae), Heliophila (Brassicaceae) and low succulent shrubs (Aizoaceae). Diagnosis and relationships: Lapeirousia macrospatha can be recognised by its large flower with a long, slender perianth tube mm long, widening noticeably towards the apex and white to beige tepals red on the outside and with red markings on the lower tepals. The redbrown, bell-shaped corms bear fine marginal, downward directed teeth and are virtually identical with those of L. arenicola, which has similarly coloured flowers, much smaller in size and with a ± cylindric perianth tube mm long. The larger flower of L. macrospatha is reflected in a dorsal tepal ± 20 7 mm whereas the smaller flowers of L. arenicola have a dorsal tepal mm. The longer anthers of L. macrospatha, 3 5 mm long, also reflect the larger flowers than those of L. arenicola, in which the anthers are mm long. The two species appear to grow in an identical habitat, deep red sand, L. arenicola extending along the Atlantic coast from near Klawer in Western Cape through the sand plains and coastal dunes of western Namaqualand to the southern Richtersveld on the Anenous flats in Northern Cape, slightly overlapping but largely complementing the more northern distribution of L. macrospatha. The DNA sequence-based phylogeny supports the hypothesis that L. arenicola and L. macrospatha are closely allied, and are in current analyses, sister species (Forest et al. 2014). History: described by J.G. Baker in 1896, Lapeirousia macrospatha was based on a collection made by Harry Bolus in 1883 on his only expedition to Namaqualand. It was until recently represented in herbaria by limited material, much of it poorly preserved, and this led Goldblatt (1972) to regard the species as merely a longer-tubed variant of then equally poorly understood L. barklyi. Examination of living plants in the southern Richtersveld of Namaqualand has now made it clear that the two are separate species (Goldblatt & Manning 1994). Their flowers,

71 S T R E L I T Z I A 35 (2015) 63 seen alive, are quite different, those of L. barklyi mauve-pink rather than ivory with pink reverse and have a relatively wide floral cup formed by the upper part of the perianth tube and the tepal bases. We no longer believe that they are even immediately related. The corms also differ, those of L. macrospatha resembling those of L. arenicola in their bell-shape with fine, marginal teeth, whereas those of L. barklyi are triangular in profile with laterally extended spines like those of L. fabricii and its allies. Selected specimens SOUTH AFRICA. NORTHERN CAPE (Oranjemund): north of Port Nolloth at Holgat River, ( DD), 6 Sept. 1987, Germishuizen 4795 (PRE); 12 Sept. 1961, Hardy 603 (PRE) (Vioolsdrif): 1 km west of Eksteenfontein, ( CD), 1 Sept. 1986, Williamson 3608 (NBG); Eksteenfontein, in sand, 29 Aug. 1980, Venter 8062 (PRE) (Port Nolloth): 8 km east of Port Nolloth, ( BD), 13 Sept. 1974, Rösch & Le Roux 670 (PRE) (Springbok): near Abbevlakte, Aug. 1883, H. Bolus 6576 (BOL); road to Eksteenfontein, 2 4 km north of Port Nolloth road, ( AB), 8 Sept. 1980, FIGURE 16. Lapeirousia macrospatha, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

72 64 S T R E L I T Z I A 35 (2015) FIGURE 17. Distribution of Lapeirousia arenicola, ; L. macro spatha,. Goldblatt 5716 (M, MO, NBG); Farm Grasvlakte, sandy flats, ( AB), 19 Sept. 1989, Viviers 2042 (NBG), 28 Aug. 1991, Goldblatt & Manning 9138 (K, MO, NBG, PRE, M); Grootmis [Grootmist] to Springbok, 28 Aug. 1937, (CA/CB), Verdoorn & Dyer 1880 (K, PRE); sandveld just east of Grootmis at turnoff to Port Nolloth, ( CA), 25 Aug. 1993, Goldblatt & Manning 9655 (MO, NBG); ± 48 km west of Springbok, ( CB), Thompson 1056 (NBG); sandveld ± 55 km westnorthwest of Komaggas, Nuttrabooi 199, 0.5 km south of Buffelsrivier, ( CB), 27 Aug. 1999, Desmet 294 (NBG). 18. Lapeirousia arenicola Schltr. in Botanische Jahrbucher für Systematik 27: 105 (1900); Goldblatt: 54 (1972). Type: South Africa, [Western Cape], Atties [near Vredendal], 30 July 1896, Schlechter 8329 (B, holo.!; BM!, BOL!, COI!, E!, G!, GRA!, K!, MO!, P!, PH!, PRE!, S!, UPS!, iso.). [arenicola growing in sand, referring to the habitat, deep sandy ground.] Plants ( 150) mm high, often ± tufted, usually branched mainly at base, stem compressed and 2-angled, angles often weakly winged, slightly serrated, normally without cormlets at below ground nodes. Corm bell-shaped, mm below ground, ± 6 mm diam.; tunics red-brown, smooth, basal rim lightly serrated, teeth directed downward. Leaves 3 6( many), lowermost usually conspicuously longer than others, linear and ascending to falcate, 2 3 mm wide, strongly ribbed, upper cauline leaves shorter, usually resembling bracts and not or barely ribbed, usually subtending branches. Inflorescences 2 6-flowered spikes, lax or crowded; outer bracts green, leaf-like, mm long, lanceolate, acute, keeled throughout or at least distally, unifacial apically for 3 5 mm, keels often red, minutely serrated, margins hyaline, inner bract transparent with 2 green keels, 1 / 2 to 2 / 3 as long as outer, forked at tip. Flower zygomorphic, cream to biscuit-pink, lower tepals each with red mark near base; perianth tube 15 25( 33) mm long, cylindric, curving slightly outward at apex; tepals subequal or dorsal slightly larger, lanceolate and acute or obtuse, mm, usually spreading ± horizontally when fully open, all in same plane. Stamens unilateral, erect; filaments ± 7 mm long, exserted ± 3.5 mm from tube; anthers contiguous, mm long, violet, pollen mauve. Style dividing opposite upper half of anthers, style branches ± 1.5 mm long, divided for ± half their length. Capsules oblong, (8 )10 12 ± 6 mm, apices of locules forming auriculate lobes decurrent on wing-like locular ridges in upper half. Seeds globose, flattened at chalazal end, ± 1.4 mm diam., dark shiny brown. Chromosome number 2n = 16. Flowering time: August to September, occasionally in early October. Figure 18. Distribution and biology: restricted to the South African west coast and near interior, Lapeirousia arenicola extends from near Vanrhynsdorp and Vredendal in Western Cape northward through the sandveld and low coastal hills of Namaqualand as far north as the Anenous flats at the southern edge of the Richtersveld (Figure 17). It favours deep sandy soils, especially the rather fine-grained, red-tinged sands of lowland and coastal Namaqualand. It is commonly found in association with the spiny perennial grass Eragrostis spinescens, succulent-leaved Trachyandra falcata and T. bulbinifolia (Asphodelaceae) and strandveld annuals such as Heliophila (Brassicaceae), Manulea and Lyperia tristis (Scrophulariaceae), and Cotula (Asteraceae). It is a selfcompatible species, but probably facultatively outcrossing early in the flowering season when plants produce fairly large, long-tubed flowers with stronger colouring than later in the season. Towards the end of the flowering season the flow-

73 S T R E L I T Z I A 35 (2015) 65 ers of some populations have weakly contrasting markings, are smaller, and have a perianth tube mm long (compared with ± 25 mm on the same plants earlier in the season). We assume these late-blooming flowers are primarily autogamous. L. arenicola co-occurs at several localities with the florally almost identical L. simulans. Diagnosis and relationships: low-growing and typically a small plant, Lapeirousia arenicola can readily be distinguished from most other species of Lapeirousia by its small size, pale pinkishbeige, slender-tubed flowers and narrow tepals. Perhaps most easily confused with L. macrospatha, which has similarly coloured flowers, L. arenicola has a uniformly narrow, cylindric perianth tube mm long and tepals mm, whereas L. macrospatha has a wider tube, mm long, expanded toward the apex and considerably larger tepals, the uppermost largest and ± 20 7 mm. Although the two species appear similar on casual examination, they are quite distinct and do not intergrade although their ranges overlap slightly. Both have been found growing together just north of the Port Nolloth road below Anenous Pass and here they maintain their differences. Perianth tube length may coincide but the tepals of L. arenicola are always smaller and the tube more often than not is substantially shorter than in L. macrospatha. As discussed above under L. macrospatha, we regard L. arenicola as immediately related to this longer-tubed species, an association supported by the DNA-based phylogeny (Figure 1). C A B FIGURE 18. Lapeirousia arenicola, Goldblatt & Manning A, flower and corm; B, flower; C, capsules. Scale bar: 10 mm. Artist: John Manning.

74 66 S T R E L I T Z I A 35 (2015) Although Lapeirousia arenicola bears a superficial resemblance to the Western Cape L. anceps in flower colour and in the narrowly cylindric perianth tube, these similarities are evidently due to convergence as the molecular phylogeny (Figure 1) shows them to be distantly related. More puzzling is the striking resemblance between L. arenicola and L. simulans, which co-occur at several sites along the west coast, where their nearly identical flowers make a strong impression. Vegetatively, L. arenicola is always more compact in habit and often has a somewhat shorter perianth tube, so that they can be distinguished when seen growing side-by-side. More significant differences are underground: the corms of L. arenicola lie mm below the surface and the underground part of the stem rarely produces cormlets whereas corms of L. simulans are usually mm below ground and always have a fairly large cormlet at each of two underground nodes. Apart from an apparently close relationship to the northern Namaqualand L. macrospatha, and possibly with L. simulans, the affinities of L. arenicola are uncertain. Molecular data place L. arenicola closest to L. macrospatha in the sect. Lapeirousia clade. The relationship with L. jacquinii suggested by Goldblatt (1972), largely on the basis of their similar corms and additionally by similar capsules with auriculate apical lobes, is no longer tenable. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): Eksteenfontein road, 3 km north of Port Nolloth road, ( AB), 8 Sept. 1980, Goldblatt 5715 (MO, NBG); Spektakel, sand dunes, ( DA), 20 July 1967, Van der Schijff 6970 (PRE); c. 6 km south of Wildepaardehoek Pass, ( DC), Thompson 1077 (BOL, STE); sandy flats below Wildepaardehoek Pass, ( DC), 9 Sept. 1980, Goldblatt 5760 (MO) (Hondeklipbaai): between Hondeklipbaai and Wallekraal, ( AD), 4 Sept. 1932, Leighton s.n. (BOL30718), 28 Sept. 1976, Goldblatt 4242 (MO); road to Soebatsfontein from Buffels River, ( BB), 6 Sept. 1945, Leighton 1209 (BOL); near Soebatsfontein, ( BB), Sept. 1945, Lewis 1384 (SAM); Grootvlei, ( BB), 7 Sept. 1941, Barker 3710 (NBG); ± 25 km east of Wallekraal toward Garies, ( DA), 17 Sept. 1991, Goldblatt 9228 (MO); Farm Hardekoppie northwest of Kotzesrus on road to Groen River Mouth, 180 m, ( DC), 9 Sept 1987, Reid 1296 (PRE) (Kamiesberg): Eenkokerboom, ( CC), Sept. 1897, Schlechter (GRA). WESTERN CAPE (Vanrhynsdorp): ± 26 km north of Vanrhynsdorp, ( BC), 24 July 1948, Compton (NBG); Koekenaap to Graafwater, Farm Kommandokraal, ( CA), 10 Sept. 2008, Goldblatt & Porter (MO); 3 km northeast of Vredendal, ( DA), 7 Aug. 1970, Hall 3714 (NBG), 3784 (NBG); north of Klawer, ( DA), 2 Sept. 1951, Barker 7401 (NBG); Sept. 1945, Lewis 1406 (SAM); red sand flats 10 km north of Vanrhynsdorp, ( DA), 14 Sept. 1992, Goldblatt & Manning 9453 (MO, NBG). 19. Lapeirousia simulans Goldblatt & J.C.Manning in Novon 4: 341 (1994). Type: South Africa, [Western Cape], sandveld ± 11 km north of Vanrhynsdorp, 14 Sept. 1992, Goldblatt & Manning 9454 (NBG, holo.!; K!, MO!, PRE!, WAG!, iso.). [simulans imitating, for the close resemblance to L. arenicola, with which it was confused until 1994.] Plants mm high, 2 3( 5)-branched (rarely simple), branching ± divaricate, bearing a cormlet at each of 2 subterranean nodes, stem compressed and 2-angled, angles minutely serrated. Corm bell-shaped, 7 8 mm diam., mm below ground; tunics dark brown, glossy, margins lightly serrated, teeth directed downward. Leaves 3 or more, lowermost longest, usually ± twice as long as stem, linear, mm wide, corrugately ribbed, remaining leaves progressively shorter, becoming ± bract-like and usually subtending branch. Inflorescence lax spikes of (2)3 5 flowers; outer bracts green, transparent on veins, with slightly serrate keels decurrent on stem angles, mm long; inner bracts ± 1 / 2 as long as outer. Flowers zygomorphic, ± white, reddish-brown on reverse, lower 3 tepals each with small, claw-like tooth near base and with small basal and larger median dark red spot, unscented; perianth tube cylindric, mm long, slightly wider and curved near apex; tepals unequal, upper 3 slightly larger than lower 3, when fully open all held ± horizontally at right angles to tube, ± 11 4 mm, lower 3 held closely together, mm. Stamens unilateral, erect; filaments 5 6 mm long, exserted ± 3 mm from tube; anthers mm long, purple; pollen light purple. Style erect, adaxial to stamens, dividing between mouth of tube and anther bases, style branches ± 2 mm long, divided for slightly > 1 / 2 their length. Capsules oblong, 8 12 mm long, slightly warty in upper 1/2, without auriculate apices or locular ridges. Seeds globose, light brown, mm diam. Chromosome number unknown. Flowering time: early to late September. Figure 19.

75 S T R E L I T Z I A 35 (2015) 67 B A C FIGURE 19. Lapeirousia simulans, Goldblatt & Manning A, plant; B, capsules; C, seed. Scale bar: A, B, 10 mm; C, 0.5 mm. Artist: John Manning.

76 68 S T R E L I T Z I A 35 (2015) Distribution and biology: restricted to southern Namaqualand, Lapeirousia simulans extends from the Groen River west of Garies to Vanrhynsdorp in the south (Figure 20). Originally thought to be restricted to deep red sands at the southern margins of the Knersvlakte, it has now been recorded more widely in Namaqualand. Plants often co-occur with L. arenicola, which it closely resembles, and sometimes with L. anceps and L. fabricii, which have larger but similarly coloured flowers with nearly identical markings. We infer a shared pollination system using long-proboscid flies for L. simulans, but there are no records of pollinator visits to the species, which blooms somewhat earlier than its presumed pollinators are on the wing. Self-compatibility and autogamy, which we determined by pollinator exclusion, assure reproduction by seed in the absence of insect-mediated pollen transfer. Vegetative reproduction is ensured by production of cormlets at the underground nodes of the deeply buried stem. Diagnosis and relationships: the small corms with glossy dark brown tunics and a basal rim of minute, downward directed teeth closely resemble those of L. arenicola and several other members of sect. Lapeirousia. Although remarkably similar in general appearance to L. arenicola, the immediate affinities of L. simulans probably do not lie with that species, which is not surprising given that they are sometimes sympatric. Their flowers are so similar in tube length and general size and markings that they can be distinguished only with difficulty. Their flowering times overlap, but that of L. simulans is generally somewhat later in the season, usually mid- to late September, by which time the last flowers of L. arenicola are often depauperate and have shorter tubes and duller colouring than earlier in the season. The two are most readily distinguished by the underground part of the stem. While the corms of L. arenicola lie mm below the surface on a straight stem that lacks axillary cormlets, the underground stem of L. simulans is mm long so that the corm lies at a greater depth than in L. arenicola. Each of the two underground nodes consistently bears a large cormlet in L. simulans, a feature fairly rare in the genus, although also known in some other species including L. anceps and L. divaricata. Molecular studies show L. simulans to be a member of the sect. Lapeirousia clade, but not immediately allied to L. arenicola and L. macrospatha. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Hondeklipbaai): Farm Steenkampskraal, 9 km north of foot of Killians Pass, ( BA), 12 Aug. 1997, Goldblatt & Manning (MO); slopes between Soebatsfontein and Skilpad, ( BB), 15 Aug. 2005, Steyn 718 (PRE). WESTERN CAPE (Lepelfontein): Katdoringvlei, sandy slopes, ( BB), 10 Sept. 2009, Goldblatt & Porter (MO); Brand-se-Baai, strandveld, ( BD), 14 Sept. 1991, Van Rooyen 2030 (PRE) (Vanrhynsdorp): between Koekenaap and the coast, deep red sands, ( CA), Goldblatt, Manning & Porter (MO, NBG); 1.5 km northwest of Vredendal (Cohen s farm), ( DA), 8 Sept. 1970, Van der Merwe 151 (BOL, NBG); ± 10 km north of Vanrhynsdorp, ( DA), 20 Aug. 1993, Goldblatt & Manning 9609 (MO, NBG), 10 Sept (fr.), Goldblatt & Manning 9701 (MO, NBG); Vanrhynsdorp, ( DA), 2 Sept. 1951, Martin 825 (MO, NBG). 20. Lapeirousia spinosa (Goldblatt) Goldblatt & Manning in Novon 4: 343 (1994). L. divaricata var. spinosa Goldblatt: 66 (1972). Type: South Africa, [Northern Cape], ± 68 km west of Steinkopf to Port Nolloth, 24 Sept. 1970, Goldblatt 567 (BOL, holo.!). [spinosa spiny, referring to long, spinelike, rigid teeth that extend outwards from the corm base.] FIGURE 20. Distribution of Lapeirousia simulans, ; L. spinosa, ; L. tenuis,.

77 S T R E L I T Z I A 35 (2015) 69 Plants 50 90( 140) mm high; stem erect, sometimes branched at base, somewhat compressed but not often angled; without cormlets at nodes below ground. Corm ± bell-shaped (to ± obconic), 9 15 mm diam., mm below ground; tunics blackish, often glossy, basal rim with prominent horizontal spines. Leaves 3 4 (or more if plants branched), lowermost usually much the longest and usually shortly exceeding stem, linear to narrowly lanceolate, ascending or falcate, 2 4 mm wide, strongly corrugately ribbed; remaining leaves shorter and progressively more bract-like above, ribs weakly or not developed. Inflorescences spikes of (3 )5 9 flowers, usually moderately crowded; outer bract green or flushed reddish below, firm, 12 20( 35) mm long, enclosing slender part of tube, veins transparent, margins narrowly hyaline; channelled below, lightly keeled in upper 1 / 2, keels minutely serrated or crisped; inner bract up to 1 / 2 as long as outer, 2-keeled, with broad transparent margins. Flowers zygomorphic, white or flushed pink to lilac (more so with age and when dry), lower tepals yellow below and each with one or two purple median marks near limb base, bearing basal toothlike ridge ± 2 mm long, lightly sweet scented; perianth tube 9 12 mm long (shorter in dried material), narrow below, curved and widening above; tepals unequal, lanceolate, acute, dorsal largest, mm, erect, held apart from others, lower margins undulate or lightly crisped, upper lateral tepals united with lower 3 for ± 5 mm, directed forward below, reflexed distally, lower 3 tepals horizontal below, limbs mm, flexed sharply downward at ± 45, margins undulate. Stamens unilateral and arcuate; filaments 9 12 mm long, exserted 7 9 mm from tube; anthers parallel, 4 6 mm long, white; pollen white. Style arching over stamens, dividing between base and middle of anthers; branches 4 5 mm long, forked for slightly more than 1 / 2 their length, recurved. Capsules ovoid, 6 7 mm long, with auricular lobes above locules. Seeds globose, ± 1.2 mm diam., flattened at chalazal end, smooth, brown. Chromosome number 2n = 16. Flowering time: mid-august to mid-september. Distribution and biology: restricted to westcentral and northern Namaqualand, Lapeirousia spinosa extends from the Orange River at Arris Drift southward to Wallekraal, east of Hondeklipbaai (Figure 20). Although not yet recorded in Namibia, L. spinosa almost certainly occurs there, as it grows close to the south bank of the Orange River, the border of that country. Plants are usually found in compacted sandy soil in places where the ground is covered by small quartz pebbles, but on the plains west of Anenous Pass, L. spinosa occurs in its thousands in light clay devoid of stones. The species is most common in the north of its range, from the Anenous plains to the Orange River, but populations occur below the main Namaqualand escarpment as far south as Wallekraal. The lightly scented flowers appear to be pollinated by anthophorid bees, but pollinator activity, observed only in the Anenous Pass area, is low and more observations are needed. It seems likely that noctuid moths also visit and pollinate the species, as they do in L. divaricata, which has nearly identical flowers. Diagnosis and relationships: early collections of Lapeirousia spinosa were first included in the broadly similar L. divaricata and Gold blatt (1972) treated it as var. spinosa of that species. The flowers of the two are alike in their white (or palest pink) perianth, short perianth tube, lower three tepals united with the upper laterals for several mm, and tooth-like median ridges on the lower tepals. Goldblatt & Manning (1994) raised var. spinosa to species rank due to their quite separate geographic ranges, different habitats and differences in flower size and corm shape. Flowers of L. spinosa are larger, have a perianth tube 9 12 mm long and a dorsal tepal mm long compared with a tube 7 10 mm long and dorsal tepal mm long in L. divaricata. The latter, restricted to moist, sandy habitats in the Bokkeveld, Cederberg and Oli fants River Mtns, usually seasonally waterlogged, is also taller, has relatively lax spikes, bell-shaped corms and corm tunics with margins minutely serrated. L. spinosa occurs on dry, well-drained sandy flats, is a fairly compact plant, and has obconic corms and tunics with prominent, laterally directed spines. History: an early collection of Lapeirousia spinosa at the Kew Herbarium, without date, locality or collector but annotated Herb Forsyth indicates its source William Forsyth s private herbarium, auctioned by Sotheby s in 1835 and purchased for the Kew Herbarium. The collector may have been William Paterson who collected in Namaqualand in the late 1700s. Forsyth supported Paterson for a time, which explains how he acquired some of

78 70 S T R E L I T Z I A 35 (2015) Paterson s collections. The species was recollected in August 1883 by Harry Bolus east of Port Nolloth on his only expedition to Namaqualand, which he had reached by ship from Cape Town. The species remained confused with the superficially similar L. divaricata until treated as var. spinosa of that species by Goldblatt (1972). Later collections made it clear that the taxon merits separate species rank (Goldblatt & Manning 1992). Although floral morphology suggests the two species are related, they have different corms, those of L. divaricata lacking the prominent lateral spines found in of L. spinosa. Molecular studies retrieve the species in separate clades and we now place L. spinosa in sect. Lapeirousia, whereas L. divaricata is nested in sect. Chasmatocallis. Specimens examined SOUTH AFRICA. NORTHERN CAPE (Oranjemund): stony gravel flats near Arrisdrif, 31 Aug. 1925, ( DA), Marloth 12389B (PRE) (Vioolsdrif): 5 km east of Eksteenfontein, quartzite plain, ( CD), 1 Sept. 1986, Williamson 3610 (MO, NBG) (Port Nolloth): between Port Nolloth and Holgat, Sept. 1929, ( BB), M. Schlechter s.n. (PRE, NBG); between Port Nolloth and Eleven Mile Station, 22 Aug. 1883, ( BD), H. Bolus 6577 (BOL) (Springbok): Richtersveld, Anenous Flats, Farm Grasvlakte, ( AB), 15 Sept. 1992, Goldblatt & Manning 9456 (MO); road to Eksteenfontein, 4 km north of Port Nolloth highway, deep sand, ( AB), 8 Sept. 1980, Goldblatt 5730 (K, MO, NBG, PRE, S, WAG); 8 km west-northwest of Vaalheuvel, Stryd River valley, 24 Aug ( AD), Acocks (PRE); stony river valley near Harras House, ( AD), 24 Aug. 1980, Van Berkel 237 (MO, NBG); 5.7 km from N7 on road to Kommagas, from Wildepaardehoek Koingnaas intersection, common on quartzite pebble areas, ( CD), 21 Aug. 2002, Goldblatt & Porter (MO, NBG, PRE); Komaggas, ( CD), 9 Sept. 1950, Barker 6675 (NBG) (Hondeklipbaai): 4 km from Wallekraal to Kamieskroon, stony ground, ( BC), 28 Sept. 1976, Goldblatt 4246 (MO, NBG, PRE); Wallekraal, ( BC), 30 Aug. 1935, Compton 5420 (BOL, NBG). 21. Lapeirousia tenuis (Goldblatt) Goldblatt & Manning in Novon 4: 344 (1994). L. divaricata var. tenuis Goldblatt: 66 (1972). Type: South Africa, [Northern Cape], between Eleven Mile Station and Oograbies Poort (Port Nolloth and the mountains of the interior), 23 Aug. 1883, H. Bolus 6575 Lapeirousia divaricata Baker (BOL, holo.!; G, K!, SAM!, iso.). [tenuis slender or thin, referring to the flowers with narrow, almost linear tepals.] Plants ( 220) mm high, mostly 2 3-branched or unbranched; branches produced well above ground level; stem lightly compressed, oval in section, occasionally lightly or obscurely angled, angles smooth. Corm broadly obconic (triangular in outline), mm diam., up to 40 mm below ground; tunics dark brown to blackish, basal margins with fine laterally extended teeth 1 3( 4) mm long, often massively accumulating. Leaves 3 4, only lowermost well-developed, straight and linear or falcate, usually ± as long as to twice as long as stem, rarely slightly shorter, 2 3( 4) mm wide, corrugately ribbed; remaining leaves < 1 / 2 as long as lowermost and progressively more bract-like above. Inflorescences spikes of 9 16 flowers, lateral spikes with fewer flowers, 2-ranked in bud, becoming spiral after anthesis; outer bract grey-green, transparent on veins, acute, 7 11 mm long, folded on midline but not keeled, apices acute and curving upward, inner bract 1 / 2 to 1 / 3 as long as outer, transparent with two green keels, forked at tip. Flowers zygomorphic, pale lilac-pink, lower tepals each with darker blotch or streak in midline and claw-like cusp near base, cusp ± white surrounded basally by red-purple zone, occasionally upper lateral tepals also lightly marked in lower 1 / 2, lightly sweet scented; perianth tube mm long, slender below, curving outward and wider in upper 1 / 3 ; tepals narrowly lanceolate to ± linear, dorsal mm, ± erect, upper laterals recurving from base, lower 3 united for ± 2 mm, extending horizontally below, flexed downward distally. Stamens unilateral, ± erect; filaments ± 6 mm long, exserted ± 3 mm from tube; anthers ± 3 mm long, parallel, light purple; pollen ± white. Style straight, adaxial to stamens, dividing between middle and apex of anthers; branches ± 2.5 mm long, deeply divided and recurved. Capsules globose-trigonous, ± 4 5 mm long. Seeds globose, mm diam., smooth, brown, glossy. Chromosome number unknown. Flowering time: mainly in July, occasionally midto late August, and once recorded in bloom in early September. Distribution and biology: poorly known and evidently rare, Lapeirousia tenuis appears to be restricted to coastal central Namaqualand (Figure 20). All but two of the few known collections are from the coastal plain north and east of Kleinzee suggesting a very local distribution. The

79 S T R E L I T Z I A 35 (2015) 71 remaining two collections lack precise locality data, but at least one is from some distance to the north, inland from Port Nolloth. The Namaqualand coast between Port Nolloth and Hondeklipbaai is not well collected and the distribution of L. tenuis may be wider than the present record suggests. The species favours stony gravel slopes and is most often found in fine sandy clay mixed with granite and quartzite pebbles. At one site we have seen it growing close to L. spinosa and L. macrospatha, and a clear distinction was evident in the habitat preferences of the three. L. tenuis grew on sloping ground in pebbly places, L. spinosa grew on flat, less stony ground, and L. macrospatha was restricted to deep sands on flat ground. Diagnosis and relationships: although when first accorded formal taxonomic rank, Lapeirousia tenuis was regarded as a variety of L. divaricata (Goldblatt 1972), additional collections of the latter, and of the related L. spinosa, also regarded by Goldblatt as a variety of L. divaricata, make it clear that three species are involved. They share spiny corm tunic rims and comparatively small bracts but the flowers differ markedly. In both L. divaricata and L. spinosa the lower tepals are united with the upper laterals for some distance and the lower tepals have large, tooth-like median ridges. This is not the case in L. tenuis, the narrow tepals of which spread ± uniformly from the apex of the perianth tube. The lower tepals appear to be united for a short distance but are not fused to the upper tepals at all, and they have claw-like basal cusps rather than tooth-like median ridges. History: discovered by Harry Bolus in August 1883 east of Port Nolloth, and recollected much later by Rudolf Marloth in 1925, Lapeirousia tenuis was at first identified as L. setifolia (the name then used for L. divaricata). The narrow tepals and distinctively obconic corm with spiny basal margins prompted Goldblatt (1972) to treat the four collections then known as a separate varieties of L. divaricata. As the latter has become better known, it was evident that L. divaricata is restricted to the northwestern Cape, and that the two Namaqualand variants deserve species rank. The spiny corm tunics of L. tenuis appear to most closely resemble those of L. dolomitica and L. lewisiana rather than L. divaricata and molecular studies retrieve them as only distantly related. L. tenuis and L. spinosa are retrieved as a terminal species pair in our molecular phylogeny. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): 15 km north of Kleinzee, ( CA), 24 July 1983, Van Wyk s.n. (BOL); 8 km north of Grootmis [Grootmist], ( CA), 20 Aug. 1952, Hall 579 (NBG); (K, MO, NBG, PRE); sandveld between Grootmis and Springbok, ( CA/CB), 28 July 1937, Verdoorn & Dyer 1882 (PRE); Spektakel Kleinzee road, 1 mile [1.5 km] east of Langhoogte Mine, ( CB), 25 Aug. 1993, Goldblatt & Manning 9651 (K, MO, NBG, PRE), 17 Aug. 1980, Van Berkel 196 (MO, NBG); 62 km west of Springbok, road to Grootmis, ( CB), 21 Aug. 1994, Goldblatt & Manning 9935 (MO, NBG); ± 6 km north of Uguabis [not Uhabis, northeast of Vioolsdrif in Namibia], 6 Sept. 1925, ( DA), Marloth (PRE). 22. Lapeirousia violacea Goldblatt in Contributions from the Bolus Herbarium 4: 59 (1972). Type: South Africa, [Western Cape], Biedouw Valley, 26 Aug. 1970, Goldblatt 569 (BOL, holo.!; K!, MO!, iso.). [violacea violet, the flower colour.] Plants mm high, simple or 1 4-branched from near base, stem lightly compressed, generally 1-angled, angles decurrent on leaves or bract keels. Corm broadly obconic, 8 10 mm diam.; tunics dark brown, with light vertical ridging, basal rim with spreading spines 2 5 mm long. Leaves 4 6 (or more when stem branched), lowermost longest, generally shorter than stem and ascending, narrowly lanceolate or falcate, 5 7 mm wide, corrugately ribbed other leaves 1 / 2 as long or less, only lightly ribbed, upper of these often bract-like. Inflorescences (2)3 or 4-flowered spikes; outer bracts pale green, firm, 15 22( 30) mm long, folded and keeled in midline, keels minutely crisped, often 2 3 mm wide near apex; inner bracts mostly green with broad transparent margins, 2 / 3 to ± as long. Flowers zygomorphic, dark violet, lower tepals each with dark purple to red spearshaped basal mark and white lines along lower margins; when fully open tepals held ± in one plane, lowermost or all three lower tepals bearing a small basal claw-like cusp; perianth tube mm long, slender, widening and curving toward apex; tepals unequal, narrowly lanceolate, dorsal largest, ± 10 5 mm wide, sepa-

80 72 S T R E L I T Z I A 35 (2015) FIGURE 21. Lapeirousia violacea, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

81 S T R E L I T Z I A 35 (2015) 73 rated from others, upper lateral tepals reflexed, lower 3 tepals joined basally for 3 4 mm, free parts held closely together, 7 8 ± 4 mm. Stamens unilateral, ± erect; filaments ± 9 mm long, exserted ± 5 mm from tube; anthers ± 2.5 mm long, purple; pollen purple. Style dividing opposite base or middle of anthers, style branches ± 2 mm long, deeply forked and recurved. Capsules oblong-trigonous, remaining enclosed in bracts, 9 11 mm long, without locular ridges. Seeds globose, slightly domed outward at chalazal end, mm diam., light brown. Chromosome number 2n = 16. Flowering time: August to mid-september. Figure 21. Distribution and biology: Lapeirousia violacea is restricted to the mountains and intermontane valleys of northwestern Western Cape and adjacent Northern Cape, where it extends from the valleys of the Biedouw and Doorn rivers in the south through Botterkloof as far north as Papkuilsfontein south of Nieuwoudtville (Figure 22). Plants grow in sandy soil, mostly in stony ground, and tend to be scattered and seldom common. Surprisingly, they seem to occur with the same frequency in valley bottoms as on exposed hilly sites. Plants are often found growing sympatrically with L. jacquinii, which has flowers nearly identical in colour, shape and size. They share the same pollinators, Prosoeca sp. and P. peringueyi (Nemestrinidae), long-proboscid flies that pollinate several species of Lapeirousia and Babiana that have dark blue to purple, violet or red, long-tubed flowers (Goldblatt et al. 1995). Despite their sympatry and sharing the same pollinators, no hybrids between the two are known. Both L. violacea and L. jacquinii are self-compatible and facultatively autogamous, but are, we assume, frequently outcrossing, for both species are regularly visited by Prosoeca species in the wild. As in most Lapeirousia species, the flowers close at night and in cool weather. Diagnosis and relationships: the dark violet flowers with reddish to purple and white markings on the lower tepals of Lapeirousia violacea are so like those of L. jacquinii that plants must be examined closely to tell them apart, especially as they sometimes grow together. The two differ in their floral bracts and corms and examination of these organs makes them easy to distinguish. The corms of L. jacquinii are bellshaped and lack the long spiny, laterally projecting teeth typical of the obconic corms of L. violacea, which has inflated outer bracts folded or slightly keeled, thus very different from the shorter, typically 2-keeled bracts of L. jacquinii. Despite the similarity of their flowers, L. violacea and L. jacquinii are not immediately related and are here assigned to separate sections of the genus, an action supported by our molecular phylogeny. The bracts and especially the obconic, basally toothed corms of L. violacea suggest a close relationship with L. dolomitica and its allies of Namaqualand and southern Namibia as well as L. purpurea from the Swartruggens Mtns of Western Cape. History: evidently recorded in 1897 by Rudolf Schlechter, who gave his collection the manuscript name L. affinis, Lapeirousia violacea remained undescribed until 1972 when it was formally named (Goldblatt 1972). Schlechter s epithet was most likely intended to refer to the similarity of the flowers to those of the common and sometimes sympatric and co-blooming L. jacquinii. Representative specimens SOUTH AFRICA. NORTHERN CAPE 3119 (Calvinia): Nieuwoudtville Escarpment, Farm Papkuilsfontein, ( AC), 5 Sept. 1995, Goldblatt & Manning (MO, FIGURE 22. Distribution of Lapeirousia dolomitica, ; L. lewisiana, ; L. purpurea, ; L. violacea,.

82 74 S T R E L I T Z I A 35 (2015) NBG); Lokenburg, ( CA), Barker 1308 (NBG), Leistner 324 (PRE); Karoskloof, Ceres Karoo, ( DC), 27 Aug. 1983, Bayer 3507 (NBG). WESTERN CAPE 3119 (Calvinia): top of Botterkloof Pass, among sandstone rocks and pavement, ( CC), 31 Aug. 2006, Goldblatt & Porter (MO); 1 mile [1.5 km] south of Doorn River bridge, ( CC), Lewis 2345 (SAM) (Wuppertal): Biedouw Valley, Welbedacht (growing with L. jacquinii), ( AA), 5 Sept. 1991, Goldblatt 9197 (MO); Lammkraal, (?AA), 14 Aug. 1897, Schlechter (BOL, GRA, K, MO). 23. Lapeirousia purpurea Goldblatt & J.C.Manning, sp. nov. TYPE. South Africa, Western Cape, 3219 (Wuppertal): Swartruggens, Gansfontein, rocky sandstone slopes, ( DA), Sept. 2008, Manning & Roux 3179 (NBG, holo.; K, MO, PRE, S, iso.). Plants mm high, simple or fewto many-branched, mostly from base, stem compressed and 1- or 2-angled. Corm triangular in outline, basal rim prominently serrated, mm diam. at base. Leaves several, the basal ribbed, 1 / 3 to ± as long as stem, to 5 mm wide, upper leaves progressively smaller above, smooth, the uppermost resembling floral bracts. Spike 6 10-flowered; bracts green, outer mm long, with shallow, slightly serrated keel, inner ± 1 / 3 to 1 / 2 as long as outer, membranous with two darker keels, notched at apex. Flowers blue-purple, lower tepals with V-shaped white mark above base surrounded by darker violet to ± black zone, lower median tepal with a small claw-like tooth above base, unscented; perianth tube slender below, widening and curving outward in upper 1 / 3, mm long; tepals unequal, dorsal erect, ovate, ± 15 6 mm, upper laterals slightly smaller, recurved, lower tepals united basally for ± 3 mm, curving downward, ± 8 4 mm. Stamens unilateral, arcuate; filaments ± 7 mm long, exserted ± 3 mm from mouth of tube; anthers suberect, prominently tailed, ± 4 mm long; pollen yellow. Style arching above stamens, dividing opposite lower 1 / 3 to middle of anthers, style branches ± 3.5 mm long, divided for slightly > 1 / 2 their length. Capsules ovoid, ± 12 mm long, showing outline of seeds, with moderately well-developed locular ridges. Seeds unknown. Chromosome number unknown. Flowering time: September. Figure 23. Distribution and biology: Lapeirousia purpurea is known from just two collections from interior valleys of the Swartruggens that drain into the Tanqua Basin (Figure 22). Plants grow in sandy soil among sandstone rocks. The intensely blue-purple flowers with white markings on the lower tepals and long perianth tube exhibit the typical adaptations for the Prosoeca peringueyi long-proboscid fly pollination system (Goldblatt & Manning 2000). Floral biology of the species remains to be investigated. Diagnosis and relationships: first collected in September 2008, Lapeirousia purpurea is a surprising discovery, and emphasises the pattern of local endemism of so many members of the Cape flora. The corms, triangular in profile, have fine teeth projecting from the base, and resemble closely corms of L. dolomitica, L. lewisiana and L. violacea, also long-tubed species. L. violacea can be distinguished by the rich violet colour of the perianth with the upper lateral tepals strongly reflexed and the lower three tepals with a V- shaped white mark above the base surrounded by a dark violet to black zone. The lower median tepal bears a small claw-like cusp near the base. In L. violacea all three lower tepals usually bear a small tooth near the base, but such teeth are lacking in L. dolomitica and L. lewisiana. Additional specimens SOUTH AFRICA. WESTERN CAPE (Worcester): Swartruggens, De Mond to Farm Elandsdrif, Kaggakamma Nature Reserve ( DA), Sept. 2007, Harrower 3590 (NBG). 24. Lapeirousia dolomitica Dinter in Repertorium specierum novarum regni vegetabilis 29: 255 (1931); Goldblatt: 61 (1972). Type: Namibia, Alicetal, pump station near Pomona, June 1929, Dinter 6432 (B, holo.!; BM!, BOL!, G!, GH!, K!, iso.). Plants (50 ) mm high, simple or 1 2-branched from base, stem terete or compressed and 2-ridged to 2-winged below first flower. Corm mm diam., broadly conic (triangular in outline); tunics dark brown, rugose, basal rim with well-developed, laterally projecting spines. Leaves 1 3, lowermost basal

83 S T R E L I T Z I A 35 (2015) 75 A C D B E and longest, straight to falcate, mm long, usually ± as long as stem, mm wide, strongly ribbed, smooth or minutely papillate along rib margins, remaining leaves much smaller, resembling floral bracts. Inflorescences spikes of (5 )8 15 flowers, initially 2-ranked, becoming spiral; outer bract mm long, pale to dark green, margins hyaline or red, semi-transparent over veins, inflated, ± round in section, abaxial surface weakly keeled, keels usually restricted to distal half, inner bract 1 / 2 to 1 / 3 as long as outer, with two broad green keels. Flowers zygomorphic, pale blue to violet or lilac, lower 3 tepals partly or entirely yellow near base and darker blue or purple in median upper half, sometimes with short median ridges near base usually curving downward, sometimes sweetly scented; perianth tube slender, ascending, arching outward near apex, 25 35( 40) mm long (in severely stressed plants occasionally less than 25 mm); tepals unequal, lanceolate and acute, dorsal longest and erect, (12 ) mm, narrowed below, upper laterals lightly shorter, spreading or reflexed, lower 3 tepals united for 2 3 mm, slightly narrowed toward base, held closely together, (10 )12 14 mm long. Stamens unilateral, ± erect; filaments ± 10 mm long, exserted 2 4 mm, arching outward; anthers mm long, parallel, purple or cream, pollen lilac or cream. Style arching adaxial to stamens, usually dividing opposite middle 1 / 3 of anthers (rarely below anther bases in northern populations), dark violet; style branches 3 4 mm long, recurved, divided for 1 / 2 their length. Capsules ovoid-oblong, weakly trigonous, mm long, without locular ridges, usually warty or papillose. Seeds globose, chalazal end somewhat flattened, ± 1.8 mm diam., glossy dark brown. Chromosome number 2n = 16. Flowering time: June to late July, sometimes in August. FIGURE 23. Lapeirousia purpurea, Manning & Roux A, plant; B, flower; C, half flower; D, outer (right) and inner (left) bracts; E, capsule. Scale bar: 10 mm. Artist: John Manning. Distribution and biology: restricted to the most arid parts of the southern African winter rainfall zone, Lapeirousia dolomitica is most common in northern Namaqualand but extends north into southwestern Namibia as far as Pomona (Figure 22). It is probably fairly common in southern Namibia although the number of collections from there is scant owing to the rough terrain and few passable roads. Despite the epithet dolomitica the species does not normally grow in dolomite or any other type of limestone

84 76 S T R E L I T Z I A 35 (2015) but it seems to favour rocky situations where the parent rock is at least partly quartzite and does not often grow in the granite-derived soils that predominate in Namaqualand. Although variable in flower colour, perianth tube length and even for presence of fragrance, Lapeirousia dolomitica is most probably consistently pollinated by the long-proboscid nemestrinid fly, Prosoeca peringueyi. We wonder whether floral fragrance, notable only in populations with pale blue flowers, is merely an added attraction, or signifies an alternative pollinator here, then most likely moths. Diagnosis and relationships: Lapeirousia dolomitica is recognised by its distinctive, obconic corms with blackish, rough-surfaced tunics bearing prominent spines extending outward from the base and the rather inflated bracts with only weakly developed keels. When seen alive the bracts are unusual in having alternating longitudinal bands of pale translucent and dark opaque colouring. The flowers are pale blue, lilac, or said in some collections to be violet, always with pale yellow markings at the bases of the lower tepals and sometimes short, tooth-like basal ridges. Among the long-tubed species of Lapeirousia, it has a shorter than usual perianth tube, mostly mm long. In overall appearance flowers of L. dolomitica are particularly reminiscent of the central Namaqualand species, L. tenuis but this species has a perianth tube mm long and very well-developed, tooth-like ridges on the lower tepals. The two also have similar corms. In an earlier treatment of Lapeirousia dolomitica, Goldblatt (1972) recognised two subspecies, the northern subsp. dolomitica and the central and southern Namaqualand subsp. lewisiana. Additional collections of both, then poorly collected, have shown that their differences are consistent and of greater degree than usual for subspecies and we now recognise the latter as a separate species, L. lewisiana. The two share similar corms with spiny basal margins but that character has been documented in L. spinosa, L. kamiesmontana and L. violacea as well. Another shared feature, not until now recognised, are particularly long style branches, up to 4 mm long. However, smooth leaves in L. dolomitica versus sparsely hairy leaves in L. lewisiana plus differences in capsule surface texture, perianth tube length, perianth colour and markings, and shape of the lower tepals emphasise their differences (Table 5). L. dolomitica always has paler coloured flowers with a somewhat shorter tube, mostly mm long, whereas L. lewisiana has larger, deep magenta red flowers with a tube mostly mm long. The bracts of L. dolomitica are only lightly inflated and generally have weakly developed keels but L. lewisiana has strongly inflated bracts lacking keels. Lastly, the capsules of L. dolomitica are often papillose to warty but the character is not consistent in the species. Capsules of L. lewisiana are always smooth as are those of other species of the genus. Representative specimens NAMIBIA (Bogenfels): Sperrgebiet, east of Pomona, ( AB), 3 Aug. 2001, Smook (MO, PRE, WIND); Klinghardt Mtns, west-facing slopes, ( BC), 26 July 1977, Müller 660 (M, WIND) (Witputz): Sperrgebiet, Auros Mtns, eastern side of highest peak, 905 m, quartzite slope, ( CB), 10 Aug. 2001, Smook (PRE); Auros Mtns, ( CB), 11 Aug. 2001, Smook (MO, PRE); Zebrafontein 87, rocky ground, ± m, ( DA), 11 Sept. 1973), Giess (M, NBG, PRE, WIND); 22 Aug. 1983, Venter 8918 (PRE); Witputs, ( DA), July 1939, Herre s.n. (BOL 30721); 12 km west of Lorelei Mine, riviergeröll, ( DD), 30 Aug. 1963, Merxmuller & Giess 3318 (M, WIND) (Grünau): Karas District, Aurus Mtns, saddle overlooking Roter Kamm, rock cracks, ( CB), 27 Sept. 1996, Mannheimer & Mannheimer 334 (PRE, WIND) (Oranjemund): Rooilepel summit, July 1989, ( BA), Williamson 4262 (NBG); quartz ridges north of Obib, ( DA), 1 Sept. 1963, Merxmuller & Giess 3420 (BR, M, WIND). SOUTH AFRICA. NORTHERN CAPE (Oranjemund): Daveras Diamond Area 1, ( BB), June 1977, Williamson 2551 (BOL); Richtersveld, Numees, ( BD), Aug. 1983, Jurgens 20 (PRE); gravel flats at Arrisdrif, 70 m, ( DA), 31 Aug. 1925, Marloth 12389a (PRE) (Vioolsdrif): between Eksteenfontein and Little Helskloof, rocky outcrop among quartz pebbles, ( CA), 9 Aug. 1979, Snijman 163 (NBG); Stinkfontein Mtns, near foot of Cornelsberg, ( CA), 22 Aug. 1994, Goldblatt & Manning 9949 (MO, NBG, PRE); 10 km west of Eksteenfontein, ( CC), 9 Aug. 1986, Williamson 3539 (BOL, NBG); 20 km west of Eksteenfontein, ( CD), 1 Sept. 1986, Williamson 3606 (MO, NBG); east of Eksteenfontein in open quartz pebble sites, ( CD), July 1989, Williamson 4276 (NBG) (Springbok): Lekkersing, ( AA), 230 m, 24 Aug. 1925, Marloth (PRE); Karuchab Poort south of Lekkersing, ( AA), 18 Aug. 1970, Tölken s.n. (BOL 30720); mountains north of Grasvlakte on Eksteenfontein road, ( AB), Aug. 1992, Goldblatt & Manning 9308 (MO, NBG, PRE).

85 S T R E L I T Z I A 35 (2015) Lapeirousia lewisiana B.Nord. in Botaniska Notiser 123: 434 (1970). L. dolomitica subsp. lewisiana (B.Nord.) Goldblatt: 64 (1972). Type: South Africa, [Western Cape], Komkans, 11 Aug. 1962, Nordenstam 965 (NBG, holo.!; S!, iso.). Plants (50 ) mm high, simple or 2-branched from base, stem terete or compressed and 2-ridged to 2-winged below first flower. Corms mm diam., broadly obconic; tunics dark brown, rugose, basal rim with well-developed, slender, laterally projecting spines. Leaves 1 3, lowermost basal and longest, straight to somewhat falcate, mm long and usually ± as long as stem, mm wide, strongly ribbed, sparsely and irregularly white-hairy on rib edges at least in lower half, remaining leaves small, resembling floral bracts. Inflorescences spikes of up to 14 flowers; outer bract ± 20 mm long, inflated, usually rounded and without keels except occasionally near tips. Flowers reddish purple to magenta, lower tepals with broad ± white median streak in lower, narrow half and darker red-purple blotch in upper centre; perianth tube (35 )40 48 mm long; tepals unequal, lanceolate to ± spathulate or obovate, apex acute or rounded, dorsal ± mm, lower 3 united for ± 3 mm, narrow at base for 2 3 mm, held at ± 45º to horizontal, ± mm. Stamens unilateral, ± erect; filaments exserted ± 3.5 mm; anthers 3 4 mm long, dark violet. Style arching over stamens, between middle and apex of anthers, style branches 3 4 mm long, recurved, divided for 1 / 2 their length. Capsules ovoid-oblong, weakly trigonous, mm long, without locular ridges, smooth. Seeds globose, ± flattened at chalazal end, mm diam., glossy dark brown. Flowering time: mid-july to mid-august. Figure 24. Distribution and biology: never common, Lapeirousia lewisiana is recorded from several collections from central and southern Namaqualand. It is best known from the area between Garies and Bitterfontein, but also occurs locally to the west toward the coast (Figure 22). The type locality is in the extreme southern end of its range near Komkans, west of Nuwerus. An isolated population from near Komaggas is the most northern record. Diagnosis and relationships: described in 1970 and shortly thereafter reduced to subspecies rank in Lapeirousia dolomitica (Nordenstam 1970; Goldblatt 1972), the status and affinities of L. lewisiana have proven difficult to assess. With several additional records of both taxa now available, we consider it most useful to treat them as separate species. Although they share virtually identical obconic corms with prominent, spreading marginal spines, and long tubed flowers, several differences between them have been confirmed in all new collections. These include different flower colour and markings, perianth tube length and leaf vestiture. L. lewisiana has magenta red (also sometimes described as purple) flowers, the lower three tepals with prominent white, median streaks in the proximal 1 / 3, and the perianth tube is typically mm long (Table 5). Unusual for Lapeirousia, the basal foliage leaf is finely, but sparsely, hairy in all specimens we have examined. In contrast L. dolomitica has smooth leaves, a pale blue to violet perianth with paired white streaks on the lower tepals, a perianth tube mm long, rarely slightly longer. Capsules are poorly known in L. dolomitica, but are slightly warty or papillose where recorded, whereas L. lewisiana has smooth capsules. There are also slight differences in the bracts of the two species, although somewhat inflated in both: TABLE 5. Comparison of critical taxonomic features of Lapeirousia dolomitica, L. kamiesmontana and L. lewisiana Species Leaf surface Perianth colour Lower tepal markings L. dolomitica smooth pale blue to pale lilac white-edged blue to violet Perianth tube, mm Capsule surface 25 35( 40) often warty or papillose L. kamiesmontana ± hairy white red smooth L. lewisiana ± hairy dark magenta-red white-edged dark magenta (35 )40 48 smooth

86 78 S T R E L I T Z I A 35 (2015) FIGURE 24. Lapeirousia lewisiana, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

87 S T R E L I T Z I A 35 (2015) 79 bracts of L. dolomitica have a weakly developed keel in the distal half, whereas those of L. lewisiana are rounded or become slightly folded only along the midline near the tips. Despite its close relationship to Lapeirousia dolomitica, L. lewisiana is most easily confused with the common Namaqualand species, L. silenoides, which it resembles in its long-tubed, darkly pigmented magenta-red flowers. The latter has the bell-shaped corms without spiny projections of any sort that place it in sect. Chasmatocallis, and the superficial resemblance to L. lewisiana is evidently due to convergence for the same pollinator, the long-proboscid fly, Prosoeca peringueyi. Also closely related to Lapeirousia lewisiana but readily distinguished from it, L. kamiesmontana has similar vegetative features, including the corm tunics with spiny margins and slightly hairy foliage leaf. Flowering later in the season, usually two to three weeks after L. lewisiana, even at the same elevation, L. kamiesmontana has white flowers, the lower tepals marked with red, the tube and the reverse of the tepals flushed dark red, and a perianth tube mostly mm long. We have considered that the two species are closely related but adapted for pollination by different species of long-proboscid flies. An immediate relationship is not, however, supported by the molecular phylogeny (Figure 1). Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): Wolfberg-Komaggas, 800 m, ( CB), 23 Aug. 1981, Van Berkel 364 (MO) (Hondeklipbaai): Kotzesrus, granitic soil, ( DD), Aug. 1980, Van Berkel 172 (NBG); 10 km south of Garies on road to Bitterfontein, ( CA), 27 Aug. 1970, Goldblatt 538 (BOL), 29 Aug. 1991, Goldblatt & Manning 9198 (K, M, MO, NBG, PRE); 8.5 km north of Garies on the road to Leliefontein, ( CA), 19 Sept., 1991 (fr.), Goldblatt 9269 (MO, NBG, PRE); 30 km south of Garies, stony east-facing slope, ( CC), 2 Aug. 1994, Goldblatt & Manning 9893 (MO, NBG). WESTERN CAPE (Vanrhynsdorp): 4 km north of Farm Komkans, Geelkop, stony quartzite, ( AA), 7 Sept (fl. & fr.), Nordenstam & Lundgren 1715 (MO, NBG, S). 26. Lapeirousia kamiesmontana Goldblatt & J.C.Manning, sp. nov. TYPE. South Africa, Northern Cape, 3018 (Kamiesberg): between De Kom and Garies, ( AC), 27 Aug. 1976, Goldblatt 3980 (NBG, holo.; K, MO, PRE, S, iso.). Plants mm high, simple or 2-branched from base; stem round to slightly compressed. Corm mm diam., broadly obconic; tunics dark brown, rugose, basal rim with long, slender, spreading spines. Leaves 1 3, lowermost basal and longest, straight to somewhat falcate, mm long, sometimes exceeding flowers, mm wide, strongly ribbed, sparsely woolly (rarely ± smooth), remaining leaves much smaller, resembling floral bracts. Inflorescence spikes of (5 )8 10 flowers, initially 2-ranked, becoming spiral; outer bract mm long, acute, ± inflated, rounded, without keels, inner bract ± half as long, transparent with 2 green keels. Flowers white (pale yellow when dry), flushed red on reverse of tepals and tube, lower tepals with red blotch in lower 1/ and paired, red lines near base 3 running into throat (purple when dry); perianth tube mostly mm long, ± uniformly wide throughout; tepals unequal, obovate and markedly narrowed in basal 2 mm, dorsal (7 )9 10 mm, lower tepals held ± horizontally, united basally for ± 1.5 mm, ± mm. Stamens ± 10 mm long, exserted 2 3 mm; anthers mm, violet, pollen purple. Style dividing opposite anther tips, style branches ± 3.5 mm long, divided for half their length, recurved, white. Capsules and seeds unknown. Flowering time: late August to early October. Figure 25. Distribution and biology: Lapeirousia kamiesmontana is restricted to the southern Kamiesberg where it has been recorded from the flats around the Farm Doornkraal at ± 600 m and in the Langkloof at elevations of up to at least 950 m (Figure 26). The rugged area is poorly explored except alongside the road from Doornkraal to Welkom and we suspect the species has a wider range than is documented. Plants grow in decomposed granitic soil and occasionally in granite outcrops. The form and colour of the long-tubed flowers suggest pollination by longproboscid flies, but no insect visitors have been recorded for the species. Diagnosis and relationships: the broadly obconic corm with dark brown, rugose tunics spiny along the rim of Lapeirousia kamiesmontana are shared with L. dolomitica and L. lewisiana. Observations in the field show that the

88 80 S T R E L I T Z I A 35 (2015) B C A FIGURE 25. Lapeirousia kamiesmontana, Goldblatt & Manning A, plant and corm; B, half flower; C, outer (left) and inner (right) bracts. Scale bar: 10 mm. Artist: John Manning.

89 S T R E L I T Z I A 35 (2015) 81 Gladiolus fabricii (D.Delaroche) Thunb.: 186 (1800). Ovieda fabricii (D.Delaroche) Spreng.: 147 (1824). Peyrousia fabricii (D.Delaroche) Sweet: 499 (1830a). Meristostigma fabricii (D.Delaroche) A.Dietr.: 597 (1833). Type: South Africa, Olifants River valley, Leipoldt s.n. [BOL17111, neo.!, designated by Goldblatt & Barnard: 305 (1970)]. FIGURE 26. Distribution of Lapeirousia fabricii subsp. compressa, ; subsp. fabricii, ; subsp. purpurascens, ; L. kamiesmontana,. three do not intergrade: relatively late-blooming L. kamiesmontana consistently has white flowers with distinctive red markings and a perianth tube mostly mm long (Table 5). In contrast, early-blooming L. lewisiana has dark red-purple to magenta flowers with white markings and a perianth tube (35 )40 48 mm long. The ranges of the two do not overlap and flowering times, even at the same elevation, are separated by at least two to three weeks, if not longer, in any particular year. We consider L. kamiesmontana to be most closely related to L. lewisiana despite its position as sister to L. fabricii in the molecular phylogenetic study. Representative specimens SOUTH AFRICA. NORTHERN CAPE (Kamiesberg): Kamiesberg, Langkloof south of Welkom, ( AC), 20 Sept. 1994, Goldblatt & Manning (MO, NBG); Kamiesberg, near Farm Naartjiesdam, south of Karas, ( CA), 1 Sept. 2002, Goldblatt & Porter (MO, NBG); Kamiesberg, Rondefontein, slopes south of Nartjiesdam, 780 m, ( CA), 1 Sept. 1975, Oliver 5969 (NBG, PRE). 27. Lapeirousia fabricii (D.Delaroche) Ker Gawl. in Curtis s Botanical Magazine 31: sub t (1810); Goldblatt: 69 (1972). Ixia fabricii D.Delaroche: 18, no. 5 (1766). Plants mostly mm high, usually 4 6-branched; stem compressed, angled and 2-winged, wings irregularly serrated. Corm mm diam., broadly obconic; tunics brown, lightly ridged vertically, basal rim with short, spreading spines 1 2 mm long. Leaves several, lowermost longest, linear to falcate, ribbed, shorter or longer than stem; remaining leaves cauline, progressively shorter, not or barely ribbed, often subtending branches, upper becoming bractlike. Inflorescences (2 )4 8-flowered spikes; outer bract green or flushed red, (10 )14 20( 25) mm long, keeled, acute, margins often reddish, keel crisped or minutely serrate; inner bract transparent with 2 green keels, 1 / 2 to 2 / 3 as long as outer, forked at apex. Flowers zygomorphic, white to pale yellow or beige, purple or rarely pink, usually red or pink on reverse of tepals and tube, lower tepals each with red spot in midline, throat marked with fine red lines inside and out, decurrent on lower part of tube; perianth tube obliquely funnel-shaped, mm long, narrow and cylindric below, abruptly expanded into wide upper part up to 10 mm long; tepals usually unequal (rarely subequal), all straight below and continuing upper part of tube as a tepal cup, then spreading horizontally, upper 3 largest, mm (subsp. fabricii), mm (subspp. compressa and purpurascens), lower 3 with claws ± 4 mm long, limbs mm, each often with short, claw-like cusp arching over limb base. Stamens unilateral, arched; filaments 6 10 mm long (subspp. compressa and purpurascens) or mm long (subsp. fabricii), exserted 3 7 mm from upper part of tube and 1 3 mm from tepal cup; anthers 3 5 mm long, dark purple; pollen purple. Style arching over stamens; branches 3 5 mm long, usually dividing near middle of anthers, forked for 1 / 2 their length, recurved. Capsules oblong, mm long, apices of locules forming earlike lobes decurrent on low locular ridges. Seeds

90 82 S T R E L I T Z I A 35 (2015) globose, flattened at chalazal end, mm diam. Chromosome number 2n = 16. Flowering time: August to October depending on subspecies, usually fairly late in spring. Figures 27 & 28. Distribution and biology: widespread in the drier, western half of Western Cape and common in Namaqualand in Northern Cape, Lapeirousia fabricii extends from near Steinkopf in the north to Malmesbury in the south and inland to the western edges of the Tanqua Basin along the eastern foothills of the Swartruggens and Cederberg (Figure 26). Plants most often grow in rocky sandstone- or granite-derived soils in mountainous areas, but occasionally in deep, hard sand, or in crevices in rocks. The long-tubed flowers of the typical subspecies are known to be pollinated by long-proboscid flies (Goldblatt et al. 1995; Manning & Goldblatt 1996) and we infer the same pollination for subsp. compressa and subsp. purpurascens, probably using a different fly species in the latter. Diagnosis and relationships: the flowers of Lapeirousia fabricii are among the largest in the genus, with its long perianth tube and often wide throat. The perianth tube ranges from mm long and is usually expanded in the upper 5 10 mm in a short to long cup or gullet. The tepals are shortly clawed, particularly the lower three, which are united basally for ± 2 mm and the erect claws form the upper part of the floral cup. When fully open the lower and sometimes the upper tepal limbs spread at right angles to the tube, or the dorsal is suberect. The lower tepal claws often bear claw-like cusps that arch forward over the limb bases; these cusps are sometimes absent and represented by thickened median ridges. Vegetatively, L. fabricii has distinctive flattened, narrowly winged stems, the wings usually minutely crisped or serrated. The well-developed outer floral bracts are ribbed and have crisped to serrated keels. Variation across the range of Lapeirousia fabricii has prompted us to recognise three subspecies, the most common and widespread of which, subsp. fabricii, is centred in the northern half of its range. It has predominantly pale yellow to beige flowers with red markings, lower tepals about half as wide as the dorsal, and a perianth tube mostly mm long with a wide, deep throat up to 10 mm long. South and west of the Olifants River Mtns, plants that we treat as subsp. compressa have white or pale pink flowers with pink markings, lower tepals not or only slightly narrower than the dorsal, a perianth tube mm long, and a short, relatively narrow throat usually only ± 5 mm long. The third subspecies, subsp. purpurascens, is restricted to the dry interior valleys of the Cederberg and northern Swartruggens and has blue-purple flowers, lower tepals ± 1 / 2 as wide as the upper and a perianth tube mm long. Its flowers also differ from the typical in having a relatively shallow gullet ± 5 mm deep and correspondingly shorter filaments 7 9 mm long. History: few southern African species have been subjected to as much confusion as Lapeirousia fabricii. It was described by D. Delaroche (also de la Roche) in 1766, without illustration and perhaps without a preserved specimen, but, nevertheless, in considerable detail. Thunberg (1784), and later Jacquin (1794: t. 269) and other contemporary botanists, confused the species with L. anceps, which has similarly coloured but smaller flowers, and this name was applied to L. fabricii even by Baker (1896). The two were not distinguished from one another by most botanists, although Lamarck (1789) and Sweet (1830b) did recognise both L. fabricii and L. anceps, naming the larger-flowered species Gladiolus denticulatus and Peyrousia aculeata respectively. Few at the time seem to have associated Pourret s (1788) L. compressa, type of the genus, with either L. anceps or L. fabricii, even when Lapeirousia (or the variant Peyrousia) was recognised. Because L. anceps and L. fabricii were regarded as the same species by Baker (1892; 1896), Schlechter (1897) described L. serrulata for the large-flowered northern form of L. fabricii. Unravelling this web of confusion, Goldblatt (1972) recognised L. fabricii as distinct from L. anceps, but did not recognise infraspecific taxa, although he pointed out the differences between the northern populations, now subsp. fabricii, from those in the south of the range, now subsp. compressa. The populations we assign to subsp. purpurascens were not known at the time.

91 S T R E L I T Z I A 35 (2015) 83 B C D A E FIGURE 27. Lapeirousia fabricii subsp. compressa, Goldblatt & Manning A, plant; B, half flower; C, capsule. L. fabricii subsp. purpurascens, Goldblatt & Manning D, flower; E, half flower. Scale bar: 10 mm. Artist: John Manning.

92 84 S T R E L I T Z I A 35 (2015) FIGURE 28. Lapeirousia fabricii subsp. fabricii, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

93 S T R E L I T Z I A 35 (2015) 85 Key to subspecies 1a Flowers with deep gullet 9 13 mm long including limbs of lower tepals; perianth usually beige to pale yellow, or ± white, flushed red on reverse, with prominent red stripes in lower throat; filaments mm long a. subsp. fabricii 1b Flowers with shallow gullet ± 5 mm long; filaments 7 9 mm long: 2a Perianth white to pale or deep pink; anthers 3 4 mm long; lower tepals only slightly narrower than upper tepals and with or without tooth-like cusps b. subsp. compressa 2b Perianth blue-purple (rarely pink-purple) with red markings; anthers 4 5 mm long; lower tepals ± half as wide as upper tepals and always with short tooth-like cusp near limb base c. subsp. purpurascens 27a. subsp. fabricii Gladiolus denticulatus Lam.: 728 (1786), nom. illegit. superfl. pro Gladiolus anceps L.f. (1782). Lapeirousia denticulata G.H.M.Lawr.: 134 (1955), as nom. nov. pro G. denticulatus nom. illegit. Type: South Africa, without precise locality or collector, Herb. Lamarck (P, holo.!). Peyrousia aculeata Sweet: t 39 (1830b) [Lapeirousia aculeata Sweet: 396 (1827), name without description]. Meristostigma aculeatum (Sweet) Steud.: 130 (1841). Ovieda aculeata (Sweet) Klatt: 777 (1864). Lapeirousia anceps var. aculeata (Sweet) Baker: 156 (1877). Type: South Africa, without precise locality or collector, illustration in British Flower Garden ser. 2: t. 39 (1830). Lapeirousia serrulata Schltr.: 432 (1897). Type: South Africa, [Western Cape], Clanwilliam, Leipoldt 383 (B, holo.!; BOL!, SAM!, iso.). Flowers pale yellow or beige to ± white, often red or pink on reverse of tepals and tube, lower tepals each with red spot in midline and clawlike projection at top of claw, throat marked with fine red lines inside and out decurrent on lower part of tube; perianth tube mostly mm long, slender and cylindric below, expanded upper part 9 13 mm long; tepals unequal, straight below and continuing upper part of tube as a cup for ± 4 mm, then spreading, upper 3 tepals largest, mm, lower 3 tepals much narrower, mm. Stamens with filaments mm long, exserted ± 7 mm from tube and ± 3 mm from tepal cup; anthers 5 6 mm long. Style dividing between base and apex of anthers, style branches ± 5 mm long. Figure 28. Distribution: subsp. fabricii ranges from near Steinkopf in northern Namaqualand along the Bokkeveld Mtns to Clanwilliam and the Biedouw Valley, and along the Atlantic coast as far south as Darling (Figure 26). The flowers are adapted for pollination by long-proboscid flies. Recorded pollinators include the horse fly, Philoliche gulosa and the nemestrinid, Moegistorhynchus longirostris (Goldblatt et al. 1995; Manning & Goldblatt 1997). Representative specimens SOUTH AFRICA. NORTHERN CAPE (Springbok): Klipfontein, ( CA), Hutchinson 923 (K); hills at Concordia L. maximiliani Schltr. n. sp., ( CB), 20 Sept. 1897, Schlechter (B, COI, E, G, K, MO); between Brakwater and Kommagas, ( CD), Barker (NBG); near top of Spektakel Pass, ( DA), 11 Sept. 1993, Goldblatt & Manning 9711 (MO) (Hondeklipbaai): 45 km south of Springbok, ( BB), 15 Oct. 1976, Goldblatt 3068 (MO); 52 km from Garies to Wallekraal, ( BC), 28 Sept. 1976, Goldblatt 4215 (MO); Garies, ( DB), Thorne s.n. (NBG61563); Farm Roodeheuwel, 9 km west of Nariep, ( DC), 28 Sept. 1987, Reid 1275 (PRE) (Kamiesberg): border of Western Cape, slopes facing Swartdoorn River, ( CC), 24 Sept. 2008, Goldblatt & Porter (MO) (Calvinia): Grasberg road, northwest of Nieuwoudtville, ( AC), 13 Nov. 1974, Snijman 30 (NBG); Botterkloof Pass, ( CC), Hall 3877 (NBG); Soutpan, Calvinia, ( CD), 20 Nov. 1961, Lewis 5755 (NBG), 13 Oct. 1983, Goldblatt 7081 (MO). WESTERN CAPE (Lepelfontein): 40 km from Lutzville to Brand-se-Baai, ( BB), 15 Sept. 2008, Goldblatt & Porter (MO, NBG) (Vanrhynsdorp): Bitterfontein, ( AB), Salter 1584 (BM); Vredendal Commonage, ( DA), 25 Sept. 1970, Hall 3845 (HUS, NBG) (Calvinia): foot of Vanrhyns Pass, ( AC), Lewis 2903 (SAM) (Clanwilliam): Clanwilliam, ( BB), Leipoldt 383 (BOL, P, SAM); between Clanwilliam and Citrusdal, near Patrysberg, ( BD), 8 Nov. 1974, Boucher 2616 (MO, NBG) (Wuppertal): Brandewyn River, ( AA), 20 Sept. 1937, Barker 253 (NBG); 13 Sept. 1947, Compton (NBG); Pakhuis Pass, ( AA), 30 Sept. 1940, Barker 1223 (NBG); Welbedacht, Biedouw, ( AA), 22 Sept. 1952, Johnson 538 (MO, NBG), 21 Sept. 1952, Lewis 2511 (SAM); Algeria Forest Station, ( AC), 6 Oct. 1976, Botha & Coetzee 1723 (PRE); Cederberg, Matjiesrivier Nature Reserve, ( AD), 5 Oct. 1997, Lechmere-Oertel 871 (NBG); Farm Rockwood (Appalto), 1 km south of homestead, ( CA), 19 Oct. 2001, Hanekom 3341 (MO, NBG) (Cape Town): Malmesbury District, near Modder River, north of Bok River, ( AD), 22 Nov. 1970, Acocks (PRE); Waylands, Darling, ( AD), Thompson 82 (NBG); ± 10 km north of Malmesbury between farms Klipfontein and Saamstaan, ( BC), 6 Nov. 1982, Goldblatt 6714 (K, MO, PRE).

94 86 S T R E L I T Z I A 35 (2015) 27b. subsp. compressa (Pourr.) Goldblatt & J.C.Manning, stat. nov. Lapeirousia compressa Pourr.: 79 (1788). Ixia lapeirousia J.F.Gmel.: 108 (1791), nom. illegit. superfl. pro L. compressa Pourr. Type: South Africa, without locality, said to have been collected by Philibert Commerson in Isle de France [Mauritius], but probably by Pierre Sonnerat, his colleague, who spent a short time at the Cape en route to Mauritius (P: Herb. Lamarck, holo.!). Like subsp. fabricii in vegetative features. Flowers white to ivory or pale (rarely dark) pink, flushed pink to red on reverse, lower tepals with red markings at limb bases; perianth tube mm long, expanded in upper ± 5 mm into wide throat; tepals extending upward for ± 3 mm as a cup, spreading distally, dorsal mm, lower tepals mm. Stamens with filaments 6 10 mm long; anthers 3 4 mm long. Style dividing opposite lower to middle 1 / 3 of anthers, branches ± 3 mm long. Flowering time: September to mid-october, occasionally in early November. Figure 27A C. Distribution: subsp. compressa occurs locally in the upper Olifants River valley, but is more common south and west of the Olifants River Mtns, extending as far south as Moorreesburg. Isolated populations occur to the east along the margins of the Tanqua Basin from the interior Cederberg south to Karoo Poort (Figure 26). Diagnosis: usually recognised by the nearly subequal tepals and almost radially symmetric flower, subsp. compressa has a white to ivory or pink (rarely almost red) perianth, the lower tepals marked with pink to red near the base of the limbs. Vegetatively plants are identical to subsp. fabricii. The two subspecies intergrade in the mid- to upper Olifants River Valley at Piekeniers Kloof and south of Citrusdal where it can be difficult to assign plants to subspecies. Further north and to the west and south, plants can readily be recognised as subsp. fabricii and have a beige to almost yellow perianth and lower tepals ± half as wide as the upper. Hybrids between Lapeirousia fabricii subsp. compressa and L. jacquinii have been recorded south of Piketberg near the old bridge over the Berg River where flowering times of the two taxa overlap (Goldblatt & Manning 9955 MO). We have also seen hybrids between subsp. compressa and L. anceps at sites where both were in flower and growing together. These are among the few interspecific hybrids known in the genus. Selected specimens SOUTH AFRICA. WESTERN CAPE (Clanwilliam): south of Lamberts Bay, ( AB), 20 Sept. 1953, Taylor 4009 (NBG); Zwartboskraal, northeast of Boekenberg, ( BC), 7 Sept. 1972, Oliver 3885 (MO, NBG, PRE); Swartrug, Piketberg, east of Engelsman-se-Baken, flats, ( DA), 10 Oct. 1977, Thompson 3550 (NBG); Piketberg, Moutonsvlei, ( DA), 21 Aug. 1984, Goldblatt 7201 (MO); road junction south of Het Kruis, ( DB), 20 Sept. 1975, Wisura 3502 (NBG); top of Piekenierskloof Pass, ( DB), 10 Sept. 1993, Goldblatt & Manning 9695 (MO, NBG); Berg River near Piketberg, ( DB), 10 Sept. 1894, Schlechter 5265 (BM, BOL, COI, G, GRA, K, MO); 11 Sept. 1981, Goldblatt 6213 (MO, PRE, S, US); Middelpos, west of Piketberg, 250 m, ( DC), 10 Oct. 1984, Morley 241 (PRE); Piketberg to Sauer, ( DC), 20 Sept. 1974, Goldblatt 2728 (MO); south bank of Berg River, west of old bridge, ( DC), 1 Sept. 1992, Goldblatt & Manning 9337 (MO, NBG, PRE) (Wuppertal): Olifants River Mtns, Dasklip Pass, rocky shale slope, ( CC), 30 Sept. 1972, Oliver 3998 (NBG); slopes of Dasklip Pass, ( CC), 8 Nov. 1976, Goldblatt 4436 (MO) (Cape Town): Oshoekkop, north of Moorreesburg, shale hillside, ( BA), 17 Sept. 1982, Van Zyl 3287 (NBG) (Worcester): De Hoek, ( AA), 28 Sept. 1952, Lewis s.n. or 2656 (MO, SAM); Barker 2571 (NBG); 2 miles [3 km] north of Karoo Poort, ( BC), 11 Nov. 1974, Snijman 12 (NBG). 27c. subsp. purpurascens Goldblatt & Manning, subsp. nov. TYPE. South Africa, Western Cape, 3219 (Wuppertal): rocky sandstone slope 2 km south of Wuppertal, ( AC), 25 Aug. 2009, Goldblatt & Manning (NBG, holo.; K, MO, PRE, iso.). Plants mostly mm high. Flowers pale to deep purple, darker on reverse of tepals and tube, lower tepals each with dark red ± triangular mark in lower midline and with short yellow, claw-like cusp near base, throat dark turquoise to violet, often with pale narrow lines; perianth tube ± straight, slender, mm long, expanded in upper ± 5 mm; tepals unequal (rarely subequal), upper three mm, spreading distally; lower 3 erect below continuing upper part of tube as tepal cup for ± 4 mm, spreading distally, mm. Stamens with filaments ± 7 8 mm long, dark

95 S T R E L I T Z I A 35 (2015) 87 turquoise, exserted 2 3 mm from tube; anthers 4 5 mm long. Style suberect, usually dividing between middle and apex of anthers; branches ± 3.5 mm long. Flowering time: mid-august to mid-september, thus early to mid-spring. Figure 27D, E. Distribution: subsp. purpurascens has a narrow range in the interior valleys of the Western Cape, extending from Wuppertal in the north through the drier valleys of the interior northern Cederberg and adjacent Swartruggens (Figure 26). The flowers, with their dark blue- or rarely red-purple colour and long perianth tube, combined with August to mid-september blooming, suggest pollination by the long-proboscid fly Prosoeca peringueyi (Goldblatt et al. 1995), but to date there are no observations on pollination of the subspecies. Diagnosis: subsp. purpurascens is distinguished by its predominantly purple flowers with darker purple markings on the lower tepals, each of which bears a short, yellow, claw-like cusp near the base, and the short, dark violet to turquoise throat ± 5 mm long. Flowers of subsp. fabricii, which occurs within the range of subsp. purpurascens but blooms two to three weeks later, are creamy yellow with bright red markings on the lower tepals. The throat in subsp. fabricii is up to 10 mm deep and the perianth tube is usually longer, mm vs mm in subsp. purpurascens. Flowering in subsp. fabricii usually begins in mid-september, continuing into October, sometimes even as late as November, so that the two subspecies are isolated by flowering time as well as pollinator preference. The habitat is essentially the same for both rocky sandstone slopes although in Namaqualand, subsp. fabricii grows on granite-derived soils and granite outcrops. We hypothesise that subsp. purpurascens is a local segregate of the widespread subsp. fabricii that has shifted pollination from the Philoliche Moegistorhynchus system to the Prosoeca peringueyi system (Goldblatt & Manning 2000). Species belonging to the latter guild have red to purple flowers with pale nectar guides, and frequently a somewhat shorter perianth tube than those pollinated by Philoliche and Moegistorhynchus, exactly the floral features of subsp. purpurascens. We know of no intermediates between subsp. fabricii and subsp. purpurascens, but have chosen subspecies rank for the two taxa because of the weak morphological differences between them, primarily perianth colour and gullet length. Representative specimens SOUTH AFRICA. WESTERN CAPE (Wuppertal): near Wuppertal, Stormvlei, stony ground, ( AA), 11 Sept. 1897, Leipoldt 542 (NBG); Wuppertal, ( AA), 20 Aug. 1959, Jackson s.n. (NBG); Ceres Karoo, Karretjies, west of Elandsvlei, ( AB), 12 Sept. 2006, Bruyns (NBG); Swartruggens, Farm Doornfontein, rocky sandstone slopes, ( DA), 12 Aug. 2009, Manning 3228 (NBG).

96 88 S T R E L I T Z I A 35 (2015) Systematics of Codonorhiza Codonorhiza Goldblatt & J.C.Manning, gen. nov. Type: C. corymbosa (L.) Goldblatt & Manning. Lapeirousia sect. Fastigiatae [as Fastigiata] Gold blatt in Contributions from the Bolus Herbarium 4: 15 (1972). [L. subg. Paniculatae Goldblatt & J.C.Manning sect. Fastigiatae (Goldblatt) Goldblatt & J.C.Manning 77: 373 (1990).] Type: L. corymbosa (L.) Ker Gawl. (= Codonorhiza corymbosa (L.) Goldblatt & J.C.Manning). Deciduous geophytes. Corm flat-based, broadly obconic (± triangular in profile), rooting from base and axillary in origin; tunics of dark brown to blackish, densely compacted fibres. Foliage leaves 2 or 3, lowermost inserted at ground level, longest plane with ± prominent main vein, upper leaves progressively smaller. Stem aerial, compressed and angled to winged, simple or repeatedly branched. Inflorescence either a spike or a ± flat-topped, false panicle with sessile flowers; bracts 2, opposed, green, firm textured, usually obtuse to ± truncate, inner ± as long as outer, sometimes notched apically. Flowers radially symmetric or zygomorphic, long-lived, salver- or bowlshaped, blue, purple, red, pink or white, usually with contrasting darker or lighter markings, sometimes sweetly scented; perianth tube ± cylindric or funnel-shaped, short or elongate; tepals subequal or unequal with outer whorl larger. Stamens symmetrically disposed or unilateral and arcuate; filaments slender, free; anthers oblong-linear, dehiscence longitudinal. Ovary globose; style filiform, branches usually forked for ± 1 / 2 their length, occasionally barely forked at tips. Capsules cartilaginous, globose to ± top-shaped and 3-lobed. Seeds ovoid( orbicular), flattened at chalazal end, with reticulate to rugulose sculpturing; funicular collar prominent, surface cells ± flat (obscurely domed). Basic chromosome number x = 10. Species: 7; restricted to Western Cape, South Africa. Key to species 1a Inflorescence a spike; flowers white to pink, lower tepals with purple or red marks in basal half; perianth tube cylindric C. falcata 1b Inflorescence a flat-topped, false panicle with ultimate branches bearing 1 5 sessile flowers; flowers predominantly blue, yellow or maroon to light brown; perianth tube cylindric or funnel-shaped: 2a Perianth tube (14 )18 20 mm long; tepals 7 8 mm long C. pillansii 2b Perianth tube 3 12 mm long; tepals other: 3a Flowers dull yellow, creamy brown or maroon; perianth tube cylindric 8 10 mm long, ± twice as long as tepals C. micrantha 3b Flowers blue to violet or pale yellow; perianth tube funnel-shaped, shorter than tepals: 4a Flowers zygomorphic with unilateral stamens; perianth blue to violet, with white or dark, blue-black, contrasting markings: 5a Flowers medium to dark blue( violet) with white, bracket-shaped band in basal 1 / 3 of lower tepals; anthers 3 4 mm long; pollen blue or grey; tepals mm long C. elandsmontana 5b Flowers deep blue( violet) with dark blue, red-purple or blackish mark (sometimes outlined in white) at base of lower tepals; anthers 3 5 mm long; pollen usually dark red-brown, blue or pale grey; tepals mm long C. azurea

97 S T R E L I T Z I A 35 (2015) 89 4b Flowers actinomorphic with stamens ascending; perianth blue (rarely white) or pale yellow: 6a Perianth light to dark blue with white, star-shaped mark in centre outlined in darker colour; tepals 6 10 mm long; filaments 5 6 mm long C. corymbosa 6b Perianth pale yellow, each tepal with small dark mark in lower midline; tepals mm long; filaments ± 10 mm long C. fastigiata Spp Codonorhiza falcata (L.f.) Goldblatt & J.C.Manning, comb. nov. Gladiolus falcatus L.f., Supplementum plantarum: 96 (1782). Lapeirousia falcata (L.f.) Ker Gawl.: 238 (1804); Goldblatt: 28 (1972). Ovieda falcata (L.f.) Spreng.: 147 (1824). Peyrousia falcata (L.f.) Sweet: 499 (1830a). Meristostigma falcatum (L.f.) A.Dietr.: 594 (1833). Type: South Africa, without precise locality, Thunberg s.n. UPS-THUNB1025 (UPS- THUNB, holo.!). Lapeirousia pappei Baker: 170 (1892). Type: South Africa, [Western Cape], Winterhoeksberg, Tulbagh, Pappe s.n. (K, holo!; BOL!, iso.). Plants short and compact to fairly slender, seldom exceeding 100 mm, simple or with up to 3 branches from near base, stem terete. Corm ± 8 mm diam., broadly obconic; tunics dark brown to black, rugose, split longitudinally at base and apex. Leaves usually 2, lowermost basal, longest, falcate, ± plane with main vein slightly raised, ± leathery, mostly 5 8 mm wide; upper leaf much shorter. Inflorescence a spike, with up to 12 flowers in 2 ranks; bracts green with red margins, outer ± 6 mm long, inner 4 5 mm long. Flowers zygomorphic, pale pink to nearly white, lower 3 tepals each with dark pink to mauve or red spot in lower midline, unscented; perianth tube ± cylindric, mm long, slightly wider in upper 3 4 mm; tepals ovate-oblong 7 8 ± 3 mm. Stamens unilateral, ± erect; filaments ± 6 mm long, exserted ± 3 mm; anthers ± 1.2 mm long, white, pollen white. Style erect, dividing shortly below apex of filaments, branches ± 2.3 mm long, deeply divided, spreading between anthers. Capsules top-shaped and 3-angled, 4 5 mm long. Seeds ovoid-orbicular, flattened at chalazal end, brown, ± mm, reticulate, funicular collar up to 0.3 mm long. Chromosome number 2n = 20. Flowering time: mainly September and October at lower elevations but as late as mid-december at high elevations. Distribution and biology: like most other members of the genus, Codonorhiza falcata has a relatively narrow range, extending from Matroosberg and Lakenvlei, east of Ceres, through the Agterwitsenberg Vlakte and Cold Bokkeveld to Elandskloof, south of Citrusdal (Figure 29). A collection, Templemann sub MacOwan in Herb. Norm. Austr. Afr. 1656, said to be from the Swartberg at Caledon, is almost certainly not from there. Plants occur exclusively on sandstone substrate and often in crevices in rock outcrops or in seeps on sandstone pavement. Most records are from middle elevations between 800 and m, but there are collections from Matroosberg at m, also the most easterly station. Additional collecting in the Hex River Mtns will likely yield additional records. Flowering is stimulated by fire but plants may be found in bloom in open sites several years after a fire. Pollination in the species has not been studied. Diagnosis and relationships: both morphology, especially the corms, and molecular data FIGURE 29. Distribution of Codonorhiza falcata, ; C. fastigiata, ; C. pillansii,.

98 90 S T R E L I T Z I A 35 (2015) confirm the relationship of Codonorhiza falcata to the C. corymbosa complex. In the DNA-based phylogeny (Figure 1), C. falcata is consistently retrieved as sister to the remaining species of the genus. Unlike all other Codonorhiza species, the inflorescence is a spike rather than a flat-topped false panicle, and we infer that a spike is the ancestral condition for the genus on the basis of outgroup comparison since all genera except Pillansia of the sister clade have spicate inflorescences. The blackish, rough-textured corm tunics, plane, falcate leaf and green floral bracts with red margins accord exactly with the C. corymbosa complex, as do the small, ± ovoid seeds with reticulate sculpturing and prominent funicular collar. The flowers are pale pink to ± white with dark pink to mauve or red markings on the lower tepals, colours that are unique in the genus. C. falcata is one of only three species with a cylindrical perianth tube, but the other two, L. micrantha and L. pillansii, are immediately distinguished by their flat-topped inflorescences and flowers of different colour. Described by the younger Linnaeus in 1782, as Gladiolus falcatus, the species was based largely, if not entirely, on a Thunberg collection and probably in part on Thunberg s manuscript notes. Early confusion with Freesia verrucosa and its illegitimate synonym, Anomatheca juncea, which is superficially similar to Codonorhiza falcata, was the result of their flowers of similar size and proportions, but they differ in almost all vegetative features. Linnaeus fil. confused them and a specimen of F. verrucosa in the Linnean Herbarium annotated by him has a question mark against the name Gladiolus falcatus. Thunberg s (1784) more detailed account in the Dissertatio de Gladiolo clearly applies only to his collection (Goldblatt 1972). G. falcatus was transferred to Lapeirousia by Ker Gawler (1804). Later in the 19th century the name L. falcata was abandoned in favour of the later synonym, L. pappei (Baker 1892). Representative specimens: SOUTH AFRICA. WESTERN CAPE 3219 (Wuppertal): Elandskloof, ( CA), Lewis s.n (BOL 30699); 29 Sept. 1944, Barker 3081 (NBG); northern Cold Bokkeveld, flats at west base of Geelberg, 2400 ft [731 m], ( CA), 2 Oct. 1972, Thompson 1547 (PRE); sandy plateau, Twenty Four Rivers Mtns, ( CC), 22 Oct. 1947, Esterhuysen (BOL, NBG); Cold Bokkeveld, near De Keur, ( CD), 1 Oct. 1944, Barker 3128 (NBG); Cold Bokkeveld, sandstone rocks between Rosendal and Liberty, ( CD), 12 Oct. 2000, Goldblatt (MO); Groenfontein, Zeekoegat, west of Riet River, ( DC), 2 Oct. 2002, Stobie 2 (NBG) (Worcester): Agterwitsenberg Vlakte, ( AA), Mauve 4078 (PRE); Gydo Pass, ( AB), 1 Oct. 1944, Barker 3128 (NBG); 20 Sept. 1993, Goldblatt & Manning 9742 (MO); Ceres Nature Reserve, top of Michell s Pass, ( AD), 29 Sept. 2009, Goldblatt & Porter (MO, NBG); Lakenvlei, ( BC), Nov. 1917, Phillips 2069 (SAM); Matroosberg, near beacon, m, ( BC), 10 Dec. 1981, Kotze 101 (NBG); Esterhuysen s.n. (MO). 2. Codonorhiza micrantha (E.Mey. ex Klatt) Goldblatt & J.C.Manning, comb. nov. Ovieda micrantha E.Mey. ex Klatt in Linnaea 32: 781 (1864). Lapeirousia micrantha (E.Mey. ex Klatt) Baker: 156 (1877); Goldblatt: 27 (1972). Types: South Africa, [Western Cape], Worcester, waterfall, Ecklon & Zeyher Irid 253 (K, lecto.!, designated here; MO!, isolecto.). [South Africa, [Western Cape], Gifberg, Drège s.n. (SAM, syn.)]. Lapeirousia manuleiflora Eckl.: 31 (1827), nom. nud. Meristostigma manuleiflorum (Eckl.) Steud.: 130 (1841). Plants mostly mm high, repeatedly branched, branches subtended by bracts (1 )2 5 mm long, these sometimes leaf-like below, stem flexuose, compressed and 2 4-winged. Corm mm diam., broadly obconic; tunics dark brown to blackish, basal rim ± crenate, splitting vertically from base. Leaves 2 or 3, lowermost basal, ± falcate, mm, margins usually undulate or slightly crisped, main vein prominent, upper leaves inserted near middle of stem, smaller than basal, not or hardly sheathing at base. Inflorescence a congested, ± flat-topped false panicle, flowers crowded at branch tips, ultimate branchlets 1 3-flowered; bracts green, ± leathery, 2 3 mm long, margins red, inner ± as long as outer, obtuse or inner slightly emarginate. Flowers weakly zygomorphic with perianth radially symmetric and stamens and style unilateral, dull yellow to buff or brown to dull maroon, acrid-sweet or clove scented; perianth tube ± cylindric, 8 10 mm long; tepals subequal, ± elliptic, obtuse, 3 5 ± mm. Stamens unilateral, ± erect; filaments ± 3.5 mm long, exserted ± 1.5 mm; anthers mm long, pollen dull yellow to buff. Style dividing between base and upper 1 / 3 of anthers, style branches ± 1 mm long,

99 S T R E L I T Z I A 35 (2015) 91 FIGURE 30. Codonorhiza micrantha, Goldblatt 10795C. Scale bar: 10 mm. Artist: John Manning.

100 92 S T R E L I T Z I A 35 (2015) usually divided for up to 1 / 2 their length or undivided. Capsules top-shaped, triangular, 5 6 mm long, with up to 6 seeds per locule. Seeds ± ovoid, dark brown, mm long, funicular appendage up to 0.5 mm long, surface reticulate. Chromosome number 2n = 20. Flowering time: late September to December. Figure 30. Distribution and biology: the most widespread species of Codonorhiza, C. micrantha, extends from the Gifberg Matsikamma mountain complex in the northwest to the Langeberg at Garcia s Pass in the southeast, thus embracing the major part of the sandstone mountains of the Cape Floristic Region (Figure 31). Despite the wide range, this inconspicuous species is seldom seen, for it rarely flowers except in the first and second season after fire. We infer settling moth pollination for the species because of the dull flower colour and heavy, often clove-like floral odour that are matched in some moth-pollinated species of Gladiolus (Goldblatt & Manning 2002). Diagnosis and relationships: Codonorhiza micrantha stands out among the species with crowded, flat-topped inflorescences in its cylindrical perianth tube, 8 10 mm long, and small, yellow to brown or maroon, strongly scented flowers. In the molecular phylogeny (Figure 1) C. micrantha is retrieved as sister to C. corymbosa. FIGURE 31. Distribution of Codonorhiza azurea, ; C. micrantha,. History: although first recorded by William Burchell in late 1814 near Riversdale, Codonorhiza micrantha was based on the later collections of J.-F. Drège (circa 1830) and Ecklon & Zeyher (before 1827). The Drège collection was distributed under the manuscript name Ovieda micrantha and was so listed by Meyer (1843) in the publication that dealt with the distribution ranges of plants collected by Drège. Klatt (1864) formally named the species based on both the Drège and Ecklon & Zeyher collections, assigning it to the illegitimate genus Ovieda (= Lapeirousia). The very apt epithet manuleiflora (the flowers do indeed closely resemble those of several, smallflowered Manulea species) was used by Ecklon (1827), who assigned the species to Lapeirousia, but without any description, the name is thus invalid. Representative specimens: SOUTH AFRICA. WESTERN CAPE (Vanrhynsdorp): Matsikamma Mtns, north of Farm Die Vlei, ( DB), 11 Nov. 1985, Snijman 959 (NBG); Gifberg slopes at Farm Colinshoek, ( DD), 11 Sept. 2009, Goldblatt & Porter (MO) (Clanwilliam): Kransvlei, Clanwilliam, ( BB), Leipoldt 4427 (BOL); Piketberg, Moutonsvlei, ( DC), Nov. 2008, Ebrahim s.n. (NBG); Piketberg, Sandleegte, ( DC), 5 Nov. 1973, Linder 93 (BOL) (Wuppertal): Biedouw Mtns, ( AA), Nov. 1929, Thode A2140 (NBG); 3 km west of Algeria, Clanwilliam road, ( AC), 16 Nov (fl. & fr.), Goldblatt 5150 (MO, S); Cederberg State Forest, research site 12, six months after fire ( AC), 10 Nov. 1981, Le Maitre 295 (PRE); Cold Bokkeveld Mtns, Keerom, ( CC), 4 Dec. 1950, Esterhuysen (BOL, MO, NBG) (Worcester): Roodezand Pass, Tulbagh, stony clay slope, ( AA), 22 Oct. 1994, Oliver (NBG); Matroosberg near Lakenvlei, ( BC), Phillips 2068 (SAM); Goudini, Farm Groenvlei, ( CB), 6 Nov. 1981, Walters 2552 (NBG); Franschhoek Pass, eastern end on south-facing slopes burned last summer, ( CC), Goldblatt 8580 (MO) (Ladismith): Langeberg slopes, near Swellendam, ( CD), 5 Nov. 1962, Taylor 4224 (NBG, PRE) (Simonstown): Kogelberg, northern slopes of main peak, after fire, 490 m, ( BB), 21 Nov. 1990, Oliver 9790 (NBG, PRE) (Caledon): Nuweberg, in two year old fynbos, ( AA), 11 Nov. 1982, Viviers 759 (NBG, PRE). Houw Hoek, ( AA), 24 Nov. 1896, Schlechter 9397 (BM, BOL, GRA, K, MO); top of Houw Hoek Pass, ( AA), 20 Nov. 1958, Lewis 5313 (NBG); slopes southeast of Villiersdorp, burned last [previous] summer, ( AB), 25 Oct. 1997, Goldblatt 10795C (MO); Vogelgat, Psoralea Gully, ( AD), 10 Nov. 1982, Williams 3355 (NBG); Hermanus, Vogelgat Nature Reserve, after fire, 3 Dec. 1975, Rourke 1508 (MO, NBG); Bredasdorp Mtns, recently burnt north slope, ( BD), 21 Nov. 1970, Barker (MO, NBG) (Riversdale): between Little Vet River and Kampscheberg, Riversdale, ( AB), Burchell 6883 (K).

101 S T R E L I T Z I A 35 (2015) Codonorhiza pillansii Goldblatt & J.C.Man ning, sp. nov. TYPE. South Africa, Western Cape, 3418 (Simonstown): Cape Peninsula, western edge of plateau west of Buffels Bay, ( AD), 22 Nov. 1921, Pillans 8928 (BOL, holo.!; MO!, iso.). Plants mm high, repeatedly branched, upper branches subtended by bracts up to 10 mm long, stem ± flexuose, compressed and 2-winged. Corm ± 10 mm diam., broadly obconic; tunics blackish, splitting vertically at base and apex. Leaves 2 or 3, lowermost basal and longest, usually falcate, mm, acute, with prominent main vein, margins plane, slightly thickened, not or hardly sheathing at base; upper leaves similar, inserted in middle of stem, mm long. Inflorescence a congested, flattopped false panicle, flowers crowded at branch tips, ultimate branchlets mostly 2- or 3-flowered; bracts 5 7 mm long, green, leathery, inner ± as long as outer or slightly longer, obtuse or inner ± emarginate. Flowers zygomorphic, dark blue, evidently uniformly coloured, scent unknown; perianth tube ± cylindric, (14 )18 20 mm long, ± 1 mm diam. at base, ± 2 mm diam. at mouth; tepals 7 8 ± 3 mm. Stamens unilateral; filaments inserted ± 4 mm below mouth of tube, 7 8 mm long, exserted 3 4 mm; anthers ± 3 mm long, pollen colour unknown. Style dividing opposite middle and apex of anthers, ± 2.5 mm long, divided for ± 1 / 2 their length. Capsules topshaped, 3-lobed, ± 5 mm long (mature capsules and seeds unknown). Chromosome number unknown. Flowering time: October and November. Distribution and biology: poorly understood, Codonorhiza pillansii is known from several plants collected in 1921 by N.S. Pillans in Cape Point Nature Reserve. The area where the plants were said to have been collected is low fynbos on sandy, stony ground. We suspect plants flower only after fire (Figure 29). The long perianth tube, up to 20 mm, suggests that the flowers are adapted for pollination by an insect with a relatively long proboscis. Diagnosis and relationships: it is with some hesitation that we recognise here a particularly unusual population of Codonorhiza said to be from the west end of the plateau west of Buffels Bay on the southern Cape Peninsula as a separate species, C. pillansii. Plants collected there by N.S. Pillans in November 1921, but incorporated into the Bolus Herbarium only after publication of a revision of Lapeirousia (Goldblatt 1972), resemble C. corymbosa in general appearance, but the flowers have a slender tube (14 )18 20 mm long. We have determined by dissection that the filaments are unilateral and that the tepals are subequal in size and evidently also in disposition. The tepals appear uniformly coloured, which accords with Pillans s comment flowers blue. These plants clearly represent a distinct variant, which we describe as C. pillansii, honouring N.S. Pillans who discovered the species. Pillans s locality was burned in the summer of 1991, providing ideal conditions for spring flowering of Codonorhiza, but we could locate only typical C. corymbosa on a visit there. Until more material can be found we remain ignorant of the biology and several details of the morphology of C. pillansii. 4. Codonorhiza azurea (Eckl. ex Baker) Goldblatt & J.C.Manning, comb. nov. Meristostigma azureum Eckl. ex Steud.: 131 (1841), nom. nud. Lapeirousia corymbosa var. azurea Eckl. ex Baker: 90 (1896) [L. azurea Eckl.: 31 (1827), nom. nud.]. Lapeirousia azurea (Eckl. ex Baker) Goldblatt in Goldblatt & Manning: 333 (1992). Type: South Africa, [Western Cape], between Paarl and Pont, Drège 8509a (K, lecto.!, designated by Goldblatt & Manning: 334 (1992); BM!, G!, MO!, P!, isolecto.). Plants ( 260) mm high, 5 7- branched, branches subtended by bracts (1 ) 2 5 mm long, those below appearing leaf-like, stem compressed and winged. Corms mm in diam.; tunics blackish, basal rim split vertically. Leaves 2, lowermost inserted at ground level, largest, falcate, ( 240) (6 )10 15 mm, margins undulate and crisped, obtuse to subapiculate, with prominent main vein and a pair of secondary veins (rarely one) on either side almost as prominent, second leaf inserted near middle of stem, mm long, oblong lanceolate, margins ± straight, not or hardly sheathing at base. Inflorescence a congested, flat-topped false panicle, flowers crowded at branch tips, ultimate branchlets 1- or 2-flowered; bracts green, firm-textured, 9 11 mm long, outer folded in midline, margins often red, inner obtuse or

102 94 S T R E L I T Z I A 35 (2015) slightly emarginate. Flowers zygomorphic, tilted backward, dark blue to violet, with triangular, dark blue-black or red-purple blotch at base of lower 3 and sometimes adjacent upper lateral tepals; perianth tube funnel-shaped, mm long; tepals elliptic-ovate, obtuse, mostly mm. Stamens unilateral, ascending or extended horizontally due to adaxial tilting of flower, mm long, exserted for 8 10 mm; anthers 3 5 mm long, pollen dark blue or redbrown. Style unilateral, dividing opposite upper 1 / 3 of anthers; branches mm long, divided for ± 1 / 2 their length. Capsules ± top-shaped and 3-lobed, ± 6 mm long, with up to 8 seeds per locule. Seeds red-brown, ovoid, truncate at chalazal end, rugulo-reticulate, ± mm, with funicular appendage mm long. Chromosome number 2n = B. Flowering time: late September to late October. Figure 32. Distribution and biology: Lapeirousia azurea is restricted to lowland sites in the southwestern Cape between Paarl, Mamre and Riebeeck Kasteel north of Malmesbury (Figure 31). It favours granite-derived gravel soils rather than the shalederived clay soils, also common in the area. Because of the relatively high fertility of these soils this habitat is now extensively cultivated, and L. azurea has become increasingly rare. In 1992 we wrote that it was probably not in immediate danger of extinction as it was known in sites away from cultivation. Some of these are now lost to urban expansion and some of its remaining sites are being developed for housing. We suspect that the dark-coloured flowers with even darker, blue-black or red-purple markings are adapted for pollination by hopliine beetles. At a site in Malmesbury large, buff-brown hopliines were the only insects we have seen on the flowers, using the flowers as sites of assembly and mating. Goldblatt et al. (1995) also reported visits to flowers of C. azurea (as Lapeirousia) by the anthophorine bee, Anthophora diversipes, suggesting a possible bimodal pollination system, using both large-bodied bees and hopliine beetles. Diagnosis and relationships: although initially treated taxonomically as a variety of Codonorhiza corymbosa, C. azurea is distinguished by its larger, zygomorphic, intensely dark blue flower bearing large black or dark red blotches on the adaxial tepals. The perianth tube is 8 12 mm long and the tepals (12 )16 20 mm long. The flower is tilted back toward the axis, especially in crowded inflorescences, when the posterior (adaxial) tepals, usually erect, are tilted out of position and come to lie horizontally, thus below the abaxial (lower) tepals. This type of floral presentation is not unique in Codonorhiza, but also occurs in C. elandsmontana. Although united with C. fastigiata (as Lapeirousia corymbosa subsp. fastigiata) by Goldblatt (1972), molecular data (Figure 1) suggest that C. azurea is not directly related to C. fastigiata despite their comparably large perianth. In the molecular phylogeny (Figure 1) C. azurea is sister to C. corymbosa + C. micrantha. History: Codonorhiza azurea was figured as early as 1791 by Jacquin in his Icones plantarum rariorum under the name Ixia corymbosa in a composite painting that included true C. corymbosa as well. The earliest extant herbarium records appear to be the collections of Ecklon & Zeyher, made in the late 1820s, and those made by J.-F. Drège at about the same time or shortly thereafter. It is not entirely clear whether Ecklon considered L. azurea distinct from L. corymbosa, for although he mentioned both names in his Topographisches Verzeichniss (1827), L. azurea without description, his collections of both species bear the latter manuscript name. C. azurea was regarded as a variety of C. corymbosa (as Lapeirousia) by Baker (1896) and as conspecific with C. corymbosa subsp. fastigiata by Goldblatt (1972). Only recently have sufficient specimens become available to help establish the status and rank of this plant. Reasons for treating C. azurea and C. fastigiata as separate species are discussed in detail under the latter. Representative specimens SOUTH AFRICA. WESTERN CAPE (Cape Town): Darling, ( AD), 27 Sept. 1890, Guthrie 2069 (NBG); Groene Kloof [Mamre], ( CB), MacOwan 2280 (SAM); commonage at Malmesbury, ( BC), 14 Sept. 1953, Lewis 3644 (SAM); Barker 8047 (NBG); 13 Sept. 1981, Goldblatt 6279 (MO); 28 Oct (fr.), Goldblatt 8016 (MO); hills around Malmesbury, ( BC), 1 Oct. 1892, Schlechter 1616 (B, G, GRA, P); Riebeeck Kasteel, ( BD), Oct. 1951, Pillans (G, MO); 18 Oct. 1941, Barker 1311 (NBG); top of Bothmaskloof, Riebeeck Kasteel, ( BD), Esterhuysen 6033 (BOL); clay flats near Paarl, ( DB), 24 Sept. 1974, Goldblatt 2740 (MO, NBG); north slopes of Paardeberg, ( DB),

103 S T R E L I T Z I A 35 (2015) 95 FIGURE 32. Codonorhiza azurea, Goldblatt Scale bar: 10 mm. Artist: John Manning.

104 96 S T R E L I T Z I A 35 (2015) Goldblatt 647 (BOL); east slopes of Paardeberg, ( DB), Pillans 7785 (BOL); Slent Farm, southeast end of Paardeberg, ( DB), 28 Sept. 1999, Goldblatt (MO); hillside near Joostenberg, ( DD), Esterhuysen (BOL). 5. Codonorhiza elandsmontana Goldblatt & J.C.Manning, sp. nov. Type: South Africa, Western Cape, 3319 (Worcester): foot of Elandskloof Mtns, Elandsberg Nature Reserve, stony ground, ( AC), 11 Sept. 2010, Goldblatt & Manning (NBG, holo.; MO, PRE, iso.). Plants mm high, several-branched, branches subtended by bracts (1 )2 5 mm long, or sometimes leaf-like below and longer; stem ± flexuose, compressed and 2-winged. Corm obconic, mm diam.; tunics blackish, splitting vertically from base and apex. Leaves 2(3), lower 1(2) basal, usually falcate (lanceolate), ( 400) mm, subacute to acute, with prominent main vein and 1 or pair of secondary veins on either side also raised; margins usually ± straight to slightly undulate, rarely slightly crisped; uppermost leaf inserted near middle of stem, mm long, narrowly oblong, not or hardly sheathing at base. Inflorescence a congested, flat-topped false panicle, ultimate branchlets 1- or 2-flowered; bracts green, leathery, 4 7 mm long, usually red distally and slightly warty, outer obtuse to truncate, keeled, inner ± obtuse to slightly emarginate. Flowers zygomorphic, tilted backward with adaxial (dorsal and upper lateral) tepals often nearly horizontal, light to dark blue with white throat and basal 1 / 2 of abaxial tepals with white, bracket-shaped band, not scented; perianth tube funnel-shaped, 8 10 mm long; tepals elliptic, obtuse, mm. Stamens ± unilateral, ± horizontal and facing upward; filaments 9 10 mm long, exserted 7 8 mm; anthers mm long, pollen blue to blue-grey, or purple. Style unilateral, dividing opposite or shortly beyond anther apices, held below stamens, style branches mm long, divided for ± 1 / 2 their length. Capsules top-shaped, 3-lobed, ± 5 mm long, with up to 6 seeds per locule. Seeds ovoid, reticulately sculptured, ± flattened to concave at chalazal end, dark brown, ± mm; funicular collar up to 0.3 mm long. Chromosome number unknown. Flowering time: late September and October. Figure 33. Distribution and biology: locally common in the Tulbagh Valley, Codonorhiza elandsmontana extends from Tulbagh eastward through the upper Breede River Valley to Botha and westward to Gouda and the lower slopes of the Elands kloof Mtns as far south as Bain s Kloof (Figure 34). Plants favour rocky ground, often in clay or clay loam. Plants at Elandsberg were actively visited by the tabanid fly, Philoliche angulata, which has a body ± 17 mm long and a proboscis ± half as long. These flies carried visible loads of the pale blue-grey pollen of C. elandsmontana and must be regarded as legitimate pollinators. Diagnosis and relationships: long confused with Codonorhiza corymbosa or sometimes with C. azurea, both of which it broadly resembles, especially in dry herbariums specimens, C. elandsmontana differs from C. corymbosa in the zygomorphic flowers with unilateral stamens and white, bracket-shaped markings only on the three abaxial tepals. Other floral details include the slightly longer perianth tube, 8 10 mm long vs. 4 7(8) mm long and the larger tepals, mm long vs mm long in C. corymbosa. Flowers of C. corymbosa are strictly actinomorphic, with all six tepals bearing white, bracket-shaped markings. C. elandsmontana may also be confused with C. azurea and C. fastigiata, which are similar in stature and have a broad basal leaf with undulate to crisped margins. The larger flowers of C. azurea are zygomorphic like those of C. elandsmontana, but have a perianth tube mm long and tepals mostly mm and the three abaxial tepals have a dark blue-black or red markings in the lower half. Possible hybrids between Codonorhiza azurea and C. fastigiata have been recorded at Piketberg Road (Gouda) [Bolus (BM, BOL, PH), Guthrie 2697 (NBG)]. Representative specimens SOUTH AFRICA. WESTERN CAPE (Worcester): Saron, renosterveld east of town, ( AA), 23 Sept. 2012, Goldblatt & Porter 13784A (MO); Tulbagh, ( AC), Grant 2427 (MO); Tulbagh, near cemetery, pebbly clay ground, ( AC), 24 Nov (fr.), Goldblatt 5226 (MO); 1 mile [± 1.5 km] south of Tulbagh Road Station, ( AC), Goldblatt 203 (BOL); rocky road verge south of Tulbagh, ( AC), Goldblatt & Porter (MO, NBG, PRE); Tulbaghskloof (77.9), ( AC), Ecklon & Zeyher Irid. 245 (G, MO); foot of the Elandskloof Mtns, Farm Elandsberg, ( AC), 23

105 S T R E L I T Z I A 35 (2015) 97 FIGURE 33. Codonorhiza elandsmontana, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

106 98 S T R E L I T Z I A 35 (2015) 6. Codonorhiza corymbosa (L.) Goldblatt & J.C.Manning, comb. nov. Ixia corymbosa L., Centuria plantarum: 4 (1756). Ovieda corymbosa (L.) Spreng.: 147 (1825). Lapeirousia corymbosa (L.) Ker Gawl.: t. 595 (1802); Baker: 90 (1896) [incl. C. azurea]; Marloth: pl. 43 (1915); Goldblatt: 20 (1972), excl. subsp. fastigiata and subsp. alta. Peyrousia corymbosa (L.) Poir.: 365 (1826). Meristostigma corymbosum (L.) A.Dietr.: 594 (1833). Type: South Africa, [Western Cape], near Camps Bay, sandy rocky ground, MacOwan s.n. in Herb. Norm. Austr. Afr. 268 (SAM, neo.!, designated by Goldblatt & Manning: 332 (1992); BM!, BOL!, G!, iso.). Ixia crispifolia Andrews: t. 35 (1798). Type: South Africa, without precise locality or collector, figure in Andrews: t. 35 (1798). Lapeirousia corymbosa var. grandiflora Klatt: 204 (1885). Type: South Africa, [Western Cape], mountains near Houw Hoek (Noord Hoek probably intended), Sept. 1883, MacOwan 2284a (K!, SAM 20988!, syn.). Lapeirousia corymbosa var. purpurea Klatt: 204 (1885). Type: South Africa, [Western Cape], sandy depressions, Fish Hoek, Sept. 1883, MacOwan 2284b (SAM, as Noord Hoek, syn.!). FIGURE 34. Distribution of Codonorhiza corymbosa, ; C. elandsmontana,. Sept. 1980, Goldblatt 5852 (MO); Gouda [Piquetberg Road Station], ( AC), Oct. 1892, Guthrie 2697 (NBG); H. Bolus (BM, BOL, PH) (mixed with hybrids with L. fastigiata); Bain s Kloof, lower west slopes ( CA), Goldblatt 590 (BOL); bottom of Bain s Kloof, Wellington, ( CA), 16 Oct. 1970, Strauss 50 (NBG); Botha, Farm Schoonuitzig, ( CB), 16 Oct. 1974, Walters 1230 (NBG). Plants mm high, 4 10-branched, branches subtended by bracts (1 )2 5 mm long, these sometimes leaf-like below and longer, stem ± flexuose, compressed and 2-winged. Corm obconic, mm diam.; tunics blackish, splitting vertically from base and apex. Leaves 2 or 3, lower 1(2) basal, usually falcate (or ± lanceolate), ( 10) mm, subacute to acute, with prominent main vein and 1 or a pair of secondary veins on either side slightly less prominent, margins ± straight, slightly undulate, rarely slightly crisped, uppermost leaf inserted in middle 1 / 3 of stem, mm long, narrowly oblong, not or hardly sheathing at base. Inflorescence a congested, flat-tipped false panicle, ultimate branchlets 1- or 2-flowered; bracts green, leathery, mm long, margins usually red, obtuse or inner slightly emarginate. Flowers actinomorphic, light to dark blue with white throat and lower half of tepals with white, bracketshaped band (rarely ± white with pale blue markings), not scented; perianth tube funnelshaped, 4 7( 8) mm long; tepals elliptic, obtuse, mm. Stamens symmetrically arranged around style; filaments 5 6 mm long, exserted 3 4 mm; anthers mm long, usually blue; pollen blue to grey, or ± white. Style dividing between base and upper 1 / 3 of anthers, often deflexed in older flowers, style branches mm long, divided for ± 1 / 2 their length. Capsules top-shaped, 3-lobed, mm long, with up to 6 seeds per locule. Seeds ovoid, reticulately sculptured, flattened to concave at chalazal end, dark brown, mm long; funicular collar up to 0.5 mm long. Flowering time: late September and October, occasionally until mid-november. Figure 35. Distribution and biology: Codonorhiza corymbosa extends from the coastal plain of Western Cape near Darling through the Cape Peninsula and Cape Flats to Bredasdorp and Elim (Figure 34). Flowering relatively late in spring, plants are usually found on sandy, well-drained soils derived from Cape Sandstone or weathered granite, but occasionally also on clay loam. Once

107 S T R E L I T Z I A 35 (2015) 99 FIGURE 35. Codonorhiza corymbosa, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

108 100 S T R E L I T Z I A 35 (2015) fairly common, it is becoming rare, especially on lowland sites, now mostly given over to farmland or urban development. It can still be found on lower mountain slopes and, especially after fires, may be common. Pollination of C. corymbosa has not been investigated (Goldblatt et al. 1995) but we infer that it has a generalist pollination strategy. Diagnosis and relationships: the crowded, ± flat-topped inflorescence of radially symmetric flowers with a pale to fairly dark blue perianth and white bracket-shaped marks in the lower middle of the tepals, distinguish Codonorhiza corymbosa from other Codonorhiza species. Although small, the upright flowers are particularly attractive with their white, star-shaped central marking. C. corymbosa is most easily confused with C. elandsmontana from the Worcester and Tulbagh districts, but that species has unilateral stamens, and although the perianth shape and colouration is similar, the flowers are larger and only the lower (abaxial) tepals have white markings. The perianth tube of C. elandsmontana is 8 10 mm long and the tepals mm long vs. a tube 4 7(8) mm long and tepals 6 10 mm in C. corymbosa. Confusion is also possible with the deep blue-flowered L. azurea, especially when specimens are poorly preserved, but the flowers and bracts of L. corymbosa are always smaller than those of L. azurea, the tepals of which are mm long. When alive the larger, dark blue, zygomorphic, abaxially tilted flowers of L. azurea with unilateral, horizontally extended stamens are unmistakable. White-flowered plants occur occasionally in populations across the range of Codonorhiza corymbosa, but at Riverlands near Malmesbury the entire population has white flowers. In other respects the plants from Riverlands appear to conform closely to typical C. corymbosa. Plants tentatively included in C. corymbosa by Goldblatt & Manning (1992) from Cape Point Nature Reserve (Pillans 8928, Nov. 1921, BOL), with slender perianth tube mm and tepals 7 8 ± 3 mm, are now regarded as a separate species, C. pillansii. The perianth tube length and shape are inconsistent with L. corymbosa, the funnelshaped perianth tube of which is 4 7 mm long. History: known to science at least since the last decade of the 17th century when it was figured by Plukenet (1696) as Caryophyllus monomatapensis, Codonorhiza corymbosa was assigned to the genus Ixia by Linnaeus in John Ker Gawler transferred the species to Lapeirousia (then spelled Lapeyrousia), where it has remained until now. Because the original specimen on which the species was based is lost, Goldblatt (1972) chose a neotype for the species. The specimen in the Linnaean Hebarium was collected by Anders Sparrman, circa 1771, some years after the publication of the protologue of Ixia corymbosa. Baker s account of Lapeirousia corymbosa in Flora capensis (1896) included as a variety the plant we here recognise as Codonorhiza azurea. Baker also regarded what we consider a second species, C. fastigiata, as a synonym of typical L. corymbosa, although he did recognise L. purpureo-lutea, the type of which is conspecific with C. fastigiata. Codonorhiza azurea and C. fastigiata were regarded by Goldblatt (1972) as a single subspecies of L. corymbosa, but were later recognised by Goldblatt & Manning (1992) as separate species, a treatment that is most consistent with their morphological and ecological differences. Representative specimens SOUTH AFRICA. WESTERN CAPE (Cape Town): Riverlands between Mamre Road and Malmesbury, 23 Oct. 1980, ( BC), Esterhuysen s.n. (MO); Lions Head, ( CD), H. Bolus 2819 (BOL, K); Table Mtn, ( CD), Ecklon s.n. or 545 L. azurea (C, G, K); Claremont flats, ( CD), Wolley Dod 389 (BOL, K); Paarl Mtn, southern end near Taal Monument, ( DB), 17 Oct. 1986, Goldblatt 7922 (MO); Jonkershoek, ( DD), 1 Oct. 1958, Werdermann & Oberdieck 353 (B, MO); 28 Oct. 1976, Kruger 211 (NBG, PRE) (Worcester): Franschhoek, between Pniel and Robertsvlei ( CC), Phillips 1314 (SAM); between Groot Drakenstein and the foot of Paarl Mtn, ( CC), Drège 8510 (E, G, MO) (Simonstown): Bergvliet Farm, ( AB), Purcell s.n. (SAM 90197, 90200); burnt lower slopes of Klaasjagersberg, opposite Cape Point Reserve, ( AD), 26 Nov. 1979, Goldblatt 5259 (MO); Smitswinkel, ( AD), Compton 8024 (NBG); Buffels Bay, ( AD), Compton 6381 (NBG); Cape Point Nature Reserve, west of Diaz Cross, ( AD), 3 Jan (fr.), Goldblatt & Manning 9499 (MO); near Sir Lowry s Pass, 250 m, ( BB), Schlechter 5362 (G, GRA, K); Lourensford Estate, Somerset West, ( BB), Parker 4208 (BOL, NBG); Steenbras River mouth, ( BB), Compton (NBG); Betty s Bay, Porter Reserve, ( BD), Ebersohn 162 (NBG) (Caledon): Caledon, 300 m, ( AB), 17 Oct. 1894, Schlechter 5569 (B, GRA, MO, Z); Hermanus, Vogelklip, ( AD), 16 Oct. 1980, Williams 511 (MO); Vogelgat near Hermanus, ( AD), 15 Oct. 1986, Goldblatt 7914

109 S T R E L I T Z I A 35 (2015) 101 (MO); 8 miles [± 12 km] from Stanford on road to Elim, ( AD), 9 Oct. 1938, Gillett 4503 (BOL, MO); Fairfield, ( BD), 5 Oct. 1993, Nänni 3 (NBG); between Baardscheerdersbos and Elim, ( DA), 15 Oct. 1983, Goldblatt 7106 (MO); 4 km west of Elim, ( DA), 9 Nov (fl. & fr.), Goldblatt & Porter (MO) (Bredasdorp): Strandkloof, 8 Oct. 1950, ( CA), Martin 601 (NBG). 7. Codonorhiza fastigiata (Lam.) Goldblatt & J.C.Manning, comb. nov. Lapeirousia fastigiata (Lam.) Ker Gawl. in König & Sims Annals of Botany 1: 238 (1804). Ixia fastigiata Lam.: 337 (1789). L. corymbosa subsp. fastigiata (Lam.) Goldblatt: 25 (1972). Type: South Africa, without precise locality or collector (P: Herb. Lamarck, holo!). Ovieda purpureolutea Klatt, Linnaea 32: 781 (1864). Lapeirousia purpureolutea (Klatt) Baker: 515 (1877); 91 (1896). Type: South Africa, [Western Cape], Tulbagh, collector unknown, Herb. Bergius s.n. [SBT, lecto.! designated by Goldblatt: 25 (1972)]. Lapeirorosea corymbosa var. rosea Suess.: 88 (19%). Type: South Africa, [Western Cape], Saron, 29 Sept. 1946, Rehm s.n. (M, holo.). Plants mm high, several-branched, branches subtended by bracts (1 )2 3 mm long, lower sometimes ± leaf-like and longer, stem ± flexuose, compressed and narrowly winged, sheathed below by 2 membranous, pale or light brown cataphylls. Corm obconic, mm diam.; tunics blackish, splitting vertically from base and apex. Leaves 2, lowermost inserted at ground level, largest, falcate, ( 250) mm, obtuse to subacute, with prominent main vein and pair of secondary veins on either side almost as prominent, margins usually undulate and crisped, rarely ± straight; upper leaf inserted near middle of stem, (12 )25 45 mm long, oblong-lanceolate, margins straight, not or hardly sheathing at base. Inflorescence a congested, ± flat-topped false panicle, ultimate branchlets (1)2 5-flowered; bracts green, often red on margins, 6 10 mm long, obtuse or inner emarginate. Flowers actinomorphic, pale yellow to nearly white with purple or brown median triangular mark in basal 1 / 3 of each tepal, sometimes flushed dull purple or brown outside, not scented; perianth tube funnel-shaped, mm long; tepals elliptic, obtuse, mm. Stamens symmetrically disposed, diverging; filaments ± 10 mm long; anthers 3 4 mm long, yellow, pollen white to pale yellow. Style dividing between base and middle of anthers, style branches mm long, divided for ± 1 / 2 their length. Capsules top-shaped and 3-lobed, 5 6 mm long, with up to 6 seeds per locule. Seeds ovoid, reticulately sculptured, red-brown, darker on ridges, mm long with funicular appendage up to 0.4 mm long. Chromosome number 2n = 20. Flowering time: late September to mid-october. Figure 36. Distribution and biology: once fairly common locally in Western Cape from Saron and Moorreesburg in the south to Piekeniers Kloof Pass in the north, Codonorhiza fastigiata is most often seen on shale soils on hills and lower mountain slopes in renosterveld shrubland, but plants are sometimes found on rocky, sandstone slopes (Figure 29). Flowering is enhanced by clearing or burning, but it also blooms in undisturbed vegetation, when plants are smaller and bear fewer flowers. A collection from Gydouw (in the Ceres valley), Leipoldt 3867, some distance from other records for C. fastigiata, requires verification [Gydouw may be a transcription error for Gouda, where the species does grow]. Pollination of C. fastigiata has been little studied, but Goldblatt et al. (1995) reported visits to the species by the large-bodied bees, Anthophora diversipes (Apidae) and Hoplitis similis (Megachilidae) (sex not recorded). We suspect C. fastigiata flowers offer only pollen as a reward as we were unable to detect even a trace of nectar at the base of the perianth tube in unbagged flowers. Diagnosis and relationships: pale yellow flowers and relatively large perianth with a tube 6 8 mm long and tepals mm long are sufficient to distinguish Codonorhiza fastigiata in the genus. Other species of the genus with flattopped inflorescences have light to dark blue or violet flowers except L. micrantha, which has a buff to brown or maroon perianth, but smaller flowers with a cylindric perianth tube. Both perianth and stamens of L. fastigiata are symmetrically arranged so that L. azurea, which has flowers of similar size and shape, can immediately be distinguished by its asymmetrically disposed stamens and perianth with markings only on the lower three tepals, as well as by its dark blue perianth. Collections from Piketberg (Guthrie 2696) and Gouda (as Piketberg Road Station) (H. Bolus

110 102 S T R E L I T Z I A 35 (2015) FIGURE 36. Codonorhiza fastigiata, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

111 S T R E L I T Z I A 35 (2015) ) that comprise plants with either yellow or blue flowers were thought by Goldblatt (1972) to represent populations of plants intermediate between Codonorhiza fastigiata and C. azurea, and based largely on these collections, Goldblatt concluded that the two were conspecific. Additional field study has failed to reveal any other intermediates while several more populations corresponding exactly to one or other of the two species have been found. We are now convinced that L. fastigiata and L. azurea are separate species and the two gatherings of intermediate plants now seem more likely to represent hybrid individuals. That the flowers of the yellow-flowered plants of these collections differ in the intensity of purple colouration on the reverse of the tepals seems to support this argument. Such variation within a population is rare except among hybrids. History: available records indicate that Codonorhiza fastigiata was first collected sometime before 1789, when the species was described by the French scientist J.P. Lamarck, then referred to Ixia. The source of several Cape species now in the Lamarck and Jussieu collections at the Paris Herbarium is uncertain, although sometimes attributed to Philibert Commerson, who did not visit the Cape (Gunn & Codd 1981). His colleague, Pierre Sonnerat, did, however, briefly collect there en route to Mauritius where he worked with Commerson. Specimens in the Bergius Herbarium in Stockholm, Sweden, assembled between 1760 and 1790, are contemporary, and their source is also unknown. Later collections made by Ecklon and Zeyher, probably in 1829, as well as the material in the Bergius Herbarium, formed the basis for Klatt s Ovieda purpureo-lutea, described in Baker transferred the species to Lapeirousia in Lapeirousia purpureo-lutea, L. fastigiata and L. azurea were united at subspecific rank under L. corymbosa by Goldblatt (1972), a decision based primarily on two collections of plants that appeared to be intermediate between the two species. Representative specimens SOUTH AFRICA. WESTERN CAPE (Clanwilliam): Piketberg, east of Farm Weltevrede, ( DA), 19 Sept. 2007, Goldblatt & Manning (MO, NBG); top of Grey s Pass [Piekeniers Kloof], ( DB), Taylor 967 (BOL); Piketberg, ( DC), Fremantle s.n. (BOL 30698); Guthrie 2696 (BOL) (partly hybrid); between Piketberg and Berg River bridge, ( DC), Barnes s.n. (NBG); 5 miles [7.5 km] north of Piketberg, ( DC), Salter 2720 (BOL); slopes of Versfeld Pass, Piketberg, ( DC), Goldblatt 671 (BOL); north slopes of Heuningberg, ( DD), Goldblatt 650 (BOL) (Wuppertal): lower slopes of the Olifants River Mtns at Cardouw Pass, ( CC), 27 Sept. 1976, Goldblatt 4211 (MO) (Cape Town): Moorreesburg, ( BA), Bachmann 775 (B); near Moorreesburg, clay soil, ( AC), H. Bolus 9983 (BOL); Porterville, ( BB), Loubser 498 (NBG) (Worcester): near Saron, ( AA), Oct. 1896, Schlechter (BM, E, G, GRA, K, MO, P, PH); Gydouw, ( AB), Leipoldt 3867 (BOL); Tulbaghskloof, ( AC), Ecklon & Zeyher Irid. 246 (G, MO).

112 104 S T R E L I T Z I A 35 (2015) Systematics of Schizorhiza Schizorhiza Goldblatt & J.C.Manning, gen. nov. Type: S. neglecta (Goldblatt) Goldblatt & J.C.Manning. Deciduous geophytes. Corm ellipsoid with small flat base, rooting from base, axillary in origin, producing several new corms in place of parent corm at end of growing season; tunics of ± corky layers. Foliage leaves several, linear to sword-shaped, evidently plane but slightly pleated, with visible main vein; lowermost often longest and inserted on stem near ground level, upper leaves progressively smaller. Stem aerial, compressed and angled, branching repeatedly, producing cormlets in leaf axils above and below ground. Inflorescence much branched, forming a ± round-topped false panicle, with sessile flowers; bracts green, firm, inner ± as long as outer, acute or emarginate. Flowers zygomorphic, long-lived, often salver-shaped, lower (abaxial) tepals with contrasting darker markings; perianth tube ± cylindric; tepals subequal, but dorsal slightly larger. Stamens unilateral; filaments slender, free; anthers oblong-linear, dehiscence longitudinal. Ovary globose, sessile; style filiform, branches forked for ± 1 / 2 their length. Capsules cartilaginous, ± top-shaped, 3-lobed above. Seeds ± globose, flattened at chalazal end, slightly wrinkled; surface cells ± flat. Basic chromosome number x = 6; 2n = 12 and 24. Species: 1, mountains of the southwestern Cape, South Africa. 1. Schizorhiza neglecta (Goldblatt) Goldblatt & J.C.Manning, comb. nov. Lapeirousia neglecta Goldblatt in Goldblatt & Manning, South African Journal of Botany 58: 335 (1992). Type: South Africa, [Western Cape], Jonkershoek, slopes near Berg River Nek, Nov. 1989, Esterhuysen (BOL, holo.!; B, BR!, E, G, K!, M, MO!, NBG!, P, PRE!, S!, SRGH!, US, WAG, iso.). L. corymbosa subsp. alta Goldblatt: 24 (1972). Type: South Africa, [Western Cape], near Steenbras Dam, Dec. 1945, Lewis 1595 (SAM, holo.!; BOL!, SAM!, iso.). Plants ( 550) mm high, laxly 5 7-branched, branches subtended by bracts 3 5( 12) mm long, those subtending lower branches becoming ± leaf-like, stem ± straight, compressed, 2-winged below, often ± square above and 4-angled to narrowly 4-winged; bearing 2 or more cormlets in axils of lower leaves. Corm mm diam., with several cormlets at base; tunics corky to somewhat papery, brown, not accumulating. Leaves 3 6, lower 2 4 inserted below ground level and largest, narrowly lanceolate, lowermost reaching to middle to shortly exceeding inflorescence, 7 15 mm wide, with prominent main and a pair of secondary veins (rarely 2 pairs) on either side ± as prominent; upper 1 3 leaves inserted in lower half of stem, progressively shorter above, not or hardly sheathing at base. Inflorescence a lax, rounded to ± flat-topped false panicle, flowers crowded distally, ultimate branchlets 1- or 2( 4)-flowered; bracts green, margins often red, 4 7 mm long, inner usually slightly longer than outer, ± membranous above at anthesis, acute or inner slightly emarginate. Flowers zygomorphic, either white, then lower tepals usually with one or a pair of purple median spots (Jonkershoek), or blue with darker blue to violet spear-shaped markings in lower midline and often white-dotted in centre of dark colour (Bain s Kloof), with slight sweet scent; perianth tube ± cylindric, 7 10 (Jon kershoek), (Bain s Kloof and Steenbras) mm long, flared from 1 mm diam. at base to 2 mm at mouth; tepals elliptic, obtuse, unequal, dorsal ± 4.5 mm, lower held closely together, ± 4 mm wide. Stamens unilateral and arcuate; filaments 5 6 (Jonkershoek), ± 9 (Bain s Kloof) mm long, exserted 3 5 mm; anthers mm long, pollen ± white. Style arching

113 S T R E L I T Z I A 35 (2015) 105 adaxial to filaments, dividing opposite lower to middle 1 / 3 of anthers, style branches ± 2 mm long, divided for ± 1 / 3 their length. Capsules topshaped and 3-lobed, 4 5 mm long, sometimes flushed red before drying; with 1 or 2(3) per locule. Seeds globose to ovoid, sometimes flattened by pressure, slightly wrinkled, ± mm, with small funicular collar. Chromosome numbers 2n = 12, 24. Flowering time: mid- November to late January. Figure 37. Distribution and biology: restricted to a 65 km portion of the north south trending mountains in the extreme southwest of Western Cape, South Africa, Schizorhiza neglecta is known from just three extended populations in Bain s Kloof, Jonkershoek and in the Steenbras Mtns to the south (Figure 38). Before 1989, when it flowered profusely at Jonkershoek near Stellenbosch, S. neglecta had been recorded only a handful of times since its discovery by R.H. Compton in November 1943 in the Jonkershoek Mtns. Its presence in Baviaans Kloof, a tributary valley of Bain s Kloof, was established shortly thereafter by Compton and G.J. Lewis in January It grows in rocky sandstone-derived soil at elevations of m, generally in sheltered, wetter sites such as slopes below cliffs and near streams where additional moisture is available. Its habit of flowering in the early summer following a fire the previous summer or autumn partly explains why it is so seldom collected. It is fairly common locally at Jonkershoek, where it occurs in several isolated sites, and can form extensive and dense populations, notably at Berg River Nek, where it carpets the slopes with white when in full bloom. Pollination of S. neglecta has not been critically studied, but Goldblatt et al. (1995) recorded visits by Xylocopa hottentotta (Apidae: Xylocopidae), a large-bodied but short-tongued bee. More study is needed before this observation can be placed in context. Diagnosis and relationships: when first described, Schizorhiza neglecta was associated with the Cape species of Lapeirousia (now Codonorhiza), but its basic chromosome number of x = 6 is discordant with those species, which have x = 10 (excluding B chromosomes). Corm morphology is likewise unlike that of Codonorhiza and is unique for Crocoideae in the ellipsoid shape with the flat base of narrower diameter than the middle of the corm. The corky to papery tunics, rather than woody (Lapeirousia) or of compacted fibres (Codonorhiza, Psilosiphon), are likewise distinctive. The large, smooth to slightly wrinkled, ± globose seeds nearly match those of Lapeirousia and Psilosiphon, but are quite unlike the smaller, ovoid, reticulate seeds of Codonorhiza. Leaf anatomy, including the absence of marginal veins or subepidermal sclerenchyma strands and closed vascular bundle sheaths conform to the condition in Codonorhiza, but the alternate main vascular bundles and slightly plicate leaf blades are unique. Pollen grains with a single banded operculum recall Codonorhiza, whereas all Lapeirousia and Psilosiphon species examined for the character have a 2-banded operculum. Morphologically, the flowers of S. neglecta appear most like those of the short-tubed P. erythranthus alliance, particularly the southern African P. sandersonii, especially in having the upper tepal reflexed to lie in the same plane as the lower tepals. The production of multiple cormlets in the lower nodes of the stem and in place of the parent corm at the end of the growing season is also unique for the Lapeirousia clade. In the DNA-based phylogeny (Figure 1) Schizorhiza neglecta is retrieved as sister to the Lapeirousia plus Psilosiphon clades (using plastid DNA sequences alone it is retrieved in a polytomy with these two clades). This result at first seemed puzzling as S. neglecta seemed to accord with L. sect. Paniculatae, now Psilosiphon, in floral and seed morphology and cytology, although discordant geographically. Careful re-examination of the species has shown that its leaf anatomy and corm morphology is unique, thus not conforming to Psilosiphon, and the pollen grains with a 1-banded operculum also differ from that genus, resembling instead those of Codonorhiza among the Lapeirousia group of genera. Variation: the type population of Schizorhiza neglecta from Jonkershoek, with white flowers and a shorter perianth tube, 8 10 mm long is diploid, 2n = 12 (Goldblatt & Manning 1992), but counts from blue-flowered plants from Bain s Kloof with a longer perianth tube, mm, are tetraploid, 2n = 24 (Table 4). These differences led us to consider whether they might be better regarded as separate taxa and in the description above, we separate the significant differences between these populations. The longer-tubed Bain s Kloof and Steenbras popula-

114 106 S T R E L I T Z I A 35 (2015) C D A B FIGURE 37. Schizorhiza neglecta. A, flowering stems; B, corm and cormlets [see text]; C, fruiting stem; D, flowers. A C, Goldblatt & Manning 9022; D, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

115 S T R E L I T Z I A 35 (2015) 107 tions also have longer floral bracts 6 7 mm long (vs. 4 6 mm) and anthers mm long (vs mm) than the Jonkershoek plants. The size differences may be explained by the ploidy difference, although chromosome number for the Steenbras population is unknown and flower colour is uncertain (residual pigmentation on herbarium specimens suggests that they were blue). We defer recognition of infraspecific taxa for this rare Cape plant, known from just three extended populations, pending additional study. History: although thought by G.J. Lewis, a specialist on the systematics of southern African Iridaceae, to be a new species of Lapeirousia when first collected in the 1940s, Schizorhiza neglecta remained unnamed until Goldblatt (1972) treated it as L. corymbosa subsp. alta. Our examination of living plants in 1989 at Jonkershoek made it clear that these populations represented a separate species, which was then described as L. neglecta (Goldblatt & Manning 1992). At the time we considered it to be not even particularly closely related to the L. corymbosa complex, and placed the species in the otherwise tropical and eastern southern Africa sect. Paniculatae, now Psilosiphon. Representative specimens SOUTH AFRICA. WESTERN CAPE 3318 (Cape Town): Jonkershoek, near waterfall, ( DD), 6 Nov. 1943, Compton (PRE); Jonkershoek Valley, wet gully along trail to Berg River Nek, ( DD), 29 Dec. 1989, Goldblatt & Manning 9022 (B, K, MO, NBG, PRE, WAG); Jonkershoek, FIGURE 38. Distribution of Schizorhiza neglecta,. Dwarsberg, upper contour path, among rocks in recently burnt firebreak, ( DD), 19 Jan. 1975, Kerfoot & Haynes 14 (MO, PRE); trail at foot of cliffs above Berg River Nek, ( DD), 29 Dec. 1989, Goldblatt & Manning 9031 (K, MO, NBG, PRE, WAG) (Worcester): Bain s Kloof, ( CA), 17 Jan. 1945, Compton (NBG); Lewis 979 (SAM); Baviaanskloof, ( CA), 31 Dec. 1992, Goldblatt & Manning 9489 (MO, NBG) (Simonstown) Jonkershoek, Swartboskloof, upper west-facing slope at base of cliffs, m ( BB), 2 Dec. 1989, Oliver 9338 (K, NBG); near Steenbras Dam, ( BB), Dec. 1945, Lewis 1595 (BOL, SAM).

116 108 S T R E L I T Z I A 35 (2015) Systematics of Psilosiphon Psilosiphon Welw. ex Goldblatt & J.C.Manning, gen. nov. Type: P. sandersonii (Baker) Goldblatt & J.C.Manning. Lapeirousia subg. Paniculatae [Paniculata] Goldblatt & J.C.Manning [sect. Paniculatae only] in Annals of the Missouri Botanical Garden 77: 373 (1990). Type: L. erythrantha (Klotzsch ex Klatt) Baker (= P. erythranthus (Klotzsch ex Klatt) Goldblatt & J.C.Manning). Deciduous geophytes. Corm flat-based, bell-shaped, rooting from base, axillary in origin; tunics of compacted fibres, outer layers decaying into loose fibrous network or vertical strips. Foliage leaves few to several, linear to sword-shaped, falcate or ± terete, plane with visible main vein; lowermost often longest and inserted on stem near ground level, upper leaves progressively smaller. Stem aerial, compressed and angled to winged, usually branched, often repeatedly. Inflorescence either much branched, forming a ± a round or flat-topped false panicle, with sessile flowers, or a simple or branched spike; bracts green and ± soft-textured, often becoming dry distally, inner ± as long as outer, often forked near apex. Flowers usually zygomorphic, or radially symmetric, long-lived, often salver-shaped or rotate, when zygomorphic, lower tepals usually with contrasting darker or lighter markings, sometimes sweetly scented; perianth tube cylindric or funnel-shaped, short to extremely long; tepals subequal or unequal with dorsal largest. Stamens symmetrically disposed or unilateral and arcuate; filaments slender, free; anthers oblong-linear, dehiscence longitudinal. Ovary globose, sessile; style filiform, branches forked for 1 / 3 to 1 / 2 their length, undivided or notched apically. Capsules cartilaginous, ± globose. Seeds ± globose, flattened at chalazal end, smooth or slightly wrinkled; surface cells usually domed or ± flat. Basic chromosome number uncertain, diploid numbers, 2n = 16, 12, 10, 8 (diploids); 2n = 14, 12, 10, 8, 6 (tetraploids). Species: 15; sub-saharan Africa, from Ethiopia to Zimbabwe, Mozambique and Angola, one species also in Nigeria, several in Botswana and Namibia, two in eastern South Africa. Key to species in southern Africa 1a Perianth tube 1 2 mm long; flowers actinomorphic; all tepals with contrasting markings: 2a Tepals (4 )5 7 mm (appearing shorter in poorly preserved specimens); flowers 1(2) on ultimate branches of inflorescence; corm tunics dark brown to blackish and decaying irregularly into vertical strips P. avasmontanus 2b Tepals mm; flowers (2)3 5 on ultimate branches of the inflorescence; corm tunics light brown, outer layers of fine reticulate fibres P. coeruleus 1a Perianth tube mm long; flowers zygomorphic with arcuate to ± erect unilateral stamens; contrasting markings when present confined to 3 lower (abaxial) tepals: 3a Perianth tube mm long; flowers white, sometimes flushed pale blue to purple outside L schimperi 3b Perianth tube 3 50 mm long; flower variously coloured: 4a Inflorescence a divaricately branched false panicle with main branches 1- or 2(3)-flowered: 5a Perianth white to pale pink; corm tunics straw-coloured and outer layers ± fibrous and reticulate; floral bracts dry before anthesis, becoming membranous, dry and light brown with tips often darker brown P. bainesii

117 S T R E L I T Z I A 35 (2015) 109 5b Perianth blue to violet, lower tepals with white markings edged dark blue to violet; corm tunics dark brown and outer layers breaking into vertical strips, not fibrous; floral bracts acute-attenuate and dark brown in distal half P. sandersonii 4b Inflorescence a ± false panicle or spike with at least some main branches 3 8-flowered: 6a Perianth tube mm long: 7a Flowers creamy white, lower tepals with purple median spear-shaped markings; perianth tube mm long; tepals mm P. otaviensis 7b Flowers blue to violet, rarely pale blue to almost white, lower tepals with blue to violet markings; perianth tube mm long; tepals mm: 8a Flowers violet, lower tepals with white median mark edged in darker violet; tepals ± 14 mm long; floral bracts 8 10 mm long; perianth tube mm long P. erongoensis 8b Flowers pale blue to white with bluish markings on lower tepals; tepals mm long; floral bracts 3 6( 7) mm long; perianth tube mm long P. gracilis 6b Perianth tube 3 25 mm long: 9a Perianth tube (15 )18 25 mm long: 10a Perianth greenish or blue-violet with red to purple and white markings on lower tepals; floral bracts 5 7 mm long; plants of Malawi, Mozambique, South Africa (Limpopo Province) and Zimbabwe P. masukuensis 10b Perianth pale blue or white; floral bracts 3 6( 7) mm long; plants of Namibia P. gracilis 9b Perianth tube (5 )7 15 mm long: 17a Dorsal tepal suberect or arched over stamens; lower tepals often (?always) with tooth-like median ridge up to 1 mm long; style branches undivided or notched at tips P. rivularis 17b Dorsal tepal reflexed, lying in same plane as lower tepals; lower tepals without toothlike median ridges; style branches divided for 1 / 3 1 / 2 their length P. erythranthus Spp Psilosiphon sandersonii (Baker) Goldblatt & J.C.Manning, comb. nov. Lapeirousia sandersonii Baker, Handbook of the Irideae: 169 (1892), 95 (1896), 7: 352 (1898, excl. specimens cited); Goldblatt: 463 (1990a). Type: South Africa, Transvaal, Sanderson s.n. [K, lecto.! designated by Goldblatt: 463 (1990a)]. [Lapeirousia bainesii var. breviflora Baker: 156 (1877), nom. nud.] [no specimen cited, but Nelson 402, K, from Rhenosterpoort is so annotated by Baker]. Plants ( 450) mm high, divaricately and often intricately branched, stem compressed, 2-angled below, 3-angled above branches, slightly winged, sheathed below by dark brown cataphylls, inner reaching shortly above ground. Corms mm in diam.; tunics dark brown, hard-leathery internally, decaying irregularly into vertical sections, seldom becoming fibrous and never netted. Leaves 2 4, ± linear to falcate, 2 3 mm wide, firm to rigid, main and secondary vein pairs ± prominent, closely set; lowermost leaf longest, sometimes exceeding inflorescence, upper leaves progressively shorter. Inflorescence a rounded to ± flat-topped false panicle but with main axis sometimes evident, ultimate branches 1-or 2(3)-flowered; bracts 5 12 mm long, green with brown tips in bud, often flushed purple, becoming dry and dark brown with age, acute-attenuate, inner sometimes longer than outer, usually notched at apex. Flowers zygomorphic, blue to violet, lower 3 tepals each with deep red to purple and white spear-shaped mark guide in lower midline; perianth tube 8 12 mm (subsp. limpopoensis), or mostly mm (subsp. sandersonii), slender, widening slightly in upper 2 3 mm; tepals subequal, lanceolate, mm, dorsal held apart from others, reflexed, lower 3 joined for ± 1 mm and held close together, when fully open all ± spreading. Filaments unilateral, ± erect, ± 5 mm long, exserted mm from tube; anthers 3 5 mm long, purple; pollen pale yellow or white. Style dividing near

118 110 S T R E L I T Z I A 35 (2015) apex of anthers or shortly (rarely much) exceeding anther tips; branches 1 2 mm long, divided for up to 1 / 3 their length, notched apically or undivided. Capsules depressed globose, mm. Seeds nearly globose, ± mm diam. Chromosome number 2n = 10. Flowering time: mainly February and March, occasionally from late December or in April. Figures 39, 40. Distribution: Psilosiphon sandersonii is native to the interior of eastern Botswana and the adjacent northern South Africa where it occurs in relatively dry habitats in North West and Limpopo provinces, as well as in Northern Cape and western Mpumalanga (Figure 41). It appears to be especially common in the Magaliesberg from Pretoria west to Rustenburg, and according to the collection record, is relatively rare in Limpopo and Botswana. The easternmost records from the cool, high, well-watered Dullstroom, Belfast and Middelburg areas of Mpumalanga are surprising in view of the rest of the range in semi-arid country, but the eastern populations differ only in their slightly longer perianth tube, mm long, from those occurring further to the west. The habitat is always rocky, and usually well drained, including hill slopes, ridges and summits. The distinctive corms are seldom collected, which probably reflects the difficulty in their extraction from rocky ground. We recognise two subspecies, subsp. sandersonii extending from eastern Botswana and adjacent South Africa as far east as Dullstroom in Mpumalanga and as far south as the Kuruman hills in Northern Cape, where some populations have a perianth tube up to 30 mm long; and subsp. limpopoensis, with a short perianth tube 8 12 mm long and most common in the Waterberg of Limpopo Province, but extending north into the Soutpansberg, where subsp. sandersonii also occurs. Diagnosis and relationships: allied to the Psilosiphon erythranthus complex of shorttubed species, P. sandersonii has the repeatedly branched, panicle-like inflorescence and blue-violet flowers that characterise most tropical African species of the genus. It can be distinguished from the related P. erythranthus by its typically longer perianth tube, mostly mm long, and ± divaricately and often intricately branched inflorescence with 1 or 2 (rarely 3) flowers on the major terminal branches. The floral bracts are often relatively long, 8 12 mm, acute-attenuate and flushed purple or brown and aging dark brown. The corm tunics too, are distinctive, being dark brown and fragmenting into vertical strips rather than becoming a fibrous network. P. erythranthus typically has fewer-branched inflorescences with 3 6 flowers per main terminal branch and a perianth tube usually 6 11 mm long, rarely up to 15 mm. The floral bracts are 4 6 mm long, shorter than those in most populations of P. sandersonii. A good illustration of P. sandersonii is in Flowering Plants of Africa 31: t (1956). The strongly asymmetric karyotype of Psilosiphon sandersonii, with 2n = 10 (Goldblatt, 1990b: one population counted), consists of a pair of very long chromosomes and 9 pairs of much shorter chromosomes. This contrasts sharply with populations of P. erythranthus, which have 2n = 8, 14 or 12 and a relatively symmetric karyotype. The variation in perianth tube length, not evident to Goldblatt (1990a), correlates with geographical distribution. Plants from Limpopo, mainly in the Waterberg and nearby, have particularly short tubes, 8 12 mm long, whereas populations in the southwest of the range, in the Kuruman hills and adjacent country in Northern Cape, stand out in having a much longer perianth tube, mm long. The regional variation is not always consistent and plants from Mpumalanga sometimes have perianth tubes almost as long as those from further west. In other respects, especially the branching pattern and rust-brown, acute-attenuate floral bracts, the long- and shorttubed plants accord with typical P. sandersonii. We recognise just two geographic subspecies to reflect the variation in tube length, referring populations from the Waterberg and nearby to subsp. limpopoensis and the remaining, longer-tubed populations to subsp. sandersonii. Subsp. limpopoensis, in particular, broadly resembles P. erythranthus in its short perianth tube, 8 12 mm long, and relatively short floral bracts, 5 6 mm long (see under subsp. limpopoensis). The plants, however, have the divaricately branched stem and brown floral bracts typical of P. sandersonii except in their shorter length. They may represent recent or ancient hybridisation between P. sandersonii and P. erythranthus, the ranges of which do not overlap today but lie less than 100 km apart in Botswana. The southernmost stations of P. erythranthus in Zimbabwe and the northernmost stations of P. sandersonii subsp. limpopoensis in South Africa lie substantially further apart.

119 S T R E L I T Z I A 35 (2015) 111 FIGURE 39. Psilosiphon sandersonii subsp. sandersonii, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

120 112 S T R E L I T Z I A 35 (2015) FIGURE 40. Psilosiphon sandersonii subsp. sandersonii, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

121 S T R E L I T Z I A 35 (2015) 113 Psilosiphon masukuensis is sometimes confused with P. sandersonii, but the inflorescence of that species is comparatively sparsely branched with the main branches 5 9-flowered, and the flowers are smaller, although perianth tube length, mm long, overlaps that of L. sandersonii. The corms of the two species also differ, those of P. masukuensis being smaller and having paler, fibrous tunics. P. masukuensis appears to favour wetter, lowland sites in contrast to the montane, rocky, well-drained habitats of P. sandersonii. History: collected first by John Sanderson, probably in 1852 when he travelled to Rustenburg and the Magaliesberg on his only journey to then Transvaal Republic (Gunn & Codd 1981), Psilosiphon sandersonii first appeared in the literature as the nomen nudum Lapeirousia bainesii var. breviflora (Baker 1877). Baker subsequently described L. sandersonii in 1892 and the species was so treated in Flora capensis (Baker 1896). In Flora of tropical Africa (Baker 1898), specimens cited as L. sandersonii are not this species, but the superficially similar, blue- or red-flowered P. erythranthus, then poorly understood. Key to subspecies 1a Perianth tube mostly mm long; floral bracts 8 12 mm long; style branches mm long... 1a. subsp. sandersonii 1b Perianth tube mostly 8 12 mm long; floral bracts 5 6 mm long; style branches mm long.... 1b. subsp. limpopoensis 1a. subsp. sandersonii Plants ( 450) mm high. Corms mm diam. Inflorescence with bracts 8 12 mm long. Flowers with perianth tube mm long; tepals mm. Stamens with filaments 5 7 mm long, exserted mm. Style dividing near apex of anthers, branches mm long, divided for up to 1 / 4 their length or barely notched at tips (or apparently undivided). Figures 39, 40 & 41. Representative specimens BOTSWANA (Dikgomo di Kae): Jwaneng TV hill, 37 km southeast of Jwaneng to Kanye, ( DD), 24 Feb. 1997, Cole 1214 (MO, PRE) (Gaborone): near Molepolole, shale, ( BC), 15 Mar. 1930, Van Son s.n. (PRE 28664); west of Gaborone Dam, ( DB), 1976, Mott 928 (SRGH, UCBG); Gaborone, near Notwane sewage ponds, ( DB), 22 Mar. 1981, Barnes 185 (PRE) (Mafeking): 6 miles [9 km] west of Kanye, ( AB), 12 Feb. 1971, Van Rensburg B4226 (PRE). SOUTH AFRICA. LIMPOPO (Waterpoort): Zoutpansberg, 1 mile [1.5 km] from Dandy Farm, road to Sand River, south slopes in rocky outcrop, ( DC), 3 Apr. 1957, Meeuse (K, M, PRE, SRGH); Farm Thornhill 743, red sand in riverine woodland, ( DD), 18 Mar. 1985, FIGURE 41. Distribution of Psilosiphon avasmontanus, ; P. sandersonii subsp. limpopoensis, ; subsp. sandersonii,.

122 114 S T R E L I T Z I A 35 (2015) Raal 434 (PRE) (Messina): Louis Trichardt, Farm Riet bok 226, among rocks in montane scrub forest, 1476 m, ( CC/CD), 25 Mar. 1982, Venter 7812 (PRE). NORTH WEST (Zeerust): Zeerust, ( CA), Jan. 1928, Thode A1503 (GH, K), Feb. 1912, Jenkins TM11660 (K, PRE); 15 m [22 km], east of Swartruggens, bushveld on ridges, 14 Feb. 1956, ( DB), Acocks (K, PRE) (Rustenburg) Rustenburg, 4000 ft [1 240 m], stony hillside near town, ( CA), 1904 (fl. & fr.), Pegler 1108 (GRA, K, PRE, SAM); 22 m west of Rustenburg, ( CA), 14 Feb. 1956, Leistner 539 (K, PRE); Rustenburg, Tierkloof, moeras by wildreservaathek, ( CA), 13 Mar. 1976, Venter 654 (K, MO, PRE, SRGH, WAG); Brits, Silikaatsnek, top of poort in shallow sand over quartzite, ( DB), Codd 736 (PRE) (Vryburg): Vryburg, ( DC), 5 Feb. 1948, Rodin 3501 (K, MO, P, PRE, S) (Delareyville): Vryburg District, Gold Ridge Farm, koppie peak in banded ironstone rocks, ( AB), 15 Feb. 1980, Gubb (PRE) (Potchefstroom): Dassiesrand, Potchefstroom, ( CA), 23 Mar. 1940, Van der Westhuizen 1114 (PRE). GAUTENG (Rustenburg): Krugersdorp, Jack Scott Nature Reserve, ( DC), 2 Feb. 1961, Wells 2305 (K) (Pretoria): Magaliesberg, Hornsnek, 12 km west of Pretoria, ( CA), 12 Jan. 1956, Schlieben 7751 (B, BR, G, HBG, K, M); 31 Jan. 1983, Perry 2017 (MO, NBG); The Willows, Pretoria, ( CA), 9 Mar. 1906, Burtt Davy 5333 (GRA, K); Cullinan, Farm Vaalwaterkrans, 25.2 km from Pretoria on Roodeplaat Dam road, ( CB), 28 Jan. 1980, Retief & Herman 159 (MO, PRE); Tierpoort Vakansieoord, east of Pretoria, sandstone rock crevices and slopes, ( CD), 8 Mar. 1977, Van Jaarsveld 1870 (NBG); Donkerhoek, 20 miles [± 30 km], east of Pretoria, ( CD), 19 Mar. 1959, Codd 9908 (K, PRE); Premier Mine, ( DA), Jan. 1919, Rogers (K, MO, P, SAM), June 1921, Rogers (B, BR, G, GRA, K, LISC, MO, P, S, SAM); Renosterkop, Bronkhorstspruit Distr., ( DB), 7 Mar. 1932, Young 2116 (K, PRE); north of Bronkhorstspruit to Groblersdal, ( DB), 1 Feb. 1994, Goldblatt & Manning 9806 (MO). MPUMALANGA (Pilgrims Rest): 5.3 miles (± 8 km] from Burgersfort on Steelpoort road, 2500 ft [762 m], ( CB), 14 Mar. 1953, Story 4067 (GRA, K, PRE, SRGH) (Witbank): Loskop Dam, Nooitgedacht, shallow ground on summit, ( AD), 9 Feb. 1967, Theron 1214 (PRE); 5 miles west of Middelburg, ( CD), 30 Jan. 1929, Hutchinson 2716 (K, PRE); Botsabelo, Renosterpoort, mountain summit, ( CD), Mar. 1878, Nelson 402 (K, PRE) (Lydenburg): hills above Dullstroom, ( AC), 23 Feb. 1937, Van der Merwe 1255 (B, K, PRE); Dullstroom, among dolerite rocks on Farm Valleispruit, 6500 [1 920 m], ( AC), 30 Jan. 1933, Galpin (K, P, PRE); Belfast, Feb. 1909, ( CA), Doidge 4800 (K). NORTHERN CAPE (Kuruman): Mansfield Farm, among ironstone rocks, top of beacon hill, ( CB), 13 Mar. 1981, Gubb (PRE); Kuruman Hills, Brancote Farm, red sandy loam, m, ( CB), 31 Mar. 1988, Venter (PRE); Klipfontein Farm, sandy ground with ironstone pebbles, ( DC), Gubb (PRE); 7 miles [10.5 km] southwest of Kuruman, ( DC), 16 Mar. 1971, Wisura 2003 (NBG). 1b. subsp. limpopoensis Goldblatt & J.C.Manning, subsp. nov. TYPE. SOUTH AFRICA, Limpopo, 2327 (Ellisras): Waterberg, between Groothoek and Vaalwater, m, ( BD), 6 Jan. 1959, Werdermann & Oberdieck 1640 (PRE, holo.!; B!, BR!, GH, K!, MO!, WAG!). Like subsp. sandersonii but corms mm diam.; tunics sometimes becoming fibrous and netted with age. Spike with floral bracts 5 6 mm long. Flowers with perianth tube 8 12 mm long; tepals mm. Stamens with filaments ± 5 mm long, exserted mm. Style dividing near or beyond apex of anthers, branches ± mm long, divided for up to 1 / 3 their length. Distribution: subsp. limpopoensis is restricted to the western Limpopo Province of South Africa in the Waterberg Mtns and surrounding country (Figure 41). As outlined above it bears a superficial similarity to the mainly tropical African Psilosiphon erythranthus because of its relatively short perianth tube. The dark, tough, non-reticulate corm tunics, intricate branching pattern and rather dark floral bracts all accord better with the more widespread P. sandersonii subsp. sandersonii. Representative specimens SOUTH AFRICA. LIMPOPO (Ellisras): Waterberg 67 miles [± 101 km] north of Vaalwater on Beauty road, ( DB), 3 Feb. 1983, Reid 656 (PRE); Waterberg, 30 km west of Vaalwater, ( BD), 12 Mar. 1990, Balkwill et al (J, MO) (Baltimore): Doornleegte Farm, 6.5 km from main gate of Lapalala Wilderness, 29 Mar. 2004, Bredenkamp 3009 (PRE) (Nylstroom): Vlakfontein, 3 miles north of P.O. Palala, ( BC), 29 Jan. 1960, Codd 9975 (PRE); 16 km from Palala on road to Bamboeskloof, ( BC), 9 Mar. 1978, Germishuizen 735 (PRE); Sterkrivier Nature Reserve, rocky slope in mixed woodland, below slabs of bedrock, ( BD), 13 Mar. 1973, Jacobsen 2794 (PRE); Nylstroom, Heuningfontein, Farm Donkerkloof, woodland on sand, ( AD), 13 Feb. 1987, Westfall 2231 (PRE). 2. Psilosiphon avasmontanus (Dinter) Gold blatt & J.C.Manning, comb. nov. Lapeirousia avasmontana Dinter in Repertorium specierum novarum regni vegetabilis 29: 256 (1931); Goldblatt: 447 (1990a). Type: Namibia, Lichtenstein, Auas Mtns., 20 Feb. 1923, Dinter 4454 (B, holo.!; GH!, S!, Z [3 sheets]!, iso., K!, [photo of specimen at B]).

123 S T R E L I T Z I A 35 (2015) 115 Plants mm high, ± divaricately branched in upper half, stem rounded to slightly triangular in section, sometimes obscurely winged or ridged at one or more angles. Corm bell-shaped, mm diam.; tunics dark brown to blackish, inner layers ± woody, outer decaying irregularly, often into vertical strands. Leaves 3 or 4, linear, 2 3 mm wide, main vein prominent, lowermost inserted near ground level and longest, usually slightly exceeding inflorescence, others inserted above ground and progressively smaller above, those subtending branches ± bract-like. Inflorescence a rounded, ± flat-topped false panicle, ultimate branches with 1(2) flowers; bracts 5 6 mm long, green below, membranous in upper half and bent outwards by tepals, becoming completely dry in fruit, inner ± as long as or slightly longer than outer. Flower actinomorphic, blue to light purple with median white heart-shaped mark feathered purple on edges in basal half of each tepal; perianth tube ± 1 mm long, cylindric below, widening in upper 1 / 2 ; tepals spreading but curving upwards near tips, subequal, lanceolate to elliptic, (4 ) 5 7 mm. Stamens symmetrically arranged; filaments 4 5 mm long, diverging distally; anthers diverging, 4 5 mm long, apices curving inward after dehiscence; pollen light blue-purple to ± white. Style straight, 6 7 mm long, dividing near anther apices, branches ± 1.2 mm long, barely notched apically. Capsules ± globose, weakly 3-lobed, showing outline of seeds, 6 8 ± 7 mm. Seeds globose, dark brown, mm diam., rounded to weakly angled by pressure, tapering toward attached funicle, surface smooth. Chromosome number 2n = 16. Flowering time: February to April. Figure 42. Distribution and biology: endemic to central interior Namibia, Psilosiphon avasmontanus is locally common on the hills and mountains around Windhoek, extending north toward Okahandja and east to Wit Vlei (Figure 41). Apparently preferring well-drained sites, it grows in open, stony, sloping ground or sometimes in light scrub. Depending on rainfall, flowering may last between 3 and 8 weeks. New branches continue to be produced from the cauline leaf axils as long as the ground remains moist, and late rainfall can stimulate production of a second flush of flowering from new branches. The pale blue flowers are visited by a variety of insects including bees, wasps and butterflies. We were unable to detect nectar in excised flowers and we assume that the only reward to the bees and wasps is pollen and that butterfly visits are exploratory. Diagnosis and relationships: the relatively large, pale blue flowers with a white centre, radially symmetric perianth and stamens, and large, ± flat-topped inflorescence distinguish Psilosiphon avasmontanus from other tropical and subtropical African species of the genus. It is most easily confused with the more widespread P. coeruleus, which has similar, but smaller, actinomorphic flowers. P. avasmontanus has dark brown ± woody corm tunics that break into vertical strips as they decay and a divaricately branched inflorescence, the terminal branches of which bear a single (rarely 2) flowers. Corm tunics in P. coeruleus are light brown, becoming finely fibrous and netted as they decay, and the inflorescence is less strongly branched with ascending branches, some of which terminate in spikes of 3 5 flowers. The flowers of P. avasmontanus have relatively larger, pale blue tepals, mm long, with heart-shaped white markings, whereas P. coeruleus has smaller tepals, 8 9 mm long, with spearshaped white markings. Although they are the only two species of Psilosiphon in Namibia with radially symmetric flowers, chromosome cytology confirms that P. avasmontanus and P. coeruleus are not immediately related. P. avasmontanus is diploid with 2n = 16 and the strongly bimodal karyotype consists of 1 long and 7 short chromosome pairs (Goldblatt 1990b), a pattern quite different from P. coeruleus, which has 2n = 8 and a karyotype of 2 long and 2 shorter chromosome pairs. Despite the apparent numerical polyploid relationship, their very different karyotypes have about the same total chromosome length (a proxy for total DNA per cell), making it clear that a polyploid relationship is not involved here. The similar corm tunics and dichotomously branched inflorescences of P. avasmontanus and the eastern southern African P. sandersonii suggest to us that the two are closely related. P. sandersonii also has a bimodal karyotype, in this case consisting of 1 long and 5 much smaller chromosome pairs. History: Psilosiphon avasmontanus was discovered in 1923 by Kurt Dinter and described by him in 1931 as Lapeirousia avasmontana. In

124 116 S T R E L I T Z I A 35 (2015) B C A FIGURE 42. Psilosiphon avasmontanus, Goldblatt & Manning A, plant and corm; B, flower; C, stamens and style. Scale bar: A, 10 mm; B, 5 mm; C, 2 mm. Artist: John Manning.

125 S T R E L I T Z I A 35 (2015) 117 the protologue he compared the species to the superficially similar P. coeruleus, which, as Dinter pointed out, grows in a different habitat and differs in several vegetative and floral features. Although P. avasmontanus was included in P. coeruleus (as Lapeirousia) in the Prodromus einer Flora von Südwestafrika by Sölch (1969). Goldblatt (1990a) treated them as separate species. Representative specimens NAMIBIA (Otjimbingwe): Farm Onduno, Hochflaeche, ( BD), 27 Feb. 1966, Meyer 115 (MO, WIND); Farm Terra Rossa, rocky slope, ( CD), 13 May 1973, Giess (K, M, NBG, PRE, WAG, WIND); Farm Friedenau, ( DB), 25 Mar. 1982, Müller & Kolberg 2039 (PRE, WIND) (Windhoek): Elisenheim, Erosberge, below Wächter, smooth shale slopes, ( AC), 28 Feb. 1974, Merxmüller & Giess (K, M, S, PRE, SRGH, WAG, WIND); near Brakwater on the Windhoek Okahandja road, ( AC), 14 Mar. 1959, De Winter & Giess 7136 (M, PRE, WIND); 5 km west of Windhoek on Daan Viljoen road, stony quartz hill slopes, ( AC), 14 Mar. 1988, Goldblatt & Manning 8798 (E, K, M, MO, NBG, PRE, S, WAG, WIND) (Gobabis): sandy grassland at Wit Nossob River between Seeis and Witvley, ( AC), 7 Mar. 1997, Germishuizen 9504 (MO, PRE). 3. Psilosiphon gracilis (Vaupel) Goldblatt & J.C.Manning, comb. nov. Lapeirousia gracilis Vaupel in Botanische Jarhbucher für Systematik 48: 548 (1912); Sölch 155: 9 (1969); Goldblatt: 465 (1990a). Type: Namibia, Great Namaqualand, Doorns, dolomite, m, Mar. 1907, Range 292 (B, holo.! [(K, photo!]; M! [fragment], iso.). Plants mm high, often slender, stem 2 several branched, main axis dominant, branches ascending, flexed at base of main spike, 2-angled and winged below first branch, 3-angled or ± winged above. Corm bell-shaped, mm diam.; tunics light brown, of layers of compacted fibres, becoming loosely fibrous with age, fibres extending upwards as short, soft spines. Leaves 2 or 3(4), lowermost inserted just below ground, linear to narrowly lanceolate, 3 5( 8) mm wide, ± as long as or somewhat longer than inflorescence, thickened around main vein, second leaf usually largest, upper leaves decreasing in size, becoming bract-like. Inflorescence a spike or somewhat panicle-like, main terminal branches 2 6-flowered; bracts 3 6( 7) mm long, soft, green below, becoming ± membranous above, sometimes flushed purple, or membranous and dry before anthesis, then usually transparent with fine brown veins, subequal or outer slightly larger. Flowers zygomorphic, pale blue to mauve or white, then often with blue tube, lower three tepals white or pale yellow in lower third, edged distally with darker blue to violet, occasionally one or more lower tepals with a small, median tooth-like ridge, lightly fragrant [also said to be unpleasantly scented]; perianth tube cylindric, slightly curved outwards at apex, (18 )25 38 mm long; tepals lanceolate, dorsal largest and held apart, suberect to reflexed, mm, margins slightly undulate, lower three slightly smaller, held close together. Stamens unilateral; filaments exserted ± 2 mm; anthers 4 5 mm long, pale lilac, blue or ± white; pollen cream. Style arching over to filaments, dividing between middle and tips of anthers, branches mm long, divided for ± 1 / 4 their length. Capsules ± globose, obtusely trigonous, mm. Seeds globose to weakly angular, ± 2 mm diam., dark brown, smooth. Chromosome number 2n = 12. Flowering time: mainly late January to April (May), also recorded in October. Distribution and biology: restricted to western Namibia, Psilosiphon gracilis extends in a relatively broad band along the western part of the country from the Fish River Canyon in the south to the Hoanib River in the Kaokoveld in the northwest (Figure 43). It favours rocky sites, often on limestone, dolomite or other calcareous rock types, but is also said to grow on sandy flats and, especially in more arid places, is often associated with springs and places where additional water is available to supplement the sparse rainfall. Flowering normally occurs at the end of summer, mostly in February and March, but sometimes as late as May. One collection, Craven 1341, from the Brandberg and blooming in October, is difficult to explain, and perhaps indicates an unusual pattern of rainfall that particular year. The spring-blooming plants do not differ in any way from those flowering in late summer. The relatively long perianth tube, up to 38 mm long, pale perianth and floral scent suggests that the flowers are adapted for pollination by moths or butterflies with relatively long probosces. Diagnosis and relationships: the slender perianth tube, typically mm long (occasionally slightly longer or shorter), either pale

126 118 S T R E L I T Z I A 35 (2015) FIGURE 43. Distribution of Psilosiphon erongoensis, ; P. gracilis,. blue or white perianth, usually short floral bracts, 3 6( 7) mm long and comparatively slender habit with up to 6 flowers per terminal branch make Psilosiphon gracilis relatively easy to recognise. It most closely resembles P. bainesii and P. otaviensis, both of which also have long-tubed flowers, but P. bainesii has white to pale pink flowers with dark red to brown markings, a tube mm long, and a more robust habit, longer floral bracts, 7 12 mm long, and the highly ramified inflorescence has only 1(2) flowers per ultimate branch. Confusion with P. otaviensis is more likely as, like P. gracilis, it has an inflorescence with the main terminal branches bearing 2 5 flowers. The flowers of P. otaviensis are unscented, larger and have a perianth tube somewhat longer, mm, and longer floral bracts, (7 )8 11 mm long. The flowers of P. gracilis are usually pale blue but fade after drying and appear white, but in a few collections the flower colour is noted as being white. Phylogenetic relationships are difficult to assess with confidence in Psilosiphon but Goldblatt (1990a) postulated that P. gracilis, P. bainesii and P. otaviensis were probably closely allied, sharing derived pale-coloured flowers with an extended perianth tube and light brown, fibrous corm tunics. Countering this is the chromosome cytology (Table 4). P. gracilis has 2n = 12 and a bimodal karyotype (Goldblatt 1990b), quite different from L. otaviensis and L. bainesii, which have 2n = 10 (or 6 in some populations of the latter), nonbimodal karyotypes and are cryptic polyploids with a DNA amount per cell ± twice that in P. gracilis. A closer relationship with the long-tubed L. erongoensis of central Namibia and L. masukuensis of eastern southern Africa now seems more likely. Chromosome number of both these species is unknown. History: evidently first collected by the German geologist, Paul Range, in 1907 near Doorns in southern Namibia, this collection formed the basis for Lapeirousia gracilis, described in 1912 by Vaupel. It remained poorly known for many years, but botanical exploration in Namibia since the 1960s has filled in its range so that its distribution now forms a coherent pattern. Representative specimens NAMIBIA (Hoanib Mouth): Hoanib river valley ± 20 km inland, rocky hills among boulders, ( BD), 20 Apr. 1985, Moss & Jacobsen K311 (PRE, WIND) (Uis): Brandberg, ( AB), 17 Oct. 1981, Craven 1341 (WIND); Brandberg, Orabeswand, m, ( AB), 6 Apr. 1964, Nordenstam 3666 (M, S) (Karibib): Farm Klein Spitzkoppe, coarse granitic sand, western slopes and below the summit, ( CC), 23 Feb. 1965, Geiss 8496 (B, K, M, MO, PRE, S, WAG, WIND) (Trekkopje): Farm Tsabichab, cracks in marble rock around spring, ( BA), 19 June 1973, Giess (K, M, MO, S, WIND) (Otjimbingwe):

127 S T R E L I T Z I A 35 (2015) 119 Namibrand, Karibib, Otjosandu, ( AA), 1 Mar. 1963, Seydel 3419 (B, C, G, GH, K, MO, S, SRGH, WAG, WIND) (Nauchas): Farm Weisenfels, ( AD), 11 Mar. 1953, Walter & Walter 1675 (B, WIND) (Mariental): Farm Haribes, dry sterile vlei, ( DA), 1 Mar. 1963, Giess et al (M, PRE, WIND) (Maltahohe): north-northwest of Maltahohe on district road D850 to Bullspoort, ( DB), 11 Mar. 1995, Smook 9337 (MO, PRE, WIND) (Helmeringhausen): Farm Chamchawib, ( DB), 23 Jan. 1974, Giess & Robinson (K, M, MO, PRE, WAG, WIND) (Gibeon): Asab, hard clay along the rail tracks, ( BD), May 1913, Dinter 3061 (SAM), 13 Mar. 1988, Goldblatt & Manning 8770 (K, MO, NBG, PRE, S, WIND) (Aus): westsouthwest of Bethanien loop road D437, ( BC), 15 Mar. 1995, Smook 9468 (PRE, WIND) (Keetmanshoop): Spitzkoppe, ( AD), 7 Apr. 1984, Craven 1666 (WIND) (Chamaites): Fish River Canyon, river camp, ( DA), 30 Mar. 1953, Walter & Walter 2263 (B, WIND). 4. Psilosiphon erongoensis Goldblatt & J.C.Manning, sp. nov. TYPE: Namibia, 2115 (Karibib): Erongo Mtns, western edge of the Erongo Mtns above Farm Ameib, ( DA), 27 Mar. 1976, Craven & Craven 115 (WIND, holo.; PRE, iso.). Plants mm high, repeatedly branched, main axis ± straight, dominant, lateral branches diverging at ± 30 ; stem weakly compressed below, triangular in middle part and rectangular above, angles slightly winged. Corm mm diam.; tunics light brown, coarsely fibrous. Leaves 4 7, ± linear, grey-green, main vein prominent, lowermost 2 basal and longest, mostly 4 5 mm wide, reaching to ± apex of inflorescence, upper leaves cauline and decreasing in size above, those subtending branches becoming ± bract-like. Inflorescence ± a rounded to flat-topped false panicle, main terminal branches with (2 )4 or 5 flowers in short spikes; bracts 8 10 mm long, soft-textured, ± green with translucent, membranous margins, lanceolate, acute, becoming dry apically, inner slightly shorter than outer. Flowers zygomorphic, violet-blue, lower three tepals with white mark near base edged dark violet distally,?unscented; perianth tube mm long, cylindric, expanded and curved in upper 3 mm; tepals subequal or dorsal slightly larger, lanceolate, ± acute, margins evidently straight, ± mm, lower 3 united for 1.5 mm. Stamens unilateral, ± erect; filaments ± 8 mm long, exserted ± 5 mm; anthers 5 6 mm long, evidently yellow; pollen yellow. Style nearly straight, dividing opposite anther tips; branches ± 3 mm long, forked for ± 1 / 3 their length, violet. Capsules and seeds unknown. Chromosome number unknown. Flowering time: March and April. Distribution and biology: Psilosiphon erongoensis is evidently restricted to the Erongo Mtns of west central Namibia (Figure 43). We suspect that it grows in pockets of soil around the granite domes that characterise the area, where additional water is available to plants in this arid terrain. Other details of its biology remain to be studied. Diagnosis and relationships: included in Psilosiphon otaviensis by Goldblatt (1990a), the violet flowers and slightly shorter perianth tube, ± 37 mm long, and shorter, narrower tepals, ± mm, serve to distinguish P. erongoensis from that species, which has creamy white flowers with violet longitudinal markings on the lower tepals, a somewhat longer perianth tube, mm long, tapering gradually from base to apex, and tepals mm. The branching of the inflorescence and general aspect of the two species differ hardy at all. However, we now think P. erongoensis may be more closely related to P. gracilis, which has pale blue or less often white flowers, a perianth tube (18 )25 38 mm long and, significantly, short, pale floral bracts 4 6( 7) mm long (vs mm in P. erongoensis). Chromosome number, useful in determining relationships in Psilosiphon, is unknown and so far we have been unable to obtain material for DNA extraction and molecular study. Additional specimen NAMIBIA (Karibib): Erongo Plateau, among granite boulders, ( DC), 1985, Lavranos (E, K, M, MO, P, PRE, S). 5. Psilosiphon rivularis (Wanntorp) Goldblatt & J.C.Manning, comb. nov. Lapeirousia rivularis Wanntorp in Svensk Botanisk Tidskrift 65: 53 (1971); Roessler: 394 (1979); Goldblatt: 448 (1990a). Type: Namibia, Karibib, Ameib Ranch, ± 25 km northeast of Usakos, granite kopje [koppie] south of track, ± 5 km east of farmhouse, 15 Apr. 1968, Wanntorp & Wanntorp 907 (S, holo.!; M!, iso.).

128 120 S T R E L I T Z I A 35 (2015) Plants mm high, laxly fewbranched, stem weakly compressed and 3 4- angled. Corm bell-shaped, 8 12 mm diam.; tunics dark brown, fibres medium-textured, reticulate. Leaves mostly 2(3), lower 1 or 2 ± basal, linear, 2 3 mm wide, reaching to ± middle of inflorescence, firm, main vein lightly raised, lowermost longest, upper leaves subtending branches, progressively shorter. Inflorescence a laxly branched spike, main axis mostly laxly 2 5(6)-flowered; bracts 4 5 mm long, green below, membranous in upper half, apices dry becoming light brown, inner bract ± as long as outer, entire or shallowly forked at tip. Flowers zygomorphic, pale blue with white nectar guides on lower three tepals outlined in blue-violet, with tooth-like median ridge up to 1 mm long, distal half of throat lined dark blue on abaxial side; perianth tube ascending, widening gradually from base, ± 1.3 mm wide at base, 3 mm at mouth, (5 )7 9 mm long, ± straight; tepals narrowly lanceolate, margins usually straight, subequal, (7 ) mm wide, dorsal tepal suberect to arching forward, held apart from others, lower tepals directed forward, nearly horizontal, united basally for ± 1.5 mm. Stamens unilateral and arcuate; filaments ± 7.5 mm long, exserted ± 4 mm; anthers parallel, contiguous, ± 4 mm long, pale blue-mauve; pollen pale bluemauve. Style suberect, adaxial to stamens, ± 15 mm long, dividing opposite upper 1 / 3 of anthers; branches ± 1 mm long, undivided or notched at apex, spreading. Capsules mm long, topshaped in profile, 3-lobed in distal half, with 2 or 3 seeds per locule. Seeds ± globose, dark brown, ± 2 mm diam. Chromosome number unknown in Namibian populations, 2n = 12 (in one Zambian population). Flowering time: mostly late February to early April. (Description based only on plants from Namibia). Distribution and biology: primarily a plant of seasonally wet depressions on rock outcrops and seasonally waterlogged grassland on shallow soils, Psilosiphon rivularis occurs across the northern half of Namibia into adjacent Angola (Figure 44). In Namibia it is particularly common in the Erongo Mtns, a range characterised by massive granite outcroppings, and also occurs on the Brandberg, the Paresis Mtns near Otjiwarongo and north of Etosha Pan. Taller, more robust plants, sometimes with larger flowers from Zambia and eastern Angola were included here by Goldblatt (1990a), but may represent a separate species. Diagnosis and relationships: an often diminutive plant, Psilosiphon rivularis is recognised by the pale blue flowers with a relatively narrow perianth tube mostly 7 9 mm long, tepals usually mm long (occasionally smaller), the lower FIGURE 44. Distribution of Psilosiphon coeruleus, ; P. erythranthus, ; P. masukuensis, ; P. rivularis,, in southern Africa.

129 S T R E L I T Z I A 35 (2015) 121 three bearing a white marking edged in darker blue and often a prominent tooth-like ridge in the lower midline. The style, ± 15 mm long, divides into short, undivided branches ± 1 mm long. Plants at the type area, the Erongo Mtns, rarely exceed 200 mm in height, but elsewhere in Namibia may reach up to 270 mm. Goldblatt (1990a) included populations from Zambia and central Angola in L. rivularis. These plants, sometimes up to 450 mm tall, have larger flowers, which lack ridges on the lower tepals, and among other differences have a longer style, mm long and style branches 2 3 mm long, divided for ± one third to half their length in contrast to the undivided style branches of P. rivularis. More field work is needed to establish the status of the Zambian populations, which may represent another taxon. Zambian plants have a diploid chromosome number of 2n = 12, suggesting a relationship with the widespread P. erythranthus, which has the same number in south-tropical African populations. Chromosome number is not known for P. rivularis in Namibia. History: Psilosiphon rivularis was described by the Swedish botanist Hans-Erik Wanntorp in 1971, although he was not the first to record the species. It appears to have been collected in Namibia by the South African biologist, K.H. Barnard, on the northern border of Namibia along the Cunene River in The type locality of L. rivularis, the Farm Ameib at the southern end of the Erongo Mtns, is its southernmost known station. Representative specimens NAMIBIA (Ondangua): Engela Mission, shallow vlei in black sticky clay, ( BD), 16 Feb. 1959, De Winter & Giess 7061 (B, K, M, PRE, SRGH, WIND); Uutapi District, Uutapi Ogongo road, in standing water, ( CA), 27 Mar. 2006, Kangombe & Kruger 269 (WIND) (Otjiwarongo): Otjiwarongo, not on summit but high up, 16 Mar (BC), Craven 1129 (WIND) (Uis): Brandberg, Sonuseb, common on top of Sonuseb saddle at water hole, ( BA), 12 Apr. 1985, Craven 2277 (WIND) (Karibib): granite koppies south of Gross Spitzkoppe 71, wet granite rocks, ( CC), Sullivan 418 (WIND); Farm Ameib, below Jatow cave, somewhat marshy, granite soil, ( DC) 19 Mar. 1963, Giess (B, M, PRE, WIND); 22 Mar. 1965, Giess 8452 (K, M, MO, WAG, WIND); 17 Mar. 1963, Giess et al (B, M, PRE, WIND); Farm Ameib, coarse sandy, marshy ground below granite outcrops at the Devils Pulpit, ( DC), 18 Mar. 1968, Giess (M, WIND); 5 Apr. 1974, Merxmüller & Giess (B, M, K, PRE, SRGH, WAG, WIND). [Exsiccatae assigned to Psilosiphon rivularis from Angola and Zambia are cited in Goldblatt (1990a).] 6. Psilosiphon coeruleus (Schinz) Goldblatt & Manning, comb. nov. Lapeirousia coerulea Schinz in Verhandlungen des botanische Vereins der Provinz Brandenburg 31: (1890); Goldblatt: 445 (1990a). Type: Namibia, (Upingtonia), Ombale, südost Ondonga, Mar. 1886, Schinz 13 (K, lecto.!, designated by Goldblatt: 445 (1990a); COI!, isolecto.). Ixia dinteri Schinz: 14 (1900). Type: Namibia, Karibib, Spitzkop, marshy ground, in 1898, Dinter 22 (32 on the type) (Z, holo.!). Plants (120 ) mm high, repeatedly branched, stem compressed, 2- or 3-angled. Corm bell-shaped, mm diam., light brown; tunics of pale compacted fibres decaying to become coarsely to finely fibrous and netted. Leaves 2 4, only lowermost inserted at base, this longest and reaching to at least middle, and sometimes shortly exceeding inflorescence, ± linear, 2 4( 5) mm wide, remaining leaves decreasing in size above. Inflorescence a laxly branched, round-topped false panicle, larger ultimate branches (2)3 5-flowered; bracts 3 4 mm long, subequal, green below, membranous distally, becoming completely dry in fruit, inner ± as long as outer. Flower actinomorphic, ± rotate, blue to light purple with white, spear-shaped mark outlined in dark blue to purple on basal 1 / 2 of each tepal; perianth tube mm long, narrowly funnel-shaped; tepals spreading below, curving upward distally, subequal, lanceolate, mm. Stamens symmetrically arranged; filaments united basally for < 0.5 mm by an obscure corona-like ring, erect, ± 4 mm long, nearly contiguous around style; anthers diverging, mm long, curving inwards after anthesis; pollen pale blue-purple (white to yellow when dry). Style erect, 6 7 mm long, dividing near anther tips, style branches ± 1.2 mm long, shallowly notched apically. Capsules obovoid globose, weakly 3-lobed, ± 4 mm long; seeds dark brown, globose to weakly angled by pressure, mm diam., tapering near attached funicle. Chromosome number 2n = 8. Flowering time: (January)February to April, occasionally in December. Figure 45.

130 122 S T R E L I T Z I A 35 (2015) Distribution and biology: typically a species of seasonally wet habitats, Psilosiphon coeruleus is widespread throughout the summer rainfall part of Namibia, extending locally into northwestern Botswana, where it has only occasionally been reported (Figure 44). Plants grow in seeps, seasonal marshes, poorly drained grassland and shallow soil in rock outcrops where water accumulates in the wet season. The pollination biology is unknown. The flowers produce no detectable nectar and the only potential pollinators we have seen are small bees (family unknown) nestling in the open flowers. D C FIGURE 45. Psilosiphon coeruleus, Goldblatt & Manning 8811A. A, plant and corm; B, flower; C, stamens and style; D, capsule. Scale bar: A, D, 10 mm; B, C, 5 mm. Artist: John Manning. A B Several collections are accompanied by notes indicating that corms of Psilosiphon coeruleus were in the past eaten by the indigenous population, particularly the!kung, for whom the corms may have been a staple, eaten after roasting in hot ash (Story 6121), or sometimes raw (Rodin 1985). Diagnosis and relationships: Psilosiphon coeruleus, with its radially symmetric flower and short perianth tube mm long, and undivided style branches appears to be taxonomically isolated in the genus. The chromosome number, 2n = 8, recorded in seven populations, and small corm with dark brown, fibrous tunics match other blue-flowered species including P. erythranthus (some populations 2n = 12) and P. rivularis (2n = 12) suggesting a relationship with this group of species, which have zygomorphic, bilaterally symmetric flowers, often of the same colour, although sometimes red in P. erythranthus. One other species of Psilosiphon has radially symmetric flowers, P. avasmontanus, but this more robust species has a different inflorescence with the ultimate branches bearing 1 or 2 larger flowers, and dark brown corm tunics that fragment into vertical strips, like those of P. sandersonii. Its chromosome number, 2n = 16, with a karyotype consisting of one long and seven short chromosome pairs make P. avasmontanus an unlikely close relative of P. coeruleus. The differences between Psilosiphon coeruleus, a species of wet low-lying places, and P. avasmontanus, which grows in well-drained, stony banks and hills, are discussed in detail under P. avasmontanus. The morphological differences, combined with their disparate habitat preferences and chromosome cytology, compel their separation despite the alternative treatment by Sölch (1969) uniting them.

131 S T R E L I T Z I A 35 (2015) 123 The simple flower structure is deceptive radially symmetric flowers are specialised and derived in the genus. Chromosome number in P. rivularis from Namibia, retrieved as sister to P. coeruleus in the plastid and combined plastid and nuclear DNA-based phylogeny (Figure 1), is unknown. History: Psilosiphon coeruleus was first collected in 1879 by the Swedish traveller-explorer, Ture Een, on his second expedition to Namibia (Gunn & Codd 1981). This early record received no attention, and later collections made independently after 1885 by Hans Schinz, August Lüderitz and Waldemar Belck formed the basis for the protologue of Lapeirousia coeruleus published by Schinz in Schinz (1900) also described Ixia dinteri, a species undoubtedly conspecific with P. coeruleus, based on specimens collected by Kurt Dinter in Representative specimens BOTSWANA (Aha Hills): Xangwe, ( CB), Mar. 1961, Gibson 186 (MO, WIND); Dobe region, north of Aha Hills near Namibian border, dry pan, ( CB), 25 Apr. 1980, Smith 3496 (MO, PRE, SRGH); hard clay in dry pan, Quangwa River catchment, ( CB), 23 Apr. 1981, Smith 3674 (BR, PRE) (Kobe): pan on Farm 102, Kuki, ( AC), 21 Feb. 1970, Brown & Brown 8723 (C, PRE, SRGH) (Okwa): 53 km northeast of Ghanzi on road to Maun, seasonally flooded limestone outcrops, ( AB), 29 Dec. 1977, Skarpe S-213 (K, MO, PRE, UCBG). NAMIBIA (Ondangua): 3 km south of Oshikango, ( BD), 3 Apr. 1973, Rodin 9203 (K, M, MO, PRE, WIND) (Okahakana): Etosha Pan, large vlei 5 miles [± 7.5 km] west of Okondeka, ( DD), 27 Mar. 1963, Giess et al (M, WIND) (Tsumeb): 25 km from Tsumeb towards Ondangua, ( BA), 14 Feb. 1983, Lavranos & Pehlemann (WIND); Farm Toevlug, lime marl, ( BA), 5 Feb. 1971, Giess (K, MO, PRE, SRGH, WAG, WIND); Farm Goab, spring meadows, ( CA), 29 Jan. 1978, Giess (M, MO, PRE, WAG, WIND) (Grootfontein): Grootfontein, red loamy flats, ( CA), Jan. 1935, Schoenfelder 35 (K, PRE) (Tsumkwe): pans at Tsumkwe, ( DA), 14 Jan. 1971, Giess et al (M, S, PRE, WIND); Tsumkwe, ( DA), 14 Jan. 1958, Story 6121 (M, PRE, SRGH) (Otjiwarongo): 30 km south of Otjiwarongo, ( CB), 1985, Lavranos (MO) (Waterberg): omuramba [Herero: ancient river bed] at Omam, ( CA), 11 Mar. 1940, Volk 2845 (WIND) (Karibib): Ohere-Oos, granite flats, ( BA), 14 Feb. 1958, Merxmüller & Giess 1583 (M, PRE, WIND); Ameib Ranch, granite rock flushes at Bull Parties, ( DC), 15 Mar. 1988, Goldblatt & Manning 8811A (M, MO, PRE, WAG, WIND) (Steinhausen): Farm Wilhelmsruhe, ( DC), 10 Feb. 1982, Rauh (WIND) (Trekkopje): Okongawa, Granitbankberg, ( BB), 4 Feb. 1934, Dinter 6957 (B, BOL, G, K, HBG, M, PRE, S, WIND, Z) (Okahandja): Farm Omatako View, ( BA), 15 Feb. 1974, Woortman 116 (M, PRE, WIND); Farm Omongongua, shallow depressions in omuramba, ( DB), 27 Mar. 1960, Seydel 2203 (B, K, HUH, MO, WIND); Farm Okambahe, ± 16 miles [24 km] west of Okahandja, road to Karibib, ( DC), 21 Dec. 1963, Giess et al (M, MO, PRE, WIND). 2215: (Otjihorongo): Farm Goreis, red loam sand, 17 Feb ( BB), Giess 11248A (K, M, PRE, WIND) (Windhoek): Neudam Experimental Farm, ( AD), 22 Mar. 1960, Van Vuuren 1027 (K, M, PRE, SRGH, WIND); Farm Otjikundua 67 miles [± 100 km] west-southwest of Steinhausen, open vlei, ( BA), 19 Feb. 1955, De Winter 2409 (K, M, NBG, PRE, WIND); Farm Aris, south of Windhoek, omuramba, ( CA), 1 Mar. 1953, Walter & Walter 1550 (B, BR, WIND) (Nauchas): Farm Göllschau, ( BC), 27 Jan. 1972, Giess & Hübsch (M, PRE, WIND) (Rehoboth): Farm Gravenstein, ( BC), 20 Nov. 1956, Volk (M, MO, WIND); Farm Tsumis, sandy soil, ( CA), Müller 1396 (M, PRE, WIND) (Keetmanshoop): Spitzkoppe, ( AD), 7 Apr. 1984, Craven 1533 (WIND). Without precise locality: Damaraland, 1879, Een s.n. (BM). 7. Psilosiphon erythranthus (Klotzsch ex Klatt) Goldblatt & J.C.Manning, comb. nov. Ovieda erythrantha Klotzsch ex Klatt in Peters, Naturwissenschaftliche reise nach Mossambique. Botanik 6(2): 516, t. 58 (1864). Lapeirousia erythrantha (Klotzsch ex Klatt) Baker: 155 (1877); Geerinck et al.: (1972), excluding var. teretifolia and var. welwitschii; Goldblatt: 451 (1990a). Type: Mozambique, Boror, Caboceira, Peters s.n. (B, holo.!). [See full synonymy in Goldblatt (1990a)]. Plants (150 ) mm high, branched repeatedly; stem compressed, 2-angled below, 3-angled above. Corm ± bell-shaped, 8 16 mm diam.; tunics light to dark brown, of densely compacted fibres, outer layers becoming coarsely fibrous, ultimately forming a matted mass. Leaves 3 or 4, lower 2 or 3 longest and 1 / 3 to 2 / 3 as long as stem, ± linear to lanceolate or ± falcate, (2 ) 4 8( 11) mm wide, main vein prominent when alive; upper leaves becoming progressively bractlike. Inflorescence a several to many branched false panicle, often ± flat-topped, ultimate branches mostly (2)3 6-flowered, flowers crowded terminally; bracts 3 6 mm long, green below, dry and membranous above, becoming entirely

132 124 S T R E L I T Z I A 35 (2015) dry and transparent, brownish in upper 1 / 3, outer obtuse, inner acute or apically forked, usually slightly shorter than outer. Flowers zygomorphic, either blue-violet with spear-shaped white mark outlined in dark blue to purple on lower tepals or crimson red, then without markings or lower tepals each with white or dark red median streak; perianth tube (6 )7 11( 15) mm long, slender below, expanded and slightly curved near throat; tepals subequal, lanceolate-spathulate, acute to obtuse, (6 ) ( 4) mm, dorsal held apart, reflexed to lie ± same plane as lower tepals; lower 3 tepals narrowed at base thus ± clawed, set closely together and joined for ± 1 mm. Stamens unilateral, ± erect; filaments 4 6 mm long, exserted mm from tube; anthers mm long, pale bluish to white, purple or red; pollen ± white. Style arched to erect, adaxial to stamens, dividing opposite middle to apex of anthers, ± 15 mm long; branches mm long, forked for 1 / 3 1 / 2 their length, recurved. Capsules ± globose in profile, 3-lobed, 3 4 mm long, showing outline of seeds. Seeds 2 5 per locule, ± globose, mm diam., dark brown, ± smooth to rugulose. Chromosome number 2n = 14, B (polyploids), 8 (diploids). Flowering time: late January to March, occasionally from mid-december. Figure 46. Distribution and biology: although restricted in southern Africa to eastern Botswana (Figure 44), Psilosiphon erythranthus has a wide distribution across tropical Africa (Goldblatt 1990a), extending from eastern Angola and southern Democratic Republic of the Congo, through Zambia to Zimbabwe, Malawi, western and southern Tanzania, and central Mozambique. A disjunct series of populations also occur in northeastern Nigeria. The most common species of Psilosiphon in central Africa, P. erythranthus, can frequently be found in rocky outcrops throughout its range, or less often in open grassland or woodland. Its frequency in thin soils in rocky sites suggests that the corms are protected in such habitats while in deeper ground they are accessible to rodents, porcupines and other herbivores. Cultivated plants grow well in almost any type and depth of soil. The small flowers, conspicuous in the manyflowered inflorescences, are visited and presumably pollinated by a variety of bees, wasps and even butterflies, which forage in large numbers in warm weather for nectar and/or pollen. Annotations on two specimens indicate that Psilosiphon erythranthus is edible (Simpathu 90, Victoria Falls; Buchanan 426, Shire Highlands eaten in time of great famine ). However, there appears to be no record of the species being significant in the regular diet of any human populations. Diagnosis and relationships: one of the several species of Psilosiphon with a strongly ramified, ± round- or flat-topped inflorescence, P. erythranthus can be distinguished by its relatively small flowers, typically (but not always) softtextured leaves with a conspicuous main vein, and flowers with a short perianth tube seldom exceeding 12 mm. The flowers are somewhat crowded at the branch tips, which are in effect spikes of up to 6 sessile flowers. The orientation of the tepals is also distinctive the tepals lie in ± the same plane, with the three lower tepals joined together for ± 1 mm and held closely together, while the upper tepal is reflexed and held apart from the others. Similar flowers also characterise P. sandersonii and P. masukuensis, both of which have a longer perianth tube, mostly at least 15 mm long and in some populations of P. sandersonii up to 30 mm long, and which also has different corms tunics and an inflorescence in which the ultimate branches have only one or two flowers. Notably P. sandersonii subsp. limpopoensis has a perianth tube 8 12 mm long, thus comparable with that of P. erythranthus, but the subspecies can immediately be distinguished by the branching pattern and different corm tunics. Less closely related, Psilosiphon rivularis is almost indistinguishable from P. erythranthus when pressed, but they have different flowers. In P. rivularis the upper tepal is arched forwards over the stamens and style branches. This important difference between the two species is generally obscured in herbarium specimens. P. rivularis grows in seasonally wet sites from central Zambia, across southern Angola to northern Namibia, and its range thus partly overlaps that of P. erythranthus. Other species of the Psilosiphon erythranthus complex outside southern Africa, including P. abyssinicus, P. setifolius and P. teretifolius, were discussed in depth by Goldblatt (1990a). History: Psilosiphon erythranthus was first collected by the German physician and explorer

133 S T R E L I T Z I A 35 (2015) 125 FIGURE 46. Psilosiphon erythranthus, no voucher. Scale bar: 10 mm. Artist: John Manning.

134 126 S T R E L I T Z I A 35 (2015) Wilhelm Peters in the years when he travelled in Mozambique under the Prussian King Friedrich Wilhelm IV s patronage. Later, the Scottish explorer, Sir John Kirk, who accompanied David Livingstone on his Zambezi expedition in 1858, also recorded the species in the lower Shire valley in Mozambique. The species was described by F.W. Klatt (in Peters 1864), who saw only the Peters collection, in the illegitimate genus Ovieda. Baker (1877) transferred O. erythrantha to Lapeirousia and in Flora of Tropical Africa he reserved L. erythrantha for red-flowered plants and referred those with blue flowers to L. sandersonii (Baker 1898). Later, plants from Zimbabwe currently included here, were assigned to L. rhodesiana by N.E. Brown (1906), while those from Mozambique were treated by Vaupel (1912) as L. graminea and L. plagiostoma. Other species, now regarded as synonyms of P. erythranthus include L. spicigera Vaupel from Angola and L. nigeriensis R.C.Foster (= L. montana Hutch.) from Nigeria (Goldblatt 1990a). The species deserves additional study, especially the larger-flowered populations from Zimbabwe that were treated as L. rhodesiana. Representative specimens BOTSWANA (Nata): Bakalaka area, between Francistown Maun road and Marapong, ( AC), 26 Jan. 1967, McClintock K73 (K); between Francistown and Nata River, on Maun road, ( AC), 21 Jan. 1959, West 3898 (BR, K, LISC, PRE) (Plumtree): Ramakqwebane area near Tsessebe Station, ( DC), 6 Jan. 1974, Ngoni 243 (MO, SRGH); border near Plumtree, sandy flat roadside, ( DC), 7 Mar. 1961, Richards (K, SRGH) (Palapye): Selebi, ( BB), Jan. 1978, Kerfoot 8007 (PRE). [Exsiccatae from tropical Africa are cited by Goldblatt (1990a).] 8. Psilosiphon masukuensis (Vaupel & Schltr.) Goldblatt & J.C.Manning, comb. nov. Lapeirousia masukuensis Vaupel & Schltr. in Vaupel, Botanische Jahrbucher für Systematik 48: 545 (1912); Goldblatt: 461 (1990a). Type: Mozambique. Inhambane: Masuku forest [Insilva Maseku], 10 Feb. 1898, Schlechter (B, holo.!; BR!, COI!, G!, K!, P!, PRE!, SAM!, iso.). Plants mm high, stem usually several-branched, compressed and 2-angled to winged below, 3-angled and slightly winged above branches. Corm bell-shaped, ± 15 mm diam.; tunics light brown, of ± leathery layers with reticulate surface, becoming loosely fibrous and netted with age. Leaves 4 6, lowermost inserted close to ground and longest, usually shortly exceeding inflorescence, linear-lanceolate, central vein prominent, 3 5 mm wide, remaining leaves progressively shorter above. Inflorescence a branched spike or round-topped false panicle, main axis ± straight, dominant, major ultimate branches 5 9-flowered, often crowded distally with bracts overlapping and (1.0 ) internodes long; bracts 5 7 mm long, green at anthesis, later dry and membranous, then either pale below or throughout or brownish above, streaked brown on veins, inner ± as long as outer or slightly shorter. Flowers zygomorphic, either dark blue to violet or greenish cream, lower tepals each with purple to red and white spear-shaped median streak in basal half; perianth tube cylindric, (15 )20 25 mm long, slightly expanded in upper 4 mm; tepals subequal with dorsal slightly larger, lanceolate, 8 10 ± 3 mm, spreading at right angles to tube and lying in ± same plane with dorsal reflexed and held apart. Stamens unilateral, ± erect; filaments ± 5 mm long, exserted mm; anthers 3.5 mm long, probably pale yellow when alive. Style unilateral, arching over stamens, dividing opposite middle to upper 1 / 3 of anthers; branches ± 2.2 mm long, divided for ± 1 / 2 their length. Capsules ± globose, slightly 3-lobed, mm diam., showing outline of seeds. Seeds globose, ± 2 mm diam. Chromosome number unknown. Flowering time: February to April. Distribution and biology: restricted to southeastern Africa, Psilosiphon masukuensis extends from central and southern Mozambique in the provinces of Inhambane and Maputo [Sul do Save] and adjacent Malawi south and west into southeastern Zimbabwe and lowland Limpopo and Mpumalanga of South Africa (Figure 44). The presence of P. masukuensis in Malawi was uncertain when Goldblatt s (1990a) account was published but we have now seen collections from there that match the species well. Collection data make it clear that P. masukuensis prefers relatively wet habitats during its growing season. Most specimens are recorded as growing in vleis, the edge of marshes, or in seasonally waterlogged flats. Pollination biology is unknown.

135 S T R E L I T Z I A 35 (2015) 127 The Mozambique populations are all coastal and probably from sandy habitats. Diagnosis and relationships: the comparatively tall stature, usually over 450 mm high, alternate branching pattern with the main axis dominant, ultimate inflorescence branches bearing 5 9 apically crowded flowers and the long straight perianth tube, usually mm long, are the main diagnostic characters of Psilosiphon masukuensis. Plants from the Mozambican coast have greenish flowers with red markings (Bandeira et al. 2007), unusual in the genus. In eastern South Africa, Zimbabwe and Malawi the flowers are shades of blue-mauve or violet with the white and dark blue markings typical of the P. erythranthus group, which we believe includes P. masukuensis. The long perianth tube and general appearance of the flowers, including their colour, suggest a relationship with P. sandersonii of eastern South African and Botswana, but this is typically a shorter plant, rarely exceeding 350 mm, with a divaricate, often intricately branched habit, and the ultimate branches of the inflorescence bearing 1 or 2, rarely 3 flowers. The corms of the two species also differ, those of P. masukuensis being relatively small and having the outer tunic layers fibrous and reticulate whereas the larger corms of P. sandersonii have dark brown tunics that decay irregularly into vertical strips. The corm tunics of P. masukuensis resemble closely the brown, fibrous and reticulate tunics of P. erythranthus to which it is probably most closely allied. Plants from South Africa and Zimbabwe are more variable than those from Mozambique, particularly in the length of the perianth tube and the number of flowers on the terminal branches of the inflorescence. Generally plants from Malawi, South Africa and Zimbabwe have 5 6 flowers per main branch compared with 6 9 in Mozambique. A notable example is Mauve 4326, 10 miles north of Abel Erasmus Pass, which has some flowers with a perianth tube just 15 mm long, while others have a tube up to 22 mm. Such apparently gross variation in perianth tube length may be due to ecological stress during the growing season or possibly some flowers could have opened while in a press without developing further. This degree of variability in tube length is sometimes encountered in collections of other tropical African Psilosiphon, particularly in P. bainesii. When more collections are available from Mozambique, the possibility that these represent a separate taxon from populations in South Africa and Zimbabwe should be reconsidered. The interior populations occur at higher elevation and in different habitats. History: discovered in 1898 by the widely travelled German botanist and prolific collector Rudolf Schlechter, at a site Schlechter called Insilva Maseku, near Massinga and north of Inhambane in central Mozambique, the collection formed the basis of Lapeirousia masukuensis, described by Vaupel & Schlechter in Psilosiphon masukuensis appears to be comparatively rare, particularly in Malawi, Mozambique and Zimbabwe where it has been gathered only a few times. An early collection made by French missionary, Henri Junod at Shilouvane in Limpopo Province, South Africa, bears the manuscript name L. junodii N.E.Br. Evidently Brown at one time considered the species distinct from what was then L. sandersonii, to which early collections were often referred. Representative specimens SOUTH AFRICA. LIMPOPO (Pafuri): Kruger National Park, Klopperfontein, ( CA), 21 Jan. 1953, Van der Schijff 1858 (PRE) (Tzaneen): Hans Merensky Nature Reserve, mopane veld in damp earth along sloot, ( DA), 11 Feb. 1971, Oates 371 (PRE); Hans Merensky Nature Reserve, waterlogged clay loam in vlei, 2000 ft [620 m], ( DA), 15 Mar. 1977, Zambatis 737 (PRE) (Phalaborwa): Mahlangeni, 4 km from Nsihana windpomp, ( AC), Wentzel 163 (PRE) (Pilgrims Rest): Shilouvane, ( AB), Jan. 1919, Junod 4139 (G, M, PRE); Shilouvane plain, L. junodii N.E.Br., ( AB), Junod 736 (G, K); 10 m [15 km], north of Abel Erasmus Pass, grass in bushveld, ( DA), 16 Dec. 1964, Mauve 4326 (K, PRE) (Acornhoek): Farm Grootdraai, stony flats, 1500 ft [457 m], ( AA), 15 Nov. 1973, Zambatis 543 (MO); Klaserie Private Nature Reserve, Farm Sark, seasonal seep in sandy clay, ( AC), 28 Jan. 1982, Zambatis 1345 (PRE); Kruger National Park, Pumbe, Satara, sandy ground, ( BB), Mar. 1967, Van Wyk 4783 (PRE). MPUMALANGA (Carolina): Eerstehoek, Songimvelo Nature Reserve, Doornhoek, south of Komati River, ( BB), 31 Jan. 1992, Stalmans 2618 (J, MO). [Exsiccatae from Malawi, Mozambique, and Zimbabwe are cited in Goldblatt (1990a).]

136 128 S T R E L I T Z I A 35 (2015) 9. Psilosiphon otaviensis (R.C.Foster) Goldblatt & J.C.Manning, comb. nov. Lapeirousia otaviensis R.C.Foster in Contributions from the Gray Herbarium 127: (1939); Goldblatt: 467 (1990a). Type: Namibia, Farm Auros near Otavi, 11 Feb. 1925, Dinter 5577 (GH, holo.!; B!, G!, PRE!, SAM!, Z (3 sheets)!, iso.). Plants mm high, repeatedly branched, stem with main axis ± straight and dominant, lateral braches diverging at 45 80, stem weakly compressed below, triangular in middle, ± rectangular above, angles lightly winged. Corm bell-shaped, mm diam.; tunics light brown, firm-leathery at first, becoming coarsely fibrous with age, sometimes ultimately finely fibrous. Leaves 4 7, ± linear( narrowly lanceolate), glaucous, main vein prominent, lower 2 4 basal and longest, 5 9 mm wide, exceeding inflorescence by mm, upper leaves decreasing in size above, becoming ± bract-like. Inflorescence a rounded to ± flat-topped false panicle, main terminal branches 3 5-flowered; bracts soft-textured, green, lanceolate, acute, often flushed red above, becoming dry from tips toward end of flowering, (7 )8 11 mm long, inner often slightly longer than outer. Flowers zygomorphic, white to cream, sometimes flushed pale lilac, lower three tepals with broad violet or purple median streak, with darker, paired purple to red lines near base, rarely upper 3 tepals with pale purple median line, unscented; perianth tube mm long, straight, gradually flared from base, ± 1.2 mm diam. below, ± 2.2 mm diam. at mouth; tepals unequal, dorsal slightly larger than others, lanceolate, ± acute, margins straight, mm, reflexed ± at right angles to tube. Stamens unilateral, ± erect; filaments 5 6 mm long, exserted for 2 4 mm, white; anthers 5 6 mm long, grey-purple; pollen light yellow-brown. Style nearly straight, adaxial to filaments, style usually dividing between apex of anthers to 3 mm beyond apex; branches ± 2 mm long, forked for 1 / 3 to 1 / 2 their length, diverging, but barely or not at all recurved. Capsules ± globose, 3-lobed, ± 7 mm long. Seeds not known. Chromosome number 2n = 10. Flowering time: February to April. Figure 47. Distribution and biology: with a relatively restricted geographic range, Psilosiphon otaviensis extends in a wide arc from the hills west and south of Etosha Pan in western Namibia to the Otavi Hills in the northeast and locally also in adjacent southern Angola (Figure 48). It is apparently confined to rocky sites, usually growing in granite but in the Otavi Hills, the type area, it occurs on shale outcrops and not, as far as we know, in dolomite, the most common rock type in the area. Nothing is known of its pollination biology, but the long-tubed, unscented flowers that open in mid-morning have stereotypical characteristics of those adapted for pollination by long-proboscid flies. Diagnosis and relationships: the relatively large flower with a perianth tube mm long, white to lilac perianth with lanceolate tepals and ultimate branches of the inflorescence with up to 5 flowers (rarely less than 2) distinguish Psilosiphon otaviensis. It has often been confused in herbaria with similarly long-tubed P. bainesii, which can be separated by a series of characters including flowers with a slightly shorter perianth tube, mm long, often flushed pink to redpurple outside, and white to pale pink tepals narrowed below into claws and with undulate margins. The inflorescence is divaricately branched and the ultimate branches have 1, rarely 2 flowers. The habitats of the two species also differ: P. otaviensis is evidently restricted to rocky sites, mostly granite or shale, whereas P. bainesii occurs in deep, hard sand in savanna or bushveld. Goldblatt (1990a) included blue-violet flowered plants from the Erongo Mtns in P. otaviensis, but we now consider these a separate species, P. erongoensis, distinguished in addition to perianth colour by a shorter perianth tube ± 37 mm long, and shorter, narrower tepals, ± 12 3 mm (vs mm). History: although first collected in 1879 in Damaraland by Ture Een, the traveller and botanical explorer, Psilosiphon otaviensis was only described in 1939 by the American, R.C. Foster, from a gathering made in 1925 by Dinter at the Farm Auros near Otavi. Dinter s collection of P. otaviensis was the first to be distributed widely to herbaria, and appears to have been recognised as a distinct species by accident, as Foster compared it to the very different, short-tubed P. avasmontanus (as Lapeirousia) in the protologue. The reason for this is obscure and may have been because he had misidentified material of the latter. Foster was unaware at the time of the

137 S T R E L I T Z I A 35 (2015) 129 A B FIGURE 47. Psilosiphon otaviensis, Goldblatt & Manning A, plant and corm; B, flower from front. Scale bar: 10 mm. Artist: John Manning.

138 130 S T R E L I T Z I A 35 (2015) similarity between L. otaviensis and L. bainesii, with which it was later confused. In the Prodromus Einer Flora von Südwestafrika, Sölch (1969) treated L. otaviensis and L. bainesii as conspecific and distinguished the two in the key by the several flowered ultimate inflorescence branches (a feature only of P. otaviensis). Sölch reserved the name L. vaupeliana for what is now P. bainesii (Goldblatt 1990a). Representative specimens NAMIBIA (Posto Velho): granite slope at Ombepera, rock crevices, corms edible, ( BD), 10 Apr. 1957, De Winter & Leistner 5481 (B, K, M, PRE, WIND) (Kamanjab): 8 km north of Kamanjab Nord, granite domes, ( DB), 18 Mar. 1974, Merxmüller & Geiss (K, M, PRE, WAG, WIND); Kamanjab, granite hills in rock crevices, ( DB), 2 Mar. 1957, De Winter & Leistner 5132 (B, K, M, PRE, WIND) (Tsumeb): Farm Auros, shale hills near farmhouse, ( DA), 21 Mar. 1988, Goldblatt & Manning 8837 (E, K, M, MO, NBG, PRE, S, WAG, WIND) (Otjihorongo): 90 km from Omaruru to Fransfontein, Table Mtn, by Ozondati, ( BB), 11 Apr. 1968, Wanntorp & Wanntorp 840 (K, S) (Otjiwarongo): Paresis Mtns, ( AD), Barnard 201 (SAM). ) (Trekkopje): Okongava, Kalkberg, granite slope, ( BB), 4 Feb. 1934, Dinter 6962 (B, BM, G, HBG, K, M, PRE, S, WIND, Z). Without precise locality: Damaraland, 1879, Een s.n. (BM). [Exsiccatae from Angola are cited by Goldblatt (1990a).] 10. Psilosiphon bainesii (Baker) Goldblatt & J.C.Manning, comb. nov. Lapeirousia bainesii Baker in Journal of Botany 14: 338 (1876); 352 (1898); Sölch: 8 (1969) [name misapplied to L. otaviensis]; Goldblatt: 469 (1990a). Type: Botswana, Kobe Pan [as inter Koobie et N Shaw valley ], Baines s.n. (K, lecto.!, effectively designated by N.E.Brown on specimen). Lapeirousia vaupeliana Dinter: 436 (1922). Sölch: 8 (1969). Type: Namibia, Gameros (cultivated at Okahandja), Dinter 3087 [B, lecto.!, designated by Sölch: 8 (1959)]. Plants mm high, usually repeatedly branched, stem compressed and 2 winged below, 3 4-angled and winged above, wings sometimes slightly crisped or finely serrated. Corm bell-shaped, mm diam.; tunics midto dark brown, ± leathery internally with veins sharply outlined, becoming ± fibrous, fibres coarse and wiry. Foliage leaves narrowly lanceolate to linear, main vein prominent and a pair of lateral veins on either side evident, 5 7 mm wide, usually slightly exceeding inflorescence. Inflorescence a rounded, divaricately branched false panicle, ultimate branches 1(2)-flowered, axes sharply 3-angled; bracts green below, becoming dry and membranous at anthesis, later dry and light brown, apices often darker brown, FIGURE 48. Distribution of Psilosiphon bainesii, ; P. otaviensis,, in southern Africa.

139 S T R E L I T Z I A 35 (2015) ( 12) mm long, inner slightly larger than outer. Flowers zygomorphic, ± white, often flushed or fading pale pink, tube pale pink to purple outside, lower 3 tepals each usually marked with red to brown streak in basal 1 / 2 and with dark mark at base, top of throat red on abaxial side, sweetly scented; perianth tube cylindric, slightly expanded in upper 5 mm, (25 )32 40 mm long, usually inclined; tepals subequal or dorsal slightly larger, ± spathulate, ± clawed (rarely with a tooth-like ridge in lower midline of lower tepals, margins undulate, dorsal ± upright to reflexed at right angles to tube, 9 12( 15) 3 4 mm, lower 3 tepals held close together, forming lip. Stamens unilateral, straight; filaments exserted mm, white; anthers mm long, pale purple; pollen ± white. Style arching over stamens, dividing between middle and apex of anthers or sometimes exceeding them, branches spreading, ± 2 mm long, ± undivided or notched apically. Capsules obovoid to globose, 5 6 mm long. Seeds globose, ± 2 mm diam. Chromosome number 2n = 10, 6. Flowering time: January to April, rarely late November or December. Figure 49. Distribution and biology: a species of the dry plains of central interior southern Africa, Psilosiphon bainesii extends from near Windhoek in central Namibia north into southern Angola and across the Kalahari to eastern Botswana and northwestern Limpopo in South Africa (Figure 48). It favours hard flat sandy ground where the corms may be lodged up to 300 mm below the surface. We have never seen it in rocky terrain, the habitat preferred by the related and morphologically similar P. otaviensis. The flowers open in the late afternoon and last through the night and into the following day, but rapidly become wilted in the dry and hot daylight conditions. The freshly opened flowers have a strong sweet and pleasant odour and produce a fair quantity of nectar and this combined with the pale perianth colour suggest adaptation to pollination by hawk moths. Like several other species of Psilosiphon in Namibia and elsewhere (see Ethnobotany pp. 20), the corms of P. bainesii are eaten raw or roasted by the indigenous population (Rodin 1985). Diagnosis and relationships: the divaricately branched inflorescence with branches bearing 1 or rarely 2 flowers, long perianth tube, and ± white perianth with pink to red markings near the base of the lower tepals distinguish Psilosiphon bainesii. The perianth tube is usually mm long, rarely up to 40 mm or less than 20 mm. The corm tunics are composed of light brown, densely compacted fibres, the outer layers of which become increasingly netted with age. These corm tunics, combined with the pale, long-tubed flower and the basic chromosome number of x = 5, suggest a close relationship with P. otaviensis and P. schimperi, which have similar karyotypes (Goldblatt 1990b). The similarity in the size and general shape of the flowers of P. bainesii and P. otaviensis led to their confusion in the past, but our field studies in Namibia have shown convincingly that these are separate species with different ranges and habitat preferences as well as different floral and inflorescence morphology. Populations of P. bainesii from west-central Namibia have a derived chromosome number, 2n = 6, most likely the result of Robertsonian fusion of chromosomes to yield large metacentric pairs (Goldblatt & Takei 1995). This number is, incidentally, the lowest recorded in the Iridaceae, coincidentally shared with some species of Crocus L. Specimens from eastern Botswana near Macheng, Hansen 3357, represent an unusual form of Psilosiphon bainesii: the flowers have a short tube, mm vs mm in most other collections of the species. In other respects the plants accord with P. bainesii, including the fairly distinctive translucent, pearly perianth colouring. The collection may represent a hybrid population, and then most likely with P. sandersonii, which also occurs in eastern Botswana. One of the few collections that we have seen from Angola, Barbosa & Moreno from Chibia, is unusual in having a wine red perianth tube. The reverse of the outer tepals is a similar dark colour, although the condition in the living plants may have been somewhat different. We cannot properly assess the significance of this variant until more material from Angola becomes available. History: apparently first collected in 1863 by the nineteenth century English landscape artist Thomas Baines and named in his honour, Psilosiphon bainesii was described by J.G. Baker in 1876, as Lapeirousia. The species was based on Baines s collection, but also on a second gathering from the Transvaal made by a certain Mr. Todd, about whom nothing seems to be

140 132 S T R E L I T Z I A 35 (2015) FIGURE 49. Psilosiphon bainesii, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

141 S T R E L I T Z I A 35 (2015) 133 known. Baines s plants were collected in western Botswana near Kobe Pan, an area still not well-known botanically. Plants from west central Namibia were described in 1922 as L. vaupeliana by Dinter, based on three of his own collections. The distinction that Dinter made between this and the avowedly related L. bainesii was the redviolet flower with darker veins in his species compared to the white flowers of L. bainesii with dark brown markings. These differences did not seem significant to Goldblatt (1990a) and we endorse his decision to include L. vaupeliana in P. bainesii. Psilosiphon otaviensis was regarded as a synonym of Lapeirousia bainesii by Sölch (1969) in his account of Iridaceae for Prodromus Einer Flora von Südwestafrika. However, he upheld L. vaupeliana, distinguishing it in the diagnostic key by the ultimate branches of the inflorescences each bearing a single flower in contrast to P. bainesii in which the ultimate branches were said to have two or more sessile flowers. The type specimen of P. bainesii has, however, single-flowered ultimate inflorescence branches and we have no doubt it is conspecific with the later L. vaupeliana whereas P. otaviensis has 3 5 flowers on the ultimate branches of the inflorescence and also has fairly different flowers. Representative specimens BOTSWANA (Aha Hills): Aha hills, 110 km west of Nokaneng, ( CB), Wild & Drummond 6919 (BM); Dobe, 26 km north of Aha Hills, SWA [Namibia] border, ( CA), Wild & Drummond 7202 (K, PRE) (Koanaka Hills): near SWA [Namibia] border fence, sandy grassland, (?), Smith 3330 (MO) (Kwebe Hills): Ngamiland, Kwebe, Lugard & Lugard 179 (GRA, K, SAM) (Mompse): Odiakwe, savanna, west of Francistown just north of Mkarikari Pan, ( AB), Wild & Drummond 6826 (K) (Lothlekane): Ngamiland, near Bachakuru, white and chocolate, ( DD), Lugard 242 (K) (Serowe): Kweneng, Serowe, roadside, sandy grass veld, ( BC), 2 Mar. 1991, Barnard 639 (PRE) (Mahalapye): Mahalapye Exp. Morale, shallow gritty sandy loam, ( BB), Yalala 356 (K, LISC, PRE) (Ellisras): 10 miles west of Macheng towards Mahalapye, woodland, ( AB), Hansen 3357 (C, K, PRE, WAG) (Gaborone): 12 km south of Gaborone, Mmokolodi, ( DD), 24 Mar. 1994, Cole 546 (PRE) (Mochudi): Mochudi, ( AC), Jan. 1915, Harbor sub Rogers 6569 or Harbor s.n. (G, K, PRE, SAM). NAMIBIA (Ondangua): 100 km east Oshikango, corm eaten raw or roasted, ( BD), Rodin 9295 (K, M, MO, PRE, WIND) (Rundu): 14.6 m [22 km] east of Runtu on Sambiu road, ( DD), De Winter & Marais 4558 (K, M, WIND) (Karakuwise): Cigarette, northeast of Karakuwise, ( DC), Maguire 2381 (NBG, PRE) (Gobaub): north of Outjo, between Farms Grensplaas and Tsabis, stony sandy hill slope, ( AD), 12 Mar. 1997, Germishuizen 9692 (PRE); Farm Norabis 387, west of Otavi, thornveld on red sand, ( DD), Goldblatt & Manning 8826 (MO, PRE, WIND) (Tsumeb): Otavi, ( CB), Dinter 5755 (B, G, PRE, Z); Grootfontein, Otjirukaku, ( DB), Seydel 2068 (B, BR, C, GH, K, M, MO, S, WAG, WIND) (Grootfontein): Farm Oliewenhof, sehr häufig auf der Palmflache, ( CB), Merxmuller & Giess (K, M, WIND, PRE, S, SRGH, WAG) (Tsumkwe): 6 km east of Tsumkwe on the road to Botswana, white sand, ( DA), Giess et al A (WIND); 2 km west of Kremetartkop, open grassland, ( DD), 26 Feb. 1986, Hines 587 (PRE) (Okahandja): Farm Omatako View, ( BA), Woortman 152 (M, WIND) (Waterberg): Quickborn, sand, ( AA), Bradfield 196 (K, PRE) (Otjosondu): 48 km along Kapps Farm road from Steinhausen, compact sand, ( DC), Goldblatt & Manning 8808 (E, K, M, MO, NBG, PRE, S, WAG, WIND) (Steinhausen): Farm Mex, 45 km north of Witvlei, red sand, ( AB), Germishuizen 2662 (PRE, WIND) (Tses): Kalaharirand, Tutara, Farm Okamatangara, ( DB), Seydel 2549 (BR, K, M, MO, SRGH, WAG). SOUTH AFRICA. LIMPOPO (Waterpoort): 10 miles [15 km] southeast of Dongola Reserve, Farm Coila, ( AD), Codd & Dyer 3780 (E, K, PRE); Dongola, Soutpansberg, ( BC), Pole Evans 4305 (K, MO, PRE, SRGH) (Ellisras): Potgietersrus, 2 miles northeast of Tomburke, red gritty flats, ( BB), Codd 6614 (K, PRE, SRGH) (Baltimore): Farm Dansfontein, red, sandy loam, ( AB), 31 Jan. 1996, Du Toit 4965 (PRE). NORTH WEST (Mafikeng): Mafikeng, Farm Panvlakte 3, thornveld, sandy soil, ( DC), 22 Jan. 1985, Knollys 3 (GRA). [Also in southern Angola (Goldblatt 1990a).] 11. Psilosiphon schimperi (Aschers. & Klatt) Goldblatt & J.C.Manning, comb. nov. Tritonia schimperi Aschers. & Klatt in Klatt in Linnaea 34: 697 (1866b). Acidanthera unicolor Hochst. ex Baker: 160 (1877), 359 (1898), nom. superfl. pro Tritonia schimperi Aschers. & Klatt. Lapeirousia schimperi (Aschers. & Klatt) Milne-Redh.: 307 (1934); Goldblatt: 472 (1990a). Acidanthera schimperi (Aschers. & Klatt) R.C.Foster: 42 (1936). Type: Ethiopia. Tigray: woods and thickets near Goelleb on the river Tacazze, 4000 ft [1 220 m], Schimper 2304 (B lecto.! designated by Goldblatt: 472 (1990a); B!, G!, K!, P!, isolecto.).

142 134 S T R E L I T Z I A 35 (2015) Anomatheca angolensis Baker: 337 (1876). Lapeirousia monteiroi Baker: 355 (1898), illegitimate superfluous name for A. angolensis Baker (1876). Freesia angolensis (Baker) Klatt: 187 (1894). Lapeirousia angolensis (Baker) R.C.Foster: 48 (1936). Type: Angola, without precise locality, cult. Hort. Kew, Monteiro s.n. (K, holo.!). Lapeirousia fragrans Welw. ex Baker: 272 (1878). Type: Angola. Huila: ad Morro de Lopollo, 5200 [1 585 m], stony and rocky places, Dec. 1859, Welwitsch 1552 (K, holo.!; BM!, C!, COI!, G!, P!, iso.). Lapeirousia cyanescens Welw. ex Baker: 272 (1878). Type: Angola. Huila: hills near Humpata, Apr. 1860, Welwitsch 1553 (K, lecto.! designated by Goldblatt: 472 (1990a); B!, BM!, COI!, G!, iso.). Lapeirousia edulis Schinz, Bull. Herb. Boissier 4, Appendix 3: 49 (1896). Type: Namibia, Amboland, west of Olukonda, 5 Apr. 1893, Rautanen 106 (Z, lecto.!, designated by Goldblatt: 472 (1990a); K!, P (2 sheets)!, Z (3 sheets)!, isolecto.). Lapeirousia porphyrosiphon Baker: 353 (1898). Type: Botswana, Ngamiland, Kalahari Desert near Mamunwe, 26 Feb. 1897, Lugard 338 (K, holo.!). Lapeirousia erythreae Chiovenda: 139 (1911). Type: Eritrea, Accheli Guzai (Ocule Cusai) near Loggo Sarda, Deggahen, m, 15 Sept. 1902, Pappi 1414 (G, lecto.!, designated by Goldblatt: 472 (1990a); GH!, MO!, isolecto.). Lapeirousia montaboniana Chiovenda: (1911). Type: Eritrea, Bogos near Cheren, 24 Aug. 1906, Pappi 7087 (? FI, not seen). Lapeirousia dinteri Vaupel: (1912). Type: Namibia, Damaraland, Aukos Kreyfontein, 28 Dec. 1908, Dinter 810 (B, holo.!; B!, SAM (2 sheets)!, iso.). Lapeirousia uliginosa Dinter: 13, 14 (1912). Type: unknown, none cited. Plants (200 ) mm high, usually laxly few(several)-branched, stem rounded below to nearly square and 4-angled to 4-winged above. Corm bell-shaped, mm diam.; tunics of compacted fibres, light to dark brown, outer layers becoming loosely fibrous and netted. Foliage leaves 3 or more, lower 2 largest, usually slightly exceeding inflorescence, decreasing in size above, narrowly lanceolate to ± linear, 5 10( 15) mm wide, main vein slightly raised. Inflorescence a lax false panicle or branched spike, ultimate branches 1 3-flowered; bracts (10 )20 35( 45) mm long, green, becoming membranous to ± dry and papery above, then light to dark brown, tips dark brown. Flowers zygomorphic, white to cream, rarely pale violet, when whitish sometimes fading lilac on drying especially on tube; opening at sunset and then sometimes scented; perianth tube cylindric, slender, ( 150) mm long; tepals lanceolate, extended ± at right angles to tube, 18 22( 25) 6 8( 11) mm. Stamens unilateral, ± erect; filaments exserted 5 7 mm from tube; anthers 6 7 mm long, cream or purple; pollen cream or blue. Style filiform, arching over filaments, dividing opposite anther apices or to up to 3 mm beyond anthers; branches ± 2 mm long, forked for ± 1 / 3 their length. Capsules obovoidoblong, 8 12 mm long, partly enclosed in bracts. Seeds globose to slightly angular, mm in diam. Chromosome number 2n = 10. Flowering time: mid- to late summer, December to March, in central and south tropical Africa [north of the equator mostly in September and October but in Kenya and Ethiopia also April to June]. Figure 50. Distribution and biology: with a remarkably wide distribution across Africa, Psilosiphon schimperi extends from southwestern Angola and north-central Namibia across northern Botswana, southern Zambia and Zimbabwe in an almost continuous belt (Figure 51). Further north its distribution is scattered, and it occurs in northern Tanzania, eastern Kenya, southern Ethiopia and then again in northern Ethiopia, Eritrea, Somalia and the Jebel Marra and Jebel Gurgeil highlands of northwest Sudan. It is almost certainly absent from the Democratic Republic of the Congo, Uganda, central and western Kenya, and southern Tanzania. A plant from Yemen in the southern Arabian Peninsula is cited in the protologue, but we have not been able to confirm its presence there. P. schimperi grows in locally moist sites in otherwise largely arid country, typically in washes, the omurambas [ancient river beds] of Namibia, dambo margins, stream banks and seasonal marshes as well as in damp grassland. The flowers open in late afternoon and last through the night, but gradually wilt the next day. Some collections note a sweet odour when the flowers are open but live plants from Namibia that we have examined were scentless during the day and early evening hours, indicating variation for scent. The corms are edible, and eaten either raw or roasted by the Ovambo of northern Namibia (Rodin 1985).

143 S T R E L I T Z I A 35 (2015) 135 FIGURE 50. Psilosiphon schimperi, Goldblatt & Manning Scale bar: 10 mm. Artist: John Manning.

144 136 S T R E L I T Z I A 35 (2015) Diagnosis and relationships: the large white flower with a perianth tube mm long and the laxly branched inflorescence readily distinguish Psilosiphon schimperi from other species in the genus. Its immediate relationships probably lie with the Namibian and Angolan longtubed species, P. bainesii and P. otaviensis, with which it shares a similar karyotype and diploid chromosome number of 2n = 10 (Goldblatt 1990b). These two species, centred in northern Namibia, also have pale flowers with relatively long perianth tubes, up to 45 mm long, but more compact inflorescences and they are not likely to be confused with P. schimperi. Unusually long-tubed white flowers characterise another tropical African species, Lapeirousia odoratissima, but this species has the corrugate leaves and woody corm tunics that place it firmly in Lapeirousia rather than Psilosiphon. The floral similarity is evidently due to convergence, presumably for pollination by hawk moths. P. schimperi exhibits relatively little variation for a species with such a wide distribution. Some specimens from Angola and Namibia in the southwest of its range have unusually short bracts, mm long (e.g., Giess & Loutit 14123), but at least some plants from Namibia have bracts up to 35 mm long, thus comparable with those in plants from tropical and northeast Africa in which the bracts range from mm. Although lanceolate-elliptic tepals mm are the norm, plants collected in southern Angola (Harrower et al. s.n. NBG) have unusually large, ovate tepals up to mm. Other features are consistent with P. schimperi. As noted by Goldblatt (1990a), some collections of P. schimperi from Ethiopia and Sudan (e.g., Schimper 2304, 431) also have particularly short bracts, mm long, and either pale or fairly dark brown. These plants have a perianth tube only mm long and may be depauperate owing to a poor growing season or they may represent an ecotype or race constant for these features. History: although not particularly variable, Psilosiphon schimperi has a long and confused history, largely due to its wide distribution rather than to a conviction that any of the later synonyms represented separate species. The type locality is in northern Ethiopia where collections were made in the 1840s by the French botanists Quartin Dillon and Petit and later by Georg Schimper in It was named in Schimper s honour by Ascherson & Klatt (in Klatt 1866b), who referred it to the southern African genus Tritonia. Three separate collections from southwestern Angola made at this period by Joachim Monteiro and Friedrich Welwitsch were described by J.G. Baker as Anomatheca angolensis (1876), Lapeirousia cyanescens FIGURE 51. Distribution of Psilosiphon schimperi in southern Africa,.

145 S T R E L I T Z I A 35 (2015) 137 (1877), and L. fragrans (1878) respectively, without reference to the Ethiopian species. Later collections from Namibia were named independently by Hans Schinz as L. edulis in 1896 and Friedrich Vaupel as L. dinteri in Plants from Botswana, collected by E.J. Lugard were assigned to L. porphyrosiphon (Baker 1898) while in 1911 A. Chiovenda described L. erythreae and L. montaboniana based on contemporary collections from Eritrea. Tritonia schimperi was transferred to Lapeirousia in 1934, replacing the name L. porphyrosiphon, until then used for the species in south tropical Africa. Representative specimens BOTSWANA (Livingstone): south side of Chobe River, above Kasane, (?), May 1972, Sheppe 167 (SRGH); Chobe National Park, Savuti, Gogha Hills, ( CC), 13 Mar. 1994, Roodt 110 (PRE) (Andara): southwest end of Tsodilo Hills, open savanna, ( DD), 11 Mar. 1985, Long (E) (Maun): Khwai/Maxwe road, Moremi Wildlife Reserve, ( BC), 16 Mar. 1977, Smith 1936 (BR, K, SRGH) (Rakops): Rakops, Makuba Dukuwe Fence, marsh, ( AB), 2 March 1991, Barnard 661 (PRE). NAMIBIA (Ondangua): 100 km east of Oshikango, ( BD), 19 Apr. 1973, Rodin 9295A (M, MO, PRE, WIND) (Kuring Kuru): Katui-Tui, maize land in sandy loam, ( AD), 15 May 1965 (fr.), Barnard 191 (WIND); Kuring Kuru, Okavango River bank in forest, 28 Mar. 1966, ( DA), Soni s.n. (WIND 3938) (Runtu): swampy marshes below Runtu, 31 Jan. 1956, ( AD), De Winter & Marais 4469 (K, M, PRE, WIND, Z) (Katima Mulilo): Katima Mulilo, vlei, ( AD), 30 Jan 1975, Vahrmeijer & Du Preez 2496 (MO, PRE) (Namutoni): Etosha National Park, Bigales Huh, ( C), 15 Feb. 1974, Le Roux 644 (PRE, WIND) (Karakuwisa): grass flats, Karakuwisa, 4 Mar. 1958, ( DC), Merxmüller & Giess 1795 (BM, K, M, PRE, WIND) (Andara): Kaprivi side of the river at Andara Mission, crevices on rocky outcrops, 23 Feb. 1956, ( AB), De Winter & Marais 4820 (K, PRE, WIND); Okavango River, 19 km north of Shakawe on Botswana border, ( BA), 16 Mar. 1965, Wild & Drummond 7093 (K, LISC, M, PRE, SRGH) (Okakuejo): Etosha Pan, black peat soil on the road to Ombika, ( BB), 4 Apr. 1980, Giess (M, MO, PRE, WIND); Etosha National Park, 2.5 km east of Okakuejo Ombika road, ( BD), 5 Mar. 1976, Giess & Loutit (K, M, MO, PRE, WIND); grey-black peat flats north of Ombika, ( BD), Le Roux 321 (PRE, WIND) (Gobaub): Etosha Game Park, near Homob water hole, ( AA), 11 Feb. 1966, Tinley 1285 (M, PRE, WIND); Farm Neidaus North 78, heavy black clay with limestone, seasonally waterlogged, ( DC), 19 Mar. 1988, Goldblatt & Manning 8831 (MO, WIND) (Tsumeb): Farm Malta, ( AB), 5 Feb. 1971, Giess (M, PRE, WIND); Farm Kumkauas, large colonies among tough grasses, ( CA), 30 Jan. 1971, Giess (M, WAG, WIND), 9 Mar. 1974, Merxmüller & Giess (K, M, S, PRE, SRGH, WAG, WIND). New combinations for Psilosiphon species not represented in southern Africa Psilosiphon abyssinicus (R.Br. ex A.Rich.) Goldblatt & J.C.Manning, comb. nov. Lapeirousia abyssinica (R.Br. ex A.Rich.) Baker in Journal of the Linnean Society, Botany 16: 155 (1877). Geissorhiza abyssinica R.Br. ex A.Rich.: 308 (1850) [Geissorhiza abyssinica R.Br. in Salt: App. 1 (1814), nom. nud.]. Type. Ethiopia, Maigoigoi ad Dobre Sina, Quartin Dillon & Petit s.n. (P, lecto.! designated by Goldblatt: 459 (1990a); P!; BR!, isolecto.). Montbretia abyssinica Hochst. ex A. Rich.: 308 (1850) [Montbretia abyssinica Hochst., Flora 24(1 Intell.): 21 (1841), nom. nud.]. Tritonia abyssinica (Hochst.) Walp.: 613 (1852). Type: Ethiopia, hills near Adoa, October (fl. & fr.), Schimper 329 (P, lecto.!, designated by Goldblatt: 459 (1990a); BM!, BR!, G!, K!, M!, P!, isolecto.). Psilosiphon setifolius (Harms) Goldblatt & J.C.Manning, comb. nov. Lapeirousia setifolia Harms in Botanische Jahrbucher für Systematik 30: 278 (1901). Lapeirousia erythrantha var. setifolia (Harms) Geerinck et al.: 337 (1972). Type: Tanzania, Eastern Livingstone Mtns, Ubena, Tsausingewe district, m, March 1899, Goetze 812 (B, holo.!; BR!, E!, iso.). Psilosiphon teretifolius (Geerinck et al.) Goldblatt & J.C.Manning, comb. nov. Lapeirousia teretifolia (Geerinck et al.) Goldblatt in Annals of the Missouri Botanical Garden 77: 457 (1990a). Lapeirousia erythrantha var. teretifolia Geerinck, Lisowski, Malaisse & Symoens: 342 (1972). Type: Zaire [Democratic Republic of the Congo], Shaba, Plateau de la Manika, env. 2 km west de Katema, 20 Jan. 1969, Lisowski, Malaisse & Symoens 182 (LSHI, holo., BR!, BRVU, K, POZG, iso. [the lectotypification by Goldblatt: 457 (1990a) was made in error]. Psilosiphon zambeziacus (Goldblatt) Goldblatt & J.C.Manning, comb. nov. Lapei

146 138 S T R E L I T Z I A 35 (2015) rousia zambeziaca Goldblatt in South African Journal of Botany 57: 226 (1991), nom. nov. pro L. angolensis Goldblatt: 458 (1990a), illegitimate homonym. Type: Angola, Moxico, a few miles west of River Kaperu (Kapelu), boggy grassland, 10 Jan. 1938, Milne-Redhead 4037 (K, holo.!; BM!, BR!, LISC!, P!, PRE!, iso.). Excluded species Lapeirousia welwitschii Baker, Handbook of the Irideae 168 (1892). Type: Angola: Malange, Pungo Andongo, Jan. 1857, Welwitsch 1531 (BM, annotated as lectotype in unknown hand; C, COI, G, K, P, isolecto.). L. erythrantha var. welwitschii (Baker) Marais ex Geerinck, Lisowski, Malaisse & Symoens: 340 (1972), applied to L. erythrantha as L. briartii. [The type specimens are poorly preserved, especially the flowers, and are most likely either Psilosiphon erythranthus or P. rivularis. The type locality must be visited to resolve this question. Neither species has otherwise been recorded at Pungo Andongo in north-central Angola and P. rivularis is only known from Angola only in the south of the country.]

147 S T R E L I T Z I A 35 (2015) 139 References ANDERSON, B., COLE, W.W. & BARRETT, S.C.H Specialized bird perch aids cross-pollination. Nature 435: ANDREWS, H.C Ixia crispifolia. The botanist s repository 1: t. 35. BAKER, J.G New Gladioleae. Journal of Botany 14: BAKER, J.G [as 1878]. Systema Iridearum. Journal of the Linnean Society, Botany 16: BAKER, J.G Report on the Liliaceae, Iridaceae, Hypoxidaceae, and Haemodoraceae of Welwitsch s Angolan Herbarium. Transactions of the Linnean Society, London, Botany, ser. 2, 1: BAKER, J.G Handbook of the Irideae. George Bell & Sons, London. BAKER, J.G Irideae. In W.T. Thiselton-Dyer (ed.) Flora capensis 6: Reeve & Co., London. BAKER, J.G Irideae. In W.T. Thiselton-Dyer (ed.) Flora of tropical Africa 7: Reeve & Co., London. BAKER, J. G Diagnoses Africanae; XIV. Bulletin of Miscellaneous Information, Royal Botanic Gardens, Kew 1906: BANDEIRA, S,. BOLNICK, D. & BARBOSA, F Wild flowers of southern Mozambique. Universidade Eduardo Mondlane, Maputo. BROWN, N.E Diagnoses Africanae; XVII. Bulletin of Miscellaneous Information, Royal Botanic Gardens, Kew 1906: BROWN, N.E The South African Iridaceae of Thunberg s herbarium. Journal of the Linnean Society, Botany 48: CHIOVENDA, E Plantae novae vel minus notae e regione aethiopica. Annali di Bontanica 9: DELAROCHE, D Descriptiones plantarum aliquot novarum. J. & H. Verbeek, Leiden. DEMISSEW, S., NORDAL, I. & STABBETORP, O.E Flowers of Ethiopia and Eritrea: Aloes and other Lilies. Shama Books, Addis Ababa. DE WAAL, C., ANDERSON, B. & BARRETT, S.C.H The natural history of pollination and mating in bird-pollinated Babiana (Iridaceae). Annals of Botany 109: DIETRICH, A.G Species plantarum. Nauck, Berlin. DINTER, K Die vegetabilische Veldkost Deutsch-Südwest-Afrikas. K. Dinter, Okahandja. DINTER, K Index der aus Deutsch-SudwesAfrika bis zum Jahre 1917 bekannt bewardenen. Pflanzenarten. XII. Repertorium specierum novarum regni vegetabilis 18: DINTER, K Diagnosen neuer südwestafrikanischer Pflanzen. Repertorium specierum novarum regni vegetabilis 29: ECKLON, C.F Topographisches Verzeichniss der Pflanzensammlung von C. F. Ecklon. Esslingen. ENGLER, A. & PRANTL, K Die natürlichen Pflanzenfamilien ed. 2, 15a. W. Engelman, Leipzig, Germany. FOREST, F., GOLDBLATT, P., MANNING, J.C., BAKER, D., COLVILLE, J., DEVEY, D.S., JOSE, S., KAYE, M. & BUERKI, S Pollinator shifts as triggers of speciation in painted petal irises (Lapeirousia: Iridaceae). Annals of Botany (London) 113: FOSTER, R.C Notes on nomenclature in Iridaceae. Contributions from the Gray Herbarium of Harvard University 114: FOSTER, R.C Studies in the Iridaceae 1. Contributions from the Gray Herbarium of Harvard University 127: FOX, F.W. & YOUNG, M.E.N Food from the Veld. Delta, Johannesburg. GEERINCK, D., LISOWSKI, S., MALAISSE, F. & SYMOENS, J.J Le genre Lapeirousia Pourr. (Iridaceae) au Zaïre. Bulletin de la Societé Royale de Botanique de Belgique 105: GMELIN, J.F Systema Naturae, vol. 2. Emanuel Beer, Leipzig. GOLDBLATT, P Cytological and morphological studies in the southern African Iridaceae. Journal of South African Botany 37: GOLDBLATT, P A revision of the genera Lapeirousia Pourret and Anomatheca Ker in the winter rainfall region of South Africa. Contributions from the Bolus Herbarium 4: GOLDBLATT, P Lapeirousia oreogena Schltr. ex Goldbl. Flowering Plants of Africa 42: tab GOLDBLATT, P Revision of the tropical African genus Zygotritonia (Iridaceae). Bulletin du Museum d Histoire Naturelle, 4 sér., sect. B, Adansonia 11: GOLDBLATT, P. 1990a. Systematics of Lapeirousia (Iridaceae Ixioideae) in tropical Africa. Annals of the Missouri Botanical Garden 77: GOLDBLATT, P. 1990b. Cytological variability in the African genus Lapeirousia (Iridaceae: Ixioideae). Annals of the Missouri Botanical Garden 77:

148 140 S T R E L I T Z I A 35 (2015) GOLDBLATT, P Nomenclatural notes on African Iridaceae. South African Journal of Botany 57: 226. GOLDBLATT, P. & BARNARD, T.T The Iridaceae of Daniel de la Roche. Journal of South African Botany 36: GOLDBLATT, P. & MANNING, J.C Leaf and corm structure in Lapeirousia (Iridaceae Ixioideae) in relation to phylogeny and infrageneric classification. Annals of the Missouri Botanical Garden 77: GOLDBLATT, P. & MANNING, J.C Systematics of the southern African Lapeirousia corymbosa (Iridaceae Ixioideae) complex (sect. Fastigiata) and a new species of sect. Paniculata. South African Journal of Botany 58: GOLDBLATT, P. & MANNING, J.C New taxa and revisions to the taxonomy of southern African Lapeirousia subgenus Lapeirousia (Iridaceae subfamily Ixioideae). Novon 4: GOLDBLATT, P. & MANNING, J.C Phylogeny and speciation in Lapeirousia subgenus Lapeirousia (Iridaceae subfamily Ixioideae). Annals of the Missouri Botanical Garden 83: GOLDBLATT, P. & MANNING, J.C The long-proboscid fly pollination system in southern Africa. Annals of the Missouri Botanical Garden 87: GOLDBLATT, P. & MANNING, J.C Evidence for moth and butterfly pollination in Gladiolus (Iridaceae: Crocoideae). Annals of the Missouri Botanical Garden 89: GOLDBLATT, P. & MANNING, J.C Radiation of pollination systems in the Iridaceae of sub-saharan Africa. Annals of Botany 97: GOLDBLATT, P. & MANNING, J.C Floral biology of Babiana (Iridaceae: Crocoideae): adaptive floral radiation and pollination. Annals of the Missouri Botanical Garden 94: GOLDBLATT, P. & MANNING, J.C The Iris family: natural history and classification. Timber Press, Portland. GOLDBLATT, P. & MANNING, J.C New species of Geissorhiza (Iridaceae: Crocoideae) from the southern African winter-rainfall zone, nomenclatural changes, range extensions and notes on pollen morphology and floral ecology. Bothalia 39: GOLDBLATT, P. & MANNING, J.C Systematics and biology of the Cape genus Sparaxis (Iridaceae). Strelitzia 32. GOLDBLATT, P. & MARAIS, W Savannosiphon gen. nov., a segregate of Lapeirousia (Iridaceae Ixioideae). Annals of the Missouri Botanical Garden 66: GOLDBLATT, P. & TAKEI, M Chromosome cytology of the African genus Lapeirousia (Iridaceae: Ixioideae). Annals of the Missouri Botanical Garden 80: GOLDBLATT, P., BARI, A., & MANNING, J.C Sulcus and operculum structure in the pollen grains of Iridaceae subfamily Ixioideae. Annals of the Missouri Botanical Garden 78: GOLDBLATT, P., MANNING, J.C. & BERNHARDT, P Pollination biology of Lapeirousia subgenus Lapeirousia (Iridaceae) in southern Africa: floral divergence and adaptation for long-tongued fly pollination. Annals of the Missouri Botanical Garden 82: GOLDBLATT, P., MANNING, J.C. & BERNHARDT, P Adaptive radiation of pollination mechanisms in Sparaxis (Iridaceae: Ixioideae). Adansonia sér. 3, 22: GOLDBLATT, P., MANNING, J., DAVIES, T.J., SAVOLAIN- EN, S. & REZAI, S. 2004a. Cyanixia, a new genus for the Socotran endemic Babiana socotrana (Iridaceae Crocoideae). Edinburgh Journal of Botany 60: GOLDBLATT, P., NÄNNI, I., BERNHARDT, P. & MAN- NING, J.C. 2004b. Floral biology of Hesperantha (Iridaceae: Crocoideae): how minor shifts in floral presentation change the pollination system. Annals of the Missouri Botanical Garden 91(1): GOLDBLATT, P., DAVIES, T.J., MANNING, J.C., VAN DER BANK, M. & SAVOLAINEN, V Phylogeny of Iridaceae subfamily Crocoideae based on combined multigene plastid DNA analysis. Aliso 22: GOLDBLATT, P., MANNING, J.C. & DEMISSEW, S Two new species of Zygotritonia Mildbr. (Iridaceae: Crocoideae) from eastern tropical Africa. South African journal of Botany 96: GUNN, M. & CODD, L.E Botanical Exploration of Southern Africa. Balkema, Cape Town. HARMS, H In Engler, A., Beiträge zur Flora von Afrika. XXII. Botanische Jahrbucher für Systematik 30: HOCHSTETTER, C.F Ankündigung verkäuflicher Pflanzen des Würtembergischen Reisevereins. Flora 24. Intelligenzblatt 1(2): JACQUIN, N.J. VON Ixia corymbosa. Icones Plantarum Rariorum. Wappler, Vienna. JACQUIN, N.J. VON Collecteana botanica, 4. Wappler, Vienna. JACQUIN, N.J. VON Gladiolus silenoides. Icones Plantarum Rariorum. Wappler, Vienna. JACQUIN, N.J. VON Galaxia plicata. Icones Plantarum Rariorum. Wappler, Vienna. KER GAWLER, J Lapeyrousia corymbosa. Leveltopped Lapeirousia. Curtis s Botanical Magazine 16: t KER GAWLER, J.B Ensatorum ordo. König & Sims Annals of Botany 1: KER GAWLER, J.B Lapeyrousia fissifolia. Leafyspiked Lapeyrousia. Curtis s Botanical Magazine 31: sub t KLATT, F.W Revisio Iridearum. Linnaea 32: KLATT, F.W. 1866a. Revisio Iridearum (Conclusio). Linnaea 34: KLATT, F.W. 1866b. Revisio Iridearum (Addenda, Emendata et Corrigenda). Linnaea 34:

149 S T R E L I T Z I A 35 (2015) 141 KLATT, F.W Ergänzungen und Berichtigungen zu Baker s Systema Iridacearum. Abhandlungen der Naturforschenden Gesellschaft zu Halle 15: KLATT, F.W Determination and description of Cape Irideae. Transactions of the South African Philosophical Society 3: KLATT, F.W Iridaceae. In Th. Durand & H. Schinz (editors), Conspectus Florae Africae 5: LAMARCK, J.P.B. DE Encyclopédie méthodique. Botanique 2. Panckoucke, Paris. LAMARCK, J.P.B. DE Encyclopédie méthodique. Botanique 3. Panckoucke, Paris. LAWRENCE, G.H.M Notes on Lapeirousia in cultivation. Baileya 3: LEFFERS, A Gemsbok bean & Kalahari truffle. Mac- Millan Education Namibia, Windhoek. LEWIS, G.J Iridaceae. Iridaceae. Pp in R.S. Adamson & T.M. Salter (eds), Flora of the Cape Peninsula. Juta, Cape Town. LEWIS, G.J Plantae novae Africanae. Annals of the South African Museum 40(1): 1 9. LINNAEUS, C Species Plantarum. Salvius, Stockholm. LINNAEUS, C Centuria Plantarum 2. Salvius, Stockholm. LINNAEUS, C. (fil.) [as 1781]. Supplementum plantarum. Orphanotropheus, Brunswick. MANNING, J.C. & GOLDBLATT, P The Prosoeca peringueyi (Diptera: Nemestrinidae) pollination syndrome in southern Africa: long-tongued flies and their tubular flowers. Annals of the Missouri Botanical Garden 83: MANNING, J.C. & GOLDBLATT, P The Moegistorhynchus longirostris (Diptera: Nemestrinidae) pollination guild: long-tubed flowers and a specialized long-proboscid fly pollination system in southern Africa. Plant Systematics and Evolution 206: MANNING, J.C. & GOLDBLATT, P Radiation of pollination systems in the Cape genus Tritoniopsis (Iridaceae: Crocoideae) and the development of bimodal pollination strategies. International Journal of Plant Science 166: MANNING, J.C. & GOLDBLATT, P. & MANNHEIMER, C Here today, gone tomorrow: Gladiolus halophilus (Iridaceae: Crocoideae), a remarkable new species from southern Namibia facing imminent extinction. South African Journal of Botany 77: MARLOTH, R Flora of South Africa 4: plate 43. Juta, Cape Town. MARSHALL, L The!Kung of Nyae Nyae. Harvard University Press, Cambridge. MEYER, E Zwei pflanzengeographische Documente von J.F. Drège. Flora 26 (Beigabe): MILNE-REDHEAD, E.W.B.H Tropical African plants. XII. Bulletin of Miscellaneous Information. Royal Botanic Gardens, Kew [1934]: NORDENSTAM, B Notes on South African Iridaceae: Lapeirousia and Babiana. Botaniska Notiser 123: PAUW, A., STOFBERG, J. & WATERMAN, R.J Flies and flowers in Darwin s race. Evolution 63: PERSOON, H Synopsis Plantarum, Vol. 1. C.H. Cramer, Paris. PETERS, W.C.H Naturwissenschaftlike Reise nach Mossambique. Botanik. 6 (2). Berlin. PLUKENET, L Almagestum Botanicum (Phytographiae Pt. 4) 87 & fig. 275, f. 1. London. POIRET, J.L.M In G.-F. Cuvier, Dictionnaire des sciences naturelles 39: Levrault, Strassbourg. POURRET, P.A Description de deux genres de la famille des Liliacées, désignés sous le nom de Lomenia et de Lapeirousia. Histoire et Mémoires de l Académie Royale des Sciences, Toulouse 3: RENDLE, A.B A contribution to the flora of eastern tropical Africa. Journal of the Linnean Society, Botany 30: RICHARD, A Tentamen Florae Abyssinicae, 2. Bertrand, Paris. RODIN, R The ethnobotany of the Kwanyama Ovambos. Monographs in Systematic Botany from the Missouri Botanical Garden 9. ROEMER, J.J. & SCHULTES, J.A Systema vegetabilium, vol. 1. J.G. Cotta, Stuttgart. ROESSLER, H Iridaceae. In H. Roessler & H. Merxmüller, Neue Pteridophyten un Liliifloren aus Sudwestafrica. Mitteilungen der botanische Staatssammlung, München 15: RUDALL, P.A Anatomy of the Monocotyledons. Iridaceae. Clarendon Press, Oxford. RUDALL, P. & GOLDBLATT, P Leaf anatomy and phylogeny of Ixioideae (Iridaceae). Botanical Journal of the Linnean Society 106: SALT, H A voyage to Abyssinia etc. Reprinted, Cass Library of African Studies. Travels and Narratives 16. Frank Cass & Co., London. SCHINZ, H Beiträge zur Kenntnis der Flora von Deutsch-Südwest-Afrika und der angrenzenden Gebiete. IV. Verhandlungen des botanische Vereins der Provinz Brandenburg 31: SCHINZ, H Die Pflanzenwelt Deutsch-Südwest-Afrikas. Bulletin de l Herbier Boissier 4. App. III SCHINZ, H Beiträge zur Kenntnis der Afrikanischen Flora. (Neue Folge). XII. Mémoires de l Herbier Boissier 20: SCHLECHTER, R Decades plantarum novarum austro-africanum. Journal of Botany 35: SCHLECHTER, R Plantae schlechterianae novae vel minus cognitae describuntur II. Botanische Jahrbucher für Systematik 27: SCHULZE, W Beitrage zur Pollenmorphologie der Iridaceae und ihre Bedeutung fur die Taxonomie. Feddes Repertorium 82: SÖLCH, A Iridaceae. In H. Merxmüller (editor), Prodromus einer Flora von Südwestafrika 155: J. Cramer, Lehre. SPRENGEL, K.P.J Anleitung zur Kenntnis der Gewächse, edition 2, 2. Kummel, Halle.

150 142 S T R E L I T Z I A 35 (2015) SPRENGEL, K.P.J Systema vegetabilium, vol. 1. Dieterich, Gottingen. STEUDEL, E.T Nomenclator botanicus, ed. 2. J.G. Cotta, Stuttgart. SUESSENG, K In K. Suessenguth & H. Merxmüller, Species et varietates novae in Africa australis et centrali lectae. Mitteilungen der Botanische Staatssammlung, Munchen 1: SWEET, R Hortus britannicus. Ridgeway, London. SWEET, R. 1830a. Hortus britannicus, ed. 2. Ridgeway, London. SWEET, R. 1830b. Peyrousia aculeata. British Flower Garden ser. 2: t. 29. THUNBERG, C.P Dissertatio de Gladiolo. Edman, Uppsala. THUNBERG, C.P Prodromus plantarum capensium. Edman, Uppsala. VALENTE, L.M., MANNING, J.C., GOLDBLATT, P., VAR- GAS, P Did pollination shifts drive diversification in Southern African Gladiolus? Evaluating the model of pollinator-driven speciation. American Naturalist 180: VAUPEL, F Iridaceae Africanae novae. Botanische Jahrbucher für Systematik 48: WALPERS, G.G Ordo CCXLIX. Iridaceae Lindl. Annales Botanices Systematicae 3: WANNTORP, H.-E Lapeirousia rivularis, a new species of Iridaceae from South West Africa. Svensk Botanisk Tidskrift 65: WATT, J.M. & BREYER-BRANDWIJK, M.G Medicinal and poisonous plants of southern and eastern Africa, ed. 2. Livingstone Ltd., Edinburgh. WIJNANDS, D.O., WILSON, M.L. & TOUSSAINT VAN HOVE, T Jan Commelin s monograph on the Cape Flora. University of Stellenbosch, Stellenbosch. WILLDENOW. C.L Species plantarum 1. Nauk, Berlin.

151 S T R E L I T Z I A 35 (2015) 143 Index Acidanthera schimperi (Aschers. & Klatt) R.C.Foster Acidanthera unicolor Hochst. ex Baker Anomatheca angolensis Baker Chasmatocallis R.C.Foster Chasmatocallis macowanii R.C.Foster Codonorhiza Goldblatt & J.C.Manning Codonorhiza azurea (Eckl. ex Baker) Goldblatt & J.C.Manning Codonorhiza corymbosa (L.) Goldblatt & J.C.Manning Codonorhiza elandsmontana Goldblatt & J.C.Manning Codonorhiza falcata (L.f.) Ker Gawl Codonorhiza fastigiata (Lam.) Ker Gawl Codonorhiza micrantha (E.Mey. ex Klatt) Goldblatt & J.C.Manning Codonorhiza pillansii Goldblatt & J.C.Manning Freesia angolensis (Baker) Klatt Galaxia plicata Jacq Geissorhiza abyssinica R.Br. ex A.Rich Geissorhiza abyssinica R.Br. in Salt Gladiolus anceps L.f Gladiolus bracteatus Thunb Gladiolus denticulatus Lam Gladiolus fabricii (D.Delaroche) Thunb Gladiolus falcatus L.f Gladiolus fissifolius Jacq Gladiolus setifolius L.f Gladiolus silenoides Jacq Ixia corymbosa L Ixia crispifolia Andrews Ixia dinteri Schinz Ixia fabricii D.Delaroche Ixia fastigiata Lam Ixia heterophylla Willd Ixia lapeirousia J.F.Gmel Ixia plicata L Ixia pyramidalis Lam Lapeirousia Pourr sect. Lapeirousia sect. Chasmatocallis (R.C.Foster) Goldblatt & J.C.Manning sect. Sophronia (Licht. ex Roem. & Schult.) Goldblatt & J.C.Manning sect. Fastigiatae [Fastigiata] Goldblatt sect. Paniculatae [Paniculata] subg. Paniculatae [Paniculata] Goldblatt & J.C.Manning subg. Sophronia (Licht. ex Roem. & Schult.) Baker Lapeirousia abyssinica (R.Br. ex A.Rich.) Baker Lapeirousia aculeata Sweet Lapeirousia anceps (L.f.) Ker Gawl Lapeirousia anceps var. aculeata (Sweet) Baker Lapeirousia angolensis (Baker) R.C.Foster Lapeirousia angolensis Goldblatt

152 144 S T R E L I T Z I A 35 (2015) Lapeirousia angustifolia Schltr Lapeirousia arenicola Schltr Lapeirousia avasmontana Dinter Lapeirousia azurea (Eckl. ex Baker) Goldblatt Lapeirousia azurea Eckl Lapeirousia bainesii Baker Lapeirousia bainesii var. breviflora Baker Lapeirousia barklyi Baker Lapeirousia bracteata (Thunb.) Ker Gawl Lapeirousia burchellii Baker Lapeirousia caespitosa (Licht. ex Roem. & Schult.) Baker Lapeirousia caudata Schinz Lapeirousia caudata subsp. burchellii (Baker) Marais & Goldblatt Lapeirousia coerulea Schinz Lapeirousia compressa Pourr Lapeirousia congesta Rendle Lapeirousia corymbosa (L.) Ker Gawl Lapeirousia corymbosa subsp. alta Goldblatt Lapeirousia corymbosa subsp. fastigiata (Lam.) Goldblatt Lapeirousia corymbosa var. azurea Eckl. ex Baker Lapeirousia corymbosa var. grandiflora Klatt Lapeirousia corymbosa var. purpurea Klatt Lapeirousia cyanescens Welw. ex Baker Lapeirousia delagoensis Baker Lapeirousia denticulata G.H.M.Lawr Lapeirousia dinteri Vaupel Lapeirousia divaricata Baker Lapeirousia divaricata var. spinosa Goldblatt Lapeirousia divaricata var. tenuis Goldblatt Lapeirousia dolomitica Dinter Lapeirousia dolomitica subsp. lewisiana (B.Nord.) Goldblatt Lapeirousia edulis Schinz Lapeirousia effurcata G.J.Lewis Lapeirousia erythreae Chiov Lapeirousia erythrantha (Klotzsch ex Klatt) Baker Lapeirousia exilis Goldblatt Lapeirousia fabricii (D.Delaroche) Ker Gawl subsp. compressa (Pourr.) Goldblatt & J.C.Manning subsp. fabricii subsp. purpurascens Goldblatt & J.C.Manning Lapeirousia fasciculata Ker Gawl Lapeirousia falcata (L.f.) Ker Gawl Lapeirousia fastigiata (Lam.) Ker Gawl Lapeirousia fissifolia Ker Gawl Lapeirousia fragrans Welw. ex Baker Lapeirousia galaxioides Baker Lapeirousia gracilis Vaupel Lapeirousia heterophylla (Willd.) R.C.Foster Lapeirousia homoidea Klatt Lapeirousia jacquinii N.E.Br Lapeirousia juttae Dinter Lapeirousia kalahariensis Goldblatt & J.C.Manning Lapeirousia kamiesmontana Goldblatt & J.C.Manning Lapeirousia lacinulata Vaupel Lapeirousia lewisiana B.Nord Lapeirousia littoralis Baker subsp. caudata (Schinz) Goldblatt Lapeirousia macrospatha Baker... 61

153 S T R E L I T Z I A 35 (2015) 145 Lapeirousia manuleiflora Eckl Lapeirousia masukuensis Vaupel & Schltr Lapeirousia micrantha (E.Mey. ex Klatt) Baker Lapeirousia montana Klatt Lapeirousia monteiroi Baker Lapeirousia montaboniana Chiov Lapeirousia montana Hutch Lapeirousia neglecta Goldblatt Lapeirousia odoratissima Welw. ex Baker Lapeirousia oreogena Schltr. ex Goldblatt Lapeirousia otaviensis R.C.Foster Lapeirousia pappei Baker Lapeirousia pentheri Baker Lapeirousia plicata (Jacq.) Diels subsp. effurcata (G.J.Lewis) Goldblatt subsp. plicata subsp. foliosa Goldblatt & J.C.Manning subsp. longifolia Goldblatt Lapeirousia porphyrosiphon Baker Lapeirousia purpurea Goldblatt & J.C.Manning Lapeirousia purpureolutea (Klatt) Baker Lapeirousia pyramidalis (Lam.) Goldblatt subsp. pyramidalis subsp. regalis Goldblatt & J.C.Manning Lapeirousia ramosissima Dinter Lapeirousia rivularis Wanntorp Lapeirousia sandersonii Baker Lapeirousia schimperi (Aschers. & Klatt) Milne-Redh Lapeirousia serrulata Schltr Lapeirousia setifolia (L.f.) N.E.Br Lapeirousia setifolia Harms Lapeirousia silenoides (Jacq.) Ker Gawl Lapeirousia simulans Goldblatt & J.C.Manning Lapeirousia speciosa Schltr Lapeirousia spinosa (Goldblatt) Goldblatt & J.C.Manning Lapeirousia stenoloba Vaupel Lapeirousia streyi Suess Lapeirousia tenuis (Goldblatt) Goldblatt & J.C.Manning Lapeirousia teretifolia (Geerinck, Lisowski, Malaisse & Symoens) Goldblatt Lapeirousia uliginosa Dinter Lapeirousia vaupeliana Dinter Lapeirousia verecunda Goldblatt Lapeirousia violacea Goldblatt Lapeirousia welwitschii Baker Lapeirousia zambeziaca Goldblatt Meristostigma A.Dietr Meristostigma aculeatum (Sweet) Steud Meristostigma anceps (L.f.) A.Dietr Meristostigma azureum Eckl. ex Steud Meristostigma bracteatum (Thunb.) A.Dietr Meristostigma corymbosum (L.) A.Dietr Meristostigma fabricii (D.Delaroche) A.Dietr Meristostigma falcatum (L.f.) A.Dietr Meristostigma fissifolium (Ker Gawl.) A.Dietr Meristostigma heterophyllum (Willd.) A.Dietr Meristostigma manuleiflorum (Eckl.) Steud Meristostigma silenoides (Jacq.) A.Dietr Montbretia abyssinica Hochst. ex A.Rich

154 146 S T R E L I T Z I A 35 (2015) Montbretia abyssinica Hochst Ovieda Spreng Ovieda aculeata (Sweet) Klatt Ovieda anceps (L.f.) Spreng Ovieda bracteata (Thunb.) Spreng Ovieda corymbosa (L.) Spreng Ovieda fabricii (D.Delaroche) Spreng Ovieda falcata (L.f.) Spreng Ovieda fasciculata (Ker Gawl.) Spreng Ovieda fissifolia (Ker Gawl.) Spreng Ovieda micrantha E.Mey. ex Klatt Ovieda purpureolutea Klatt Ovieda silenoides (Jacq.) Spreng Peyrousia Poir Peyrousia aculeata Sweet Peyrousia anceps (L.f.) Poir Peyrousia bracteata (Thunb.) Sweet Peyrousia corymbosa (L.) Poir Peyrousia fabricii (D.Delaroche) Sweet Peyrousia falcata (L.f.) Sweet Peyrousia fasciculata (Ker Gawl.) Sweet Peyrousia fissifolia (Ker Gawl.) Poir Peyrousia silenoides (Jacq.) Poir Psilosiphon Welw. ex Goldblatt & J.C.Manning Psilosiphon abyssinicus (R.Br. ex A.Rich.) Goldblatt & J.C.Manning Psilosiphon avasmontanus (Dinter) Goldblatt & J.C.Manning Psilosiphon bainesii (Baker) Goldblatt & J.C.Manning Psilosiphon coeruleus (Schinz) Goldblatt & J.C.Manning Psilosiphon erongoensis Goldblatt & J.C.Manning Psilosiphon erythranthus (Klotzsch ex Klatt) Goldblatt & J.C.Manning Psilosiphon gracilis (Vaupel) Goldblatt & J.C.Manning Psilosiphon masukuensis (Vaupel & Schltr.) Goldblatt & J.C.Manning Psilosiphon otaviensis (R.C.Foster) Goldblatt & J.C.Manning Psilosiphon rivularis (Wanntorp) Goldblatt & J.C.Manning Psilosiphon sandersonii (Baker) Goldblatt & J.C.Manning subsp. limpopoensis Goldblatt & J.C.Manning subsp. sandersonii Psilosiphon schimperi (Aschers. & Klatt) Goldblatt & J.C.Manning Psilosiphon setifolius (Harms) Goldblatt & J.C.Manning Psilosiphon teretifolius (Geerinck, Lisowski, Malaisse & Symoens) Goldblatt Psilosiphon zambeziacus (Goldblatt) Goldblatt & J.C.Manning Schizorhiza Goldblatt & J.C.Manning Schizorhiza neglecta (Goldblatt) Goldblatt & J.C.Manning Sophronia Licht. ex Roem. & Schult Sophronia caespitosa Licht. ex Roem. & Schult Tritonia abyssinica (Hochst. ) Walp Tritonia schimperi Aschers. & Klatt Witsenia pyramidalis (Lam.) Pers

155

156