Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 43,2

October 2013

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN

NATIONAL BIODIVERSITY INSTITUTE (SANBI) PRETORIA

Obtainable from SANBI, Private Bag XI 01, Pretoria 0001,

RSA. A catalogue of all available publications will be issued on

request.

BOTHALIA

Bothalia is named in honour of General Louis Botha, first

Premier and Minister of Agriculture of the Union of South

Africa. This house journal of SANBI, Pretoria, is devoted to

the furtherance of botanical science. The main fields covered

are taxonomy, ecology, anatomy and cytology. Two parts of

the journal and an index to contents, authors and subjects are

published annually.

Bothalia Contents: five booklets containing a list of authors

and titles, and an index to taxa and keywords: a) to vols 1-20

(1921-1990); b) to vols 21-25 (1991-1995); to vols 26-30

(1996-2000); d) to vols 31-37 (2001-2007); to vols 38^1

(2008-2011).

FLORA OF SOUTHERN AFRICA (FSA)

A taxonomic treatise on the flora of the Republic of South

Africa, Lesotho, Swaziland, Namibia and Botswana. Contains

descriptions of families, genera, species, infraspecific taxa,

keys to genera and species, synonymy, literature and limited

specimen citations, as well as taxonomic and ecological notes.

Project discontinued.

FSA contributions 1-22 appear in Bothalia:

1 : Aquifoliaceae. S. Andrews. 1994. Bothalia 24: 163-166.

2: Asphodelaceae/Aloaceae, 1029010 Chortolirion. G.F. Smith.

1995. Bothalia 25: 31-33.

3: Asphodelaceae/Aloaceae, 102S010 Poellnitzia. G.F. Smith

\995. Bothalia 25: 35,36.

4: Agavaceae. G.F. Smith & M. Mdssmer. 1996. Bothalia 26:

31-35.

5: Buxaceae. H.F. Glen. 1996. Bothalia 26: 37-AO.

6: Orchidaceae: Holothrix. K.L. Immelman. 1996. Bothalia 26:

25^0.

7: Verbenaceae: Vitex. C.L. Bredenkamp & D.J. Botha. 1996.

Bothalia 26: 141-151.

8: Ceratophyllaceae. C.M. Wilmot-Dear. 1997. Bothalia 27:

125-128.

9: Onagraceae. P. Goldblatt &. PH. Raven. 1997. Bothalia 21:

149-165.

10: Trapaceae. B. Verdcourt. 1998. Bothalia 28: 11-14.

1 1 : Zingiberaceae. R.M. Smith. 1998. Bothalia 28: 35-39.

12: Plantaginaceae. H.F. Glen. 1998. Bothalia 28: 151-157.

13: Ulmaceae. C.M. Wilmot-Dear. 1999. Bothalia 29: 239-247.

14: Cannabaceae. C.M. Wilmot-Dear. 1999. Bothalia 29:

249-252.

15: Piperaceae. K.L. Immelman. 2000. Bothalia 30: 25-30.

16: Sphenocleaceae. W.G. Welman. 2000. Bothalia 30: 31-33.

17: Casuarinaceae. C.M. Wilmot-Dear. 2000. Bothalia 30:

143-146.

18: Salicaceae s. str. M. Jordaan. 2005. Bothalia 35: 7-20.

19: Asteraceae: Anthemideae: Eumorphia. N. Swelankomo.

201 1.5o//?a/m 41: 277-282.

20: Asteraceae: Anthemideae: Inezia. N. Swelankomo. 2012.

Bothalia 42: 247-250.

21 : Connaraceae. M. Jordaan. 2013. Bothalia 43: 99-106.

22. Asteraceae: Calenduleae: Garuleum. N. Swelankomo. 2013.

Bothalia 42: 167-178.

STRELITZIA

A series of occasional publications on southern African flora and

vegetation, replacing Memoirs of the Botanical Survey of South

Africa and Annals of Kirstenbosch Botanic Gardens.

• The Memoirs are individual treatises usually of an ecological

nature, but sometimes dealing with taxonomy or economic

botany. Published: Nos 1-63 (many out of print). Discontin-

ued after No. 63.

• The Annals are a series devoted to the publication of mono-

graphs and major works on southern African flora. Published:

Vols 14-19 (earlier vols published as suppl. vols to the Jour-

nal of South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA {FPA)

This serial presents colour plates of African plants with

accompanying text. The plates are prepared mainly by the

artists at SANBI. Many botanical artists have contributed to the

series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian

Condy, Betty Connell, Stella Gower, Rosemary Holcroft,

Kathleen Lansdell, Cythna Letty (over 700 plates), Claire

Linder-Smith and Ellaphie Ward-Hilhorst. The Editor is pleased

to receive living plants of general interest or of economic value

for illustration.

From Vol. 55, twenty plates are published at irregular intervals.

An index to Vols 1^9 is available.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeoflora on a pattern comparable to that of the Flora

of southern Africa. Much of the information is presented in

the form of tables and photographic plates depicting fossil

populations. Now available:

• Molteno Formation (Triassic) Vol. 1 . Introduction. Dicroidi-

um, 1983, by J.M. & H.M. Anderson.

• Molteno Formation (Triassic) Vol. 2. Gymnosperms (exclud-

ing Dicroidium), 1983, by J.M. & H.M. Anderson.

• Prodromus of South African Megafloras. Devonian to Lower

Cretaceous, 1985, by J.M. & H.M. Anderson. Obtainable from:

A.A. Balkema Marketing, Box 317, Claremont 7735, RSA.

• Towards Gondwana Alive. Promoting biodiversity and stem-

ming the Sixth Extinction, 1999, by J.M. Anderson (ed.).

• Heyday of the gymnosperms: systematics and biodiversity of

the Late Triassic Molteno fructifications, 2003, by J.M. An-

derson & H.M. Anderson. Strelitzia 15.

• Brief history of the gymnosperms: classification, biodiver-

sity, phytogeography and ecology, 2007, by J.M. Anderson,

H.M. Anderson & C.J. deal. Strelitzia 20.

• Molteno ferns: Late Triassic biodiversity in southern Africa,

2008, by H.M. Anderson & J.M. Anderson. Strelitzia 21.

SANBI BIODIVERSITY SERIES

A series of occasional reports on projects, technologies,

workshops, symposia and other activities initated by or executed

in partnership with SANBI.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 43,2

Scientific Editor; J.C. Manning

Production Editor; Y. Steenkamp

SANBI

Biodiversity for Life

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI 01, Pretoria 0001

ISSN 0006 8241

October 2013

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Archer, Mrs C., South African National Biodiversity Institute, Pretoria, RSA.

Bernhardt, Prof P., St Louis University, St Louis, USA.

Coetzer, Dr L.A., ex University of Pretoria, Pretoria, RSA.

Cupido, Dr C., South African National Biodiversity Institute, Cape Town, RSA.

Forest, Dr F., Royal Botanic Gardens, Kew, UK.

Geerinck, Dr D., Meise Herb., Jardin Botanique National, Meise, Belgium.

Germishuizen, Mr G., ex South African National Biodiversity Institute, Pretoria, RSA.

Goldblatt, Prof. R, Missouri Botanical Garden, St Louis, USA.

Grobler, Mrs A., South African National Biodiversity Institute, Pretoria, RSA.

Herman, Mr P.P.J., South African National Biodiversity Institute, Pretoria, RSA.

Jones, Prof C.S., University of Connecticut, Storrs, USA.

Kissling, Dr J., University of Neuchatel, Neuchatel, Switzerland.

Klopper, Mrs, R.R., South African National Biodiversity Institute, Pretoria, RSA.

Le Roux, Dr M., South African National Biodiversity Institute, Pretoria, RSA.

Leistner, Dr O.A., ex South African National Biodiversity Institute, Pretoria, RSA.

Magee, Dr A.R., South African National Biodiversity Institute, Cape Town, RSA.

Manning, Dr J.C., South African National Biodiversity Institute, Cape Town, RSA.

Martinez-Azorin, Dr M., Universidad de Alicante, Spain.

Musselman, Prof L.J., Old Dominion University, Norvolk, USA.

Nickrent, Prof D.L., Southern Illinois University, Carbonedale, USA.

Nordenstam, Prof. R.B., Swedish Museum of Natural History, Stockholm, Sweden.

Pelser, Dr P.B., University of Canterbury, Christchurch, New Zealand.

Phillipson, Mr P.B., Missouri Botanical Garden, St Louis, USA.

Proches, Dr S., University of KwaZulu-Natal, Durban, RSA.

Raimondo, Ms D.C., South African National Biodiversity Institute, Pretoria, RSA.

Retief, Dr E., ex South African National Biodiversity Institute, Pretoria, RSA.

Singh, Dr Y., South African National Biodiversity Institute, Durban, RSA.

Snijman, Dr D.A., South African National Biodiversity Institute, Cape Town, RSA.

Steyn, Ms H.M., South African National Biodiversity Institute, Pretoria, RSA.

Stirton, Dr C., 4 Nelson Villas, Bath, BAl 2BD, Avon, UK.

Verboom, Prof T., University of Cape Town, Cape Town, RSA.

Date of publication of Bothalia 43,1: 1 May 2013.

Be aware that it has eome to SANBI’s attention that there is an online website purporting to be a website for on-line submissions

I'ortheSANBI house journal Bothalia.

Please note that the bogus website www.bothalia.eom has no conneetion or relevance to SANBI’s house Journal Bothalia, and

that manuscripts submitted to this bogus website will not be processed for publication in SANBI’s Bothalia.

SANF3I has no relationship with the owners of this website and would like to caution anyone from visiting this website or

submitting any content to it in the mistaken belicrihat it is a genuine site linked to the SANIil journal Bothalia.

CONTENTS

Bothalia 43,2

1. The Cape genus Micranthns (Iridaceae: Crocoideae), nomenclature and taxonomy. P. GOLD-

BLATT, J.C. MANNING and R.E. GEREAU 127

2. New taxa of Hesperantha (Iridaceae; Crocoideae) from the southern African winter rainfall

region and a review of the H. pilosa complex. P. GOLDBLATT and J.C. MANNfNG . . . 145

3. Eight new species of Moraea (Iridaceae) from southern Afriea with range extensions and mor-

phological notes in the genus. P. GOLDBLATT and J.C. MANNING 153

4. ESA Contribution 22; Asteraceae; Calenduleae; Gariileiim. N. SWELANKOMO 167

5. The native and naturalised species of Peltocalathos and Rammciilus (Ranunculaceae; Ranun-

culeae) in southern Afriea. J.C. MANNING and P. GOLDBLATT 179

6. A new infrageneric classification for Mesembiyanthemiim (Aizoaceae; Mesembryanthemoid-

eae). C. KLAK and P.V. BRUYNS ^ 197

7. Notes on African plants;

Acanthaceae & Lamiaceae. Nomenclatural corrections in Justicia and Leonotis. J.C. MAN-

NING 221

Asteraceae. Senecio pseiidolongifolius, a new name for the misapplied S. linifolius. J. CAL-

VO, J.C. MANNING, F. MUNOZ GARMENDIA and C. AEDO . 227

Fabaceae. A new species of Lessertia (Galegeae) from KwaZulu-Natal, South Africa. T.

NKONKI, A.M. NGWENYA and B.-E. VAN WYK 222

Gentianaceae. The taxonomic and conservation status of Sebaea fourcadei. O. BALOYI,

J.E. VICTOR and N. SWELANKOMO 225

Geraniaceae. Pelargonium pachypodium (sect. Otidia), a new species from Northern Cape,

South Africa. J.P. ROUX 230

Geraniaceae. Validating Pelargonium sect. Reniformia. J.P. ROUX 233

Hyacinthaceae. Five new combinations in Omithogaloideae in southern Africa and a rec-

ommendation for optional combinations in the subfamily. J.C. MANNING and P.

GOLDBLATT 229

Iridaceae. A new species of Hesperantha from the Overberg, Western Cape, and observa-

tions on a novel mode for pollen transfer in the genus and family by a hesperid but-

terfly. P. GOLDBLATT, J.C. MANNING and O.E. CURTIS 211

Iridaceae. Taxonomic notes on Aristea (Aristeoideae) in tropical and eastern southern Af-

rica. P. GOLDBLATT and J.C. MANNING 207

Isoetaceae. Isoetes aemulans, a new species from South Africa. J.P. ROUX 218

Polygalaceae. The reinstatement of Polygala affinis and the identity of Polygala scabra. E.

FIGUEIREDO, J. PAIVA and G.F. SMITH .' 216

Santalaceae. The minor genera Kunkeliella and Thesidium included in Thesium. F. FOREST

and J.C. MANNING 214

8. South African National Biodiversity Institute; publications 1 January 2012 to 31 December

2012. Compiler; Y. STEENKAMP 235

9. Guide for authors to Bothalia 241

New combinations, names, sections, series, species and statuses in Bothalia 43,2 (2013)

Albuca chartacea (Mart.-Azorin et alj J.C. Manning & Goldblatt, comb, nov., 229

Albuca comosa (Welw. ex Baker) J.C. Manning & Goldblatt, comb, nov., 229

Albuca craibii (Mart.-Azorin et al.j J.C. Manning & Goldblatt, comb, nov., 229

Albuca recurva (Oberin.) J.C. Manning & Goldblatt, comb, nov., 230

Aristea gerrardii Weim., syn. nov., 210

Hesperantha ciliolata Goldblatt, syn. nov., 149

Hesperantha dolomitica Goldblatt & J.C. Manning, sp. nov., 145

Hesperantha kiaratayloriae Goldblatt & J.C. Manning, sp. nov., 211

Hesperantha laxifolia Goldblatt & J.C. Manning, sp. nov., 146

Hesperantha pilosa subsp. bracteolata (R.C. Foster) Goldblatt & J.C. Manning, comb, et stat. nov., 149

Hesperantha secunda Goldblatt & J.C. Manning, sp. nov., 150

Isoetes aemulans J.P.Roiix, sp. nov., 218

Justicia tubulosa subsp. late-ovata (C.B. Clarke) J.C. Manning & Goldblatt, comb, nov., 221

Kunkeliella W.T.Stearn, syn. nov., 215

Leonotis pentadentata J.C. Manning & Goldblatt, nom. nov. pro, 222

Lessertia amajubica Nkonki, sp. nov., 222

Mesembryanthemum sect. Brownanthus (Schwantes) Klak, comb, et stat. nov., 202

Mesembryanthemum sect. Callistigma (Dinter & Schwantes) Klak, comb, et stat. nov., 201

Mesembryanthemum seet. Eurystigma (L. Bolus) Klak, comb, et stat. nov., 202

Mesembryanthemum sect. Phyllobolus (N.E.Br) Klak, comb, et stat. nov., 203

Mesembryanthemum sect. Volkeranthus (Gerbaulet) Klak, comb, et stat. nov., 206

Micranthus cruciatus Goldblatt & J.C. Manning, sp. nov., 139

Micranthus filifolius Goldblatt & J.C. Manning, sp. nov., 137

Micranthus simplex Goldblatt & J.C. Manning, sp. nov., 133

Micranthus thereianthoides Goldblatt & J.C. Manning, sp. nov., 134

Moraea doleritica Goldblatt & J.C. Manning, sp. nov., 160

Moraea eburnean Goldblatt & J.C. Manning, sp. nov., 160

Moraea filamentosa Goldblatt & J.C. Manning, sp. nov., 154

Moraea lazulina Goldblatt & J.C. Manning, sp. nov., 158

Moraea petricola Goldblatt & J.C. Manning, sp. nov., 158

Moraea singularis Goldblatt & J.C. Manning, sp. nov., 154

Moraea striata Goldblatt & J.C. Manning, sp. nov., 153

Moraea thermarum Goldblatt & J.C. Manning, sp. nov., 156

Ornithogalum tanquanum (Mart.-Azorin & M.B. Crespo) J.C. Manning & Goldblatt, comb, nov., 230

Osteospermwn sonchifolium var. subpetiolatum Harv., syn. nov., 168

Pelargonium pachypodium J.P.Roux, sp. nov., 230

Pelargonium sect. Reniformia (R.Knuth) Dreyer ex J.P.Roux, comb, et stat. nov., 233

Ranunculus J.C. Manning & Goldblatt, nom. nov. pro, 190

Ranunculus multifidus Forssk., syn. nov., 185

Ranunculus pinnatus var. hermannii DC., syn. nov., 185

Ranunculus pubescens var. glabrescens Burtt Davy, syn. nov., 185

Ranunculus pubescens var. harveyanus Burtt Davy, syn. nov., 185

Senecio pseudolongifolius Sch.Bip. exJ.Calvo, nom. nov. pro, 228

Thesium canariensis (W.T.Stearn) J.C. Manning & F.Forest, comb, nov., 216

Thesium confusum J.C. Manning & F.Forest, nom. nov. pro, 215

Thesium fruticulosum (A.W.Hill) J.C. Manning & F.Forest, comb, nov., 215

Thesium glohulosum A. DC., syn. nov., 2 1 6

Thesium minus (A. W.Hill) J.C. Manning & F.Forest, comb, nov., 215

Thesium retamoides (A. Santos) J.C. Manning & F.Forest, comb, nov., 216

Thesium sect. Kunkeliella (W.T.Stearn) J.C. Manning & F.Forest, stat. et comb, nov., 216

Thesium subsucculentum (Kdnvner) J.C. Manning & F.Forest, comb, nov.,216

IV

Bothalia 43,2: 127-144(2013)

The Cape genus Micranthus (Iridaceae: Crocoideae), nomenclature

and taxonomy

P. GOLDBLATT' \ J.C. MANNING-^ & R.E. GEREAU-*

Keywords: Cape flora, Iridaceae, Micranthus, new species, nomenclature, taxonomy

ABSTRACT

The genus Micranthus (Pers.) Eckl., has traditionally been treated as comprising three species, all with virtually identi-

cal, bilaterally symmetric, deep or pale blue to white flowers arranged in crowded, 2-ranked spikes and with divided style

branches, but differing in their foliage. Examination of plants in the field and herbarium shows that there are four additional

species. M. filifolius Goldblatt & J.C. Manning, from the Caledon District of the southwestern Western Cape, has up to six,

filiform leaves, the blades of at least the lowermost terete and cross-shaped in section, and usually pale blue-mauve flowers.

M. simplex Goldblatt & J.C. Manning from high elevations on Zebrakop, Piketberg, has the smallest flowers in the genus,

white but tinged lilac as they age, linear leaves up to 1 ,5 mm wide, and undivided style branches. M. cruciatus Goldblatt

& J.C. Manning, from the northern Cedarberg and Bokkeveld Mtns, has up to four leaves, the lower with linear or terete

blades with heavily thickened margins and central vein and relatively large flowers, unusual in having the style dividing at

the mouth of the perianth tube into particularly long branches, these deeply divided as is typical of the genus. M. thereian-

thoides Goldblatt & J.C. Manning, from the Paardeberg south of Malmesbury, is unique in the genus in having flowers with

an elongate perianth tube. We also document the occurrence of large populations of putative hybrids at some sites. We provide

a complete revision of Micranthus with original observations on leaf anatomy, pollen morphology and reproductive biology

and discuss its confused taxonomic and nomenclatural history and that of the three common species of the genus, known for

over 1 50 years. In so doing, we neotypify Gladiolus alopecuroides L. ( 1 756) [= Micranthus alopecuroides (L.) Eckl. ( 1 827)],

type of the genus, and choose lectotypes for M. plantagineus Eckl. var.jiinceus Baker (1892) and Gladiolus fistulosus Jacq.

Now with seven species, Micranthus remains endemic to the Cape flora region, extending from its extreme northern limit

in the Bokkeveld Mtns south-eastwards to Port Elizabeth. We also deal with the genera Paulomagnusia Kuntze and Beilia

Kuntze with w'hich Micranthus has sometimes been associated, although both are nomenclatural synonyms of Thereianthus

G.J. Lewis, a genus close allied to Micranthus.

INTRODUCTION

Micranthus (Pers.) Eckl., endemic to the Cape flora

region of South Africa, was first recognized as a genus

when Ecklon (1827) raised Gladiolus subgen. Micran-

thus Pers. to generic rank. He admitted three species to

the genus, in this order: M. plantagineus Eckl., M. alo-

pecuroides (L.) Eckl. (based on Gladiolus alopecuroides

L. ), and M. fistulosus Eckl. The names M. fistulosus and

M. plantagineus appear to be implicit references respec-

tively to Gladiolus fistulosus Jacq. (1797) and Ixia plan-

taginea Alton (1789), the latter a superfluous name for

G. alopecuroides. An indirect reference opposite the

genus name "Watsonia Link’, leads to Link (1821), in

which Watsonia plantaginea Ker Gawl. (1803) is listed,

and this work in turn cites Ixia plantaginea. We thus

treat M. plantagineus Eckl. as a new name in Micran-

thus with its type that of Ixia plantaginea. We find no

such indirect reference for M. fistulosus and the name

must continue to be regarded as a nomen nudum and

therefore invalid.

' B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P. O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt(^'mobot.org.

“ Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. E-mail: j.manning@

sanbi.org. za.

^ Research Centre for Plant Growth and Development, School of Bio-

logical and Conservation Sciences, University of KwaZulu-Natal, Pie-

termaritzburg, Private Bag XOl, Scottsville 3209.

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166,

USA, Email: roy.gereau(2;mobot.org.

Until now, Micranthus has included just three spe-

cies (Lewis 1950; Goldblatt & Manning 2000); M.

alopecuroides, M. tubulosus (Burm.f.) N.E.Br. (1929)

[with Gladiolus fistulosus a heterotypic synonym],

and M. junceus (Baker) N.E.Br. (1929) [a combination

based on M. plantagineus xm. junceus Baker (1892)

and a later name for M. plantagineus Eckl.]. Micran-

thus (Pers.) Eckl. (1827), itself a later homonym, is con-

served against Micranthus J.C.Wendl. ( 1798), a genus of

Acanthaceae, with Gladiolus alopecuroides L. (M. alo-

pecuroides (L.) Eckl.) as its conserved type (Rickett &

Stafleu 1959: 241; McNeill et al. 2006: 272).

These three species of Micranthus are mostly read-

ily distinguished by their leaf morphology: M. alo-

pecuroides has plane, ± lanceolate to falcate leaves with

an evident main vein (Figure lA); M. tubulosus has two

or more short, inflated, terete, hollow (fistulose), falcate

leaves, usually half as long as the stem (Figure IE); and

M. plantagineus is a taller plant with long, terete, hol-

low, straight foliage leaves (Figure IF). All three spe-

cies have apparently identical, small, mid to deep blue

(sometimes described as violet), blue-mauve or occa-

sionally white, bilabiate flowers arranged in congested,

2-ranked spikes subtended by dry, brittle bracts with

broad membranous margins. The conns, capsules, and

specialized, narrow, 3-sided, elongate seeds are also sim-

ilar in all three species. Apart from their leaf differences,

each species shows a modest preference for a different

habitat: M. alopecuroides is most often found on sandy

ground; M. tubulosus on dry, usually shale- or granite-

derived soils; and M. plantagineus in wet habitats, often

in marshes, seeps or along streams, most often in sandy

or peaty soils. That said, we have seen two or even all

128

Bothalia 43,2 (2013)

three species growing together locally with only very

small habitat differences, if any, so habitat preferences

are far from absolute.

Several additional populations of Micranthus extend

the range of leaf morphology in the genus. Plants at

high elevations in the Piketberg (Goldblatt & Man-

ning 10172, MO, NBG) have narrow, straight to falcate

leaves ± 1 mm wide with one or more strongly thick-

ened veins, small, white flowers fading pale lilac, and

undivided style branches. These plants grow in an unu-

sual habitat for Micranthus, crevices and shallow pock-

ets of soil on wet sandstone rocks.

A second series of populations (e.g. Goldblatt 10438

MO, NBG) from the northern Cedarberg and Bokkeveld

Mtns has long, slender, linear leaves with a heavily

thickened main vein and equally thick margins, thus

often cross-shaped in transverse section (Figure ID).

These slender, often tall, plants also stand out in having

the style dividing at the mouth of the perianth tube, with

unusually long style branches divided for less than a

FIGURE I . — Leaf anatomy in Micrcmiliiis. A, M. alopecwvicJes, Elandsberg Nature Reserve, Goldhia/l & Manning I36I6\ B, M.filifoliiis, Kogel-

berg, no voucher; C, M. filifolius. Drayton, Gokihlall Manning 13623', D, M. crncialiis, Pakhuis, Golctblall & Porter 13766', E, M. liilni-

losus, Roncicbosch Common, Goldhtall & Manning 13620', F, M. planlagineiis, Drayton, Goldblatt di Manning 13632. Scale bar: 500 gm.

Bothalia 43,2 (2013)

129

quarter of their length. They grow in seasonally marshy

sites in peaty soil, often in moss on sandstone pavement,

and appear to flower particularly well after fire.

A third series of populations from the Caledon Dis-

trict of Western Cape has up to 6 leaves with linear to

filiform blades, often cross-shaped in section (Figure IB,

C), with the bases persisting in a well-developed fibrous

neck. These plants grow on stony, loamy clay or sandy

soils, usually in well drained sites that are dry at flower-

ing time.

Lastly, a population from the Paardeberg south of

Malmesbury, only discovered in 2012, has hollow leaves

reminiscent of those of Micranthus tubidosus but is

unique in the genus in having dark blue flowers with an

elongate perianth tube, 20-22 mm long, thus more than

twice as long as in any other species of Micranthus.

Consistent treatment of the genus suggests that

these divergent populations should logically be rec-

ognized as separate species. The circumscriptions of

the existing species cannot be expanded to accom-

modate these plants. We describe these new species as

Micranthus cruciatus Goldblatt & J.C. Manning, M.

filifolius Goldblatt & J.C. Manning, M. simplex Gold-

blatt & J.C. Manning and M. thereianthoides Goldblatt

& J.C. Manning. Other variants, which we believe are

interspecific hybrids, occur locally and we discuss these

below. One of them, evidently Micranthus plantagineus

X M. tubidosus, is particularly common at the foot of the

Elandskloof Mtns. Plants have a flexuose stem, terete,

hollow leaves and short spikes of up to 10 flowers, and

appeared at first to be a separate species, so different

were they from their putative parents.

We review the nomenclature of Micranthus, choose

types for the two species currently lacking designated

types, and present a systematic revision, thus dealing

with collections that do not accord with the current cir-

cumscriptions of the three species included in the genus.

We also deal with Paulomagnusia Kuntze (1891). When

described, Paulomagnusia included two species, one a

Micranthus and the other, P. spicatus (L.) Kuntze, now

the type species of Thereianthus G.J. Lewis (1941). Our

revision includes new observations on leaf anatomy, pol-

len morphology, and reproductive biology and pollina-

tion; these presented following the generic description

and nomenclature.

TAXONOMIC HISTORY AND RELATIONSHIPS

Micranthus is most closely allied to Thereianthus,

also endemic to the Cape flora region. Lewis (1950) first

pointed out an unusual, specialized feature shared by

the two genera, namely that the lowermost foliage leaf

is inserted on the flowering stem as it is in Lapeirousia

Pourret. also in tribe Watsonieae Klatt. rather than on

the corm. This means that the corm tunics are formed

solely from the cataphylls, without any contribution

from the leaf bases as is found in Watsonia Mill, and

some other members of the tribe. Molecular systematic

studies of plastid DNA sequences confirm the immedi-

ate relationship of the two genera, which together are

sister to Watsonia plus Pillansia L. Bolus (Reeves et al.

2002; Goldblatt et al. 2008), with Lapeirousia (sens

lat. ) retrieved as member of a second clade of the tribe,

which includes Cyanixia Goldblatt & J.C. Manning and

Savannosiphon Goldblatt & Marais. The close relation-

ship ot Micranthus and Thereianthus is reflected in their

largely shared taxonomic and nomenclatural history.

Although Micranthus was maintained by most

authors dealing with the genus since its recognition at

generic rank by Ecklon (1827), species now recognized

as Thereianthus have had a more chequered history,

beginning with Ecklon (1827), who placed two spe-

cies of that genus in "Beilia', then lacking a validating

description. Although species of Micranthus and There-

ianthus had first been referred respectively to Ixia L.

or to Gladiolus L., they were included by Ker Gawler

(1804) in Watsonia, largely because they share divided

style branches with that genus. Their nomenclature

subsequently became intertwined. Heynhold (1847)

included the two species of Thereianthus known at that

time in Micranthus as M. spicatus (E.) Heyn. and M.

triticeus (Thunb.) Heyn. The British botanist and spe-

cialist in the taxonomy of Iridaceae, Baker ( 1877), rec-

ognized Micranthus in its current sense and included

one species of Thereianthus in Watsonia unranked Beilia

Eckl. ex Baker, as W. punctata (Andrews) Ker Gawl.

(now Thereianthus bracteolatus (Lam.) G.J. Lewis).

Baker ( 1892) later formalized Watsonia unranked Beilia,

then with several species, as Watsonia subgen. Beilia

(Eckl. ex Baker) Baker. His German contemporary, Klatt

(1882), completely misunderstood the situation, and in

his worldwide account of the Iridaceae, he included two

species of Thereianthus in Micranthus. as M. spicatus

(E.) Klatt (evidently referring to what is now T. spicatus

(L.) G.J. Lewis) and M. triticeus (Burm.f [sic]) Klatt [he

was evidently unaware of combinations in Micranthus

for these species by Heynhold in 1847; we also assume

that the basionym attributed to Burman fil. was an error

for Thunberg, as Ixia triticea Burm.f is a very different

species, currently Tritoniopsis triticea (Burm.f) Gold-

blatt]. Klatt (1882) also included one species of There-

ianthus, T. juncifolius (Baker) G.J. Lewis, in Anomatheca

Ker Gawl. as A. calamifolia Klatt, and several more in

Watsonia unranked Beila. Usually astute, Klatt appar-

ently made no reference to any species we now regard

as belonging to Micranthus but partly corrected the error

when he recognized M. plantagineus with one variety,

\diV. junceus (Klatt 1894). Thereianthus spicatus, how-

ever, remained in Micranthus.

Kuntze (1891) included both what are now Micran-

thus alopecuroides and Thereianthus spicatus in his new

genus Paulomagnusia, evidently intended as a nomen

novum for the later homonym, Micranthus (Pers.) Eckl.

1827 (non Micranthus J.C.Wendl. 1798). This genus

has now been conserved, although not against Paulo-

magnusia. Kuntze (1898) validated Ecklon ’s "Beilia'

at generic rank as Beilia Kuntze, and included in the

genus only B. spicata (L.) Eckl. ex Kuntze, which is

thus its type. Unfortunately Beilia is superfluous because

Kuntze listed the valid Paulomagnusia in synonymy. It

remained for Lewis (1941), over a century after Eck-

lon (1827) used the invalid "Beilia'', to erect the valid

genus Thereianthus in which she placed the two species

of Ecklon’s "Beilia' and several more then included in

130

Bothalia 43,2 (2013)

Watsonia. Thereianthus now has 11 species (Manning &

Goldblatt 2011).

SYSTEMATICS

Micranthus (Pers.) Eckl., Topographisches Verzeich-

niss der Pflanzensammlung von C.F. Ecklon: 43 (1827),

name conserved, non J.C.Wendl. (1898, Acanthaceae).

Gladiolus subg. Micranthus Pers.; 46 (1805). Hebea

subg./unranked Micranthus (Pers.) R.Hedw.: 24 (1806).

Type (conserved): Gladiolus alopecuroides L. (= M. alo-

pecuroides (L.) Eckl.).

Paulomagnusia Kuntze: 702 (1891). Type: P.

alopecuroides (L.) Kuntze [= Micranthus alopecuroides

(L.) Eckl.)], lectotype designated by Goldblatt & Man-

ning: 133 (2008).

Note: although Persoon’s (1805) infrageneric taxa

appear at first to be unranked, the preface (ix) to his Syn-

opsis has the following statement: Melius autein judi-

cavi, eas species (nonnullis forte tamen excipiendis) ab

aliis leviter in charactere aberrantes, imprimis si genus

minus amplum sit, sub divisione peculiari aut SUBGE-

NERE, quo etiam nonnula Botanicorum recentiiim gen-

era relata sunt, comprehendere, ne ultra necessitatem

genera multiplicentur. This is a clear statement that his

infrageneric taxa are subgenera. [We have judged it bet-

ter to include those species (with some exceptions) that

in their character(s) are only slightly different from oth-

ers, especially if the genus is not very large, under the

‘particular division’ or subgenus (which are some of

genera of recent botanists), so that genera are not multi-

plied beyond necessity.]

Deciduous geophytes. Conn axillary in origin, sub-

globose, rooting from below; tunics coarsely fibrous.

Leaves few, the lower 2 or 3 cataphylls, lowermost foli-

age leaf longest, inserted on stem above corm, blades

either plane with a definite main vein and falcate or lan-

ceolate with margins moderately to heavily thickened, or

± tubular and hollow, or terete and ± solid with heavily

thickened central vein and margins separated by narrow

longitudinal grooves. Stem erect and straight or ± flexu-

ose, simple or few- to several-branched. Inflorescence

a congested, 2-ranked spike, usually weakly rotated;

bracts short, overlapping, with leathery or dry central

portion and broad membranous margins, inner forked

apically and shorter than to ± as long as outer. Flowers

zygomorphic, lasting several days, blue to violet, mauve,

white or flushed lilac, scentless or pleasantly scented,

with nectar from septal nectaries; perianth tube short,

curving outward, ± cylindric below, flaring in upper half;

tepals ± equal, dorsal slightly larger and arching over

stamens, lower tepals extended ± horizontally. Stamens

unilateral and arcuate; filaments slender, free; anthers

oblong, held under the dorsal tepal, splitting longitudi-

nally. Ovary ovoid, sessile; style branches slender, usu-

ally deeply divided and recurved, or barely notched at

apex. Capsules woody, small, narrowly ovoid-ellipsoid

or urn-shaped with ovules in lower fourth. Seeds 2^(5)

per locLile, 3(4)-sided below, elongate, widest at micro-

pylar end with micropylar crest and micropyle above

base, tapering and pointed at chalazal end, surface

slightly wrinkled. Basic chromosome number x = 10.

The diagnostic features of Micranthus are the crowded,

2-ranked spike; small, bilaterally symmetric, tubular flow-

ers; distinctive dry outer floral bracts with broad membra-

nous margins; basal leaf inserted on the stem above the

level of the corm (shared with Lapeirousia and Thereian-

thus); and small, narrow capsules, eaeh locule containing

up to four slender seeds almost as long as the locules and

with a micropylar crest at the proximal end. Micranthus

is unique among subfamily Crocoideae in having zonasul-

cate pollen grains (Figure 2). The sulci are distal as seen

at the tetrad stage (S. Nilsson, pers. comm. Oct. 1996) and

the zonasulcate condition in Micranthus is thus derived

from the basic monosulcate grain by extension of the sul-

cus until it encircles the grain. Exine sculpturing is retic-

ulate, grading to tectate-perforate close to the aperture

margin. Among Crocoideae, only a few species of There-

ianthus also have reticulate exine sculpturing (Manning

& Goldblatt 2011). Most genera of Crocoideae, includ-

ing Thereianthus, have sulcate pollen grains, with a pair

(sometimes solitary) of narrow bands of exine (elongated

opercula) lying parallel to one another along the long axis

of the aperture. Other more complex apertures are known

in Geissorhiza Ker Gawl. (Goldblatt & Manning 2009).

Cyanixia and Zygotritonia have trisulculate grains. All

these pollen types have tectate-perforate exine with small

SLipratectal spinules.

Leaf marginal anatomy in species with plane leaves

conforms to the norm for Watsonieae in combining

unspecialized marginal epidermal cells and a marginal

vein with a sclerenchyma cap below the epidennis. This

condition prevails in Watsonia and the Lapeirousia clade

(excluding L. cojymbosa (L.) Ker Gawl. and its imme-

diate allies), but notably not in Thereianthus or Pillan-

sia, both of which lack a marginal vein or sclerenchyma

strand below the unspecialized marginal epidermis (Gold-

blatt & Manning 1990; Rudall & Goldblatt 1991; Gold-

blatt et al. 2004; Manning & Goldblatt 2011).

Chromosome cytology, the basic chromosome

number for Micranthus is .x = 10. One population each

of the four speeies counted, namely M. alopecuroides,

the new M. filifolius (reported as M. junceus), M. plan-

tagineus (as M. junceus) and M. tubulosus, are diploid,

2/7 = 20 (Goldblatt 1971; Goldblatt & Takei 1997). The

base number and karyotype, consisting of one long and

nine short chromosome pairs, are matched exactly in

Thereianthus. The related genus Watsonia has x = 9 and

a derived karyotype with two long chromosome pairs.

Pillansia, the fourth and last genus of this lineage of

Watsonieae, also has x= 10, with its single species tetra-

ploid, 2/7 = 40 (Goldblatt 1977; not 44 as originally pub-

lished by Goldblatt 1971).

Reproductive system, compatibility and pollination-.

virtually nothing has been reported about the reproduc-

tive system in Micranthus, but we infer that self-incom-

patibility and compatibility are important in the evolu-

tion and distribution of the genus. It is notable that three

species, M. alopecuroides, M. plantagineus and M. tub-

ulosus typically have all flowers produeing a full com-

plement of capsules and we infer self-compatibility and

facultative autogamy for these species. In eontrast, M.

filifolius and M. thereianthoides exhibit lower capsule

production and we infer self-incompatibility for these

species. We are unable to infer compatibility relations

Bothalia 43,2 (2013)

131

FIGURE 2. — Pollen morphology of Micranthus plantagineiis. Piket-

berg, Mannuig 2093. Scale bar: 20 pm.

for M cniciatiis and M. simplex as good fruiting mate-

rial is not available. Significantly, the putatively self-

compatible species M. alopecuroides, M. plantagineus

and M. tubulosus have the widest ranges in the genus,

with M. plantagineus occurring over the entire range

of the genus. M. filifolius has a modest range, entirely

within the Caledon District of Western Cape but M.

cniciatiis, M. simplex and M. thereianthoides are local

endemics, the latter two currently known from only one

or few populations. Self-incompatibility is believed to

be ancestral for Iridaceae (Goldblatt & Manning 2008).

In genera that we have studied, we have found rela-

tively few species to be facultatively autogamous, and

we assume such species are specialized. Reversals from

self-compatibility to incompatibility are believed to be

unlikely. Thus in Micranthus we infer that the self-com-

patible M. alopecuroides, M. plantagineus and M. tubu-

losus are derived for this character. The latter two spe-

cies are also derived in their hollow leaves as outgroup

comparison indicates that plane, isobilateral leaves are

the plesiomorphic condition (present in most members

of the family and universal in sister genus Thereian-

tlius). Reproduction through aerial cormlets that replace

flowering on the spike axis is also known only in these

three species. The arrangement of species in our account

reflects our belief that the self-incompatible species are

closer to the ancestral stock of Micranthus.

The small flowers of all species except Micranthus

thereianthoides are so similar in size, shape and colour

(perianth tube 3~5 mm long) that they almost certainly

share the same generalist pollination ecology, though we

have only recorded insect visitors for M. alopecuroides,

M. plantagineus and M. tubulosus. Goldblatt & Manning

(2006) regarded these three species as having a general-

ist pollination strategy and insect visitors to these spe-

cies include large-bodied bees (Apidae), bee-flies (Bom-

byliidae), hopbine beetles (Scarabaeidae; Hopliini) and

butterflies. Among the latter are Pieris helice (Pieridae)

(M. plantagineus) and Cynthia cardui and Colias electo

(Pieridae) (M. tubulosus). New observations confirm to

the generalist pattern with anthophorine bees and wasps

including Delta cf. caffra (Eumenidae), a species of

Sphecidae visiting M. plantagineus. The longer perianth

tube of M. thereianthoides, 22-25 mm long, suggests a

specialized pollination system using a long-proboscid

pollinator, possibly a long-proboscid fly species or large

butterfly. The nectar reward, retained in the lower part of

the perianth tube, is only accessible to pollinators with

a proboscis at least 18 mm long, thus excluding access

to smaller butterflies, bees, bee-flies and wasps that visit

flowers of other Micranthus species.

Key to the species

Note: plants with the lower part of the spike bearing

smaller, paler floral bracts subtending one or more small

cormlets may be Micranthus jiinceits or M. tubulosus or

may be hybrids involving these two species or with M.

alopecuroides and are not accommodated in the key.

la Leaf blades either plane with evident central vein or one or

two prominently thickened veins, or terete and 4-grooved

with thickened margins, not hollow; style variously divid-

ing between mouth of perianth tube to opposite middle of

anthers;

2a Lowermost leaves plane and linear to lanceolate or falcate,

(2-)4-12 mm wide; style branches divided ± halfway . .

1 . M alopecuroides

2b Lowermost leaves either plane and linear or terete and

cross-shaped in section, up to 2 mm wide; style branches

divided up to halfvvay or undivided:

3a Flowers white, outer tepals tipped palest lilac, fading

slightly darker lilac; style branches undivided or barely

notched at apex 2. M simplex

3b Flowers pale blue, blue-mauve or deep blue or white;

style branches divided up to halfway;

4a Style dividing at mouth of perianth tube opposite

middle of filaments; style branches ± 2.5 mm long,

divided less than one third; leaves at least 1.5 mm

wide 6. M cniciatiis

4b Style dividing between base and middle of anthers;

style branches usually ± 1.0 mm (up to 1.5 mm)

long, divided up to halfvvay; leaves < 1 mm wide . . .

5. M. filifolius

lb Leaf blades tubular and hollow, round or ± compressed in

section; style dividing opposite base to middle of anthers

(rarely just below anther bases):

5a Perianth tube elongate and ± twice as long as bracts, 22-25

mm long; bracts 8-15 mm long 3. M thereianthoides

5b Perianth tube shorter than bracts, < 10 mm long; bracts

5-7 mm long:

6a Flowering stem ± flexed at each aerial node; stem with

prominent, coarsely fibrous collar around base . . .

M. plantagineus x A/, tubulosus

6b Flowering stem straight, stiffly erect; stem with or with-

out collar of fibres around base:

132

Bothalia 43,2 (2013)

7a Blade of lowennost leaf terete or oval in section, ±

straight, green at flowering and smooth when fresh

(with evident thickened veins when dry); stem with-

out collar of fibres around base; capsules narrowly

ovoid to ± urn-shaped 7. M plantagineiis

7b Blade of lowermost leaf tubular and inflated, falcate

with prominent apical mucro, often dry at flower-

ing, without prominent veins when alive or dry; stem

with collar of fibres around base; capsules narrowly

ovoid 4. M. tubulosiis

1. Micranthus alopecuroides (L.) Eckl., Topo-

graphisches Verzeichniss der Pflanzensammlung von

C.F. Ecklon: 43 (1827). Gladiolus alopecuroides L.:

5 (1756). Ixia alopecuroides (L.) L.f.; 92 (1782). Ixia

plautagiuea Alton: 59 (1789), nom. illeg. superfl. pro

Gladiolus alopecuroides L. [see note 1]. Paulomagnu-

sia alopecuroides (L.) Kuntze: 702 (1891). Type: South

Africa, [Western Cape], Somerset West, 19 Nov. 1944,

Barker 3384 (NBG, neo., here designated; PRE isoneo.)

[see note 2].

Watsonia compacta Lodd.: t. 1577 (1830),

nom. nud.

FIGURE 3. — Distribution of Micranthus alopecuroides,

• ; M simplex, o; M. thereianthoides, A.

but have also been recorded on sandy ground. The Oli-

fants River Valley populations grow in thin clay or

sandy gravel, often over rocky pavement that is totally

dry even before flowering commences.

Plants mostly 200^50 mm high, base usually

sheathed with collar of short fibres. Conn mostly 10-12

mm diam., tunics of dark brown, relatively coarse, retic-

ulate fibres, drawn into short bristles above. Stem usu-

ally simple or 1- or 2(3)-branched, when unbranched

often with one or more scales below base of spike, usu-

ally bearing 1 or more cormlets in axil of lowermost

foliage leaf. Leaves 2^(5), lowermost 1 or 2 plane,

broadly to narrowly falcate (occasionally ± lanceolate)

or linear, (2-)5-10(-l 5) mm wide, with moderately

prominent main vein; margins slightly or occasionally

heavily thickened {De Vos 2288), hyaline when dry;

upper I or 2(3) leaves largely sheathing. Spike mostly

40-80-flowered, often much congested, with inter-

nodes 1. 5-3.0 mm long; bracts 5-7 mm long, outer with

broad to narrow brown centre and translucent mem-

branous margins, inner ± as long as outer, notched api-

cally, translucent with 2 dark veins slightly broader

toward base; lower or all nodes sometimes vegetative

and then bracts paler in colour and subtending one (or

more) cormlets in each axil. Flowers usually dark blue,

sometimes pale blue, often lower third to fourth of tepals

paler blue or white, distally edged with a thin darker

blue line, unscented; perianth tube ± 5 mm long; tepals

subequal, elliptic, 7-8 x ± 3 mm, with short narrow,

claw-like base. Stamens with filaments ± 5 mm long,

diverging in upper half; anthers oblong, 3^ mm long,

pale mauve; pollen white to pale blue. Style ± 7 mm

long, mostly dividing between upper third of filaments

and lower third of anthers; branches ± 1 .2-1 .6 mm long,

divided for ± half their length. Capsules oblong to nar-

rowly ovoid, ± 5 mm long but ± 4 mm long when dry.

Seeds angular-elongate, 3. 5-4.0 mm long, 3 or 4 per loc-

ule. Flowering time: October in the north, November to

December in the south.

Distribution: centred in the southwestern Western

Cape, Micranthus alopecuroides has a relatively narrow

range, extending from the Cape Peninsula north into the

Olifants River Valley and east locally to Flermanus and

Swellendam (Figure 3). Plants typically grow on well-

drained clay or loamy, seasonally wet, slopes and fiats

Diagnosis: Micranthus alopecuroides is distinctive

in its plane (Figure lA), sometimes very broad basal

leaves; the blades lanceolate to linear or falcate, some-

times up to 12 mm wide or exceptionally to 15 mm in

plants from the Roman’s River area of the upper Breede

River Valley. Exceptions are numerous and there are col-

leetions with ± linear leaves 3-5 mm wide and only up

to 25 mm long (notably Purcell s.n., NBG). The flowers

are typical of the genus, usually dark blue, with a peri-

anth tube ± 5 mm long. The spikes of 40 or more flowers

are often unusually congested with the internodes ± 1.5

mm long.

Populations from the Olifants River Valley and flow-

ering in October, at least three weeks earlier than else-

where, stand out in their relatively lax spikes with inter-

nodes 2. 5-3.0 mm long (vs. ± 1. 5-2.0 mm elsewhere)

and relatively short, straight leaves, 5-8 mm wide with

partieularly prominent mucronate tips. The outer bracts

of these plants also differ from those in populations to

the south in their broader translucent margins, thus with

a significantly narrower central band of green tissue

(brown when dry). These plants are typically restricted

to thin clay or light sandy soils over rocky pavement and

represent a distinctive race of the species.

Hybrids: certain collections from sandy flats south

of Malmesbury constitute a puzzle. They consist of

plants with abnormally elongated spikes up to 120 mm

long, more than 3/4 of their length bearing small, pale

bracts each enclosing not a flower but a small cormlet.

Only the top fourth of the spikes have properly formed,

dark brown bracts subtending either pale blue flow-

ers (e.g. Goldhlatt & Manning 10431, MO, NBG, with

tubular leaves) or deep blue flowers {Goldhlatt & Man-

ning 10432, MO, NBG, with plane leaves). The leaves,

either tubular or plane with a central vein, correspond to

Micranthus tuhulosus or M. alopecuroides respectively.

We conclude that these plants constitute hybrids or a

hybrid swarm with M. alopecuroides as one parent and

M. tuhulosus or possibly M. plantagineiis as the other.

Microscopic examination of the pollen shows some

apparently normal grains and others smaller than normal

Bothalia 43,2 (2013)

133

and evidently sterile. The available collections of this

putative hybrid were made too early in the life cycle to

have capsules, which if developed, would have appeared

later in the season.

Similar specimens from other sites show the same

striking feature (e.g. Goldblatt 8711 MO, from Greyton;

Leighton 722 BOL from Camp Ground, Rondebosch;

and Williams 1195 MO, NBG from near Vogelgat, Her-

manns). This last collection consists of plants with the

sterile part of the spikes 180-250 mm long and the fer-

tile part 20-30 mm long. Populations at Elandsberg near

Bo-Hermon, with plane, narrowly lanceolate leaves (e.g.

Goldblatt & Manning 13616) have the inflorescence

sterile throughout and bearing a cormlet in all bract axils

as do many individuals of the species from Rondebosch

Common, Cape Town (Goldblatt & Manning 13619).

The status of these plants is uncertain but we suspect

them to have a hybrid origin.

[Note 1 . Daniel Solander, the unacknowledged author

of Hortus kewensis published under William Aiton’s

name (1789), described the new species Ixia plantag-

inea [folds linearibus strictis, spica disticha imbricata',

based on a collection of Francis Masson and, for rea-

sons that are obscure, at the same time cited Linnaeus’s

Gladiolus alopecuroides in synonymy. The epithet plan-

taginea, alluding to the similarity of the inflorescence

to that of Plantago L., is no more apt than Linnaeus’s

recalling the resemblance to the grass, Alopecurus L.,

and constitutes an illegitimate superfluous name. Never-

theless, the broad-leaved species remained known by the

later epithet plant agineits, until well into the 20th cen-

tury (e.g. Lewis 1950) despite the leaves being described

as narrow and linear in the protologue. The epithet plan-

tagineus was applied to M. alopecuroides by, among

others, Ker Gawler (1803), who evidently did not realize

it applied to two different species. Baker (1892, 1896),

who also used the name M. plantagineus for M. alo-

pecuroides, compounded this error and recognized M.

plantagineus var. junceus not realizing that the type of

the species was in fact identical with his new variety.]

[Note 2. Described in 1756 by Linnaeus as Gladiolus

alopecuroides, with the brief diagnosis [folds linearibus,

spica disticha imbricata, ^ the species was transferred to

Micranthus by Ecklon (1827), when he raised Persoon’s

Gladiolus subg. Micranthus to generic rank. Of the three

sheets identified as G. alopecuroides in the Linnaean

herbarium, one [LINN 59.13] is a Sparrman collection

post-dating the protologue, and the other two cannot be

unambiguously related to the name. One [LINN 59.15]

is M. tubulosus and the other [LINN 59.14] may be M.

alopecuroides but is atypical in its large size, numer-

ous branches and particularly broad leaves that hardly

accord with the protologue [leaves linear]. We prefer

to choose a neotype: Barker 3384, which has relatively

narrow leaves and conforms exactly to the protologue.

This action unambiguously preserves the current appli-

cation of the name to the plane-leaved species (Lewis

1950; Goldblatt & Manning 2000).]

[Note 3. Gladiolus minutiflorus Schrank (1822) has

been associated with Micranthus alopecuroides, which

Schrank also recognized (as Gladiolus), but the descrip-

tion is vague (flowers small, secund, tepals subequal)

and we are unable to determine the plant to genus with

confidence, let alone to species. Schrank did, however,

explicitly describe the leaves as short, striate and narrow,

the lower ± 5 inches ( 125 mm) long. No authentic mate-

rial has been located either at the Munich (M) or Brus-

sels (BR) Herbarium, the institutions where the types of

Schrank’s species, where they exist, are believed to be

located.]

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): clay hillside S of

Algeria turnoff on Clanwilliam-Citrusdal road (N7), (-BD), 13 Oct.

1974, Goldblatt 3030 (MO). 3220 (Wuppertal): near Citrusdal on

old Clanwilliam road, (-CA), 11 Oct. 1984, Bean & Viviers 1504

(BOL); N of Citrusdal, (-CA), 16 Oct. 1935, Taylor 1224 (BOL);

clay slope near Farm Klaweivlei on road to Algeria, (-CA), 11 Oct.

2011, Goldblatt & Porter 13864 (MO, NBG, PRE). 3318 (Cape

Town): Cape Peninsula, Wynberg Hill, (-CD), Nov. 1950, Pillans

10208 (BOL, MO); Wynberg Hill, Edinburgh Drive, (-CD), 25 Jan.

2011 (fr.), Goldblatt & Manning 13631 (MO, NBG); fields near Cape

Town, (-CD), Aug.-Nov., H. Bolus 2829 (BOL); Devil’s Peak above

Vredehoek, clay slopes, (-CD), 30 Oct. 1982, Goldblatt 6637 (MO);

slopes of Lions Head, (-CD), 20 Nov. 1938, Penfold s.n. SAM53I59

(SAM); Jonkershoek, Bosboukloof, (-DD), 27 Nov. 1973, Smith 140

(NBG); Jonkershoek Valley, (-DD), 27 Nov. 1975, Kruger 84 (NBG).

3319 (Worcester): Grootwinterhoek, (-AA), without date, Pappe s.n.

SAM21101 (SAM); Mostertshoek, (-AC), 8 Dec. 1973, De Vos 2288

(NBG); slopes at Wabooms R., foot of Waaihoek Peak, (-AD), 11

Dec. 1948. Esterhuysen 14822 (BOL); Bo-Hermon, Elandsberg Nature

Reserve, entrance to Bosplaas, (-AC), 22 Jan. 2011 (fr. and sterile);

Goldblatt & Manning 13616 (MO, NBG); Wemmershoek, (-CC),

2 Nov. 1947, Barker 4903 (BOL, NBG). 3418 (Simonstown): Cape

Peninsula. Bergvliet Farm, E of sandpit, (-AB), 22 Nov. 1918, 5 Dec.

1818, Purcell s.n. (NBG); Helderberg, Somerset West, (-BB), 2 Dec.

1944, Parte;- 3959 (BOL, NBG). 3419 (Caledon): Elgin Basin, Ariesk-

raal, well drained clay ground, (-AA), 5 Dec. 1994, Rode (& Boucher

0207 (NBG); Elgin, (-AA), 19 Nov. 1944, Barker 3369 (NBG). 3420

(Bredasdorp): Swellendam, Bontebok Park, (-AB), 2 Nov. 1965,

Grobler 552 (NBG).

2. Micranthus simplex Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape, 3218 (Clanwilliam): Piket-

berg, southwestern slopes of Zebrakop, (-DB), shallow

soil on sandstone pavement, 4 Jan. 1995, Goldblatt &

Manning 10172 (NBG, holo; K, MO, PRE, iso.).

Plants ( 100-) 140-200 mm high, base sheathed by

short collar of brittle fibres. Conn tunics of dark brown,

reticulate fibres. Stem usually simple, rarely 1 -branched.

Leaves (2)3, plane, linear or falcate, ± 1 mm wide, usu-

ally with 1 or 2 prominent veins, margins thickened,

hyaline when dry. Spike 16- to 40-flowered; bracts pur-

ple-brown with broad translucent, brown-flecked mem-

branous margins, ± 5 mm long, inner bracts ± as long as

outer, membranous with 2 dark keels, notched at apex.

Flowers white fading to lilac, outer tepals tipped pale

lilac, with subapical brown ridge on reverse; perianth

tube ± 3 mm long; tepals oblong, ± 4 x 1.2 mm. Stamens

with filaments ± 2.5 mm long; anthers oblong, ± 2.5 mm

long. Style ± 7 mm long, dividing opposite middle of

anthers; branches ± 1 mm long, barely notched at apex.

Capsules oblong, slightly warty in distal half, ± 4 mm

long. Seeds elongate-angular, ± 3 mm long. Flowering

time: December to at least mid-January.

Distribution', known only from the slopes of Zebra-

kop, highest peak in the Piketberg, Micranthus simplex,

like M. cruciatus, grows in shallow soils in moss or in

rock crevices on wet sandstone rocks (Figure 3). The

134

Bothalia 43,2 (2013)

habitat remains moist as late as January when the spe-

cies blooms.

Diagnosis: unusually small for the genus, stems of

Micranthiis simplex rarely exceed 180 mm and the white

flowers with lilac-tipped outer tepals are distinctive,

other species having flowers in shades of deep to pale

blue or blue-mauve, or occasionally white. The inflo-

rescence has the appearance of being relatively lax, the

bracts of the lower flowers of the spike not overlapping

those above them, but the upper bracts are as closely set

as in other species. It is one of two species of Micran-

thus with consistently plane leaves; the other, M. alo-

peciiroides, is a taller plant with congested spikes of 40

to 80 flowers and broader leaves mostly 5-12 mm wide.

The flowers of M. simplex are the smallest in the genus,

the perianth tube just 3 mm long and the short anthers

± 2.5 mm long. The short, undivided style branches,

± 1 mm long, are likewise unusual for Micrantlms, other

species of which normally have the style branches some-

what to considerably longer and divided for at least one

third their length.

Additional specimens

WESTERN CAPE. — 3218 (Clanwilliam): Piketberg, Zebrakop, (-

DB), in moist sand, 800 m, 3 Jan. 1973, Linder 193 (BOL).

3. Micranthus thereianthoides Goldblatt &

J.C. Manning, sp. nov.

TYPE. — Western Cape, 3318 (Cape Town): Paarde-

berg, Vondeling, (-DB), rock cracks and sands along

stream, 9 Jan. 2013, Nicolson 995 (NBG, holo.; K, MO,

iso.).

Plants 300-800(-1200) mm high, base weakly

sheathed by fine fibres. Conn 10-15 mm diam., tunics of

fine to moderately coarse, dark brown, reticulate fibres.

Stem simple or rarely branched, with solitary cormlet in

axil of second leaf and sometimes also third leaf. Leaves

(4)5 or 6, green or drying at flowering, lowermost 2 or

3 longest, blades 100-300(-800) mm long, 2. 5-5.0

(-15) mm diam., tubular and hollow, sometimes inflated,

acute-mucronate, upper leaves progressively shorter and

narrower, uppermost bract-like and entirely sheathing.

Spike 10- to 40(-70)-flowered, bracts brown with broad

translucent membranous margins, 8-ll(-15) mm long,

as long as 1 .5-2.0 spike internodes, inner bracts slightly

shorter than outer, forked apically, membranous with 2

dark keels broadened toward base. Flowers suberect,

dark violet or purple, unscented; perianth tube ± cylin-

dric, 22-25 mm long, tepals oblong, 5-6 x 1.5-2. 5 mm,

reverse of outer tepals with prominent subapical ridge.

Stamens with filaments 6-8 mm long, exserted ± 3 mm;

anthers oblong, 3. 5-4.0 mm long. Style 24-27 mm long,

dividing between middle and slightly beyond anthers,

branches ± l.5(-2.0) mm long, divided for ± half their

length. Capsules ovoid, smooth, 5-6 mm long, with ±

4 seeds per locule. Seeds elongate-angular, tapering to

points at both ends, ± 3.5 mm long. Flowering time: Jan-

uary. Figure 4.

Distribution: a highly local endemic, Micranthus

thereianthoides is restricted to the Paardeberg near

Malmesbury (Figure 3), where it grows at mid to upper

altitudes along the banks of seasonal streams, the conns

usually wedged among granite rocks, sometimes in

humic loam, where the plants are more robust. Plants

are locally plentiful along several streams on the range.

The long-tubed, violet flowers are evidently adapted to

pollination by long-proboscid flies. The incomplete fruit

set in wild plants suggests that M. thereianthoides is an

obligate outcrosser.

Diagnosis: Micranthus thereianthoides closely resem-

bles M. plantagineus and some forms of M. tubulosus

in its cylindrical leaves but is unique in the genus in the

relatively large floral bracts, 8-11 mm long, and most

strikingly in its dark violet flowers with elongate, cylin-

drical perianth tube 22-25 mm long, thus ± twice as

long as the bracts (Figure 4). The species appears never

to develop cormlets in the floral bract axils.

The long-tubed flowers suggest the genus Thereian-

thus, but the hollow leaves, the small, obtuse tepals, and

the bracts with broad, membranous margins are charac-

teristic for Micranthus. The zonasulcate pollen grains

with reticulate exine conform exactly to those of other

species of Micranthus, leaving no doubt as to its generic

placement.

This extraordinary species was discovered in Janu-

ary 2012 by local plant enthusiasts Greg Nicholson and

Dewan Roets during a botanical survey of the Paarde-

berg.

Additional specimens

WESTERN CAPE. — 3318 (Cape Town): Paardeberg, between

Wellington and Malmesbury, Paardeberg Nature Reserve next to

Malmesbury Dam, (-DB), rocky crevices near water, 10 Jan. 2012,

Nicolson & Roets 788 (NBG); Vondeling, (-DB), Feb. 2012 (fruiting),

Nicolson 994 (MO, NBG).

4. Micranthus tubulosus (Burm.f.) N.E.Br. in Bul-

letin of Miscellaneous Information, Royal Botanic

Gardens, Kew 1929: 133 (1929). Cladiolus tubulosus

Burm.f: 2 (1768). Ixia cepacea Basseporte ex DC. in

Redoute: t. 96 ( 1804), nom. nov. in Ixia, non /. tubulosa

Burm. f (= Babiana tubulosa (Burm.f) Ker Gawk).

Type: South Africa, without precise locality or collector

(G: Herb. Burman, holo., image seen).

Cladiolus fistulosus Jacq.: 8 (1797). Ixia fis-

tulosa (Jacq.) Sims: t. 523 (1801), horn, illegit. non

Andrews (1799) [= Hesperantha radiata (L.f) Ker

Gawl.]. Micranthus fistulosus (Jacq.) Eckl. ex Baker:

179 (1892), nom. superfl. pro C. tubulosus Burm.f

Type: South Africa, without precise locality or collector,

illustration in Jacq.: t. 16 (1797), left hand plant, lecto-

type designated here.

Watsonia spicata Sol. ex Ker Gawl.: sub t.

553 (1803). nom. superfl. pro C. spicatus L. (1753), C.

tubulosus Burm.f (1768) et C. fistulosus Jacq. (1797).

[The citation Watsonia spicata (L.) Ker Gawl. in Annals

of Botany (Konig & Sims) 1: 229 (1804) is an error].

Type: South Africa, without precise locality, illustration

in Curtis’s Botanical Magazine 15: t. 523 dxia fistulosa'

(1801).

Ixia teretifolia Herb. Banks ex Sims: t. 523

(1801), nom. nud. pro syn.

Bothalia 43,2 (2013)

135

FIGURE 4. — Micraruhiis thereianthoides. Paardeberg, Nicolson 995 (NBG). A, flowering plant; B, flower; C, outer (left) and inner (right) bract; D,

capsule; E, seed. Scale bar: A, 1 0 mm; B-D, 2.5 mm; E, 1 .25 mm. Artist: John Manning.

136

Bothalia 43,2 (2013)

Micranthiis fistulosus Eckl.: 44 (1827), nom.

nud. [Probably intended as a combination but basionym

not cited.]

Plants (70-)150-350(-600) mm high, base sheathed

by collar of short, stiff, bristly fibres. Conn 14-18 mm

diam., tunics of coarse, dark brown, reticulate fibres.

Stem simple or branched, occasionally with cormlets

in axil of lowermost leaf and of uppermost cataphyll.

Leaves (2)3-5, usually dry at flowering, lowermost

1 or 2 longest, blades 50-200 mm long (to 300 mm in

sterile plants), 4-7 mm diam., inflated, tubular and hol-

low, apex obtuse-mucronate, upper leaves progressively

shorter and narrower. Spike 16- to 40-flowered; bracts

mid to dark brown, 5-6 mm long, outer with broad

translucent membranous margins, inner bracts ± as long

as outer, forked apically, membranous with 2 dark keels

broadened toward base; lower nodes sometimes veg-

etative with one or more cormlets and bracts then pale.

Flowers pale or dark blue or white, sweetly scented;

perianth tube 5-6 mm long; tepals oblong, 5-6(-10) x

1.2-2. 5 mm, reverse of outer tepals with prominent

subapical ridge. Stamens with filaments 6-8 mm long;

anthers oblong, 3. 5-4.0 mm long. Style 7-8 mm long,

dividing opposite or slightly below base of anthers,

branches ± 1.5(-2.0) mm long, divided for ± half their

length. Capsules ovoid, smooth, 4-5 mm long, with 3

or 4 seeds per locule. Seeds elongate-angular, tapering

to points at both ends, ± 3 mm long. Flowering time:

November to December.

Distribution', typically a species of lower slopes usu-

ally on clay and granite-derived soils but also on sand-

stone, Micranthiis tubulosus is restricted to the west-

ern half of Western Cape. It extends north of the Cape

Peninsula as far as the northern Cedarberg, where an

early (1923) collection documents its occurrence at

Heuningvlei, and no further east of the Peninsula than

Suurbraak near Swellendam and the Agulhas Peninsula

(Figure 5). Like other species of the genus, it blooms

late in the season when the hollow, inflated leaves are

often dry and brown. Plants from the Pakhuis Mtns

growing in moist, sandy ground are exceptional in

their small size (leaves up to 100 mm long) and require

additional study. A much dwarfed fragment of Micran-

tlnis tubulosus, said to be from Caries (Caporn s.n., ex

hort. Kirstenbosch (as Nat Bot Card. 915/15) in BOL)

is unlikely to be from there as no other records of the

genus from Namaqualand exist.

Diagnosis', the inflated, tubular, falcate leaves (Figure

1 E) are diagnostic for the species, the spikes and flow-

ers of which differ hardly at all from those of Micran-

thiis alopecuroides. A particularly distinctive feature of

the leaves is the prominent brown mucro at the obtuse to

± truncate apices. The leaves are often ± dry at flower-

ing time — Marloth’s (1915: plate 41) has a particularly

apt illustration of the species. As in M. planiagineus, one

or more cormlets may be produced in the lower axils of

the spike, a phenomenon first noted by Ker Gawler (in

Sims 1801) and later confirmed by Lewis (1950). The

condition is more frequent, although not consistent, in

M. planiagineus. Despite its apparently preferred habitat

on relatively dry slopes, M. tubulosus can occasionally

be found on moist sandy flats, sometimes co-occurring

with M alopecuroides and M. planiagineus (Wurts 519

ly 18° 20° 22° 24° 26° 28“

FIGURE 5. — Distribution of Micranthiis tubulosus.

consists of just such a mixture, M. tubulosus and M.

planiagineus evidently found growing in close proxim-

ity). Hybrids between these two species at shared sites

blur their usually clear foliar differences. We discuss

putative hybrids between Micranthiis tubulosus and M.

alopecuroides or M. planiagineus in more detail below.

Plants collected near Saron (e.g. Schlechter 10618)

are unusually small, mostly 100-150 mm but some just

70 mm tall, and have shorter, fewer-flowered spikes than

usual. They appear linked to taller, more robust speci-

mens by a range of intennediates. In contrast, plants

from Gouda {Barker 9861), nearby, are exceptionally

robust, up to 600 mm tall, with leaves almost as long,

and the white flowers have tepals 1 0 mm long, the outer

2.5 mm wide.

A curious feature of Micranthiis tubulosus is that

populations may consist of a mixture of some plants

with entirely fertile spikes and others with the lower part

of the spike sterile (e.g. Ecklon & Zeyher Irid 192 and

190). All three specimens of Goldblatt 8711 and several

of Purcell 43 have spikes sterile in the lower half Par-

ticularly short leaves in Goldblatt 8711 are also puzzling

but not unique.

History'. Long known as Micranthiis fistulosus (Jacq.)

Eckl. (e.g. Baker 1896), based on Gladiolus fistulosus

Jacq. (1797), that combination was in fact not valid,

though it was used as M. fistulosus Eckl. (a nomen

nudum assumed to be a valid combination) by Baker

(1892, 1896). By citing the basionym. Baker’s use of

the name M. fistulosus becomes a valid (albeit unin-

tended) combination, also superfluous through his citing

of valid earlier synonyms, including Gladiolus tubulo-

sus Burm.f (1768). Jacquin’s illustration of G. fistulosus

has two plants: we designate as lectotype the left hand

one, which has dark blue flowers and leaves typical of

M. tubulosus. The right hand plant, which has pale blue

flowers and the upper leaf more typical of M. plantag-

ineiis, may be a hybrid with that species. The sterile

lower nodes of the spike, bearing silvery bracts, are

more typical of M. plantagineiis and represent at least a

different genotype from the right hand plant.

Brown (1929) identified the type of Gladiolus tubu-

losus among specimens in Burman’s herbarium, and

realizing that it was an earlier name for M. fistulosus,

provided the combination M. tubulosus. A fine il lustra-

Bothalia 43,2 (2013)

137

tion of the species in Redoute’s Les Liliacees (1804),

as Ixia cepacea, a name coined by the artist, Madeleine

Fran9oise Basseporte, shows that M tubidosus was cul-

tivated in France in the late 18th and early 19th cen-

turies. The Basseporte painting, which indeed repre-

sents M tiibiilosiis, is part of an unpublished collection

of velins (paintings on parchment), which document

plants and animals in the Jardin Royal in Paris and the

Menagerie Royale in Versailles, now in the Bibliotheque

Centrale of the Museum National d’ Histoire Naturelle

in Paris. We treat the name as having been validated

by De Candolle in 1804 in the Redoute volume. De

Candolle’s citation of the earlier Gladiolus tubulosiis

Burm.f. appears to render his epithet superfluous, but

the name Ixia tubulosa Burm.f (now Babiana tubulosa

(Burm.f) Ker Gawl.) prevents transfer of Gladiolus tub-

ulosus to Ixia. Ixia cepacea must be regarded as legiti-

mate and a new name in Ixia for G. tubulosiis.

Hybrids: the following interspecific hybrids are

known involving Micranthiis tubulosiis: M. tubulosiis x

M. alopeciiroides and M. tubulosiis x M. plantagineus.

Those with M. plantagineus can form large populations

locally, probably backcrossing with one other parental

species. We discuss these separately at the end of the

species account.

Additional specimens

WESTERN CAPE. — 3218 (Clanwilliam): Goedverwacht, Piket-

berg, (-DC), 23 Nov. 1982, Koutnik 1037 (BOL, MO); Piketberg,

top of Versfeld Pass, (-DC), 2 Nov. 2011 (in bud), Goldbiatt & Por-

ter 13707 (MO, NBG). 3219 (Wuppertal): Cedarberg. Heuningvlei,

sandy vlakte, (-AA), 23 Oct. 1923, Pocock 586 (NBG); Olifants River

near Villa Brakfontein [Citrusdal], (-?CA), Nov., Ecklon & Zeyher

Irid 190 (MO, SAM); Elandskloof, bridge ± 10 miles [± 16 km] SE

of Citrusdal, (-CA), 21 Sept. 1952 (sterile), Maguire 1832 (NBG).

3318 (Cape Town): near Hopefield, (-AB), 19 Oct. 1932, Lavis s.n.

(BOL); Malmesbury, clay slope, (-BC), 3 Nov. 1986, Goldbiatt 8048

(MO); Cape Town, Camps Bay, (-CD), Barker 7191 (NBG); Signal

Hill, (-CD), Nov. 1939, Lewis 707 (SAM); Rondebosch Common, dry,

hard ground, (-CD), 24 Jan. 2011, Goldbiatt <& Manning 13620 (MO,

NBG); Wynberg Hill, (-CD), Nov. 1922, L. Bolus s.n. (BOL17188):

Langverwacht, Kuils River, main kloof, (-DC), 14 Dec. 1973. Oliver

4820 (NBG); Stellenbosch Mtn, S of Paradys Kloof, (-DD), 3 Dec.

1989. Buys 132 (NBG); Muldersvlei, (-DD). Nov. 1916, Diithie 352

(BOL). 3319 (Worcester): near Saron, 800’ [244 m], (-AC), Oct.

1896. Schlechter 10618 (MO); Elandsberg Estate, Vangkraal road near

Mountain road, (-AC), 22 Jan. 2011 (fr.), Goldbiatt dc^Manning 13610

(MO); Gouda, (-AC), 6 Dec. 1962 (white flowers, very robust). Barker

9861 (NBG); Worcester [District], Waterfall, (-CC), Nov., Ecklon &

Zeyher Irid 192 (MO); Ceres, Schurfdeberg. lower slopes, (-AD),

Dec. 1944, Leivis 863 (SAM). 3320 (Montagu): Swellendam, hill

below Eleven O’Clock Mtn, (-CD), 25 Nov. 1952, IVurts 519 (mixed

with M. plantagineus) (NBG). 3418 (Simonstown): Bergvliet Farm,

flats near sand pit, (-AB), Nov. 1915. Purcell 43 (SAM); Helderberg

Nature Reserve, (pale blue or white), (-BB), 23 Dec. 1993, Runnals

647 (NBG). 3419 (Caledon): Greyton Nature Reserve, dry flats, 1000'

[305 m], 5 Dec. 1987, Goldbiatt 8711 (MO). 3420 (Bredasdorp): Suur-

braak, Middelplaas, (-BA), 5 Dec. 1982 (sterile), Viviers 274 (NBG);

Struisbaai to Elim near Springfontein turnoff, hard sandy gravel, (-

DB). 9 Nov. 2011 (in bud), Goldbiatt & Porter 13738 (growing with

M. plantagineus) (MO, NBG); 4 km W of Elim (growing with M. fiU-

folius), (-DA), 9 Nov. 2011, Goldbiatt & Porter 13738 (MO, NBG).

Unknown locality: Leeufontein, burned veld, 28 Nov. 1908, Pearson

3185 (BOL).

5. Micranthus filifolius Goldbiatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape, 3419 (Caledon): Akkedisberg

Pass, sandy hillside, (-AC), 18 Nov. 2011, Goldbiatt &

Porter 13370 (NBG, holo.; MO, PRE, iso.).

Plants 180-300 cm high, base sheathed with sparse to

well-developed collar of fibres. Corm 12-16 mm diam.,

tunics of relatively soft, fine or thicker fibres. Stem

unbranched or rarely with single short branch, without

cormlets in leaf axils. Leaves 4-6, green or beginning

to dry from tips at flowering time, lowermost longest,

reaching to middle of spike to shortly exceeding it, blade

either ± terete and ± 1 mm diam. or ± plane and ± 2 mm

wide, with heavily thickened central vein and margins,

separated when dry by narrow longitudinal grooves,

upper leaves shorter, with sheaths overlapping, upper-

most 1 or 2 leaves sheathing for most of their length,

with short free tips. Spike mostly 1 8-50-flowered,

closely congested, lower bracts always subtending flow-

ers; bracts mid to dark brown, ± 5 mm long, outer with

broad translucent membranous margins, inner slightly

shorter than outer, with 2 dark keels broadened toward

base, notched apically. Flowers pale mauve or mid-blue,

unscented; perianth tube ± 5 mm long; tepals oblong, ±

5 X 2. 2-2. 8 mm. Stamens with filaments ± 5 mm long,

exserted ± 2.5 mm; anthers oblong, ± 3 mm long. Style

± 7 mm long, dividing between base and middle of

anthers; branches 1.0- 1.6 mm long, divided for up to

half their length, rarely only notched at apex. Capsules

narrowly ovoid, smooth, 4-6 x ± 2 mm, with up to 4

seeds per locule. Seeds elongate-angular, mostly 3-sided,

tapering to points at both ends, 3-5 mm long. Flowering

time: mid-November to late February. Figure 6.

Distribution: centred in the Caledon District of West-

ern Cape, Micranthus filifolius is largely coastal with

populations recorded from Steenbras and Cape Hang-

klip eastward to Hermanns and inland to Shaw’s Mtns,

the lower slopes of Caledon Swartberg and east to Akke-

disberg Pass and Elim (Figure 7). Collections are mostly

from clay and clay-loam soils, occasionally from sandy

sites, but even collections from the Klein River Mtns

above Hermanns at elevations of up to 400 m are from

a shale band. The species is particularly abundant after

fire (e.g. Drewe 495, HOI) but will flower in unbumed

veld unless shaded out by taller vegetation. Plants bloom

unusually late in the season, with most flowering collec-

tions made after mid-January, and two (Gillett 520', Lev-

yns II269) were in mid- to late February. We have con-

firmed late flowering at near-coastal sites ourselves but

inland populations, as from Drayton Siding, east of Cal-

edon, and Akkedisberg Pass, flower from mid-November

to early January and are in fruit before any coastal popu-

lations come into flower. We suggest that this early flow-

ering is due to warmer and drier conditions well inland

of the coast. The coarser corm tunic fibres and collar of

fibres around the base of the stems in these populations

are perhaps adaptations to the drier habitat. Plants some-

times co-occur with or grow close to M. plantagineus,

which is in fruit when M. filifolius begins to bloom, two

or three weeks after the last flowers of M. plantagineus

have faded, both at the coast and at inland sites. We have

also found M. filifolius growing together with M. tubu-

losiis.

Diagnosis: with its narrow leaves, the lowennost

of which is linear or terete (Figure IB & C), Micran-

thus filifolius is most like M. plantagineus in general

appearance. It differs, however, from that species in sev-

eral respects, particularly in the solid leaf blades of the

138

Bothalia 43,2 (2013)

FIGURE h.-Micnmihus filifolius, Akkedisberg Pass, GolcIhlaU & Porter 13370 (NBG), A, flowering plant; B, flower; C, outer (lower) and inner

(upper) bract. Seale bar; A, 10 mm; B, C, 1 .25 mm. Artist; .lohn Manning.

Bothalia 43,2 (2013)

139

lowermost and sometimes the upper leaves. The leaves

closely overlap one another and sheath the stem up to

the base of the spike. Plants typically have four or five

leaves but several specimens have six leaves, the upper

one or two largely to entirely sheathing. Unlike M plan-

tagineiis, plants do not produce cormlets in the leaf axils

and the corm tunics are usually soft-textured with the

base enclosed by a collar of fibres, whereas M. plan-

tagineus has coarser corm tunics, lacks a basal collar

of fibres, and cormlet production is conspicuous in the

lowermost and sometimes other leaf axils. In addition,

the flowers, although typical of Micranthus, are some-

what smaller than in M plantagineus, having a perianth

tube mostly 4-5 mm long and tepals ± 4 mm long, and

are more often pale mauve (drying white), although the

western populations are dark blue (drying blue), the

flower colour in most populations of M. plantagineus.

Typical M. plantagineus has been recorded close to most

localities of M filifolius, flowering in November and

December, supporting our conclusion that M. filifolius is

not a local variant but a different species, flowering later,

sometimes in the same habitats as M. plantagineus, or in

drier sites.

Additional specimens

WESTERN CAPE. — 3418 (Simonstown): Steenbras, (-BB), Sept.

1944 (late fr.), Stokoe s.n. S.4M68012 (SAM); Buffels River dam area

[near Rooiels], (-BD), 19 Feb. 1972, Boucher 1822 (NBG. PRE);

Cape Hangklip, peaty marsh, (-BD), Le\yns 10220 (BOL); Kogelberg

Nature Reserve, (-BD), 19 Mar. 1983, Kroon 10200 (PRE); Betty's

Bay, sandy slopes, (-BD), 13 Feb. 1962, Le\yns 11269 (BOL). 3419

(Caledon): Drayton Siding, pale blue (-AB), 16 Dec. 1968, Goldblatt

395 (BOL); field E of Drayton siding, (-BA), 25 Jan. 2011, Gold-

blatt & Manning 13623 (MO, NBG); Kleinmond, near Palmiet River

mouth, (-AC), 31 Jan. 1933, Gillen 615 (NBG); Kleinmond, road to

reservoir, (-AC), 27 Jan, 1947, De Vos 485 (NBG); Hermanns, (-AC),

Jan. 1920, Burn Davy 18711 (BOL); top of Shaw's Pass, (-AD), Jan.

1957, Lewis 2904 (SAM); Shaw's Pass, east side, (-AD), 29 Dec.

1955, Lewis 4454 (SAM); Hemel-en-Aarde, mountain side, (-AD),

15 Jan. 1933, Gillen 520 (NBG); Vogelgat, Hermanns, Vogelpool to

Femkloof, S slopes on shale band, (-AD), 2 Jan. 1979, Williams 2710

(NBG); Femkloof, Hermanns, 350 m, clay area. 1 year after fire, (-

AD), 17 Jan. 1987, Drewe 495 (MO); 400 m, shale band, after fire, 25

Jan. 1996, Drewe 1101 (MO); 4 km W of Elim. stony clay in renos-

terveld, (growing with M. tubulosus), (-DA), 9 Nov. 2011 (in bud),

Goldblatt & Porter 13745 (MO, NBG).

6. Micranthus cruciatus Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape, 3219 (Wuppertal): Pakhuis

Mts, trail to Heuningvlei, (-AA), local in wet seep on

rocky sandstone slope, 19 Dec. 1995, Goldblatt 10438

(NBG, holo.; MO, iso.).

Plants 300-450 mm high. Corm globose, 8-10 mm

diam., tunics of brown, soft membranous layers not

accumulating. Stem simple or 1 -branched, usually with

1 or 2 small cormlets in lowermost leaf axil. Leaves

(3)4 or 5, lower 3 linear to subterete, ± 1.5 mm wide,

margins and midrib heavily thickened with narrow lon-

gitudinal grooves between (often cross-shaped in sec-

tion with 4 narrow longitudinal grooves), reaching to

base or middle of spike, uppermost leaf sheathing stem

almost to base of spike, with short free portion. Spike

up to 70-flowered; bracts brown or straw-coloured with

broad translucent membranous margins, ± 4 mm long,

inner ± as long as outer, membranous with 2 dark keels,

apically notched. Flowers pale blue-mauve (drying ±

16” 18” 20' 22' 24” 26' 28'

FIGURE 7. — Distribution ol Micranthus filifolius, •; M. cruciatus, o.

white) or deep blue, perianth tube ± 3 mm long, tepals

oblong, ± 4 X 1.5 mm. Stamens with filaments ± 5 mm

long; anthers oblong-linear, ± 3 mm long. Style ± 4 mm

long, dividing ± at mouth of tube opposite middle of

filaments, branches ± 2.5 mm long, divided for ± one

third their length. Capsules oblong, smooth, 5. 0-5. 5 mm

long. Seeds elongate-angular, mostly 3-sided, tapering to

points at both ends, ± 3 mm long. Flowering time: mid-

November to late December, possibly lasting into Janu-

ary.

Distribution: restricted to the northern Cape flora

region, Micranthus cruciatus is known from the north-

ern Cedarberg immediately south of Pakhuis Pass and

in the Bokkeveld Mtns southwest of Nieuwoudtville

(Figure 7). Plants grow on rocky slopes, in seeps on

thin sandy soil over sandstone pavement, flowering in

December as the habitat dries out in the hot weather. No

doubt the species is rare but the very few collections are

probably due to its midsummer flowering when little

plant collecting is undertaken. We suspect that M. cru-

ciatus occurs in suitable sites between its few stations,

thus in the southern Bokkeveld Mtns and the Gifberg/

Matsikamma Mtn complex and perhaps elsewhere in the

Cedarberg. First collected by the late Elsie Esterhuysen

in 1941 according to available records, M. cruciatus has

elicited no attention until now and was assigned to the

broadly similar M. plantagineus (as M. jitnceus) in her-

baria.

Diagnosis: linear- to terete-leaved Micranthus crucia-

tus is immediately distinguished by its solid, narrow leaf

blades, ± 1.5 mm wide, with heavily thickened veins

and margins separated by narrow longitudinal grooves

(Figure ID). Leaves are either linear or terete becom-

ing cross-shaped in section distally with only the mar-

gins and central vein thickened. Plants broadly resemble

M. plantagineus although they are more slender than is

usual in that species, which has hollow leaves 2-3 mm

diam. and is conspicuous in the production of cormlets

in the lowermost and sometimes upper leaf axils. In con-

trast, M. cruciatus has no more than one or two small

cormlets, these borne in the axil of the lowermost leaf.

The pale mauve-blue or sometimes dark blue flowers

are typical of the genus in shape but notable in the short

perianth tube, ± 3 mm long, in the style dividing at the

mouth of the perianth tube and in the unusually long

style branches up to 2.5 mm long, divided for up to one

140

Bothalia 43,2 (2013)

third their length. Most species of Micranthus have the

style dividing opposite the base to middle of the anthers

and style branches typically less than 1.6 mm long. The

narrow, heavily thickened leaf in M. cniciatus is conver-

gent with that in M. filifolius, but in other critical details

the two are very different, the latter with coarsely fibrous

corm tunics and a collar of fibres around the base of the

stem.

It is noteworthy that typical, hollow-leaved M. plan-

tagineus with dark blue flowers also occurs in the

Pakhuis and Bokkeveld Mtns (e.g. Leipoldt 3596 BOL,

NBG, PRE) as well as in the Cedarberg, but it has not

been recorded growing near M. cniciatus and they evi-

dently have somewhat different habitat preferences.

Additional specimens

NORTHERN CAPE. — 3119 (Calvinia): sandstone slope between

Nieuwoudtville and Vanrhyns Pass on road to Keyserfontein, (-AD),

27 Nov. 1985, Goldblatt 7399 (MO, PRE).

WESTERN CAPE. — 3218 (Clanwilliam): Zandfontein, Farm Ver-

keerde Vley (Klip-op-mekaar), 12 km N of Pakhuis Pass, (-BB), July

2013 (fr.), Helme 111?, (NBG). 3219 (Wuppertal): Cedarberg, Pakliuis

to Heuning Vlei (-AA), 28 Dec. 1941, Esterhiiysen 7436 (BOL);

Pakhuis Pass, Kliphuis campsite, wet seep on sandstone pavement,

(-AA), 15 Nov. 201 1 (in bud), Goldblatt & Porter 13766 (MO, NBG).

7. Micranthus plantagineus Eckl., Topographisches

Verzeichniss der Pflanzensammlung von C.F. Ecklon:

43 (1827), nom. nov. pro Ixia plantaginea Alton: 59

(1789), nom. illeg. superfl. pro Gladiolus alopecuroides

L. Watsonia plantaginea Ker Gawk: t. 553 ( 1803), nom.

nov. pro Ixia plantaginea Aiton et nom. illeg. superfl.

pro G. alopecuroides L. Gladiolus plantagineus Pers.:

46 (1805), nom. nov. pro Ixia plantaginea Aiton et

nom. illeg. superfl. pro G. alopecuroides L. Type: South

Africa, without precise locality, Masson s.n. BM922008

(BM, holo.! — narrow-leaved specimens mounted

with Nelson 1777 with broad, flat leaves [= M. alo-

peciiroides]).

Phalangiiim spicaturn Burm.f.: 3 (1768), nom.

nud. [cited illustration, Plukenet: t. 310, f 1 (1694) lacks

text or figure analysis; it probably represents Ixia scilla-

ris L.; specimen in G: Herb. Burman is Micranthus and

designated the ‘type’.]. Micranthus spicatus (Burm.f)

N.E.Br.: 138 ( 1929), nom. inval.

Phalangium spicaturn Houtt.: 1 1 5 (1780). Type:

South Africa, without precise locality or collector, illus-

tration in Houtt., Nat. Hist. ed. 2, 12: t. 80 f 2 (1780).

Micranthus plantagineus var. junceus Baker:

179 (1892). Micranthus junceus (Baker) N.E.Br.: 138

(1929). Type: South Africa, [Western Cape], Groenek-

loof and vicinity, Zeyher 1611 (K., lecto!, designated

here, K000320508; PRE!, isolecto.; other collections

numbered Zeyher 1611 in PRE and SAM are from Klip-

fontein or Tulbagh, thus not type material).

Plants 200~400(-650) mm high, base without col-

lar of fibres. Conn globose, 12-15 mm diam., tunics of

dark brown, medium-textured, reticulate fibres drawn

into fine points above. Stem erect, simple or with up to

9 short branches, with cormlets in axil of lowermost

leaf and sometimes of upper cataphyll and rarely other

leaf axils. Leaves (2)3(4), green at flowering, lower-

most leaf longest, blades terete or oval in section, hol-

low, 2. 0-3. 5 mm diam., smooth when fresh with trans-

lucent veins, when dry, veins appearing thickened with

homologue of marginal vein pair more prominent, usu-

ally reaching to middle of spike to shortly exceeding

spike, uppermost I or 2 leaves sheathing for most of

length, with free part often slightly longer than sheath.

Spike (16-)40-100-flowered, lower bracts sometimes

subtending cormlets; bracts mid- to dark brown, ± 6 mm

long but slightly smaller if subtending cormlets, outer

with broad translucent margins, apices sharply acute

and ultimately curved outward, inner bracts ± as long as

outer, with 2 dark keels broadened toward base. Flow-

ers usually dark blue, occasionally pale blue or white,

evidently sometimes slightly sweetly scented, perianth

tube 6-7 mm long, tepals oblong, with thickened sub-

apical ridge on reverse, (4-)6-7 x ± 1.2 mm. Stamen

filaments ± 5 mm long; anthers oblong, 3-4 mm long.

Style ± 7 mm long, dividing between lower one third and

middle of anthers (rarely ± 1 mm below anther bases),

branches 1.0- 1.8 mm long, divided for one third to half

their length. Capsules smooth, ± um-shaped or narrowly

ovoid, (3)4-5 x 2-3 mm, with (2)3 or 4 seeds per locule,

5 mm long. Seeds elongate-angular, 3(4)-sided, taper-

ing to points at both ends. Flowering time: October to

December (rarely in May).

Distribution: Micranthus plantagineus has a wide

range across the Cape flora region, extending from the

Bokkeveld Plateau near Nieuwoudtville south to the

Cape Peninsula and east to Port Elizabeth (Figure 8). An

isolated population from the Anysberg Nature Reserve

in the Little Karoo {Vlok 2545) appears typical of the

species except for the shorter perianth tube, ± 4 mm

long. Plants typically grow in seasonally wet habitats,

often in marshy sites, along streams, or at least in places

that are waterlogged in the winter months.

Diagnosis: the elongate inflorescence with up to 100

flowers and a perianth that is often deep blue, but some-

times pale blue or white, are unexceptional for the genus

and identification of Micranthus plantagineus depends

on leaf morphology. The two to four leaves are straight,

stiffly erect, hollow and terete to oval in section, ± 2-3

mm diam., and reach or shortly exceed the spike (Fig-

ure 1 F). When alive the leaves are smooth with the veins

evident as paler, translucent lines. On drying, the veins

appear hyaline and the veins at the adaxial and abaxial

poles are somewhat more prominent. In addition, the

stem is often branched, and as many as four (exception-

ally nine) short branches may be produced shortly below

the base of the main spike, these seldom exceeding half

the length of the main spike. Lewis ( 1950) noted that the

lower flowers of the spike are often replaced by cormlets

[as many as five may be present in an axil]. That feature

is not universal and many othewise typical plants may

have normal flowers and capsules from base to apex of

the spike. A second characteristic feature of M. planta-

gineus is the presence of one or more cormlets in the

lowermost leaf axil (not invariably present in other spe-

cies) and occasionally in the axils of the upper cataphyll

and one or more of the upper leaves. Plants lack a col-

lar of fibres around the base (in contrast to superficially

similar M. filifolius and M. tubiilosiis).

Bothalia 43,2 (2013)

141

16° 18° 20° 22' 24° 26° 28-

FIGURE 8. — Distribution of Micranthiis plantagineiis.

There are two, somewhat poorly defined, morphs of

the species. One has narrow leaves, ± 2 mm diam. when

alive, and slender capsules, ± 4.3 x 1.5 mm. Plants with

mature capsules have 2( 1 ) fusiform-angular seeds per

capsule, ± 3. 7^.0 mm long. The second morph has

broader leaves, 2-3 mm diam. when alive, and urn-

shaped capsules ± 5 x 2-3 mm. Capsules have 4 seeds

per locule, these flattened-angular, 3. 0-3. 5 mm long.

Collections of Micranthiis plantagineiis from

Grootwinterhoek Forestry Station exemplify this situa-

tion: Goldblatt 10451 with dark blue flowers represents

the robust morph of the species, but slender-leaved

plants {Goldblatt 10452), growing adjacent to stands

of the robust morph, have white flowers. Other slen-

der-leaved M plantagineiis (Goldblatt 104 5 3) growing

nearby have blue flowers. These last two collections

consist of shorter, less robust plants, 200-250 mm tall,

and have more slender, but still hollow leaves ± 1.5 mm

diam. No other Micranthiis species were found in the

area making the possibility that hybridization has played

a role in this pattern of variation unlikely.

We must also mention a collection made by T.M.

Salter in May 1935 from Viljoen’s Pass. The morphol-

ogy conforms closely to Micranthiis plantagineiis in

the three terete and hollow foliage leaves, stems with-

out a collar of fibres at the base, cormlets in axils of all

leaves and in floral dimensions. Thus the only differ-

ence we see is the flowering time, noted as anomalous

on the specimen label by Salter. The marshy habitat like-

wise conforms to the species. A search for the popula-

tion in May 2013 failed to find any Micranthiis species

in bloom at this time of year, but the site may simply be

lost to farming activity or dam construction.

Putative hybrids between Micranthiis plantagineiis

and M. alopeciiroides are discussed under the latter spe-

cies.

Related species’, until now, several collections of

plants with filiform-linear and plane or terete, but not

hollow, leaves with a thickened central vein and mar-

gins (cross-shaped in section) have been included in

Micranthiis plantagineiis (as M. jiinceiis) in herbaria.

We believe these are separate species. The several south-

ern Western Cape populations with this leaf type, here

referred to M. filifolius, always have four or five, rarely

six leaves, the blades ± 1 mm wide vs, mostly three

(rarely two or four) leaves in M. plantagineiis. These

plants have flowers with a consistently shorter perianth

tube ± 4 mm long and shorter tepals, also ± 4 mm long

vs, both perianth tube and tepals mostly ± 6 mm long in

M. plantagineiis and often a pale blue mauve to almost

white (less often deep blue) perianth. M. filifolius is

rarely branched and we have seen no specimens with

the lower flowers aborted and replaced by cormlets, both

common but not universal traits of M. plantagineiis.

Other collections with this derived leaf type are

known from the northern Cedarberg and Bokkeveld

Mtns and are here treated as the new Micranthiis crii-

ciatiis Goldblatt & J.C. Manning. These plants have

only four, rarely five leaves, the two basal with linear

or terete, four-grooved blades, ± 1.5 mm wide, similar

to but broader than those of M. filifolius. An important

associated character is the style, which divides at the

mouth of the perianth tube into unusually long branches

2. 0-2. 5 mm long, divided for ± one third their length.

Other species of Micranthiis have the style dividing

between the base and middle of the anthers and the style

branches never exceed 1.5 mm.

Histoiy. long known as Micranthiis jiinceiis (Lewis

1950; Goldblatt & Manning 2000), M. plantagineiis

was evidently first recognized as a distinct species,

called Phalangiiim spicatiim by Burman ( 1768), at least

as to the specimen in his collection (now at the Deles-

sert Herbarium, Geneva). Burman provided no validat-

ing description, instead merely citing Plukenet’s (1694)

illustration in part 3 of the Phytogeographia. There

is no accompanying text or even polynomial identi-

fying the illustration, plate 310, f. 1., nor does the fig-

ure constitute a validating illustration with analysis.

Phalangiiim spicatiim Burm.f. is thus a nomen nudum

and invalid. Plukenet’s illustration is of a broad-leaved

plant and does not, in our opinion, represent any species

of Micranthiis but is probably Ixia scillaris L. Even if

any text associated with this illustration is found and if

the name is lectotypified on the specimen rather than the

Plukenet illustration, the combination M. spicatiis (L.)

Heyn. (1847) (= Thereianthiis spicatiis) prevents the use

of Bunnan’s epithet at species rank in Micranthiis.

Curiously, Phalangiiim spicatiim Houtt. (1780), typi-

fied by a good illustration, marks this as the first valid

naming of M. plantagineiis. Although seeming to refer

to Butman’s P. spicatiim, Houttuyn makes it clear this is

his species (Phalangiiim scapis spicatis mihi, i.e. Hout-

tuyn) and that the Plukenet figure cited by Burman is

an entirely different plant. As noted above, Heynhold’s

combination M. spicatiis (L.) Heyn. bars transfer of

Houttuyn’s epithet to Micranthiis.

In Aiton’s (1789) Hortiis Kewensis, Daniel Solan-

der, the unacknowledged author of the species in this

work, described Ixia plantaginea based on a collection

of Francis Masson. The sheet at BM includes two plants

with narrow, stiffly erect, centric leaves (the Masson col-

lection) and three specimens with shorter, plane leaves

(Nelson 1777) that are M. alopeciiroides. The Masson

specimens conform to the diagnosis, foliis linearibiis

strictis, spica disticha imbricata [leaves linear, straight

and upright] and constitute the holotype. The name is

142

Bothalia 43,2 (2013)

unfortunately superfluous as Gladiolus alopecuroides

was cited in synonymy and, likewise, transfers of Ixia

plantaginea to Watsouia (Ker Gawler 1803), and Gladi-

olus Pers. (1805), are superfluous as both authors cited

Gladiolus alopecuroides as synonyms. Ecklon (1827),

however, intended to transfer the species to Micranthus,

where it becomes valid and is treated as a new name

from that date rather than a new combination based on

/. plantaginea Aiton. Ecklon (1827) recognized M alo-

pecuroides as a separate species, the first author to dif-

ferentiate it from M. plantagineus, but whether deliber-

ately or by accident is uncertain.

Baker (1892) described Micranthus plantagineus van

junceus, citing no specimens, but later listed several

exsiccatae (Baker 1896), all of which were available to

him in 1892. We choose a lectotype from among these,

Zeyher 1611, a specimen in good condition and repre-

sentative of the species. The taxon was raised to spe-

cies rank by Brown ( 1929), who, at the time also identi-

fied Phalangium spicatum Burm.f as the same species,

at least as to the specimen in Bunnan’s herbarium. M

junceus, a name used until now for this plant, becomes a

synonym of M. plantagineus.

Additional specimens

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville waterfall,

damp washes along stream, on sandstone, (-AC), 5 Dec. 1996, Man-

ning 2129 (NBG); Nieuwoudtville Escarpment, small vlei in arid fyn-

bos, (-AC), 28 Nov. 1993, MacGregor s.n. (NBG153534); Oorlogsk-

loof Nature Reserve, (-AC), 14 January 2000, Pretorius 664 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Matsikammaberge,

among sandstone rocks, (-DB), 11 Nov. 1985, Van Jaarsveld & Bod-

enstein 8283 (NBG); top of Gifberg Pass, Fann Van Taakskom, (-DD),

11 Nov. 1985, Snijman 946 (NBG). 3218 (Clanwilliam); Piketberg,

road to Sun Mtn, (-DA), 16 Nov. 1993, Manning 2093 (NBG). 3219

(Wuppertal): Pakhuis Mtns above 3500 ft [1 065 m], (-AA), 30 Dec.

1940, Leipoldt 3596 (BOL, PRE), Nov. 1929, Thode A2I41 (PRE);

Biedouw Valley, (-AA), 25 Nov, 1955, Middlemost 1897 (MO, NBG);

Wuppertal, (-AA), Oct. 1929, Thode A2083 (NBG); Driehoek Vlei,

Cedarberg, (-AC), 3 Dec. 1934, Compton 4798 (NBG); banks of the

Olifants River at Citrusdal, sandy ground, (-CA), 5 Nov. 1982, Gold-

blatt 6707 (MO), Feb. 1982, Goldblatt 6556 (fr.) (MO); Citrusdal,

Farm Kleinplaas, moist hillocks in loamy clay among restios, (-CA),

11 Dec. 1997, Hanekom 2972 (MO, NBG, PRE); Gonnafontein, sea-

sonally damp sand, (-CB), 3 Dec. 2000, Pond 254 (NBG); Leeu River,

Ceres, (-CD), 18 Dec. 1944, Compton 16741 (BOL, NBG). 3318

(Cape Town): Darling Flora Reserve, (-AD), 17 Nov. 1964, Thomp-

son 76 (NBG); 13 Nov. 1956, (-AD), Winkler 166 (BOL); Kenilworth

Racecourse, low lying areas wet in winter, (-CD), 5 Jan. 1970 (fr.),

Esterhuysen s.n. (MO); Devil’s Peak, 300 ft [± 90 m], Dec., Pappe

s.n. (SAM). 3319 (Worcester): Groot Winterhoek Forest Station, rocky

sandstone flats, (-AA), 27 Dec. 1995. Goldblatt 10451 (MO, NBG);

Keerom hills at foot of Twenty Four Rivers Mtns, (-AA), 3 Dec. 1950,

Esterhuysen 17869 (BOL, PRE); wet flats 9.4 miles [± 14 km] NE of

Hermon Station, (-CC), I80ct. 1959, /IcocCv 20744 (MO, PRE). 3320

(Montagu): Anysberg Nature Reserve, deep loamy sand, edge of seep,

12 Oct. 1991, (BC), Vlok 2545 (MO); Swellendam, hill below Eleven

o’clock Mtn, (-DC), 25 Nov. 1952, Wiirts 519 (mixed with M. tiibiilo-

siis) (NBG); Langeberg between Lemoenshoek and Naauwkranz, Farm

Strawberry Hill, (-DD), II Jan. \951 , Stokoe s.n. (NBG). 3321 (Ladi-

smith): Garcias Pass, 1300 ft, (-CC), Dec. \9[)4, Luyt s.n. (BOL). 3323

(Oudtshoorn): Saasveld, George, (-DC), I Dec. 1985, Vhk 1299 (MO,

NBG). 3418 (Simonstown): Bergvliet Farm, E of sand pit, (-AB), 5

Dec. 1918, Purcell s.n. (SAM90I06)\ Cirkels Vlei, Cape Peninsula, (-

AB), 15 Jan, 1946, Barker 3954 (NBG), Lewis 1495 (SAM); Betty’s

Bay, (-BB), 5 Jan. 1962, Tijmans 25B1962 (NBG); Cape Hangkiip,

marsh (with M. filifolius ), (-BB), 25 Jan. 201 1, Goldblatt cS Manning

13625 (MO, NBG). 3419 (Caledon): Riviersonderend Bridge, foot of

Franchhoek Pass, (-AA), I Jan. 1936, Barker s.n. (/JOL45075); Nuwe-

berg Forest Reserve, below Forestry offices, (-AA), 31 Dec. 1989,

Goldblatt 9035 (MO); Viljoen’s Pass, in marsh, (-AA), 4 May 1935,

Salter 5255 (BOL, K). Drayton siding, Caledon, near stream, (-BA),

25 Jan. 2011, Goldblatt & Manning 13622 (MO, NBG); Femkloof

Nature Reserve, Hermanus, deep sand, (-AD), 5 Dec. 1975, Orchard

349 (MO, NBG); Fairfield Farm, W of Napier, clay ground, (-BD), 9

Dec. 1994, Kemper 1PC750 (NBG). 3421 (Riversdale): Stilbaai, Farm

Klipfontein, shale ground near water, (-AD), 26 Nov. 1990, Bohnen

9152 (NBG). 3422 (Mossel Bay): Mossel Bay, grassy plains, (-AA),

Jan. 1926, Taylor 316 (BOL); inland of Oubaai, George, (-AB), 3 Jan.

1994, Victor 558 (BOL); Belvedere, churchyard, (-BB), 30 Dec. 1928,

Duthie s.n. STE29795 (NBG). Without precise locality, as Stellen-

bosch, Somerset [West] and Hottentots Holland, without date, Ecklon

& Zeyher Irid 193 (83) (SAM).

EASTERN CAPE. — 3324 (Steytlerville): Honeyville Farm, 10

km along Humansdorp-Hankey road, (-DC), 9 Feb. 2009, Van Wyk

FBG293/CR3761/ (NBG); 'Galgebosch, Uitenhage’ [near Hankey],

(-DD), 1935, MacOwan s.n. (SAM). 3325 (Port Elizabeth): Loerie

Forest Reserve, (-CC), 21 Dec. 1933, Long 1 (NBG); Uitenhage Divi-

sion, between Vanstadensberg and Bethelsdorp, (-CD), 1840, Drege

8445 (K); between Port Elizabeth and Thornhill, (-CD), 31 Dec. 1939,

Barker 604 (NBG). 3424 (Humansdorp): Witte Els Bosch, flats, (-

AA), Dec, 1920, Fourcade 1025 (BOL, NBG, SAM); Humansdorp, (-

BB), Jan. 1932, Wagner s.n. STE17114 (NBG). Without precise local-

ity, as ‘Uitenhage,’ Dec., Ecklon & Zeyher Irid 194 (MO, SAM).

Hybrids

Interspecific hybrids are not uncommon in Micran-

thus and are likely to occur when two or more species

co-occur. Most striking of the hybrids is that between M.

plantagineus and M. tubulosiis. The two species flower

together at the foot of the Elandskloof Mtns in Elands-

berg Nature Reserve and present a remarkable sight. The

hybrids are locally very common growing with typical

M. tubulosiis and are always slightly shorter than the

parent, 100-150 cm high, and like it have a well-devel-

oped collar of fairly coarse fibres around base. The other

parent is less common, but present in small clumps, rec-

ognized by its erect habit, straight leaves and pale blue

flowers. The hybrid is evidently fertile (plants in fruit

have well developed capsules with apparently normal

seeds) and stand out in having a slightly flexuose stem

and narrower leaves than either parent. Unlike M. tub-

ulosus, which they otherwise most closely resemble,

hybrid individuals bear small cormlets at aerial nodes

and sometimes at the base of the spike. We have seen

similar hybrid plants near Elim where M. tubulosus and

M. plantagineus also grew side-by-side.

WESTERN CAPE. — 3319 (Worcester): Elandsberg Estate, foot of

the Elandskloof Mtns, (-AC), 2 Mar. 2000 (sterile), Goldblatt & Man-

ning 1 1281 (MO, NBG), I36I7(fr.) (MO, NBG); Jan. 2011 (fr.; grow-

ing with M. plantagineus and M. tubulosiis), Goldblatt d Manning

13605 (MO, NBG), 22 Jan. 2011 (sterile), 13609 (MO, NBG, 11 Nov,

201 1, Goldblatt & Manning 13751 (MO, NBG, PRE).

Less common are hybrids between Micranthus tubu-

losus and M. alopecuroides, but at Elandsberg Nature

Reserve we noted both species growing close to one

another with apparent hybrids among them. The putative

hybrids have short, plane leaves, in outline like those

of M. tubulosus but not round in section, although the

leaves have an airspace between the two surfaces and

lack the visible main veins of M. alopecuroides.

WESTERN CAPE. — 3319 (Worcester): Elandsberg Estate, foot of

the Elandskloof Mtns, Vangkraal road, (-AC), 22 Jan. 2010 (sterile),

Goldblatt & Manning I361I (MO, NBG).

ACKNOWLEDGEMENTS

We thank Elizabeth Parker and Lendon Porter for

their assistance and companionship in the field; Mary

Stiffler, Research Librarian, Missouri Botanical Gar-

Bothalia 43,2 (2013)

143

den, for providing copies of needed literature; and Clare

Archer, SANBI (Pretoria) for help with several ques-

tions relating to collections at PRE. We also thank Nick

Helme for helping with field observations of M plan-

tagineus. Collecting penriits were obtained from the

Nature Conservation authorities of Western Cape, South

Africa. We thank the curators of the following herbaria

for access to their collections or loans of types and other

specimens: BOL, MO, NBG, PRE, SAM.

REFERENCES

AITON, W. 1789. Hortiis Kewensis, vol. 1. George Nicol, London.

ANDREWS, H. 1799. Ixia fistulosa. The botanists repositoty 1,3; t.

59.

BAKER, J.G. 1877 [as 1878]. Systema iridearum. Journal of the Lin-

nean Society. Botany 16: 61-180.

BAKER, J.G. 1892. Handbook of the Irideae. George Bell and Co.,

London.

BAKER, J.G. 1896. Iridaceae. In W.T. Thiselton-Dyer, Flora Capensis

6: 7-71. Reeve, Ashford.

BROWN, N.E. 1929. The Iridaceae of Burman’s Florae capensis pro-

dronms. Bulletin of Miscellaneous Information. Roval Botanic

Gardens. Ke^v 1929: 129-139.

BURMAN, N.E. 1768. Prodromus florae capensis. Cornelius Haak,

Leiden.

ECKLON, C.F. 1827. Topographisches Verzeichniss der Pfianzen-

sammlung von C.F. Ecklon. Reiseverein, Esslingen.

GOLDBLATT, R 1971. Cytological and morphological studies in the

southern African Iridaceae. Journal of South African Botanv 37:

317^60.

GOLDBLATT, P. 1977, Chromosome number in Pillansia (Iridaceae).

.Annals of the Missouri Botanical Garden 64: 36, 37.

GOLDBLATT, P. & MANNING, J.C. 1990. Leaf and corm tunic struc-

ture Lapeirousia (Iridaceae-lxioideae) in relation to phylogeny

and infrageneric classification. Annals of the Missouri Botanical

Garden 77: 365-374.

GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa Strelitzia 9. National Botani-

cal Institute of South Africa, Cape Town and Missouri Botanical

Garden, St. Louis, Missouri.

GOLDBLATT, P. & MANNING, J.C. 2006. Radiation of pollinations

systems in the Iridaceae of sub-Saharan Africa. .Annals of Botany

(London) 97: 317-344.

GOLDBLATT, P. & MANNING, J.C. 2008. The Iris Family: natural

history and classification. Timber Press, Oregon.

GOLDBLATT, P. & MANNING, J.C. 2009. New species ofGeissorhiza

(Iridaceae: Crocoideae) from the southern African winter-rain-

fall zone, nomenclatural changes, range extensions and notes on

pollen morphology and floral ecology. Bothalia 39: 123-152.

GOLDBLATT, P. & TAKEl, M. 1997. Chromosome cytology of Iridace-

ae, base numbers, patterns of variation and modes of karyotype

change. .Annals of the Missouri Botanical Garden 84: 285-304.

GOLDBLATT, P., MANNING, J.C., DAVIES, T.J., SAVOLAINEN,

V. & REZAI, S. 2004. Cyanixia. a new genus for the Socotran

endemic Babiana socotrana (Iridaceae-Crocoideae). Edinburgh

Journal of Botanv 60: 517—532.

GOLDBLATT, P., RODRIGUEZ, A., POWELL, M.P., DAVIES, T.J.,

MANNING, J.C., VAN DER BANK, M. & SAVOLAINEN, V.

2008. Iridaceae ‘out of Australasia’? Phylogeny, biogeography,

and divergence time based on plastid DNA sequences. System-

atic Botany 33: 495-508.

HEDWIG, R.A. 1806. Genera Plantarum. Reclam, Leipzig.

HEYNHOLD, G. 1847 [as 1846]. Alphabetische imd synonymische

Aufzdhlung der Gewdchse. Dresden & Leipzig.

HOUTTUYN, M. 1780. Natuurlyke Historie, ed. 2, 12. F. Flouttuyn,

Amsterdam.

JACQUIN, N.J. 1797. Plantarum rariorum horti caesarei schoenbrun-

nensis I . C.F. Wappler, Vienna.

KER GAWLER, J. 1801 . Ixia fistulosa. Hollow-leaved Watsonia. Cur-

tis's Botanical Magazine 15: t. 523.

KER GAWLER, J. 1803. Ixia plantaginea. Small-flowered Watsonia.

Curtis’s Botanical Magazine 16: t. 553.

KER GAWLER. J. 1804 [as 1805]. Ensatorum ordo. Koenig & Sims

Annals of Botany 1 : 219-247.

KLATT, F.W. 1882. Erganzungen und Berichtigungen zu Baker’s Sys-

tema Iridacearum. Abhandlimgen der Naturforschenden Gesells-

chaft zu Halle 15: 44-404.

KLATT, F.W. 1894. Irideae. In T. Durand, T. & H. Schinz, Conspectus

florae africae 5: 143-230. Charles Van de Weghe, Brussels.

KUNTZE, O. 1891. Revisio Generum Plantarum. vol. 2. Felix, Leipzig.

KUNTZE, O. 1898. Revisio Generum Plantarum. vol. 3, part 3. Felix,

Leipzig.

LEWIS, G.J. 1941. Iridaceae. New genera and species and miscellane-

ous notes. Journal of South African Botany 7: 19-59.

LEWIS, G.J. 1950. Iridaceae. In R.A. Adamson & T.M. Salter (eds).

Flora of the Cape Peninsula, pp. 271-265. Juta, Cape Town.

LINK, J.H.F. 1821. Enumeratio plantarum. etc. Reimer, Berlin.

LINNAEUS, C. 1753. Species Plantarum. Salvius, Stockholm.

LINNAEUS, C. 1 756. Centuria Plantarum II. Hdjer, Uppsala.

LINNAEUS, C, fil. 1782 [as 1781]. Supplementum plantarum. Orph-

anotropheus, Braunschweig.

LODDIGES, G. 1830. Watsonia compacta. The Botanical Cabinet 16.

MANNfNG, J.C. & GOLDBLATT, P. 2011. Taxonomic revision of the

genus Thereianthus G.J. Lewis (Iridaceae: Crocoideae). Bothalia

41: 239-267.

MARLOTH, R. 1915. Flora of South Africa. 4. Monocotyledons. Juta,

Cape Town.

MCNEILL, J., BARRIE, F.R., BURDET, H.M., DEMOULIN, V,

HAWKSWORTH, D.L., MARHOLD, K., NICOLSON, D.H.,

PRADO, J., SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. 2006. International Code of Botanical Nomen-

clature (Vienna Code) adapted by the 17th International Botani-

cal Congress, Vienna, Austria, July 2005. Regnum Vegetabile

146: i-xviii, 1-568. Koeltz, Konigstein.

PERSOON, C.H. 1805. Synopsis plantarum 1. Cramer, Paris.

PLUKENET, L. 1694. Phytogeographia. Part 4. London.

REDOUTE. P.J. 1804. Les Liliacees 3. Didot Jeune, Paris.

REEVES, G., CHASE, M.W., GOLDBLATT, R, RUDALL, R, FAY,

M.F., COX, A.V., LEJEUNE, B. & SOUZA-CHIES, T. 2002.

Molecular systematics of Iridaceae: evidence from four plastid

DNA regions. American Journal of Botany 88: 2074-2087.

RICKETT, H.W. &. STAFLEU, F.A. 1959. Nomina generica conserv-

anda et rejicienda spermatophytorum. Tccxon 8: 213-243.

RUDALL & GOLDBLATT, P. 1991. Leaf anatomy and phylogeny of

Ixioideae (Iridaceae). Botanical Journal of the Linnean Society

106: 329-345.

SCHRANK, F. DE P. DE. 1822. Commentarius in Irideas capenses.

Denkschriften der Kdniglich-Baierischen Botanischen Gesells-

chaft in Regensburg 2: 165-224.

SIMS, G. 1801. Ixia fistulosa. Hollow-leaved ixia. Curtis's Botanical

Magazine 15: t. 523.

WENDLAND, J.C. 1898. Botanische Beobachtungen. Gebriidem

Hahn, Hannover.

Hi,

VI ■ ■' ■':'‘.^i-'. -s

y^\y^

’■■■■ v*i .'j- 1. ../< , V'/ ,6»v,‘, '.V"*y*¥ ..|H||pj^i

IWM;' ^

t yi ■ •■ ..r . ’ ■|■■^^'^«»■|^«»(’A' 1 "■ ■(«•■:. ^ '. ‘’^)sf

Pt'S xf

i^tii i*i/«»

• •l^ci»il)(‘tt»saBI*' Vi-' hi*! *■ I

' ■■5‘! (U h‘>!' ' 1

:*'* JI, “^iv.

"*' ' " ''<)i- ‘ - '' 4 ' '‘'

"||( . ■ ‘/•fr’.; «;.i4ii#«u9i(j|M ■;*'• ‘‘ ’ T't

';-? V. s * • *?% • V i-. -!»»*■. ,'

m ■- iV „Vdl^

r%»*- <j(. 1; *‘-'i\Si.if

ly n .) v*%

k4

‘»v .rifi 'n

4'

■v>^ - Tl- <*■■

99-.1

r$j»V«Vi(i

,l.il ^il r J*l. ft ■■-4! -

■ ■••»*-y. -•vji.ivMipi

, .'' ", ..' '. fjfrf-' '•

'■'<!!

»?-'l

X

■<■'*•■', rJi- • i-.-«

Vi

.’■s.

i f%UMah

iF»Wl(^ ■

•Wijk* a’H »*''fvt\V,

■ #.;

III I m >,4 irt* if-j

.-' •%,

.»: Hff *v-., \\iiro'A M

•■ ..^A4i4.ie iij’.

'‘i* xitw

'•»>r

s:;^*Wu,-

; tMrl r ■ .. ■-.■ *-» iV *Vf

jit ..^,j*^'/ri4u:j',y‘ ■ tV -i*v-J‘4 ■'■.'» ...i-

n ♦' «-(>*,». I , 'Al< ,ri-

'' ' ''

*'

|:r

U aU 4

#/' '

\ ■'*'*-*«»ht»4rjB

**■’ ■- '' ' JL> .-. ’ ■ '.I-' 'lii' *.»-l *■•

" ' ' ■ ‘ ' Ki! '

.. k,t rfMfe‘1'-

4

««• ’ I '4-'. -p T

>'*-t■^^i^-; “.Alf

. i ■ ' t l-*«' V 'i

|r$4fe^kv<w 'i'-

£»4

'•'<Mi. 4VT

'■:'i. ' 4i ,

,,fiv -*«*•

' ••! ‘* *W«

If ><m«-^fi

It A.i

.«i»-*Lr«i|

-% . •;

-Iti' Hi

. V. I-., , SL

'■'J' ■■■»

> ,.(31(10 -'I®-'--^! -jCr-iflifij.* ' I ■

■-,■■■ ' "•k"

W:--,, '

it"

Bothalia43,2: 145-151 (2013)

New taxa of Hesperantha (Iridaceae: Crocoideae) from the southern

African winter rainfall region and a review of the H, pilosa complex

P. GOLDBLATT' - and J.C. MANNING-^

Keywords: Iridaceae, Hesperantha, new' species, southern Africa, taxonomy, winter rainfall zone

ABSTRACT

The southern and tropical Africa genus Hesperantha Ker Gawl., now with 85 species, is distinguished in subfam. Cro-

coideae by the style dividing into relatively long, usually laxly spreading style branches at or shortly below the mouth of

the perianth tube (rarely well within the tube or above the mouth of the tube) and, with a few exceptions, by hard, woody

corm tunics. We describe three new species here. H. dolomitica Goldblatt & J.C. Manning, a narrow endemic of limestone

outcrops on slopes north of the Vars River in the Knersviakte, Western Cape, has the bell-shaped corms characteristic of the

small sect. Hesperantha but is distinctive in the section in its pure white perianth with relatively long tube and soft-textured,

falcate to distally trailing leaves. H. laxifolia Goldblatt & J.C. Manning from the Pakhuis Mtns, Western Cape, stands out in

sect. Hesperantha in its prostate, somew hat succulent foliage leaves, and spikes of 2-5 white flowers with unusually short

filaments less than 1 mm long and particularly short anthers, ± 4 mm long. The short style branches, ± 4 mm long, remain

suberect rather than laxly spreading. H. secunda Goldblatt & J.C. Manning from the Roggeveld Escarpment, Northern Cape,

has until now been included in H. pilosa but differs in its secund spike of nodding flowers with short style branches, and

leaves with broadly winged margins. We also recognize a new subsp. bracteolata (R.C. Foster) Goldblatt & J.C. Manning

of H. pilosa (L.f ) K.er Gawl. for populations of plants with diurnal flowers with usually blue or purple (occasionally white)

tepals lacking dark pigmentation on the reverse. With additional material to hand, w'e reduce blue-flowered H. ciliolata Gold-

blatt to synonymy in subsp. bracteolata and report range extensions for H. pilosa subsp. pilosa. now' recorded as far east as

the Langeberg near Cloete’s Pass.

INTRODUCTION

Hesperantha Ker Gawl., now with 85 species (Gold-

blatt 1984, 2003; Goldblatt & Manning 2007a) is one

of the larger genera of Iridaceae subfam. Crocoideae

Burnett. Its range extends from the southwestern Cape

and Namaqualand, South Africa, through eastern south-

ern Africa to East Africa, Ethiopia and Cameroon,

but is centred in the southern African winter rainfall

zone. Hesperantha is distinguished by the style divid-

ing shortly below the mouth of the perianth tube (rarely

within or well above the mouth of the tube) into rela-

tively long, ± straight, usually laxly spreading style

branches and, with a few exceptions, by hard, woody

corm tunics (Goldblatt & Manning 2008). Here we

describe three new species of the genus, two from the

western half of Western Cape and one from Northern

Cape. H. dolomitica Goldblatt & J.C. Manning, a nar-

row endemic of limestone outcrops in the Knersviakte

of southern Namaqualand, stands out in its long, nar-

row, falcate leaves, and pure white flowers with rela-

tively long tube and tepals remaining cupped when

fully expanded. H. laxifolia Goldblatt & J.C.Manning

from the Pakhuis Mtns of northwestern Western Cape

has prostrate, soft-textured basal leaves and among the

smallest flowers in the genus, the perianth tube about 5

mm long, tepals 10 mm long and filaments less than 1

' B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P. O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt@mobot.org.

^ Research Centre for Plant Growth and Development, School of Life

Sciences, University of KwaZulu-Natal. Pietermaritzburg, Private Bag

XOl, Scottsville 3209, South Africa.

^Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. E-mail: j.manning@

sanbi.org.za.

mm long. Both H. dolomitica and H. laxifolia have bell-

shaped corms with a flat base and margins finely ser-

rated, placing them in sect. Hesperantha of the genus

(Goldblatt 1982). We provide a revised key to the sec-

tion, now with eleven species. In the H. pilosa (E.f.)

Ker Gawl. complex of sect. Concentricae Goldblatt, we

recognize the new subsp. bracteolata (R.C. Foster) Gold-

blatt & J.C.Manning for populations of H. pilosa (L.f.)

Ker Gawl. with diurnal flower phenology and mostly

blue or purple (rarely white) flowers (vs. usually flowers

white with dark pigmentation on the reverse of the tepals

and crepuscular floral phenology). New collections of

subsp. bracteolata have rendered the distinction between

it and blue-flowered H. ciliolata Goldblatt untenable and

we reduce the latter to synonymy. Range extensions for

H. pilosa subsp. pilosa show that it occurs near Swellen-

dam and in the eastern Langeberg near Cloete’s Pass,

well to the east of its recorded stations. Lastly, we recog-

nise the new species H. secunda (H. sect. Concentricae)

for a collection from near Middelpos until now included

in H. pilosa but differing from that species in its secund

spike of nodding flowers with curved perianth tube and

relatively short style branches ± 4 mm long, and leaves

with winged leaf margins and raised main vein.

NEW SPECIES OF SECT. HESPEH4NTHA

1 . Hesperantha dolomitica Goldblatt &

J.C.Manning, sp. nov.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Kners-

viakte, Fann Varsche Rivier 260, 300 m north of Vars

River, south-facing limestone crevices, ledges and moss

pads, 76 m, (-BC), 25 July 2012, Helme 7425 (NBG,

holo.).

Plants mostly 180-250 mm high. Corm bell-shaped

with flat base; tunics dark brown, softly woody, outer

146

Bothalia 43,2 (2013)

layers irregularly broken, margins minutely toothed.

Stem unbranched, glabrous. Leaves 4, lower two basal,

± linear, flaccid, ± falcate, third leaf inserted shortly

above ground and sheathing stem for most of its length,

uppermost leaf bract-like, inserted on upper third of

stem, entirely sheathing, basal leaf blades 60-80 x 3^

mm. Spike 2^-flowered, outer bract mostly 12-15 mm

long, pale green, evidently becoming dry and pale straw-

coloured during flowering, inner slightly shorter, with 2

green keels, narrowed distally and sometimes recurved

above. Flowers radially symmetric, white without darker

colouring on reverse, tepals remaining cupped when

fully open, sweetly scented; perianth tube narrowly fun-

nel-shaped, 12-14 mm long; tepals subequal, elliptic,

12-14 X 4 rnm. Stamens with filaments ± 3 mm long;

anthers 6-8 mm long, pale yellow; pollen yellow. Ovaiy

ovoid, 2. 5-3.0 mm long; style dividing at mouth of tube,

branches ± 7 mm long, diverging. Capsules and seeds

unknown. Flowering time'. July, probably also August;

flowers begin to open ± 12:30 and are fully open later in

the day, time of closing not known.

Distribution and ecology, local in the Knersvlakte

in southern Namaqualand, Flesperantha dolomitica is

restricted to limestone outcrops north of the Vars River

(Figure 1). Mostly wedged in cracks in the rocks, plants

are also occasionally found in loamy, red soils at the

base of the south-facing limestone cliffs. We regard the

species as EN (Endangered), in light of its narrow range,

very particular habitat requirements, small total popula-

tion, and potential for the mining of its limestone habi-

tat.

Diagnosis'. FI. dolomitica has the bell-shaped corms

with a flat base that are characteristic of sect. Hesperan-

tha (Goldblatt 2003). Within the section, //. dolomitica

is recognized by its relatively large, pure white flowers

with tepals remaining cupped even when fully expanded

and without the darker pigmentation on the outside of

the tepals that is a common feature in white-flowered

species of the genus. The perianth tube and tepals are

about the same length, 12-14 mm long, relatively large

for sect. Hesperantha. Additional observations are

needed to confirm the ultimate orientation of the tepals,

which may not occur until after dark, but we believe

the tepals remain cupped as they achieved this orienta-

tion by early afternoon and had remained unchanged by

17:00 (N. Helme pers. comm.).

Hesperantha dolomitica is one of a growing number

of very narrow endemic plant species discovered

on limestone substrates in the Knersvlakte. In Iri-

daceae these now include Babiana carminea Gold-

blatt & J.C. Manning (2007b), Ixia acaulis Goldblatt &

J.C. Manning (1993) and Moraea deserticola Goldblatt

& J.C. Manning (1986) as well as H. dolomitica. All

these species, and up to fourteen additional endemics

in other families, are endangered if plans for mining the

limestone substrate on which these plants grow come to

fruition.

Additional specimen seen

WE.STERN CAPE— 3118 (Vanrhynsdorp): Knersvlakte, Farm

Varsehe Rivier E,xtension A 227, sleep, S-facing creviees in limestone

ridge, (-BC), 25 .Inly 2012, Helme 7424 (MO, NBC).

FIGURE I. — UxstnbnUon of Hesperantha dolomitica. A;//, laxifolia,

• ; and H. seciinda, o.

2. Hesperantha laxifolia Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape, 3219 (Wuppertal): moun-

tains north of Pakhuis Pass, occasional among sandstone

rocks on south-facing slopes and banks, (-AA), 12 Oct.

2012, Goldblatt & Porter 13877 (NBG, holo.; K, MO,

PRE, iso.).

Plants 90-160 mm high. Conn bell-shaped with

flat, sometimes oblique base, 10-15 mm diam.; tunics

dark brown, inner layers ± woody, outer layers becom-

ing soft and broken, margins minutely serrated. Stem

unbranched, glabrous. Leaves 3(4), lower 2 leaves usu-

ally prostrate (suberect in deeply shaded sites), fleshy,

flaccid, obtuse, 50-70 x 4-5 mm, with main vein raised,

margins pale, slightly thickened, third leaf sheathing

lower 1/3 of stem, slightly inflated, fourth leaf (when

present) on upper part of stem, bract-like and entirely

sheathing. Spike mostly 2-3(5 )-flowered, outer bract

8-9 mm long, green flushed red distally, inner slightly

shorter to ± as long, partly membranous with 2 green

keels, notched apically. Flowers white, outer and some-

times inner tepals red to brown outside, rose-scented;

perianth tube narrowly cylindric, slightly expanded near

apex, ± 5 mm long; tepals subequal, elliptic, ±10x4

mm. Stamens with filaments ± 0.5 mm long; anthers ±

4 mm long, erect, pale yellow; pollen white. Ovary

globose, ± 2.5 mm long; style dividing ± 1 mm below

mouth of tube, branches ± 2.5 mm long, arching upward,

reaching to slightly below middle of anthers. Capsules

and seeds unknown. Flowering time'. September to late

October; opening ± 18:00, closing ± 21:00-21:30. Fig-

ure 2.

Distribution', so far known from a single collection in

the Pakhuis Mtns (Figure 1 ), in peaty and loamy sand in

rocky sites, mainly on shady, south-facing slopes. As far

as we can determine, Hesperantha laxifolia has a very

limited range in the ridges north of Pakhuis Pass, and

has not been recorded south of the Pass, an area that has

Bothalia 43,2 (2013)

147

FIGURE 2. — Hesperantha laxifolia, Goldblatt & Porter 13877 (NBG). A, flowering plant; B, flower detail; C, capsules. Scale bar: A, C, 10 mm;

B, 2 mm. Artist: J.C. Manning.

148

Bothalia 43,2 (2013)

been moderately well botanized. The narrow range of

the species falls within a declared Wilderness Area.

Diagnosis: with white, crepuscular flowers typical of

the genus, Hesperantha laxifolia can be distinguished by

its usually small flowers, the tepals ± 10 mm long and

the tube only half as long. Particularly unusual, the short

stamens have filaments only about 0.5 long and anthers

± 4 mm long. The style divides just below the mouth of

the tube into style branches only 2.5 mm long, likewise

unusually small for the genus. The two basal leaves are

slightly succulent and normally prostrate (the feature for

which the species is named), although these leaves may

be ascending to suberect in damp, shady situations. The

bell-shaped corms with the tunic margins finely toothed

are typical of sect. Hesperantha (Goldblatt 1984), but

we have no hypothesis concerning the immediate rela-

tionships of H. laxifolia. The only other species of the

genus with prostrate leaves, H. montigena Goldblatt,

from the Hex River and Riviersonderend Mtns, has

much larger flowers with a perianth tube 12-15 mm long

and tepals 14-16 mm long, thus about as long as the

tube, and the corms are asymmetric, placing it in sect.

Concentrica Goldblatt.

Key to species of sect. Hesperantha

(species with corms triangular in outline to bell-shaped,

with a flat base horizontal or obliquely oriented)

1 . Corm base usually with prominent radiating spines or mar-

gins toothed; flowers diurnal, closing in late afternoon;

perianth white or pink to reddish purple, rarely yellow:

2. Perianth tube 6-1 1 mm long H. pauciflora G.J. Lewis

2’. Perianth tube 15-35 mm long:

3. Plants acaulescent; perianth white with dark markings at

base of tepals, tube (15-)30-45 mm long; anthers 3-5

mm long H. liiticola Goldbl.

3’. Plants usually caulescent; perianth unifonniy deep pink,

tube 15-25 mm long; anthers (4.0-)5.5-7.0 mm long . .

H. lalifolia M.P.deVos

1 ’. Corm base with small teeth or scarcely serrated and usually

without prominent spines; flowers crepuscular, opening

in late afternoon or after sunset; perianth white, cream or

pale yellow:

4. Style dividing at or below middle of perianth tube; style

branches and sometimes anthers partly or completely

included in perianth tube:

5. Flowers 20-25 mm diam. with tepals 10-12 mm long;

stamens and style branches fully included in perianth

tube H. cedannontana Goldblatt

5'. Flowers ± 18 mm diam. with tepals 9-10 mm long;

anthers exserted, but style branches reaching only to

mouth of tube H. satdanhae Goldblatt

4’. Style dividing just below mouth of perianth tube; anthers

and style branches fully exserted and filaments usually at

least partly exserted from perianth tube:

6. Flowers small, secund on straight spike; perianth tube

slightly curved, 4-6 mm long; tepals 4-7 mm long;

leaves either plane and sometimes with crisped margins,

or terete to ovoid in cross section and hollow; seeds with

sharp angles (minute wings) and seed coat with whitish

spongy cells /7. .syji/cYJto (Burm.f ) N.E.Br.

6'. Flowers small to large, not obviously secund, on straight

or flexuose spike; perianth tube straight, (5-)7-14 mm

long; tepals 6-18 mm long; leaves plane with straight

margins; seeds globose or sides lightly flattened by pres-

sure, seed coat dark brown:

7. Plants with 3 basal leaves and without cauline leaves;

perianth tube 12-16 mm long; tepals 6-10 mm long,

always shorter than tube; flowers pale yellow

H. sufflava Goldblatt

7’. Plants with 2 or more basal leaves and usually an addi-

tional 1 or 2 sub-basal or cauline, largely sheathing

leaves; perianth tube 5-14 mm long; tepals (9-) 12-1 8

mm long, usually longer than tube; flowers white,

cream or yellow:

8. Foliage leaves 2, usually prostrate, semi-succulent; fila-

ments < 1 mm long

H. laxifolia Goldblatt & J.C. Manning

8’. Foliage leaves 3, erect or falcate, not noticeably suc-

culent; filaments > 2 mm long:

9. Tepals uniformly white on reverse, aseending and

cupped when fully open; perianth tube 12-14 mm

long, ± as long as tepals; leaves falcate to trailing . .

H. dolomitica Goldblatt & J.C. Manning

9’. Outer or all tepals flushed red to brown outside,

spreading horizontally when fully open; perianth

tube 4—12 mm long, shorter than tepals; leaves fal-

cate to trailing:

10. Leaves narrow, 1-2 mm wide; perianth tube 10-12

mm; plants of shallow soils on sandstone pave-

ment H. lithicola J.C. Manning & Goldblatt

10’. Leaves broader, 4—8 mm wide; perianth tube 4—9

mm long; plants of deeper clay or sandy soils:

1 1 . Bracts green, rounded to truncate and often with

reddish margin; leaves usually at least 4; flow-

ers usually remote from leaves, borne on upper

third of the stem

H.falcata (L.f.) Ker Gawk: typical form

1 1 ’. Bracts green or becoming membranous and dry

above, and then ± acute; leaves often only 3;

flowers usually borne close to leaves and from

about middle of stem H. falcata:pentheri Baker

and trifolia Baker forms (Goldblatt 2003).

NEW TAXA IN THE HESPERANTHA PILOSA COMPLEX (SECT.

CONCENTRICAE)

Relatively widespread in the southern African winter-

rainfall zone, Hesperantha pilosa was recorded by Gold-

blatt (1984, as subsp. pilosa) from the Bokkeveld Mtns

in Northern Cape south to the Cape Peninsula and east

to Bredasdorp and Riviersonderend in Western Cape. H.

pilosa subsp. latifolia Goldblatt was raised by Goldblatt

(1987) to species rank as H. pseudopilosa Goldblatt and

is excluded from this discussion. As currently circum-

scribed then, H. pilosa consists of two distinct morphs.

Those from the Bokkeveld Mtns, Roggeveld and Klein

Roggeveld extending south to Touws River have diurnal

flowers with a blue to purple (rarely white) perianth with

the reverse of the tepals not strongly marked in contrast-

ing colour; whereas those from the central Cedarberg

south to southern Western Cape have crepuscular flow-

ers with a white (rarely pale to deep lilac) perianth with

the tepals coloured brown to dull red or purple outside.

Goldblatt (2003) drew attention to the two colour

morphs and noted some minor differences; particu-

larly that the flowers of blue to purple flowered plants

Bothalia 43,2 (2013)

149

have ascending to suberect anthers and often some-

what longer style branches than those of white flow-

ered plants, which have the anthers spreading hori-

zontally. Although Goldblatt then suggested that the

two morphs should be recognized taxonomically, he

expressed uncertainty that the blue- to purple-flowered

morphs were monophyletic given that some plants with

pale lilac flowers had been recorded in the Darling area

of Western Cape; thus nested within the range of the

white-flowered plants. Differences in pigmentation pat-

terns between the blue-flowered southern and northern

populations now lead us to consider that the populations

with diurnal flowers and mostly a blue or purple peri-

anth from the Bokkeveld Mtns and Roggeveld are most

likely monophyletic, and it seems useftil to recognize

them as a separate taxon. Two names at species rank are

available for these populations, H. puberula Schltr. ex

R.C. Foster from the Bokkeveld Mtns and H. bracteo-

lata R.C. Foster from near Sutherland (Foster 1948). We

now treat H. pilosa as comprising two subspecies, subsp.

pilosa and subsp. bracteolata (R.C. Foster) Goldblatt

& Manning, preferring subspecies rank because of the

weak morphological differences between them.

Our review has brought to attention the morphologi-

cally similar Hesperantha ciliolata Goldblatt, which was

distinguished from blue-flowered H. pilosa by its nar-

rower, strongly ribbed leaf blades, oval in cross section,

with short, spreading cilia. Typical H. pilosa was distin-

guished by its leaves with rather longer, softer cilia. New

collections of what we are now calling subsp. bracteo-

lata have bridged the differences between the two taxa.

A collection from Driefontein-se-Berg, southwest of

Calvinia [Goldblatt & Manning 13999 (NBG)], has the

ribbed leaf blades typical of H. ciliolata but bearing the

scattered fine, long hairs typical of H. pilosa, and several

recent collections from the Roggeveld and Bokkeveld

Mtns are intermediate, with plane, weakly ribbed,

shortly ciliate leaves. This expanded circumscription of

H. pilosa makes it impossible to continue to recognize

H. ciliolata.

Among collections identified as Hesperantha pilosa

from the Roggeveld, one collection {Goldblatt 5810)

from near Middelpos was noted by Goldblatt as unusual

in its secund spike of nodding flowers, and leaves with

winged leaf margins and raised main vein. After exam-

ining the specimens again while reviewing variation in

H. pilosa we note that the flowers have a curved peri-

anth tube, creamy white perianth, and style branches ±

4 mm long, thus half to two-thirds as long as other H.

pilosa subsp. bracteolata in the Roggeveld and else-

where. We describe this population as a separate species,

H. seciinda.

3. Hesperantha pilosa subsp. bracteolata

(R.C. Foster) Goldblatt & J.C. Manning, comb, et stat.

nov. H. bracteolata R.C. Foster in Contributions from

the Gray Herbarium 166: 6 (1948). Type: South Africa,

[Northern Cape], Farm Uitkyk, Marloth 9907 (B, holo.!;

PRE, iso.!).

Hesperantha puberula Schltr. ex R.C. Foster: 22

(1948). Type: South Africa, [Northern Cape], Oorlogs-

kloof, Schlechter 10952 (B, lecto.l designated by Gold-

blatt: 57 (1984); B!, BOLi, Gi, GRAI, Kl, PREI, US!,

iso.).

Hesperantha ciliolata Goldblatt: 59 (1984),

syn. nov. Type. South Africa. Northern Cape, Roggeveld

Escarpment, Farm Geelhoek, 21 Sept. 1953, Acocks

77/76 (PRE, holo.!).

Plants 70-200(-300) mm high. Corm ovoid, ± 6

mm diam; tunics woody, concentric. Stem unbranched,

glabrous or sparsely hairy. Foliage leaves 3, lower 2

linear to narrowly sword-shaped, blades plane or with

main vein and margins raised, or oval in cross section

and ribbed, 1^ mm wide, pubescent or minutely cili-

ate, cilia horizontal on edges of ribs or margins, margins

sometimes ± winged, upper leaf partly sheathing lower

half of stem, usually ribbed; with minute, scale like bract

on upper 1/3 of stem. Spike (2)3-5-flowered, bracts sub-

equal, 8-10 mm long, outer green or becoming ± pale

and membranous, inner membranous with 2 green keels,

forked at apex or entire. Flowers blue or purple (rarely

white), without darker pigmentation on reverse, salver-

shaped, unscented or with acrid, sour odour, opening

10:00 to 11:00 and closing mid to late afternoon; peri-

anth tube 6-8 mm long; tepals (11-)12-16 x 3-5 mm,

inner slightly shorter and wider than outer. Stamens with

filaments 2. 0-2. 5 mm long; anthers (4)5-7 mm long,

usually ascending to suberect, pale yellow or white; pol-

len yellow to white. Style dividing 1-2 mm below mouth

of perianth tube, branches ultimately spreading, (7)8-

9( 1 1 ) mm long, reaching to upper third of anthers, some-

times exceeding them, ± white, rarely purple. Capsules

sub-globose, 5-6 x 4. 5-5.0 mm. Seeds angular, ± 1 mm

long. Flowering time: mid-August to late September.

Distribution: subsp. bracteolata extends from the

Bokkeveld Mtns northwest of Nieuwoudtville through

the Roggeveld south into the Klein Roggeveld and

locally into the Voetpadsberg east of Touws River (Fig-

ure 3), usually in sandy, stony ground.

Diagnosis: distinguished from subsp. pilosa by the

blue or purple (rarely white) perianth, the outer or all

tepals not or only slightly more darkly pigmented on

the reverse, subsp. bracteolata also usually has particu-

larly long style branches, up to 11 mm long and reaching

the upper third of the anthers when lined up together, or

even exceeding them. Some plants may have relatively

prominently ribbed leaves, the rib edges with short cilia

directed horizontally as well as long hairs on the veins,

a feature not seen in subsp pilosa. Flowers are occasion-

ally white or pale blue to almost white but are always

diurnal, open during the morning and closing after

12:20, sometimes after 14:00. In contrast, flowers of

subsp. pilosa are crepuscular, opening in the evening,

after 17:00 or at sunset, and then strongly scented

(Goldblatt et al. 2004) and the tepals are white (creamy-

white), rarely pale mauve, with the outer or all tepals

darkly pigmented red to brown or purple. Populations

with pale mauve tepals occur at scattered sites in the

southwestern Cape (e.g. Barker 10653 (NBG), Swart-

water Farm near Darling; Acocks 24314 (MO, PRE),

Lucasfontein, Malmesbury; Goldblatt 2501 (MO, NBG),

Theewaterskloof, Caledon). Tepals of subsp. pilosa are

mostly 9-15 X 2-4 mm and the style branches are 5-9

mm long, these dimensions overlapping those of subsp.

bracteolata, which has tepals (11-)12-16 x ± MS and

style branches (7-)8-9(-l 1 ) mm long.

150

Bothalia 43,2 (2013)

FIGURE 3. — Distribution of Hesperantha pilosa subsp. pilosa, •; and

subsp. bracteolata, o.

As noted by Goldblatt (2003) the type of subsp.

bracteolata consists of short plants about 70 mm

high, but a later collection from near the type local-

ity includes taller, more robust plants, some up to 150

mm high, evidently growing in a wetter season. Foster

( 1948) named the species for the tiny scale in the upper

half of the stem but that feature is actually typical of H.

pilosa and some related species. The type plants match

several other collections from the southern Roggeveld

in their purple perianth. The type of H. pubenda from

the Bokkeveld Mtns south of Nieuwoudtville has the

blue perianth characteristic of other collections from

the area. There are also a few records of white-flowered

plants within the range of subsp. bracteolata but these,

like those with blue to purple flowers, have a diurnal

flower phenology and lack red to dark brown pigmenta-

tion on the outside of the outer or all tepals. Flowers of

subsp. bracteolata are blue in the Bokkeveld Mtns and

the Roggeveld as far south to Ganagga Pass, purple west

of Sutherland close to the escarpment and pale blue to

white in the Klein Roggeveld and nearby. Plants from

sites between Matjiesfontein and Touws River have

blue flowers, thus matching those from the Bokkeveld

Mtns. Flowering phenology has been recorded as open-

ing 7:30, closing 12:30 in the Bokkeveld Mtns, closing

mid-afternoon west of Sutherland and after 14:00 in the

Klein Roggeveld.

Additional collections seen

NORTHERN CAPE.— 3119 (Calvinia): Bokkeveld Escarpment,

W of Farm Grasberg, (-AC), I Sept. 2005, Goldhiall 12682 (MO,

NBG); Grasberg road, NW of Nieuwoudtville, (-AC), 13 Sept. 1981,

Goldhiall 6262 (MO), 12 Oct. 1983 (IV.), 7072A (MO); ± 5 km N of

Papkuilsforitein, (-AC), 27 Aug. 2009, Davidson II595 (MO); Farm

Dricfontein, dolerite foothills at E end of Driefontein-se-Berg, (-DA),

16 Sept. 2009, Goldhiall. Manning Porler 13382 (MO, NBG). 3220

(Sutherland): Roggeveld Escarpment, Just above top of Ganagga Pass,

(-AA), I Sept. 1993, Goldhiall & Manning 9680 (MO, NBG); Farm

I lottentotskloof, Bo-Visrivier road, W of Sutherland, wet clay flats, (-

AC), 2 Oct. 1999, Goldhiall & Ncinni 1 1191 (MO, NBG); Farm Voel-

fontcin, valley S of farmhouse, wet places, (-AD), Goldhiall 6316

(MO); Roggeveld Escarpment, Farm Blesfontein, SW of Sutherland.

(-CD), 24 Sept. 2011, Goldhiall & Manning 13669A (MO, NBG); top

of Verlate Kloof, S of Sutherland, (-DA), 2 Oct. 1999, Goldhiall d

Nanni 11189 (MO, NBG); Klein Roggeveld, Farm De Hoop, (-DC),

30 Aug. 2007, Goldhiall & Porler 12920 (MO, NBG).

WESTERN CAPE. — 3220 (Sutherland): top of valley N of Farm

Fortuin, (-DC), 26 Aug. 2006, Goldhiall & Porler 12707 (MO, NBG);

78 km S of Sutherland, S-slope in small valley, (-DC), Goldhiall &

Manning 9662 (MO, NBG); Klein Roggeveld, Farm De Hoop, (-DC),

30 Aug. 2007, Goldhiall & Porler 12920 (MO). 3320 (Montagu): foot

of the Voetpadsberg, 21.5 km E of Touws River, foot of sandstone

slope, (-AB), 3 Oct. 1999 (fr.), Goldhiall & Nanni 11200 (K, MO,

NBG, PRE); II Sept. 2001, Goldhiall & Porter 11877 (K, MO, NBG,

PRE, WAG); Matjiesfontein to Touws River, sandstone outcrop, (-AB),

27 Aug. 2006, Goldhiall & Porler 12712 (MO).

4. Hesperantha seeunda Goldblatt & J.C. Manning,

sp. nov.

TYPE. — [Northern Cape], 3119 (Calvinia): Rog-

geveld Escarpment, 56 km from Calvinia on Blomfon-

tein road to Middelpos, (-DD), 17 Sept. 1980, Goldblatt

5810 (NBG, holo.; K, MO, PRE, S, US, iso.).

Plants 100-180 mm high. Conn ovoid, 7-8 mm

diam; tunics woody, concentric. Stem unbranched, gla-

brous, with minute, membranous, scale like bract up

to 3 mm long on upper third of stem. Foliage leaves 3,

lower two basal, linear to narrowly sword-shaped, fal-

cate or straight, finely hairy, blades 2. 5^.0 mm wide,

margins raised and winged at 90” to surface, main vein

also raised and with narrow wings parallel to surface,

uppermost leaf sheathing lower part of stem for all

or part of its length, several ribbed, finely hairy. Spike

mostly 3-7-flowered, bracts subequal, 7-10 mm long,

outer green or becoming ± pale and membranous, inner

membranous with 2 green keels, forked at apex. Flowers

creamy white, nodding, tepals spreading at right angles

to tube, presence of scent unknown; perianth tube cylin-

dric, curving outward, 6-7 mm long; tepals 10-11 x

2. 5-3. 5 mm, inner slightly shorter and wider than outer.

Stamens with filaments ± 3 mm long; anthers 5-6 mm

long, evidently spreading, pale yellow; pollen yellow.

Style dividing ± 1.5 mm below mouth of perianth tube;

branches ascending, ± 4 mm long, reaching to base or

lower quarter of anthers. Capsules and seeds unknown.

Flowering time: September.

Distribution: known from one population from the

Roggeveld Escarpment west of Middelpos (Figure 1 ),

Hesperantha seeunda is recorded on shallow, fine-

grained soil over rocky sandstone pavement.

Diagnosis: when first collected, Hesperantha seeunda

was referred to H. pilosa with comments on its unusual

winged leaf margins and main vein and nodding flowers

on a secLind spike (Goldblatt 1984). Reviewing the vari-

ation in H. pilosa in light of the many new collections

made since then, this collection remains unique and we

conclude that it represents a separate species, evidently

allied to H. pilosa but distinguished by vegetative and

floral morphology. All collections of H. pilosa have

plane to ribbed leaves, the margins slightly thickened

but rarely winged, and upright flowers with a straight

perianth tube. Details of the flowers of H. seeunda are

also unusual for H. pilosa, including the relatively short

tepals, 10-11 mm long vs. 9-15 mm in H. pilosa and

short style branches, ± 4 mm long and reaching the base

or lower quarter of the anthers when lined up together

Bothalia43,2 (2013)

151

(the style branches diverge in the open flower). In H.

pilosa the style branches reach to the middle or upper

third of the anthers or even exceed them and are 7-9(-

1 1 ) mm long. An illustration of the species appears in

Goldblatt (1984: 55 as H. pilosa).

RANGE EXTENSION

Hesperantha pilosa subsp. pilosa: until now the

recorded range of subsp. pilosa is from the central

Cedarberg at Middelberg and Uitkyk Pass to the Cape

Peninsula and east to the Great Swartberg at Bles-

berg and from Caledon east to Riviersonderend and

Bredasdorp (Goldblatt 1984, 1987). There are now

records from near Swellendam and from Herbertsdale at

the foot of Cloete’s Pass in the eastern Langeberg (Fig-

ure 3). The latter represents a range extension of nearly

200 km east of Riviersonderend, previously the most

easterly recorded station. Cloete’s Pass specimens are

consistent with subsp. pilosa, having crepuscular flowers

with a white perianth with dark brown on the reverse of

the outer tepals, a perianth tube ± 10 mm long and tepals

± 10 mm long.

WESTERN CAPE. — 3420 (Bredasdorp): S of Swellendam on road

to Bontebok Reserve, (-AB), 26 Aug. 2000, Goldblatt 11433 (MO,

NBG); 3421 (Mossel Bay): 2 km from Herbertsdale to Cloete's Pass,

(-BA), 26 Sept. 2003, Goldblatt & Porter 12599 (MO, NBG); N of

Herbertsdale at the foot of Cloete's Pass, (-?AB), 30 Sept. 2004, Gold-

blatt & Porter 13599 (MO, NBG).

ACKNOWLEDGEMENTS

We extend our gratitude to Elizabeth Parker and Len-

don Porter for their assistance and companionship in the

field; Nick Helme for bringing his discovery of the new

Hesperantha dolomitica to our attention; and Michelle

Smith for preparing the maps. Collecting permits were

provided by the Nature Conservation authorities of

Western and Northern Cape Provinces.

REFERENCES

FOSTER. R.C. 1948. Studies in Iridaceae V. Some new or noteworthy

species of Hesperantha. Contributions from the Gray Herbarium

166:3-27.

GOLDBLATT, P. 1982. Corm morphology in Hesperantha (Iridaceae,

I.xioideae) and a proposed infrageneric taxonomy. Annals of the

Missouri Botanical Garden 69: 370-378.

GOLDBLATT, P. 1984. A revision ol Hesperantha (Iridaceae) in the

winter rainfall area of southern Africa. Journal of South African

Botany 50: 15-141.

GOLDBLATT, P. 1987. New species and notes on southern African

Hesperantha (Iridaceae). South African Journal of Botany 53:

459^63.

GOLDBLATT, P, 2003. A synoptic review of the African genus Hesper-

antha (Iridaceae: Crocoideae). Annals of the Missouri Botanical

Garden 90: 390-443.

GOLDBLATT, P. & MANNING, J.C. 1986. Convergent evolution of the

Homeria flower type in six new species of Moraea (Iridaceae) in

southern Africa. Annals of the Missouri Botanical Garden 73:

102-116.

GOLDBLATT, P. & MANNING, J.C. 1993. Ixia acaulis. a new acau-

lescent species of Iridaceae: Ixioideae from the Knersvlakte,

Namaqualand, South Africa. Novon 3: 148-153.

GOLDBLATT, P. & MANNING, J.C. 2007a. New species and notes

on Hesperantha (InddiCtSLe) in southern Africa. Bothalia 37:177-

182.

GOLDBLATT, P. & MANNING, J.C. 2007b. A revision of the southern

African genus Babiana. Iridaceae: Crocoideae. Strelltzia 18.

GOLDBLATT, P. & MANNING, J.C. 2008. The Iris family: natural

history and classification. Timber Press: Portland, London.

GOLDBLATT, R, NANNI, L, BERNHARDT, P. & MANNING, J.C.

2004. Floral biology of Hesperantha (Iridaceae: Crocoideae):

how minor shifts in floral presentation change the pollination

system. Annals of the Missouri Botanical Garden 9 1 , 1 : 1 86-206.

;!hw ♦jnr.-rf',^4jr»'»v*ll *»t f'»*e» fh'^lNC tfi-nl' ,f •♦

' 't- ’ ■'LJ?' "4- ■

4.^^i<h-r '.,^49' n 41.'^‘t twili Ji.x/i'-itati-'* -- ’

V . ! .. 1 ;) li -. jajji 4 ,*)it if

' " ’*#' ’*<•• • ’• • f' ‘ '

*4;Mif!i*»-» -Jl .'

■ ■■ '“' »«** *<^'' ■■ ' 'r ^

w^ia;''iiM*i'(» . w. ' . ■ - '%' ^

^

. , y-‘;i»ti^ wv^- ’ ' |SaS

ii'i ,1 r

'-.i

*• lil*

■^»5»

/ »r^ ^ )|jT^

-^* • '■ / '*v,i l,,.)MMMai!3r:' '

f^HPiV--*

4IU^

■;^4SiWI(i|r'

Bothalia 43,2: 153-165 (2013)

Eight new species of Moraea (Iridaceae) from southern Africa with

range extensions and morphological notes in the genus

P. GOLDBLATP-^’and J.C. MANNING'’-^

Keywords: Iridaceae, Moraea L., new species, range extensions, southern Africa, taxonomy

ABSTRACT

We describe eight new southern African species in the largely sub-Saharan African genus Moraea L., one from the winter-

dry Great Karoo and the others from the winter-rainfall zone of the southwest of the subcontinent. M. striata Goldblatt &

J.C. Manning (subg. Visciramosae Goldblatt) from the southern foothills of the Great Swartberg Mtns has khaki-green flowers

with refiexed tepal limbs streaked with dark lines. The acaulescent M. singularis Goldblatt & J.C. Manning (subg. Umbel-

latae Goldblatt & J.C. Manning) with terete leaf blades is restricted to the dry, interior foothills of the Kamiesberg in Northern

Cape. M. filamentosa Goldblatt & J.C. Manning, subg. Acatiles (Baker) Goldblatt & J.C. Manning, is described from the

dry interior of Namaqualand in Northern Cape. The species is closely allied to the widespread M. falcifolia Klatt. In subg.

Polyanthes (Goldblatt) Goldblatt & J.C. Manning, we describe M. thermarum Goldblatt & J.C. Manning (sect. Hexaglottis

(Vent.) Goldblatt) from southwestern Namibia, allied to M. brevituba (Goldblatt) Goldblatt, and M. lazulina Goldblatt &

J.C. Manning (sect. Pseudospicata Goldblatt & J.C. Manning) from the Little Karoo, until now confused with M. exiliflora

Goldblatt but differing in its larger flowers with strongly reflexed tepals. The new M. petricola Goldblatt & J.C. Manning

(subg. Viensseiixia (D.Delaroche) Goldblatt), is restricted to higher elevations of the Pakhuis Mtns of northern Western Cape.

Lastly, in subg. Homeria (Vent.) Goldblatt & J.C. Manning, we recognize M. doleritica Goldblatt & J.C. Manning with pale

yellow flowers and anthers exceeding the style branches, restricted to dolerite outcrops in the Great Karoo, and M eburnea

Goldblatt & J.C. Manning from the northern foothills of the Klein Swartberg, which has a single, basal leaf and pale yellow-

flowers with filaments free distally. Moraea now has 222 species, including 203 in southern Africa, of which 200 are endemic

to the region. In addition to describing new species, the paper brings to light range extensions of a further eight species: M.

barnardii L. Bolus, M. elliotii Baker, M. exiliflora Goldblatt, M. falcifolia Goldblatt & J.C. Manning, M. fenestralis (Goldblatt

& E.G.H. Oliver) Goldblatt, M. loidsabolusiae Goldblatt, M. tulbaghensis L. Bolus and M. imgidculata Ker Gawl. Each of the

species falls into one of the subgenera mentioned for the new species e.xcept M. fenestralis which belongs in subg. Galaxia.

INTRODUCTION

The African and Eurasian Moraea Mill. (Iridaceae:

Iridoideae) is centred in the southern African win-

ter rainfall region in the southwest of the subcontinent

where the genus is both most diverse and most spe-

cies rich (Goldblatt et al 2002; Schnitzler et al. 2011).

Of the 1 1 subgenera recognized in a revised classifica-

tion of the genus (Goldblatt el al. 2013), all occur in the

region and all but one of them are endemic or centred

there. Patterns of speciation in the winter rainfall region,

an area of diverse soils and varied, but often low, rain-

fall, has resulted in the evolution of many extremely

local endemics, several known from just one or very few

populations. It is not surprising, therefore, that most of

the eight new species described here are narrow endem-

ics, and that five of them were discovered only in the

past five years in spite of many years of field explora-

tion by the authors as well as others. Just one, M. fila-

mentosa Goldblatt & J.C. Manning, first collected in

1958, has been known for some time but was until now

referred to M. falcifolia Klatt. Including these novelties.

“ B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P.O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt{S^mobot.org.

Research Centre for Plant Growth and Development, School of Life

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

XOl. Scottsville 3209, South Africa.

' Compton Herbarium. South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. E-mail: j.manning(g

sanbi.org.za.

Moraea now comprises 222 species, all but two of them

restricted to sub-Saharan Africa. The species described

here are arranged following the new the classification of

the genus by Goldblatt et al. (2013).

NEW SPECIES

Subg. Visciramosae Goldblatt

Sect. Miiltifolieae Goldblatt & J.C. Manning

1. Moraea striata Goldblatt & J.C. Manning, sp. nov.

TYPE. — South Africa, Western Cape, 3322 (Oudts-

hoom): foothills of the Great Swartberg, Rust-en-Vrede,

road from entrance to waterfall, (-BD), 8 Oct. 2012,

Goldblatt & Porter 13858 (NBG, holo.; MO, iso.).

Plants 450-600 mm high. Conn including tunics

20-25 mm diam.; tunics brown, sticky internally, accu-

mulating as coarse matted fibres forming collar around

base of stem. Stem 2-5-branched from upper nodes,

sticky below nodes; branches short, arching outward

above sheathing leaf, erect distally; sheathing leaves

30^0 mm long, acute-attenuate. Foliage leaves 3, all

basal, linear, falcate, often trailing distally, channelled,

V-shaped in cross section with pronounced keel, 4-5

mm wide when opened flat. Rhipidial spathes grey-

green, becoming dry distally, acute-attenuate, inner

30^0 mm long, outer ± half as long. Flowers dull

khaki, tepal limbs fully reflexed, with several longitu-

dinal dark brown-grey lines along entire length, outer

tepal limbs with pale yellow nectar guides at bases;

154

Bothalia 43,2 (2013)

outer tepals 25-30 x 10-11 mm, claws ± 7 mm long,

with dark basal nectary; inner tepals 22-25 x ± 8 mm.

Stamens with filaments ± 1 0 mm long, free but contigu-

ous in lower 6 mm, diverging in distal 4 mm; anthers

± 4 mm long, buff-yellow, shortly exceeding stigmatic

lobes; pollen orange-red. Ovary’ ± cylindric, 5-6 mm

long; style branches ±6x5 mm, crests erect, broadly

wedge-shaped, 6-7 mm long, outer margins lightly ser-

rated, with broad, entire petaloid stigmatic appendage

± 4 mm long. Capsules and seeds unknown. Flowering

time: October, probably also November; flowers opening

14:00-15:00 and fading at sunset. Figure 1.

Distribution: evidently a local endemic of the Oudts-

hoorn section of the Great Swartberg (Figure 2), Moraea

striata was recorded twice in 2012 on clay slopes at the

southern foot of the range. There appear to be no earlier

records of the species.

Diagnosis: the sticky stems, distinctive branching

pattern and stigmatic appendage place Moraea striata

in subg. Visciramosa where the unequal floral bracts

and more than two foliage leaves correspond with sect.

Midtifolieae, one of the two sections of the subgenus.

Flowers of M. striata are predominantly dull greyish

green or dull khaki, the outer tepal limbs with pale yel-

low nectar guides at the base and finely streaked with

dark grey longitudinal lines running almost the length

of the limbs. The anthers are buff-yellow and the pollen

bright orange. The tepal limbs of both whorls are fully

reflexed and the filaments are contiguous in the lower

half but free to the base, a feature also characteristic of

subg. Visciramosae. A particularly unusual feature of

M. striata is that the leaf blades are V-shaped in cross

section with a sharply angled keel. Similar leaves are

known in very few species of the genus, none of subg.

Visciramosae, in which rounded keels are typical.

Moraea striata most closely resembles M. biibal-

ina Goldblatt, a species of the western, winter rainfall

Karoo, which also has three leaves but flowers with

broad, diverging tepal claws forming a wide cup, the

tepal limbs buff to brown and reflexed to about 45°. M.

bubalina also has a short, acute stigmatic appendage up

to 2 mm long, very different from the large, petaloid

stigmatic appendage about 4 mm long in M. striata.

Flowers of M. striata bear a superficial resemblance to

those of M. inconspicua Goldblatt in their fully reflexed

tepals but the flowers of that species are much smaller,

the outer tepals 12-18 mm long (vs. 25-30 mm in M.

striata), and it consistently has two foliage leaves and

subequal, finely veined rhipidial spathes.

Additional specimen

WESTERN CAPE.— 3321 (Ladismith): foothills of the Great

Swartberg, road to Calizdorp W of turnoff to Swartberg Pass, (-DB), 8

Oct. 2012, Goldhiall & Porter 13855 (MO, NBG).

Subg. Umbellatae Goldblatt & J.C. Manning

2. Moraea singularis Goldblatt & J.C. Manning, sp.

nov.

TYPE. — South Africa, Northern Cape, 3018

(Kamiesberg): Vaalputs, NECSA site, (-BA), Platbak-

kies Succulent Karoo, 2 Aug. 2011, Van Rooyen 2827

(PRU, holo.).

Plants small, acaulescent 30^0 mm high. Corm

spindle-shaped, ± 10 mm diam., tunics of grey-brown,

wiry, reticulate fibres. Stem with 1-4 congested branches

at ground level. Leaves 3, clustered at ground level,

lowermost longest, inserted on stem at ground level,

falcate, terete, 50-80 mm long, 0. 8-1.0 mm diam.,

upper 2 leaves much shorter, recurved, bifacial, chan-

nelled or with short terete blade up to 20 mm long.

Rhipidial spathes green, outer leaf-like, recurved dis-

tally, 25-30 mm long, inner ± as long. Flowers rotate,

pale to golden yellow, limbs of all tepals with brown,

spear- or lozenge-shaped nectar guide near base, tepal

claws ascending, forming cup enclosing base of fila-

ment column, limbs spreading, inner usually twisted

through 90° distally; tepals united for 2 mm, claws ± 1

mm long, limbs obovate-pandurate, truncate, outer ± 12

X 6 mm, inner slightly smaller. Stamens with filaments

± 5 mm long, connate in a smooth, cylindric column ±

4 mm long, free and diverging in upper 1 mm; anthers

± 3 mm long, yellow; pollen yellow. Ovary cylindric, ±

10 mm long, included; style branches divided almost to

base into terete-involute arms diverging on either side of

each anther, ± 1.5 mm long, stigmatic at apices, crests

reduced, ± 1 mm long. Capsules and seeds unknown.

Flowering time: late July to early August. Figure 3.

Distribution and ecology: known from two localities

along the eastern foot of the Kamiesberg massif in cen-

tral Namaqualand (Figure 2), Moraea singularis grows

on gravel flats derived from granite bedrock. The species

is too poorly known to assess its conservation status, but

we provisionally suggest a listing of RARE. We see no

threat to its continued existence in this remote and thinly

populated area of minimal importance to farming activ-

ity. So inconspicuous is the species that it is easily over-

looked unless in bloom. Plants flower for just a few days

in any year and unusually early after the first rains of the

season. We suspect the species has a wider range than is

presently recorded.

Diagnosis: an acaulescent species, Mor-aea singu-

laris is recognized by its underground stem and short,

terete leaf blades. The yellow flowers, with both inner

and outer tepals limbs bearing brown nectar guides,

have tepal claws only ± 1 mm long forming a narrow

cup including the base of the smooth filament column.

The tepal limbs are obovate-pandurate, the inner three

slightly smaller than the outer and twisted through 90°.

The asymmetric corms with brown corm tunics accord

best with subg. Umbellatae as do the outer rhipidial

spathe, which sheaths the inner spathe only in the lower

half and arches outward distally, and also the darkly

veined tepals and brown nectar guides. Unique to the

subgenus is the terete blade of the basal leaf, while the

upper leaves are either channelled entirely or terete dis-

tal I y.

Additional specimen

NORTHERN CAPE.— 3018 (Kamiesberg): NE of Kamieskroon,

near Rooifontein, (-AB), 27 Aug. 2006, Gwvime-Evans s.n. (NBG,

photo.).

Subg. Acaules (Baker) Goldblatt & J.C. Manning

3. Moraea filamentosa Goldblatt & J.C.Mannirrg,

sp. nov.

Bothalia 43,2 (2013)

155

FIGURE 1. — Moraea striata (Goldblatt & Porter 13858). A, flowering plant; B, androecium and style. Scale bar; A, 10 mm; B, 2 mm. Artist;

John Manning.

156

Bothalia 43,2 (2013)

FIGURE 2. — Distribution of Moraea filamentosa, o; M. thennanmi. A;

M. singiilaris, ▲; M striata, •.

TYPE. — South Africa, Northern Cape, 3019 (Loer-

iesfontein): 22 miles [± 34 km] N of Loeriesfontein, (-

DA), May 1958, Hall 1385 (NBG, holo.).

Plants acaulescent, ± 20 mm high including flow-

ers. Conn ± 12 mm diam.; tunics of medium-textured,

shaggy fibres. Stem with congested branches at ground

level. Foliage leaves several, blades falcate, channelled,

up to 10 mm long, up to 1.2 mm wide when opened

flat, with thickened, hyaline main vein on abaxial sur-

face, apex obtuse, margins hyaline, ciliolate. Rhipidial

spathes: outer not distinct from leaves, with broad,

membranous sheaths, inner ± membranous, floral bracts

membranous, dry, elongate, often irregularly torn, ± as

long as leaves. Flowers possibly white, markings and

orientation of tepals not known; outer tepals lanceolate,

±14x4 mm, claws ± 5 mm, limbs probably spreading,

inner tepals ±12x2 mm, possibly erect. Stamens with

filaments ± 4.5 mm long, united in lower half; anthers

± 2.5 mm long, evidently yellow. Ovaiy elongate, fertile

part club-shaped, ± 5 mm long; style branches broad, ± 4

mm long, crests triangular, ± 4 mm long, erect. Capsules

and seeds unknown. Flowering time: May.

Distribution: known only from a single collection

from north of Loeriesfontein (Figure 2). The habitat is

not known.

Diagnosis: Moraea filamentosa is a dwarf, acaules-

cent species reaching no more than 20 mm high includ-

ing the flowers. Despite their small size, the leaves are

distinctive, having broad, membranous sheaths and fal-

cate, channelled blades with obtuse apices, ciliolate mar-

gins, and a thickened ridge in the midline of the abaxial

surface. A remarkable feature not known in any other

species of Moraea is that the floral bracts are elongate,

membranous, and about as long as the leaves, thus vis-

ible as pale filamentous threads among the leaf blades.

The flowers are small, with outer tepals ± 14 mm long,

but otherwise typical of Moraea, with filaments united

FIGURE 3. — Moraea singiilaris (Van Rooyen 2827 and Gnynne-Evans

s.n.). A, flowering plant; B, flower. Scale bar; 10 mm. Artist; John

Manning.

in the lower half, broad style branches, and prominent

erect crests. The species appears to be most closely

related to M. falcifolia, which has a broadly similar habit

but is a larger plant with larger flowers, the outer tepals

mostly 18-22 mm long, and long, tapering leaves up to

100 mm long with acute tips.

Although discovered in 1958, the species has

remained unnamed until now. The hope that additional

material would come to hand has not been fulfilled and

we have decided to describe the species from the two

plants that comprise the collection, hoping its formal

naming will stimulate a search for more plants.

Subgen. Polyanthes (Goldblatt) Goldblatt &

J.C. Manning

Sect. Hexaglottis (Vent.) Goldblatt

4. Moraea thermarum Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Namibia, 2817 (Vioolsdrif): Karas, Ai-Ais

Hotsprings Game Park, along Orange River, (-AA), on

cliff face, 13 Sept. 2012,^. Burke 12035 (WIND, holo.).

Plants 30-100 mm high. Conn 10-15 mm diam.;

tunics of dark brown to black fibres, extending upward

as short stiff bristles. Stem simple or 1- or 2-branched;

branches shortly stalked; sheathing leaves 1 or 2, 11-17

mm long. Cataphylls reddish brown. Foliage leaves 3

or 4, all ± basal, trailing or pendant, 80-120 x 1-3 mm,

undulate and loosely twisted, margins crisped, with

1 conspicuous lateral vein per side. Rhipidial spathes

green or becoming dry from tips, acute, inner 15-25 mm

long, outer ± half as long. Flowers fugaceous, yellow.

Bothalia 43,2 (2013)

157

Stellate, tepals connate into short cylindrical tube ± 1 .5

mm long, limbs spreading, elliptic, ± 15 x 5 mm, shortly

clawed, claws 0. 5-1.0 mm long. Stamens with filaments

united only at base (seeming free), ± 5 mm long; anthers

± 3 mm long, straight and suberect before dehiscence,

yellow. Ovaiy included in spathes, style arms paired,

filiform, spreading, 4-5 mm long. Capsules and seeds

unknown. Flowering time: mid-September to October.

Figure 4.

Distribution: known from two collections fringing

the Huib Hoch Plateau in southern Namibia, one from

the summit of the Aurusberg and the other along the

Orange River near Ai-Ais (Figure 2). Plants are confined

to shaded cliff faces, growing wedged in cracks in the

rocks, recorded in one instance as granite.

Diagnosis: resembling Moraea namaquana (Gold-

blatt) Goldblatt in its trailing, undulate or crisped leaves

and stalked lateral inflorescences, M. thermarum is dis-

tinguished by its narrower, linear leaves, 1-3 mm wide

with a single lateral vein on each side and somewhat

smaller flowers with a distinct perianth tube ± 1 .5 mm

long, the ovary included in the spathes. M namaquana,

which grows in open rocky ground, is a more robust spe-

cies, with leaves 5-11 mm wide with 2 conspicuous lat-

eral veins per side, and the tepals are free to the base.

FIGURE 4.— Mjraea thermantm (Burke 12055 and Williamson & Hammer 4564). A, flowering plant; B, flower; C, androecium and style.

Scale bar: A, B, 10 mm; C, 5 mm. Artist: John Manning.

158

Bothalia 43,2 (2013)

Additional specimen

NAMIBIA. — 2716 (Witpiitz): Karas, summit of Aurusberg, ±

1 000 m, (-CB), rock cracks in shade on southern aspect, 2 Nov. 1992,

G. Williamson & S. Hammer 4564 (NBG).

Sect. Pseudospicatae Goldblatt & J.C. Manning

5. Moraea lazulina Goldblatt & J.C. Manning, sp.

nov.

TYPE. — South Africa, Western Cape, 3322 (Oudts-

hoom): Schoemanshoek, Kranskloof, eastern foot of

rocky hill, in stony sandstone ground, (-AC), 8 Oct.

2012, Goldblatt & Porter 13859 (NBG, holo.; MO,

PRE, iso.).

Plants 160-350 mm high. Conn ± conical, 12-18 mm

at widest diam., with numerous small cormlets around

base; tunics of medium-textured, matted, dark grey

fibres. Stem flexuose, 1-5-branched, rarely simple. Foli-

age /eq/' solitary, linear, longer than stem, erect to fal-

cate, narrowly channelled, 2-3 mm wide when opened

flat. Rliipidial spathes with brown, attenuate tips, inner

33-45 mm long, outer ± half as long. Flowers fuga-

ceous, blue, outer tepal limbs with white nectar guides

sometimes edged with dark blue, limbs of both whorls

reflexed >45°, inner often more so than outer, unscented;

outer tepals ± lanceolate, 22-27 x g-9 mm, limbs 15-18

mm long, claws 7-8 mm long, inner tepals 17-20 x

5-7 mm. Stamens with filaments 8 mm long, united in

lower half; anthers ± 5 mm long, grey-blue; pollen white

or pale blue. Ovaty ± cylindric, 5-6 mm long; style

branches ±7x5 mm, crests narrowly wedge-shaped,

7-8 mm long. Capsules and seeds unknown. Flower-

ing time: mid-September to late October; flowers open ±

1 5:00 and collapse at dusk. Figure 5.

Distribution: relatively widespread in the Little

Karoo, Moraea lazulina extends from near Barrydale

(J.J. Vlok pers. comm.) and the Rooiberg south of Cal-

itzdorp to De Rust, east of Oudtshoom and Perdepoort

in the upper Long Kloof (Figure 6). Plants occur in

rocky sandstone ground and on limestone slopes near

the Cango Caves on the lower southern slopes of the

Great Swartberg.

Diagnosis: blue-flowered Moraea lazulina has until

now usually been associated with a second Little Karoo

species, M. exiliflora Goldblatt, which also has a soli-

tary, narrowly channelled basal leaf, and sometimes

even with multi-leaved M. bipartita L. Bolus. M. lazulina

has significantly larger flowers than M. exiliflora and

is typically a taller plant, usually with a moderately

branched stem. The flowers are always blue with large,

white nectar guides and outer tepals 22-27 x g-9 mm,

inner tepals 17-20 x 7 mm and filaments ± 8 mm long,

whereas M. exiliflora has smaller, pale blue, mauve or

white flowers with yellow (or white) nectar guides and

outer tepals 14-17 x 4. 5-6.0 mm, inner tepals 13-15

X 4. 5-6.0 mm, and filaments ± 4.5 mm long. Other

features of M. lazulina are likewise larger than in M.

exiliflora. The form of the flowers also differs to some

extent, the tepal limbs of M. lazulina being reflexed

more than 45°, the inner sometimes more so than the

outer. In contrast, the tepal limbs of M. exiliflora are

laxly spreading to reflexed up to 30°.

Additional specimens

WESTERN CAPE. — 3321 (Ladismith): Oudshoom-Calitz-

dorp road, 30.6 km E of Vanwyksdorp turnoff, (-DB), 16 Aug. 1985,

Steiner 952 (NBG). 3322 (Oudtshoom): limestone slopes near Cango

Caves, (— AC), 8 Oct. 2012, Goldblatt & Porter 13862 (MO). Perde-

poort, N of Camfer, sandstone slope burned last summer, (-CD), 28

Sept. 2004, Goldblatt & Porter 12574 (MO).

Subg. Vieusseuxia

6. Moraea petricola Goldblatt & J.C. Manning,

sp. nov.

TYPE. — South Africa, Western Cape, 3220 (Wupper-

tal): Pakhuis peaks, ± 5 km NW of Kliphuis, Rheboks-

vlei Extension 185, 925 m, (-AA), 25 Oct. 2008, Helme

5726 (NBG, holo.).

Plants ± 30 mm high. Conn globose, ± 12-16 mm

diam.; tunics of grey-brown, medium-textured, net-

ted fibres, accumulating in fibrous collar around base

of stem. Stem three intemodes long above ground,

unbranched, sheathing leaves with dry attenuate apices,

35-40 mm long. Foliage leaf solitary, basal, linear, nar-

rowly channelled, 2/3 to ± as long as stem, ± 2 mm wide

opened flat. Rhipidial spathes with dry, attenuate apices,

inner ± 40 mm long, outer ± half as long. Flowers last-

ing 3 days, purple, outer tepal limbs with white, wedge-

shaped nectar guides at bases, laxly spreading to weakly

reflexed; outer tepals ± 20 mm long, claws 5-6 mm

long, ascending; inner tepals ± 9 mm long, limbs spread-

ing, ± 4 mm long, 3-lobed with long, tapering central

cusp and short, rounded, obtuse lateral lobes, claws ± 5

mm long, ascending, tapering from narrow base to wide

apex. Stamens with filaments ± 8 mm long, united in

smooth cylindrical column in lower ± 3 mm, diverging

distally; anthers ± 5 mm long, dark purple; pollen red.

Ovaty ± cylindric, ± 8 mm long, exserted; style ± 5 mm

long, branches ±8x2 mm long, ascending, held above

subtending limbs, crests narrowly wedge-shaped, ± 4

mm long. Capsules and seeds unknown. Flowering time:

October to mid-November.

Distribution: as far as we know, Moraea petri-

cola (Latin: “growing in rocky sites” and also recall-

ing Kliphuis [Stone house] Peak nearby) is restricted

to higher elevations in the mountains to the north of

Pakhuis Pass in the northern Cedarberg (Figure 6).

Known from just one collection, it almost certainly has a

wider range than the present record indicates but is, nev-

ertheless, a fairly local endemic. Available information

makes it impossible to accurately assess its conservation

status, although its range is presently in a pristine habi-

tat, disturbed only by occasional fires. We provisionally

suggest a status of RARE.

Diagnosis: collected just once in flower in 2008 and

subsequently found in vegetative state in 2012, Moraea

petricola is a typical member of subg. Vieusseuxia

in its solitary basal leaf, 3-lobed inner tepals and, we

infer, long-lived flowers. The purple flowers have nar-

row, white, wedge-shaped nectar guides edged with

dark purple on the outer tepal limbs. The inner tepals

have unusually broad, 3-lobed limbs with a prominent,

linear, reflexed central cusp and rounded lateral lobes.

The flowers in general recall M. decipiens Goldblatt

& J.C. Manning, particularly in colour and shape, but

Bothalia 43,2 (2013)

159

have much longer filaments, ± 8 mm long (vs. 3. 5^.0

mm in M. decipiens), a style ± 5 mm long (vs. ± 2 mm)

and shorter tepal claws, 5-6 mm (vs. 8-9 mm). The

relatively long style branches are held well above the

subtending outer tepals limbs, whereas in M decipiens

the style branches are appressed to the tepal claws and

the inner tepals curve upward rather than having reflex

limbs with the central cusp directed downward. Addi-

tional collections of this rare species are needed to con-

firm our current observations based on limited material.

FIGURE 5. — Moraea lazidina (Goldblatt & Porter 13859). A, flowering plant; B, androecium and style. Scale bar: A, 10 mm; B, 2 mm. Artist: John

Manning.

160

Bothalia 43,2 (2013)

FIGURE 6. — Distribution of Moraea doleritica. A; M. eburnea, o; M.

laziilma, •; M. petricola, A.

Subg. Homeria (Vent.) Goldblatt & J.C. Manning

Sect. Stipanthera (Goldblatt) Goldblatt &

J.C. Manning

7. Moraea doleritica Goldblatt & J.C. Manning, sp.

nov.

TYPE. — South Africa, Western Cape, 3221 (Mer-

weville): Great Karoo, ± 52 km S of Fraserburg to Leeu

Gamka, low dolerite outcrops below Great Escarpment,

(-CB), 27 Sept. 2012, Goldblatt & Porter 13806 (NBG,

holo.; MO, PRE, iso.).

Plants 350^50 mm high. Conn unknown. Stem with

( 1 )2 or 3 branches from penultimate node, bearing a

single sheathing leaf 45-55 mm long, attenuate, ± dry

and light brown midway through flowering. Cataphylls

dry, brown and broken at flowering. Foliage leaf soli-

tary, linear, fibrotic, falcate, longer than stem, narrowly

channelled, 2-3 mm wide when opened flat. Rhipidial

spathes pale grey-green, dry and light brown in upper

half, attenuate, with fine white nerves evident, inner

50-65 mm tong at anthesis, outer ± half as long, inner

elongating after flowering to enclose ripening capsules.

Flowers fugaceous, pale yellow, outer tepal limbs with

deeper yellow nectar guides at base with fine dark dots;

outer tepals obovoid-pandurate, 25-27 mm long, limbs ±

20 mm long, claws of both whorls ± 6 mm long, ascend-

ing to form narrow cup including lower half of filament

column, limbs spreading and twisted through ± 45° dis-

tally, inner tepals ± 22 x 7 mm. Stamens with filaments

7-8 mm long, united for most of their length in smooth,

cylindric column, free and diverging in upper 0.7 mm;

anthers ± 8 mm long before anthesis, 5-6 mm after

releasing pollen, then tips arching inward, exceeding

style branches, yellow, pollen yellow. Ova?y cylindric,

usually exserted, ± 12 mm long; style branches ± 4 mm

long, crests ± linear, erect, ± 1.5 mm long. Capsules and

seeds unknown. Flowering time'. mid-September and

October; flowers opening early morning and collapsed

by 11:00. Figure 7.

Distribution', known only from the Great Karoo

between Leeu Gamka and Fraserburg in Western Cape

(Figure 6), Moraea doleritica is evidently restricted to

dolerite outcrops. Plants of both populations that we

encountered grew only among large, partly buried dol-

erite rocks, their corms tightly wedged in crevices in

the bedrock. The species is so far known from just two

populations only a few kilometres apart and south of the

Nuweveld Mtns that comprise this section of the Great

Escarpment. It is the only Great Karoo endemic species

of the genus.

Diagnosis', the pale yellow flowers with relatively

well developed style branches and short, erect crests

and, in particular, the filaments, free distally for a short

distance, are somewhat unusual for subg. Homeria

and recall M. pallida (Baker) Goldblatt and M. cookii

(L. Bolus) Goldblatt as well as two Namaqualand spe-

cies, M. schlechteri (L. Bolus) Goldblatt and M. kners-

vlaktensis Goldblatt. The latter two species have multi-

ple foliage leaves whereas M. cookii and M. pallida as

well as M. doleritica have a single foliage leaf and we

suggest that M. doleritica may be most closely related

to these two species. The spreading tepals twisted dis-

tally through about 45° of M. doleritica are also found

in M. cookii and M. pallida, another reason to consider

them related. Both M. cookii and M. pallida have a rela-

tively broad leaf clasping the lower part of the stem, thus

rather different to the particularly long, narrowly chan-

nelled, and relatively rigid, tough, fibrotic leaf of M. dol-

eritica. The range of M. doleritica in the Great Karoo is

complimentary to that of M. cookii and M pallida. M.

cookii is widespread in southern Africa, extending from

the Cedarberg in Western Cape through the mountains of

the Karoo to Lesotho and M. pallida occurs across the

Upper Karoo through Free State to Mpumalanga.

8. Moraea eburnea Goldblatt & J.C. Manning,

sp. nov.

TYPE. — South Africa, Western Cape, 3321 (Ladi-

smith): Seweweekspoort to Laingsburg near Fann Vlei-

land, northern foothills of the Klein Swartberg Mtns, (—

AC), 25 Sept. 2004, Goldblatt & Porter 12561 (NBG,

holo.; K, MO, PRE, iso.).

Plants 150-200 mm high. Conn unknown. Stem

mostly 2^-branched, flexed above sheaths of basal

and cauline leaves, sheathing leaves 30^0 mm long,

attenuate, dry and light brown in distal half Cataphylls

dry, brown, ± fibrous at flowering. Foliage leaf soli-

tary, basal, linear, falcate, ± twice as long as stem, nar-

rowly channelled, ± 2 mm wide when opened flat, often

dry and trailing distally. Rhipidial spathes dry and light

brown in distal half, attenuate, inner 30-35 mm long,

outer almost half as long. Flowers fugaceous, pale

ivory-yellow, limbs of all tepals with deeper yellow

nectar guides at bases, spreading; outer tepals narrowly

obovate, 20-25 x 7-8 mm, limbs 18-22 mm long, claws

of both whorls ± 2 mm long, ascending as a narrow cup

including lower half of filament column, inner tepals

18-21 X 6-7 mm. Stamens with filaments ± 5 mm long,

united in smooth, cylindric column 4 mm long, free and

Bothalia 43,2 (2013)

161

FIGURE 7. — Moraea doleritica (Goldblatt & Porter 13806). A, flowering plant (without corm); B, androecium and style; C, style. Scale bar: A,

mm; B, 2 mm; C, 1 mm. Artist: John Manning.

162

Bothalia43,2 (2013)

diverging in upper 1 mm; anthers ± 3 mm long before

anthesis, ± 2 mm after releasing pollen, exceeding style

branches, yellow; pollen yellow. Ovaty spindle-shaped,

usually exserted, ± 5.5 mm long; style branches flat-

tened, 2.5 X 1.5 mm long, with broad stigmatic lobes,

crests ± linear, erect, ± 1 mm long. Capsules obovoid, ±

10 mm long. Seeds unknown. Flowering time: Septem-

ber to mid-October.

Distribution: known only from the Farm Vleiland

north of the Klein Swartberg Mtns (Figure 6).

Diagnosis: the relatively small, pale yellow flowers of

Moraea eburnea have moderately well developed, flat-

tened style branches terminating in unusually broad stig-

matic lobes and short, erect crests somewhat unusual in

subg. Homeria, as are the filaments, united in a smooth,

cylindric column about 4 mm long, but free and diverg-

ing on the upper 1 mm. The distally free filaments and

moderately well developed style branches suggest that

M eburnea may be most closely related to the M. pal-

lida group (sect. Stipantherae) as most other species of

subg. Homeria have the filaments completely united.

If this assignment is correct, M. eburnea is by far the

smallest member of the alliance. The slender stem and

narrow leaf place M. eburnea in an isolated position in

the group. Relationships in subg. Homeria are difficult

to assess by phenetic comparison (Goldblatt 1981) and

molecular study using plastid DNA sequences have so

far provided no useful insights except that the M. pallida

group (including M. cookii, M. pallida and M reflexa

Goldblatt) is retrieved as a clade sister to the remaining

species of subg. Homeria (Schnitzler et al. 2011; Gold-

blatt et al. 2013). Of the 37 species of subg. Homeria,

only M eburnea and M. doleritica remain to be included

in any molecular analysis.

RANGE EXTENSIONS AND MORPHOLOGICAL NOTES

1. Moraea barnardii L. Bolus (subg. Vieusseuxia):

has small, white outer tepals flecked with dark blue and

undulate margins, bright red pollen and lacking inner

tepals. The distinctive M. barnardii is known for certain

from Shaw’s Pass, south of Caledon (Goldblatt 1976),

where it has been collected repeatedly. It is not common

there but plants are scattered on the sandstone slopes

east and west of the summit of the Pass and bloom well

only after a veld fire. There is also an unconfirmed and

somewhat doubtful record from near Gansbaai (Gillett

43 H BOL), some distance to the southeast. Because of

its narrow range, M. barnardii is currently regarded as

Critically Rare (CR) (Raimondo et al. 2009). We report

here a possible new locality for the species, from near

the mouth of the Palmiet River, Kleinmond (Gillett

4250 BR, MO). The collection, made by J.B. Gillett in

1938, and until now identified as M. tripetala, conforms

closely to type of M. barnardii. The flowers lack inner

tepals entirely, the outer tepal limbs have undulate edges

and the filaments, ± 8 mm long, are united for ± 1 .5 mm.

The dark-coloured anthers, the original colour of which

is no longer evident, are only 3 mm long, thus conform-

ing closely to M. barnardii, in which the anthers are 3^

mm long in the type population. Like Gillett’s collection

from Gansbaai, the Palmiet River record requires confir-

mation. If correct, the conservation status of the species

will require revision.

2. Moraea elliotii Baker (subg. Polyanthes): as cur-

rently circumscribed, single-leaved and usually blue-

flowered, M. elliotii extends through eastern southern

Africa from the Grahamstown area of Eastern Cape

through KwaZulu-Natal to Limpopo and Swaziland and

with an outlying record from Malawi (Goldblatt 1986,

1993). New collections from west of Grahamstown

expand the geographic range of M. elliotii and call into

question its circumscription. Plants matching the type,

which is from Mpumalanga, have ascending tepal claws

and spreading to ultimately half reflexed inner and outer

tepal limbs (the form is illustrated in Goldblatt 1986),

filaments usually 3. 5^. 5 mm long united in the lower

1.5-2. 0 mm and style branches 8-10 mm long. Plants

of the two Eastern Cape collections have somewhat dif-

ferent flowers, the inner tepals ultimately spread from

the base and are somewhat twisted in propeller fash-

ion, the filaments are united only basally and the long

style branches lie appressed to the outer tepals claws for

almost their entire length. The leaves are also unusual,

having tightly inrolled margins and seeming terete and

are inserted above ground level. Flowers of the Elands-

berg population are white with purple-edged nectar

guides, but those from the Suuranys Mtns have pale,

grey blue flowers. The floral form in these two collec-

tions is a good match for flowers of M. elliotii illustrated

in Pooley (1998), Wild Flowers of KwaZulu-Natal,

which also have a white perianth. The circumscription

of M. elliotii needs re-examination. We note here that at

least the synonym M. juncifolia N.E.Br. (Brown 1929),

based on plants from the Saddleback, Barberton, Mpu-

malanga, has a terete leaf inserted on the stem above

ground level. We are unable to resolve the question as to

whether M. elliotii, as presently interpreted, comprises

two species, with the terete-leaved plants, often with

white to pale grey-blue flowers a separate taxon, but

wish to draw attention to the question.

EASTERN CAPE. — 3324 (Steytlerville); Suuranysberg, (-CC),

Nov. 2011, Smith s.n. (NBG); Elandsberg, pass from Patensie to

Melkhoutboom via Mimosa Vale, crest of hill ‘white with mauve

markings,’ (-DB), 22 Oct. 1993, Beau & Viviers 3057 (BOL).

3. Moraea exiliflora Goldblatt (subg. Polyanthes):

when described (Goldblatt 1986) Moraea exiliflora was

known from only two collections, both from Towerkop

in the Klein Swartberg near Ladismith in the Tittle

Karoo. New collections have expanded its range consid-

erably: there are records from the Touwsberg, foothills

of the Kammanassie Mtns, southwest of Eadismith, and

from the upper Long Kloof southwest of Uniondale.

Plants at the Long Kloof site are restricted to relatively

dry, west-facing, rocky sandstone slopes and flower

in unburned, dry fynbos vegetation dominated by Res-

tionaceae and Bobartia. M. exiliflora is clearly not a rare

endemic of the Little Karoo but is a moderately wide-

spread, although poorly collected, species. Its apparent

rarity is evidently due in part to its small flowers with

for example, outer tepals 14-18 x 4. 5-6.0 mm, inner

tepals 13-15 x 2. 5-3. 5 mm and anthers 3. 0-3. 5 mm

long. It also has an unusual flowering phenology: flow-

ers open in the late afternoon; those of the population

near Uniondale opened after 16:00 and collapsed by

18:00, a pattern that probably reflects that for the entire

species. M. exiliflora is most easily confused with M.

lazulina, described in this paper, which also has a single

Bothalia 43,2 (2013)

163

foliage leaf and a similarly shaped flower but is a taller,

more robust plant with larger flowers. The similarly pro-

portioned flowers of M. lazulina have outer tepals 22-27

X 7-9 mm (limbs 14-18 mm long), inner tepals 17-20 x

7 mm and anthers ± 5 mm long. The tepals are strongly

reflexed, at least to 50°, the inner tepals often more

strongly reflexed than the outer, whereas tepals of M.

exiliflora are laxly spreading to slightly reflexed, mostly

< 15° below horizontal.

WESTERN CAPE. — 3321 (Ladismith): NE slopes of Touvvsberg

above Farm Boerboonfontein 115, (-CA), 7 Oct. 1993, Snijman 1394

(NBG). 3322 (Oudtshoom): roadside near Mons Ruber Winery, (-CB),

8 Sept. 2002, Goldblatt & Porter 12556A (MO); southern foothills

of Kammanassie Mtns, between Koutjie and Scheeperskraal, (-DA),

Vlok & Schutie 274 (MO); Avontuur to Herold, sandstone slopes above

Farm Kykoe, (-DD), 19 Sept. 2010, Goldblatt & Porter 13558 (MO,

NBG), 26 Sept. 2012, 13802 (MO. NBG).

4. Moraea falcifolia Goldblatt & J.C. Manning (subg.

Acaules): widespread in arid and semi-arid southern

Africa, M. falcifolia extends from southern Namibia and

Namaqualand across Bushmanland and the Upper and

Great Karoo to Alexandria in Eastern Cape and to Kuru-

man and Kimberley in Northern Cape (Goldblatt 1976,

Goldblatt & Manning 2009). Here we report a signifi-

cant range extension of the species from Jacobsbaai on

the Atlantic coast of Western Cape near Saldanha, some

120 km north of Cape Town. The most southwesterly

records until now are from Vredendal and Vanrhynsdorp,

some 160 km to the north. This record is notable also

for perianth colour: predominantly white with yellow

nectar guides at the bases of the outer tepal limbs and

large dark marks at the apices of the inner tepal claws,

features typical of nearly all populations of M. falcifolia,

but the style crests also have large dark pigmentation on

the lower half, rarely present in the species.

We also draw attention to a collection of Moraea fal-

cifolia from the plateau of the Kamiesberg of central

Namaqualand, above 1 000 m. Plants of this population

are the most robust in the species, the leaves reaching

to 100 mm long and the flowers are the largest so far

recorded. For example, the outer tepals are 25-26 mm

long, the inner ± 24 mm long, whereas until now outer

tepals have been described as 15-22 mm long, the inner

13-20 mm long; the length of the anthers and ovary,

4-5 mm long are the same as recorded for M. falcifolia.

The inner tepals of this population lack the characteris-

tic purple marks otherwise, as far as we know, univer-

sal in the species, and when collected the inner tepals

were suberect, whereas the limbs are spreading in other

populations. The features of this population significantly

expand the circumscription of M. falcifolia. Typical M.

falcifolia has been recorded near Kamieskroon not far

distant from the unusual population we describe here but

at significantly lower elevation, ± 400 m. Apart from the

size of most features, the Kamiesberg population con-

forms in general aspect to M. falcifolia.

NORTHERN CAPE. — 3018 (Kamiesberg): Namaqualand,

Kamiesberg, plateau above Kamieskroon, along Buffels River on road

to Pedroskloof, deep sand, (-AA), 17 Sept. 2001, Goldblatt & Porter

7/907 (K, MO, NBG, PRE).

WESTERN CAPE.— 3217 (Vredenburg): Saldanha District,

Jacobsbaai, limestone ridge, (-DD), July 2007, Claassens s.ti. (NBG,

photo only).

5. Moraea fenestralis (Goldblatt & E.G.H. Oliver)

Goldblatt (subg. Galaxia): originally described as Gal-

axia fenestralis (Goldblatt & Oliver 1993) and trans-

ferred to Moraea by Goldblatt (1998), M. fenestra-

lis was then known from granite outcrops in eastern

Namaqualand near Kliprand in Northern Cape. A new

record from the granite hills east of Okiep near Spring-

bok (Helme 6619, NBG), made in July 2010, is there-

fore surprising. Plants from this site, flowering in July,

have very pale pink flowers with yellow anthers, fila-

ments united in a column, diverging in upper ± 2 mm

and leaves with a translucent central line, and thus cor-

respond closely to the type and other collections of the

species. M. fenestralis is inconspicuous even in bloom

and we suspect there are other populations awaiting

discovery in the area between Kliprand and Springbok-

Okiep area, a distance of over 125 km.

NORTHERN CAPE. — 2917 (Springbok): Namaqualand, granite

ridge top 6 km E of Okiep, (-DB), 17 July 2010, Helme 6619 (NBG).

6. Moraea loiiisaboliisiae Goldblatt (subg. Home-

ria): described in 1981 as Homeria boliisiae Goldblatt,

Moraea loitisaboliisiae was then known from northern

Western Cape in the Clanwilliam District in the hills

near Kransvlei and the lower slopes of the Nardouw

Mtns and from Northern Cape on the Bokkeveld escarp-

ment at Lokenburg. Plants matching the species have

now been recorded in the Kamiesberg on the slopes of

Rooiberg in central Namaqualand {Goldblatt & Man-

ning 9767). Plants from the Kamiesberg differ very lit-

tle from those already known. They have a single foliage

leaf inserted above ground level, a stem flexed above

the sheaths of the foliage and sheathing leaves and yel-

low flowers with short, erect tepal claws and filaments

largely united in a slender smooth column but free dis-

tally. The free portion of the filaments is ± 1 mm long

vs. 0.5 mm in plants from elsewhere and the style crests

are ± 1.5 mm long vs. 1 mm long in earlier collections.

The Kamiesberg record is significant as it extends the

range of M. loiiisaboliisiae some 160 km to the north of

its next nearest known station and outside the Core Cape

flora region (Manning & Goldblatt 2013).

NORTHERN CAPE. — 3018 (Kamiesberg): Kamiesberg, Farm

Karas [Welkom], low spurs of Rooiberg, (-AD), 26 Sept. 1993, Gold-

blatt & Manning 9767 (MO, NBG).

7. Moraea tiilbaghensis L. Bolus (subg. Vieiis-

seiixia): the southwestern Cape Moraea tiilbaghensis,

now including M. neopavonia R.C. Foster (Goldblatt &

Manning 2002), is allied to M. villosa (Ker Gawl.) Ker

Gawk, the two sharing villous leaves and stems and

flowers with broad, ± suborbicular to broadly ovate

outer tepal limbs up to 38 mm long with prominent, con-

trasting, dark-coloured nectar guides (Goldblatt 1976,

1986). Most populations of M. tiilbaghensis have bright

orange or occasionally red flowers with deep blue to iri-

descent green nectar guides. The species differs from

M. villosa. which has purple or mauve, rarely green-

ish white or even orange flowers, outer tepals 22-38 x

20-35 mm, anthers 6-10 mm long, not exceeding the

relatively long style crests (2. 5)5. 0-8.0 mm long. In M.

tiilbaghensis the outer tepals are mostly 1 5-20 x 1 5-20

mm and the anthers, 7-10 mm long, and always exceed

the short style crests, 1-2 mm long. Plants resembling

these two species, discovered on the western slopes of

Riebeek Kasteel Mtn, have white tepals with dark blue

nectar guides, a colour combination most consistent with

164

Bothalia 43,2 (2013)

M villosa, but the anthers, ± 9 mm long, exceeding the

style crests, only ± 2 mm long. These dimensions are

consistent with M tulbaghensis as are other features of

the flower except for the somewhat broader outer tepal

limbs (up to 23 mm wide vs. 15-20 mm) (Table 1). We

conclude that the Riebeek Kasteel population represents

a striking colour morph of M tulbaghensis with slightly

broader outer tepals not before recorded in the species

but differing in no other taxonomically significant fea-

tures. The population also represents a modest range

extension for the species, which has been recorded from

the Tulbagh Valley and in the Berg River Valley and

adjacent hills from the outskirts of Paarl in the south to

Korningberg near Piketberg.

WESTERN CAPE. — 3318 (Cape Town): western flank of Riebeek

Kasteel Mtn, clay slope, (-BD), 25 Sept. 2009, Goldblatt & Porter

/i445(MO,NBG, PRE).

8. Moraea imguiciilata Ker Gawl. (subg. Vieus-

seiixia): across its wide range, from Steinkopf in north-

ern Namaqualand to Graaf Reinett and Port Elizabeth,

M imguiciilata has prominently trilobed inner tepals,

the central, linear-attenuate lobe coiled inward and the

lateral lobes broader, obtuse and shorter than the cen-

tral (Goldblatt 1976; Goldblatt & Manning 1995). A

population referred here from the top of the Langkloof

in the southern Kamiesberg in Northern Cape {Gold-

blatt & Manning 10425, MO, NBG), flowering 18 Nov.

1995, stands out in having ± linear-filiform inner tepals

± 20 mm long (the tightly coiled distal half makes exact

measurement impossible), with slightly wider membra-

nous tissue in the middle. In addition, the filaments are

united in a column ± 5.5 mm long but free in the upper

2.5 mm and the anthers, ± 4 mm long, have an unusu-

ally prominent apiculus ± 1 mm long. Until now we

have regarded this as a local morph of the species and of

trivial significance. Typical M. imguiciilata is, however,

now known to occur throughout the upper Kamiesberg

including the Langkloof and the hills near Leliefon-

tein, a short distance to the north (e.g. Goldblatt & Por-

ter 13576 MO, NBG: Goldblatt & Porter 13692, MO,

NBG). These plants have the prominently trilobed inner

tepals exactly conforming to the type. The filaments in

these Kamiesberg populations are united in a column up

to 10 mm long and free for ± 1 .3 mm and the anthers are

± 6 mm long, with a short apiculus ± 0.5 mm long. Vari-

ation in populations of M. imguiciilata in Namaqualand

needs to be carefully compared with the divergent pop-

ulation before we can assess the significance of its dis-

cordant features. Provisionally we prefer not to expand

the circumscription of M. imguiciilata and the divergent

population is not accommodated in keys to the genus.

ACKNOWLEDGEMENTS

We thank Nick Helme of Cape Town for bringing to

our attention his discovery of Moraea petricola and the

occurrence of M. fenestralis near Okiep; Helga Rosch

of Middelpos, for alerting us to the existence of M. sin-

gularis; Jan Vlok of Oudtshoom for sharing his knowl-

edge of Moraea in the Little Karoo; Patrick Fraser for

directing us to his discovery of the white morph of M.

tulbaghensis', and Clare Archer for providing answers to

our several questions about holdings at the Pretoria her-

barium. We are also grateful to and Elizabeth Parker and

Len Porter for assistance in the field.

REFERENCES

BROWN, N.E. 1 929, Contributions to a knowledge of the Transvaal Iri-

daceae. Part 1. Transactions of the Royal Society of South Africa

17: 341-352.

GOLDBLATT, P, 1976 [1977]. The genus Moraea in the winter-rainfall

region of southern Africa. Annals of the Missouri Botanical Gar-

den 63:63:657-786.

GOLDBLATT, P. 1981. Systematics and biology of Homeria (Iridace-

ae). Annals of the Missouri Botanical Garden 68: 413-503.

GOLDBLATT, P. 1986. Convergent evolution of the Homeria flower

type in six new species of Moraea (Iridaceae) in southern Africa.

Annals of the Missouri Botanical Garden 73: 102-1 162,

GOLDBLATT, P. 1 993. Iridaceae. In G.V. Pope (ed.). Flora Zambesiaca

12,4: 1-106. Flora Zambesiaca Managing Committee, London.

GOLDBLATT, P. 1998. Reduction of Barnardiella, Galaxia, Gynandr-

iris, Hexaglottis. Homeria and Roggeveldia in Moraea (Iridaceae:

Irideae). Novon 8: 371-377.

GOLDBLATT, P. & MANNING, J.C. 1995. New species of southern

African Moraea (Iridaceae: Iridoideae), and the reduction of Rhe-

ome. Novon 5: 262-269.

GOLDBLATT, P. & MANNING, J.C. 2002. Notes and new species of

Moraea (Iridaceae: Iridoideae) from the southern African win-

ter-rainfall zone. Novon 12: 352-359.

GOLDBLATT, P. & MANNING, J.C. 2009. New species of Moraea

(Iridaceae: Iridoideae), with range extensions and miscellaneous

notes for southern African species. Bothalia 39: 1-10.

GOLDBLATT, P, & OLIVER, E.G.H. 1993. A new species of Galaxia

(Iridaceae subfamily Iridoideae) from Namaqualand, South Afri-

ca. Novon 3: 404-407.

TABLE I . — Comparison of critical taxonomic features of Moraea tulbaghensis and M. villosa

Bothalia 43,2 (2013)

165

GOLDBLATT, P„ SAVOLAINEN, V., PORTEOUS, 0., SOSTARIC,

I., POWELL, M„ REEVES, G., MANNING, J.C., BARRA-

CLOUGH, T.G. & CHASE, M.W. 2002. Radiation in the Cape

flora and the phytogeny of peacock irises Moraea (Iridaceae)

based on four plastid DNA regions. Molecular Phylogenetics

and Evolution 25: 341-360.

GOLDBLATT, R, MANNING, J.C. & SCHNITZLER, J. 2013. A

revised infrageneric classification and synopsis of the Afro-Eur-

asian genus Moraea (Iridaceae: Irideae). Bothalia 43: 29-41.

MANNING, J. & GOLDBLATT, P. 2013. Plants of the Greater Cape

Floristic Region 1 : the Core Cape flora. Strelitzia 29. South Afri-

can National Biodiversity Institute, Pretoria.

POOLEY, E. 1998. A field guide to wild flowers of KwaZulu-Natal and

the eastern region. Natal Flora Publications Trust, Durban.

RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E.,

HELME, N.A., TURNER, R.C., KAMUNDl, D.A. & MANYA-

MA, P.A. 2009. Red List of South African Plants. Strelitzia 25.

South African National Biodiversity Institute, Pretoria.

SCHNITZLER, J., BARRACLOUGH, T.G., BOATWRIGHT, J.S.,

MANNING, J.C., GOLDBLATT, R, POWELL, M.P, REBELO,

T. & SAVOLAINEN, V. 2011. Causes of plant diversification in

the Cape biodiversity hotspot of South Africa. Systematic Biol-

ogy 60: 343-357.

KHm •

liiCl

■•FI ,>*1''^. •; H'-^f|»l4 i Ji^'lt-^ :'•'' *»:>*

^ r ■ i-,^jS^.

»»

.,. ■; f i.. .

W’5 5*«0*'i»j- ^,* .♦‘mfr ^

^ r*.-’-_

‘ < W>» .|l_^

1 ' •

:i*

& m

W£Piwi|(.v-'.^/fct • ■

wtiL, 1^ ‘ ■ •"■4..?*jufci • S

■^;.,, i|

> '‘;#i*lV’''‘^ ■*. ■> •■ ■ ''''' ■*■■■■ ' *

VV. , rl

J j&t ;■; .„»,' ’-»'5 ■ ^ _ ^ ;‘ I

i'. ' ■■«-• »' ^ X

“ f ' • -'Vrf, -..ft ftiilllAmM' • " ■-‘i.C'yMnH:^^^ ^

Bothalia43.2: 167-178 (2013)

FSA Contribution 22: Asteraceae: Calenduleae: Garuleum

N. SWELANKOMO*

INTRODUCTION

The genus Garuleum Cass, is a member of the tribe

Calenduleae in the subfamily Asteroideae of the family

Asteraceae (Bremer 1994). Calenduleae comprises 12

genera, of which 1 1 occur naturally in southern Africa

(Nordenstam 2007). Only one genus. Calendula L., is

confined to the northern hemisphere, but some of its spe-

cies are cultivated or occur as weeds in southern Africa.

The tribe is economically important as some species are

highly palatable to livestock and game, e.g. Chrysanthe-

moides incana (Burm.f) Norl. (Van Breda & Barnard

1987), while others, e.g. Dimorphotheca cimeata DC.,

are poisonous to stock (Watt & Breyer-Brandwijk 1962).

Garuleum can be distinguished from other genera in

the tribe by its unique style. The style of the disc florets

is deeply bifurcate, with linear lobes covered with papil-

lae to well below point of bifurcation; whereas in other

genera the style of disc florets is shallowly bifid with a

short apical collar of hairs (Norlindh 1943; Goldblatt &

Manning 2000; Herman et al. 2000). In Garuleum, fruits

develop from the ray florets only, except in G. bipinna-

tum, in which the disc florets are bisexual and the fruits

that develop from these disc florets are winged, evi-

dently as an adaptation to wind dispersal (Swelankomo

2010). The ray cypselae in this species vary in shape

from obpyramidal-triangular with small acute angles to

narrowly obovate or conical, and the disc cypselae are

flattened and cordate, as in many species of Dimorplio-

theca Moench.

Garuleum currently includes eight species and two

subspecies, all endemic to southern Africa (Merxmiiller

1967). Although widespread in southern Africa, Garu-

leum is absent from Botswana, Swaziland and the Lim-

popo and North-West Provinces of South Africa (Her-

man et al. 2000, 2003).

No complete taxonomic revision exists for Garuleum,

a situation that is rectified here with a detailed taxo-

nomic treatment of the genus. Lectotypes are designated

as required and full descriptions, a key to species, illus-

trations, and distributions are provided for all the spe-

cies.

MATERIAL AND METHODS

Morphological characters in Garuleum were studied

from herbarium specimens housed in PRE, NH, GRA

and NBG (acronyms after Holmgren et al. 1990). The

length of the corolla, length and width of the involu-

cre, length of the style and ovary, number of ray florets

(where possible) and length of cypselae were measured

' Biosystematics and Collections Division, South African National

Biodiversity Institute, Private Bag XlOl. Pretoria, 0001. E-mail:

N.Swelankomo(gisanbi.org.za.

with the aid of a Bausch & Lomb StereoZoom® Series

microscope. Two cypselae from each taxon (except in G.

album and G. tanacetifolium) were examined by means

of a scanning electron microscope (ISl SX 25). All sam-

ples were dry and were not chemically treated before

being sputter-coated with gold-palladium. Anthers were

studied under a dissecting and light microscope (Olym-

pus Vanox-S).

TAXONOMY

Garuleum Cass, in Bulletin de la Societe Philoma-

tique de Paris 1819; 172 (1819); Harv.: 92 ( 1865); Norl.:

33 (1943); B.Nord.: 375 (1994); P.P.J. Herman et al.\ 139

(2000). Type species; Osteospermum caeruleum Jacq.

[= G. pinnatifidum (L’Herit.) DC.], lecto., designated by

Pfeiffer: 1411 (1873-1874).

Perennial herbs, shrublets or shrubs, sometimes vis-

cid, scabrid or pilose. Leaves alternate, sessile, pin-

natifid or toothed, sometimes pungent. Capitula radiate,

solitary or corymbose, tenninal, pedunculate. Involucre

hemispherical-campanulate. Involucral bracts in 2 or 3

rows, ovate to linear, usually ciliate, sometimes reflexed

and sometimes pungent. Receptacle convex, epaleate.

Ray florets female, fertile; corolla blue, purple, mauve,

white or yellow, lamina linear or oblong, 3-toothed, up

to 3 X as long as tube. Style branches linear, sub-acute

with stigmatic papillae along margins. Ray cypselae tri-

quetrous or somewhat compressed with entire or crenate

angles, glabrous. Pappus absent. Disc florets bisexual or

functionally male, yellow; corolla tube sub-campanulate

above, with 5 lanceolate lobes, lobes glandular, some-

times hairy on back. Anthers minutely caudate, with

ovate apical appendages; endothecial tissue polarized.

Style terete, deeply bifurcate with linear lobes, covered

with acute papillae to well below point of bifurcation or

stigmatic papillae along sides. Disc cypselae (develop-

ing only in G. bipinnatum) obovate or obcordate in out-

line, flattened, marginally winged with thickened rim,

glabrous. Pappus absent.

8 spp., southern Africa

Etymology: although the derivation of the name is not

explained by Cassini (1819), it is most likely a corrup-

tion of the Latin caeruleus, alluding to the blue ray flo-

rets, as suggested by Jackson ( 1990).

Key to species

1 a Leaves toothed or pinnatifid, divided up to ± halfway to midrib:

2a Leaves discolorous, white-woolly beneath

LG. album

2b Leaves not strongly discolorous, green and sparsely pilose

to glandular beneath:

3a Involucral bracts sub-biseriate, all ± equal in length

2. G. sonchifolium

168

Bothalia43,2 (2013)

3b Involucral bracts 3-seriate, outer much shorter than inner

3. G. latifolium

lb Leaves pinnatisect or pinnate, divided to midrib:

4a Capitula borne on short, inconspicuous peduncles obscured

by apical leaves 4. G. tanacetifolhim

4b Capitula borne on prominent, sparsely leafy peduncles:

5a. Involucral bracts ovate, apices obtuse to acute; capitula

usually solitary; stem very rough with conspicuous

glands 5. G. woodii

5b. Involucral bracts narrowly lanceolate, apices acuminate;

capitula in loose corymbs:

6a Subshrubs with annual flowering stems from gnarled

woody base; disc florets female-fertile, fruit develop-

ing from both ray and disc florets 6. G. bipinnatum

6b Shrublets with softly woody stems; disc florets female-sterile:

7a All leaves pinnate; stem minutely glandular; cypselae

3. 0-3. 5 mm long 7. G. pinna! ifidum

7b Lower leaves irregularly bipinnatipartite or almost

tripinnatipartite; stem glandular-hispid; cypselae

4.0 mm long 8 G. schinzii.

1. Garuleum album S. Moore in Journal of Botany

London 16: 133 (1878); C.Claassen 40; t. 1593 (1970).

Type: [Eastern Cape], ‘Kaffraria, Shawbury,’ 1800' [590

m], June [without year], Baur 226 (K000273563 —

ALUKA image!, lecto., designated here; K000273562 —

ALUKA image!, isolecto.). [K000273563 is selected as

lectotype as it is the most complete specimen, with asso-

ciated dissections, and bears Baur’s handwritten original

collecting label].

Viscid aromatic shrub, 0.6-0. 9 m high, abundantly

leafy and branched, with white-woolly indumentum

especially on abaxial surface of leaves, on peduncles

and on involucre. Leaves sessile, alternate, lyrato-pinnat-

ifid, 50-95 mm long, white-woolly on abaxial surface,

green and very scabrid above, margin serrate with pun-

gent teeth. Capitula radiate, several clustered towards

ends of branchlets, peduncles white-woolly, 15-30 mm

long. Involucre campanulate, 10 x 12-15 mm. Involu-

cral bracts 3-seriate, imbricate, oblong-linear, apices

reflexed, white-woolly abaxially, especially towards api-

ces, outer bracts 5. 0-5. 5 mm long, middle bracts 8. 5-9.0

mm long, inner bracts 12.0-12.5 mm long. Ray florets

± 26-37, female-fertile, purplish-blue; corolla limb

oblong-elliptic, 9-15 x 2.5 mm, somewhat glandular

towards base; corolla tube 4-5 mm long, glandular. Style

8. 5-9.0 mm long, swollen at the base, glabrous, cylin-

dric, branched, style branches linear, ± 2 mm long, suba-

cute. Ovary glabrous, obovoid, ± 3-angled, 3 mm long.

Mature cypselae not seen. Disc florets functionally male,

yellow, 6-7 mm long; lobes narrowly ovate, 1.5 mm

long, somewhat hairy on back. Anthers 3 mm long, lin-

ear, minutely caudate with a small apical, ovate append-

age. Style, 8-9 mm long, terete swollen at base; style

branches 1 mm long, narrowly ovate, hairy on outer sur-

face. Ovary narrowly obovate in outline, compressed,

narrowly winged, 4-5 mm long, glabrous. Flowering

//we.' August-February. Figure lA.

Distribution and habitat: Garuleum album is a range-

restricted species known only from a small area between

Lusikisiki and Port St Johns in the Eastern Cape Prov-

ince (Figure 3), occurring in a densely populated area

where it is declining due to ongoing habitat loss and

degradation. It is recorded from rocky, dolerite soils on

hill-slopes at ca. 600 m.

Diagnosis: Garuleum album is distinguished by the

soft, white-woolly indumentum, especially on the under-

surface of the lyrato-pinnatifid leaves, the peduncle, and

the involucre.

Additional specimens

EASTERN CAPE. — 3129 (Port St Johns): Tabankulu, Qala above

the road to Gomo Forest, (-AB), 15 Mar. 1995, Wopida 153 (NH);

10 miles [16 km] N of Lusikisiki, (-BC), II Aug. 1953, Codd 7992

(PRE); Mlengana, between Umtata and Port St Johns, (-CA), Feb.

1896, Flanagan 2532 (PRE); Port St Johns, Mlengana, (-CA), 18 Jan.

1910, Pegler 1569 (PRE); Near the Khoweni Forest, Nkanga location,

Libode area, (-CA), 26 Apr. 2006, Styles 3056 (NH).

2. Garuleum sonchifolium (DC.) Norl. in Studies in

the Calenduleae 1: 425 (1943) (and cf p. 30); Hilliard:

518 (1977). Osteospermum sonchifolium DC.: 465

(1838); Harv.: 440 (1865); Type: Eastern Cape, between

the Umtata and Umzimvubu Rivers, Drege 5109

(G-DC — microfiche! [PRE — photo.!], holo.; E, K, iso.)

Osteospermum sonchifolium var. subpetiola-

tum Harv.; 440 (1865), syn. nov. Type: [Eastern Cape],

‘Keiskammahoek’, Cooper 227 (TCD — ALUKA

image!, lecto., designated here; K — ALUKA image!,

NH!, PRE!, isolecto. [The TCD collection is more

complete and plentiful and includes dissected capitula

attached (in the capsule). The specimen also includes a

good drawing of a disc floret], [Syntype: [Eastern Cape],

Kreili’s country’, Bowker 337 (TCD — digital image!)].

Glandular-pubescent, viscidulous, aromatic, perennial

herb, suffrutex or dwarf-shrub, 0.3-1 .2 m high. Leaves

alternate, sessile, ear clasping at bases, membranous,

netted-veined, inciso-pinnatifid; the lobes short, sharply

few-toothed, all teeth acuminate. Capitula pedunculate,

radiate. Peduncle mostly solitary, rarely in a corymb.

Involucre campanulate, 9-10 x 3 mm. Involucral bracts

sub-biseriate, lanceolate-acuminate, 8.0-10.0 x 2-3 mm.

Ray florets ± 15-20, female, fertile, purple-blue, violet-

blue, blue; corolla limb oblong-elliptic, 8-10 x 4 mm;

corolla tube cylindric, up to 3 mm long, glandular. Style

up to 4 mm long, linear branehed, style branches ± 2.2

mm long. Ovary up to 2 mm long, glabrous, obovate

in outline. Disc florets functionally male, yellow; lobes

ovate, 0.2 mm long; corolla limb 5 mm long. Style 5

mm long, branched, style branches 0.2 mm long, linear

with pollen sweeping hairs, which cover style below the

point of bifurcation, for same length as that of branches.

Anthers linear, minutely caudate with ovate apical

appendages. Ovaty up to 3 mm long. Cypselae 5.5 mm

long, glabrous, three-angles, obpyramidial, strongly

rugose. Flowering time: December- July. Figure 1 B, 2A.

Distribution and habitat: Garuleum sonchifolium

occurs in KwaZulu-Natal from around Underberg and

Mooi River southwards to Kokstad and Fort Beaufort

Division in Eastern Cape, (Figure 4), between altitudes

of 450-1 860 m. G. sonchifolium has been recorded

along river banks and on hillsides, mountain slopes and

Bothalia 43,2 (2013)

169

ridges, along the road, and along forest margins, on

stony soils, rocky outcrops and grassland.

Diagnosis: resembling G. latifolinm in its alternate,

sessile, membranous leaves, coarsely toothed or more

deeply cut halfway to midrib but distinguished by the

sub-biseriate vs. 3-seriate involucre.

Additional specimens

KWAZULU-NATAL.— 2929 (Underberg): Lundy’s Hill, (-DB),

26 Mar. 1901, Wood 8116 (PRE); Tillietudlem, (-DB), 9 Feb. 1947,

Huntley I3I (NH). 2930 (Pietermaritzburg): Mooi River, Warley

Common, (-AA), 17 Mar. 1920, Mogg 7179 (PRE); Mooi River, near

Mooi River, (-AA), 19 Dec. 1885, Wood PRE 44110 (PRE); Notting-

ham road, (-AC), Mar. 1939, McClean 922 (PRE); Richmond, Byrne,

FIGURE 1. — Variation in shape and size of the leaves of Gandeiim species. A, G. album, Codd 7992; B, G. sonchifolium, McClean 922; C, G.

latifolium, Hitchins 54; D, G. bipinnatum, Esterhuysen 5286; E, G. tanacetifolium, Acocks 16812; F, G. woodii, Devenish 491; G, G. pin-

natifidum, Hanekom 908; H, G. schinzii subsp. schinzii, Giess, Volk & Bleissner 7176 ; I, G. schinzii subsp. crinitum, Mexmuller 826. Scale

bars: A-I, 20 mm.

170

Bothalia 43,2 (2013)

FIGURE 2. — SEM micrographs of the cypselae of Ganileiim species. A, G. sonchifoUum, Flanagan 599, ray cypsela; B, G. latifoliiim, Galpin

13374, ray cypsela; C, G. hipinnaliim, Rodriguez-Onbina c6 Cruces 2107, ray cypsela; D, G. hipinnalum, Rodriguez-Ouhina & Cruces 2107,

disc cypsela; E. G.woodii, Jordaan 2763, ray cypsela; F, G. pinnalifidum, Hanekom 1954, ray cypsela; G, G. schinzii subsp. schinzii, Oliver

& Sleenkamp 6306, ray cypsela; H, G. schinzii subsp. crinitum, Mer.xniuller 826, ray cypsela. Scale bars: A, B, E, H, 49 1 .5 1 2 pm; C, 385.238

pm; D, G, 509.065 pm; F, 475. 128 pm.

Bothalia 43,2 (2013)

171

FIGURE 3. — Distribution of Ganileum album. FIGURE 4. — Distribution of Ganileum sonchifolium.

(-CC), 24 Apr. 1932, Galpin 11913 (PRE). 3029 (Kokstad): Alfred,

Weza, Zuurberg, (-BC), 3 Mar. 1974, Hilliard 5465 (PRE); Alfred,

Flarding, Rooval Farm, (-DB), 2 Mar. 1983, Hilliard & Burn 16754

(PRE).

EASTERN CAPE. — 3029 (Kokstad): Griqualand East, Malowe

Mountain, (-BD), Feb. 1885, Tyson 1047 (PRE). 3129 (Port St Johns):

Umtakatyi River, head of Umtakatyi River N of Libode, (-CC), 16

Sept. 1947, Acocks 13809 (PRE); On road from Tabankulu to Lusiki-

siki, near the Florseshoe, (-AB), 22 June 1995, Phillipson, Bold &

Cloete 4326 (PRE). 3226 (Fort Beaufort): Stockenstromm, Katberg,

(-BC), 1898, Sim 19812 (PRE); Seymour, Hogsback, Auckland For-

est, (-DB), 14 Aug. 1943, Giffen 838 (PRE); Amatole Mountains, bot-

tom of Hogsback Pass, (-DB), 13 Aug. 1984, Phillipson 874 (PRE).

3227 (Stutterheim): Keiskammahoek, Keiskamma River, Boma Pass,

(-CA), 20 Sept. 1942, Acocks 9098 (PRE); King Williams Town, Pirie,

(-CB), 24 Jan. 1936, Taylor 1753 (PRE); Stutterheim, 8.4 miles [13.4

km] from Stutterheim on Keiskamma Hoek road, (-CB), 22 Sept.

1954, Marais 526 (PRE); Keiskammahoek, 11.5 miles [18.4 km] from

Keiskammahoek bridge on road to Debe Nek, (-CC), 18 Mar. 1948,

Story 3374 (PRE); Isedenga Forest near King William’s Town, (-CD),

9 Mar. 1964, Batten. A.U. 1 Plate 54 (NBG); Komga, Valley near

Komga, (-DB), Feb. 1891, Flanagan 599 (PRE). 3228 (Butterworth):

Willowvale, Ngqaqini, (-AD), Nov. 1983, Van Eeden B369 (PRE); Kei

River near Kei River. (-CA), Flanagan 599A (PRE). 3327 (Peddie):

East London, (-BB), 28 Sept. 1909, Rattray 7878 (PRE).

3. Garuleum latifolium Har\’. in Flora Capensis 3:

92 (1865); Hilliard; 516 (1977). Type; [Kwazulu-Natal],

‘Natal, Tongaat River’, Gerrard s.n. K000273564. —

ALUKA image!, lecto., designated here; TCD0002908

(TCD. — ALUKA image!, isolecto.). [K000273564 was

selected as the lectotype over the available syntype,

TCD0002908 as it is of better quality and this specimen

closely resembles G. latifolhtm compared to the TCD

specimen].

Osteospennum calendulaceitm Harv. in Flora

Capensis 3; 440 (1865), horn, illegit. non L.f. (1782).

Type; [KwaZulu-Natal], ‘Port Natal’ Giieinziits s.n.

MEL2357571 (MEL— digital image!, holo.).

Slender, leafy, soft-wooded sub-shrub up to 1.2 m

tall; stem and leaves minutely glandular-pubescent, aro-

matic. Leaves alternate, membranous, sessile or some-

times narrowed to a petiole-like base often with a pair

of stipule-like basal auricles, up to 60 x 30 mm, oblan-

ceolate or oblong in outline, coarsely toothed or more

deeply cut halfway to midrib, lobes toothed, teeth sub-

ulate. Capitula radiate, pedunculate. Peduncles up to

15 mm long arranged in lax terminal corymbs. Involu-

cre and upper part of peduncle loosely woolly at first,

later glandular-hispid. Involucre campanulate. Involit-

cral bracts in 3 rows, imbricate, ovate-linear, pungent,

outer 2. 5-3.0 x 1.5 nim, inner 8. 5-9.0 x l.) mm. Ray

florets numerous, female, fertile, ± 10 mm long, blue or

white; corolla limb up to 6 mm long; corolla tube up to

4 mm long, glandular. Style up to 4 mm long, branched,

linear, style branches 2 mm long, subacute. Ovary up to

1.8 mm long, glabrous, obovate in outline, angled. Disc

florets functionally male, yellow; the corolla limb 3.0-

3.5 mm long, glandular, 5-lobed; the lobes ovate 0.2 mm

long; lower part of corolla 1 mm long. Anthers linear,

minutely caudate, with an ovate apical appendage. Sty4e

up to 5 mm long, branched, linear, style branches 0.1

mm long with stigmatic papillae along margins. Ovar\>

2 mm long. Cypselae 4-5 mm long, glabrous, nar-

rowly obovate, somewhat 3-angled, obscurely winged,

strongly transversely wrinkled and warty, reddish-

brown. Flowering time: February-July. Figure 1C, 2B.

Distribution and habitat. Garuleum latifolium is

recorded from the Lebombo Mountain in the north,

down to Pietermaritzburg in Inanda District, (Figure 5),

up to 656 m. It grows in sandy-loam soil, outcrops, on

the margins of bush clumps. Natal Group Sandstone.

Diagnosis: resembling G. sonchifolium in the alter-

nate, membranous, coarsely toothed or more deeply cut

leaves, but distinguished by the involucral bracts in 3

series, ovate-linear, with the outer bracts much shorter

than the inner.

Hilliard (1977) synonymised Osteospennum calendu-

laceum under Garuleum latifolium.

Additional specimens

KWAZULU-NATAL. — 2732 (Ubombo): Mkuze, top of Mtn, (-

CA), 27 Aug. 1932, Galpin 13374 (PRE); Hlabisa, Makhowe, (-CC),

29 May 1971, Lavranos 8690 (PRE). 2831 (Nkandla): Mhlabatini,

Townlands, (-AB), 1 May 1942, Gerstner 4669 (PRE, NBG). 2832

172

Bothalia 43,2 (2013)

FIGURE 6. — Distribution of Ganileiim tanacetifolium.

(Matubatuba): Hlabisa, Hluhluwe Game Reserve, (-AA), 20 July

1961, Hitchins 54 (PRE). 2930 (Pietermaritzburg): Greater Durban

Metropolitan Area, Hammarsdale area. Hector (ESKOM) Substation

Site, (-DC), 29 Mar. 1995, Ward 12972 (PRE); Inanda, (-DB), Nov.

1883, Wood 160 [PRE).

4. Garuleum tanacetifolium (MacOwan) Norl. in

Studies in the Calenduleae 1: 30 (1943), Osteospermum

tanacetifolium MacOwan: t. 1839 (1889). Type: [East-

ern Cape], ‘Kagaberg and Boschberg’, MacOwan 1382

(GRA— ALUKA image!, holo.; K— ALUKA image!,

PRE!, iso.).

Shrub with simple and leafless stem at the base,

branched above. Leaves alternate, scabrid-glandular,

sessile, very densely crowded at ends of branches,

recLirved-spreading, interruptedly bipinnatipartite with

sharply incised lobes, and pectinate rachis on both sides.

Capitula radiate, solitary, shortly pedunculate. Involu-

cre campanulate, 10 mm in diam. Involiicral bracts in

2-series, more-or-less equal, oblong, slightly acute with

membranous margins; outer ± 7.0 mm long; inner ± 7.5

mm long. Ray florets (10-)12, female, fertile, dark pur-

ple, mauve, pale lilac or white, oblong-elliptic; corolla

limb 12.0-12.5 mm long, narrowly linear; corolla tube

up to 4 mm long, glandular. Style up to 5.5 mm long, lin-

ear, branched, style branches ± 2 mm long. Ovary up

to 2.2 mm long, glabrous, obovate in outline. Mature

cypselae not seen. Disc florets functionally male, yellow,

5-lobed; lobes ovate, 0.2 mm long; corolla limb 2. 5-3.0

mm long, glandular; corolla tube 1 .2 mm long, glandu-

lar. Style 3. 0-3. 2 mm long, branched, style branches ±

I mm long, linear with pollen sweeping hairs. Anthers

linear, minutely caudate, with ovate apical appendages.

Ovary up to 2.5 mm long. Flowering time: May to June.

Figure I E.

Distribution and habitat'. Garuleum tanacetifolium is

endemic to the Sneeuberg Centre of Endemism, (Figure

6), I 200 to I 800 m. It was found along edges of forests

and on mountain ridges, abundant on sandstone soils.

Until now Garuleum tanacetifolium was known from

just two collections on the Boschberg and was treated

as DDT (Data Deficient-Taxonomically problematic)

in conservation assessments (Raimondo et al. 2009).

Recent collections by R. Clark, during field work for a

checklist of Boschberg, have shown that the species is in

fact abundant in the Nardousberg area of the Sneeuberg,

certain south-facing slopes above 1 600 m, and occurs

locally on the Boschberg in montane fynbos. It also

apparently occurs on the Great Winterberg-Amatolas but

has not been collected from there (R. Clark pers. comm.,

January 2013).

Diagnosis: Garuleum tanacetifolium is distinguished

from other species by recurved-spreading, bipinnati-

partite leaves with sharply incised, pectinate lobes, and

heads borne on a short, inconspicuous peduncle.

Additional specimens

EASTERN CAPE. — 3225 (Somerset East); Sneeuberg, Asante

Sana Private Game Reserve, on the mountains behind old town of

Petersburg (-AC), 6 Oct. 2008, Clark & Coombs 703, (GRA, NBG);

Boschberg, edge of forest, (-DA), 1 May 1953, Acocks 16812 (PRE);

Boschberg Nature Reserve, upper slopes of Boschberg along path, in

forest, (-DA), 5 Dec. 2008, Clark, Daniels, Fabricus & le Roux 243

(GRA); Groot-Bruintjieshoogte, Labuschagne’s Kraal, 57 Farm, sum-

mit, (-CB), 11 Dec. 2008, Clark, Daniels, Fabricus & le Roux 452

(GRA).

5. Garuleum woodii Schinz in Bulletin de L’Herbier

Boissier 3: 440 (1895); Hilliard 518 (1977). Type:

[KwaZulu-Natal], valley of Buffalo River near Char-

lestown, [1 525-1 830 m], 6 Dec. 1892, Wood 4840 (Z,

lecto., designated here; BOL, K, PRE!, isolecto.). [Wood

4840 was selected as the lectotype over the available

syntype. Wood 4496 as it is of better quality.]

Viscid, aromatic, well-branched shrublet, up to 600

mm high; stems and leaves rough with glandular hairs.

Leaves sessile, alternate, oblong in outline, up to 50 x 20

mm, once or twiee pinnatisect, the segments very nar-

row, sometimes lobulate, sub-acute, base half-clasping.

Capitula solitary, showy, on long remotely leafy or brac-

teate peduncles terminating upper branches. Involucre

campanulate. Involiicral bracts in ± 4 series, imbricate;

outer shorter 3. 5-^.0 mm long; middle up to 5 mm long;

inner 5. 5-6.0 mm long, all ovate, gland-dotted, apices

obtuse to acute. Ray florets female, fertile, blue or white;

corolla limb 6. 5-7.0 x 2 mm, oblong-elliptic; corolla

Bothalia 43,2 (2013)

173

FIGURE 7. — Distribution of Gandeum woodii.

tube up to 2.8 mm long, glandular, cylindric. St}’le up

to 2.8 mm long, branched, linear, style branches 2.2

mm long. Ovary up to 2 mm long, glabrous, obovate in

outline. Disc florets functionally male, yellow, 5-lobed;

lobes ovate, 0.2 mm long; corolla limb glandular, 2.5-

3.0 mm long; corolla tube 1 mm long, glandular. Stymie

4. 0-4. 5 mm long, branched, style branches 0.2 mm long,

linear with pollen sweeping hairs, which cover style

below point of bifurcation. Anthers linear, minutely cau-

date, with ovate apical appendages. Ovajy up to 2.5 mm

long. Cypselae ± 6 mm long, glabrous, narrowly obo-

vate, somewhat 3-angled, obscurely winged, obscurely

transversely wrinkled, light yellowing-brown. Flowering

time: January-May. Figure IF, 2E.

Diagnosis', distinguished from other species by the

stem and leaves that are rough with glandular hairs, and

ovate involucral bracts.

Distribution and habitat'. Ganileuin woodii is

recorded from Lesotho to Ficksburg and Platberg out-

side Harrismith, northwards along the Drakensberg to

Wakkerstroom, and inland to Suikerbosrand Nature

Reserve in Gauteng, (Figure 7), 1 645-2 745 m. It

occurs on hillsides and mountain slopes along streams

and rivers, on cliff faces and along roads and ridges, on

sandstone or clay soils in stony or rocky situations.

Additional specimens

GAUTENG. — 2628 (Johannesburg); Klipriviersberg, S of Roset-

tenville, (-AC), 8 Apr. 1945, Wasserfatl 912 (PRE); Suikerbosrand

Nature Reserve, Kareekloof, (-CA), 16 Oct. 1971, Bredenkamp 154

(PRE).

MPUMALANGA. — 2630 (Carolina): Kalkoenkrantz, Elands-

berg Farm, (-CC), 6 Nov. 1984, Turner 342 (PRE). 2729 (Volksrust):

Ermelo, Amersfoort, Maquiabe Farm of Koos Wessels, (-BB), 8 Jan.

2000, Siebert & Du Plessis 1225 (PRE); Amersfoort, Sterkfontein

Farm, (-BB), 30 Mar. 1987, Turner 1597 (PRE); Volksrust Highlands,

(-BD), 7 May 1920, Mogg 7504 (PRE). 2730 (Vryheid); Wakker-

stroom, (-AC), Feb. 1917, Beeton HWB 204 (PRE); Wakkerstroom,

Tafelkop Farm, (-AC), 3 Feb. 1994, Jordaan 2763 (PRE).

KWAZULU-NATAL.— 2729 (Volksrust): Charlestown, near Char-

lestown, (-BD), 25 Feb. 1895, Wood 5542 (PRE); Charlestown, near

Charlestown, (-BD), 10 Jan. 1894, Wood 5185 (NH). 2730 (Vryheid):

Utrecht, Naauwhoek, (-AD), 19 Jan. 1961, Devenish 491 (PRE);

Wakkerstroom, Oshoek Farm, on edge of escarpment, (-AD), 21 Feb.

\91^,Smook 1209 (?KE).

FREE STATE. — 2729 (Volksrust): Vrede, Berlin Farm, at Slang

River en route to ascent 16 km NW of Vrede, (-AC), 12 Mar. 1991,

Eckhardt 12 (PRE); Driekoppe Darm; 42 km SE of Vrede on road to

Verkykerskop, base of Aasvoelkop, beginning of kloof, (-CD), 4 Feb.

1987, Relief 1987 (PRE); Verkykerskop, Nonpareil Farm, at foot of

hill, (-CD), 14 Mar. 1990, Saaiman 664 (PRE). 2826 (Brandfort): Near

Roodepoort boundary, Syferfontein Farm, ± 2 miles [3.2 km] SSW of

Greylingstad on road to Rooiwal and Leeuwspruit, northwestern slopes

of koppie, (-DC), 20 Mar. 1967, Scheepers 1591 (PRE). 2828 (Beth-

lehem): Ficksburg. Moolman’s Hoek Peak, summit of peak. (-CA),

30 Oct. 1934, Galpin 13924 (PRE), 2829 (Harrismith): Loskop, (-

AA), 18 Mar. 1970, Siam 420 (PRE); Platberg, (-AC), 14 Dec. 1976,

Jacobsz 3054 (PRE); Platberg, (-AC), 8 Sept. 1983, Roux 1262 (PRE,

NBG); Platberg, One Man Pass, (-AC), 13 Dec. 1976, Hilliard & Burn

9507(PRE); Harrismith, (-AC), Dec. 1912, Thode 2895 (PRE).

LESOTHO. — 2927 (Maseru): E of Maseru, (-AD), 25 Oct. 1963.

Cooley & Kofler 10926 (PRE); Makheka, Roma Valley, on top of

Makheka, (-BC), Sept. 1974, Schmitz 4190 (PRE); Mafeteng. Thaba

Tsoeu, (-CD), 10 Mar. 1917, Dieterlen 1273 (PRE), 2928 (Maraka-

beis): Monia Hania, hillslope, (-CA), 10 Jan. 1954, Jacot-Guillarmod

1711 (PRE); Orange River Valley, 26 km from Taung between Mashai

and Sehonghong, (-DB), 1 Dec. 1977, Killick 4294 (PRE). 2929

(Underberg): Mokhotlong, (-AC), Mar. \9A9, Jacot-Guillarmod 1136

(PRE); Mokhotlong, (-AC), 27 Feb. 1949, Compton 21539 (PRE,

NBG).

6. Garuleum bipinnatum (Thiinb.) Less., Synopsis

generum compositarum: 194 (1832); DC.: 309 (1836);

Harv.: 92 (1865); Norl.: 377 (1977). Osteospermiim

bipinnatum Thunb: 167 (1800); Thunb.: 717 (1825);

Juel: 717 (1918). Type; [Northern Cape], Hantam, Thiin-

berg s.n. UPS-THUNB20814 (UPS-THUNB— micro-

fiche!, holo.).

Dimorphotheca multifida DC.: 73 (1838);

Harv.: 422 (1865). Type: [Eastern Cape], ‘Albany’, Eck-

lon & Zevher (G-DC — microfiche!, lecto., designated by

Norlindh: 378 (1977).

Aromatic, viscid, subshrub, 0.6-1. 5 m tall. Stems

arising from woody rootstock, up to 3 cm in diam.,

erect, usually branched, striate or slightly furrowed,

glandular. Leaves alternate, glandular, about 10-40

mm long, bipinnatipartite. Capitiila radiate, peduncu-

late, solitary at ends of stem and branches or arranged

in loose corymbs. Peduncles vary greatly in length, up

to 120 mm long, and have a few small, dentate or entire

bracts. Involucre campanulate 5-8 mm high, ± 10 mm

broad when pressed. Involucral bracts in 2 or 3 series,

imbricate, ± glandular, outer bracts narrowly lanceo-

late, ± 3. 0^.0 mm long, inner bracts narrowly oblong-

lanceolate, ± 5. 0-6.0 mm long, with whitish-scarious

margins and often violet tips. Ray florets female, fertile,

strap shaped, varying a lot in colour, including white,

blue, mauve, purple and violet, and are about as long as

involucres; corolla limb up to 10-15 x 3 mm, elliptic;

corolla tube up to 3 mm long, glandular, cylindric. Style

up to 3 mm long, branched, linear, style branches 2 mm

long, subacute. Ovaty up to 2.1 mm long, glabrous, obo-

vate in outline, angled. Cypselae: compressed triangular-

obpyramidal, ± 4. 0-5.0 mm long, glabrous. Disc florets

bisexual, yellow and about same size as involucres;

corolla limb 3 mm long, widening upwards, glandular,

5-lobed; lobes ovate, 0.2 mm long; lower part of corolla

tubular, 1.8 mm long, glandular. Style up to 5 mm long,

branched, curved, bulbous at the base, style branches 2.5

mm long with stigmatic papillae along sides. Anthers 2

174

Bothalia 43,2 (2013)

FIGURE 8 — Distribution of Garulewn bipinnalum.

mm long, linear, with an ovate apical appendage, cau-

date. Ovan> 3 mm long. Cvpselae: outer fertile, obovate

and slightly emarginate at the tip or almost obcordate,

marginal wing with stiff, thickened margin, up to 5.0 x

3.5 mm, inner sterile narrowly obovate, 5-6 x 2 mm,

pellucid. Flowering time: throughout the year. Figure

ID, 2C, 6.

Distribution and habitat: Garuleum bipinnatum

occurs from Fort Beaufort Division in Eastern Cape

westwards through the Little Karoo to Montagu in West-

ern Cape and Vanrhynsdorp in Northern Cape (Figure

8), at altitudes of 305-1 220 m. The species grows in

various substrates and habitats, including dolerite rocks,

clay soil, shale soil, stony sandy loam soil, stony loam

soil, sandy clay gravel on shale soil, on dry stony kop-

pies [hillocks], mountain slopes and rocky outcrops,

in dry stream-beds, on floodplains. Karoo plains and

river banks, in xeric succulent thicket, false macchia,

spekboom veld and broken karroid veld.

This species is palatable to livestock and is aromatic.

The flowers have a range of colours from white to blue,

mauve, purple and violet and has horticultural poten-

tial as it is easily cultivated (Swelankomo 2010). The

roots were used as snake bite remedy among early set-

tlers. Farmers have also used a decoction or a tincture

of it in various diseases of the chest, and in the Cape the

root has been used in preparing a mouth wash (Watt &

Breyer-Brandwijk 1962).

Diagnosis: Garuleum bipinnatum is the only spe-

cies where both ray and outer disc florets develop fertile

cypselae; the leaves are twice-pinnate, decreasing in size

and degree of lobing towards the inflorescence; inner

bracts have whitish-scarious margins and often violet

tips, and the plant is glabrous. It can be easily confused

with Garuleum schinzii, but in G. schinzii the fruits

develop from ray florets only, and the plant is glandular-

hispid.

Additional specimens

NORFHERN CAPE.— 3118 (Vanrhynsdorp): Vanrhynsdorp, (-

DA), 25 .luly 1941, Esterhuysen 5286 (PRE). 3119 (Calvinia): Koker-

boomkop, (-AB), 24 Aug. 1926, Acocks 18969 (PRE); Vanrhynsdorp,

Van Rhyns Pass, (-AC), 25 July 1941, Compton 11124 (PRE); Nieu-

woudtville Reserve, (-AC) 25 July 1983, Peny & Snijman 2171

(PRE); Vanrhynsdorp, Van Rhyns Pass, (-AC), 25 July 1941, Ester-

huysen 5286 (PRE); Hantam Mountains, (-BC), Thiinberg PRE 44089

(PRE); Niewoudtville Wildflower Reserve, W-facing hill at circle

route hiking trail, (-AC), 3 Sept. 2007, Hitchcock. Cowell & Bennett

MSBP 3418 (NBG); Vanrhynsdorp, halfway down Vanrhyn’s Pass, (-

AC), 8 Sept. 2006, Hetme 4250 (NBG). 3220 (Sutherland): 17 km N

of Matjiesfontein next to national road, (-BA), 13 Mar. 1981, Van Wyk

513 (PRE). 3221 (Merweville): Fraserburg, Layton, Springbok camp,

(-BB), 20 June 1985, Shearing 1033 (PRE).

WESTERN CAPE. — 3123 (Victoria West): Richmond, 36 km from

Richmond to Beaufort West, on N1 road near Rietfontein West, (-DA),

28 Aug. 1995, Rodriguez-Oubina & Cruces 2115 (PRE); Murraysburg,

(-DD), Dugmore 2060 (PRE). 3221 (Merweville): Fraserburg, Layton,

Eselrand, SE of mountain, (-BB), 28 May 1967, Shearing 89 (PRE);

Prince Albert, 10 km beyond Prince Albert road on road to Merweville,

(-DC), 7 Dec. 1976, Venter 1430 (PRE). 3222 (Beaufort West): Bul-

water P.O.; 20 miles [32 km] NW of Bulwater P.O., (-BC), 17 Mar.

1948, Acocks 14148 (PRE); Karoo National Park, (-BC), 6 Sept.

1983, Braack 63 (PRE); Prince Albert, Klein-Waterval farm, ± 2 km

before farmhouse on road from Seekoegat, (-CD), 7 Oct. 1983, Relief

6 Reid 193 (PRE). 3320 (Montagu): Laingsburg, Whitehill ridge, (-

BA), 20 Sept. 1943, Compton 14904 (PRE); Josephskraal, (-BA), 7

July 1966, Van Breda 2127 (PRE); 17 km N of Matjiesfontein next to

national road, (-BA), 13 Mar. 1981, Van Wyk 513 (NBG); Laingsburg,

7 km from Matjiesfontein on N1 road to Laingsburg, (-BB), 27 Aug.

1995, Rodriguez-Oubina & Cruces 2107 (PRE); Swellendam, Barry-

dale near Barrydale at Meintjes Kraal, (-DC), Sept. 1913, Muir 1037

(PRE); Swellendam, Barrydale, Meinjieskraal, (-DC), 20 Sept. 1913,

Muir 9426 (PRE); Warmwatersberg, I mile [1.6 km] NE of Warm-

watersberg Hot Springs, (-DD), 29 June 1948, Acocks 14564 (PRE).

3321 (Ladismith): Laingsburg, (-AB), 24 Jan. 1941, Esterhuysen PRE

44105 (PRE); Seweweekspoort to Laingsburg near Vleiland farm, (-

AC), 25 Sept. 2004, Goldblatt & Porter 12560 (NBG); Prince Albert,

Bosluiskloof, between 7 Weeks Poort & Prince Albert, (-AD), 16 July

1954, Lewis SAM 68695 (PRE); Laingsburg, Bosluiskloof Pass, (-BC),

14 Sept. 1955, Leistner 246 (PRE); Karoo South, Gamkapoort Nature

Reserve, Witpoort, (-BC), 4 Nov. 1982, Cattell & Cattell 220 (PRE);

Rietfontein Farm, N slope of hill 2.4 km S of bam taking right turn just

before second gate, (-CA), 26 Aug. 1982, Laidler 339 (PRE, NBG);

Little Karoo, Noukloof Nature Reserve, 1.9 km from dam on eircular

drive, (-CA), 6 July 1982, Laidler 59 (PRE, NBG); Ladismith, (-CB),

22 Mar. 1976, Hugo 193 (PRE, NBG); Oudtshoom, Gamka Mountain

Reservoir, (-CB), 17 May 1982, Cattell & Cattell 19 (PRE); George,

Gouritzrivier; (-DC), Dec., Ecklon & Zeyher 232 (PRE); Little Karoo,

Kleinfontein Farm, mid-slope 30 miles [48 km] N of road near top

of small pass, (-DD), 8 Aug. 1984, Laidler 576 (PRE). 3322 (Oudts-

hoom): Prince Albert, (-AA), Sept. 1921, Marloth 10588 (PRE);

Prince Albert Road, (-AA), June 1921, Marloth 10142 (PRE); Prince

Albert in hills, (-AA), Dec. 1905, Bolus 11525 (PRE); Prince Albert

near cemetery to Weltevrede hill N of road, (-AA), 5 July 1975,

Wisura 3396 (NBG); Swartberg Pass, (-AC), Dec. 1943, Stokoe PRE

44094 (PRE); De Rust; P.M.K. le Roux’s Farm, 3 miles [4.8 km] E of

De Rust, (-DA), 19 Oct. 1970, Dahlstrand, 1461 (PRE).

EASTERN CAPE. — 3224 (Graaff-Reinet): Aberdeen, S of Arber-

deen, (-CA), Mar. 1929, Dyer 1940 (PRE); Kendrew, (-DA), Mar.

1929, Dyer 1953 (PRE); between Graaff-Reinet and Jansenville, ± 50

km N of Jansenville, (-DA), 1 June 2005, Meyer 4451 (PRE); GraatT-

Reinet, (-DA), 15 Mar. 1930, Galpin 1151 (PRE); Jansenville, (-DC),

5 Dec. 1947, Compton 20320 (NBG). 3225 (Somerset East): Cradock,

Cradock Mountains, (-BA), 19 Nov. 1953, Henrici 4575 (PRE). 3226

(Fort Beaufort): between Fort Beaufort & Grahamstown, (-DC), 18

July 2001, Koekemoer 2077 (PRE); Fort Beaufort, (-DC), 21 Feb.

1938, Britten 6632 (PRE). 3326 (Grahamstown): Albany, Fish River,

between Breakfast VIei & Grahamstown, (-BB), 1 May 1955, Lewis

SAM 68531 (PRE); Albany, (-BC), May 1928, Dyer 1536 (PRE).

7. Garuleum pinnatifidum (L'Herit.) DC., Pro-

dromus 5: 309 (1836); Harv.: 92 (1865). Osteosper-

mum pinnatifidum L’Her.: 11 (1785); Thunb.: 167

(1800); Thunb.: 717 (1825). G. viscosum Cass.: 172

(1819), nom. illegit. pro O. pinnatifidum L’Her. Type:

‘Cap. Bonae Spei’, without precise locality or col-

lector (G-DC — microfiche!, lecto., designated here).

[L’Heritier’s herbarium was acquired by A.P. de Can-

Bothalia 43,2 (2013)

175

FIGURE 9. — Line drawing of Gani I eiim pinnatifidum. A, habit; B, leaf; C, ray floret; D, disc floret, Hanekom 1954; E, fruit, Zietsman & Zietsman

329. Scale bar; A, 1 cm; B, 0.5 cm; C-E, 0.25 cm. Artist; Gillian Condy.

176

Bothalia 43,2 (2013)

FIGURE 10, — Distribution of Garuleum pinnatifidum.

dolle after his death (Stafteu & Cowan 1981). Of the

four specimens labeled O. pinnatifidum in De Can-

dolle’s herbarium, the two left hand specimens are from

L’Heritier’s herbarium: the first bears a label to that

effect and the second the appropriate locality label. Both

are thus syntypes. We designate the right hand specimen

as lectotype as it bears the correct locality label, is the

most complete, and closely matches the illustration of

the species published by L’Heritier (1785) in the proto-

logue.

Osteospermiim caendeum Jacq.: 78 (1786);

Aiton: 215 (1789); Harv.: 92 (1865) [as O. coeriileum

Alton], Lecto: Illustration in Jacq. leones plantatum rari-

orum 1: plate 179 (1781-1786), designated here.

Aromatic, viscid, much branched shrub, 0.3-1. 5 m

tall. Stem thick, terete, leafy throughout, dividing into

flower-bearing peduncles above. Leaves alternate, ses-

sile, pinnate, minutely villous and ciliate ± 50 mm long;

pinnae oblong, irregularly serrate, somewhat obtuse,

diminishing in size. Capitida radiate, pedunculate.

Peduncle solitary. Involucre campanulate, up to 10 mm

long. Jnvolucral bracts in 2-series, more-or-less equal,

thick, acute; outer 3.5-A.O x 1 mm; inner 4. 0-5.0 x 1

mm. Ray florets ± 18, female, fertile, strap-shaped, blue,

mauve; corolla limb up to 9 x 3 mm, elliptic; corolla

tube 2. 8-3.0 mm long, glandular, cylindric. Style up to 3

mm long, linear, branched, style branches 2.2 mm long,

sub-acute. Ovary up to 2.1 mm long, glabrous, obo-

vate in outline. Disc florets functionally male, yellow,

5-lobed; lobes ovate, 0.2 mm long; corolla tube very

hairy. Style 3. 5^.0 mm long, branched, style branches

0.2 mm long, linear with pollen sweeping hairs, which

cover the style below the point of bifurcation for the

same length as that of the branches. Anthers minutely

caudate, linear with ovate apical appendages. Ovary

up to 2 mm long. Cypselae 3. 0-3. 5 mm long, glabrous,

3-ribbed, narrowly obovate and the surface is rugose and

papillate. Flowering time: October-May. Figure IG, 2F

& 9.

Distribution and habitat: Garuleum pinnatfidum

is recorded throughout the Great Karoo, from Graaff-

Reinet to Cradock and into the Free State as far north as

Bloemfontein, Thaba Nchu and Rouxville (Figure 10),

765-1 705 m. The species grows on mountain and hill

slopes, along the road, in loamy and sandy soils in grass-

land and False Karoo.

Diagnosis: distinguished from other species by hav-

ing 1 -pinnate leaves and firm involucral braets; covered

in glandular-scabrid hairs.

Additional specimens

FREE STATE. — 2826 (Brandfort); Bloemfontein, Glen Agriculture

College, (-CD), 22 Mar. 1951, Mosterl 9 (PRE); Bloemfontein, Glen

Agriculture College, (-CD), 18 Mar. 1985, Zietsman & Zietsman 329

(PRE). 2827 (Senekal): Winburg, Willem Pretorius Game Reserve,

(-AC), 10 Apr. 1962, Leistmr 2998 (PRE); Willem Pretorius Game

Reserve, (-AC), 20 Jan. 1972, Midler 835 (PRE); Doomkop, (-DD),

2 Dec. 1931, Goossens 730 (PRE). 2925 (Jagersfontein): Edenburg,

Bethanie, (-DB), 1 Feb. 1996, Peyper 2013 (PRE). 2926 Winterval-

ley, N of Bloemfontein underneath trees below S slope of koppie, (-

AA), 26 Mar. 1968, Midler 204 (PRE); Thaba Nchu, Adelaide, (-BB),

4 Dec. 1963, Roberts 2911 (PRE); Thaba Nchu, Dakpoort Flats, (-

BD), 4 Dec. 1963, Roberts 2923 (PRE); Dewetsdorp, (-DA), 15 Apr.

1950, Steyn 926 (NBG). 2927 (Maseru): Wepener, (-CA), 29 Jan.

1945, Acocks 11169 (PRE). 3026 (Aliwal North): Gryskop, 24 km N

of Rouxville on slope of hill, (-BB), 8 Feb. 1986, Welman 719 (PRE).

EASTERN CAPE.— 3124 (Hanover): Middelburg, Carlton hills,

(-BD), 27 Mar. 1947, Acocks 13541 (PRE); Richmond, Rhenoster-

fontein, S of Richmond, (-CA), 27 Apr. 1950, Acocks 15835 (PRE);

Richmond, Roelofsfontein, (-DA), 25 Nov. 1972, Hanekom 1954

(PRE); Graaff-Reinet, Blue Gum House Farm, in front of house,

(-DD), 14 Oct. 1983, Retief & Reid 533 (PRE). 3223 (Rietbron):

Willowmore,Van Rooyenskraal Farm, along tributary of Hopsrivier, (-

DD), 11 Oct. 1983, Retief & Reid 441 (PRE). 3224 (Graaff-Reinet):

Moordenaarsrivier; Houd Constant Pass S of farm to river, (-AA), 7

Nov. 1974. Oliver 5268 (PRE); Graaff-Reinet, Wimbledon, (-BC), 8

May 1954, Henrici 4991 (PRE); Graaff-Reinet, 9 miles [14.4 km] N

of Graaff-Reinet, (-BC), 10 Feb. 1955, Adamson D.211 (PRE). 3225

(Somerset East); Cradock, Mountain Zebra National Park, (-AB), 26

Mar. 1952, Biynard 13 (PRE); Cradock, Mountain Zebra National

Park, near dam at house, (-AB), 4 May 1963, Liebeiiberg 7239 (PRE);

Cradock; Mountain Zebra National Park, (-AB), 21 Apr. 1969, Muller

628 (PRE); Cradock, Mountain Zebra National Park, (-AB), 25 Jan.

1966, Liebenberg 7635 (PRE); Cradock, Mountain Zebra National

Park. (-AD), 30 Oct. 1985, Zietsman & Zietsman 1286 (PRE); Moun-

tain Zebra National Park, (-AD), 12 Dec. 2005, Bester 6309 (PRE);

Uniondale, 7 miles [11.2 km] from Uniondale along road from Union-

dale to Willowmore, (-CB), 23 June 1961, Van Breda 1190 (PRE).

3324 (Steytlerville): Steytlerville, (-AD), 9 Nov. 1950, Theron 908

(PRE).

8. Garuleum schinzii O.Hoffm. in Bulletin de

L’Herbier Boisseir 1; 74(1893); Merxm. 139: 71 (1967);

Merxm.: 34 (1954). Type: Namibia, Tiras, 21 Apr. 1875,

Schinz 698 (Z — digital image!, hole.).

Viscid, aromatic, much branched subshrub from

a woody stem, 0. 2-1.0 m high. Stem 3-5 mm thick,

glandular-hispid, ribbed. Leaves alternate, sessile, glo-

bose glands, lower leaves irregularly bipinnatipartite

or almost tripinnatipartite with short linear segments,

upper ones reduced, simply pinnatifid. Capitula radiate,

pedunculate. Peduncles up to 90 mm long, mostly soli-

tary, ribbed, shortly hirsute, bracteate, some hairs with

glandular tips. Involucre hemispherical, up to 15 mm

diameter when pressed. Involucral bracts in 2-series,

imbricate, finely pubescent, narrowly lanceolate, acumi-

nate; outer ones 4. 5-5.0 x 1.0 mm; inner ones 6. 0-7.0 x

1.1 mm with coloured apex. Ray florets female, fertile,

blue; corolla limb up to 15.0 x 1.5 mm, narrowly lin-

Bothalia 43,2 (2013)

177

FIGURE 11. — Distribution of Gandewn schinzii subsp. schinzii •,

G schinzii subsp. crinitnm ▲ .

ear; corolla tube up to 4 mm long, glandular, cylindric.

Sty’le 4. 5-5.0 mm long, branched, linear, style branches

2.2 mm long. Ovaiy up to 2.2 mm long, glabrous, obo-

vate in outline. Disc florets functionally male, yellow,

5-lobed; lobes ovate, 0.2 mm long; corolla limb 4. 0^.2

mm long; corolla tube 2. 5-3.0 mm long, glandular. Style

3. 8^.0 mm long, bifid bearing sweeping hairs all way

to below division but lacking papillae, linear. Anthers

minutely caudate, linear with ovate apical appendages.

Ovaiy up to 2.0 mm long. Cypselae 4.0 mm long, gla-

brous, slightly curved, three-angled, obpyramidial,

strongly rugose. Flowering time: throughout the year.

Diagnosis', distinguished from other Gandewn spe-

cies by its bipinnate leaves and loose involucral bracts

covered with glandular-pilose hairs.

Merxmiiller (1967) separated schinzii in into two sub-

species. In Gandewn schinzii subsp. schinzii, the ray

florets are blue, this species prefers limestone, shale and

calcareous soils. It occurs from Bethanien District, Aus

down to Klein Karas (Namibia) and from Langeberg

Range to Barkly West as far south as Belmont (Northern

Cape Province). In Gandewn schinzii subsp. crinitwn,

the ray florets are white; this species prefers dolomite

soils and it occurs from Windhoek and Rehobot as far

south as Bullesport Mountains (Namibia).

Key to subspecies:

la. Ray florets blue; peduncles up to 90 mm long

8a subsp. schinzii

lb. Ray florets white; peduncles 40-60 mm long

8b. subsp. crinitwn

8a. subsp. schinzii

Lower leaves irregularly bipinnatipartite or almost

tripinnatipartite with short linear segments, upper leaves

reduced, simply pinnatifid. Peduncles up to 90 mm

long. Ray florets blue. Cypselae 4.0 mm long, glabrous,

slightly curved, three-angled, obpyramidial, strongly

rugose. Flowering time: throughout the year (Figure IH

&2G).

Distribution and habitat', found in Namibia (Bethan-

ien District, Aus down to Klein Karas) and Northern

Cape Province, from Langeberg Range to Barkly West

as far south as Belmont, (Figure 11), I 069-1 145 m,

occurring on hill slopes, along roads, in valleys, on lime-

stone, shale, calcareous sands and dry riverbeds.

Additional specimens

NAMIBIA. — 2217 (-CC), Windhoek, hills between Windhoek &

Rehoboth. 14 Mar. 1988, Goldblatt & Manning 8797. 2516 (Helm-

eringhausen): Kunjas, W of Helmeringhausen, (-DC), 9 May 1976,

Oliver. Midler & Steenkamp 6478 (PRE); Maltahohe, dry riverbed

near road to Maltahohe, (-DD), 16 Apr. 1949, Kinges 2103 (PRE);

Bethanien, Goais, (-DD), 19 May 1965, Giess 8816 (PRE). 2616

(Aus): Frisgewaagd, (-BA), 3 Apr. 1968, Giess 10274; Keetmanshoop,

Aroab 3 miles [4.8 km] W of Aroab, (-DC), 2 May 1955, De Winter

3368 (PRE). 2718 (Griina); Great Karasberg. Genadendal, S end of

the Great Karasberg slopes in main kloof, (-BC), 5 May 1976, Oliver

& Steenkamp 6306 (PRE); Great Namaqualand, Klein Karas, (-CA),

8 Apr. 1931, Ortendahl 34 (PRE); Klein Karas, (-CA), 2 Aug. 1923,

Dinter 4810 yWE).

NORTHERN CAPE.— 2822 (Glen Lyon): about 24.8 km west of

Olifantshoek on Pearson’s Hunt road in a poort in Langeberge, (-BA),

10 Aug. 2000, Smook 10965 (PRE); Olifantshoek, 24.8 km NW of

town on road to Pearson's Hunt, (-BA), 10 Aug. 2000, Mey’er 2887

(PRE); Langeberg Range, Dunmurray; W slopes of Langeberg Range,

(-BC), 22 Mar. 1920, Pole Evans 87 (PRE); Hay, Floradale, foot of

hills, (-BD), Apr. 1940, Esterhuysen 2325 (PRE); Postmasburg, Klip-

bank, (-BD), 17 Jan. 1958, Repton 4777 (PRE). 2823 (Griekwastad):

Hay, Swartkop, Black Jasper Hills, (-CD), 31 July 1936, Acocks 544

(PRE); Herbert, Campbell, 8.5 miles [13.6 km] NE of Campbell, (-

DC), 27 Sept. 1957, Leistner 914 (PRE). 2824 (Kimberley): Barkly

West, Kransfontein, (-AC), Aug. 1936, Eaton MCG.M.3737 (PRE);

Herbert, Schmidtsdrift, 9 miles NW of Schmidtsdrift slopes, (-CA),

25 Aug. 1961, Leistner & Joynt 693 (PRE); Caerwinning, (-DA), 7

Oct. 1937, Ferrar 31884 (NH). 2922 (Prieska): Marydale, 6.5 miles

[10.4 km] S of Marydale, (-AC), 10 May 1946, Story> 1224 (PRE);

Niekerkshoop, (-BD), Venneiden TRV 23453 (PRE); 15 miles NW

of Prieska, (-DA), 13 May 1961, Schlieben 8793 (PRE); 60 km from

Marydale on Prieska road, (-DA), 8 June 1977, Smook & Harding 719

(PRE); Prieska valley and lower hills, (-DA), 18 Feb. 1920. Btyant

J40 (PRE). 2924 (Hopetown): Kimberley, Belmont, Fort Richmond, (-

AC), Sept. 1950, Waylands PRE 44121 (PRE).

8b. subsp. crinitum (Dinter) Merxm. in Mitteilun-

gen der Botanischen Staatssammlung Miinchen 2: 34.

(1954); Merxm. 139: 71 (1967). G. crinitwn Dinter in

Feddes Repertorium 30: 184 (1932); Merxm. 139: 71

(1967). Type: Namibia, (Hereroland), Auas Mountain,

Dinter 4654 (B, lecto., designated by Merxmiiller: 34

(1954); PRE!, isolecto).

G. bipinnatum auct. Dinter: 308 (1921), non

(Thunb.) Less. (1832).

Leaves pinnatifid, 8-12 x 3^ cm long. Peduncles

40-60 mm long. Ray florets white. Cypselae 3.5 mm

long, glabrous, obpyramidial, bluntly triangular and the

surface is glandular and rugose. Flowering time. Febru-

ary-May. Figure 1 1, 2H.

Distribution and habitat'. Garideum schinzii subsp.

crinitum is endemic to Namibia, from Windhoek and

Rehobot as far south as Bullesport Mountains (Figure

1 1 ), preferring mountain slopes, dolomite soils.

Additional specimens

NAMIBIA. — 2217 (Windhoek): Avis Dam near Windhoek, (-CA),

21 Feb. 1959, Seydel 1715 (PRE). 2317 (Rehoboth): Bergland, Farm

264, (-AC), 19 Dec. 1957, Merxmiiller 826 (PRE). 2416 (Maltahohe):

Naukluft plateau. W of Kapokvlakte, track to Beacon 1949, (-AA), 23

Mar. 1995, Burke 9564 (PRE); Naukluft, (-AB), 16 May 1978, Muller

178

Bothalia 43,2 (2013)

& Tilson 860 (PRJE); Rehoboth, Buellsport Mtn, (-AB), 26 Mar. 1948,

Strey 2305 (PRE).

ACKNOWLEDGEMENTS

My colleagues from the South African National Bio-

diversity Institute, namely P. Herman for guidance and

discussions, Hester Steyn for producing the distribution

maps, and Gillian Condy for kindly preparing the line

drawing. Dr Leistner for the translation of Latin text.

Referees for their valuable comments. Curators and

staff of NH, GRA and NBG herbaria, for making mate-

rial available on loan. The author would also like to

express gratitude to the curator and staff of Melbourne

herbarium and Trinity College Dublin for sending digital

image of type material.

REFERENCES

AITON, W. 1 789. Hortiis Kewensis. Vol. III. Diadelphia-Cn’ptogamia.

George Nicol, London.

BREMER, K. 1994. Asteraceae: cladislics and classification. Timber

Press, Portland, Oregon.

CANDOLLE, A.P. DE. 1836. Compositae. Prodromus 5. Treuttel &

Wiirtz, Paris.

CANDOLLE, A.P. DE. 1838. Compositae. Prodromus 6. Treuttel &

Wiirtz, Paris.

CASSINI. A.H.G. DE. 1819. Description des nouveaux genres Garii-

leum et Phaganalon; par M. Henri Cassini. Bulletin de la Sociele

Philomatique de Paris November: 1 72.

CASSINI, A.H.G. DE. 1819. Dictionnaire des sciences naturelles 18:

163. Ed. F.Cuvier. edn 2. LeNormant, Paris.

CLAASSEN, C. 1970. Asteraceae. Ganileum album. Flowering plants

of Africa. Vol. 40: t. 1593.

DINTER, K. 1921. Index, der aus Deutscli-Siidwestafrika bis zum Jahre

1917 bekannt gewordenen Pflanzenarten. IX. Feddes Reperto-

rium 17: 308

DINTER, K. 1932. Diagnosen neuer Siidwestafrikanischer Pflanzen.

Feddes Repertorium 30: 184.

GOLDBLATT, P. & MANNING, J. 2000. Cape Plants. A conspectus of

the Cape flora of South Africa. Strelitzia 9. National Botanical

Institute of South Africa, Cape Town.

HARVEY, W.H. 1865. Compositae. Flora capensis 3: 44—530.

HERMAN, P.P.J., RETIEF, E., KOEKEMOER, M. & WELMAN, W.G.

2000. Asteraceae. In O.A. Leistner, Seed plants of southern Afri-

ca: families and genera. Strelitzia 10: 101-170. National Botani-

cal Institute, Pretoria.

HERMAN, P.P.J., WELMAN, W.G., RETIEF, E., KOEKEMOER,

M. & NETNOU, N. 2003. Asteraceae. In G. Germishuizen &

N. L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 217. National Botanical Institute, Pretoria.

HILLIARD. O.M. 1977. Compositae in Natal. University of Natal

Press, Pietemiaritzburg, South Africa.

HOFFMANN, O. 1893. Compositae. Garuleum schinzii. In H. Schinz,

Bulletin de L 'Herbier Boissier 1 : 74.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index herbariorum, part I: the herbaria of the World. New York

Botanical Garden, New York.

JACKSON, W.P.U. 1 990. Origins and meanings of names of South Afri-

can plant genera. Pteridophytes, gymnosperms and monocoty-

ledonous angiosperms. University of Cape Town, South Africa.

JACQUIN, N.J.B. 1786. Osteospermum caeruleum. Collectanea ad

Botanicum, Chemiam, Historiam Naturalem. Vol. 1. pg. 78, 79.

Vindobonae.

JUEL, H.O. 1918. Plantae thunbergianae. Uppsala.

LESSING, C.F. 1832. Asteraceae. Garuleum bipinnatum Less. Synopsis

generum compositarum . Duncker & Humblot, Berlin, pp. 1 94.

L’HERITIER DE BRUTELLE, C. 1785. Stirpes novae aut minus cog-

nitae. pp. 11 tab 6.

LINNAEUS, C. VON, f. 1782. Asteraceae. Osteospermum calendu-

laceum L.f Supplementum Plantarum: 386.

MACOWAN, P. 1889. Compositae, Tribe Calendulaceae. Osteosper-

mum tanacetifolium. Hooker 's leones Plantarum 1 9, Plate 1 839.

MERMULLER, H. 1954. Compositen-Studien 4: Die Compositen-Gat-

tungen Siidwestafrikas. Mitteilungen der Botanischen Staatssa-

mmlung Miinchen 2: 34.

MERXMULLER, H. 1967. Asteraceae. Prodromus einer Flora von

Siidwestafiica 139: 71, 72.

MOORE, S. LE M. 1878. Alabastra diversa. 16: 133.

NORDENSTAM, B. 1994. Tribe Calenduleae. In K. Bremer, Asterace-

ae, cladistics and classification: 365-376. Timber Press, Oregon.

NORDENSTAM, B. 2007. XIII. Tribe Calenduleae Cass. (1819). In

K. Kubitzki. The families and genera of vascular plants. VIII.

Flowering Plants-Eudicots, Asterales (eds J.W. Kadereit & C.

Jeffrey): 241-245. Springer, Berlin.

NORLINDH, T. 1943. Studies in the Calenduleae. I. Monograph of the

genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and

Chiysanthemoides. Gleerup, Lund.

NORLINDH, T. 1977. Garuleum subgenera Rutidocarpaea, a mono-

typic subgenus showing achene dimorphism. Botaniska Notiser

130: 377-380.

PFEIFFER, L.K.G. \^14. Nomenclator Botanicus.Nol 1: 1411-1412.

Kassel, Fischer.

RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E„

HELME, N,A. TURNER, R.C., KUMUNDI, D.A. & MANYA-

MA, P.A. 2009. Red List of South African plants. Strelitzia 25.

South African National Biodiversity Institute, Pretoria.

SCHINZ, H. 1895. Compositae: Garuleum woodii. Bulletin de

L ’Herbier Boissier 3: 440.

STAFLEU, F.A. & COWAN, R.S. 1981. Taxonomic literature. A selec-

tive guide to botanical publications and collections with dates,

commentaries and types. Vol. 3.

SWELANKOMO, N. 2010. Garuleum bipinnatum (Thunb.) Less.

http://www.plantzafrica.com/plantefg/garulbip.htm.

THUNBERG, C.P. 1800. Prodromus plantarum capensium 2. Uppsala,

pp. 167.

THUNBERG, C.P. 1825. Flora capensis. Ed. Schultes. Stuttgart, pp.

717.

VAN BREDA, P.A.B. & BARNARD, S.A. 1987. Veld plants. A.9.

Githoutjie. Farming in South Africa, Pamphlet: Winter rainfall

region. Department of Agriculture and Water Supply, Pretoria.

WATT, J.M. & BREYER-BRANDWIJK, M.G. 1962. The medicinal

and poisonous plants of southern and eastern Africa. Living-

stone, Edinburgh.

Bothalia43,2: 179-195 (2013)

The native and naturalised species of Peltocalathos and Ranunculus

(Ranunculaceae: Ranunculeae) in southern Africa

J.C. MANNING'-^and P. GOLDBLAir-^

Keywords: Peltocalathos Tamaru, Ranunculaceae. Ranunculus L., southern Africa, taxonomy

ABSTRACT

The five native and naturalised species of Ranunculus L. and Peltocalathos Tamaru (Ranunculaceae: Ranunculeae) from

southern Africa are review'ed, with full descriptions and nomenclature, taxonomic history, ecological notes, and distribution

data. All species are illustrated. Peltocalathos. a monotypic genus based on R. bauhi MacOwan, has been overlooked in the

regional literature until now. The new name R. dregei J.C. Manning & Goldblatt is proposed as a replacement name for the

later homonym R. meyeri Harv'., non Lowe. We conclude that the poorly known R. capensis Thunb. represents juvenile forms

of the widespread and variable R. tnultifidus Forssk. and we synonymise the name, along with the three trivial epithets R.

pinnatus var. herniannii DC., R. pubescens van glabrescens Burtt Davy and R. pubescens van han’ey’anus Burtt Davy, which

have been overlooked in the recent literature. Based on number (mostly < 40 achenes per head) and size (mostly 1 .2-1 .8 mm

long) of achenes, we conclude that the southern African material of the aquatic species recently re-identified as R. rionii Bag-

ger is more appropriately treated as R. trichophyllus Chaix.

INTRODUCTION

Ranuculaceae are a large family of ± 2 150 spp. in ±

60 genera distributed worldwide, showing a diversity of

floral organisations and fruit types (Tamaru 1993). Sev-

eral classifications of the family have been proposed

over the years, most recently by Tamaru (1995), who

recognised three subfamilies and eleven tribes, based

largely on chromosome base number and carpel and

fruit types, including placentation. Tribe Ranuculeae

DC. of subfamily Ranuculoideae Hutch, includes ± 650

spp. occurring on all continents, and is diagnosed by

carpels with a solitary, unitegmic and usually ascending

ovule, and petals with at least one nectary near the base

(Tamaru 1993, Emadzade et al. 2010). Molecular phy-

logenetic studies suggest that the tribe is monophyletic

but that parallel, adaptive and convergent evolution of

morphological characters is rife, hindering the devel-

opment of a morphology-based classification of the

tribe. The recent generic classification of Ranunculeae

derived by Emadzade et al. (2010) from their molecu-

lar analysis largely corroborates the narrow generic

circumscriptions proposed by Tamaru (1995) [excep-

tions are the inclusion of Aphanostemma A.St.-Hil. and

Gamsoceras Steven in Ramtnciihts but the exclusion of

Coptidium (Prantl.) Beurl. ex Rydb. and Ficaria Guett.

as separate genera], ultimately recognizing 18 genera in

the tribe, of which Ramtncithts L. (500-550 spp.) is by

far the largest. This phylogeny also establishes that the

aquatic members of R. sect. Batrachium DC. are deeply

' Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, Claremont 7735, South Africa. E-mail: J.Manning(@

sanbi.org.za.

- Research Centre for Plant Growth and Development, School of Life

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

XOl, Scottsville 3209, South Africa.

^ B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P.O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blattfS-mobot.org.

nested in the genus in a clade that includes wetland spe-

cies such as R. repens L., affirming that their segregation

at the higher taxonomic levels of genus (Gray 1821) or

subgenus (Gray 1886) is not justified.

Rancunculeae are poorly represented in sub-Saharan

Africa, with only a handful of species of Ramtncithts

recorded as native, mainly from the temperate eastern

highlands of the continent (Oliver 1868; Milne-Redhead

& Turrill 1952; Exell & Milne-Redhead 1960). The

generic diversity is higher in southern Africa, with two

indigenous genera. In addition to Rammcitlus itself, the

subcontinent is home to the recently described endemic

monotypic genus Peltocalathos Tamaru (1992), based

on R. baiirii MacOwan and separated from Rammcitlus

by its peltate leaves and longitudinally ribbed achenes

lacking a sclerenchyma layer in the carpel wall, the ribs

corresponding to the vascularisation of the carpel walls.

In addition to the native taxa, the European Myosunis

minimus L. is naturalised in parts of Western and North-

ern Cape.

The last regional revision of Ranunculus in southern

Africa is that of Harvey (1860), in which six species

were recognised. The application of three of these names

is now known to have been incorrect. Current checklists

for the subcontinent list seven species (including R. bait-

rii, now P. baurii), with R. capensis Thunb. regarded

as a doubtful species and R. plebeiits R.Br. ex DC. as a

doubtful determination (Winter 2006). The description

of the monotypic Peltocalathos for R. baurii has been

overlooked in all of the regional literature.

A modem review of the southern African species

is clearly overdue, both to establish the taxonomic and

nomenclatural status of the native species as well as the

distribution and identification of all species recorded for

the region. Here we review the genera Peltocalathos and

Ranunculus in southern Africa, providing full descrip-

tions and nomenclature, taxonomic history, ecological

notes, and distributions for the five native and natural-

ised species that we recognise in the region.

180

Bothalia 43,2 (2013)

MATERIALS AND METHODS

All relevant types were examined, as well as all her-

barium material from BOL, GRA, NBG, PRE and SAM

(acronyms after Holmgren et al. 1990), the primary col-

lections of southern African material. All species with

the exception of Rammciilus muricaliis were also stud-

ied in the field.

TAXONOMY

Key to genera of Ranimculeae in southern Africa

la. Scapose annuals with linear or linear-oblong leaves; sepals

with spur-like projection at base; petals long-clawed;

receptacle greatly elongated and tail-like in fruit; achenes

with strong dorsal ridge; ovule pendant, anatropous

Myosiirus [not treated further here]

lb. Annuals or perennials with peltate or variously lobed

leaves; sepals without spur-like projection; petals short-

clawed; receptacle not greatly elongated in fruit; achenes

with or without inconspicuous dorsal ridge; ovule ascendant:

2a. Leaves peltate, with finely toothed, reddish margins

and hollow petioles; inflorescence a subscapose

cyme; petals 11-17; achenes ellipsoid, longitudinally

ribbed Peltocalathos

2b. Leaves not peltate, coarsely toothed to pinnate; petals

5 or 6(-10); inflorescence various; achenes ellipsoid

or compressed, smooth, tuberculate or transversely

rugose Rammcidus

Peltocalathos Tamarii in Acta Phytotaxa Geobotanica

43: 139 (1992). Type species: Peltocalathos haiirii

(MacOwan) Tamaru (= Ranunculus haurii MacOwan).

Perennial, rhizomatous herbs, subglabrous. Roots

fleshy, without a taproot. Stems erect. Leaves spiral,

basal, petiolate, stipules adnate to base of petiole and

sheathing; blades simple, peltate, finely toothed. Flow-

ers usually in a scapose, paniculate cyme, rarely soli-

tary and terminal, bisexual, actinomorphic, all parts spi-

ral. Sepals 5, reflexed, caducous. Petals 11-17, yellow,

glossy adaxially, narrowed at base into short claw, with

lunate, nectar-secreting pit on adaxial surface at top of

claw. Stamens numerous. Carpels free, numerous, with

1 basal ovule. Achenes broadly ellipsoid, longitudinally

ribbed, shortly beaked, glabrous, leathery; pericarp with-

out inner sclerenchyma layer.

1 sp., endemic to the eastern highlands of South

Africa, Lesotho and Swaziland.

Ranunculus haurii was established as the monotypic

genus Peltocalathos by Tamaru (1992) on the basis of

its peltate leaves and longitudinally veined and ribbed

achenes lacking a sclerenchymatous inner layer to the

carpel wall. Phylogenetic analysis of plastid and nuclear

gene sequence data of tribe Ranunculeae supports this

decision, placing the species well outside of Ranuncu-

lus s. str. in a clade of small, mainly New World gen-

era, where it is resolved as sister to the southern South

American genus llamacltyas Comm, ex Juss. (Emad-

zade et al. 2010). The strong geographical isolation of

the genera in this clade is presumed to have fostered the

evolution of distinct autapomorphies among its mem-

bers, which must be regarded as either relictual survi-

vors of ancient radiations or as lineages that have failed

to diversify (Emadzade et al. 2010).

1. Peltocalathos baurii (MacOwan) Tamaru in Acta

Phytotaxa Geobotanica 43: 139 (1992). Ranunculus bau-

rii MacOwan in MacOwan & Bolus: 390 (1881); Oliv.:

t. 1503 (1886b); Tolken: t. 1508 (1965). Type: South

Africa, Eastern Cape, ‘in graminosis subhumidus summi

montis Bazija in Terra Kaffrorum (Ngangelizwe)’, 4000'

[1 200m], [Nov.], R. Baur [Ao. 544] (SAM, leeto.!,

designated here; K, isolecto. [2 sheets K000076103 &

5] — Aluka images!]. [The SAM collection is chosen as

lectotype as being most likely the specimen on which

MacOwan based his description and in displaying insect

damage (to the foliage) as reported in the protologue].

Ranunculus cooperi Oliv.: t. 1502 (1886a). Type:

Lesotho, ‘Basutoland’, 1861, T. Cooper 928 (K, syn. [2

sheets K00076106 & 7] — Aluka images!].

Tufted, rhizomatous perennial to 600 mm; stems con-

densed, covered with brown fibrous remains of old leaf

sheaths. Roots few-several, thong-like. Leaves radical,

usually 3 or 4, peltate, glabrous or sparsely villous when

young, petiole 10-450 mm long with stipules adnate and

sheathing for 20^0 mm, hollow, blade orbicular, leath-

ery, laterally folded when young, (5-)30-150(-300) mm

diam., crenulate-dentate with apical tooth largest, dull

dark green adaxially with irregular pale markings along

veins, margins reddish and cartilaginous with callus

at apex of each serration. Flowers (1)2-18 in a pedun-

culate cyme, (18-)20-30 mm diam.; peduncle 2-3 x

as long as leaves, (30-) 100-600 mm long, 1.5-10.0

mm diam. at base, hollow, usually with apical fringe of

short, obtuse hairs at base of sepals; bracts ovate-sub-

orbicular, cucullate, finely or irregularly toothed with

reddish, cartilaginous margins, rarely lowermost leaf-

like and amplexicaul. Sepals (4)5, reflexed, caducous,

ovate to suborbicular, cueullate, 4-6 mm long, glabrous

or obtusely hispidulous beneath towards base, mar-

gins irregularly toothed, yellowish or with reddish mar-

gins. Petals (1 1)12-15(-17), narrowly obovate-cuneate,

8-15 X 2. 5^.0 mm, varnished bright yellow adaxially

but dull and paler beneath, narrowed into short fleshy

claw in basal ± 1 mm, truncate to acute, veins branched,

nectary lunate. Stamens ± 50 in ± 6 series, filaments ±

5 mm long, anthers ± 0.5 mm long. Carpels ± 50-100,

glabrous. Achenes in subglobose head, broadly ellip-

soid, 5-7 X 1, 5-2. 5 mm, longitudinally ribbed, shortly

beaked; receptacle pubescent. Flowering time'. (Sept.-)

Oct.-Dec.(-Jan.). Figure 1.

Distribution and ecology: a montane species endemic

to the southern African Drakensberg, ranging from

the Katberg in Eastern Cape through KwaZulu-Natal,

Lesotho, Swaziland and Mpumalanga, with a single

collection from the headwaters of the Olifants River in

Limpopo (Figure 2).

Peltocalathos baurii is restricted to damp gullies,

streamsides and the base of wet cliffs, from 1 000-

3 000 m.

Diagnosis: a distinctive species recognised by its

peltate leaves with hollow petioles and scapose inflores-

cence of moderately large flowers with numerous petals

and caducous sepals. The leaf blades are suborbicular

Bothalia 43,2 (2013)

181

FIGURE 1 . — Peltocalathos baurii, Naude's Nek, no voucher. A, flowering plant plus detached leaf and inflorescences; B, sepal; C, petal; D, stamen;

E, pistil; F, carpel; G, achene [Dieterlen 883], Scale bar: A, 10 mm; B-G, 1 mm. Artist: John Manning.

182

Bothalia 43,2 (2013)

FIGURE 2. — Distribution of Peltocalathos baurii.

and finely toothed with attractive pale green markings

along the primary veins, conspicuous against the dull,

dark green upper surface.

Plants in sheltered situations tend to be much

larger than those growing in shallower soils or in

more exposed conditions on banks or in alpine seep-

ages. Although the two extremes are striking, we have

encountered a complete range of variants on wet banks

and seepages on Naude’s Nek in the southern Drakens-

berg, ranging from small, solitary-flowered plants with

leaves less than 30 mm in diameter to well-grown indi-

viduals over 300 mm high with multiple flowers and

leaves to 80 mm diam.

Plants collected early in the season at high altitudes

around 3 000 m in the Drakensberg Mtns, most com-

monly on the summit plateau of Mt aux Sources, are

exceptionally dwarfed, with solitary (rarely two) flowers

on scapes 30-70 mm long and leaves either just emer-

gent or with reduced blades 5-20 mm diam. Tolken

(1965), who also remarked on this form, was uncertain

how to treat it. Although these populations may rep-

resent a distinct alpine ecotype, the species as a whole

exhibits such marked phenotypic plasticity that they are

more likely merely extreme morphotypes, especially

since early-flowering individuals from lower altitudes

elsewhere may be similarly diminutive.

History: The species was described in 1881 by

Peter MacOwan (1830-1909), principal of Shaw Col-

lege in Grahamstown and expert on the regional flora.

from a collection made near Baziya, west of Umtata,

by the missionary Leopold Richard Baur (1825-1889).

The collection is undated but was presumably made

after 1873 since it was during this year that Port Eliza-

beth businessman and amateur botanist Russell Hal-

lack (1824-1903) visited Baur and encouraged him to

botanise the area and send his collections to MacOwan.

MacOwan associated Baur’s plants with a collection

from Lesotho made in 1861 by the professional plant

collector Thomas Cooper (1815-1913), who had recog-

nised that his find represented a new species. This col-

lection (BOL, SAM) is the voucher for seeds and rhi-

zomes that Cooper despatched to his employer, W.W.

Saunders of Reigate, for cultivation, where they were

successfully grown and flowered. A brief description

by Kew botanist D. Oliver of the herbarium specimens

pressed from these cultivated plants formed a footnote to

MacOwan ’s description of R. baurii.

With the arrival of a duplicate of Baur’s collection at

Kew in 1885, Oliver lost no time in having both it and

Cooper’s specimens lithographed for Hooker’s leones

plantarum (Oliver 1886a, b), and in describing the lat-

ter under the name R. cooperii. The differences between

the two were slight, ‘chiefly in dimensions and number

of flowers’ (Oliver 1886a) and were followed by the

caveat, or presentiment, that the two taxa might later

have to be reduced to a single species as it was ‘not

improbable that connecting forms may be found’. Both

names [the former sometimes misspelled as ‘'baueriP]

continued to be applied to plants from KwaZulu-Natal

Bothalia 43,2 (2013)

183

for some time (Wood 1907; Bews 1921), but the two

were eventually united (Tolken 1965).

Additional specimens seen

LIMPOPO. — 2430 (Pilgrim’s Rest): Olifants River, 5400' [1 650

m], (-AA), R. Schlechter 3819 (BOL).

MPUMALANGA. — 2430 (Pilgrim's Rest): Graskop Spruit, (-

DD), 16 Dec. 1937, E. Galpin s.n. (BOL, PRE); Graskop, marshy

slopes, (-DD), 22 Nov. 1951, i. Codd 6739 (PRE). 2530 (Lydenburg):

Dullstroom, Caravan Park, (-AC), 21 Nov. 1985, B. Clarke 1460

(PRE); Dullstroom, Farm Driehoek, (-AC), 8 Nov. 2001, A. Hankev

1428 (PRE); Belfast, (-CA), Dec. 1909, J. Williams s.n. (BOL); 7 Dec.

1909, R. Leendertz 7961 (PRE). 2531 (Komatipoort): White River,

Mr Kay’s Farm, (-CC), Oct. 1931, L. Liebenberg 2815 (PRE). 2630

(Carolina): near Carolina, 5600' [1 707 m], (-AA), Dec. 1905, H.

Bolus 11678 (BOL); Ermelo, Nooitgedacht, (-CB), Jan. 1928, M. Hen-

rici 1739 (PRE); Ermelo, Spitskop, (-CB), Nov. 1915, R. Pott 5125

(PRE). 2730 (Vryheid): Wakkerstroom, Farm Oshoek, (-AD), 24 Nov.

1960, N. Devenish 268 (PRE); Kwa-Mandlangampisis, Farm Groot-

hoek, (-BA), 26 Nov. 1980, L. dii Toit 68 (PRE); Piet Retief, (-BB),

Dec. 191 1, T. Jenkins 10972 (PRE).

FREE STATE. — 2828 (Bethlehem): gully behind Three Witches, (-

DB), 20 Dec. 1983, Dove 54 (NBG). 2829 (Harrismith): Swinburne,

Rensburgskop, (-AC), 10 Dec. 1962, M Jacobsz 15 (PRE)

SWAZILAND. — 2631 (Mbabane): Mbabane. Forbes Reef road.

4500' [1 400m], swamp, (-AC), 6 Nov. 1958, R.Compton 28280

(NBG, PRE); Mbabane, Kirkhill, 3500', [1 070m], (-AC), moist

streamsides, 12 Oct. 1956, R. Compton 26067 (NBG, PRE).

KWAZULU-NATAL.— 2828 (Bethlehem): Natal National Park,

(-DB). 13 Dec. 1928, E. Galpin 10164 (PRE). 2829 (Harrismith):

Van Reenen’s Pass. 5-6000' [1 500-1 800 m] (-AD), 18 Dec. 1891,

J.M.Wood 955 (BOL, GRA. PRE, SAM); Van Reenen, waterfall, (-

AD), 25 Nov. 1941, E. Dillon s.n. ABG402/39 (NBG). 2929 (Under-

berg): Highmoor Forest Station, (-BB), stream bank, 15 Jan. 1966.

D. Killick 3628 (PRE); [Kamberg] Farm Culvers, (-BC), Dec. 1923

[71925], F.A. Rogers 28264 (GRA); Giant’s Castle, (-CB), Nov. 1914,

R. Symons 173 (PRE); Bushman’s River Pass, 9000' [2 750m], moist

places on summit. (-CD), Dec. 1891, J. Thode s.n. (NBG); Bushman’s

River Valley, (-CD), 23 Oct. 1907, J. Wylie com. J. Medley Wood

10654 (PRE).

LESOTHO. — 2828 (Bethlehem): Mamalapi, S-slopes, 9000'

[2 750m], (-CD), 28 Dec. 1948, R. Compton 21329 (NBG); Ler-

ibe, slope Mechachaneng, 5-6000' [1 500-1 800 m], (-DA), Jan.

1913, A. Dieterlen 883 (PRE, SAM); slopes of Mt au.x Sources,

7-8000' [2 100-2 400 m], (-DD), 1894, H. Flanagan 2124 (PRE),

2125 (BOL, SAM); Mt aux Sources, 3 100 m, (-DD), Oct. 1897, G.

Mann sub Marloth 2880 (BOL); 9500' [3 000 m], 25 Oct. 1897, A.

Bolus sub Guthrie 4770 (BOL); 21 Nov. 1930, H. Schweickerdt 689

(PRE); 10500' [3 200 m], 3 Oct. 1950, Sidey-2013 (PRE); Sept. 1963,

Trauseld 53 (PRE); Mothae Mtns, (-DD), 8 Jan. 1958, J. Coetzee 816

(PRE). 2927 (Maseru): Maluti Mtns, Makheke’s Pass. (-DB), without

date, R. Staples 179 (PRE). 2928 (Marakabei): Little Bokong Valley, (-

AA), 5 Jan. 1947, /I. Guillarmod 317 (GRA. PRE). Imprecise locality:

‘Basutoland’, T. Cooper s.n. (BOL. SAM).

EASTERN CAPE.— 3027 (Lady Grey): Witteberg, Ben Mac-

Dhui, damp ground under rocks, 9200-9600' [2 800-2 900 m], (-DB),

11 Mar. 1904 (fruiting/sterile), E.E. Galpin 6567 (BOL, GRA, PRE,

SAM). 3028 (Matatiele): Ramatsiliso’s Beacon, moist cliffs, 2 325

m, (-BB), 12 Oct. 1976, M. Boardman 134 (PRE). 3127 (Lady Frere):

Saalboom Nek, (-BA), 15 Jan. 1959, J. Acocks 20205 (PRE); R56 ±

17 km from Elliot to Ugie, moist slopes, (-BD), 3 Nov. 2000, E. Nien-

aber 986 (PRE). 3128 (Umtata): Mjika, above Mhlahlane Forestry Sta-

tion, 1 300 m, (-BC), 8 Sept. 1984, A. Hutchings 1228 (PRE). 3226

(Fort Beaufort): under krantzes, summit of Great Katberg Mtn, (-BC),

Nov. 1884, W.C. Scully 143 (SAM); top of Katberg Pass, 1 720 m, wet

places at base of cliff, (-BC), T. Dold 1246 (GRA); Amatole Mtns,

Elandsberg, 5000' [1 500 m], S-facing cliffs, (-DB), 28 Oct. 1981, P.

Phillipson 432 (PRE).

Ranunculus L. in Species plantarum 1: 548 (1753).

Type species: Ranunculus acris L. (Jarvis 2007).

Ranunculus sect. Batrachium DC.: 232 (1817). Bat-

rachium (DC.) S.F.Gray: 270 (1821). R. subg. Batra-

chium (DC.) A. Gray: 363 (1886). Type species: Ranun-

culus hederaceum L.

Annual or perennial herbs, sometimes aquatic, gla-

brous or pubescent. Roots fibrous to fleshy, usually with-

out a taproot. Stems erect or creeping, sometimes stolo-

niferous. Leaves spiral, cauline and/or basal, petiolate,

stipules adnate to base of petiole and sheathing; blades

simple and usually palmately lobed or pinnatisect (rarely

peltate in R. lyallii from New Zealand), or compound,

trifoliolate or imparipinnate, in aquatic species often

decompound into filifonn or linear segments. Flowers

solitary and terminal or in cymes, bisexual, actinomor-

phic, all parts spiral. Sepals usually 5(3), spreading or

reflexed. Petals [also termed honey-leaves] 5 or 6 (-10),

yellow or white, often glossy adaxially, narrowed at

base into short claw, with nectar-secreting pit on adaxial

surface at top of claw often covered by laterally adnate

scale or flap forming nectary-pocket. Stamens numerous,

rarely 5 or less. Carpels free, usually numerous, with 1

basal ovule. Achenes smooth, tuberculate or transversely

rugose, glabrous or pubescent, with or without persist-

ent style-beak; pericarp with inner sclerenchyma layer

present.

500-550 spp., cosmopolitan but mainly temperate

Northern Hemisphere; 4 spp. in southern Africa, with

3 spp. native and 1 sp. naturalised. Some species are

reported to be poisonous to livestock when fresh (Eich-

ler & Walsh 2007) but Burtt Davy (1926) reported that

the leaves of R. multifidus are eaten in spring and at

other times when grass is scarce.

Key to species

la. Submerged and floating aquatic; leaf blades finely dissected

into linear or hair-like segments; petals white with yellow

claw; achenes transversely rugose, not beaked

R. trichophyllus

lb. Terrestrial but often growing in damp places; leaves vari-

ously simple or compound but never finely dissected; pet-

als yellow; achenes smooth, ribbed or tuberculate/muri-

cate, beaked:

2a. Dw arf, creeping, stoloniferous perennial rooting from

nodes; leaf blades cordate-reniform and crenate-

dentate, mostly 5-20 mm diam.; flowers solitary,

scapose; petals ligulate, acute-attenuate, ± 1 mm

wide, veins unbranched R. dregei

2b. Erect or sprawling, tufted annuals or perennials; flow-

ers mostly in cymes; leaf blades variously lobed or

parted, 15-70 mm long; petals obovate, obtuse, 3^

mm wide, veins branched:

3a. Perennial, ± pubescent; leaves temately pinnate

or bipinnate; achenes smooth or tuberculate,

2. 0-2. 5 mm long, shortly beaked with beak ±

0.5 mm long R. multifidus

3b. Annual, ± glabrous; leaves simple or 3-5-parted;

achenes muricate, 5-8 mm long, strongly

beaked with blade-like beak 2-3 mm long

R. muricatus

Sect. Ranunculus

1 . Ranunculus muricatus L. in Species plantarum 1 :

555 (1753); Eichler & Walsh: 347 (2007). Type: ‘Habitat

in Europae australis fossis & humentibus’, L1NN71 5.66

[LfNN — digital image!, lecto., designated by Lourteig:

184

Bothalia 43,2(2013)

487 (1951)].

[IR. sceleratus sensu Harv.: 6 (1860), non L.

(1753)]

Erect or sprawling annual, mostly 100-200(-500)

min tall, stems simple or branched in upper parts, gla-

brous or pilose with hairs to 2 mm long. Roots fibrous or

narrowly thong-like. Leaves basal and cauline, petioles

of basal leaves 50-150 mm long with stipules adnate

and sheathing for 5-30 mm, glabrous or thinly pilose,

blade suborbicular in outline, unlobed or 3-5-lobed to

± midway, 15-50 mm diam., base truncate to cordate,

coarsely crenate-toothed with callus at tip of each serra-

tion, usually glabrous but sometimes thinly adpressed-

pilose on both surfaces, cauline leaves ± cuneate. Flow-

ers in simple or compound cymes, 10-15 mm diam.

Sepals 5, strongly reflexed, ovate, cucullate, ± 5 mm

long, pilose beneath, yellowish. Petals 5, obovate, 5-8

X 3^ mm, varnished bright yellow above, dull and paler

beneath, narrowed into short claw in basal ± 1 mm,

obtuse, veins branched, nectary pocket-like. Stamens

15-20 in 2 or 3 series, filaments ± 3 mm long, anthers

± 1 mm long. Carpels 8-20, glabrous. Achenes in a glo-

bose or hemispherical head, broadly obovoid or subcir-

cular, 5-8 mm long (inch beak), muricate with numerous

acute tubercles, margin keeled and grooved, beak stout,

blade-like, 2-3 mm long; receptacle pubescent. Flower-

ing time: Sept.-Nov. Figure 3.

Distribution and ecology, native to the Mediterranean

region but adventive in the southwestern Cape, where

it has been recorded from the Cape Peninsula to Klein-

mond and Swellendam (Figure 4), in marshy ground and

riverbanks. It has been collected so far only near settle-

ments and has not been recorded in undisturbed veg-

etation. The species is widely naturalised in Australia

and parts of the continental United States, mainly as a

weed of damp waste places and croplands but occasion-

ally along lake and stream margins in native vegetation

(DiTomaso & Healy 2007; Eichler & Walsh 2007).

Diagnosis: readily distinguished by the mostly sub-

glabrous, simple or 3-5-lobed, orbicular leaves and echi-

nate achenes, 5-8 mm long, with a stout, blade-like or

hooked beak 2-3 mm long.

Ranunculus muricatus was not mentioned by Harvey

(1860) by name [but see discussion below] and the first

documented South African records are almost contem-

poraneous collections from the late nineteenth century,

from Rondebosch on the Cape Peninsula and from Zuur-

braak [Suurbraak], a mission station near Swellendam

established in 1812 by the London Missionary Society.

The species has been collected only sporadically since

then, initially from elsewhere on and around the Penin-

sula but by the middle of the century from Stellenbosch

and Somerset West, and most recently from Kleinmond

and Caledon. It has not been recollected on the Cape

Peninsula since 1942 and does not appear to be invasive.

It is possible that Harvey’s (1860) reference to

Ranunculus sceleratus L. is relevant here. In his account

of the genus in southern Africa, he reported the occur-

rence of R. sceleratus in ditches near Cape Town but

neglected to preserve any specimens. Native to Europe,

FIGURE 3. — Rammculus miiricalus. Camp’s Bay, Saxton 135 (NBG).

Achene, lateral and dorsal view. Scale bar: 1 mm. Artist: John

Manning.

R. sclereratus is a subglabrous annual with lobed and

crenate basal leaves, deflexed sepals, and elongate-con-

ical receptacles bearing numerous small, ± 1 mm long,

ovoid, faintly rugose achenes (Cook 1964). No plants of

R. sceleratus have ever been collected in southern Africa

and the identity of Harvey’s plants remains unclear. A

collection by Ecklon and Zeyher from the banks of the

Zwartkops River listed by them under this name is in

fact R. miiltifidus and although it is possible that Har-

vey (1860) made a similar error we consider this to be

unlikely given his familiarity with that species on the

Cape Peninsula. R. sceleratus is vegetatively much more

similar to R. muricatus than to R. miiltifidus and if con-

fusion occurred, it is more likely to have been between

the former two species, especially without fruits.

Additional specimens seen

WESTERN CAPE. — 3318 (Cape Town): railway at Rondebosch,

(-CD), 5 Sept. 1896, WoUey-Dod 1604 (BOL); Rondebosch, (-CD),

Sept. 1898, H. Bolus 7991 (BOL); Rondebosch Common, (-CD), 9

Oct. 1938, R.S. Adamson 2122 (PRE); Maitland, (-CD), Nov, 1904,

H. Bolus s.n. (BOL); Camps Bay, (-CD), Nov. 1907, W.T. Saxton 135

(NBG); Lion’s Rump, (-CD), Nov. 1924, R. Marlolh 6305 (NBG);

Klipfontein Road, 4 mi [6.4 km] E of Mowbray, (-CD), 12 Oct. 1913,

N.S. Pillans 2157 (BOL); Mowbray, (-CD), Oct. 1915, R. Marlolh

724 7 (PRE); Camp Ground, (-CD), 16 Oct. 1942, R.H. Compton

13909 (NBG); Tygerberg Nature Reserve, under trees near stream, (-

DC), 2 Oct. 1975, J. W. Loubser 3416 (NBG); Stellenbosch, Elsenburg

Agricultural College, (-DD), without date, P. le Roux s.n. (PRE); Stel-

lenbosch, Goedvertrouw, riverbank, (-DD), 23 Sept. 1994, P. Nel & C.

Boucher 414 (NBG, PRE); Banhoekweg, marshy ground, (-DD), 19

Sept. 1946, £)../. Lomv s.n. (NBG). 3418 (Simonstown): Youngsfield,

(-AB), 5 Nov. 1935, R.H. Compton 5955 (NBG); Cape Flats, (-BA),

25 Oct. 1925, R. Young 26414 (PRE); Somerset Strand, (-BB), 14

Nov. 1946, R. Sirey 737 (PRE); Somerset West, garden weed, (-BB),

Bothalia 43,2 (2013)

185

FIGURE 4. — Distribution of Rammcnlus muricatiis, o; R. dregei, •, in southern Africa.

5 Nov. 1949, R.N. Parker 4464 (BOL, NBG). 3419 (Caledon): 1 km S

of Caledon, just E of rifle range, marshy area, 245 m, (-AB), 13 Oct.

2001, N.A. Helme 2300 (NBG); Kleinmond, (-AC), without date, M.P

de Vos 12 (NBG). 3420 (Swellendam): Zuurbraak [Suurbraak], near

watermill, (-BA), 12 Oct. 1897, £. Galpin 3738 {PRE}.

2. Ranunuculus multifidus Forssk. in Flora aegyp-

tiaco-arabica: 102 (1775), non Pursh: 736 (1813), hoin.

illegit.; Milne-Redhead & Turrill: 19 (1952); Exell &

Milne-Redhead; 97 (1960); Cook: 241 (2004). R. for-

skoehlii DC.; 303 (1817), nom. illegit superfl. Type: Ara-

bia. near Taas, Forsskal s.n. (C, holo., not seen).

R. capensis Thunb.: 94 (1800), syn. nov. Type:

South Africa, precise without locality, Thimberg UPS-

THUNB 13126 (UPS-THUNB— microfiche!, holo.).

R. pubescens Thunb.; 94 (1800). Type: South

Africa , without precise locality, Thimbeig s.n. UPS-

THUNB 73756 (UPS-THUNB— microfiche!, holo.).

R. pinnatus Poir. in Lam.: 126 (1804). Type:

‘Les Indes’, Sonnerat s.n. (P-LAM, holo., not seen).

R. pinnatus var. hermannii DC.: 42 (1824),

syn. nov. Type: ‘Cap. Bonae Spei’, Hermann s.n. (G-DC

[000130084], holo. — digital image!).

R. pubescens var. glabrescens Burtt Davy: 342

(1921), syn. nov. Type: South Africa, [Gauteng], ‘Ver-

eeniging Dist., Burttholm, Uitgevallen 197’, 25 Apr.

1918, Bunt Davy 17682 (K [777706076092]— Aluka

image!, holo.).

R. pubescens var. harveyanus [as "harveianus']

Burtt Davy: 109 (1926), syn. nov. Type: South Africa,

[Eastern Cape], ‘Graaf Reinet Div., southern slopes

of the Schneeuwberg [Sneeuberg], Burke s.n. (PRE

[PRE04 1 8732-0]— A\uka image!, holo.).

[7?. plebeius sensu Harv.: 6 (1860), non DC.

(1817).]

[See Perrier de la Bathie (1950) and Exell & Milne

Readhead (1960) for additional synonyms from tropical

Africa]

Tufted perennial, mostly 100-500 but much dwarfed

in exposed situations and up to 1 000 mm in protected

places, developing new shoots at base; flowering stems

erect or sprawling, simple or well-branched in upper

parts, densely or sparsely patent- or adpressed-hirsute

or -pilose below with hairs to 2 mm long and adpressed-

pubescent above, rarely glabrescent or subglabrous.

Roots several to many, thong-like. Leaves basal and

cauline, petioles of basal leaves 20-150(-300) mm long

with stipules adnate and sheathing for 5-30(-40) mm,

usually pilose, sometimes subglabrous, blade ovate

in outline, ternate or ternately pinnate to bipinnatisect

(rarely tripinnatisect) with 1 or 2 lateral pairs of leaf-

lets, (20-)30-90(-130) X (15-)20-70(-100) mm, ulti-

mate segments coarsely and irregularly toothed, with

callus at tip of each serration, usually adpressed-pilose

on both surfaces, rarely glabrescent, cauline leaves pro-

gressively smaller, ultimately subsessile and lanceolate;

186

Bothalia 43,2 (2013)

juvenile leaves suborbicular and lobed to ternate. Flow-

ers in simple or compound cymes, 10-18 mm diam.

Sepals 5, strongly reflexed, ovate, cucullate, 2. 5-5.0

mm long, hispid beneath, yellowish. Petals 5, obovate,

4-9 X 3-7 mm, varnished bright yellow above, dull and

paler beneath, narrowed into short claw in basal ± 1 mm,

obtuse, veins branched, nectary pocket-like. Stamens ±

30 in 2 or 3 series, filaments 1-3 mm long, anthers ± 1 .5

mm long. Carpels 30-50(-70), glabrous. Achenes in an

ovoid head, broadly obovoid or subcircular, 2. 0-2. 5 mm

diam., usually verrucose or tuberculate but sometimes

± smooth, margin keeled and grooved, shortly beaked,

beak acute, ± 0.5-0. 8 mm long; receptacle 2-5mm long,

pubescent. Flowering time: Aug.-Jan. Figure 5.

Distribution and ecology: widespread through south-

ern and east tropical Africa to Ethiopia, Arabia and

Madagascar. In southern Africa, Ranunculus multifidiis

is largely restricted to the more mesic southern and east-

ern parts (Figure 6), from near sea level to 3 000 m, and

is largely absent from the western half of the subconti-

nent apart from some isolated populations in higher or

locally moist situations.

Plants typically occur in moist or damp, open grassy

habitats and floodplains, along streams and bogs, some-

times partially submerged, and in open patches in tem-

perate forest; it is sometimes ruderal in gardens or along

ditches and around dams. The species is self-fertile; pro-

ducing numerous fruits, and has a tendency to become

weedy in damp waste places, suburban gardens and

parklands, especially in the southern and southwest-

ern Cape, where it is commonly associated with human

activity. Local variants are readily established.

Diagnosis: a widespread and variable species, espe-

cially in the degree of dissection of the leaves, which

range from ternate to tripinnatisect, development and

density of the vestiture on the stems and leaves, size of

the flowers, and development of tubercles on the fruits,

which vary from scarcely evident to almost spinulate.

Plants from high altitudes and exposed or trampled situ-

ations are typically dwarfed and compact or almost pros-

trate and those from sheltered locations larger and more

lush, but the species is always readily distinguished from

other sub-Saharan species by its ± appressed pubescent,

ternately pinnate or bipinnate leaves with one or two

pairs of lateral leaflets, and ± tuberculate achenes. The

flowers are small to moderately sized, with reflexed,

hispid sepals and 5 obovate petals, 4-9 mm long.

History: The correct identification of the South Afri-

can material of this species took some time to establish.

The first collections of the species on the subcontinent

were made in the late eighteenth century by Carl Thun-

berg ( 1 743-1 828), who described them under the names

R. capensis Thunb. (1800) and R. puhescens Thunb.

(1800). Subsequently Harvey (1860), in his account

of the genus for Flora capensis, segregated the mate-

rial available to him among three different names: typi-

cal plants with pinnate or bipinnate leaves, including

R. puhescens Thunb., were treated by him, and later by

Oliver (1868), under the name R. pinnatus Poir.; smaller

plants with subsimple radical leaves were treated as R.

capensis Thunb.; and a collection from the Sneeuberg

with ternate leaves and smooth achenes was referred to

the Australian species R. plebeius DC., a mistake that

was later corrected by Exell & Milne-Redhead (1960).

The priority of the name R. multifidus Forssk. (1775)

over R. pinnatus Poir. (Lamarck 1804), under which

name the species was better known in southern Africa,

was established by Milne-Redhead & Turrill (1952).

Collections of subglabrous plants from Mpuma-

langa were described as var. glabrescens by Burtt Davy

(1921). The protologue clearly designates Burtt Daw

17682 (K) as the [holo] type, with Burtt Davy 17164

included among the paratypes, but this was somewhat

confused when Burtt Davy (1926) later identified both

collections as types [syntypes]. This is incorrect. He also

segregated Burke’s Sneeuberg collection [previously

treated as R. plebeius by Harvey (I860)] plus some oth-

ers as his new var. haweyanus Burtt Davy (1926). Both

variants fall within the range of variation of Ranuncu-

lus multifidus and we see no justification for upholding

them. We have seen no authentic material of R. plebeius

from southern Africa.

The identity of Ranunculus capensis Thunb. (1800),

based on a collection of three small plants — two of them

just coming into flower and the third sterile — deserves

further consideration and it was treated as a doubtful

species by Winter (2006). The name was associated by

Harvey (1860) with two or three additional collections

from the Cape Peninsula and Clanwilliam, notably Eck-

lon & Zeyh. En. No. 13 (SAM) from Green Point and the

latter is indeed a perfect match with Thunberg’s type.

After close study we conclude that the species represents

a seedling variant of R. multifidus with juvenile, ternate

leaves. This conclusion was also reached by Adamson

(1950), who suggested that the taxon was a dwarf state

of the variable R. puhescens (now R. multifidus). The

pubescent stems and petioles, the hirsute leaf blades,

the relatively small flowers with reflexed, hispid sepals,

and the pubescent receptacle with small, compressed

achenes with short beak are all consistent with R. mul-

tifidus. Additional collections from Green Point (Ecklon

s.n. SAM13992 and Thode s.n. SAM9260) are perfectly

intermediate with more typical R. multifidus and we

accordingly formally synonymise the name here.

The relationship between Ranunculus multifidus

and R. pinnatus has proven remarkably troublesome

to resolve. Following Harvey (1860), the South Afri-

can material was treated under the name R. pinnatus

(e.g. Wood 1909, Bews 1921) until Burtt Davy (1921)

queried the conspecificity of R. puhescens with R. pin-

natiis. Although evidently of the opinion that the two

were probably the same, the type locality of R. pinnatus,

given by Poiret (1804) as ‘Les hides’, was problemati-

cal to him and he therefore elected to treat the southern

African material under the name R. puhescens until this

could be resolved. Certainly, both Milne-Redhead &

Turrill (1952) and Exell & Milne-Redhead (I960) con-

sidered that the tropical African material was not con-

specific with R. pinnatus Poir., although without explicit

justification, and this opinion is still followed by some

authors (Teketay & Edwards 2000). Perrier de la Bathie

( 1950), in his account of the genus in Madagascar, how-

ever, treated the two as conspecific under the name R.

pinnatus Poir. This remains the current situation until

formally reversed.

Bothalia 43,2 (2013)

187

Additional specimens seen

NAMIBIA. — 1820 (Tarikora): swampy area below Tamso Camp

in Omuramba Khaudum, (-DA), 16 Feb. 1956, B, de Winter & W.

Marais 4687 (PRE). 1917 (Tsumeb): Otavi, (-CB), 30 Nov. 1908,

Dinter 909 (SAM); Otavifontein, (-CB), 3 Feb. 1960, Giess & Smook

10620 (PRE). 1918 (Grootfontein): Grootfontein, (-CA), without date,

Schoenfelder 293 (PRE); Grootfontein, (-CA), 3 Mar. 1960. R. Seydel

2066 (NBG). 2017 ( Waterberg): Waterberg Plateau, (-CA), Dec. 1935,

J. Boss s.n. (PRE); Gross Waterberg, (-CA), 12 Nov. 1947, R.J. Rodin

2590 (BOL, PRE); Okosongomingo, (-CA), 4 Feb. 1911, Dinter 1766

(SAM). Uncertain locality: Okasewa, alluvium along Nossob, 24 Jan.

1913, Dw/e;- 2739 (SAM).

BOTSWANA. — 2525 (Mafeking); Kanye, Majana village along

banks of Kolobeng River, (-BA), 16 Nov. 1948, Hillary & Robertson

572 (PRE).

FIGURE 5. — Ranunculus multifidus. Cape Town, Manning 3405 (NBG). A, flowering plant; B, seedlings; C, sepal; D, petal; E, stamen; F, pistil; G,

carpel; H, receptacle; I, achene, lateral and frontal view. Scale bar: A, B, 10 mm; C-E, G-1, 1 mm; F, 2 mm. Artist: John Manning.

Bothalia 43,2 (2013)

FIGURE 6. — Distribution of Rammciilus multifidus in southern Africa.

LIMPOPO. — 2229 (Waterpoort); N of Fogwells, (-DD), 23 Dec.

1935, Smuts & Gillett 3162 (PRE). 2230 (Messina): Venda, Smokey

Irrigation Scheme, Nzhelele (Dzanani), (-CB), E. Netsgingane 632

(PRE); Schikunda-Milema, along Mutail River, (-DA), 7 Oct. 1981,

A. Van Wyk & G. Theroii 4963 (PRE). 2329 (Polokwane): Pieters-

burg [Polokwane], (-CD), Feb. 1904, H. Bolus 11035 (BOL). 2330

(Tzaneen): 5 km from Elim on road to Bandolierskop, (-AA), 6 Nov.

1985, G. Germishuizen 2446 (PRE); Zoutpansberg, Tshakoma, (-

AB) , Nov. 1931, A. Obenneyer 993 (PRE); Woodbush, (-CC), with-

out date, T. Jenkins 10450 (PRE). Woodbush, Pypkop, (-CC), 29

Dec. 1944, A. Mogg 17449 (PRE). 2428 (Nylstroom): Naboomspruit,

Mosdene, Nylsviei, (-DA), 29 Oct. 1919, E. Galpin 465 (PRE). 2429

(Zebediela): Potgietersrust, (-AA), 2 Nov. 1908, R. Lendertz 6207

(PRE); Sekukuniland, Motoka, Farm Ironstone, (-DD), 9 Jan. 1939,

Mogg & Barnard 808 (PRE); Farm Schoonraad, along water furrows

and vleis, (-DD), without date, W. Barnard 14 (PRE).

NORTH-WEST.—2526 (Zeerust): Zeerust, (-CA), Jan. 1912, R.

Pott 4186 (PRE); Marico Dist., Wonderfontein, (-CC), 18 Nov. 1907,

./. Burtt-Davy 7239 (PRE). 2527 (Rustenburg): Beestkraal, (-AD),

Dec. 1908, T. Jenkins 6946 (PRE); Rustenburg, (-CA), Dec. 1918,

F. Rogers 22354 (PRE); Rustenburg Kloof, (-CA), 27 Dec. 1949, M.

Sleyn 879 (NBG). 2625 (Delareyville): Mafeking Dist., Setlagoli, (-

AC) , 13 Nov. 1911, J. Burtt-Davy 11047 (PRE). 2626 (Klerksdorp):

Klerksdorp, (-DC), Feb. 1918, L. Kretzschmar s.ii. (PRE). 2627

(Potchefstroom): Potchefstroom, (-CA), 21 Mar. 1910, R. Leendertz

7993 (PRE); abandoned park near Mooivallei suburb, (-CA), 31 Oct.

2011, D.M. Komape, LI. Mabe c6 S.J. Siebert KMS125 (NBG); Klip-

drifl E of Potchefstroom, (-CB), 4 Jan. 1935, J. Theron 1115 (PRE);

Vereeninging, (-CB), 20 Feb. 1917, J. Burtt Davy 17164 (BOL).

2724 (Taung): Taung, Kaap Plateau, 1 200 m, riverine, (-DA), 16

Dec. 1977, Peeters, Gericke & Burelli 448 (PRE). 2725 (BloemhoO;

Wolmaransstad, Boskuil, (-BD), 9 July 1929, J. Sutton 173 (PRE);

Bullfontein, Oppcrmansdrif Dam, I 220 m, (-CB), 14 Dec, 1974, T.

O'Connor 88 (PRE); Christiana, (-CC), 22 Dec. 1911,7. Burtt-Davy

s.n. (PRE).

GAUTENG.- 2527 (Rustenburg): Hennops River, (-DD), 10

Nov. 1951, L. Pimser 1628 (PRE). 2528 (Pretoria): Koedoespoort,

(-CA), 19 Oct. 1905, R. Leendertz 646 (BOL, PRE); Groenkloof Val-

ley, (-CA), 1 Nov. 1917, 7. Pole-Evans 167 (PRE); Fountains Valley,

(-CA), 22 Oct. 1928, 1. Verdoorn 514 (PRE); Muckleneuk, (-CA), 10

Oct. 1930, A. Goossens 27 (PRE); Colbyn Nature Reserve, (-CA), 14

Nov. 2000, J. Meyer 3084 (PRE); Pretoria, Silverton, Moreleta River,

(-CB), 6 Nov. 1980, A. Balsinhas 3494 (PRE). 2627 (Potchefstroom):

Roodepoort, Botanic Garden, (-BB), 19 Nov. 1982, C.M. Behr 149

(NBG); Vereeniging, Leeuwkuil Pasture Research Station, (-DB), 5

Nov. 1935, R. Stoiy 44 (PRE). 2628 (Johannesburg): near Kempton, (-

AA), Nov. 1926, C. Smith 3505 (PRE); Crown Mines, (-AA), 14 Nov.

1961, A. Lucas 35242 (PRE). Without exact locality: Johannesburg,

Dec, 1908, R. Leendertz 6856 (PRE).

MPUMALANGA. — 2430 (Pilgrims Rest): Shiluvane, (-AB),

withou date, H. Junod 25202 (PRE); Mount Sheba, damp forest floor

along streamlet, 6400' [1 900 m], (-DC), 29 Nov. 1980, J.P. Kluge

2319 (NBG, PRE). 2431 (Acomhoek): near river at Skukuza, (-DC),

Nov. 1833, E. Cholmondeley s.n. (PRE). 2529 (Witbank): Mid-

delburg, Doornkop, (-CB), 23 Oct. 1968, C. du plessis 819 (PRE).

2530 (Lydenburg): Spekboom River, 4 mi [6.4 km] N of Lydenburg,

stream bank, (-AB), 19 Nov. 1933, R. Young 464 (PRE); Dullstroom,

(-AC), 27 Nov. 1980, B. Drews 175 (PRE); 16 Nov. 1984, H. Cam-

eron 390 (PRE); SchoemaniTs Kloof, (-AD), 14 Nov. 1933, R. Young

319 (PRE); Ml Anderson, W of Anderson Pass, (-BA), 25 Dec. 1932,

Smuts & Gillett 2447 (PRE); Schagen, (-BD), Dec. 1934, 7. Lieben-

berg 3279 (PRE); Rosehaugh, (-BD), 10 Jan. 1938, A. Mogg 13670

(PRE); Witkiip, in viei, (-BD), 3 Jan. 1974, J. Kluge 405 (PRE);

15.5 mi [25 km] N of Belfast, (-CA), 16 dec. 1956, 7. Prosser 2035

(PRE). 2531 (Komatipoort): Barberton, Plaston, (-AC), Nov. 1931,

W. Holt 123 (PRE); Moodies, (-CC), Sept. /Oct. 1889, E. Galpin 587

(PRE); Barberton, (-CC), Nov. 1909, A. Williams 10446 (PRE). 2629

(Bethal): Bethal, (-AD), Dec. 1910, R. Pott s.n. (BOL); Ermelo, (-

DB), 18 Dec. 1949, M. Steyn 848 (NBG). 2630 (Carolina): Ermelo, (-

CD), 2 Nov. 1926, M. Henrici 111 (PRE). 2729 (Volksrust): Volkstrusl,

(-BD), Jan. 1911, T. Jenkins 10687 (PRE). 2730 (Vryheid): Utrecht,

(-AB), Dec. 1915, A. Wahl 15387 (PRE); Wakkerstroom, (-AB), 20

Nov. 1926, Watt (& Brandwijk 1617 (PRE).

FREE STATE. — 2627 (Potchefstroom): Sasolburg, 1 463 m, {-

DD), 14 Oct. 1977, N. Kroon 43 (PRE). 2727 (Kroonstad): Hillbrow,

Bothalia 43,2 (2013)

189

(-BD), 21 Jan. 1931, A. Goossens 492 .(PRE); Kroonstad, (-CA),

without date, D. Chennells 81 (BOL); Kroonstad, near Vais River

and Bloemspruit, (-CA), Oct.-Nov. 1927, J. Pont 201 (PRE). 2728

(Frankfort): Farm Rietspruit, 29 km from Frankfort on road to Vrede,

(-BC), 28 Jan. 1983, E. Relief 1095 (PRE). 2828 (Bethlehem): Beth-

lehem, (-AB), 3 Dec. 1919, E. Phillips 3080 (GRA, PRE); Fouries-

burg, (-CA), 8 Jan. 1918, G. Potts 3245 (PRE); Golden Gate National

Park, Oshoek, (-DA), 10 Dec. 1988, Gertenbach & Groenewald 8838

(PRE); Witzieshoek, (-DB), Oct. 1909. J. Thode s.n. (PRE). 2829

(Harrismith): Flarrismith Botanic Garden, 1 880 m, (-AC), 30 Oct.

1970, A. van der Zeyde 459 (NBG); small dam E of garden estate, (-

AC), 2 Dec. 1974, A. van der Laarse s.n. (NBG); Van Reenen’s Pass,

(-AD), 13 Dec. 1931, M. van Wyk 107 (PRE); Swinburne. (-AD), Nov.

1961, M. Jacobsz 20 (PRE). 2925 (Jagersfontein): Fauresmith, (-CB),

25 Nov. 1931, 7. Verdoorn 910 (PRE). 2926 (Bloemfontein): Winter-

valley, N of Bloemfontein, (-AA), 8 Oct. 1968, D. Muller 339 (PRE).

2927 (Maseru): Thaba'Nchu, 1 650 m, (-AA), 12 Dec. 1977, Peelers.

Gericke & Bitrelli 319 (PRE). 3025 (Colesberg): Phillipolis Road Sta-

tion, (-AB), 5 Jan. 1928, C. Smith 5219 (PRE); Trompsberg, Rietpoort

Noord, (-BA), 30 Oct. 1943, P. Kies 269 (PRE); Colesberg, (-DA), 13

Dec. 1971, Anderson 210 (PRE). 3026 (Aliwal North): Haasfontein

Farm, 40 km W of Smithfield, (-AC), 24 Jan. 1990, T. Saaiman 555

(PRE); Cliftonvale Farm ± 12 km SE of Bethulie, 1 280 m, (-CA), 18

Dec. 1983. H. Burrows 2219 (PRE); Bergplaas Farm ± 12 km SE of

Bethulie, 1 280 m, damp places near stream, (-CA), 21 Apr. 1984, 77.

Burrows 2401 (PRE). 3027 (Lady Grey): Zastron, (-AC), Apr. 1926, J.

Maree 62 (PRE).

SWAZILAND. — 2631 (Mbabane): Mbabane, Poliniane River, (-

AC), 1 1 Oct. 1963, B. Dlamini s.n. (PRE).

KWAZULU-NATAL.— 2731 (Ubombo): Mkuzi, 150' [46 m], (-

CA), 4 Sept. 1932, E. Galpin 13329 (PRE). 2828 (Bethlehem): Tugela

Valley, Natal National Park, (-DB), 26 Aug. 1950, B.E. Martin 467

(NBG). 2829 (Harrismith): Fort Mistake. (-BB), 25 Oct. 1977, 77.

Nicolson 1793 (PRE). 2830 (Dundee): Dundee, (-AB), 21 Mar. 1926,

D. Gordon Truscott 52 (PRE). 2832 (Mtubatuba): Hluhluwe Game

Reserve, (-AA), 31 Oct. 1961, P. Hitchins 117 (PRE). 2929 (Under-

burg): Bergville, (-AB), 16 Dec. 1928, E. Galpin 10791 (PRE); 20

Oct. 1950, D. Killick 1052 (PRE); Tabamhlope, (-BA), 9 Dec. 1937,

O. West 481 (PRE); Estcourt, 5000[1 500 m], (-BB), 26 Nov. 1937,

O. West 445 (BOL, GRA); Biggarsberg, (-BD), 12 Oct. 1990, W. Vos

141 (PRE); top of Sani Pass, '9500' [2 900 m], (-CB), 6 Nov. 1973,

O. Hilliard & B. Burtt 7105 (PRE); Underburg, (-CD), Mar. 1938, A.

McClean 737 (PRE); Giant's Castle, (-DD), Nov. 1914, R. Symons 157

(PRE). 2930 (Pietemaritzburg): Mooi River, (-AA), 1 Nov. 1918, A.

Mogg 3195 (PRE); Pietermaritzburg, Town Hill. (-CA), 20 Oct. 1952,

R.H. Compton 23725 (NBG); Taylo'r’s Halt, (-CB), 19 Oct. 1939, A.R.

Fail-all 54 (NBG). 2931 (Stanger): Umhloti, (-CA), 14 Mar. 1972, C.

Musil 80 (PRE); Durban, (-CC), Aug. 1883, J. Medley Wood 36 (BOL,

PRE). 3030 (Port Shepstone): l.xopo, (-AA), 7 Oct. 1918, A. Mogg

2379 (PRE).

LESOTHO. — 2828 (Bethlehem): Leribe, (-CC), without date, A.

Dieterlen 141 (PRE). 2927 (Maseru): Mafeteng, (-CD), 9 Aug. 1926,

Watt & Brandwijk 7226 (PRE); Morija, 5500' [1 700 m], (-DA), April

1919, M. Page s.n. (BOL). 2928 (Marakabei): Mamalapi, (-AC), 27

Dec. 1948, A. Jacot Guillarmod 648 (GRA, PRE); Ntibokho Valey,

(-AD), 2 Jan. 1947, A. Jacot Guillarmod 301 (PRE); Senqunyane

River near Marakabei, (-CA), 24 Nov. 1997, C. Boucher 6199 (PRE);

Cheche's Pass, 2 500 m, common in bog. (-CB), 30 Nov. 1977, D. Kil-

lick 4243 (PRE); Sehlabathebe, (-CC), Jacot Guillarmod. Getliffe &

Mzamane 212 (GRA, PRE). 2929 (Underberg): Mokhotlong, (-AC),

Jan. 1953, L. Liebenberg 5753 (PRE); Sani Valley, 2 740 m, abundant

in sponges, (-CB), 14 Jan. 1977, D. Killick 4112 (PRE).

NORTHERN CAPE.— 2816 (Oranjemund): S bank of Orange

River 1.5 km from coast, (-CB), 12 Sept. 1984, O’Callaghan & Van

Wyk 51 (NBG, PRE). 2823 (Griekwastad): Danielskuil, (-BA), 24 Mar.

1939, G.J. Lewis 434 (SAM); Griquatown Commonage, 4100' [1 250

m], (-CC), moist calcareous ground around spring, 22 Jan. 1960, O.

Leistner 1623 (BOL); banks of Riet River near Blaauwkrantz, (-DD),

10 Oct. 1935, A. Hafstrom 887 (PRE). 2824 (Kimberley): Barkly

West, Holpan, (-DA), 7 May 1936, J. Acocks 373 (PRE); Kimberley,

along Vaal River, (-DB), Dec. 1885, R. Marloth 820 (PRE); Riverton,

(-DB), Oct. 1918, J. Moran 19230 (PRE). 2924 (Hopetown): Mod-

der River, (-BA), 4 Aug. 1908, H.H.W. Pearson 1655 (NBG, SAM);

Modder River at Ritchie, (-BA), 13 Feb. 1926, C. Smith 2353 (PRE).

3018 (Kamiesberg): Kamiesberg, stream bank ± 10 km N of Lelie-

fontein, (-AC), 4 Nov. 1982, P. Goldblatt 6681 (MO, NBG), Rourke

1789 (NBG). 3024 (De Aar): Colesberg, Doomkloof Nature Reserve,

(-BD), 10 Nov. 1982, A. Hahndiek 83 (GRA). 3025 (Colesberg): near

Colesberg, (-CA), Nov. 1939, C. Thorne s.n. SAM54456 (SAM). 3026

(Aliwal North): Burgersdorp, (-CD), 1892, 77. Flanagan 1539 (PRE).

3119 (Calvinia): Oorlogskloof Nature Reserve, 578 m, (-AC), 8 Oct.

2000, W.A.J. Pretorius 609 (NBG). 3124 (Hanover): Vlakplaats [Vlak-

plaas], (-BA), Oct. 1914, 77. Bolus 13766 (BOL). 3125 (Steynsburg):

Middelburg Dist., Grootfontein Farm, (-AC), 20 Feb. 1947, G. Theron

777(PRE).

WESTERN CAPE. — 3123 (Victoria West): Murraysburg, (-DC),

Sept. 1879, W. Tyson 90 (SAM); July 1954, D.M. van Heerden s.n.

(NBG). 3221 (Merweville): Nuweveldberg, ± 60 km S of Fraser-

burg, Bok se Plaas, seepages, 1 620 m, (-BA), 26 Feb. 1986, Moffett

& Steensma 3983 (NBG). 3222 (Beaufort West): foot of Nuweveld

Mtns, along stream, 1 000m, (-BD), 15 Feb. 1978, Gibsb Russell.

Robinson. Herman & Downing 238 (PRE). 3318 (Cape Town): Green

Point flats, (-CD), Aug. 1887, J. Thode s.n. SAM9260 (SAM); Devil’s

Peak, E Buttress, (-CD), 5 Jan. 1978, B. Durand 344 (NBG); Devil’s

Peak, (-CD), 26 Sept. 2004, F. Forest. P. Goldblatt. L. Porter & I.

Ndnni 600 (NBG); above Camp's Bay, (-CD), 28 Oct. 1943, R. Adam-

son 3537 (BOL); Orange Kloof, (-CD), 24 Oct. 2000, CN. Cupido

91 (NBG); Fisantekraal, Mosselbank River floodplain, 69 m, (-DA),

29 Oct. 2008, C. Boucher 7567 (NBG); field by Maitland, (-DC), 2

Oct. 1897, Wolley Dod 3163 (BOL); Jonkershoek, Biesiesvlei, 1100'

[340 m], (-DD), 10 Sept. 1945, 77.7?. Rycroft 908 (NBG); Assegaai-

bosch. by damme, (-DD), Oct. 1969, C. van der Merwe 1241 (PRE).

3319 (Worcester): Tulbagh, Grootwinterhoek, Sneeugat Valley, (-AA),

Nov. 1916. E.P. Phillips 1699 (SAM); Ceres, Baviaansberg, 5000'

[1 500 m], (-BA), 2 Jan. 1942, R.H. Compton 12861 (NBG)'; Botha,

below dam near Groenrivier, (-CB), 26 Jan. 1982, 7.5. Walters 2526

(NBG); Franschhoek, (-CD), 30 Oct. 1913, E.P. Phillips 8266 (SAM).

3320 (Montagu): Montagu-Barrydale Rd. (-CC), Sept. 1923, M. Lev-

yns 530 (BOL); Tradouw Pass, river bank at old farm, (-DC), 28 Jan.

1969, J. Marsh 1124 (NBG). 3322 (Oudshoom): Wilderness, (-DC),

3 Nov. 1929, A. Mogg 11623 (PRE); Knysna Dist., Karatora Village,

(-DD), Dec. 1922, J. Keet 1164 (PRE). 3418 (Simonstown): Raapen-

berg VIei, (-AB), 10 Nov. 1897, A. Wolley Dod 3612 (BOL); Raap-

enberg, (-AB), 1898, Guthrie 423 (BOL); Mowbray, garden weed, (-

AB), 20 Dec. 2012, J. Manning 3405 (NBG); stream between Retreat

and Muizenburg vlei, (-AB), 14 Nov. 1897, Wolley Dod 3689 (BOL);

Table Mt slopes N of Kirstenbosch, (-AB), 12 Sept. 1928, J.B. Gillett

396 (NBG); Rondebosch, (-AB), Nov. 1885, 77. Bolus 7020 (BOL);

Rondebosch, University grounds, (-AB), 12 Oct. 1931, M. Levyns

3452 (BOL); Constantia, (-AB), 3 Nov. 2011, Ramjukadh C-LR103

(NBG); Constantiaberg, shade of forest, (-AB), 8 Mar. 1940, R.H.

Compton 8560 (BOL); Buffels Bay, damp places, (-AD), 23 Aug.

1943, R.H. Compton 14668 (NBG, PRE); Somerset West, (-DA),

20 Nov. 1947, R.N. Parker s.n. (BOL). 3419 (Caledon): Kleinmond,

bank of vlei, (-AC), 15 Oct. 1949, M.P. de Vos 1492 (NBG); Oudebos,

Riviersonderend, (-BA), Dec. 1928, C. Thorne s.n. SAM45749 (SAMf

Gansbaai, Baviaansfontein, (-CA), Aug. 1940, T.P Stokoe 7603

(BOL). 3420 (Bredasdorp): Heidelberg, Grootvadersbosch, ground

layer in moist or wet forest, (-BB), 6 Dec. 1953, 5. Maguire 1037a

(NBG); Bredasdorp, Nachtwacht, (-CA), Nov. 1926, C. Smith 3048

(PRE). 3421 (Riversdale): 8 mi [13 km], W of Riversdale, (-AA), 7

Oct. 1928, J.B. Gillett 1165 (NBG); bank of Kafferkuilsrivier, 80 m,

(-AD), 22 Dec. 1980, P. Bohnen 7785 (NBG, PRE). 3423 (Knysna):

Brenton, shady places, (-AA), 5 Jan. 1922, A. Duthie 722 (NBG).

3423 (Knysna): Keurbooms River Bridge, disused forest track, (-AB),

27 Sept. 1967, J.77. Marsh 6/9 (NBG, p'rE).

EASTERN CAPE.— 3026 (Aliwal North): Elandshoek, (-CC),

Oct. 1903 , 77. Bolus 152 (BOL). 3028 (Matatiele): Rhodes Dist.,

Naude's Nek, 2 610 m, (-CB), 11 Dec. 1999, M. Mothogoane 185

(PRE). 3029 (Kokstad): Kokstad, Palmiet, (-AB), 21 Jan. 1957, L.E.

Taylor 5524 (NBG); between Flagstaff and Kokstad, (-CB), 4 Dec.

1928, J. Hutchinson 1789 (PRE); Fort Donald, (-DC), forest, summer

1915, Sr. Stephany 635 (BOL). 3126 (Queenstown); Sterkstroom. Pen-

hoek, (-BC), 21 Dec. 1942, WE Barker 2139 (NBG); Queenstown,

river banks, (-DD), Oct. 1895, E. Galpin 1937 (PRE); Andriesberg,

(-DD), Dec. 1896, E. Galpin 2223 (GRA, PRE). 3127 (Lady Frere):

25 km from Cala tunoff on Engcobo road, (-DB), 12 Jan. 1997, C.

Bredenkamp 1149 (PRE). 3128 (Umtata): Baziya, (-BB). without date,

R. Baur 277 (SAM). 3129 (Port St Johns): Goss Point, bog, (-BD), 10

Nov. 1970, R. Strey 10146 (PRE); Port St Johns, (-DA), Oct. 1909,

77. Swinny & F. Baker 25124 (PRE). 3223 (Rietbron): Richmond Dist.,

Vlakplaats, (-CA), Oct. 1914, 77. Bolus 15292 (PRE). 3225 (Somerset

East): 21 mi [34 km], W of Cradock, Chalmers, (-AB), 2 Dec. 1950,

S. M. Johnson 691 (BOL), 5. Maguire 691 (NBG); Mountain Zebra

National Park, (-AD), 12 Dec. 2005, S. Bester 6319 (GRA). 3226

190

Bothalia 43,2 (2013)

(Fort Beaufort): Katberg, (-BC), Drege 10/1132 (BOL); Buffelshoek

Nek on road to Cradock, (-CA), 27 Oct. 1945, R. Story 82 (PRE);

Hogsback, (-DB), Jan. 1919, G. Rattray 15736 (BOL); Hogsback

Forest Reserve, ± 800 m, (-DB), 25 Nov. 1969, K. Dahlstrand 1832

(GRA, NBG); University of Fort Hare, Sandile’s Kop, (-DD), 5 Nov.

1936, M Giffen 592 (PRE). 3227 (Stutterheim): Dohne Research Sta-

tion, (-CB), 23 Oct. 1942, J. Acocks 9228 (PRE); grassy valleys near

Komgha, (-DB), Nov. 1891, H. Flanagan 1109 (BOL, GRA, PRE);

East London, (-DD), 1888, J. Thode s.n. (PRE); East London, damp

places, (-DB), Aug. 1963, A. Batten 4-Pl. 98 (NBG). 3228 (Butter-

worth): near Butterworth, (-AC), 26 Nov. 1945, R.H. Compton 17705

(NBG); Qora River Mouth near Mazeppa Bay, (-BC), without date, C.

miner 419 (PRE); Dwessa Nature Reserve, (-BD), 24 Nov. 1978, H.

Under 1890 (PRE); Morgan’s Bay, (-CB), 17 Jan. 1951, A.M. Wilman

1084 (BOL, PRE). 3323 (Willowmore): Deepwalls, (-CC), 1924, J.

Phillips s.n. (GRA, PRE); Gouna, Lilyvlei Nature Reserve, ± 1000'

[300m], (-CC), 19 Oct. 1971, H.C. Taylor 8000 (NBG); Lilyvieibush,

wet places, (-CC), 22 Nov. 1977, C.J. Geldenlmys 413 (NBG); Kny-

sna, forest near Blaauwkranz River, (-DC), 20 Oct. 1959, T.M. Wnrts

2007 (NBG). 3324 (Steytlerville): Baviaanskloof 3 km from Geel-

houtbos, (-CB), 23 Sept. 2004, P. Goldblatt & L.J. Porter 12537 (MO,

NBG); E of Gamtoos River drift, (-CD), (BOL). 3325 (Port Eliza-

beth): Winterhoek Mtns, (-CA), 12 Sept. 1930, C. Fires, T. Norlindh

& H. Weimarck 790 (GRA, PRE); Walmer, (-DC), Oct. 1909, T. Pater-

son 817 (GRA). 3326 (Grahamstown): Belmont Valley, near Clarke’s

Farm, (-BC), 13 Sept. 1961, M. Wells 2802 (GRA, PRE); Alexandria

Forest, Forester’s Cottage, (-CB), 9 Dec. 1953, S. Johnson 831 (PRE);

top end of Howieson’s Poort, (-DA), swampy place, 28 Feb. 1932, J.

& B. Rennie 577 (BOL); Kowie Dist., road to Three Sisters, (-DB),

25 Sept. 1918, L. Britten 730 (PRE). 3424 (Humansdorp): Oudebos

Flats, (-AA), Oct. 1920, H. Fourcade 959 (BOL); Tsitsikamma Strand,

(-AA), 30 Jan. 1982, H. Venter 8653 (PRE); Clarkson, (-AB), Oct.

1926, Thode 7J7(PRE).

Sect. Hecatonia (Lour.) DC. in Prodromus systematis

naturalis regni vegetabilis 1: 30 (1824).

3. Ranunculus dregei J.C. Manning & Goldblatt,

nom. nov. pro. Ranunculus meyeri Harv. in Harvey &

Sonder in Flora capensis 1: 7 (1860), horn, illegit., non

Lowe: 74 (1857), nom illegit. superfl. pro R. grandi-

folius E.Mey. in Ledebour (1830); Cook: 240 (2004).

Type: South Africa, [Eastern Cape], ‘Katberg’, Nov.

[1832], Drege s.n. (K [K00n076101]—Muka image!,

lecto., designated here; BM, K, P, PRE, TCD, iso. —

Aluka images!). [Harvey cited duplicates at K and TCD

in the protologue and we select the K specimen as lecto-

type as being the more complete. This specimen has the

collecting date 1838 inscribed on it but this is impossible

as Drege left South Africa in 1834. Drege collections of

the species in P are dated 9 Nov. 1832, which corres-

ponds with the date of his arrival at the Katberg Pass on

12 November 1832 (Glen & Germishuizen 2010) and

this evidently represents the true collection date.]

R. meyeri var. transvaalensis Szyszyl.: 102

(1887). Type: South Africa, [Gauteng], ‘Hohes feld

prope Henopsriver’, Relvnann (?Z, holo., not located).

[fide Exell & Milne-Redhead (I960)].

[/?. meyeri var. rogersii Burtt Davy nom. nud.

in Burtt Davy: 109 (1926). Specimen: Rogers 19591

(Herb. Rogers)].

[Ficaria radicans E.Mey. nom. nud. in Drege:

184 (1843-1844). Specimen: Drege s.n. {K00l)076101 ,

TCD0001712)].

[Ranunculus volkensii var. meyeri (Harv.)

T.Duncan ms. on Dieterlen s.n. (BM 00056 1551 6)\

Prostrate, mat-forming, perennial; stems condensed,

corm-like, covered with fibrous remains of old leaf

sheaths, producing creeping runners rooting at nodes,

runners glabrous, 0.5-1.5(-2.0) mm diam., intemodes

10-140 mm long. Roots few, thong-like. Leaves all

basal, in distant tufts of 2-5 along runners, petiole

(5-)10-100(-200) mm long with stipules adnate and

sheathing for 2-10 mm, glabrous or thinly villous with

hairs to 1 mm long, blade ovate to cordate or reni-

form, (2-)5-20(-25) mm diam., weakly or strongly

4-12(-16)-crenate or dentate (rarely ± entire), with

reddish callus at the tip of each serration, leathery, gla-

brous but usually with scattered hairs on margins, or

adpressed-pilose adaxially. Flowers solitary at nodes,

(5-)8-14 mm diam.; peduncle naked, scapose, ± as long

as leaves, (5-)10-60 mm long, 0. 5-1.0 mm diam. Sepals

5, weakly to strongly reflexed, ovate to suborbicular,

cucullate, (2-)3-4 mm long, glabrous, yellowish. Pet-

als 5-10, linear-lanceolate or ligulate, (3-)4-7 x 1.0-1. 5

mm, pale yellow, narrowed into short claw in basal ±

1 mm, acute or attenuate, veins 3, unbranched, nectary

pocket-like with acute flap. Stamens 11-30 in ± 2 series,

filaments 0. 5-2.0 mm long, anthers ± 0.5 mm long. Car-

pels 6-20(-30), glabrous. Achenes in subglobose head,

broadly ellipsoid, ± 2 x 1.5 mm, smooth, keeled, shortly

beaked; receptacle glabrous. Flowering time: Nov.-Jan.

(-Mar.). Figure 7.

Distribution and ecology: distributed mainly along

the eastern escarpment, from Gaika’s Kop in the Ama-

tola Mtns in Eastern Cape through KwaZulu-Natal and

Lesotho to Standerton and Vaalbank in Mpumalanga and

the Leolo Mountains in Limpopo (Figure 4), extend-

ing north of South Africa into the eastern highlands of

Zimbabwe and Mozambique (Exell & Milne-Redhead

1960), with a remarkable southern outlying station near

the summit of the Swartberg Mtns in Western Cape.

Although there are early records further inland, from

Parys in the Free State, Pretoria and Johannesburg in

Gauteng, and Potchefstroom in North-West, the species

has not been collected there since the first half of the

twentieth century and is presumably extinct here.

Ranunculus dregei restricted to peaty seeps or boggy

places at higher altitudes, 1 500-3 000m, where it fonns

mats in open spaces.

Diagnosis: a distinctive species, readily recognised

by its dwarf, creeping habit, simple leaves with soli-

tary flower per leaf tuft, and narrow, ligulate petals with

unbranched veins.

The species was first recorded from north of the Lim-

popo River Valley, in the highlands of Mozambique and

Zimbabwe, in 1956 (Exell & Milne-Redhead 1960) and

annotations on some herbarium specimens by T. Dun-

can indicate an intention to reduce it to a variety of the

tropical African Ranunculus volkensii Engl. Although

the two taxa are similar in habit, R. volkensii has notice-

ably thicker stolons, ± 2-3 mm diam., often lanceolate

leaves, and more critically, obovate petals with branch-

ing venation, quite unlike the ligulate, 3-nerved petals of

R. dregei. It also has mostly ± twice as many carpels as

R. dregei.

Histoiy: first collected in Eastern Cape by J.F. Drege

(1794-1881) sometime in the early summer of 1832,

the specimens themselves are not localised, Ranuncu-

lus dregei was listed as being collected on the Katberg

Bothalia 43,2 (2013)

191

in Drege’s Zwei pflanzengeographische Documente.

These collections were annotated with the manuscript

name Ficaria radicans by Ernst Meyer, who contrib-

uted the botanical text to the Documente, but the spe-

cies was only formally described by Harvey (1860),

who named it after Meyer. Harvey (1860) unfortunately

overlooked the fact that the name R. meyeri had already

been proposed by R.T. Lowe (1857) as a replace-

ment name for the East-Central Asian R. grandifolius

E.Mey. (1830) under the misapprehension that the lat-

ter was synchronous with his R. grandifolius Lowe

(1831) from Madeira. In fact it is R. grandifolius Lowe

that is the illegtimate later homonym and thus requires

a new name, and R. meyeri Lowe becomes an illegiti-

mate superfluous name for R. grandifolius E.Mey. This

renders R. meyeri Harv. an illegitimate later homonym

requiring a new name, for which we propose R. dregei in

honour of the original collector of the species.

A collection by Anton Rehmann (1840-1917) from

Hennopsrivier, west of Pretoria, with glabrous petioles

was distinguished as var. transvaalensis by Szyszylow-

icz (1887), but the name was synonymised by Exell &

Milne-Redhead ( 1960). We have been unable to trace the

type specimen in the Schinz herbarium in Zurich (H.P.

Linder, pers. com. 3 Jan. 2013) but have no reason to

question this decision. Other collections from Gauteng,

viz. Repton 3430 from Rietvlei Reserve near Pretoria

and Gilfillan 162 from near Johannesburg probably rep-

resent the same variant, being almost entirely glabrous,

with scattered hairs only on the sheathing leaf bases.

Additional specimens seen

LIMPOPO. — 2430 (Pilgrim’s Rest): Sekhukuniland, Leolo Mtns,

18 km from Schoonoord Police Station, (-CA), seepage area, (-CA), 4

Dec. 1999, A.E van Wyk <&. SJ. Siebert 13459 (PRE).

NORTH-WEST. — 2627 (Potchefstroom): Potchefstroom, common

in marshes bordering Mooi River, (-CA), 26 Dec, 1903, J. Burtt Davy

1029 (BOL, PRE); Gerhardusminnebron Eye, (-CA), 26 Oct. 1946, W.

Louw 1520 (PRE).

GAUTENG. — 2528 (Pretoria): Rietvlei Reserve, (-CA), 4 Oct.

1947, J. Repton 3430 (PRE). 2628 (Johannesburg): vleis near Johan-

nesburg, (-AA), Nov./Dec. 1898, D. GilfUIan 162 (GRA, PRE).

MPUMALANGA. — 2529 (Witbank): Middelburg, Wonderhoek,

pond borders, (-CD), 15 Nov. 1922, H. Rudatis 2647 (PRE); Vaal-

bank, between Middelburg and Pretoria, (-CD), 4500' [1 400 m]. Sept.

1886, H. Bolus 7639 (BOL). 2530 ( Lydenburg): 15 km from Belfast

on road to Dullstroom, Farm Pineglades, along dam, 2 000m, (-CA),

17 Apr. 1981, 1. dll Toil 333 (PRE). 2629 (Bethal): Standerton, (-CD),

16 Oct. 1893, R. Schlechter 3469 (BOL, GRA, PRE). 2630 (Carolina):

Carolina, (-AA), 20 Oct. 1932, E. Galpin s.n. (BOL); Ennelo, Spit-

skop, (-CB), Dec. 1915, R. Pott-Leendertz s.n. (PRE). 2730 (Vryheid):

Wakkerstroom, 6200' [1 900 m], (-AD), Nov. 1916, H.W. Beeton 35

FIGURE 1 .—Ranunculus dregei, Naude’s Nek, no voucher. A, flowering plant; B, two sepals; C, petal, dorsal and lateral view; D, stamen; E, pistil

and detached carpel; F, fruit and detached achene. Scale bar: A, 10 mm; B, C, E, F, 1 mm; D, 0.5 mm. Artist; John Manning.

192

Bothalia 43,2 (2013)

(SAM); Pongola Bush Nature Reserve, viei in stream, 1 400m, (-BC),

9 Oct. 1990,7/. Gle}i 2360 (PRE).

FREE STATE. — 2828 (Bethlehem); Bestersviei, near Wotzieshoek,

(-BD), 1893, H. Flanagan 2093 (PRE). 2925 (Jagersfontein): Parys,

(-CC), 20 Oct. 1906, F. Rogers 5074 (PRE).

KWAZULU-NATAL.— 2729 (Volksrust): Charlestown, (-DB), 6

Dec. 1892, J. Medley-Wood 4690 (PRE). 2929 (Underberg): Estcourt,

Ntabamhlope Pasture Reserve, vIei, (-BA), 9 Nov. 1939, O. West

1513 (PRE); Mooi River, (-BB), 26 Oct. 1918, A. Mogg 3203 (PRE);

Polela, near Underberg, in stream, (-CB), 6 Dec. 1935, R.A. Dyer 3289

(PRE); 13 mi. [21 km] from Dargle to Mpendhle, viei, (-DB), 26 Feb.

1964, EJ. Moll 673 (PRE); Underberg, Watermead Farm, (-DB), 23

Sept. 1989,/?. Williams 333 (PRE).

LESOTHO. — 2828 (Bethlehem): Leribe, (-CC), 5-6000'

[1 500-1 800 m], Feb. 1913, E.R Phillips 786 (SAM); Butha-Buthe,

9 500' [2 900 m], (-CD), 2 Feb. 1954, B. Maguire 399 (NBG). 2928

(Marakarei): Little Bokong at Mosalamane Pass crossing, (-AA),

22 Dec. 1946, A. Jacot-Gullarmod 99 (GRA, PRE); headwaters

of Bakong River, wet fen, (-AB), 14 Jan. 1996, P. Phillipson 4645

(GRA); Mamalapi, (-AC), 28 Dec. 1948, G.D. Morris 21429 (NBG);

Mamathes, (-BB), 14 Nov. 1948, A. Jacot-Guillarmod 389 (PRE).

2929 (Underberg): Mokhotlong, bog near Thabana Ntlenyana, (-AC),

20 Jan. 1955, J. Coetzee 582 (PRE); 2 Feb. 1988, C. Schwabe 13b

(PRE); Sehlabathebe, 2 300-2 500m, (-CC), 4-14 Jan, 1973, Jacot-

Guillarmod. Getliffe & Mzamane 255a (GRA, PRE). Imprecise local-

ity: ‘Basutoland, swampy ground near Buffels River Waterfall’, 15

Mar. 1904, E.E. Galpin 6565 (BOL, GRA, SAM).

WESTERN CAPE.— 3322 (Oudshoorn): Swartberg, 8 miles W

of top of Swartberg Pass, (-BC), 5000' [1 500 m], 20 Nov. 1954, T.P.

Stokoe s.n. SAM70I33 (SAM); Swartberg, marsh at head of stream

E of pass, (-BC), 19 Jan. 1961, E. Esterhuysen 28836 (BOL, PRE);

upper slopes of Blesberg, (-BC), 5950' [1 800m], 15 Dec. 1986, J.//.,/.

Vlok / 775 (NBG).

EASTERN CAPE. — 3027 (Lady Grey): Witteberg, Beddgelert, (-

DA) , 2 Dec. 1981, O. Hilliard B. Burtt 14631 (PRE); S slopes of

Ben MacDhui Mtn, Tiffendell Ski resort, 2 730 m, (-DB), 6 Jan. 1997

[fruiting], T. Bold 3483 (GRA). 3028 (Matatiele): Barkly East, 3 km

SE of Cairntoul Police Hut, wet places, (-CA), 19 Dec. 1982, P. Phil-

lipson 709 (PRE); Maclear, Woodcliffe Trails, riverine grassland, 2

300 m, (-CC), 15 Nov. 1992, A. Abbott 5865 (PRE). 3126 (Queens-

town): 19 km from Dordrecht to Queenstown, 1 311m, (-DB), 13 Jan.

1997, G. Germishuizen 8915 (PRE). 3127 (Lady Frere): Dordrecht,

5300' [1 600 m], swampy ground, (-AD), Jan. 1899, T.R. Sim s.n.

(SAM). 3225 (Somerset East): Boschberg, (-DA), Feb. without year,

MacOwan 1555 (BOL, GRA). 3226 (Fort Beaufort): summit of Great

Winterberg, locally common in seeps on dolerite, 2 250 m, (-AD),

17 Feb 2013, N.A. Helme 7715 (NBG); Katberg, Effingham, (-BC),

26 Dec. 1912, E. Galpin 8337 (PRE); Hogsback, foot of Gaika’s Kop

(-DB), Jan. 1919, G. Rattray s.n. (BOL); Hogsback, Hunterstown, (-

DB) , 9 Apr. 1944, M. Gijfen s.n. (PRE). 3227 (Stutterheim): Keiskam-

mahoek, Gxulu Mtn [Cata Peak], (-CA), 26 Jan. 1949, R. Story 3734

(GRA); small river near Kologha Forest, (-CB), 28 Nov. 1956, G.

Theron 2126 ( PRE).

Sect. Batrachiiiin DC. in Regni vegetabilis systema

naturale 1 : 323 (1817).

4. Ranunculus trichophyllus Chaix in Villars: 335

(1786); Cook: 126 (1966). Batrachiitm trichophyllus

(Chaix.) Bosch: 5 (1850); Eichler & Walsh: 312 (2007).

Type: ‘Frequentissimus in rivulis quietis, fossisque aqua

plenis’, Haller 1162 (G, neo., designated by Lourteig

(1951 ), not seen).

R. drouetii F.W.Schultz ex Godron: 10 (1842).

Type: Flora Galliae el Germaniea exsicattae it. 404 (not

seen).

|/Q aciitalilis sensii Harvey: 6 (1860), non L.

(1753)]

[See Cook ( 1966) for additional synonyms.]

Submerged aquatic annual (?or perennial), rooted in

water up to 500 mm deep; stems hollow, 2-3 mm diam.,

rooting at lower nodes. Roots fibrous. Leaves cauline,

petiole 25^0 mm long with stipules adnate and form-

ing oblong to ovate sheath 3-6 mm long, glabrous,

blade globose or obconical in outline, 15^5 mm diam.,

repeatedly tri- or dichotomously divided into rigid or

flaccid capillary segments spreading in three dimen-

sions. Flowers solitary at nodes, emergent, (4-)8-15

mm diam.; peduncle naked, scapose, 10-50 mm long,

0. 5-1.0 mm diam., usually recurved in fruit. Sepals 5,

spreading, ovate, cucullate, 2. 5-3. 5 mm long, glabrous.

Petals 5, obovate, (3.0)3.5-5.5(-6.0) x 2-5 mm, white

with yellow claw, narrowed into short claw in basal ± 1

mm, obtuse or truncate, veins branched, nectary lunate.

Stamens 9-15 in ± 2 series, filaments 1.0-1. 5 mm long,

anthers ± 0.75 mm long. Carpels 16-35, hispid. Achenes

in subglobose head, ovoid to obovoid, not compressed,

1.20-1.50(-1.75) mm long, transversely rugose, mostly

sparsely hispid subapically along lower keel, rarely gla-

brous, not beaked; receptacle densely hispid. Flowering

time: dependent on rainfall and coinciding with the wet

season, mainly Aug.-Sept. in the winter-rainfall zone

and Oct.- Jan. in the summer rainfall zone. Figure 8.

Distribution and ecology’’, distributed widely through

Furasia and North Africa, North America, Australia, Tas-

mania, and southern (but not tropical) Africa, where it

is recorded so far only from Fesotho and South Africa

(Figure 9), and reportedly also Botswana (Cook 2004)

but not Namibia (Craven 1999). In South Africa the spe-

cies is scattered through the southern and eastern interior

of the county along the Vaal/Orange Rivers and tributar-

ies, and along the southern escarpment from Calvinia to

Queenstown, with a few coastal stations in the south-

western and southern Cape around Velddrif, Cape Town,

Bredasdorp and Uitenhage.

Plants grow submerged in freshwater seasonal pools

and ponds and in slow, intermittent or perennial streams,

furrows or irrigation ditches up to 50 cm deep, persist-

ing only briefly along the muddy edges as the water

dries up. It has been recorded from sea level to well over

2 000 m. In southern Africa R. trichophyllus has occa-

sionally been regarded as troublesome in the past, viz. as

a ‘serious impediment in the irrigation canals’ near Cra-

dock in the 1950s [Barnes s.n. (GRA)].

The natural occurrence of the species in the South-

ern Hemisphere is worth closer examination given the

large disjunction to Furasia and North America. The

early collection of the species in pristine habitats in Aus-

tralia suggests that it is probably native there (Fichler &

Walsh 2007), and similar considerations apply in south-

ern Africa, where it was first documented (under the

name Ranunculus aquatilis F.) in the early 1800s from

the Eastern Cape, around Uitenhage and Graaff Reinet,

by C.F. Ecklon (1795-1868), C.L. Zeyher (1799-1858)

and I.L. Drege (1853-1921). Documentation of R. tri-

chophyllus on the Cape Peninsula only dates from the

end of the 19"' century, which is unusually late for such a

distinctive species.

Diagnosis’, the only aquatic species in the subre-

gion, distinguished at once by its habit, finely dissected

leaves, white petals with yellow claws, and small, ovoid.

Bothalia 43,2(2013)

193

FIGURE 8. — Raminciilus trichophylhis, Kareehoutrivier, Snijman 771

(NBG). Achene, lateral and dorsal view. Scale bar: 1 mm. Artist:

John Manning.

transversely rugose achenes, usually with a few, scat-

tered bristles along the lower keel below the remains of

the style but sometimes glabrous or ± densely bristly.

The transverse folds on the mature achenes are due to

the presence of styloid crystals in the carpel walls (Cook

1966).

consistent with R. trichophyllus and not R. rionii. Some

specimens, however, including Comins 800 (BOL)

from Tarkastad and others from along the Vaal River,

have heads of 40-60 achenes, each achene ± 1 mm

long, thus potentially R. rionii. Although the difference

between the two taxa in achene number and size as ini-

tially treated by Cook (1963) was absolute, the ranges of

variation in the two characters have gradually converged

(Cook 1966, 1993; Pizarro 1995) to the stage where it is

difficult to assign ‘intermediate’ specimens. It may also

be relevant here that R. rionii has been treated as con-

specific with R. trichophyllus in the past at subspecific

or varietal rank (Cook 1963). The size of the achenes

may bear relation to the number of (developing) carpels,

as seen in Wilman 21494 (PRE), in which heads with

over 40 ripened carpels have the individual achenes ± 1

mm long whereas those with few fertilised carpels have

achenes ± 1.3 mm long. Significantly, no authors have as

yet proposed that more than one aquatic species occurs

in the subcontinent, although it is not impossible that

there has been more than one introduction from Europe.

The matter deserves attention but until then we adopt a

conservative approach and provisionally treat all of the

southern African material under the older name R. tri-

chophylhis. This name has also been applied to the mate-

rial from Ethiopia (Teketay & Edwards 2000).

Additional specimens seen

History’: The recent determination of the southern

African material as Rammculus rionii (Glen 2003; Win-

ter 2006) follows the identifications of the southern Afri-

can material by C.D.K. Cook (1966, 2004), a specialist

in the taxonomy of Ramincuhis sect. Batrachiurn. Prior

to this, the South African collections had been identified

as R. trichophyllus (Adamson 1950) or its synonym R.

drouetii (Burtt Davy 1926). The two species R. rionii

and R. trichophyllus are sympatric in Europe and differ

solely in the number and size of their achenes, with R.

rionii characterised by smaller, more numerous achenes

(Cook 1964, 1966, 1993; Pizarro 1995). The number of

achenes per fruiting head ranges from 60-90 in R. rio-

nii and from 16-35 in R. trichophyllus (Cook 1966).

Dimensions for the achenes in R. rionii have been vari-

ously given as up to 1 mm long (Cook 1964, 1966,

1993) or 1.0-1. 2 mm long (Pizarro 1995) and for R. tri-

chophyllus as more than 2 mm long (Cook 1964), more

than 1.5 mm long (Cook 1966, 1993), or 1.20-1.75 mm

long (Pizarro 1995). R. trichophyllus is a very wide-

spread species, occurring throughout Eurasia, including

the Mediterranean Basin, North America and Australia,

where it is probably also native, but R. rionii has a more

restricted distribution in Central and Eastern Europe and

the eastern Mediterranean into Asia Minor (Cook 1966).

The illustration of the fruiting head of Ranunculus

rionii given by Cook (2004) in his guide to Aquatic

and wetland plants of southern Africa certainly con-

forms to that species in its numerous achenes, as does

the illustration of the individual achene, with a length

of less than 1 mm, but the source of the original mate-

rial for these drawings is not given. The majority of the

southern African material that we have examined has

fewer than 40 achenes per fruiting head, with individual

mature achenes measuring 1.2-1. 8 mm long, excluding

the stylar remnants (immature achenes are smaller), thus

GAUTENG. — 2627 (Potchefstroom): Vereeniging, Vaal River, (-

CB), Nov. 1911,/?. Leendertz 3892 (PRE).

MPUMALANGA. — 2629 (Bethal): near Ermelo. still pools in

river, (-DB), 25 Aug, 1904, J. Burtt Deny /«75 (BOL, PRE).

FREE STATE. — 2627 (Potchefstroom): Parys, irrigation ditch

along Vaal opposite the ‘Small Island', (-CD), Sept. 1932, C. Smith

6309 (PRE). 2727 (Kroonstad): Kroonstad, Vais River, (-CC), Dec,

1927, J. Pont 268 (PRE). 2827 (Senekal): Eicksburg, at Gansfontein,

(-DD), 24 Oct. 1884, E. Galpin s.n. (BOL, PRE). 2828 (Bethlehem):

‘Kroonspruit’, (-AD), 22 Sept. 1976, J. Blom 305 (PRE). 2925 (Jag-

ersfontein): 12 mi [19 km] E of Eauresmith, (-CC), 3 Sept. 1925, C.

Smith 479 (PRE); Farm ‘Driefontein West’, in Kaffirrivier, (-DB), 6

Oct. 1972, /?. Marshall M72/053/C (PRE); Eauresmith. Boomplaats,

in spruit, (-DC), 3 May 1934, 1. Verdoorn 1383 (BOL). 3026 (Aliwal

North): 'A mi [0.4 km] S of Loftes Siding, (-BA), 15 Nov. 1969, D.

Edwards 4163 (PRE).

LESOTHO. — 2828 (Bethlehem): Hlotse River at Makokoane, (-

BC), 23 Oct. 1909, A. Dieterlen 808 (BOL, PRE, SAM). 2928 (Mar-

akabei): Semongkong, (-CC), 12 Jan. 1954, A. Jacot-Guillarmod 1754

(PRE). 2929 (Underberg): Maluti Mtns, near Mokhotlong, 2 300 m, (-

AC), 22 Feb. 1987, M. Panagos 87 (PRE).

NORTHERN CAPE. — 2824 (Kimberley): Riverton, (-DA), Nov.

1919, M. Wilman 21494 (PRE). 2922 (Prieska): Prieska, Orange River,

(-DA), Oct. 1933, E.G. Bisach 963 (NBG). 2923 (Douglas): Griqua-

land West, [Douglas], Mazeisfontein, (-BA), Oct. 1919, E. Anderson

609 (BOL). 2924 (Hopetown): Modder River, (-BA), 4 Aug. 1908,

H.H. W. Pearson 1650 (NBG). 3024 (De Aar): 40 km from Philipstown

on road to Colesberg, pool in small stream, (-DB), 19 May 1989. H.

Burrows 2991 (GRA). 3025 (Colesberg): near Colesberg, (-CA), Nov.

1939, C. Thorne s.n. SAM54455 (SAM). 3119 (Calvinia): along Karee-

houtrivier, 24 km S of Bo-Downes homestead, (-DD), 27 Oct. 1983,

D. Snijman 771 (NBG). 3220 (Sutherland): Roggeveld, Soekop, Wit-

fontein, (-AA), permanent free water, 16 Sept. 2006, H. Rosch 552

(NBG); Roggeveld Escarpment, pools on Farm Blesfontein, (-AD),

4 Nov. 2012, P. Goldblatt & J. Porter 13715 (NBG); Verlatekloof

Pass, pools in stream, (-DA), 7 Sept. 1926, M. Lexyns 1587 (BOL);

8 Sept. 1988, M. Crosby 988 (PRE); 4 km from Klein Roggeveld via

Komsberg on way down pass, (-DA), 1 300 m, 6 Sept. 1986, 1. Cloete

& W. Haselau 245 (NBG); road to Komsberg Pass, ± 4-5 km SE of

Earm Tonteldoosfontein, (-DB), 15 Sept. 2004, D.A. Snijman 1946

(NBG). 3221 (Merweville): Phisante River, in stream, (-AC), 11 May.

1976, M.F. Thompson 3015 (NBG, PRE); Bok se Plaas, ± 60 km S of

194

Bothalia 43,2(2013)

FIGURE 9, — Distribution of R. trichophylhis in southern Africa.

Fraserburg, I 530 m, (-BA), Moffett d Steersma 4022 (NBG); Layton,

Rietviei, (-BB), 914 m, D. Shearing 187 (PRE).

WESTERN CAPE.— 3218 (Clanwilliatn): Velddnft, (-CA), with-

out date, C. Gaigher 5 (NBG). 3318 (Cape Town): Sea Point, (-CD),

July 1895, MacOwan 1701 (SAM); pools near Salt River junction, (-

CD), 3 Nov. 1891, Wolley-Dod 3659 (BOL); Vygekraal River, (-CD),

30 Nov. 1891, A. Wolley-Dod 3623 (BOL, PRE); Rietviei, (-CD), 10

Oct. 1965, A. Mauve 4384 (PRE); Bellville, pool in granite outcrop,

(-DC), 8 Sept. 1938, E. Cohen s.n. (NBG). 3321 (Ladismith): Prince

Albert, Gamka Poort, (-BC), Nov. 1935, C. Thorne s.n. SAM51869

(SAM). 3418 (Simonstown): Noordhoek, salt pan, (-AB), 31 Jan.

1971, I. Waher 45 (BOL); Ottery, (-BA), 18 Oct. 1935, R. Adam-

son 2170 (BOL, SAM); Cape Flats, (-BA), no date, E. Stephens s.n.

(BOL); Cape Flats near Phillipi, Edith Stevens Reserve, (-BA), in tem-

porary pools up to 0,5 m deep, 26 Sept. 1994, J.R Rourke 2065 (NBG,

PRE). 3420 (Bredasdorp): Bredasdorp, Nachtwacht, vlei, (-CA), Nov.

]926, C. Smith 3047 (PRE).

EASTERN CAPE. — 3124 (Hanover): Wapadsberg Pass, running

stream, (-DD), 26 Nov. 1977, O. Hilliard & B. Biirtt 10679 (NU,

PRE). 3126 (Queenstown): Broughton, near Molteno, 6300' [I 900

m], (-BC), Dec. 1892, H.G. Flanagan 1563 (PRE, GRA, SAM). 3222

(Beaufort West): Beaufort West, Nieuweveld Mtns, 1 676 m, (-BA),

16 Apr. 1978, B. Gibbs Russell, Robinson & Herman 448 (PRE);

Mountain View Farm, (-BD), 16 Apr, 1978, Gibbs Russell. Robinson

& Herman 448 (GRA). 3223 (Rietbron): Murraysburg, Roode Poort

[Rooipoort], (-BC), Sept. 1879, 4500', W. Tyson 66 (SAM). 3224

(Graaff Reinet): Graaff Reinet, near Zontag’s River [Sondagsrivier],

(-BA), Nov. 1865, H. Bolus s.n. (GRA). 3225 (Somerset East): Cra-

dock. Junction Farm. (-AA), Dec. 1952, //. Barnes s.n. (GRA); Cra-

dock. Farm Zuurfontein near Nardouwsberg, (-AA), 1 550 m, local in

permanent stream, 14 Jan. 1990, H P. Linder 5085 (NBG). 3226 (Fort

Beaufort); Great Winterberg, S of Tarkastad, Fairfield Farm, 5500',

fl 700 m|, (-AD), 24 March 1954, D. Comins 800 (BOL); Shiloh

prope Queenstown, (-BB), without date, R. Baur 919 (SAM). 3325

(Port Elizabeth): in stagnis prope Uitenhage, (-CD), Ecklon 1766

(SAM). 3326 (Grahamstown): Grahamstown, Stowan Farm, (-AD), 17

Apr, 1934, L. Britten 5 9 78 (GRA).

ACKNQWLEDGEMENTS

We are grateful to Cameron McMaster, Elizabeth

Parker and Rosemary Smuts for their support in the

field; to Roy Gereau, Missouri Botanical Garden, for

nomenclatural advice, and to Mary Stiffler from the

same institution for her help in obtaining literature.

REFERENCES

ADAMSON, R.S. 1950. Ranunculaceae. In R.S. Adamson & T.M.

Salter, Flora of the Cape Peninsula: 401—403. Juta, Cape Town

& Johannesburg.

BEWS, J.W. 1921. An introduction to the flora of Natal. Pietermaritz-

burg.

BOSCH, R.B. van den. 1850. Prodromus florae hatavae, vol. 1. Sum-

tibus.

BURTT DAVY, J. 1921. New or noteworthy South African plants IV.

Kew Bulletin of Miscellaneous Information 1921: 335-343.

BURTT DAVY, J. 1926. A manual of the flowering plants and ferns of

the Transvaal with Swaziland. South Africa. Longmans, Green

& Co., London.

CANDOLLE, A.P. de. 1 8 1 7 [ 1 8 1 8]. Regni vegetabilis systema naturale,

vol. 1. Treuttel & Wiirz, Paris.

CANDOLLE, A.P. de. 1824. Prodromus systematis naturalis regni veg-

elabilis, vol. 1. Treuttel & Wiirz, Paris.

COOK, C.D.K. 1964. Ranunculus subg. Batrachium. In T.G. Tutin,

V.H. Heywood, N.A. Burgess, D.H. Valentine, S.M. Walters &

D.A. Webb (eds). Flora europaea, vol. 1 Lycopodiaceae to Pla-

tanaceae: 237, 238. Cambridge University Press, Cambridge.

COOK, C.D.K. 1966. A monographic study of Ranunculus subge-

nus Batrachium (DC.) A. Gray. Milteilungen der Botanische

Staat.ssammlung Miinchen 6: 47—237.

COOK, C.D.K, 1993. Ranunculus subg. Batrachium. In T.G. Tutin,

N.A. Burges, A. O. Chater, J.R. Edmonson, V.H. Heywood, D.M.

Bothalia 43,2 (2013)

195

Moore, D.H. Valentine, S.M. Walters & D.A. Webb (eds). Flora

europaea, vol. 1 Lycopodiaceae to Platanaceae, 2"'* ed,: 285, 286,

Cambridge University Press, Cambridge.

COOK, C.D.K. 2004. Ac/uatic and wetland plants of southern Africa.

Backhuys, Leiden.

CRAVEN, P. (ed.). 1999. A checklist of Namibian plant species. South

African Biodiversity Network Report No. 7. SABONET, Pre-

toria.

DITOMASO, J.M. & E.A. HEALY. 2007. Weeds of California and

other western states. University of California, Oakland.

DREGE, J.F. 1843-1844. Zwei pfianzengeogrpahische documente.

Besondere Beigabe zur Flora 1943 Band II.

EICHLER, H. & WALSH, N.G. 2007. Ranunculus. In J.G. Wilson (ed.).

Flora of Australia 2: 313-354, ABRS/CSIRO, Melbourne.

EMADZADE, K. LEHNEBACH, C„ LOCKHART, P. & HORANDL,

E. 2010. A molecular phylogeny, morphlogy and classification

of genera of Ranunculeae (Ranunculaceae). Taxon 59: 809-828.

EXELL, A.W. &. MILNE-REDHEAD, E. I960. Ranunculaceae. In

A.W. Exell & H. Wild (ed). Flora zambesiaca 1(1): 89-102.

Crown Agents for Overseas Governments and Administrations,

London.

FORSSKAL, P. \115. Flora aegyptiaco-arabica. Mbller, Kjobenhavn,

GLEN, H.F. 2003. Ranunculaceae. In G. Germishuizen & N.L. Meyer.

Plants of southern Africa: an annotated checklist'. 809. 810,

Strelitzia 14. South African National Botanical Institute, Preto-

ria.

GLEN, H.F. & GERMISHUIZEN, G. 2010. Botanical exploration of

southern Afi'ica, Ed. 2. Strelitzia 26. South African National Bio-

diversity Institute, Pretoria.

GRAY, A. 1886. Contributions to American botany. Proceedings of the

American Academy of Arts and Sciences 21 : 363-413.

GRAY, S.F. 1 821 . 4 natural arrangement of British plants. London.

HARVEY, W.H. I860. Ranunculaceae. In W.H. Harvey & O.W. Sender,

Flora capensis, vol. 1 . L. Reeve & Co., Kent.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index Herbariorum. Part. 1: The Herbaria of the World. New

York Botanical Garden, New York.

JARVIS, C. 2007. Order out of chaos: Linnean plant names and their

types. Linnean Society of London & Natural History Museum,

London.

LAMARCK, J.B.A.P.M. de. 1804. Encyclopedic methodique. Bota-

nique, vol. 6. Pancoucke, Paris.

LEDEBOUR, C.F. 1830. Flora altaica, vol. 2. Dorpat.

LINNAEUS, C. 1753. Species Plantarum. Salvius, Stockholm.

LOURTEIG, A. 1951. Ranunculaceas de Sudamerika Templada. Dar-

winiana 9: 397-604.

LOWE, R.T. 1831. Primitiae faunae et florae Maderae et Portus Sancti.

Transactions of the Cambridge Philosophical Society 4: 1-70.

LOWE, R.T. 1857, On Ranunculus creticus, cortu.saefolius and gran-

difolius of authors. Hooker s Journal of botany and Kew garden

miscellany 9: 65-75.

MACOWAN, P. & BOLUS, H. 1881. Novitates capenses. Jonrna/ of the

Linnean Society (Botany) 18: 390-397.

MILNE-REDHEAD, E. & TURRILL, W.B. 1952. Ranunculaceae. In

W.B. Turrill & E. Milne-Redhead (ed). Flora of tropica! East

Africa Ranunculaceae. Corwn Agents for the Colonies, London.

OLIVER, D. 1868. Ranunculaceae. Flora of tropical Africa 1: 4-11.

Reeve, London.

OLIVER, D 1886a. Ranunculus cooperi. Hooker's leones Plantarum

16: t. 1502.

OLIVER, D 1886b. Ranunculus baurii. Hooker's leones Plantarum 16:

t. 1503.

PERRIER DE LA BATHIE, H. 1950. Renonculacees. In H. Humbert

(ed), Flore de Madagascar et des Comores 16'. 1-31. Museum

National d'Histoire Naturelle, Paris.

PIZZARO, J, 1995. Contribution al estudio taxonbmico de Ranunuc-

ulus L. subgen. Batrachium (DC.) A. Gray (Ranunculaceae).

Lazaron 15: 21-113.

PURSH, F.T. 1813. Flora Americae septentrionalis, vol. 2. London.

SCHULTZ, F.W. 1842. Archives de la flore de France et d'Allemagne.

C. Billot & Deux Ponts, Palatinat.

SZYSZYLOWICZ, I. von. 1887. Polypetalae thalamiflorae Rehman-

nianae. Krakow.

TAMARU, M. 1992. A new classification of the family Ranunculaceae

4. Acta phytotaxonomica et geobotanica 43 : 1 39-146.

TAMURA, M. 1993. Ranunculaceae. In K. Kubitzki, J.G. Rohwer& V.

Bittrich, The Families and Genera of Vascular Plants II Flow-

ering platits-Dicotyledons-Magnoliid. Hamamelid and Catyo-

phyllid families'. 563-583. Springer, Berlin.

TAMURA. M. 1995. Angiospemtae: Ordnung Ranunculales, Fam.

Ranunculaceae, Ranunculus. In P. Hiepko, Die Natiirlichen

Pflanzenfamilien 1 7a IV: 223^96. Duncker & Humblot, Berlin.

TEKETAY, D. & EDWARDS. S. 2000. Ranunculus. In S. Edwards, M.

Tadesse, S. Demissew & I. Hedberg. Flora of Ethiopia and Eri-

trea 2( 1 ): 24—29. National Herbarium, Addis Ababa & Uppsala

University, Uppsala,

THUNBERG, C.P. 1800. Prodromus plantarum capensium, vol. 2.

Edman, Uppsala.

TOLKEN, H.R. 1965. Ranunculus baurii. The flowering plants of Afri-

ca 37: t. 1508.

VILLARS, D. 1 786. Histoire des plantes de Dauphine, vol. 1 . Grenoble.

WINTER. P.J.D. 2006. Ranunculaceae. In G. Germishuizen, N.L.

Meyer, Y. Steenkamp & M. Keith (eds), A checklist of South

Afi'ican Plants. South African Biodiversity Network Report No.

41: 725-727. SABONET, Pretoria.

WOOD, J.M. 1907. Handbook to the flora of Natal. Pietermaritzburg.

ll

5t'

'' ^*'* ’1 I

__t.,_,hj(.i ->">■ i

-•-vn Jli ■■' «*£#*«. .; *■ , *.*.

<iK - ' ■ ■*.i

3I« f . --■ ii '

^ ••”'*• ViVvif^ l«nigi^- -.. ^

■/**■■ 'M *

r:;,^ ■ .. ,

, r»».ijifc ,.. ' . il I

;. I--^K ' '*'' ' ' "-' "

»r rf JE

•, ‘

- » I ■ ' >• ■ • -'-‘S '

S<i> • ,• ;if4vjft ’4

„ .. ..(••'wrj f , ,

■i rf *

i^tAJubl#. Vo- 'iio «. -'

'-4 S'" , '. i|,.4yS

W' ■''■■'jMait

i4^4w • a 44^<i5

iH ♦' ‘

IK|«|

>fa. «>■

HI,,* a I'

$nv ■!»■„ A'|4f

> . ,■ r nyrf»w»r^/ 1.S1 I§|^f W' ■-]

. - ' <ji .iy >kpi»" - -.■ ■

h »J< ■■■,‘:4 f"*** l.'.'if

'-^nWR-l V- «(<.^ .■'•m ',^()»v4W»tV\-iSsi*'i

JVlV ,- ... ,. t,M« i j|.‘ 1

i>w>( A.

• '•» 4)»iX ttri -H '!/■«! ,’

■ * ■-■* ■ •« Iff '^4

■ , if /V- ■■‘•■, ^ - *-"^V

'T'-/ '« • .aj;-

S «, i»,< J

■ »>J( ,A '»

:sss?/;.

M»... ■■.■. ' (n, "Is,

r’Mi Wfc '* :..|«|-Ii4il

Bothalia 43,2: 197-206(2013)

A new infrageneric classiHcation for Mesembryanthemum (Aizoaceae:

Mesembryanthemoideae)

CORNELIA KLAK' ' & PETER V. BRUYNS'

Keywords: Aizoaceae, classification, Mesembiyanihennim, section, subgenus

ABSTRACT

We present a new infrageneric classification for Mesembryanthemimi L. (Aizoaceae: Mesembryanthemoideae), based on

a phylogeny ot the genus published in 2007. We re-affirm that a broad generic circumscription for Mesembiyanthemum (in-

cluding all 103 species of the Mesembryanthemoideae) is the only group in the subfamily that can be recognized unambigu-

ously by synapomorphies. We divide Mesembiyanthemum into five subgenera, of which two are new: M. subg. Mesembiyan-

themum, M. subg. Ciyophytum (N.E. Br.) Bittrich, M. subg. Opophytum (N.E. Br.) Bittrich, M. subg. Phyllobolus (N.E. Br.)

Klak and M. subg. V'olkeranthus (Gerbaulet) Klak. Furthermore, we recognize five sections in subg. Mesembiyanthemum,

six sections in subg. Phyllobolus and two sections in M. subg. Ciyophytum. Descriptions and keys to all infrageneric ta.xa

are provided.

INTRODUCTION

The Mesembryanthemoideae is one of four sub-

families currently recognized in Aizoaceae (Klak et al.

2003) and includes 103 species (Klak et al. 2007). The

Mesembryanthemoideae include both annuals and per-

ennials, leaf- as well as stem-succulents, geophytes,

highly compact, dwarf shrubs as well as woody shrubs

that may exceed 1 m in height. Many members of this

subfamily have flattened, mesomorphic leaves and, in

many of them, both the leaves and the stems are covered

with conspicuous bladder cells. The defining features

of the subfamily are koilomorphic nectaries, axile pla-

centation and the presence in the fruits of purely septal

expanding keels that reach from the central columella

to the tips of the valves (e.g. Bittrich & Hartmann 1988;

Hartmann 1991 ).

Most of the genera in the Mesembryanthemoideae

have been revised (e.g. Gerbaulet 1996a, 1996b, 1996c,

1997; Klak & Linder 1998; Klak et al. 2006) and sev-

eral were re-circumscribed on the basis of morphologi-

cal characters (e.g. Ihlenfeldt & Bittrich 1985; Bittrich

1986; Gerbaulet 1995). Over the last 40 years, generic

concepts have been modified frequently within the sub-

family (Table 1). Bittrich (1986) was the first to con-

sider the relationships between the genera and species

in detail and this led him to put forward broad generic

concepts for Phyllobolus and Mesembiyanthemum, in

which he incorporated several of the previously recog-

nized genera (Table 1 ). Although he made no formal

taxonomic changes, he proposed subdivision of the

enlarged genus Phyllobolus into five subgenera (P. subg.

Aridaria, Phyllobolus, Prenia, Sceletium, and Sphal-

manthus). Since Gerbaulet (1995) was unable to find a

synapomorphy for Bittrich ’s broad concept of Phyllo-

bolus, she recognized the genera Aridaria, Phyllobolus,

Prenia and Sceletium and provided putative synapo-

morphies for each. In addition, she modified the generic

' Bolus Herbarium, Department of Biological Sciences, University of

Cape Town, 7701 Rondebosch, South Africa. E-mail: tcomelia.klak(g

uct.ac.za (corresponding author).

concepts of Aridaria and Prenia and included the mono-

typic genera Amoebophyllum and Dactylopsis N.E.Br.

in Phyllobolus (Gerbaulet 1995). More recent investiga-

tions showed that Dacylopsis, together with Aspazoma,

is sister to Brownantlms (Klak et al. 2003), so that it is

not closely allied to Phyllobolus. Consequently it was

reinstated as a monotypic genus (Klak et al. 2006).

In Mesembiyanthemum, Bittrich (1986) included all

the annual species of the Mesembryanthemoideae (apart

from Synaptophylhim) and recognized several of the

earlier genera as subgenera (M. subg. Ciyophytum and

subg. Opophytum), but he was not able to find synapo-

morphies for Mesembryianthemum.

A preliminary molecular study by Klak et al. (2003),

which included 16 species of the Mesembryanthemoi-

deae, showed that Mesembiyanthemum was polyphyletic

and that the generic limits needed re-investigation. This

was followed by a densely sampled molecular assess-

ment of the Mesembryanthemoideae, which confirmed

the polyphyly of Mesembiyanthemum (Klak et al. 2007).

It showed, in addition, that Phyllobolus was not mono-

phyletic and that one of its species, P. splendens (=

Mesembry’anthemum splendens), was much more closely

related to the Aptenia- and 5ce/e/fi/w-groups than to the

remainder of Phyllobolus (Klak et al. 2007).

Klak et al. (2007) found that there were exceptionally

high levels of homoplasy among the Mesembryanthe-

moideae. In particular, the M. aitonis-c\adQ (M. aitonis

and M. longistyliim DC.) and the P. splendens-c\ade (P.

splendens only) were new entities that can only be cir-

cumscribed using homoplasious characters (as discussed

in detail by Klak et al. 2007). Placing these clades in

separate genera would have made them extremely dif-

ficult to identify and any newly discovered species

may be placed in such a generic framework only using

molecular techniques. Consequently, a new classification

of the Mesembryanthemoideae was proposed by Klak et

al. (2007) with only the single genus Mesembryanthe-

mum. Although not all relationships between the species

were resolved, many of the clades in the Mesembryan-

themoideae were well supported and these clades were

198 Bothalia 43,2 (2013)

TABLE 1. — Classification of Mesembryanthemoideae over the last four decades showing the fluctuation in the number of genera recognized

(in bold)

also recovered in an analysis including both molecular

and morphological characters (Klak et al. 2007) so that

they were recognized as informal groups.

Recently, these changes have been challenged and it

has been claimed that the results of Klak et al. (2007)

can easily be accommodated within the ‘traditional clas-

sification’ by reinstating a couple of older genera and

describing one new genus, Volkeranthiis, for the clade

consisting of M aitonis and M. longistyhim (Gerbau-

let 2012). This resulted in a new classification for the

Mesembryanthemoideae involving 16 genera and revert-

ing to a generic system similar to that of Herre (1971,

Table 1 ).

Gerbaulet (2012: 189) based much of her discussion

and conclusions on a ‘new’ and ‘simplified cladogram

derived from Klak et al. (2007) and with new assess-

ments of character states’. The data and methods used

to generate this cladogram are not given, nor is it clear

on what material the new assessments of characters are

based. The new cladogram differs from that in Klak et

al. (2007) in that the Aptenia-, Ariclaria-, Phylloholiis-,

Prenia- and .S'c(?/e(/»//;-groups form a polytomy, whereas

in Klak et al. (2007), M splenciens (= Phylloholiis

spleiulens) was much more closely related (with high

statistical support) to the Aptenia- and Sceletiinn-groups

than to the Phyllobolus-^rowp. Gerbaulet (2012; 191)

erroneously interprets Phylloholiis as ‘unresolved’, since

she was not able to find any synapomorphy to support

the position of M. splenciens (= Phylloholiis splenciens)

as sister to the Aptenia- and Sceletiiim-gxoups.

Apart from our objections to the use of this ‘clado-

gram’, we are unable to accept Gerbaulet’s conclusions

for several other reasons:

( 1 ) The arguments for the creation of a new genus,

Volkeranthiis, for M. aitonis and M. longistyhim, are

flawed. Firstly, Gerbaulet (2012: 188) claimed that

Bittrich (1986) had already identified these two spe-

cies as the ‘M aitonis group’. Bittrich (1986) had

principally distinguished two groups in M. subg.

Meseinhryantheiniiin and provided morphologi-

cal circumscriptions of these two groups, citing a

‘primitive leaf shape and brown papillate seeds’ as

characterstic of the M. aitonis-group and ‘slender,

cylindrical leaves with a central water-storing tissue

and smooth seeds’ as characteristie of the M. nodi-

florum-gvoup. However, he never explicitly stated

which other species belonged to the M. aitonis-group

and which belonged to the M nodiflonim-group. M.

longistyhim is intermediate between the two groups,

since it possesses almost cylindrical, slightly chan-

Bothalia 43,2 (2013)

199

FIGURE 1. — Subgenera and sections mapped onto a simplified version of the Baysian tree by Klak et al. (2007: Figure 3) based on combined

chloroplast, nuclear and morphology data. The support for the relationships between the subgenera and sections in Mesembryanihemtim are

indicated (numbers indicate Posterior Probability values).

200

Bothalia 43,2 (2013)

nelled leaves with central water-storing tissue and

papillate seeds (Gerbaulet 2001b). Thus, Gerbaulet’s

(2012) reference to Bittrich’s ‘M aitonis-group'' as

consisting of M. aitonis and M. longistylum is incor-

rect. Secondly, Gerbaulet (2012: 194) claimed that

‘epidermal cells with a central papilla at the base of

the stamens’ characterizes the new genus Volkeran-

thus (M aitonis and M. longistydiim). She referred to

Bittrich (1986) for evidence of this synapomorphy.

However, Bittrich (1986: 26) did not state in which

species this feature is present, apart from M. aitonis.

Gerbaulet (2012) presented no new data and conse-

quently her claim that ‘epidermal cells with a central

papilla at the base of the stamens are unique to M

aitonis and M. longistylum' is not justified. The fail-

ure to present such data is particularly problematic,

since this was the only feature that characterized M.

aitonis and M. longistylum as a distinct genus.

(2) A further example of a hypothesis for which

no data were provided is the character ‘bladder cells

very large, especially on the receptacle’ (Gerbau-

let 2012: 194), which is given as a synapomorphy

for Ciyophytum . However, no evidence for it exists

either in Bittrich (1986) or in any other paper by

Gerbaulet.

(3) By placing M. splendens once more in Phyl-

lobolus, Gerbaulet (2012) created a paraphyletic

Phyllobolus. This contradicts the principles of Back-

kind & Bremer ( 1998), who argued that only mono-

phyletic groups should be recognized as genera.

We therefore maintain that a broad generic circum-

scription for Mesembryanthemiim (including all 103

species of the Mesembryanthemoideae) is the only

arrangement that accurately reflects the most recently

acquired information on the subfamily and is the only

group that can be recognized unambiguously by synapo-

morphies. The aim of this paper is therefore to formal-

ize the arrangement of species into groups by Klak et

al. (2007) as an infrageneric classification for Mesem-

hryanthemum. The major lineages within Mesemhyan-

themum are recognized as subgenera, while the clades

within these lineages are recognized as sections. The

subgenera and sections have been created here for ease

of reference but, due to the high levels of homoplasy in

the Mesembryanthemoideae, not all are easy to recog-

nize. This applies particularly to M. subg. Volkeranthus

and M. sect. Splendentia. Therefore, the keys to the sub-

genera and sections provided below are not easy to use

and often have exceptions. Nevertheless they provide

a first step in the identification of groups of species in

the Mesembryanthemoideae. Where possible, the sec-

tions consist of groups that were previously recognized

as genera. All of the subgenera and sections are mono-

phyletic, except for M. sect. Canaliculata, which forms

a polytomy together with sections Vesperifolia (mono-

phyletic) and Flavi/lora (monophyletic). Resolution

obtained so far is inadequate to argue for or against the

monophyly of sect. Canaliculata, but we recognize this

group of taxa at the rank of section for ease of identi-

fication. The subgenera and sections have been mapped

onto a simplified version of the eladogram of Klak et al.

(2007: Figure 3), which shows the support for and rela-

tionships between them.

Haworth (1795) appears to have been the first to

split up the large Linnaean genus Mesembryanthemum

(which then already included 162 speeies) into groups,

but without assigning them a formal rank. He intended

to provide ‘a numerical index of the speeies and varie-

ties, arranged after [my] own method’ (Haworth 1795:

473). Several years later, Haworth (1803) arranged the

species into 13 seetions and provided short diagnoses

{‘characteres sectionum') for each of them. Most of

the 13 sections were further subdivided but without an

indication of the rank of these segregates. Salm-Dyck

(1820:13) refined Haworth’s system and further subdi-

vided Mesembryanthemum Csubdivisionis generis'),

providing names and diagnoses for his divisions.

Although Haworth (1821:71) refered to Salm-Dyck’s

subdivisions as ‘the Prince’s divisions and sections’,

Salm-Dyck (1820) assigned no formal rank to them. Due

to the growing number of species in Mesembfyanthe-

mum, Haworth (1821 ) eonsidered it necessary to modify

the divisions and he recognized eight sections, which

were further broken into 69 numbered but unranked

subdivisions. Haworth (1821:79) noted at the end of his

sections that several of his groups could be regarded as

‘true and natural genera’. Sinee his 69 subdivisions were

unranked, their names need not to be taken into consid-

eration for nomenclatural purposes.

The species currently placed in the Mesembryan-

themoideae were included in five of the 13 sections of

Haworth (1803), namely Siibacaulia, Planifolia, Canal-

iculata, Vesperifolia and Flaviflora. Whereas sections

Subacaulia and Flaviflora each included only a single

species from the Mesembryanthemoideae, most species

of the Mesembryanthemoideae were placed in sections

Planifolia and Canaliculata. All five sections had many

subdivisions, except for section Vesperifolia, which

included only M. defoliatum Haw., M. noctiflorum L.

and M. stramineum Haw. Haworth ( 1821 ) placed all spe-

cies of the Mesembryanthemoideae in section Papulosa

Cfoliis fere semper plus minus papulosis'), except for

the more-or-less stemless, clump-forming, highly succu-

lent M. digitatum, which he included in section Acaiilia.

However, section Papulosa also included many taxa

which belong to the Ruschioideae, such as various spe-

cies of Delosperma and Cleretum.

Candolle (1828) based his classification on the ear-

lier works of Haworth and Salm-Dyck, and used many

of Haworth’s ‘sections’ at the rank of ‘subdivision’ and

Haworth’s groupings within a section at the rank of ‘see-

tions’. He recognized eight ‘subdivisions’ and 54 ‘sec-

tions’, including some new sections. Candolle (1828)

made many changes to the membership and circumscrip-

tion of the groupings of Haworth and Salm-Dyck, but

regarded two species, i.e. M. ciliatum Ait. (= M. vagi-

natum Lam.) and M. corallinum Thunb. as insufficiently

known and did not plaee them in any section (1828:

45 1 ). These two species were later included by Sonder

( 1 862) in sect. Juncea.

Most sections listed below were thus previously val-

idly published, but not typified. As their membership has

ehanged repeatedly since they were set up, they are typi-

fied below and are more accurately cireumscribed.

Bothalia 43,2 (2013)

201

TAXONOMIC TREATMENT

Mesembryanthemum L., Species plantamm 1: 480

(1753), nom. cons. Type; Mesembry’anthernum nodiflo-

nim L., typ. cons.

Key to subgenera of Mesembtyanthemum

1. Seeds whitish to light brown, rarely dark brown and sculptured ... 2

1. ’ Seeds dark brown, much sculptured (whitish and smooth in M

inachabeme but then stigmas red) 3

2. (1) Annuals; leaves highly succulent, cylindrical and 7-15 mm

diam 3. M subg. Opophytum

2’. Annuals or perennials, if annuals then leaves flat or if ± cylin-

drical then 2-5 mm diam 2. M subg. Mesembryanthemum

3. (1’) Plants perennial; testa cells convex or with central raised

papillae 4. A/, subg. Phyllobolus

3’. Plants annual or short-lived perennials; testa cells with central

raised papillae 4

4. (3’) Petaloid staminodes free, filamentous staminodes absent .

5. M. subg. Volkeranihus

4.’ Petaloid staminodes fused into tube, filamentous staminodes

present (rarely absent) M. subg. Ciyophytiim

1. M. subg. Cryophytum (N.E.Br.) Bittrich in Mit-

teilungen aus dem Institut fiir allgemeine Botanik Ham-

burg 21: 72 (1986). Cryophytum N.E.Br.; 412 (1925).

Lectotype (Bittrich 1986: 72): M crystallirnim L.

Callistigma Dinter & Schwantes in Schwantes:

644 (1928). Type: Callistigma itrachabense (Engler)

Dinter & Schwantes (= Mesembryanthemum irrach-

abense Engler).

Prostrate to ascending or erect annuals or short-lived

perennials, stems often angled or winged, epidermis

of stems with conspicuously raised bladder cells, roots

fibrous. Leaves flat and often very broad, decussate and

shortly connate at the base or becoming alternate in the

inflorescences, margins often undulating, epidermis with

conspicuous bladder idioblasts, without enlarged central

water storing cells. Flowers in cymes, 10-60 mm diam.,

cream to pale or rarely dark yellow, petaloid staminodes

fused into tube, petaloid and filamentous staminodes and

stamens very numerous and filiform, rarely filamentous

staminodes absent and petaloid staminodes and stamens

few (M inachabense); stigmas free or rarely basally

connate and red (M inachabense), nectaries shell-

shaped, often deep. Fruits 5-locular, valve wings erect

or indexed over valves. Seeds brown, rarely whithish,

rough with testa cells central papillose, rarely smooth.

Key to sections of M subg. Cryophytum

1. Margins of leaves not undulate; flowers 10-20 mm diam.;

filamentous staminodes absent, petaloid staminodes and

stamens few \.\. M. sect. Callistigma

1.’ Margins of leaves undulate; flowers 15-60 mm diam.; petaloid

staminodes, filamentous staminodes and stamens numerous

\.2. M. sect. Papulosa

1.1. M. sect. Callistigma (Dirrter & Schwantes) Klak,

comb, et stat. nov. Callistigma Dinter & Schwantes in

Schwantes in Gartenwelt 32; 644 (1928). Type: Mesem-

bryanthemum inachabense Engler (= Callistigma inach-

abense (Engler) Dinter & Schwantes).

Decumbent annuals, stems terete or 2-edged. Leaves

flat, oblong, margins not undulate, epidermis with con-

spicuous bladder idioblasts. Flowers 10-20 mm diam.,

petaloid staminodes pale to dark yellow, filamentous

staminodes absent, petaloid staminodes and stamens

few; stigmas basally connate and red, nectaries deep.

Fntits 5-locular, valve wings erect or inflexed over the

valves. Seeds whitish, testa smooth.

Species ( 1 ): Mesernbryanthernurrr inachabense Engler

1.2. M. sect. Papulosa Haw., Revisiones plantarum

succulentarum : 79 (1821). Type (designated here):

Mesembryanthemum crystallinurn L.

Prostrate to ascending or erect annuals or short-lived

perennials, stems often angled or winged. Leaves flat

and often very broad, margins usually undulate, decus-

sate and shortly connate at the base or becoming alter-

nate in inflorescences, epidermis with conspicuous

bladder idioblasts. Flowers 15-60 mm diam., cream to

pale yellow, with numerous filiform petaloid stamin-

odes, filamentous staminodes and stamens; stigmas free,

greenish-yellow, nectaries shell-shaped. Fruits 5-locu-

lar, valve wings inflexed over valves. Seeds dark or light

brown, rough, testa cells with central papillae.

Typification: Haworth (1803) included all species

of Mesembryanthemoideae and Ruschioideae with flat

leaves and a prostrate to decumbent habit in section

Planifolia, including M. crystallirnim (Cryophytum-

group), M. cordifoliurn {Aptenia-gxovrp), M. expansurn

{Sceletiurn-grovyr) and M. pallens {Prenia-growp) (see

also typification for sections Planifolia and Flaviflora).

In a subsequent publication, however, all species of

Mesembryanthemoideae (except M. digitaturn) were

accommodated in section Papulosa (Haworth 1821).

Section Papulosa included annuals, biennials and per-

ennials, with more-or-less papillate leaves PSuffrutices;

annua, biennia, perermiave: folds fere semper plus

minus papulosis' (Haworth 1821). Although most mem-

bers of the Mesembryanthemoideae have distinctly pap-

illate leaves, the size of the papillae are most striking in

the Cryophyturn-gxou^. We therefore selected seetion

Papulosa to accommodate all those members of subge-

nus Cryophytum that have particularly large and striking

bladder cells. Since M. cry’stalliniirn was the only spe-

cies from this group to be known and included in section

Papulosa by Haworth, we chose this species as the type

of section Papulosa.

Species (6): M. barklyi N.E.Br., M. crystallinurn L.,

M. gariusanurn Dinter, M. giierichiarnirn Pax, M. longi-

papillosurn Dinter, M. pelliturn Friedrich

2. M. subg. Mesembryanthemum

Annuals or perennials, prostrate, decumbent or erect

shrubs, rarely geophytes, stems cylindrical, often with

ephemeral leaves and surface remaining green and pho-

tosynthetic, roots fibrous or rarely thickened, epidermis

of stems with conspicuously raised bladder cells, or

bladder cells reduced and flattened. Leaves deciduous or

marcescent, epidermal bladder cells distinct or flattened,

without or with central enlarged water storing cells.

Flowers in cymes or solitary, 5-20(-30) mm diam.,

sepals remaining erect during anthesis or spreading, pet-

202

Bothalia 43,2 (2013)

aloid staminodes, filamentous staminodes and stamens

not particularly numerous, with or without filamentous

staminodes, petaloid staminodes white, cream, pink or

pale yellow, rarely much reduced, nectaries shell-shaped

or tubular, rarely absent. Fruits (3)4 or 5-locular, rarely

nut-like, valve wings indexed over valves or reflexed

and fused in pairs, locules deep or shallow. Seeds

smooth or testal cells with central raised papillae, light

coloured (rarely brown), rarely with crest.

Key to the sections of M. subg. Mesemhyanthemum

1 .Annuals 2

1 .’Perennials 4

2.(1) Flowers 5-30 mm diam.; filamentous staminodes absent;

petaloid staminodes and stamens not particularly numer-

ous 2.4. M. sect. Mesembryanthemum

2. ’ Flowers ± 60 mm diam.; petaloid staminodes, filamentous

staminodes and stamens very numerous

2.2. M. sect. Ewystigma

3. ( 1’) Leaves free, with tubular stem-clasping sheath

2.5. M. sect. Siibacaidia

3. ’ Leaves free or shortly connate, without tubular stem-clasping

sheath 4

4. (3’) Epidermis of stems with cylindrical, closely packed, xero-

morphic bladder cells resembling cobblestones

2.1. M sect. Brownanihus

4.’ Epidermis of stems with much flattened epidermal bladder

cells, smooth, sometimes with widely-spaced, rounded or

hair-like bladder cells 2.3. M. sect Jimcea

2.1. M. sect. Brownanthus (Schwantes) Klak, comb,

et stat. nov. Brownanthus Schwantes in Zeitschrift fur

Sukkulentenkunde 3: 20 (1927). Type: Mesembryanthe-

mum vaginatum Lam. (= Brownanthus vaginatiis (Lam.)

Chesselet & M.Pignal.

Pseudobrownanthus Ihlenf. & Bittrich:

319 (1985). Type: Pseudobrownanthus nucifer

Ihlenf.&Bittrich. (= Mesembiyanthemum nucifer (Ihlenf.

& Bittrich) Klak).

Decumbent to erect perennial shrubs, at least young-

est stems succulent and green, articulate, epidermis of

stems with closely packed, xeromophic bladder cells.

Leaves decussate, deciduous or marcescent, epidermal

bladder cells mesomorphic, with central enlarged water

storing cells. Flowers solitary or in cymes, 5-10(-20)

mm diam., sepals remaining erect during anthesis, peta-

loid staminodes cream to white, free, filamentous stami-

nodes absent, nectaries narrow. Fruits 3-5-locular, valve

wings indexed over valves, lower part of fruit shal-

low, occasionally with seed bags, rarely nut-like. Seeds

brown to cream, testa rough or ± smooth, rarely with

crest.

Typification: Mesembiyanthemum vaginatum is

the type of Brownanthus and thus the type for section

Brownanthus.

Species (14): M. arenosum Schinz, M. corallinum

Thunb., M. glareicola (Klak) Klak, M. kuntzei Schinz,

M. marlothii Pax, M. namibense Marloth, M. napierense

Klak, M. neglectum (Pierce & Gerbaulet) Klak, M. nuci-

fer (Ihlenf & Bittrich) Klak, M. pseudoschlichtianum

(Pierce & Gerbaulet) Klak, M. schenckii Schinz, M.

springbokense Klak, M. tomentosum Klak, M. vagina-

tum Lam.

2.2. M. sect. Eurystigma (L. Bolus) Klak, comb, et

stat. nov. Eurystigma L. Bolus in Notes on Mesembrian-

themum and allied genera 2: 179 (1930). Type: Mesem-

biyanthemum eurystigmatum Gerbaulet (= Eurystigma

clavatum L. Bolus).

Decumbent annuals to 20 cm tall, intemodes terete, to

4 mm diam. Leaves cylindrical, to 70 x 10 mm, bladder

cells flattened, with central enlarged water storing cells.

Flowers in cymes, deeply funnel-shaped, to 60 mm

diam., petaloid staminodes straw-coloured to yellow,

very narrow, connate into tube, petaloid staminodes, fila-

mentous staminodes and stamens very numerous; stig-

mas basally shortly connate with ovary, thick and broad,

subulate, or rather filiform, yellow, nectaries tubular.

Fruits 5-locular, valve wings indexed over valves. Seeds

whitish to light brown, testa smooth.

Species ( 1 ): M euiystigmatum Gerbaulet

2.3. M. sect. Juncea Haw. ex DC., Prodromus

systematis naturalis regni vegetabilis 3: 446 (1828).

[Unranked] Juncea Haw.: 175 (1821). Lectotype (desig-

nated here): Mesembiyanthemum juceum Haw.

Psilocaulon N.E.Br.: 433 (1925). Type: Psilo-

caulon articulatum (Thunb.) N.E.Br. (= Mesembryanthe-

mum articiilatum Thunb.).

Caulipsolon Klak: 364 (1998). Type: Caulip-

solon rapaceum (Jacq.) Klak (= Mesembryanthemum

rapaceum Jacq.).

Synaptophylluin N.E.Br.: 412 (1925). Type:

Synaptophyllum juttae (Dinter & A. Berger) N.E.Br. (=

Mesembryanthemum juttae Dinter & A. Berger).

[Unranked] Articulata Salm-Dyck: 24 (1820).

Small to large prostrate to erect shrubby perennials

or rarely annuals or geophytes (M rapaceum), at least

youngest stems succulent and green, usually articulate,

epidermis of stems and leaves similar, with much flat-

tened bladder cells. Leaves cylindrical to slightly trigo-

nous, rarely flat and broad, deciduous, without central

enlarged water storing cells. Flowers in cymes, rarely

solitary, 5-25 mm diam., petaloid staminodes free or

very shortly connate at base, filamentous staminodes and

stamens conically collected, rarely petaloid staminodes

or filamentous staminodes absent, nectaries narrowly

shell-shaped or absent. Fruits 4 or 5-locular, valve wings

indexed over valves, rarely reflexed and fused in pairs,

usually with deep locules. Seeds usually light brown to

ochre, rarely whitish, testa slightly sculptured, with cen-

tral raised papillae, rarely smooth.

Typification: Salm-Dyck ( 1 820) created subdivision

''Articulata'’ (without formal rank) to include three spe-

cies from the Psilocaulon-%xo\i\), which Haworth (1821)

placed in his subdivision ''Juncea’ (without formal

rank) within section Papulosa'. M. junceum, M. parvi-

fiorum (=M micranthon Haw., =M. tenue Haw.) and

M. rapaceum. Constricted green stems and deciduous

leaves were mentioned by both authors in the diagnoses

Bothalia 43,2 (2013)

203

of their respective subdivision, although these charac-

teristics are also found in sections Brownanthus and

Geniculiflora. Candolle (1828) treated Haworth’s subdi-

vision "Jimcea' as a section, thereby validating it for the

first time at this rank. In addition, he only included taxa

currenly placed in the Psilocaii Ion-group, making it the

correct name for this group at the rank of section. Since

M. jimceum is the eponymous species of this group we

selected it as the type of sect. Jimcea.

Species (15): M. articiilatiim Thunb., M. bicorne

Sonder, M. coriarium Burch, ex N.E.Br., M. dimorphum

Welw. ex Oliver, M. dinteri Engler, M. gessertiamim

Dinter & A. Berger, M. gramilicaule Haw., M. jimceum

Haw., M. jiittae Dinter & A. Berger, M leptarthron

A.Berger, M. ueofoliosiim Klak, M. pai-viflonim Jacq.,

M. rapaceum Jacq., M. salicornioides Pax, M. siibnodo-

sum A.Berger

2.4. M. sect. Mesembryanthemum

Annual, prostrate or erect shrublets, intemodes terete.

Leaves ± cylindrical or somewhat channelled to nar-

rowly obovate or oblong or almost truncate, epidermal

bladder cells distinct or inconspicuous, with enlarged

central water storing cells. Flowers in cymes, petaloid

staminodes free or connate into short tube, 5-30 mm

diam., petaloid staminodes yellow, white to pale pink or

pink, filamentous staminodes absent, nectaries tubular.

Fruits 5-locular, valve wings reflexed and fused in pairs.

Seeds whitish to light brown, testa smooth.

Species (4); M. excavatiim L. Bolus, M. uodifloriim

L., M. stenandrum (L. Bolus) L. Bolus, M. subtnmcatum

L. Bolus

2.5. M. sect. Subacaulia Flaw, in Miscellanea natu-

ralia: 17 (1803): Lectotype (designated here): Mesem-

bryanthemum digitatum Aiton

Aspazoma N.E.Br.: 413 (1925). Type: Aspa-

zoma amplectens (L. Bolus) N.E.Br.

Dactylopsis N.E.Br.: 413 (1925). Type: Dacty-

lopsis digitata (Aiton) N.E.Br.

Dwarf, clump-forming or bushily branched, erect

shrublets, epidermis of stem with closely packed blad-

der cells. Leaves cylindrical or slightly flattened, alter-

nating, with tubular sheaths clasping one another, drying

up completely during resting period, bladder cells much

flattened or somewhat flattened, with central enlarged

water storing cells. Flowers solitary, sepals and petaloid

staminodes basally fused into short tube, petaloid stami-

nodes white to cream, filamentous staminodes absent or

present, if absent then reproductive parts conceiled, nec-

taries shell-shaped. Fruits 4 or 5-locular, valve wings

indexed over valves or reflexed and fused in pairs. Seeds

light brown to whitish, testa smooth.

Typification: Among several species of Ruschioi-

deae, only a single species of Mesembryanthemoideae

was included in section Subacaulia (Haworth 1803),

which Haworth characterized as lacking stems or with

very short stems and perennial roots C'caulibus nullis

vel brevissimus, radice perennV). Among the species

of Mesembryanthemoideae currently known a seem-

ingly stem-less habit is still unique to M. digitatum. The

closely related M. amplectens is a shrubby plant but

shares the unique leaf morphology with M. digitatum.

Species (2): M. amplectens L. Bolus, M. digitatum

Aiton subsp. digitatum, M. digitatum subsp. littlewoodii

(L. Bolus) Klak

3. M. subg. Opophytum (N.E.Br.) Bittrich in Mit-

teilungen aus dem Institut fur allgemeine Botanik Ham-

burg 21: 73 (1986). Opophytum N.E.Br.: 412 (1925).

Lectotype (Bittrich 1986: 73): Mesembryanthemum fas-

tigiatum Thunb.

Halenbergia Dinter ex H. Jacobsen: 158, 200

(1937). Type: Halenbergia hypertrophica (Dinter) Din-

ter ex H. Jacobsen (= M. hypertrophicum Dinter).

Hydrodea N.E.Br.: 412 (1925). Type: Hydro-

dea cry'ptantha (Hook.f) N.E.Br. (= M. ciyptanthum

Hook.f).

Prostrate annuals, intemodes terete, roots fibrous,

epidermis of stems with much flattened bladder cells.

Leaves highly succulent, cylindrical or globular, decus-

sate and shortly connate at base or sometimes alternate

in inflorescence, becoming dry while plants are still in

flower, with much flattened epidermal bladder cells, with

enlarged central water storing cells. Flowers 20-60(-80)

mm diam., rarely much reduced and only 10 mm diam.,

petaloid staminodes filiform, sepals and petaloid stami-

nodes connate into short tube, petaloid staminodes, fila-

mentous staminodes and stamens numerous, nectaries

tubular. Fruits 5-locular, valve wings reflexed and fused

in pairs. Seeds ochre, smooth.

Species (3): M. ciyptanthum Hook.f, M. fastigiatum

Thunb., M. hypertrophicum Dinter

4. M. subg. Phyllobolus (N.E.Br.) Klak, comb, et stat.

nov. Phyllobolus N.E.Br. in Gardener’s Chronicle 78:

413 (1925). Type: Mesembryanthemum resurgens Kensit

(= Phyllobolus resurgens (Kensit) Schwantes).

Prostrate or erect shmbs or geophytes, stems herba-

ceous or corky or woody, roots fibrous or thickened, epi-

dermis of stems with mesomorphic bladder cells, mostly

distinct. Leaves subcylindrical to somewhat flattened or

flat, decussate and shortly connate at base or becoming

alternate in inflorescence, or alternate throughout, decid-

uous or marcescent or old dry leaves ‘skeletonized’, ±

retaining shape through lignified veins, epidermal blad-

der cells distinct or flattened. Elowers in cymes or soli-

tary, white, yellow, pink, greenish or greenish-blue,

10^0 mm diam, sepals spreading during anthesis. peta-

loid staminodes, filamentous staminodes and stamens

not particularly numerous, with or without filamentous

staminodes, reproductive parts sometimes concealed,

nectaries shell-shaped. Fruits 4 or 5-locular, valve wings

indexed over valves or reflexed and fused in pairs, rarely

absent, locules deep or shallow. Seeds dark brown, with

testa cells convex or testa with central raised papillae,

with or without a crest.

Key to the sections of M. subg. Phyllobolus

i. Old dr\' leaves ‘sceletonized’ (retaining their shape ± fully

through lignified veins) 4.4. sect. Planifolia

204

Bothalia 43,2 (2013)

1. ’ Old dry leaves sometimes becoming spiny, but not

"sceletonized’ 2

2. (r)Epidermis of stems with tall, cylindrical, closely packed,

xeromorphic bladder cells 4. 5. sect. Geniculiflora

2. ’ Epidermis of stems with much flattened bladder cells 3

3. (2') Epidermis of leaves with thick wax layer, bladder cells flat-

tened 4

3. ' Epidermis of leaves with inconspicuous wax layer, bladder

cells rarely flattened 5

4. Stems herbaceous or only weakly lignified; lower part of fruits

conspicuously longer than upper part 4.2. sect. Flaviflora

4. 'Stems woody, leaves strictly decussate; lower part of fruit ± as

long as upper part 4.3. sect. Vesperifolia

5. (3’) Large woody shrubs, to 1 m tall, rarely cushion-like,

leaves persistent, never marcescent or spiny, decussate

becoming alternate in the inflorescences . . .4.1. sect.

Splendentia

5.’ Decumbent or erect shrubs or prostrate to decumbent geo-

phytes, if decumbent or erect shrubs, leaves usually decidu-

ous, marcescent or spiny, rarely persistent, if persistent, new

leaves tutted in old and dry leaves 4.6. sect. Canaliculata

4.1. M. sect. Splendentia DC., Prodromus systematis

naturalis regni vegetabilis 3: 445 (1828). Lectotype (des-

ignated here): Mesembjyanthemum spleudens L.

Erect perennial shrubs to I m tall, stems woody, roots

not thickened. Leaves variable in shape, from subcylin-

drical to narrowly ovate, persistent, epidermal bladder

cells small, central water storing cells indistinct. Flowers

in cymes, 30^0 mm diam., petaloid staminodes cream,

pale pink, pale salmon or pale yellow, filamentous stami-

nodes present, reproductive parts not concealed. Fruits

5-locular, valve wings inflexed over valves. Seeds black

or dark brown, with rough testa, crest distinct, indistinct

or absent.

Typification: Most taxa included in sect. Splendentia

by Candolle (1828) are now considered to be conspe-

cific with M. splendens, but the section also included M

longistyliirn (now placed in subg. Volkeranthiis) and two

species of Delosperma (Ruschioideae).

Species ( 1 ); M. splendens L. subsp. splendens, M.

splendens suhs^. pentagoniim (L. Bolus) Klak

4.2. M. sect. Flaviflora Flaw, in Miscellanea natu-

ralia: 19 (1803). Lectotype (designated here): Mesem-

hryanthemwn tetragonum Thunb.

Prenia N.E.Br.: 412 (1925). Type: Prenia pal-

lens (Alton) N.E.Br. (= M. pallens Alton).

Prostrate to rarely ascending (M tetragonum) peren-

nials, stems weakly lignified, roots fibrous. Leaves ± flat

and obscurely triquetrous, obtusely trigonous or almost

cylindrical, persistent, epidermal bladder cells much

flattened, without enlarged central water storing cells.

Flowers in cymes, 20^0 mm diam., petaloid stamin-

odes white yellow or pink, sepals and petaloid stamin-

odes connate into short tube, filamentous staminodes

present, nectaries shell-shaped. Fruits 4 or 5 locular, loc-

ales deep, valve wings inflexed over valves. Seeds black,

rough, testa cells central papillose, with or without indis-

tinct crest.

Typification: Haworth (1803) included several species

of Ruschioideae and only one species of Mesembryan-

themoideae in section Flaviflora, i.e. M. tetragonum

Thunb. We take this species as the type of this section.

It was included by Haworth on the ‘authority of Thun-

berg’, who thought it was related to M. mode Alton (=

Malephora mollis (Alton) N.E.Br.). Section Flaviflora

was characterized by slightly woody, often erect stems,

triquetrous leaves and yellow, orange, or deep red pet-

aloid staminodes Ccaulibus suffrutescentibiis saepe

erectis, foliis triquetris saepe brevissimis, petalis flavis,

aurantiis, vel coccineis') (Haworth 1803: 19).

Species (6): M englishiae L. Bolus, M pallens subsp.

lanceum (Thunb.) Klak, M pallens subsp. luteum

(L. Bolus) Klak, M. pallens subsp. namaqiiense (Ger-

baulet) Klak, M. pallens Alton subsp. pallens, M. radi-

cans (L. Bolus) Klak, M sladenianum L. Bolus, M

tetragonum Thunb., M. vanrensburgii (L. Bolus) Klak

4.3. M. sect. Vesperifolia Haw. in Miscellanea natu-

ralia: 18 (1803). Type : MesemFyanthemum noctiflorum

L.

Aridaria N.E.Br.: 433 (1925). Type: Aridaria

noctiflora (L.) Schwantes (= M noctiflorum L.).

Low to tall erect shrubs to 1 m high, roots thickened.

Leaves subcylindrical, decussate, deciduous, bladder

cells small and much flattened, with enlarged central

water storing cells. Flowers in cymes, large (to 40 mm

diam.), diurnal or nocturnal, petaloid staminodes white

or pink inside, suffused with various shades of yellow,

copper, pink, or red on reverse, sepals and petaloid

staminodes connate into short tube, filamentous stami-

nodes absent, nectaries shell-shaped. Fruits 4-locular,

valve wings inflexed over valves, breaking off easily

with stalks becoming spiny. Seeds brown or dark brown,

rough, testa cells central papillose, with or without crest.

Typification: Haworth (1803) included three species

in sect. Vesperifolia (i.e. M. noctiflorum, M. stramineum

and M. defoliatum). The last two are now subspecies of

M. noctiflorum, which therefore becomes the type of the

section.

Species (4): M. brevicarpum (L. Bolus) Klak, M noc-

tiflorum subsp. defoliatum (Haw.) Klak, M noctiflorum

L. subsp. noctiflorum, M. noctiflorum subsp. strami-

neum (Haw.) Klak, M. occidentale Klak, M. serotinum

(L. Bolus) Klak

4.4. M. sect. Planifolia Haw. in Miscellanea natu-

ralia: 18 (1803). Lectotype (designated here): Mesem-

btyanthemum tortuosum L.

Sceletium N.E.Br.: 412 (1925). Type: Sceletium

tortuosum (L.) N.E.Br. (= M tortuosum L.).

Prostrate to erect small perennials, often scram-

bling in other bushes, roots fibrous. Leaves flat, ovate,

dry leaves persistent and ‘skeletonized’ through ligni-

fied veins, without enlarged central water storing cells.

Flowers in few-flowered cymes, more rarely solitary,

stalked or sessile, 20^0 mm diam., petaloid stamin-

odes white, various shades of yellow, pale pink, or pale

salmon, sepals and petaloid staminodes basally connate

Bothalia 43,2 (2013)

205

into short tube, filamentous staminodes present, some-

times concealing reproductive parts. Fruits 4— 6-locular,

with or without valve wings, valve wings indexed over

valves, breaking off easily to be dispersed by wind.

Seeds rough, testa cells convex and arranged in concen-

tric rows, with distinct or indistinct crest.

Typification: Haworth (1803) placed all species of

Mesembryanthemoideae and Ruschioideae with flat

leaves and a prostrate to decumbent habit in sect. Plani-

folia Cfoliis plains saepe subcarinatis, caulibits saepe

decumbenti— prostrates ejfiisis' . From Mesembryanthe-

moideae, he included M. ciystallinum, M. cordifolium,

M. expansmn, M. emarcidum, M. tortiiosum, M. var-

ians and M. pallens. Although all of these species fit the

description of the section well, they are unrelated and

cannot be placed in the same section. Since most species

listed fall into the Sceletiiim-groui^ we elected to associ-

ate sect. Planifolia with this group, by selecting M. tor-

tiioswn as the type of the section.

Species (8): M. archeri Klak, M. crassicaiile Haw.,

M. emarcidiim Thunb., M e.xalatum (Gerbaulet) Klak,

M. expansum L., M. ladismithiense Klak, M tortuosum

L., M varians Haw.

4.5. M. sect. Geniculiflora DC. in Prodromus sys-

tematis naturalis regni vegetabilis 3: 445 (1828). Type:

Mesembryanthemum geniculiflorum L.

Aptenia N.E.Br.: 412 (1925), Type: Aptenia

cordifolia (L.f.) Schwantes (= M. cordifoUum L.f).

Platythyra N.E.Br.: 412 (1925), Type:

Platythyra haeckeliana (A. Berger) N.E.Br. (= M haeck-

elianum A. Berger).

M sect. Cordifolia DC.: 449 (1828). Type:

Mesembiyanthemiim cordifoUum E.f.

Prostrate to scandent or climbing perennials, stems

with persistent green succulent cortex, intemodes

4-angled or terete, with closely packed, xeromorphic

bladder cells, roots fibrous, rarely fleshy. Leaves flat or

rarely almost cylindrical, cordate, ovate, lanceolate or

linear, petiolate or sessile, bladder cells mesomorphic

and loosely arranged, without enlarged central water

storing cells. Flowers solitary or in few-flowered cymes,

shortly stalked, 15-35 mm diam., petaloid staminodes

white, pale yellow or pink, basally connate into short

tube, with or without filamentous staminodes, nectar-

ies shell-shaped. Fruits 4-locular, with or without valve

wings, valve wings indexed over valves. Seeds rough,

testa cells central papillose, without a crest.

Typification: Candolle (1828) included only a single

species in sect. Geniculiflora, thus its type. Sect. Cor-

difoliae DC. (1828), described only a few pages after

section Geniculiflora, is included here in sect. Genicu-

liflora. Four sepals and papillate leaves are among the

diagnostic characters for sect. Geniculiflora (Candolle

1828) and his mention of 5 sepals for sect. Cordifoliae is

evidently incorrect (Gerbaulet, 2001a).

Species (4): M. cordifoUum L.f, M. geniculiflorum

L., M. haeckelianum A. Berger, M. lancifolium (L. Bolus)

Klak.

4.6. M. sect. Canaliculata Haw. in Miscellanea natu-

ralia: 18 (1803). Lectotype (designated here): Mesem-

bryanthemum canaliculatum Haw.

Amoebophyllum N.E.Br.: 433 (1925). Type:

Amoebophyllum angiistum N.E.Br. (= M. vanheerdei

(L. Bolus) Klak).

Phylloboliis N.E.Br.: 413 (1925). Type: Phyl-

loboliis resurgens (Kensit) Schwantes (= M. resurgens

Kensit).

Sphalmanthus N.E.Br.: 433 (1925). Type:

Sphalmantlms canaliculatus (Haw.) N.E.Br. (= M.

canaliculatum Haw.).

[Unranked] Canaliculata. Spinulifera Haw.: 56

(1803).

[Unranked] Spinulifera Haw.: 176 (1821).

Perennials, prostrate to erect shrubs to minute geo-

phytes, stems corky or woody, long or much shortened,

roots often thickened. Leaves subcylindrical or flat-

tened, deciduous, marcescent or persistent, epidermal

bladder cells mostly distinct, central water storing cells

indistinct. Flowers in cymes or solitary, petaloid stami-

nodes in various shades of green, yellow, orange, red, or

pink or almost white, ( 10-)20-40 mm diam., sepals and

petaloid staminodes basally fused into short tube, fila-

mentous staminodes present and reproductive parts not

concealed or filamentous staminodes absent and repro-

ductive parts concealed, nectaries shell-shaped. Fruits

4- or 5-locular, valve wings indexed over valves. Seeds

rough, testa cells convex and arranged in concentric

rows, with or withour crest, sometimes with additional

crests on sides.

Typification: Whereas sect. Planifolia included taxa

with flat, often broad leaves, sect. Canaliculata (Haw.

1803: 18) accommodated taxa with more slender leaves

Cfoliis linearibus, jiinioribus canaliculatis subtus con-

vexis'). Taxa with slender leaves are found in unrelated

clades within Mesembryanthemoideae. Section Canal-

iculata had been subdivided further into the unranked

subdivisions '’Canaliculata. Spinulifera'’ (Haw. 1803:56)

and "Canaliculata. Varia' (Haw. 1803:58). Of these two

groups, ‘’Canaliculata. Spinulifera' incorporated sev-

eral species previously placed in Phylloboliis, whereas

"Canaliculata. Varia' included taxa that fall into various

unrelated clades. We therefore elected that sect. Canalic-

ulata should correspond to the group of taxa previously

placed in Phylloboliis. Surprisingly many species from

this group were already known to Haworth, including M.

canaliculatum, M. grossum, M. nitidum, M. spinuliferum

and M. viridfiorum. Since slightly channelled leaves are

common to all of these species, we chose M. canalicula-

tum as the type of the section.

Species (32): M. amabile (Gerbaulet & Struck) Klak,

M. baylissii Klak, M. bulletrapense Klak, M. canalicu-

latum Haw., M. caudatum L. Bolus, M. clvysophthal-

mum (Gerbaulet & Struck) Klak, M. deciduum (L. Bolus)

Klak, M. decun’atiim (L. Bolus) Klak, M. deliim L. Bolus,

M. flavidum Klak, M. gariepense (Gerbaulet & Struck)

Klak, M. grossum Alton, M. holense Klak, M. knolfon-

teinense Klak, M. latipetalum (L. Bolus) Klak, M. lignes-

206

Bothalia 43,2 (2013)

cens (L. Bolus) Klak, M. ligneum (L. Bolus) Klak, M. lil-

lipiitamim Klak, M. nitidum Haw., M. oculatum N.E.Br.,

M. oitbergense (L. Bolus) Klak, M. prasimim (L. Bolus)

Klak, M. quartziticola Klak, M. rabiei (L. Bolus) Klak,

M. resiagens Kensit, M sinuosiim L. Bolus, M. spinu-

life7'um Haw., M. suffruticosum (L. Bolus) Klak, M. temi-

iflorum Jacq., M trichotomum Thunb., M vanheerdei

(L. Bolus) Klak, M. viridiflomm A\ion.

5. M. subg. Volkeranthus (Gerbaidet) Klak, comb,

et stat. nov. Volkeranthus Gerbaulet in Bradleya 30: 196

(2013). Type: Volkeranthus aitonis (Jacq.) Gerbaulet (=

Mesembryanthemum aitonis Jacq.).

Prostrate to ascending, annual to biennial, herbaceous

plants, epidermis of stems with mesomorphic, rounded

bladder cells, roots fibrous. Leaves flat and broad or

almost cylindrical to flattened, slightly channelled, lin-

ear to narrowly oblong, decussate and shortly connate

at base or becoming alternate in inflorescences, epider-

mis with mesomorphic, rounded bladder cells, with or

without enlarged central water storing cells. Flowers in

cymes, to 25 mm diam., white, sometimes suffused pink

or yellowish, sepals shortly connate, petaloid staminodes

narrowly lanceolate, filamentous staminodes absent,

petaloid staminodes and stamens free, nectaries as deep

as locules and tubular. Fruits 5-locular, valve wings

reflexed and fused in pairs or free and incurved over

valves. Seeds brown with rough testa, without crest.

Species (2): M. aitonis Jacq., M. longistylum DC.

REFERENCES

BACKLUND, A. & BREMER, K.. 1998. To be or not to be— princi-

ples of classification and monotypic plant families. Taxon 47:

391-tOO.

BITTRICH, V. 1986: Untersuchungen zu Merkmalsbestand, Glieder-

ung und Abgrenzung der Unterfamilie Mesembryanthemoideae

(Mesembryanthemaceae Fenzl.). Mitteihmgen aus dem InsUtut

fiir allgemeine Botanik Hamburg 21: 5-116.

BITTRICH, V. & HARTMANN, H.E.K. 1988: The Aizoaceae— a new

approach. BotanicalJournal of the Linnean SocieW 97: 239-254.

BOLUS, H.M.L. 1930. Notes on Mesembrianthemum and allied gen-

era, part 2: 1 79. Bolus Herbarium, University of Cape Town.

BROWN, N.E. 1925. Mesembryanthemum and some new genera sepa-

rated from it. Gardeners’ Chronicle, ser. 3, 78: 412, 413, 433.

CANDOLLE, A.R de. 1 828. Ficoideae. Prodromus systematis naturalis

regni vegetabiUs 3: 415-456. Treuttel & Wiirz. Paris.

GERBAULET, M, 1995. Phyllobolus N.E.Br. emend. Bittrich

(Aizoaceae): A reassessment of generic boundaries. Botanische

Jahrbiicher 117: 385-399.

GERBAULET, M. 1996a. Revision of the genus Sceletium N.E.Br.

(Aizoaceae). Botanische Jahrbiicher 118: 9-24.

GERBAULET, M. 1996b. Revision of the genus Prenia N.E.Br.

(Aizoaceae). Botanische Jahrbiicher 1 18: 25^0.

GERBAULET, M. 1996c. Revision of the genus Aridaria N.E.Br.

(Aizoaceae). Botanische Jahrbiicher 118: 41-58.

GERBAULET, M. 1997. Revision of the genus Phyllobolus N.E.Br.

(Aizoaceae). Botanische Jahrbiicher 119: 145-211.

GERBAULET, M. 2001a. Aptenia. In H.E.K. Hartmann (ed.), Illus-

trated handbook of succulent plants: Aizoaceae A-E: 66-68.

Springer, Berlin.

GERBAULET, M. 2001b. Mesembiyanthemum. In H.E.K. Hartmann

(ed.). Illustrated handbook of succulent plants: Aizoaceae F-Z:

143-153. Springer, Berlin.

GERBAULET, M. 2012. One or many genera in Mesembryanthemoi-

deae (Aizoaceae)? Discussion of a conflict in genus perception.

Bradleya 30: 187-198.

HARTMANN, H.E.K. 1991. Mesembryanthema. Contributions from

the Bolus Herbarium 13: 75-157.

HAWORTH, A.H. 1795. Observations on the genus Mesembryanthe-

mum 2: 473^78. The author, London.

HAWORTH, A H. 1803: Miscellanea naturalia. J.Taylor. London.

HAWORTH, A.H. 1821. Revisiones plantarum succulentarum. London.

HERRE, H. 1971. The genera of Mesembiyanthemaceae. Tafelberg,

Cape Town.

IHLENFELDT, H.-D. & BITTRICH, V. 1985. Morphologic, Gliederung

und Abgrenzung der Gattung Psilocaulon N.E.Br. sJ. (Mesem-

bryanthemaceae). Botanische Jahrbiicher 105: 289-322.

JACOBSEN, H. 1937. Halophyten — Pseudosukkulenten. Kakteen und

andere Sukkulenten 10: 156-159, 199,200.

KLAK, C. & LfNDER, H.P. 1998. Systematics oi Psilocaulon N.E.Br.

and Caulipsolon Klak gen. nov. (Mesembryanthemoideae,

Aizoaceae). Botanische Jahrbiicher 120: 301-375.

KLAK, C., BRUYNS, P.V. & HEDDERSON, T.A.J. 2007. A phylogeny

and new classification for Mesembryanthemoideae (Aizoaceae).

Taxon 56: 737-756.

KLAK. C„ KHUNOU, A., REEVES, G., & HEDDERSON, T.A.J.

2003. A phylogenetic hypothesis for the Aizoaceae (Caryophyl-

lales) based on four plastid DNA regions. American Journal of

Botany 90:1433-1445.

KLAK, C., NOWELL, T.L. & HEDDERSON, T.A.J. 2006. Phylogeny

and revision of Brownanthus and its close allies Aspazoma and

Dactylopsis (Aizoaceae) based on morphology and four DNA

regions. Kew Bulletin 61 : 353-400.

LINNAEUS, C. 1753. Species plantarum, edn 1: 480. Salvius, Stock-

holm.

SALM-REIFFERSCHEIDT-DYCK, J.F. 1820. Obsen’ationes botani-

cae in horto Dyckensi notatae: 1-35. Thiriart, Koln.

SCHWANTES, G. 1927. Zur Systematik der Mesembrianthemen.

Zeitschrift fur Sukkulentenkunde 3: 14—30.

SCHWANTES, G. 1928. Mesembriaceen-Studien l-III. Gartenwelt 32:

644.

SONDER, O.W. 1862. Mesembiyanthemum. In W.H. Harvey & O.W.

Sonder, Flora capensis 2: 395^60.

Bothalia 43,2: 207-233 (2013)

Notes on African plants

VARIOUS AUTHORS

IRIDACEAE

TAXONOMIC NOTES ON ARISTEA (ARISTEOIDEAE) IN TROPICAL AND

EASTERN SOUTHERN AFRICA

The Afro-Madagascan Aristea Ait., only genus of Iri-

daceae subfam. Aristeoideae Goldblatt, comprises some

58 species of rhizomatous, evergreen perennials. The

genus is currently subdivided into three subgenera: two

of them, subg. Aristea (18 spp.) and subg. Pseudaristea

(Pax) Goldblatt (9 spp.) are restricted to the southern

African winter-rainfall zone in the southwest of the sub-

continent (Goldblatt 2012), whereas subg. Eucapsiilares

(Goldblatt) Goldblatt (31 spp.) occurs widely across

sub-Saharan Africa, where we now recognize 24 species.

There are also eight species in Madagascar, one of them,

A. goetzei Harms, shared with highlands of southern

Tanzania (Goldblatt et al. 2013). Five species are shared

between tropical and southern Africa, five are exclu-

sively tropical African, and 13 species are restricted to

southern Africa, three of them restricted to the southern

African winter rainfall zone (Table 1 ).

The systematics of Aristea is relatively well under-

stood in tropical Africa as a result of accounts of the

genus for Flora Zambesiaca (Goldblatt 1993), Flora of

East Tropical A frica (Goldblatt 1996a), Flora of Ethio-

pia (Goldblatt 1997) and Flore d’Afiique Centrale

(Geerinck 2005). Nevertheless, some taxa require com-

ment and clarification. Aristea djalonis Hutch., from

the West African highlands, has been confused with A.

angolensis Baker (e.g. Hepper 1968), but we consider

it quite distinct from that species and most likely more

closely related to the eastern and southern African A.

torulosa Klatt (better known by its later synonym A.

woodii N.E.Br.).

The taxonomy of a second species, A. abysswica

Pax, requires modification: A. beqiiaertii De Wild, was

treated as a subspecies of A. alata Baker by Weimarck

(1940) in a revision of the genus, but Geerinck (2005)

included it without comment in A. abyssinica. We agree

that A. bequaertii is more closely related to A. abyssini-

ica than to the East African A. alata, but some features

of the taxon suggest it is better treated as a separate spe-

cies. The types of A. bequaertii and of A. homblei De

Wild, share with A. abyssinca and A. alata a stem con-

sisting of a long basal intemode, but whereas the latter

two species have a single subterminal leaf subtending

the single or pair of inflorescences (always binate rhip-

idia in the genus but called flower clusters here), A. beq-

uaertii consistently has two cauline leaves, one inserted

a short distance below the upper. This feature is not seen

in A. abyssinica and in conjunction with some unusual

floral features indicates to us that these plants represent a

separate species.

Lastly, Aristea gerrardii Weim. (Weimarck 1940),

recognized by Goldblatt (1993) in his account of the

genus in Flora Zambesiaca, is, we now conclude, a later

name for A. compressa Baker (1877), and we accord-

ingly reduce it to synonymy under that species. The

two taxa share a multi-branched compound inflores-

TABLE 1 . — Distribution of species of Aristea occurring in tropical Africa with ranges by country

208

Bothalia 43,2 (2013)

cence and compressed, 2-winged stems, and cited differ-

ences between them, especially in spathe and bract size,

seem to us negligible once features of a third species,

A. parviflora Baker, which was mistakenly included in

A. compressa by Weimarck (1940) and Vincent (1985),

are removed from consideration. After examining the

type and other material of A. parviflora. a montane, local

endemic of Eastern Cape, we conclude it is correctly a

separate species and we provide a description of the spe-

cies below, raising it from synonymy in A. compressa.

1. Aristea djalonis: the West African Aristea djalo-

nis, named for the Fouta Djalon highlands in Guinea and

southern Senegal, was first collected by Auguste Cheva-

lier, the French botanist and plant explorer. Chevalier

gave his collection the manuscript name Anthericum

djalonis and published just the name in 1920 in a list of

his plant collections (Chevalier 1920). The species was

treated as Aristea djalonis in a brief (and taxonomically

invalid) entry in Flora of West Tropical Africa (Hutch-

inson & Dalziel 1936), where it was diagnosed in the

key with Hutchinson listed as author. A valid, extended

description was provided later (Hutchinson 1939).

Superficially, collections of the species from Guinea,

Nigeria and Sierra Leone resemble the widespread A.

angolensis, and A. djalonis was included in that species

by Weimarck (1940) and subsequently by Hepper (1968)

in the second edition of Flora of West Tropical Africa.

One of four species of the genus that occur in West

Africa (Table 1 ), Aristea djalonis can readily be dis-

tinguished by the flowering stem, which is simple or

few-branched and compressed, conspicuously so below

the middle of the inflorescence and about 4 mm wide,

and sharply angular to narrowly winged. In A. ango-

lensis, the stem is slender and oval to terete in section.

The inflorescence spathes and bracts also differ, those

of A. djalonis being oblong, 9-11 mm long, and dark

green or brown in the centre, becoming membranous

toward the edges and light, translucent brown. Bracts of

A. angolensis are slightly shorter, dark green to black-

ish (black when dry) in the center and translucent white

on the edges. The bracts of A. djalonis recall those of

the south tropical and eastern southern African A. toru-

losa, and it is to this species that A. djalonis is probably

most closely related. A species of Aristea endemic to

West Africa is surprising, but there are at least five other

endemic Iridaceae there, including two species of Zygo-

tritonia and three of Gladiolus (Goldblatt 1989, 1996b).

Aristea djalonis A.Chev. ex Hutch, in Bulletin of

Miscellaneous Information, Royal Botanical Gardens,

Kew 1939: 246 (1939). [Hutch, in Hutch. & Dalziel,

Flora of West Tropical Africa 2; 376 ( 1936) nom. inval.

Anthericum djalonis A.Chev.: 648 (1920), nom. nud.].

Type: Guinea, Fouta Djalon, Plateau de Dalaba, ''Anthe-

ricum djalonis.' Sept.-Oct. 1907 (fr.), Chevalier 18859

(P, holo.!).

Plants 400-600 mm high. Stem compressed and

2-angled to narrowly winged, ± 4 mm wide in middle,

± 3 mm wide below first flower cluster, often simple

or l-3-branched. Leaves 4-6, narrowly sword-shaped,

mostly 6-10 mm wide, firm, basal and cauline, decreas-

ing in size above. Flower clusters several per axis in lax

spicate arrangement, sessile except terminal; terminal

mostly 4-flowered, laterals mostly 2-flowered; spathes

and bracts 8-10 mm long, dark brown in center, mem-

branous and translucent brown toward edges, becoming

tom and fringed with age. Flowers pale blue (or white),

tepals ± 12 mm long. Filaments ± 6 mm long; anthers ±

2 mm long. Ovajy subglobose, ± 3 mm long; style ± 6

mm long, 3-lobed at apex, lobes fringed. Capsules ses-

sile, ovoid, 8-11 mm long. Seeds unknown. Flowering

time-. December and January.

Distribution-. Guinea, Sierra Leone, Nigeria, in high-

land grassland.

Diagnosis and relationships: Aristea djalonis is dis-

tinguished from the other tropical African species of

the genus by the compressed stem, about 4 mm wide in

the middle, slightly angled to narrowly winged, bearing

sessile lateral flower-clusters on the main axis, which

is usually unbranched and the short, partly transparent,

entire or lightly tom spathes, 9-1 1 mm long. When the

stem is branched, the one to three branches are compara-

tively short and unusually slender. The several leaves are

mostly 6-10 mm wide, whereas leaves of A. angolensis

are usually narrower, often less than 5 mm wide.

Representative specimens

GUINEA. — Fouta Djalon, Diaguissa, 1 300-1 400 m, 18 Apr. 1905

(fr.), Chevalier 12652 (P); environs of Kindia, Benna, 1 000 m, June

1937, Jacqiies-Felix 1756 (P); Mt Loma, env. de Mali, Oct. 1937 (fr.),

Jacques-Felix 1973 (P); Dalaba, roadside near Forestry House, 23 June

\962, Adames 277 (K.).

SIERRA LEONE.— Lome Mtns, 16 Aug. 1945, Jaeger 985 (K,

MO). Loma Mtns, grassland at 5000 ft, 17 June 1966, Morton 3587

(K); Loma Mtns, Bintumane Peak, 6000 ft, 2 May, 1949, Deighion

5099 (K); Loma Mtns, Bintumane, grassland at summit of peak, 14

Aug. 1945 (fr.), Jaeger 985 (MO); Kabala, Loma Mtns, 8 Aug. 1964

Fx.),Adam 26716 (MO).

NIGERIA. — Northern: between Bukuru and Hepham, 4300 ft

[I 310 m], 4 July \92\, Lely 342 (K.).

2. Aristea bequaertir. first collected in 1912, Aris-

tea bequaertii remains a puzzling plant. Restricted to

southern Shaba Province of the Democratic Republic of

Congo (DRC), it is obviously closely allied to A. abyssi-

nica and A. alata, with which it shares a flowering stem

consisting of a long, basal internode, but unlike these

species it bears two (sometimes three) cauline leaves,

the uppemiost of these not inserted at the base of the

terminal flower cluster. Plants bear a terminal flower

cluster distant for the uppermost cauline leaf and some-

times one in each cauline leaf axil, the lower of these

more often stalked and the upper sessile. The pale blue

to almost white flowers have blue anthers and pollen

(verified from photos provided by M. Seleck), unique

in subg. Eucapsulares, in which yellow anthers and pol-

len are the rule. Also unusual are the erect stamens sur-

rounding the style. In other species of the subgenus, the

stamens diverge and the style is eccentric.

Elsewhere in Shaba and further north in the highlands

of eastern DRC, and in Burundi, Rwanda and adjacent

Zambia, typical A. abyssinica seems to be common and

can easily be distinguished from A. bequaertii. The spe-

cies was treated by Weimarck (1940) as A. alata subsp.

bequaertii, with A. abyssinica as a second subspe-

cies of ,4. alata. As currently circumscribed, A. alata is

restricted to Kenya, Uganda and Ethiopia and is as tall

Bothalia 43,2 (2013)

209

as some collections of A. beqiiaertii but it has a single

(rarely two) cauline leaves, always close to the stem

apex, the uppermost subtending the terminal flower clus-

ter. The broader leaves and prominently stalked capsules

on pedicels up to 25 mm render A. alata unmistakable.

Geerinck (2005), in Flore d'Afrique Centrale included

A. beqiiartii in A. abyssinica without comment. Addi-

tional collections of A. bequaertii with well preserved

flowers and with ripe capsules are needed to amplify our

knowledge of the species.

We provisionally include Aristea homblei, described

by De Wildeman (1913) at the same time as A. bequaer-

tii, in synonymy as did Weimarck, but note that the type

consists of rather different plants. Much shorter, the

specimens range in height from as little as 80 mm to 120

mm, bear two cauline leaves and the flattened stems are

somewhat contracted. In some plants of the collection,

the terminal flower cluster is sessile, but in others they

are stalked and distant from the uppermost cauline leaf.

More collections of Aristea in Shaba are needed to better

understand the variation in A. bequaertii and its possible

synonym, A. homblei.

Aristea bequaertii De Wild, in Repertorium spe-

cierum novarum regni vegetabilis 11: 509 (1913). A.

alata Baker subsp. bequaertii (De Wild.) Weim.: 46

(1940). Type: Congo, [Shaba], Lubumbashi [Elisabeth-

ville], 3 Apr. 1912, Bequaert 303 [BR, lecto.l, desig-

nated by Weimarck: 36 (1940)].

Aristea homblei De Wild.: 509 (1913). Type:

Congo, [Shaba], Welgelegen, 1912, Corbusier sub

Flomble 615 (BR. syn.! [3 sheets]).

Plants (80-) 180-400 mm high. Stem compressed,

2-winged, 2-A mm wide, consisting of 1 long basal

intemode and 2 or 3 shorter intemodes in upper 1/3,

bearing 2 or 3 short leaves, these often subtending a

sessile or stalked flower cluster. Leaves several, ± lin-

ear, lower 3 or 4 basal, reaching to ± middle of stem,

2. 0-3. 5 mm wide, cauline leaves 2 or 3, progressively

shorter above. Flower clusters l-2(^), lateral clusters if

present sessile or on short branches, the terminal remote

from uppermost leaf, each 2^-flowered; spathes 7-10

mm long, green with broad membranous margins, keel

sharply angled and hyaline, bracts membranous, becom-

ing ± dry, translucent light brown, slightly shorter than

spathes. Flowers blue to white, ± sessile, tepals sub-

equal, spreading, ovate, 9-12 mm long. Stamens erect

and remaining contiguous, enclosing style; filaments 4-5

mm long, purple; anthers 1.2-1. 5 mm long, pale to deep

blue. Ovar\' ± ellipsoid. ± 5 mm long before pollination,

rapidly enlarging later; style slightly exceeding anthers,

blue-purple, terminating in 3-stalked stigmatic lobes.

Capsules ovoid-ellipsoid, 7-8 mm long. Seeds unknown.

Flowering time: mostly December and January.

Distribution: Congo, Shaba, restricted to southern

Shaba, mostly (always?) in copper enriched soils, in

grassland, woodland and marshy sites. With just a hand-

ful of collections and from a small portion of Shaba, the

conservation status of Aristea bequaertii should perhaps

be regarded as RARE and possibly EN, Endangered,

given its occurrence in copper-rich soils and close to

areas under intensive mining activity.

Representative specimens

CONGO. — Shaba. Kitimbi. in swamp, Jan. 1908, Kas.sner 2326

(B, K); 14 km N of Mitwaba, 16 Jan 1960, Diivigneaud 5075 (BRLU);

Welgelegen, Corbisier 6l5h (BR); Keyberg, open forest, 9 Jan.

1957, Deiilleux 370 (BR); collines de cuivre [copper hills] de Tenke,

Kachimilumbe (2), 10 Jan. 2012, Seleck. Delhaye & Ilunga 210 (BR),

Kachimilumbe (3), 1 1 Jan. 2012, Seleck. Delhaye & Ilunga 210 (BR).

3. Aristea compressa: both A. compressa Baker

(1877) and the newly described A. gerrardii Weimarck

were recognized by Weimarck (1940) in his account

of the genus and both were also recognized by Vincent

(1985) in his treatment of Aristea for eastern south-

ern Africa. Goldblatt (1993) accordingly recognized

A. gerrardii in coastal Mozambique in Flora Zambesi-

aca. Weimarck, and later Vincent, distinguished the two

species by their inflorescence spathes, on the spurious

grounds that those of A. compressa were significantly

smaller. They are in fact virtually the same size in the

type material of both species. Part of the problem was

that several specimens included in A. compressa by both

authors are a second species, A. parviflora Baker (1906),

which has a comparably branched flowering stem to that

of A. compressa but shorter inflorescence spathes, sig-

nificantly smaller flowers and does not have a flattened

flowering stem. Restricted to northeastern Eastern Cape,

A. parx’iflora is a montane species, evidently allied to A.

angolensis.

The types of Aristea compressa and A. gerrardii, both

of which are from coastal KwaZulu-Natal, South Africa,

share a compressed flowering stem, sharply 2-angled or

narrowly 2-winged, flower clusters arranged in a com-

pound pseudopanicle with the central axis dominant,

and lanceolate spathes and floral bracts ± 5-8 mm long,

green or brown in the midline, transparent and becom-

ing tom along the edges. The small, blue flowers differ

hardly at all from those of other tropical African and

several eastern southern African species. We provide

revised descriptions of A. compressa and A. pannflora

below with adjusted nomenclature of ,4. compressa.

Aristea compressa Buching. ex Baker in Journal of

the Linnean Society, Botany 16: 111 (1877) [A. com-

pressa Buching. in Krauss, Flora 28: 309 (1845), nom.

nud.]. Type: South Africa, [KwaZulu-Natal], forest mar-

gins near Umlaas River, 1840, Krauss 358 (K, holo.l,

MO [2 sheets], iso.!).

Aristea gerrardii Weim.: 17 (1940), syn. nov.

Type: South Africa, [KwaZulu-Natal], without precise

locality or date, Gerrard 393 (K, holo.l; P, iso.!).

Plants 300-750 mm long. Stem flattened almost to

apex, narrowly winged, several- to many branched

(one order), but never dichotomously. Leaves narrowly

sword-shaped to linear, leathery, sometimes weakly

pleated, mostly 8-12 mm wide, ± 1/2 as long as stem.

Flower clusters many in compound pseudopaniculate

arrangement, lateral clusters sessile, each 2^-flow-

ered; spathes and bracts lanceolate, mostly 5-8 mm

long, green or brown in midline, transparent along

margins and upper half, becoming tom. Flowers blue,

outer tepals 10-11 x 6-7 mm, inner ±9x4 mm. Sta-

mens: filaments ± 4 mm long, anthers 1. 7-2.0 mm long.

Ovary broadly conical, ± 2 mm long, on short stalk ± 1

mm long; style ± 5 mm long, broadly 3-lobed, reaching

210

Bothalia 43,2 (2013)

mid-anther level. Capsules ovoid, ± 5 mm long, ± ses-

sile. Seeds angular-prismatic, reticulate. Flowering time:

August to January.

Distribution: Aristea compressa extends from Pebane

in coastal northern Mozambique through KwaZulu-

Natal to Port St Johns in Eastern Cape, South Africa,

occurring mainly in coastal sites, often in marshy habi-

tats, but locally inland, as at Drummond (KwaZulu-

Natal) and Mbabane (Swaziland), then often in areas

with sandstone or quartzite substrates (Figure 1). The

species is naturalized in Hawai‘i.

Representative specimens

MOZAMBIQUE. — Maputo: Matutulne, between Zitundo and Man-

hoca, 29 Nov. 1979, De Koning 7706 (BR, MO, NBG). Pebane: Pebane,

Aug. 1950, Munch 261 (SRGH).

SWAZILAND. — 2631 (Mbabane): Mbabane, (-AC), 14 Jan, 1951,

Compton 22366 (NBG), 'near houses, ?garden escape’, 27221 (NBG,

PRE); Mbabane, Poliniane River, (-AC), 6 Nov. 1956, Compton 26198

(PRE).

EASTERN CAPE.— 3129 (Port St Johns): Mkambati Nature

Reserve, hills N of airstrip, (-BD), 9 Dec. 1986, Nicholas & Smook

2335 (MO, PRE); Port St. Johns, (-DA), 10 Dec. 1938, Prance s.n.

(NBG), Nov. 1928, Dyer /577(GRA).

KWAZULU-NATAL.— 2632 (Bella Vista): Kosi Bay, National

Parks Board Camp, (-DD), Moll c6 Strey 3819 (NH, PRE). 2831

(Nkandla): Hlabisa, Palm Ridge Farm, (-BB), Harrison 216 (NH,

PRE). 2832 (Mtubatuba): St. Lucia, eastern Shores State Forest, (BA),

13 Oct. 1983, Nicholas 1602 (MO). 2930 (Pietermaritzburg): Drum-

mond, (-DA), 22 Oct. 1963, Bayliss 1729 (NBG); Clairmont, (-DD),

Oct. 1896, Wood 6139 (PRE); Pinetown, (-DD), Hafstrom & Acocks

267 (PRE), Oct. 1942, Fisher 351 (MO). 3030 (Port Shepstone):

Isipingo Beach, (-BB), Ward 527 (NU); Port Shepstone, (-CB), Nov.

1905, Rogers 494 (GRA). 3130 (Port Edward): Port Edward, (-AA),

Moss 19194 (J).

Aristea parviflora Baker in Bulletin of Miscella-

neous Information, Royal Botanic Gardens, Kew: 25

(1906). Type: South Africa, [Eastern Cape], Insizwe

Mtns, 28 Jan. 1895, Krook in Flerb. Penther 556 (K,

holo.!)

Plants 400-600 mm high. Stem slightly com-

pressed, oval in section, with low raised angles decur-

rent on cauline leaves, repeatedly branched (one or

two orders), main axis remaining dominant, branches

ascending. Leaves ± linear, leathery, basal ± 1/2 as long

as stem, mostly 4-6 mm wide, cauline leaves progres-

sively shorter above. Flower clusters many in paniculate

arrangement, sessile or stalked, each ( 1 )2^-flowered;

spathes and bracts ovate to lanceolate, mostly 3. 5^. 5

mm long, evidently translucent membranous with brown

keels and shading brown near bases, becoming torn with

age. Flowers blue, tepals ± 6 mm long, filaments ± 2.5

mm long; anthers ± 1.5 mm long. Ovaty globose-trun-

cate, ± 2 mm diam., ± sessile; style unknown. Capsules

globose, ± 3.5 mm diam., sessile. Seeds unknown. Flow-

ering time: January and February, probably also March.

Figure 1 .

Distribution: a narrow endemic, Aristea parviflora is

known only from the Insizwe and Mt. Ayliff ranges of

the southern Drakensberg of northeastern Eastern Cape.

One of the collections was made along forest margins

(Figure 1 ).

Diagnosis: described by J.G. Baker in 1906, the dis-

tinctive Aristea parviflora was included by both Wei-

FIGURE 1 . — Distribution ot' Aristea compressa (•) [excluding a record

from Pebane, north of Beira in Mozambique] and A. parviflora

(o).

marck (1940) and Vincent (1985) in A. compressa on

spurious grounds. This narrow endemic has the smallest

flowers in the genus, the tepals about 6 mm long and the

flower thus about 12 mm in diameter, and correspond-

ingly short filaments, about 2.5 mm long. Also distinc-

tive are the globose capsules up to 3.5 mm in diameter

and the stems are only slightly compressed with narrow

raised angles; whereas the stems of A. compressa are

flattened with the longer axis twice as long as the short

axis and the stems are more-or-less winged. The tepals

of A. compressa are 10-12 mm long, filaments 3. 5^.0

mm long and the capsules are ovoid and about 5 mm

long

Selected specimens

EASTERN CAPE. — 3029 (Kokstad): Griqualand East, near

Clyde.sdale, ± 915 m, (-BD), Feb, 1884, Tyson 2872 (BOL, K, PRE,

SAM); forest margins near Mt AylilT, (-BD), Feb. 1884, Tyson 1741

(BOL, SAM); Insizwa, Jan. 1 895, Schlechter 6470 (BOL, GRA).

ACKNOWLEDGEMENTS

We thank Maxime Seleck for sharing his knowledge

and collections of Aristea bequaertii and the curators of

Bothalia43,2 (2013)

211

the following herbaria for loans and access to their col-

lections: BOL, BR, K, NBG. We also thank Clare Archer

for checking holdings at the Pretoria herbarium (PRE)

for us.

REFERENCES

BAICER. J.G. 1 877. Systema Iridearum. Journal of the Linnean Society.

Botany 16: 61—180.

BAKER, J.G. 1906. II. — Diagnoses Africanae, XVI. Bulletin of Mis-

cellaneous Information [Roval Botanic Gardens. Kew] 1906:

15-30.

CHEVALIER, A. 1920, E.xploration botanique de I'Afrique occidentale

fran^aise 1. Enumeration des plantes recoltees. 1. Paris, Leche-

vallier.

DE WILDEMAN, E. 1913. Decades novarum specierum florae katan-

gensis. I -VII. Repertorium specierum novarum regni vegetabi-

lis 11: 501-524.

GEERINCK, D. 2005. Iridaceae. Flore d'Afrique Centrale. Jardin Bota-

nique National de Belgique, Meise.

GOLDBLATT, P. 1989. Revision of the tropical African genus Zygotri-

tonia (Iridaceae). Bulletin du Museum d'Histoire Naturelle, 4 sen.

sect. B. Adansonia 11:1 99-2 1 2.

GOLDBLATT, P. 1993. Iridaceae. Flora Zambesiaca 12, 4. Flora

Zambesiaca Managing Committee, London.

GOLDBLATT, P. 1996a. Iridaceae. In R.M. Polhill (td.). Flora of Tropi-

cal East Afi-ica. A.A. Balkema, Rotterdam.

GOLDBLATT, P, 1996b. Gladiolus in Tropical Africa. Timber Press,

Portland, OR.

GOLDBLATT, P. 1997. Iridaceae. In S. Edwards, S. Demissew & 1.

\\edherg{Qd%). Flora of Ethiopia &. 162-183. University of Addis

Ababa. Addis Ababa.

GOLDBLATT, P. 2012 [2013]. Aristea (Iridaceae: Aristeoideae), a sub-

generic classification. Novon 22,4 (in press).

GOLDBLATT, R, PHILLIPSON, P.B. & MANNING, J.C. 2013. The

new species Aristea farafangana (Iridaceae) from Madagascar,

biogeographic notes on Aristea in Madagascar and a revised

key to the genus. Adansonia [doi: http://dx.doi.org/10.5252/

a2013nla4],ser 3,3: 47-53.

HEPPER, N. 1968. Iridaceae, Pp. 138-142 in Flora of West Tropical

Africa, ed. 2, 3( 1 ). Crown Agents for Oversea Governments and

Administrations, London.

HUTCHINSON, J. 1939. XXVIII — Tropical African plants: XVII. Bul-

letin of Miscellaneous Information. Royal Botanical Gardens.

Xerv. 1939: 241-247.

HUTCHINSON, J. & DALZIEL, J.M. 1936. Iridaceae. P. 374-376 in

Flora of West Tropical Afi-ica 2,2. Crown Agents for the Colo-

nies, London.

KRAUSS, F. 1 845. Pflanzen des Cap- und Natal-Landes, gesammelt un

zusammengestellt von Dr. Ferdinand Krauss. Flora 28: 305-315.

VINCENT, L.P.D. 1985. A partial revision of the genus (Iridace-

ae) in South Africa, Swaziland, Lesotho, Transkei and Ciskei.

South African Journal of Botany 51 : 209-252.

WEIMARCK, H. 1940. Monograph of the genus Aristea. Lunds Uni-

versitets Arsskrift N. F. Avd. 2, 36: 1-140.

P. GOLDBLATT'■^ J.C. MANNING--^

'B.A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.

O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.goldbIatt(§

mobot.org.

-Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. E-mail: j.manning(@

sanbi.org.za.

^Research Centre for Plant Growth and Development, School of Life

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

XOl, Scottsville 3209.

MS. received: 2012-12-10

IRIDACEAE

A NEW SPECIES OF HESPERANTHA FROM THE OVERBERG, WESTERN CAPE, AND OBSERVATIONS ON

A NOVEL MODE FOR POLLEN TRANSFER IN THE GENUS AND FAMILY BY A HESPERID BUTTERFLY

INTRODUCTION

Plants of an unknown species of Hesperantha Ker

Gawk, discovered in the Ruens Hills northeast of

Bredasdorp, Western Cape, South Africa, present some

unusual features in the genus. Elesperantha, a sub-Saha-

ran African genus of some 85 species of Iridaceae sub-

family Crocoideae Burnett, extends from Western Cape

to Ethiopia and Cameroon (Goldblatt 2003; Goldblatt

& Manning 2007; Goldblatt & Manning in prep.). The

genus is recognized by ± woody corm tunics and a spi-

cate inflorescence of radially symmetric flowers with

the style branches dividing close to the mouth of the

perianth tube into long, spreading style branches. The

new speeies, H. kiaratayloriae, has diurnal, small pink,

salver-shaped flowers that are remarkable for the erect

stamens tightly enclosing the exserted style. The style

divides well above the mouth of the perianth tube near

the apices of the anthers, thus unlike any other species

in the genus, and the three style branches, which are

loosely twisted or coiled around one another, are held

well above the mouth of the perianth tube. Although the

exserted style and erect style branches are unique for the

genus, the bell-shaped conns, with an oblique flat base

and floral bracts united basally around the spike, are

consistent with sect. Radiatae Goldblatt. Three days of

observation for pollinators have shown that the flow-

ers, which begin to open at about 6:00 and close after

12:30, are visited and pollinated by the small butterfly

Tsitana tidbagha (Hesperidae), not before recorded as an

agent for pollen transfer in any sub-Saharan species of

Iridaceae (Goldblatt & Manning 2006). We describe the

new species here, discuss its relationships within Hes-

perantha. and present evidence for this novel pollination

system within Iridaceae.

TAXONOMY

Hesperantha kiaratayloriae Goldblatt &

J.C. Manning, sp. nov.

TYPE. — Western Cape, 3420 (Bredasdorp): Ruens

Hills, northeast of Bredasdorp, Farm Plaatjieskraal,

upper slopes of ridge in stony quartzite gravel over

shale, (-AC), 8 Nov. 2011, Goldblatt & Porter 13729

(NBG, holo.; K, MO, PRE, iso.).

Plants mostly 150-200 mm high, rarely to 300 mm,

erect, unbranched. Corm obliquely bell-shaped, 6-8

mm diam. at base, tunics ± woody, dark grey, fragment-

ing from lower margins into tile-like segments. Leaves

several, dry at flowering time, sheaths imbricate, blades

212

Bothalia 43,2 (2013)

linear, mostly 10-60 x 1.0-1. 5 mm, margins smooth,

uppermost 2 or 3 leaves sheathing for entire length.

Spike 2^(-7)-flowered, flexuose; bracts dry and pale

brown at flowering, outer mostly 12-15 mm long with

margins connate near base for 1-2 mm and sheath-

ing spike axis, inner slightly shorter than outer and

forked apically. Flowers diurnal, pink with white throat,

unscented, anthers bright yellow and prominently dis-

played; perianth tube cylindric, slightly expanded at

apex, 13-16 mm long, distal half exserted from bracts;

tepals subequal, spreading horizontally, elliptic, ± 12 x

4-5 mm, outer slightly larger than inner. Stamens sym-

metrically disposed, erect; filaments 2.5 mm long,

exserted ± 1 mm from mouth of tube; anthers ± 5 mm

long, dehiscing longitudinally, yellow, remaining coher-