Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 41,2

Oct. 2011

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE (SANBI) PRETORIA

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

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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 38^11

(2008-2011).

FLORA OF SOUTHERN AFRICA (FS7I)

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. See plan of FSA at back of book.

FSA contributions 1-19 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, 1028010 Poellnitzia. G.F. Smith

1995. Bothalia 25: 35, 36.

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

31-35.

5: Buxaceae. H.F. Glen. 1996. Bothalia 26: 37—40.

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

25^10.

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 27:

149-165.

10: Trapaceae. B. Verdcourt. 1998. Bothalia 28: 1 1-14.

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: C'asuarinaceae. C.M. Wilmot-Dear. 2000. Bothalia 30:

143-146.

18: Salicaceae 5'. str. M. Jordaan. 2005. Bothalia 35: 7-20.

19: Asteraceae: Anthemideae: Enmorphia. N. Swelankomo.

2011. Bothalia 4 1 : 277-282.

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. See list at

back of book.

• 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 1-49 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 Dicroidiurn ), 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.

Anderson & 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. Cleal. 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 inflated by or executed

in partnership with SANBI. See list at back of book.

With compliments from

hSANBIBooksh°P

at the National Herbarium Building. Pretoria

Tel: +27 12 843 5000 E-mail: bookshop@sanbi.org. za

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 41,2

Scientific Editors:

Technical

O.A. Leistner, G. Germishuizen

Editor: B.A. Momberg

SANBI

Biodiversity for Life

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI 01, Pretoria 0001

ISSN 0006 8241

Oct. 2011

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

Anderberg, Dr A. A. Botanical Institute, Stockholm University, Stockholm, Sweden.

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

Archer, Dr R. South African National Biodiversity Institute, Pretoria, RSA.

Boatwright, Dr J.S. South African National Biodiversity Institute, Cape Town, RSA.

Bredenkamp, Prof. G.J. Eco Agent, P.O. Box 25533, Monumentpark, 0105 Pretoria.

Browning, Dr J. Formerly: Natal University Herbarium, Private Bag X01, 3209 Scottsville, RSA.

Bruyns, Dr P. Department Mathematics, University of Cape Town, RSA.

Burgoyne, Ms P. South African National Biodiversity Institute, Pretoria, RSA.

Burrows, J.E. Box 710, 1120 Lydenburg, RSA.

Chinnock, Dr R.J. State Herbarium of South Australia, Adelaide, Australia.

Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.

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

Hoare, D. David Hoare Consulting cc. Private Bag X025, Lynnwood Ridge, Pretoria, RSA.

Jacobsen, DrN. P.O. Box 671, 6560 Wilderness, RSA.

Koekemoer, Ms M. South African National Biodiversity Institute, Pretoria, RSA.

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

Mannheimer, Dr C. P.O. Box 193, Windhoek, Namibia.

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

Meerow, Dr A.W. National Gennplasm Repository, Miami, USA.

Meve, Dr U. University of Bayreuth, Bayreuth, Germany.

Muasya, Dr M. University of Cape Town, 7700 Rondebosch, Cape Town, RSA.

Nordenstam, Prof. R.B. Naturhistoriska Riksmuseet, Stockholm, Sweden.

Oliver, Dr E.G.H. Department of Botany & Zoology, University of Stellenbosch, RSA.

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

Roux, Dr J.P. South African National Biodiversity Institute, Cape Town, RSA.

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

Schrire, Dr B. Royal Botanic Gardens, Kew, UK.

Schutte-Vlok, Dr A.L. Cape Nature, Private Bag X658, 6620 Oudtshoorn, RSA.

Smithies, Mrs S.J. South African National Biodiversity Institute, Pretoria, RSA.

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

Stewart, Dr G. New Zealand Research Centre for Urban Ecology, Christchurch, New Zealand.

Thulin, Dr M.L. Department of Systematic Botany, University of Uppsala, Uppsala, Sweden.

Van Jaarsveld, E.J. South African National Biodiversity Institute, Cape Town, RSA.

Vlok, J. Regalis Environmental Services, P.O. Box 1512, Oudtshoorn, Western Cape, RSA.

Whitehouse, Dr C.M. The Royal Horticultural Society Garden, Wisley, Woking, Surrey, England, UK.

Williams, Dr N. Dept Resource Management & Geography, University of Melbourne, Richmond, Victoria,

Australia.

Winter, P.J.D. South African National Biodiversity Institute, Pretoria, RSA.

Date of publication of Bothalia 41,1 : 3 May 201 1 .

CONTENTS

Bothalia 41 ,2

1. Ornithoglossum pulchrum (Colchicaceae: Colchiceae), a new species from southern Namibia. D.A. SNIJ-

MAN, B. NORDENSTAM and C. MANNHEIMER 231

2. Maireana brevifolia (Chenopodiaceae: Camphorosmeae), a new naturalized alien plant species in South

Africa. L. MUCINA and D.A. SNIJMAN 235

3. Taxonomic revision of the genus Thereianthus (Iridaceae: Crocoideae). J.C. MANNING and P. GOLD-

BLATT 239

4. Romulea pilosa and R. quartzicola (Iridaceae: Crocoideae), two new species from the southern African

winter rainfall region, with nomenclatural corrections including new names for R. amoena , R. neg-

lecta and R. rosea var. reflexa. J.C. MANNING, P. GOLDBLATT and A.D. HARROWER 269

5. FSA contributions 19: Asteraceae: Anthemideae: Eumorphia. N. SWELANKOMO 277

6. Review of the genus Xenoscapa (Iridaceae: Crocoideae), including X grandiflora , a new species from

southern Namibia. J.C. MANNING and P. GOLDBLATT 283

7. Notes on African plants:

Aizoaceae. New combinations in Antimima and Octopoma (Ruschioideae). C. KLAK 292

Alliaceae. Micromorphology and cytology of Prototulbaghia siebertii, with notes on its taxonomic

significance. C.G. VOSA, S.J. SIEBERT and A.E. VAN WYK 311

Amaryllidaceae. Ammocharis deserticola (Amaryllideae), a new species from Namibia and a key

to species of the genus. D.A. SNIJMAN and H. KOLBERG 308

Apocynaceae (Asclepiadoideae-Ceropegieae). First records of Orbea cooperi in Gauteng and

Mpumalanga Provinces, FSA region. S.P. BESTER and S.M. BERRUTI 295

Asteraceae. Lachnospermum neglectum (Asteroideae: Gnaphalieae), a new and overlooked species

from the Worcester Valley, Western Cape. J.C. MANNING and P. GOLDBLATT 304

Asteraceae. Biyomorphe and Dolichothrix (Gnaphalieae-Relhaniinae): taxonomy and nomencla-

ture. M. KOEKEMOER 324

Cyperaceae. New names and new combinations in Cyperus for southern Africa. C. ARCHER and

P. GOETGHEBEUR 300

Fabaceae. A new species of Xiphotheca from the Western Cape, South Africa. A.L. SCHUTTE-

VLOK 298

Fabaceae. A new species of Rhvnchosia from the northern provinces of South Africa. G. GERMIS-

HUIZEN 319

Hyacinthaceae. Albuca gariepensis (Ornithogaloideae), a new species of A. subgen. Namibiogalum

from Gordonia, South Africa, and A. prasina transferred to Ornithogalum. J.C. MANNING

and P. GOLDBLATT 314

Hyacinthaceae. Ornithogalum lebaense transferred to Albuca. J.C. MANNING and P. GOLD-

BLATT 297

Iridaceae. Gladiolus diluvialis (Crocoideae), a replacement name for the illegitimate homonym

G. halophilus. J.C. MANNING and P. GOLDBLATT 324

Lycopodiophyta: Selaginellaceae. Selaginella nivea, a new lycophyte record for South Africa, with

notes on its habitat. A. W. KLOPPER and R.R. KLOPPER 321

Nyctaginaceae. Notes on Commicarpus in southern Africa, including a new record for Namibia.

M. STRUWIG, S.J. SIEBERT and E.S. KLAASSEN 289

Scrophulariaceae. Anticharis juncea, an overlooked record for South Africa, with notes on its type

localities and flower morphology. H.M. STEYN 301

Vitaceae. A new and an overlooked record of Cyphostemma in Angola. F. DE SOUSA, E. FIGUEIRE-

DO and G.F. SMITH 294

8. Phytosociological description of norite koppies in the Rustenburg area, North-West Province and refinement

of the distribution of the Norite Koppies Bushveld on the national vegetation classification map of

South Africa. A.J.H. LAMPRECHT, S.S. CILLIERS, A.R. GOTZE and M.J. DU TOIT 327

9. The extended occurrence of Maputaland Woody Grassland further south in KwaZulu-Natal, South Africa.

S.J. SIEBERT, F. SIEBERT and M.J. DU TOIT 341

10. Floristic analysis of domestic gardens in the Tlokwe City Municipality, South Africa. C.S. LUBBE,

S.J. SIEBERT and S.S. CILLIERS 351

1 1 . Miscellaneous notes:

Tribute to Beverley Momberg, technical editor in the publications section of the South African

National Biodiversity Institute. G.F. SMITH 363

12. Obituary: Franz Sebastian Muller (1913-2010). G.F. SMITH and E. FIGUEIREDO 365

13. South African National Biodiversity Institute: publications 1 January 2010 to 31 December 2010. Com-

piler: Y. STEENKAMP 369

14. Guide for authors to Bothalia 377

New combinations, names, sections, series, species and statuses in Bothalia 41,2 (2011)

Albuca gariepensis J.C. Manning & Goldblatt sp. nov., 315

Albuca lebaensis (Van Jaarsv.) J.C. Manning & Goldblatt , comb, nov., 297

Ammocharis deserticola Snijman & Kolberg, sp. nov., 308

Antimima paripetala (L. Bolus) Klak , comb. nov., 293

Bracteolatus J.C. Manning & Goldblatt , ser. nov., 254

Brevibracteae Goldblatt & J.C. Manning, sect, nov., 246

Cyperus atriceps (Kiik.) C. Archer & Goetgh ., comb. & stat. nov., 300

Cyperus austro-afrieanus C. Archer & Goetgh., nom. nov., 300

Cyperus decurvatus (C.B. Clarke) C. Archer & Goetgh., comb. & stat. nov., 300

Cyperus palmatus (Lye) C. Archer & Goetgh., comb. & stat. nov., 300

Cyperus uitenhagensis (Steud.) C. Archer & Goetgh., comb, nov., 301

Gladiolus diluvialis Goldblatt & J.C. Manning, nom. nov., 324

Lachnospermum neglectum Schltr. ex J.C. Manning & Goldblatt, sp. nov., 304

Octopoma nanum (L. Bolus) Klak, comb, nov., 293

Ornithoglossum pulchrum Snijman, B.Nord. & Mannheimer, sp. nov., 231

Rhynchosia coddii Germish., sp. nov., 319

Romulea pilosa J.C. Manning & Goldblatt, sp. nov., 269

Romulea quartzicola J.C. Manning & Goldblatt, sp. nov., 272

Romulea rosea var. muirii (N.E.Br.) J.C. Manning & Goldblatt , comb, et stat. nov., 275

Thereianthus bulbiferus Goldblatt & J.C. Manning, sp. nov., 254

Thereianthus elandsmontanus Goldblatt & J.C. Manning, sp. nov., 263

Thereianthus intermedius J.C. Manning & Goldblatt, nom. et stat. nov., 247

Xenoscapa grandiflora Goldblatt & J.C. Manning, sp. nov., 284

Xiphotheca rosmarinifolia A.L.Schutte, sp. nov., 298

IV

Bothalia 41,2: 231-233 (2011)

Ornithoglossum pulchrum (Colchicaceae: Colchiceae), a new species

from southern Namibia

D.A. SNIJMAN*, B. NORDENSTAM** and C. MANNHEIMER***

Keywords: Colchicaceae, Colchiceae, new species, Ornithoglossum Salisb., southern Namibia, taxonomy

ABSTRACT

We describe a new species in the sub-Saharan genus Ornithoglossum Salisb. from southern Namibia. Ornithoglossum

pulchrum from near Aus, is remarkable in having bright to dark pink flowers, a feature previously unknown in the genus.

The perigone is almost concolorous apart from a contrasting, pale yellow nectary region, narrowly outlined with darker red,

near the base of each tepal. The undulate leaves together with the long filaments, which are nearly as long as the tepals,

suggest a relationship with O. undulatum, a widespread species in the western parts of southern Africa, and O. zeyheri from

Namaqualand and the northwestern Cape.

INTRODUCTION

Ornithoglossum Salisb. is a small, sub-Saharan genus

in the family Colchicaceae (Nordenstam 1998). All

but one of the eight species recognized by Nordenstam

(1982) are concentrated in the western half of southern

Africa, mostly in the winter rainfall region. Only O. vul-

gare B.Nord. is found in southern Africa’s eastern parts

and as far north as East Africa. Previously the genus was

placed in the tribe Iphigenieae (Buxbaum 1936), but in

a new classification, based on an analysis of molecular

data of the family, the genus now falls within the tribe

Colchiceae, together with Colchicum L., Gloriosa L.,

Hexacyrtis Dinter and Sandersonia Hook. (Vinnersten &

Manning 2007).

Considering its small size, the genus shows remark-

able diversity in floral morphology. The perianth is

actinomorphic or zygomorphic in form and coloured

cream to attractive yellow, green, brown or purple,

sometimes almost black, and often bicoloured. In spe-

cies such as Ornithoglossum undulatum Sweet, the

flowers are large, showy and sweetly scented, but in O.

parviflorum B.Nord. they are small, inconspicuous, dull

and unscented (Manning et al. 2002). Other variable fea-

tures are the shape of the nectaries on the basal part of

the tepals and the length and thickness of the filaments.

Several micromorphological differences in the pollen

and seeds also help to distinguish groups of species. All

the species are reported to be highly toxic to stock due to

their colchicine-type alkaloids, which accounts for their

common name ‘ slangkop ’ (Watt & Breyer-Brandwijk

1962).

While collecting material for a photographic guide to

the wildflowers of the southern Namib, the third author

found an unusual population of Ornithoglossum near the

small town of Aus, Namibia, in which the flowers were

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Cape Town. E-mail: d.snijman@sanbi.org.za.

** Swedish Museum of Natural History, Box 50007, SE- 10405 Stock-

holm, Sweden.

*** P.O. Box 193, Windhoek, Namibia. E-mail: manfam@iafrica.com.

na.

MS. received: 2011-03-15.

deep pink to crimson and almost concolorous except for

a pale yellow patch towards the base of each tepal. Due

to their undulate leaves, the plants were initially thought

to be O. undulatum (Mannheimer et al. 2008), a spe-

cies that has been recorded previously from southern

Namibia (Nordenstam 1982). Subsequent examination

of these specimens, however, has revealed further dis-

tinguishing characters, which together with the unique

flower colour, justify the description of the plants as a

new species of Ornithoglossum.

Ornithoglossum pulchrum Snijman, B.Nord. &

Mannheimer , sp. nov.

Cormus ovoideus, brevilobus. Caulis simplex gracilis

ad ± 110 mm altus. Folia 4—7, lanceolata margine con-

spicue crispato-undulato. Racemus 3-10-florus ad 40-80

mm longus. Perianthium actinomorphum, segmen-

tis 27-32 x 4-5 mm, roseis vel rubineis, luteolis prope

basim. Nectarii margo simplex. Filamenta plerumque

longitudine tepala aequantia. Antherae 2.5 x 0.5 mm.

Styli ± 25 mm longi.

TYPE. — Namibia: 2616 (Aus): riverbed, western

extent of Aus town lands, 1 386 m, (-CA), 22 Jun. 2005,

C.A. Mannheimer CM2710 (WIND, holo.).

Glabrous, cormous herb. Corm ovoid, ± 45 x 30 mm,

shallowly bilobed at base with lobes directed down-

wards, interior firm, whitish; tunics leathery, dark brown,

extended into a slender neck up to 80 mm long, sheath-

ing subterranean portion of stem, inner tunics rust-col-

oured. Cataphyll apparently absent. Stem erect, simple,

subterranean part slender, up to ± 110 mm long, as long

as leafy aerial portion, terminated by a bracteate, race-

mose inflorescence. Leaves alternate, 4-7, lanceolate

from sheathing base, recurved, 10-180 x 8-15 mm.

broadest at base, tapering distally to a narrow, mucro-

nate tip, ± conduplicate, glaucous, margins undulate,

often crisped. Inflorescence a compact, 3-10-flowered

raceme, 40-80 mm long and clustered among upper

leaves; bracts leaf-like, narrowly lanceolate, succes-

sively smaller acropetally, 30-50 x 4-6 mm, as long as

or slightly longer than pedicels, glaucous tinged pink-

ish, margins plane. Flowers suberect to spreading on

232

Bothalia 41,2 (2011)

FIGURE 1. — Ornithoglossum pitl-

chrum. A, plant in habitat

showing sheathing, undulate

leaves and compact inflores-

cence; B, detail of campanu-

late flower showing long sta-

mens and six large, concave

nectaries each with a simple

margin near base of tepals.

Photographs: C. Mannheimer.

erecto-patent pedicels 20-35 mm long, actinomorphic,

± widely campanulate, ± 40 mm diam.; tepals equally

spreading, 27-32 x 4-5 mm, claw tubular-flattened,

± 4.0 x 1 .5 mm, blade lanceolate, faintly 7-9-veined,

slightly canaliculate, bright to dark pink, with a pale

yellow nectary region narrowly outlined with darker

red, becoming paler with age; nectary concave, wide-

mouthed, ventral margin simple. Stamens slightly

spreading; filaments straight, slightly curved distally, ±

filiform but slightly thickened in proximal half, 25-28

mm long, uniformly dark pink; anthers oblong, 2.5 x 0.5

mm, slightly curved, dull yellow. Ovary oblong-globose,

5.0 x 3.5 mm, dark pink; styles free from base, spread-

ing, straight proximally, slightly curved distally, ± 25

mm long, dark pink; stigma capitate, minutely papillate.

Capsule elliptical-oblong, shortly and bluntly lobed, 7

x 4 mm (when immature but not known when mature),

erect, coriaceous. Seeds unknown. Flowering time : June

to early Aug. Figures 1 ; 2.

Distribution and ecology. Ornithoglossum pulchrum is

currently known from just one locality in the pro-Namib,

a broad, undulating plain in southern Namibia (Figure 3).

The species has been recorded in ephemeral watercourses

from the uplands near Aus, which he just below the inland

escarpment, at approximately 1 380 m. The plants grow

in coarse gravel, close to gneiss outcrops of the Namaqua

Metamorphic Complex. Lying on the border of the win-

ter and summer rainfall zones, Aus has bimodal rainfall,

averaging 85 mm per year. Most rain falls in late summer

(January to April) with a second, lower peak in June (Pal-

lett 1995). Precipitation also occurs in the form of occa-

sional fog which moves in from the coast, as well as rare

snowfalls. Winds in the region are a powerful climatic

force which can severely limit plant growth. Like most

other Ornithoglossum species from southern Africa’s win-

ter rainfall region, O. pulchrum flowers in winter, usually

from mid to late June into August.

FIGURE 2. — Ornithoglossum pul-

chrum. A, type specimen, Mann-

heimer CM2710 (WIND); B,

specimen on left shows shal-

lowly bilobed corm, specimen

on right shows undulate, crisp-

u late-edged leaves and young,

upright capsules, Mannheimer

CM4004a (WIND). Scale bars:

A, B, 100 mm.

Bothalia 41,2 (2011)

233

FIGURE 3. — Known distribution of Ornithoglossum pulchrum.

Diagnosis and relationships : Ornithoglossum pul-

chrwn is distinguished by its flowers, which are suberect

to spreading in a compact raceme that barely exceeds the

leaves, and in which the ± filiform filaments are almost

as long as the tepals. The flowers are easily distinguish-

able from those of other Ornithoglossum species by their

bright to dark pink colour (47C, 48C in R.H.S Colour

Chart 1966). This colouring is almost unbroken apart

from a pale yellow patch, narrowly outlined with dark

red, near the base of each tepal in the region of the nec-

tary. O. pulchrum shares the character of undulate, crisp-

ulate-edged leaves with four other taxa in the genus, viz.

O. undulatum, O. gracile B.Nord. and O. zeyheri B.Nord..

and also O. parviflorum var. namaquense B.Nord. Com-

pared with the floral characters, however, this veg-

etative feature is of less taxonomic importance, as quite

plane-edged leaves are known in some populations of

O. undulatum (Nordenstam 1979, 1982). Nevertheless,

the undulate leaves together with the long filaments,

which nearly equal the length of the tepals, suggest

a relationship with both O. undulatum , a widespread

species in the western parts of southern Africa, and O.

zeyheri which is confined to Namaqualand between the

Steinkopf region and the lower Olifants River Valley.

O. undulatum differs from O. pulchrum in its markedly

asymmetric flowers in which one or two tepals point

downwards and the other tepals flare upwards at anthe-

sis. These are slightly smaller than those of O. pulchrum

(16-30 x 2-5 mm vs 27-32 x 4-5 mm) and are bicol-

oured, white in the centre with reddish purple tips. In

contrast, the flowers of O. pulchrum are actinomorphic

and campanulate, features that are shared with O. zey-

heri. Unlike O. pulchrum, however, this species has few

and inconspicuous flowers which are typically produced

in May, with tepals that are short and narrow (12-15

x 1-3 mm) and coloured pale greenish with a purplish

tinge towards the base and tip. The regular symmetry of

the flowers of O. pulchrum prevent possible confusion

with any other species from southern Namibia even in

the early fruiting stages, since its withered tepals remain

evenly spread around the developing suberect to spread-

ing capsule, unlike those of O. undulatum which are dis-

tinctly reflexed from a markedly down-turned capsule.

Both O. zeyheri and O. undulatum share a simple-mar-

gined nectary with O. pulchrum , although this feature is

variable in O. undulatum, sometimes taking the form of

an entire or bifid lobe.

Ornithoglossum pulchrum has only been collected

twice so far, once when in flower and again in the early

stages of fruiting. As yet, mature fruits and seeds are not

available for comparison with other species.

Species from other families that are narrowly endemic

to the Aus area are Moraea graniticola Goldblatt (Iri-

daceae) and Oxalis ausensis R.Knuth. (Oxalidaceae),

both geophytic herbs, and the succulent shrub Juttadin-

teria ausensis (L. Bolus) Schwantes (Aizoaceae). M. gra-

niticola and J. ausensis flower only after winter rain and

when temperatures begin to rise. O. ausensis is one of

four autumn-flowering Oxalis species in the vicinity of

Aus.

Other specimen seen

NAMIBIA. — 2616 (Aus): Aus town lands, (-CA), 11 Sept. 2008,

C.A.Mannheimer CM4004a (WIND).

ACKNOWLEDGMENTS

We are grateful to Christine Swiegers and Telane

Greyling who assisted in the field and Michelle Smith

who prepared the figures. We thank both the Namibian

Ministry of Environment and Tourism for granting a col-

lecting permit to the National Botanical Research Insti-

tute, Windhoek, and the Curator of the National Herbar-

ium. Windhoek for the loan of material to the Compton

Herbarium, Claremont, Cape Town.

REFERENCES

BUXBAUM, F. 1936. Die Entwicklungslinien der Liliodeae. I. Die

Wurmbaeoideae. Botanisches Archiv 38: 213-293.

MANNHEIMER, C„ MAGGS-KOLLING, G., KOLBERG, H. &

RUGHEIMER, S. 2008. Wildflowers of the southern Namib.

Macmillan, Namibia.

MANNING, J., GOLDBLATT, P. & SNIJMAN, D. 2002. The color

encyclopedia of Cape bulbs. Timber Press, Oregon and Cam-

bridge.

NORDENSTAM, B. 1979. Ornithoglossum undulatum. The Flowering

Plants of Africa 45: t. 1799.

NORDENSTAM, B. 1982. A monograph of the genus Ornithoglossum

(Liliaceae). Opera Botanica 64: 1-51.

NORDENSTAM, B. 1998. Colchicaceae. In K. Kubitzki, The families

and genera of vascular plants, vol. 3. Flowering plants, mono-

cotyledons, Lilianae (except Orchidaceae). Springer- Verlag,

Berlin.

PALLETT, J. 1995. The Sperrgebiet Namibia’s least known wilderness.

An environmental profile of the Sperrgebiet or Diamond Area

1 in south-western Namibia. NAMDEB Diamond Corporation,

Windhoek.

R.H.S. COLOUR CFtART. 1966. Tire Royal Horticultural Society, Lon-

don.

VINNERSTEN, A. & MANNING, J. 2007. A new classification of Col-

chicaceae. Taxon 56: 171-178.

WATT, J.M. & BREYER-BRANDWIJK, M.G. 1962. The medicinal

and poisonous plants of southern and eastern Africa. Livingston,

Edinburgh.

Digitized by the Internet Archive

in 2016

j

https://archive.org/details/bothaliavolume4141unse_1

Bothalia 41,2: 235—238 (2011)

Maireana brevifolia (Chenopodiaceae: Camphorosmeae), a new natu-

ralized alien plant species in South Africa

L. MUCINA* ** and D.A. SNIJMAN*

Keywords: Australia, Chenopodiaceae, distribution, Maireana brevifolia (R.Br.) Paul G.Wilson, new naturalized species, semi-arid South Africa

ABSTRACT

We describe and discuss the distribution of a new, naturalized alien species, Maireana brevifolia (R.Br.) Paul G.Wilson

(Chenopodiaceae), a native of Australia, in the western regions of South Africa. First discovered near Worcester, Western

Cape in 1976, the species is now established in disturbed karoo shrubby rangelands, along dirt roads and on saline alluvia,

from northern Namaqualand to the western Little Karoo. In the South African flora, M. brevifolia is most easily confused

with the indigenous Bassia salsoloides (Fenzl) A.J. Scott, from which it is distinguished by the flat to cup-shaped and almost

glabrous perianth with woolly-ciliate lobes, and the hardened and winged fruiting perianth.

INTRODUCTION

Several ecological groups of Australian alien plants

have been incorporated into the South African flora,

including a number of Fabaceae, predominantly Acacia

species (most importantly A. cyclops A.Cunn. ex G.Don.

A. longifolia (Andrews) Willd., A. mearnsii De Wild.,

A. melanoxylon R.Br., A. pycnantha Benth., A. saligna

(Labill.) H.L.Wendl.), and several species of Chenopo-

diaceae. These vary from herbaceous weeds such as

Dysphania cristala (F.Muell.) Mosyakin & Clemants (=

Chenopodium cristatum F.Muell.), D. pumilio (R.Br.)

Mosyakin & Clemants (= Chenopodium pumilio R.Br.)

to weeds of karoo shrublands and disturbed sites in the

arid biomes of southern Africa, including several notor-

ious species of Atriplex, namely A. eardleyae Aellen, A.

lindleyi Moq., A. nummularia Lindl. and A. semibaccata

R.Br. (Goldblatt & Manning 2000; Makgakga 2003).

Atriplex nummularia in particular has been cultivated

widely, especially on heavy alluvial soils in South Africa

as a dry-season fodder plant (L. Mucina pers. obs.). In

many places (especially in the Little Karoo), the species

has escaped cultivation to become an established weed,

especially on alluvia of intermittent semi-desert rivers.

During the preparation of an account for the Chenopo-

diaceae in Plants of the Greater Floristic Region 2: the

Extra Cape flora (Mucina & Snijman ined.), it became

apparent that yet another adventive species of Chenopo-

diaceae, native to Australia, is present in several places in

the western regions of South Africa.

Maireana brevifolia

Maireana brevifolia (R.Br.) Paul G.Wilson is the

newest addition in Chenopodiaceae to the alien flora

of the South African semi-deserts (Figure IB). It has

been found to become established in disturbed karoo

shrubby rangelands, along dirt roads, and on saline allu-

* Curtin Institute for Biodiversity & Climate, Department of Environ-

ment & Agriculture, Curtin University, GPO Box U1987, Perth, WA

6845, Australia. E-mail: L.Mucina@curtin.edu.au.

** Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, Claremont 7735, Cape Town, South Africa. E-mail:

d.snijman@sanbi.org.za.

MS. received: 2011-03-15.

via. The latter habitat is that preferred by M. brevifolia

in its homeland — Australia. In its native range, it cov-

ers a large distribution area (Figure 1A) characterized

by Transitional’ rainfall (where the influence of western

frontal systems meets that of poorly predictable cyclonic

summer rainfall). The footprint of the distribution area

of this species marks the extent of unique temperate

Eucalyptus open woodlands that have been cleared,

especially in many parts of Western Australia, South

Australia, New South Wales, southern Queensland and

northern Victoria. The native status of this species in the

Northern Territory needs revisiting. This species has a

moderate salinity and waterlogging tolerance and prefers

heavy clay or sandy-clay alluvial soils. In Western Aus-

tralia it has been found to be one of the pioneers of re-

colonization of abandoned salinized pastures.

Maireana brevifolia is also an established weed

in Chile (Marticorena 1997), the Middle East (Danin

2000b) and the Canary Islands (Brandes 2006). Climate-

niche modelling (Figure 1C) predicts that M. brevifo-

lia could establish with high probability throughout

the Mediterranean, around the Gulf of Mexico, eastern

Argentina and neighbouring countries, as well as south-

ern China.

Maireana brevifolia might have been introduced to

South Africa when targeted as potential dry-season fod-

der or accidentally as seed admixtures with Atriplex

nummularia. The first herbarium record for the species

in South Africa, made in 1976, is from the Worcester

area, Western Cape. With her knowledge of the West-

ern Australian flora, the late Pauline Bond, formerly of

the Compton Herbarium, recognised the collection as an

alien species and was the first to identify the plants as

M. brevifolia. Since then the species has been collected

four more times, once in Namaqualand and on three sep-

arate occasions in the Western Cape, dating from the late

1990s onwards. The identity of two of these collections

has been independently confirmed by R.J. Chinnock of

the State Herbarium of South Australia, Adelaide, and H.

Freitag, Kassel, Germany.

In Australia, Maireana brevifolia has been used to

ameliorate salt-laden pastures (Runciman & Malcolm

1991; Barrett-Lennard & Malcolm 1995; Barrett-Len-

236

Bothalia 41,2 (201 1 )

FIGURE 1. — A, distribution map of Maireana brevifolia in Australia (courtesy of Australian Virtual Herbarium, http://avh.rbg.vic.gov.au/avh/);

B, distribution map of M. brevifolia in South Africa (original); C, modelled distribution of M. brevifolia using Australian localities (stored

by www.gbif.org) as basis of model. Openmodeller generated the probability distribution using Envelope Score Algorithm (see http://open-

modeller.sourceforge.net for computation details). Colour scheme (spanning hot/red colours to cool/blue colours) used in map designates a

gradient of probability of high vs low probability of occurrence of M. brevifolia under the given climate-niche scenario.

nard et al. 2003) and with less success, also mine spoil

dumps (Jefferson 2004; Jefferson & Pennacchio 2005).

Outside Australia, M. brevifolia has been introduced for

saline soil improvement and dry-season fodder produc-

tion in Pakistan (Ilyas et al. 2000), Iraq (Abdul-Halim et

al. 1990) and Israel (Danin 2000a).

Maireana brevifolia (R.Br.) Paul G. Wilson in

Nuytsia 2: 22 (1975). Type: Australia, South Australia,

Bay XI, south coast, R. Brown (BM, holo.).

Kochia brevifolia R.Br.: 409 (1810).

Succulent-leaved shrub, up to I m high, with mostly

erect, slender, striate, sparsely woolly branches. Leaves

alternate, obovoid to narrowly fusiform, ( 1— )2 — 5 mm

long, fleshy, narrowed into a short petiole, glabrous.

Flowers bisexual, solitary, ebracteolate, glabrous except

for woolly-ciliate lobes. Fruiting perianth glabrous;

tube shallowly hemispherical, thin-walled, ± 2 mm in

diam.; wings 5, rarely absent, equal, horizontal, thin,

fan-shaped, 2-3 mm long, glabrous, with delicate brown

venation when dry, with pale to dark pink tint; perianth

lobes thick and fleshy, sharply demarcated from wings.

(Description based on Wilson 1984). Flowering time :

(Aug.-)Nov. to Apr. Figure 2.

Common names in Australia'. Small-leaf Bluebush,

Eastern Cotton Bush, Yanga Bush. None is yet known in

South Africa.

Distribution in South Africa', to date Maireana brevi-

folia has been recorded from Namaqualand, between

Anenous and Komaggas, on the lower slopes of the

Bothalia 41 ,2 (2011)

237

FIGURE 2. — Maireana brevifolia. habitat and habit: A, typical habitat of M. brevifolia — disturbed edge of dirt road crossing a shallow depression

(alluvium of intermittent river) near Komaggas (Namaqualand, South Africa); B, extensively used pasture on saline alluvium near Kondinin

(Western Australia) with a bush of M. brevifolia in foreground and scattered individuals in the rear. C, D, Mucina 280407/03 MJG, NBG', from

Komaggas, South Africa: C, form with deep pink fruit; D. erect-branched habit of M. brevifolia bushes. E, Mucina 102010/03 Curtin Univ.

Herbarium from Corrigin. Western Australia: form with pale pink fruit. All photographs: L. Mucina.

eastern Cederberg Mtns bordering on the Tanqua Karoo,

the Worcester-Robertson Karoo and the western Lit-

tle Karoo. As long as disturbed habitats such as those

formed along dirt roads and on saline alluvia are avail-

able for this colonizer, M. brevifolia will persist as an

established alien in the South African flora.

Notes : the genus Maireana is native to mainland

Australia and has ± 57 species. M. brevifolia is the only

species of the genus known to have been introduced

into South Africa so far. Within the South African flora,

Maireana is most closely related to Bassia All., which

is the only native genus belonging to tribe Camphoros-

meae ( sensu Akhani et al. 2007). Maireana is distin-

guished from Bassia by flowers that are usually solitary

or in pairs in the leaf axils, the flat to cup-shaped and

mostly glabrous perianth (except in M. brevifolia where

the lobes are woolly-ciliate), and the hardened and

winged fruiting perianth. In contrast, Bassia has solitary

flowers or flowers arranged in spicate inflorescences, an

urn-shaped perianth, and the fruiting perianth has spine-

like or tubercle-like appendages or membranous, scari-

ous wings. M. brevifolia is most easily confused with

B. salsoloides (Fenzl) A.J. Scott, commonly known as

the basterinkbos, which is widespread in sandy, alluvial

soils along water courses and seasonal washes from the

Free State in the northeast, through the central Karoo to

Matjiesfontein, Western Province, in the southwest. B.

salsoloides has linear to filiform, succulent leaves that

are ± pubescent, flowers borne in tight, short, spike-like

clusters, a pubescent perianth, and a fruiting perianth

that enlarges into short, firm, brown-streaked wings,

with irregularly toothed margins.

Specimens examined

NORTHERN CAPE. — 2917 (Springbok): Namaqualand, north-

west of Komaggas, sandy-clayey saline donga, 29°17'51.76"S,

17°28T0.02"E, (-CD), 28 Apr. 2007, L. Mucina 280407/05 (MJG).

WESTERN CAPE.— 3219 (Wuppertal): Matjies River Nature

Reserve, Upland Succulent Karoo, stony soil, 760 m, 32°29.90'S,

19°20.30'E, (-AD), 21 Nov. 1999, A.B. Low 5835 (NBG); Matjies

River Nature Reserve, roadsides and valley bottom on old fields, on

sandy or loamy soils, 32°30'03"S, 19°20'18"E, (-CB), Apr. 2002, PM.

Holmes s.n. (NBG). 3319 (Worcester): Karoo Garden, Worcester, in

Euphorbia mauritanica patches, 300 m, (-CB), Mar. 1976, M.B. Bayer

149 (NBG). 3321 (Ladismith): 0.4 km east of T-junction of R327 with

connecting road to R323 just west of Langberg [Langeberg], abundant

along roadsides in Acacia karroo / Aizoaceae low shrubland, on stony

brown loam, ± 33°49’30"S, 21°28'07"E, (-CD), 21 Aug. 1998, R.J.

Chinnock 9194 (NBG).

ACKNOWLEDGEMENTS

We thank H. Freitag for the identification of a col-

lection of Maireana brevifolia from South Africa. L.

238

Bothalia 41,2 (2011)

Mucina thanks the Northern Cape Nature Conservation

Service, Kimberley for awarding a plant collecting per-

mit. G.J. Campbell-Young kindly proofread the manu-

script. M. Smith prepared the distribution map for South

Africa. We also thank two anonymous referees for their

invaluable input.

REFERENCES

ABDUL-HAL1M, R.K.., AL-BADRI, F.R., YASIN, S.H. & ALI, A.T.

1990. Survival and productivity of chenopods in salt-affected

lands of Iraq. Agriculture, Ecosystems and Environment 3 1 :

77-84.

AKHANI, H„ EDWARDS, G. & ROALSON, E.H. 2007. Diversifica-

tion of the Old World Salsoleaes./. (Chenopodiaceae): molecular

phylogenetic analysis of nuclear and chloroplast data sets and

a revised classification. International Journal of Plant Science

168: 931-956.

BARRETT-LENNARD, E.G. & MALCOLM, C.V. 1995. Saltland pas-

tures in Australia. West Australian Dept, of Agriculture Bulletin

No. 4312. Dept of Agriculture, Perth.

BARRETT-LENNARD, E.G., MALCOLM, C.V. & BATHGATE, A.

2003. Saltland pastures in Australia — a practical guide, edn 2.

CRC for Plant-based Management of Dryland Salinity, Perth.

BRANDES, D. 2006. Urbanizaciones: Die Entstehung stadtischer

Lebensraume aus der Halbwiiste. Geobotanische Kolloquien 20:

13-20.

BROWN, R. 1810. Prodromus florae Novae Hollandiae. Johnson, Lon-

don.

DANIN, A. 2000a. The inclusion of adventive plants in the second edi-

tion of Flora Palaestina. Willdenowia 30: 305-314.

DANIN, A. 2000b. Species ‘new to’ the Flora Palaestina area. Flora

Mediterranea 10: 109-172.

GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanical

Garden, Pretoria & Missouri Botanical Garden, St Louis.

ILYAS, M„ HASSAN, M„ MUHAMMAD, W. & MUHAMMAD, N.

2000. Study of yield and yield components of exotic Atriplex and

Maireana in the saline environments ofNWFP. Pakistan Journal

of Biological Sciences 3: 1 873-1 876.

JEFFERSON, L.V. 2004. Implications of plant density on the resulting

community structure of mine site land. Restoration Ecology 12:

429-J38.

JEFFERSON, L.V. & PENNACCHIO, M. 2005. The impact of shade on

establishment of shrubs adapted to the high light irradiation of semi-

arid environments. Journal of Arid Environments 63: 706-716.

MAK.GAK.GA, M.C. 2003. Chenopodiaceae. In G. Germishuizen &

N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14. National Botanical Institute, Pretoria.

MARTICORENA, C. 1997. La presencia de Maireana brevifolia

(R.Br.) Paul G. Wilson en Chile continental. Revista Gayana

Botanica 54: 193, 194.

MITCHELL, T.L. 1848. Journal of an expedition into the interior of

tropical Australia. Longman, London.

MUCINA, L. & SNIJMAN, D.A. ined. Amaranthaceae. In D.A. Snij-

man. Plants of the Greater Cape Floristic Region 2: the Extra

Cape flora. Strelitzia. South African National Biodiversity Insti-

tute, Cape Town.

RUNCIMAN, H.V. & MALCOLM, C.V. 1991. Forage shrubs and

grasses for revegetating saltland. West Australian Dept of Agri-

culture Bulletin No. 4153.

WILSON, P.G. 1975. A taxonomic revision of the genus Maireana

(Chenopodiaceae). Nuytsia 2: 2-82.

WILSON, P.G. 1984. Chenopodiaceae. In A.S. George, Flora of Australia

4: 81-330. Australian Government Publishing Service, Canberra.

Bothalia 41,2: 239-267 (2011)

Taxonomic revision of the genus Thereianthus (Iridaceae: Crocoideae)

J.C. MANNING* and P. GOLDBLATT**

Keywords: Crocoideae, Iridaceae, new species, pollen, southern Africa, taxonomy, Thereianthus G.J. Lewis

ABSTRACT

Thereianthus G.J. Lewis is revised, with full descriptions and synonomy, distribution maps, and notes on ecology and tax-

onomic history. All species are illustrated, and capsule and seed morphology are described for many of the species for the first

time. Novel characteristics of the bract, seed, and pollen operculum are used to separate the species into two sections: sect.

Brevibracteae Goidblatt & J.C. Manning is distinguished by relatively small bracts, 3-8 mm long and uniformly leathery or

soft-textured without thickened veins, seeds with filiform chalazal extension, and pollen grains with 1 -banded operculum; and

sect. Thereianthus by relatively larger bracts, (7— )8— 1 5 mm long with prominently sclerified veins, seeds without any exten-

sion to the chalazal crest, and pollen grains with ± 2-banded operculum. Species in sect. Thereianthus are further segregated

into ser. Thereianthus , with heavily ribbed leaves and suberect flowers with arcuate or erect stamens, and ser. Bracteolatus,

with plane, inconspicuously veined leaves and ± spreading flowers with declinate stamens. Eleven species are recognized in

the genus, all restricted to the southwestern portion of Western Cape. Two new species are described in sect. Thereianthus :

T. bulbiferus Goidblatt & J.C. Manning, known from three populations along the West Coast, is distinguished by the unique

development of cormels in the lower leaf axil, and by its actinomorphic perianth with white marks at the base of each tepal

and ± declinate stamens; and T. elandsmontanus Goidblatt & J.C. Manning, known from a single population in Elandsberg

Nature Reserve near Wellington, has distinctive cream-coloured, moderately long-tubed flowers with unusually narrow, linear

tepals heavily marked with purple near the base. In adddition, T. lapeyrousioides [now T. minutus] var. elatior G.J. Lewis in

sect. Brevibracteae is raised to species status as T. intermedius Goidblatt & J.C. Manning, differing from typical T. minutus

by the shorter perianth tube (10-13 vs 20-30 mm), shorter bracts (3-5 vs 6-8 mm), and smaller, ovoid capsules, 4-5 mm long.

containing ovoid seeds vs flask-shaped capsules 6-8 mm long

INTRODUCTION

Thereianthus G.J.Lewis is a small genus of deciduous

geophytes endemic to the southwestern, winter rainfall

region of Western Cape, where it is essentially restricted

to mainly montane soils derived from quartzitic sand-

stone (Manning et al. 2002). Eight species are currently

recognized, all of them flowering during the hot summer

months, mostly November to January. Most are ± pyro-

phytes, flowering most profusely in the season after the

veld has been cleared by burning.

Cladistic analysis of plastid DNA sequence data lo-

cates Thereianthus as sister to Micranthus (Pers.) Eckl.,

with these two genera together placed as sister to Watso-

nia Mill, plus Pillansia L. Bolus (Reeves et a/. 2002;

Goidblatt et al. 2008). These four genera, with Cya-

nixia Goidblatt & J.C. Manning, Lapeirousia Pourr. and

Savannosiphon Goidblatt & Marais, are now treated as

the tribe Watsonieae of subfamily Crocoideae (Goid-

blatt et al. 2008; Goidblatt & Manning 2008), charac-

terized by axillary conn development in which the cur-

rent season’s corm develops from an axillary bud at the

base of the stem rather than by expansion of the lower

stem intemodes as in other crocoid genera. Morphologi-

cal synapomorphies for Micranthus plus Thereianthus

are the superposed leaves, small, blue flowers, and ±

dry bracts with membranous margins (Goidblatt et al.

2006). The leaves in both Micranthus and Thereianthus

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town; Research Centre for

Plant Growth and Development, School of Biological and Conservation

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

X01, 3209 Scottsville, South Africa. E-mail: J.Manning@sanbi.org.za.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

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

goldblatt@mobot.org.

MS. received: 2011-02-15.

and fusiform seeds.

are all strictly cauline (Lewis 1954), with the lowermost

sub-basal, and the corm tunics are thus formed solely

from the cataphylls, without any contribution from the

bases of the lower leaves. Chromosome number, 2 n

= 20, and a karyotype of one long and nine short chro-

mosome pairs in Micranthus and Thereianthus is likely

plesiomorphic for the tribe. Watsonia, in contrast, has a

derived base chromosome number ofx = 9.

The moderately short spikes of spirally inserted flow-

ers, bracts with narrow membranous margins, and mono-

sulcate pollen distinguish Thereianthus from Micran-

thus, which has elongate, distichous spikes, bracts with

broad membranous margins, and zonasulcate pollen

(Goidblatt & Manning 2008). Our investigation of leaf

anatomy in the genus (see Results and discussion of

characters) indicates that a simple leaf margin is an addi-

tional anatomical apomorphy for Thereianthus.

Thereianthus was published by Lewis (1941), who

recognized seven species in the genus. One new species

has been described since then (Manning & Goidblatt

2004), at which time we also reduced T. lapeyrousioides

var. elatior G.J.Lewis to synonymy in T. minutus (= T.

lapeyrousioides var. lapeyrousioides). We have now had

the opportunity of examining fresh material of var. ela-

tior, which makes it clear that this taxon is in fact a dis-

tinct species. We also describe another two new species

that appeared after a bum near Tulbagh, bringing the

total number of species in the genus to eleven.

Taxonomic history, the taxonomic history of There-

ianthus is closely linked to that of Micranthus, reflect-

ing the marked similarities between the two genera and

the uncertainty that existed around their circumscrip-

tion. The first species of both genera known to science

were referred to either Ixia or to Gladiolus, following

the rather vague generic circumscriptions prevailing at

240

Bothalia 41,2 (201 1 )

the time but Ker Gawler (1804) subsequently included

all known species in Watsonici based on their divided

style branches. This treatment was not followed by Eck-

lon (1827), who concluded that the two genera were

distinct from Watsonia and who also appreciated the

differences between them. His new genus Micranthus

(Ecklon 1827), which was established for the small-

flowered species with strongly distichous spikes, was

recognized by most later authors but acceptance of a

second new genus for the larger-flowered species with

longer perianth tubes was less than universal. Eck-

lon ( 1 827) placed the two species of Thereianthus then

known. Gladiolus spicatus L. (now T. spicatus) and G.

triticeus Thunb. (now T. bracteolatus), in his new genus

Beilia but failed to provide a validating description, and

Beilia was thus not validly published. Heynhold (1847)

remained unconvinced that generic rank was appropri-

ate for Ecklon’s species of Beilia , preferring to include

them with the species of Micranthus , and he accordingly

made the new combinations M. spicatus (L.) Heyn. and

M. triticeus (Thunb.) Heyn. English bulb specialist, J.G.

Baker ( 1 877), in his account of the family, was of yet

another opinion, reverting to Ecklon’s circumscription of

Micranthus but following Ker Gawler (1804) in retain-

ing Thereianthus in Watsonia , as the unranked taxon

Beilia. Here he recognized just one species, W. punctata

(Andrews) Ker Gawk, with the earlier name, Gladiolus

spicatus L., in synonymy. At the same time he described

the third known species of Thereianthus , T. juncifolius,

in the genus Morphixia (now a synonym of Ixia). Beilia

was subsequently accorded the rank of subgenus within

Watsonia by Baker ( 1 892, 1 896), who admitted four taxa

at species rank, among them his erstwhile Morphixia

juncifolia and also W. lapeyrousioides. At this stage,

therefore, although Baker (1896) accepted both Micran-

thus and Thereianthus as currently circumscribed, only

the former was accorded generic rank.

Klatt (1882), a contemporary of Baker’s, utterly con-

fused the situation in his account of the Iridaceae. Here,

he included only two species in Micranthus , M. spicatus

(L.) Klatt and M. triticeus ( Burnt. f. [sic]) Klatt, both of

them actually species of Thereianthus , evidently una-

ware that Heynhold (1847) had already made the combi-

nations for these species in Micranthus. We also assume

that the attribution of the basionym to Burman fi 1 . was

an error for Thunberg, as Ixia triticea Burnt. f. (1768) is

a very different species [Tritoniopsis triticea (Burnt. f.)

Goldblatt] . Like Baker (1877) before him, Klatt (1882)

did not associate T. juncifolius with the other two spe-

cies, treating it instead in the genus Anomatheca as A.

calamifolia. He then spread his options still wider by

describing a fourth species of Thereianthus in Watso-

nia as W. racemosa Klatt. Inexplicably, Klatt over-

looked any species of Micranthus (as now understood)

in his account, an omission that he later partly cor-

rected (Klatt 1894) when he included M. plantagineus

[= M. alopecuroides (L.) Eckl.] in Micranthus with one

variety, var. junceus Baker [now M. junceus (Baker)

N. E.Br.]. In this work, T. spicatus remained in Micran-

thus and T. racemosus in Watsonia , where it was joined

by T. bracteolatus (as W. subulatus) but T. juncifolius

was transferred to Freesia as F. juncifolia (Klatt 1894).

This confusing species was later removed to yet another

genus by Brown (1931) as Lapeirousia juncifolia, by

which time it had been shuffled among no less than five

different genera. Lewis (1941) had no doubt that it cor-

rectly belonged in Thereianthus.

The nomenclatural tangle between Micranthus and

Thereianthus took a further turn when Kuntze (1891)

erected a new genus, Paulomagnusia, in which he

included both what are now Micranthus alopecuroides

and Thereianthus spicatus. The lectotype of Paulomag-

nusia is P. alopecuroides (now M. alopecuroides), des-

ignated by Goldblatt & Manning (2008) and the genus is

now a nomenclatural synonym of Micranthus. Ecklon’s

Beilia was raised to generic rank by Kuntze ( 1 898), who

included only B. spicata (L.) Eckl. ex Kuntze in the

genus. Unfortunately Beilia Kuntze is nomenclaturally

superfluous since Kuntze listed the validly published

Paulomagnusia in synonymy without any further expla-

nation. Kuntze also recognized var. brevifolia in his B.

spicata, and although this taxon has never been identi-

fied, its provenance from the Cape Town suburb of

Mowbray, and its broad leaves (10-20 mm) make it cer-

tain this is not a species of Thereianthus. It remained for

G.J. Lewis (1941), over a century after Ecklon proposed

Beilia, to publish the genus Thereianthus for the two

species of Ecklon’s Beilia plus an additional four more

species that were until then included in Watsonia.

Chromosome cytology, chromosome numbers are

known for just two species of Thereianthus, T. minutus

(sect. Brevibracteae) and T. bracteolatus (sect. Therei-

anthus). Both have a diploid number of 2 n = 20 and

an identical karyotype of one long and nine short chro-

mosome pairs (Goldblatt 1971 ). A report of 2n = 20 by

Goldblatt & Takei (1997) for T. spicatus is an error for

T. bracteolatus, thus representing a second count for that

species. The base number and karyotype match exactly

those of Micranthus but both are likely plesiomorphic

and thus uninformative about relationships. The two spe-

cies of Thereianthus counted, one from each section of

the genus, and the matching counts for Micranthus, sug-

gest that the basic karyotype is unlikely to differ among

the other species of Thereianthus.

Pollination systems : there are few published obser-

vations on the pollination of Thereianthus , and neither

Scott Elliot (1891) nor Vogel (1954) mentioned the

genus in their early studies of pollination in the South

African flora. Flowers of the genus are, however, diverse

and obviously adapted to a variety of mainly long-

tongued insect pollinators. At least T. ixioides has been

reported to be visited by the hopbine beetles, Hetero-

chelus sp. and Peritrichia sp. (Scarabaeidae: Hopliini).

Individuals of Peritrichia were observed moving indis-

criminately between the flowers of this species and the

remarkably similar blooms of Ixia metelerkampiae, the

pale mauve flowers of which also have a dark centre

(Goldblatt et al. 2000). These authors suggested this was

a possible example of Batesian mimicry. Although both

species have a filiform perianth tube in which the style

is tightly enclosed by the walls of the tube, T. ixioides

offers small amounts of nectar, visible at the mouth of

the tube, suggesting that the species offers a secondary

reward and may have a more generalist pollination sys-

tem than /. metelerkampiae, which does not offer nectar.

Goldblatt et al. ( 1998) also reported apparent pollination

of T. racemosus by the hopbine, Khoina bilateral is , in

Bothalia 41,2 (2011)

241

the Grootwinterhoek Wilderness area. This species, too,

produces traces of nectar and has a strong floral scent and

is likely to have alternative pollinators, probably various

bees.

Goldblatt & Manning (2006) inferred long-proboscid

fly pollination for Thereianthus longicollis, which has

a perianth tube 25-40 mm long. That prediction has

recently been validated: the long proboscid fly, Philoli-

che rostrata (proboscis ± 40 mm long), was captured

visiting flowers of this species in the mountains above

Tulbagh Waterfall in December 2010 (unpublished obs.).

Other representatives of this pollination guild at the site

included T. minutus, Geissorhiza confusa (Iridaceae)

and a long-tubed and large-flowered race of Lobelia

coronopifolia (Lobeliaceae) with a pink corolla. Visits

by P. rostrata to T. minutus (perianth tube 20-30 mm

long) were also recorded at this site. Another species, T.

elandsmontanus, also has flowers with the hallmarks of

long-proboscid fly pollination, notably a slender perianth

tube mostly 20-23 mm long and a white or pale pinkish

perianth with red markings. We predict that it is also pol-

linated by the long-proboscid flies Prosoeca guiosa and/

or P. rostrata.

We have noted the butterfly, Cynthia cardui (Nym-

phalidae), visiting the moderately long-tubed flowers of

Thereianthus spicatus at Jonkershoek. and moving from

flower to flower in a constant pattern. This insect may

be one of the legitimate pollinators of this widespread

species. We infer bee or generalist pollination for the

shorter-tubed species T. bracteolatus, T. bulbiferus and

T. intermedins. The long-tubed, violet flowers of T. mon-

tanus appear to be adapted for pollination by long-pro-

boscid flies or butterflies.

MATERIALS AND METHODS

All relevant types were examined, as well as all her-

barium material from BOL, NBG and SAM (acronyms

after Holmgren et ai. 1990), the primary collections of

species from the southwestern Cape. Most species were

also studied in the field.

For SEM work, fresh pollen grains were fixed in

FAA, critical point dried following standard procedures,

and examined at the SEM Unit, University of Cape

Town by Miranda Waldron. For operculum and sculp-

turing study, dry pollen from herbarium specimens was

rehydrated and stained in Calberla’s fluid (Goldblatt et

ai. 1991). Pollen was sampled from at least two flow-

ers per voucher. Seed shape and size was determined by

examining seeds from one or more capsules from two or

more different plants of each species.

For anatomical investigation, fresh material was

fixed in FAA and then embedded in wax before section-

ing and double-staining with alcian blue and safranin

following Rudall (1995). Mature leaf anatomy of the

following species was examined: Thereianthus race-

mosus, T. minutus, T. bulbiferus and T. spicatus. Ana-

tomical descriptions and terminology follow Rudall

(1995). Bract and bracteole anatomy was examined in T.

minutus and T. spicatus.

RESULTS AND DISCUSSION OF CHARACTERS

Leaf blade anatomy (Figures 1 A-F; 2)

Outline', surfaces smooth and ± parallel in Thereian-

thus bulbiferus, T. minutus and T. racemosus (Figures

1A-C; 2A-D), terete in T. juncifolius (Figure IF), ellip-

tical and fistulose in T. bracteolatus (Figures ID; 2E),

and abruptly raised over larger vascular bundles in T.

spicatus and surface thus ribbed or corrugated (Figures

1 E; 2F); pseudomidrib not evident except in T. minutus

but never raised in fresh leaves.

Margin : simple, i.e. without prominent marginal vein

or subepidermal sclerenchyma, rarely with small mar-

ginal bundle along one edge in T. bulbiferus', epidermal

cells unspecialized.

Mesophvll : cells ± isodiametric with no tissue dif-

ferentiation except at midrib area in T. minutus or with

evident differentiation between parenchyma and chlor-

enchyma (T. bulbiferus and T. juncifolius ) or ± radially

elongated with clear differentiation between peripheral

chlorenchyma and central mesophyll (T. bracteolatus , T.

racemosus and T. spicatus).

Vascular bundles', arranged in two rows, with larger

(primary and secondary bundles) in opposite pairs and

smaller (tertiary) bundles opposite or subopposite,

xylem poles usually separated, rarely fused in lateral

secondary bundles in T. minutus', primary bundles in T.

bulbiferus with tertiary bundle on each side contiguous

with sclerenchyma sheath; a single pair of primary bun-

dles can be identified in all species but this is most evi-

dent in T. minutus with differentiation between the pri-

mary bundle and secondary bundles less marked in other

species, particularly T. spicatus.

Bundle sheaths: bundle sheath sclerenchyma extend-

ing to epidermis as girder in primary bundles in all spe-

cies and also in secondary bundles in T. bracteolatus, T.

bulbiferus and T. spicatus, cells of outer bundle sheath

each containing solitary styloid crystal at phloem poles

in larger bundles; larger (primary) bundles with com-

plete sclerenchyma sheaths in T. bulbiferus, T. bracteo-

latus, T. juncifolius, T. minutus and T. spicatus but scari-

fication restricted to phloem caps in T. racemosus.

The most obvious distinction in leaf outline among the

species is between the ribbed/corrugated blade of There-

ianthus spicatus and the smooth blades of the other spe-

cies examined. The leaves of T. minutus and T. racemosus

are least sclerified, with only the primary bundle signifi-

cantly sclerified, compared with T. bulbiferus, T. bracteo-

latus, T. juncifolius and T. spicatus, in which some or all

of the secondary vascular bundles have completely scleri-

fied sheaths. This is most marked in T. spicatus.

Leaf blade anatomy in Thereianthus accords with the

general description for the genus (Rudall 1995) with the

significant exception that the leaf margin in all species

examined is essentially simple (i.e. lacking a prominent

marginal vascular bundle and with unspecialized epider-

mis) (Figure 2A, C-F). Rudall (1995). who examined

only T. spicatus , describes this species as possessing a

marginal vascular bundle. The outermost vascular bun-

dle is treated as marginal if it is prominent and solitary

(not paired) and oriented at right angles to the margin

with the phloem pole facing the margin. In T. minutus

and T. spicatus the outermost bundles at the leaf edges

242

Bothalia 41,2 (2011)

FIGURE I. — A-F, leaf tissue plans

of Thereianthus species: A, T.

minutus; B, T. racemosus', C,

T. bulbiferus; D, T. bracteola-

tus\ E, T. spicatus; F, T. junci-

folius. G, H, bract tissue plans:

G, T. spicatus ; H. T. minutus.

Scale bar: A-F, 500 pm; G,

H, 2 mm. Sclerenchyma solid

shading; xylem hatched. Art-

ist: John Manning.

FIGURE 2. — Leaf anatomy of Therei-

anthus species: A, T. minutus

leaf margin; B, T. minutus

pseudomidrib; C, T. racemosus

margin; D, T. bulbiferus mar-

gin; E, T. bracteolatus margin;

F, T. spicatus margin. Scale

bar: 200 pm.

Bothalia 41,2 (2011)

243

are clearly paired with the phloem poles facing the leaf

surface, and the leaves thus clearly lack marginal veins.

This is evidently also the condition in centric-leaved

T. bracteolatus and T. juncifolius but is less clear in T.

bulbiferus, in which a smaller bundle is located margin-

ally along only one edge of the leaf. In T. racemosus a

very small tertiary bundle occurs in a similar position

along one margin but is embedded within the chloren-

chyma and not located at the parenchyma/chlorenchyma

boundary as usual and is thus probably a minor branch.

A simple leaf margin is also characteristic of Pillcmsia

but not of Micranthus and Watsonia, both of which have

a well-developed marginal bundle with sclerenchyma

girder (Rudall 1995).

The presence of a marginal bundle is likely the ple-

siomorpic condition in Watsonieae (Rudall 1995). Opti-

mizing leaf margin condition on the molecular phy-

logeny of the family in Goldblatt et al. (2006, 2008)

indicates that it is most parsimonious to treat the simple

leaf margin as independently derived in Thereianthus

and Pillansia. Reversals to the simple leaf margins have

occurred several times elsewhere in the Crocoideae (e.g.

some species of Geissorhiza Ker Gawk, Hesperantha

Ker Gawl. and Lapeirousia). The character is most par-

simoniously treated as a homoplasious autapomorphy in

each instance.

Bract anatomy (Figure 1G, H)

The floral bracts of Thereianthus minutus are incon-

spicuously veined, with the vascular bundles lacking

sclerified phloem caps (Figure 1G), in contrast to the

prominently veined bracts in T. spicatus. in which the

vascular bundles have well-developed sclerenchyma

phloem caps (Figure 1H). As the bracts dry and the mes-

ophyll tissue collapses, the vascular bundles in T. spica-

tus become raised and very conspicuous whereas those

in T. minutus remain inconspicuous.

Pollen aperture and sculpturing (Figure 3)

Pollen aperture condition and exine sculpturing of

most species of Thereianthus was examined by Gold-

blatt et al. (1991). Their findings and those of the present

study covering the remaining species, are presented here

(Table 1 ). Pollen in all species is monosulcate and oper-

culate, with a tectate-columellate, sparsely scabrate tec-

tum. Operculum condition and exine sculpturing vary

in the genus. Exine sculpturing in most species is per-

forate (diameter of the lumina 0.25-1.0 * the width of

the intervening walls) (Figure 3 A, B) but is reticulate

(diameter of the lumina 2-3 x the width of the interven-

ing walls) in T. intermedins (Figure 3C, D), T. montanus

and T. racemosus , except on the operculum, where it is

perforate. The reticulate exine in these species recalls

the pollen of the sister genus Micranthus , the only other

genus of Crocoideae with reticulate exine but this genus

differs from Thereianthus and other Crocoideae in its

zonasulcate aperture (Goldblatt et al. 1991).

The operculum in most species is 1 -banded (Figure

3A, B) but is completely or incompletely 2-banded in

Thereianthus intermedius (Figure 3C), T. juncifolius ,

T. minutus , T. montanus and T. racemosus (Table 1).

A completely 2-banded (rarely 1 -banded) operculum

(the bands well separated by the apertural membrane)

is characteristic of T. intermedius, T. juncifolius and T.

FIGURE 3. — Pollen morphology of

Thereianthus species. A, B,

T. bracteolatus'. A, aperture

with 1 -banded operculum

(arrowed); B, monosulcate

grain with microperforate tec-

tum. C, D, T. intermedius'. C,

aperture with 2-banded oper-

culum (arrowed); D, reticulate

exine.

244

Bothalia 41,2 (201 1 )

TABLE 1 . — Pollen operculum type and exine sculpturing in Thereian-

ihus. Square brackets [ ] indicate closely associated operculum

bands, and curved brackets () indicate the less common opercu-

lum character state

racemosus but the operculum is incompletely 2-banded

in T. minutus and T. montanus. In these two species the

operculum is apparently 1 -banded but lacks exine along

the centre of the operculum in some or all grains, sug-

gesting incomplete fusion of two separate bands (Gold-

blatt et al. 1991 ).

A 2-banded operculum and perforate exine are likely

plesiomorphic conditions for Crocoideae, and more spe-

cifically for Watsonieae (Goldblatt et al. 1991), suggest-

ing that the species in sect. Brevibracteae with reticulate

exine as well as those species in sect. Thereiantlms with

1 -banded operculum are derived in these two respects.

MORPHOLOGY AND CLASSIFICATION

Thereiantlms is, relative to its size, quite as diverse

florally as other genera of Crocoideae (Goldblatt &

Manning 2006). Although the reproductive biology

remains poorly known, the species display a variety

of floral syndromes, including adaptations to short-

tongued, generalist, hopbine beetles, and to several

variants of long-proboscid fly or butterfly pollination

systems. Floral symmetry and perianth tube length

are valuable characters for discriminating the species.

Vegetative differences also provide critical diagnostic

characters. Four types of leaf blade are evident: linear

or lanceolate and plane with a single main or primary

vein (pseudomidrib) evident but not raised when fresh

and without evident secondary veins, the pseudomidrib

becoming raised in herbarium material through collapse

of the mesophyll tissue (71 minutus ); linear and plane

with more than one ± equally strong primary vein plus

secondary veins (71 bulbiferus, 71 intermedins, 71 minu-

tus, 71 montanus and 71 racemosus)', linear-filiform and

elliptic or terete in section without evident primary vein

(pseudomidrib) (71 bracteolatus and 71 juncifolius)', and

linear-corrugate with several equally strongly thick-

ened and raised primary veins plus secondary veins (71

elandsmontanus, 71 ixioides, 71 longicollis and 71 spica-

tus).

The striking variation in capsule shape and especially

in seed form that is evident in the genus has remained

undocumented until now. The remarkable seeds of

Thereiantlms minutus — fusiform with a long, thread-

like chalazal extension — were first noted by Manning

& Goldblatt (2004), who mistakenly interpreted the

thread-like extension as an elongated funicle. From later

examination of freshly opened, mature capsules, we are

able to confirm that the thread-like structure represents

a prolongation of the chalazal crest and not the funicle.

Similar filiform chalazal extensions of variable length

also characterize the seeds of T. intermedins, T. juncifo-

lius and T. racemosus (seeds unknown for T. montanus).

The thread-like chalazal extension is least developed in

T. racemosus, in which it is shorter than the seed body

and ± straight, but in T. intermedins , T. minutus, and T.

juncifolius the extension is two to three times longer

than the seed body and sharply bent back ± midway to

form a hooked or crook-like extension. The latter two

species have fusiform seeds and the chalazal extension

extends significantly beyond the seed body before bend-

ing back. In these two species the chalazal extensions

are largely accommodated in the elongated neck or beak

of the flask-shaped capsules. The function of the thread-

like extension is unknown.

Variation in pollen morphology in Thereiantlms was

first documented by Goldblatt et al. (1991), who discov-

ered two distinct types of operculum and exine sculptur-

ing among the species that they examined. The pollen

operculum in T. bracteolatus, T. ixioides, T. longicollis

and T. spicatus was clearly I -banded but was completely

or incompletely 2-banded in T. juncifolius, T. minutus

and T. racemosus. The latter condition was interpreted

as basic for the genus. Most species examined had

the usual microperforate tectum. The sole exception,

T. racemosus, had unusual reticulate exine sculptur-

ing, which was also found to characterize all species of

Micranthus. We have now examined pollen morphology

in all Thereiantlms species.

Thereiantlms species fall into two distinct, evidently

monophyletic groups on the basis of bract size and vena-

tion, seed shape, and pollen operculum condition: the T.

racemosus group, treated here as sect. Brevibracteae,

which is characterized by shorter bracts lacking promi-

nent veins, evidently specialized seeds with a filiform

chalazal extension (unknown in T. montanus), and pol-

len with a plesiomorphic, 2-banded operculum; and the

T. spicatus group, treated here as sect. Thereiantlms,

which has larger, prominently veined bracts, seeds with

an unspecialized chalazal crest, and apomorphic pollen

with a 1 -banded operculum. Leaf and bract anatomy, as

well as filament orientation and length, further group the

Bothalia 41,2 (2011)

245

members of sect. Thereianthus into two groups, treated

here as series. The four species with strongly thickened

and raised primary veins and shorter, erect or arcuate

filaments comprise ser. Thereianthus , and the remaining

two species with plane or smooth leaves and longer, dec-

imate filaments comprise ser. Bracteolatus.

TAXONOMY

Thereianthus G.J. Lewis in Journal of South Afri-

can Botany 7: 33 (1941). Lectotype: Thereianthus spica-

tus (L.) G.J. Lewis, designated by Goldblatt & Manning:

130 (2008).

Beilia Eckl.: 43 (1827), nom. nud.

Beilia Eckl. ex Kuntze: 305 (1898), nom. illegit.

superfl. pro Paulomagnusia Kuntze (1891) [= Micran-

thus (Pers.) Eckl., nom. cons.]. Type: Beilia spicata

(L.) Eckl. ex Kuntze (= Thereianthus spicatus (L.)

G.J. Lewis).

Watsonia unranked Beilia Eckl. ex Baker 16: 158

(1877). Type: Watsonia punctata (Andrews) Ker Gawl.

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

Watsonia subgen. Beilia Eckl. ex Baker: 177 (1892).

Lectotype: Watsonia punctata (Andrews) Ker Gawl. (=

Thereianthus bracteolatus (Lam.) G.J. Lewis), here des-

ignated.

Etymology >: from the Greek, thereios, summer, an-

thos, flower.

Deciduous geophytes with globose corm. axillary in

origin and rooting from below, tunics of coarsely net-

ted fibres. Cataphylls 2. Leaves 3 or 4(6), all cauline and

superposed, usually ± dry at flowering, lowermost sub-

basal, longest, inserted on stem above corm but below

ground level, blade falcate or linear with margins not

thickened, plane and either with definite main or primary

vein (pseudomidrib) or with several equally prominent

veins, or ± terete, glabrous or rarely scabridulous, usu-

ally dry at flowering, upper leaves progressively shorter,

uppermost sometimes entirely sheathing. Stem usually

unbranched, rarely with short branch from upper leaf

axil, terete in section. Inflorescence a crowded or rarely

lax spike, flowers spirally arranged or ± 2-ranked in bud,

entire spike loosely spirally twisted at anthesis; bracts

firm-textured, short, often overlapping, green below with

apices dry and brown or entirely dry, smooth or promi-

nently veined, inner 2-nerved and apically forked but

splitting to base as capsule matures, mostly membranous

(leathery in T. juncifolius). Flowers actinomorphic or

zygomorphic, long-lived, closing at night, usually blue

to purple, rarely white or pinkish, unmarked or lower

three tepals with darker markings in lower '/2, some-

times all tepals with white markings, usually unscented,

with nectar from septal nectaries; perianth tube short to

elongate, funnel-shaped or cylindric; tepals subequal, ±

spreading. Stamens unilateral and arcuate or declinate or

central and symmetrically arranged; filaments inserted

shortly below top of perianth tube, slender, free; anthers

linear, dehiscing longitudinally, usually purple. Ovary’

subglobose; style filiform, branches slender and deeply

divided, recurved. Capsules woody, ovoid to spindle- or

flask-shaped, tardily dehiscent. Seeds angular-elongate

with chalazal crest or narrowly ovoid-fusiform with

thread-like or filiform chalazal extension, smooth or

wrinkled, matte. Basic chromosome number, x = 10.

Eleven species, South Africa, restricted to the win-

ter rainfall zone in the southwestern portion of western

Cape, mostly in nutrient-poor, stony sandstone soils; two

Key to species

la Bracts short, 3-8 mm long, uniformly leathery or soft-textured and without prominent veins (sect. Brevibracteae ):

2a Leaf blades filiform-terete; spike lax, bracts distant (lower intemodes 3-5 * longer than bracts); inner bracts as long as or slightly longer

than outer and similar in texture, both uniformly green with dry, papery apices; tepals ovate-obovate; stamens central 5. T. juncifolius

2b Leaf blades linear to falcate, plane; spike dense, bracts ± imbricate (lower intemodes < twice as long as bracts); inner bracts shorter

than outer and membranous-papery except along veins; tepals lanceolate; stamens unilateral-arcuate:

3a Perianth tube very short, 1.0 — 1.5 mm long, included in bracts L T. racemosus

3b Perianth tube longer, 10-30 mm long, exserted from bracts:

4a Bracts 3-5 mm long; perianth tube 10-13 mm long; capsules ovoid, 4—5 mm long; plants of the Cold Bokkeveld, mountains

around Ceres, and Piketberg 2. I intermedius

4b Bracts (5-)6-8 mm long; perianth tube 20-30 mm long; plants from Grootwinterhoek to Riviersonderend Mtns and Kogelberg:

5a Leaves ± falcate, lowermost mostly 3-7 mm wide and usually much shorter than stem, with evident pseudomidrib but without

evident secondary veins, the pseudomidrib raised when dry but secondary veins never raised; perianth magenta or deep pink,

anthers pale yellow; filaments 3^1 mm long; capsules flask-shaped, 6-8 mm long 4. 71 minutus

5b Leaves linear, lowermost 2. 5-2. 8 mm wide and mostly longer than stem, with main and secondary veins raised when dry; peri-

anth and anthers purple; filaments 6-8 mm long; capsules unknown 3. 71 montanus

lb Bracts longer, (7— )8— 1 5 mm long, with prominently raised, closely set veins (sect. Thereianthus):

6a Leaves leathery without strongly raised veins; perianth tube curved and flowers thus ± nodding; filaments declinate, 6-10 mm long

(ser. Bracteolatus):

la Stem developing cluster of large cormels in axil of lowest leaf; leaves linear with evident veins; perianth tube 7-9 mm long; tepals

pale to mid-blue with white markings at base, veins not mottled, lateral longitudinal veins with numerous branches in distal half;

filaments 6-8 mm long 6. 71 bulbiferus

7b Stem without cormels; leaves terete or elliptic in section, without evident veins; perianth tube (7 — )8 — 1 0( — 1 5 ) mm long; tepals purple

to violet with prominent, darkly mottled veins, lateral longitudinal veins mostly without branches in distal half; filaments 8—10

mm long 7. T. bracteolatus

6b Leaves striate or corrugate with strongly raised veins; perianth tube straight or slightly curved distally and flowers thus suberect; fila-

ments central or arcuate, 3-5 mm long (ser. Thereianthus):

8a Perianth tube 25^40 mm long, > twice as long as tepals 1 1. 71 longicollis

8b Perianth tube 7-25 mm long, < twice as long as tepals:

9a Perianth tube filiform, tightly clasping style for most of length and then abruptly dilated at mouth; filaments central, symmetrical

and completely exposed, blocking mouth of perianth tube 9.T. ixioides

246

Bothalia 4 1 ,2 (2011)

9b Perianth tube narrowly funnel-shaped or ± cylindrical, not tightly clasping style; filaments arcuate, included within upper part of

tube and not blocking mouth:

10a Flowers usually blue, rarely white or pink, lower three tepals with white markings near base outlined distally with dark blue

chevron; tepals lanceolate, 3-5 mm wide 8. 71 spicatus

10b Flowers pale flesh-pink with purple tube, lower three tepals marked with purple in basal half; tepals linear, 1. 5-3.0 mm wide

10. T. elandsmontanus

species, Thereianthus juncifolius and T. minutus, grow in

seepages and along streams.

Sect. Brevibracteae Goldblatl & J.C. Manning,

sect. nov.

Plantae foliis filiformibus teretibusque vel line-

ari-falcatis planisque, cum vel sine nervo principali

prominenti sed nunquam fortiter nervatis marginibus

non incrassatisque; bracteis brevibus, 3-8 mm longis

uniformiter coriaceis vel modice mollibus sine costis

prominentibus; seminibus (ubi cognitis) ellipsoideo-fusi-

formibus cum appendice chalazae gracili vel filiformi,

rubro-brunneis, testa longitudinaliter rugosa vel obscure

scalariformi; grano pollinis operculo ± 2-vittato, exinio

perforato vel reticulato.

Type species: T. racemosus (Klatt) G.J. Lewis.

Leaves filiform and terete, or linear to falcate and

plane, with or without prominent main vein but never

strongly ribbed; bracts short, 3-8 mm long, uniformly

leathery or moderately soft-textured, without prominent

veins. Pollen operculum ± 2-banded, exine perforate

or reticulate. Seeds (where known) ellipsoid-fusiform

with filiform or thread-like chalazal extension, red-

dish brown, testa sculpturing longitudinally rugose or

obscurely scalariform.

I . Thereianthus racemosus (Klatt) G.J. Lewis

in Journal of South African Botany 7: 36 (1941).

Watsonia racemosa Klatt 15: 354 (1882). Type: South

Africa, [Western Cape], ‘in montibus ad Vier en Twin-

tig Rivieren’, Dec. without year, Zevher 1609 (B, holo.;

BOL!, GRA, K (2 sheets)!, PRE, SAM (3 sheets)!, iso.).

Plants 130-250 mm high. Conn globose, ± 10 mm

diam.; tunics of moderately fine-textured to coarsely net-

ted fibres, sometimes forming short neck. Cataphylls dry

and papery, reddish brown. Stem erect or flexed slightly

outward above sheath of second leaf, unbranched or

very rarely with short branch from upper leaf axil, 0.8-

1.0 mm diam. below spike. Leaves (3)4(5), lowermost

inserted below ground, blade reaching base of spike

or not, drying from tip at flowering, linear or falcate,

60—1 30(— 1 90) x l-3(-4) mm, thick-textured, plane with

main vein not thickened or raised when fresh and usu-

ally without evident secondary veins, rarely with 1 sec-

ondary vein visible on each side, upper 3 leaves cauline

with lower 2 inserted on lower 'A of stem, sheathing for

'A to 2A length with blades 20-40 mm long, uppermost

leaf entirely sheathing, well separated from spike, all

sheaths shortly imbricate or uppermost a little distant.

Spike erect, moderately densely 5-25-flowered; bracts

shortly imbricate, outer 3-5 mm long, up to 1 internode

long, acute, green below or entirely brown and leathery

with narrow membranous margins and papery tip, with-

out evident veining, inner ± as long or slightly longer,

notched apical ly, ± membranous with narrow brown

zone along veins. Flowers suberect, pale blue, lower

three tepals white at base with purple median chevron

in lower 'A, strongly honey-scented; perianth sub-actin-

omorphic, funnel-shaped; tube straight, 1.0-1. 5 mm

long, narrowly funnel-shaped, included in bract; tepals

elliptical-oblanceolate, subequal, spreading and slightly

cupped, 7-9 x 3_4 mm. Stamens unilateral but later-

als slightly spreading; filaments arcuate, 3-4 mm long,

exserted 1.5-2. 5 mm from tube; anthers 4-5 mm long,

dark blue; pollen pale blue. Ovary ovoid, ± 2 mm long;

style arching over stamens, dividing between base and

middle of anthers, branches 2. 0-2. 5 mm long, divided

for ± V2 their length. Capsules ovoid, 4-5 x 2. 5-3.0 mm.

Seeds ovoid, ± 1 mm long, with ± straight, thread-like

chalazal extension ± 0.5 mm long, reddish brown. Flow-

ering time'. Dec. Figure 4.

Distribution and ecology. Thereianthus racemosus

appears to be restricted to the Voorberg (also known as

the Twenty Four Rivers Mtns) above Porterville (Figure

5), where it occurs on seasonally moist or marshy, sandy

flats in restioid communities. Most collections are from

the Suurvlakte in the Grootwinterhoek Wilderness Area.

It is an obligate pyrophyte, flowering only in the sum-

mer following a bum.

The short-tubed, pale blue flowers are strongly honey-

scented and are evidently adapted to a variety of short-

tongued pollinators, including various Hymenoptera, Lep-

idoptera, and Coleoptera. Goldblatt et al. (1998) reported

pollination of T. racemosus by the hopbine beetle, Khoina

bilatera/is, in the Grootwinterhoek Wilderness area.

Diagnosis and relationships', the short floral bracts,

3-5 mm long, and the very short perianth tube, 1.0-1. 5

mm long, and well included within the bracts, are diag-

nostic for the species. Uniquely for the genus, the flow-

ers are fragrant and strongly honey-scented. The narrow,

leathery leaves with ± distinct main vein but usually

without evident secondary veins, the very short cap-

sules, small seeds, 1.0-1. 5 mm long, and reticulate pol-

len exine suggest that the species is most closely allied

to Thereianthus intermedius from the Cold Bokkeveld.

That species has darker blue flowers with a much longer

perianth tube, 10-13 mm long, and seeds with the chala-

zal extension longer than the seed body.

History, the species was first collected by botani-

cal collector Carl Ludwig Zeyher, who was active in

the Western Cape in the first half of the nineteenth cen-

tury. Although the type collection is not dated, Zeyher

appears to have collected on the Twenty Four Rivers

Mtns only once, in 1831, when he made a rich haul of

specimens from the mountains around Tulbagh in the

company of his sometime partner Christian Frederick

Ecklon (Glen & Germishuizen 2010), and the species

was probably discovered then. It has since been col-

lected just a handful of times on the same mountains,

always following veld fires.

Bothalia 41,2 (2011)

247

FIGURE 4. — Thereianthus racemosus. A-D, Goldblatt 10454: A, whole

plant; B, flower, front view; C, flower, side view; D, details of sta-

mens and stigma. E, F, Manning 3289 : E, capsule; F, seed. Scale bar:

A-C, 10 mm; D, 3 mm; E, 2 mm; F, 0.5 mm. Artist: John Manning.

Additional specimens examined

WESTERN CAPE. — 3319 (Worcester): Twenty Four Rivers Mtns

above Porterville, (-AA), 16 Dec. 1949, Esterhuysen 16620 (BOL);

16° 18° 20° 22° 24° 26° 28°

FIGURE 5. — Known distribution of Thereianthus racemosus, •; T.

intermedius, O.

Groot Winterhoek Wilderness Area, (-AA), 14 Dec. 1995, Steiner

3061 (NBG); near Groot Winterhoek Forest Station, (-AA), 27 Dec.

1995, Goldblatt 10454 (MO, NBG); Grootwinterhoek Wilderness

Area, Suurvlakte, (-AA), 8 Jan. 2010 (fr.). Manning 3289 (NBG).

2. Thereianthus intermedius J.C. Manning &

Goldblatt, nom. et stat. nov., T. lapeyrousioides var. ela-

tior G.J. Lewis in Journal of South African Botany 7: 38

(1941). Type: South Africa, [Western Cape], Ceres Hills,

Jan. 1892, L. Guthrie 2208 (BOL, holo.!).

Plants 200-350 mm high. Corm globose, ± 10 mm

diam.; tunics of fine-textured, netted fibres accumulating

with age and forming neck around base of stem. Cata-

phylls dry and papery, reddish brown. Stem erect, flexed

slightly outward above sheath of second leaf, unbranched,

1.0-1. 5 mm diam. below spike. Leaves (3)4, lowermost

inserted below ground, blade reaching base of spike or not,

drying from tip at flowering, linear, 70-150 x 2-3 mm,

thick-textured, with main vein ± evident but not thick-

ened or raised when fresh, plus a pair of secondary veins,

upper (2)3 leaves cauline with lowermost (1)2 inserted on

lower xh of stem, sheathing for V2 to 2h length with blades

30-60 mm long, uppermost leaf entirely sheathing or with

short blade up to 20 mm long, well separated from spike,

all sheaths shortly imbricate or uppermost a little distant.

Spike erect, moderately densely 12-40-flowered; bracts

not or scarcely imbricate, outer 3-5 mm long, up to 1

intemode long, obtuse, brown and leathery with narrow

membranous margins, without evident veining, inner ±

as long or slightly longer, notched apically but sometimes

splitting to base after fertilization by enlarging capsule, ±

membranous with narrow brown zone along veins. Flow-

ers suberect, violet (rarely pale blue), lower three tepals

whitish at base with purple or dark blue median chevron

in lower 'A; perianth subactinomorphic; tube straight or

slightly arching in upper ± 3 mm, 10-13 mm long, cylin-

drical and widening slightly in upper ± 3 mm; tepals nar-

rowly oblanceolate, subequal, spreading and slightly

cupped. 10-11 x 3. 0-3. 5 mm. Stamens unilateral; filaments

erect, ± 7 mm long, exserted ± 4 mm from tube; anthers

purple, 4—5 mm long; pollen violet. Ovary ovoid, 2. 0-2. 5

mm long; style arching over stamens, dividing below base

of anthers, branches ± 2.5 mm long, divided for ± V2 their

length. Capsules ovoid, 4-5 x 3-4 mm. Seeds ovoid, ± 1.5

x 1 .0 mm, with thread-like chalazal extension up to 3 mm

long abruptly reflexed from near middle, reddish brown,

testa striate-rugulose. Flowering time : Dec. Figure 6A-F.

248

Bothalia 41,2 (201 1 )

Distribution and biology: originally recorded from 1941 ), Thereianthus intermedins has now also been col-

two nnlocalized collections made near Ceres (Lewis lected in the Cold Bokkeveld at the headwaters of the

FIGURE 6. — A-F: Thereianthus intermedins, Hanekom & Tilney s.n. A, whole plant; B, half flower; C, outer bract; D, inner bract; E, capsule; F,

seeds. G-J: T. montanus, Oliver 12197. G, whole plant; H, flower, front view; I, half flower; J, outer (left) and inner (right) bracts. Scale bars:

A, B, G-J, 10 mm; C-E, 2 mm; F, 0.5 mm. Artist: John Manning.

Bothalia 41 ,2 (201 1 )

249

Olifants River and at high elevations on Zebrakop in the

Piketberg (Figure 5). In the north it has been collected

from the farms Elandskloof and Kunje along the eastern

foot of the Cold Bokkeveld Mtns, with another collec-

tion from Visgat on the western edge of the Skurweberg,

and in the south from Castle Rocks at the western edge

of the Hex River Mtns above Ceres. It is restricted to

seasonally waterlogged or marshy, sandy flats dominated

by restioids, between ± 500 and 1 000 m, and flowers

only after fire.

We assume that the moderately long-tubed flowers

are pollinated by various long-tongued insects, including

anthophorine bees (Apidae: Anthophorinae) and butter-

flies.

Diagnosis and relationships'. Thereianthus interme-

dins is recognized by its linear leaves, 2-3 mm wide,

elongate spike with short floral bracts, 3-5 mm long, and

violet (pale blue on Piketberg) flowers with moderately

long floral tubes, 10-13 mm long, and slightly longer

than the tepals. The leaves have an additional pair of

secondary veins, one running on each side of the central

vein close to the leaf margin, and the spike is moderately

dense, with the bracts shorter than or scarcely as long

as the intemodes. The small, ovoid capsules, 4-5 mm

long, contain distinctive seeds, with an ovoid seed body

and long, thread-like chalazal extension ± twice as long

as the seed body and sharply bent backwards along the

length of the seed.

Thereianthus intermedins is most likely to be con-

fused with T. minutus, which has similar plane leaves

(but with the pseudomidrib much less distinct), short

bracts, and long-tubed flowers. It differs most obviously

from T. minutus in its purple or blue (vs magenta or deep

pink) flowers with a shorter floral tube (10-13 mm vs

20-30 mm), shorter bracts (3-5 mm vs 6-8 mm) and in

its smaller capsules (ovoid and 4-5 mm long vs flask-

shaped and 6-8 mm long) capsules containing ovoid

(vs fusiform) seeds. The distribution of the two species

is complementary, with T. intermedins ranging from the

Cold Bokkeveld to the Skurweberg, the western Hex

River Mtns and Piketberg (Figure 5), and T. minutus dis-

tributed from the Grootwinterhoek Mtns above Tulbagh

through the Franschhoek and Du Toitskloof Mtns to the

Kogelberg and western end of the Riviersonderend Mtns

(Figure 7).

16° 18° 20° 22° 24° 26° 28°

FIGURE 7. — Known distribution of Thereianthus minutus , •; T. mon-

tanus, O.

History', the species seems to have been first col-

lected by botanist Louise Guthrie in January 1892, and

has been recorded only sporadically since then, most

recently by local landowner Louis Hanekom, who

noticed it flowering for the first time on his Farm Kunje

in Dec. 2009 after the extensive wildfire that swept the

area in the summer of 2008/09. He passed the plants on

to Jeanetta Tilney, who has been documenting the spe-

cies on her property on Middelberg Pass for some years

and who in turn brought it to our attention.

Thereianthus intermedins was first described as var.

elatior of T. lapeyrousioides [now T. minutus (Klatt)

G.J. Lewis (Goldblatt 1989)], distinguished from the

typical variety by its taller stature (240^160 mm high),

slightly longer, narrower leaves, and longer, more flor-

iferous (12-40-flowered) spike with shorter bracts ± 5

mm long (Lewis 1941 ). Subsequently, Manning & Gold-

blatt (2004) argued that collections of T. minutus from

Bainskloof displayed the full range of variation in these

features and concluded that var. elatior could not be rec-

ognized at any taxonomic level. The recent collection of

fresh material of the plants from the Farm Kunje shows

that we were wrong and that other, more significant dif-

ferences not identified by Lewis (1941) exist between

the two taxa, justifying their recognition as distinct spe-

cies. These include perianth colour, floral tube length,

shape and size of the capsule, seed shape and pollen

exine sculpturing.

Etymology : Thereianthus intermedins approaches T.

minutus in its seeds with filiform chalazal extension but

resembles T. racemosus in its short capsules and reticu-

late pollen sculpturing, and we thus propose the name T.

intermedins in recognition of its somewhat intermediate

character.

Additional specimens seen

WESTERN CAPE.— 3218 (Clanwilliam): Piketberg, SW slopes of

Zebrakop, (DA-DB). 4 Jan. 1995, Goldblatt & Manning 10168 (MO).

3219 (Wuppertal): Elandskloof, marshy places, 3000' [915 m], (-CA),

18 Dec. 1948, Levyns 9349 (BOL). 3319 (Worcester): Visgat, upper

Olifants River Valley, (-AA), 28 Dec. 1946, Esterhuysen 13435 (BOL,

NBG, PRE); Ceres, 1700' [518 m], (-AD), Jan. 1903, Bolus 8342

(BOL, PRE); Ceres, (-AD), Jan. 1929, Thode A2287 (NBG, PRE);

Ceres, (-AD), Jan. 1932, Leslie s.n. NBG46/32 (BOL); Ceres, Castle

Rocks, vlakte [flats] on NE slopes, abundant after fire, 3800' [1 158

m], (-AD), 14 Jan. 1960, Esterhuysen 28430 (BOL, MO).

3. Thereianthus montanus J.C. Manning & Gold-

blatt in Bothalia 34: 103 (2004). Type: South Africa,

Western Cape, Riviersonderend, Pilaarkop, ridge WNW

of peak, 31 Jan. 2004, E.G.H. Oliver 12197 (NBG,

holo.!; MO, iso.!).

Plants 200-350 mm high. Conn globose, 7-10 mm

diam.; tunics of fine-textured, netted fibres accumulat-

ing with age and forming neck around base of stem.

Stem erect, flexed outward above sheath of second leaf,

unbranched, 1.0-1. 5 mm diam. below spike. Cataphylls

dry and papery, reddish brown. Leaves 3 or 4, lower-

most inserted below ground, blade reaching or exceed-

ing spike, linear, 150-300 x 2. 5-2. 8 mm, thick- textured,

with main vein evident but not thickened or raised when

fresh, plus 1 or 2 evident secondary veins on each side,

upper 2 or 3 leaves cauline with lower 1 or 2 inserted on

lower 'A of stem, sheathing for V2 to 2h length with short

250

Bothalia 41,2 (2011)

blades 15-25 mm long, uppermost leaf well separated

from spike, short and entirely sheathing, all sheaths dis-

tant. Spike erect, compact, densely 7-10-flowered; bracts

imbricate, outer 6-8 mm long, obtuse to truncate, green

and leathery below, dry and brown in upper V2, ± 2 inter-

nodes long, inner ± 1.5 mm shorter than outer, notched

apically, brown and papery with narrow darker zone

along veins. Flowers suberect, purple, lower three or all

tepals each with spear-shaped, purple median streak near

base, throat and lower part of tube white; perianth sub-

actinomorphic, salver-shaped; tube straight or slightly

arching in upper ± 5 mm, 22-27 x 1.2— 1.5 mm, cylin-

drical and widening slightly in upper ± 5 mm; tepals

spreading and slightly cupped, narrowly elliptical to

lanceolate, subequal, 9-15 x 3. 5-5.0 mm. Stamens uni-

lateral; filaments erect, 6-8 mm long, exserted 3-4 mm

from tube; anthers 4-5 mm long, purple; pollen violet.

Ovary ovoid, 2. 0-2. 5 mm long; style arching over sta-

mens, dividing between base and middle of anthers,

branches ± 2.5 mm long, divided for ± V2 their length.

Capsules and seeds unknown. Flowering time : late Jan-

Feb. Figure 6G-J.

Distribution and biology: Thereianthus montanus is

known from a single population on steep, south-facing

slopes of Pilaarkop in the Riviersonderend Mtns (Fig-

ure 7). Plants are scattered in moist, loamy soil in short,

grassy fynbos at an altitude of ± 1 500 m.

The long-tubed, violet flowers appear to be adapted

for pollination by long-proboscid flies or butterflies

(Manning & Goldblatt 2004), and accumulate nectar

only in the lower few millimetres, which is consistent

with pollination by long-proboscid insects with mouth-

parts 20-25 mm long.

Diagnosis and relationships'. Thereianthus montanus

is distinguished by its linear leaves without thickened

or raised main vein when fresh, although the mesophyll

tissue collapses on drying to leave the central and sec-

ondary veins apparently raised in herbarium specimens,

and compact spikes of salver-shaped, purple flowers

with slender perianth tube 22-27 mm long and purple

anthers. It is most likely to be confused with long-tubed

T. minutus with similarly long-tubed flowers and which

has been recorded from the western Riviersonderend

Mtns, but this species is generally shorter, mostly 1 00—

200 mm tall, with a broader, mostly falcate lower leaf,

( 1 .5— )3 .0—7.0 mm wide and rarely reaching the base of

the spike, and with a prominent pseudomidrib but incon-

spicuous secondary veins. On drying, the main vein in

the leaves of T. minutus becomes raised through collapse

of the surrounding blade tissue but the secondary veins

are never pronounced. The flowers in T. minutus are a

distinctive magenta colour, with yellow anthers. Unfor-

tunately fruits and seeds are not known in T. montanus

and so cannot be compared with the distinctive flask-

shaped capsules and fusiform seeds with thread-like

chalazal extension of T. minutus. The two species differ

in pollen exine sculpturing. T. minutus has an unspecial-

ized perforate pollen exine, unlike the unusual reticu-

late pollen exine of T. montanus, which suggests that its

closest allies are actually T. racemosus from Grootwin-

terhoek and T. intermedius from the Cold Bokkeveld,

being the only other two species in the genus with retic-

ulate exine.

History', a single flowering stem of the species was

first collected by botanists Ted and Inge Oliver in Feb-

ruary 1999, and the type material was collected by Ted

Oliver from the same locality five years later, in January

2004.

Additional specimen examined

WESTERN CAPE. — 3419 (Caledon): Riviersondered Mtns,

Pilaarkop, ridge WNW of peak, ± 1 500 m, (-BB), 26 Feb. 1999,

E.G.H. <£ I.M. Oliver 11228 (NBG).

4. Thereianthus minutus (Klatt) G.J. Lewis in

Journal of South African Botany 7: 43 (1941). Watsonia

minuta Klatt: 353 (1882). Type: South Africa, [Western

Cape], Tulbagh Waterfall, Ecklon & Zeyher Irid. 189 (B,

lecto.!, designated by Goldblatt 19: 143 (1989); MO!,

S!, SAM!, isolecto.) [Syntype: South Africa, [Western

Cape], Dutoitskloofberge am Wasserfall, Drege 1551 (B

‘Herb. Liibeck’, believed destroyed)].

Watsonia lapeyrousioides Baker: 178 (1892). Thereianthus lap-

eyrousioides (Baker) G.J. Lewis 7: 38 (1941). Type: South Africa,

[Western Cape], clivis montium inter Villiersdorp & French Hoek

[Franschhoek], Nov. 1879, Bolus 5251 (BOL, lecto.!, here designated;

K., isolecto.!) [Syntype: South Africa, [Western Cape], Du Toits-

kloof, without date [‘1840' on specimen], Drege s.n. (K [K3205 1 l]l,

2S\M[SAM21067\\}.

Plants 100-200(-300) mm high. Conn globose, 7-10

mm diam.; tunics of fine-textured, netted fibres, some-

times accumulating with age and forming short neck

around base of stem. Stem usually inclined but flexed

outward above sheath of second leaf and then erect,

unbranched, 0.8-1. 5 mm diam. below spike. Cataphylls

dry and papery, reddish brown. Leaves 3 or 4(— 6), lower-

most inserted below ground, blade mostly much shorter

than spike, rarely exceeding it, mostly falcate, rarely lin-

ear-arcuate, (30-)40-100(-350) x ( 1 .5— )3— 7 mm, thick-

textured, with main vein evident but not raised when

fresh (becoming raised when dry), plus 1 or 2 evident

secondary veins on each side, upper leaves cauline with

lower I or 2 inserted on lower V3 of stem, sheathing for

V2 to 2F length with blades 20^10 mm long, uppermost

leaf well separated from spike, short and entirely sheath-

ing, all sheaths shortly imbricate or distant. Spike erect,

compact, moderately densely (3-)5-10(-20)-flowered;

bracts shortly imbricate, outer (5-)6-8 mm long, obtuse

to truncate, green and leathery below, dry and brown

in upper V2, ± 1.5 internodes long, inner ± 1.0-1. 5 mm

shorter than outer, notched apically, brown and papery

with narrow darker zone along veins. Flowers suberect,

dark pink to magenta, lower three tepals usually with

darker median mark near base, throat white; perianth

subactinomorphic, salver-shaped; tube straight, 20-30 x

1.2-1. 5 mm, widening gradually and evenly from base;

tepals spreading and slightly cupped, elliptical to oblan-

ceolate, subequal, 7-10 x 2^1 mm. Stamens unilateral;

filaments erect, 3^4 mm long, exserted ± 2 mm from

top of tube; anthers 3^4 mm long, pale yellow; pollen

cream-coloured. Ovary ovoid, 2. 0-2. 5 mm long; style

arching over stamens, dividing just below or just above

base of anthers, branches ± 4 mm long, divided for ± V2

their length. Capsules flask-shaped, 6-8 mm long. Seeds

fusiform, 1 .5-2.0 mm long, with chalazal extension up

to 4 mm long, thus ± twice as long as seed body, sharply

bent back ± midway and hooked or crook-like, some-

times twisting around adjacent seeds, reddish brown.

Bothalia 41,2 (2011)

251

FIGURE 8. — Thereianthus minutus. A-C, Goldblatl & Manning 9505:

A, whole plant; B, flower, front view; C, flower, side view. D, E,

Manning 3327 : D, capsules; E, seed. Scale bar: A-C, 10 mm: D.

2 mm; E, 0.5 mm. Artist: John Manning.

longitudinally rugulose. Flowering time : Dec.-Jan.,

rarely as early as late Oct. Figure 8.

Distribution and ecology >: endemic to the mountains

in the extreme southwestern part of Western Cape, from

above Porterville southwards to the Kogelberg and

western Riviersonderend Mtns (Figure 7). Thereian-

thus minutus is restricted to moist habitats such as rock

ledges, damp cliffs and streamsides, growing in flushes,

peaty seepages and mossy turf. Fires are necessary for

flowering to occur in plants on deeper, more heavily

vegetated soils but in exposed habitats plants flower reg-

ularly without burning.

The long-tubed, magenta flowers are consistent

with pollination by long-proboscid flies and we have

observed visits to the species by Philoliche rostrata

(Tabanidae) above Tulbagh Waterfall in December 2010.

Diagnosis and relationships'. Thereianthus minu-

tus is distinguished by its mostly falcate, relatively

broad lower leaf blade, mostly 3-7 mm wide, with a

pronounced main vein, and dark pink to magenta flow-

ers with perianth tube 20-30 mm long. On drying and

in herbarium material the pseudomidrib becomes raised

through collapse of the surrounding mesophyll tissue

but the secondary veins remain inconspicuous. The short

bracts, 5-8 mm long, without prominent veins are typi-

cal of sect. Brevibracteae but the pale yellow anthers are

unusual in the genus and found elsewhere only in T. jun-

cifolius (sect. Brevibracteae) and T. bracteolatus (sect.

Thereianthus). It is most likely to be confused with T.

intermedius and T. montanus, both of which have similar

long-tubed flowers but of the more typical blue to purple

colour and with purple stamens. The three species have

complementary ranges: T. intermedius from the Piket-

berg, Cold Bokkeveld and Skurweberg has a perianth

tube 10-13 mm long, shorter bracts 3-5 mm long, and

smaller, ovoid capsules 4-5 mm long; T. montanus from

the eastern Riviersonderend Mtns has similar bract and

perianth tube dimensions but longer filaments, 6-8 mm

long and exserted 3—4 mm from the tube, and mostly

narrower leaves, 2. 5-2. 8 mm wide, with the main and

also secondary veins becoming raised on drying and

in herbarium material. T. montanus is generally a taller

species, 200-300 mm tall, with the lower leaf longer

than the spike. In addition, the perianth tube in T. mon-

tanus is virtually cylindrical throughout, widening only

in the upper 5 mm, unlike the tube in T. minutus, which

is slightly narrower below but widening gradually and

evenly towards the top.

The fruit and seeds of Thereianthus montanus are

unfortunately not known but the flask-shaped capsules

of T. minutus are distinctive, as are the fusiform seeds

with long, thread-like chalazal extension ± twice as long

as the seed body. The placenta is restricted to the lower

portion of the capsule and the tapered neck or beak

accommodates the chalazal extensions of the seeds.

History'. Thereianthus minutus was first collected by

the botanical collectors Christian Frederick Ecklon and

Carl Ludwig Zeyher at Tulbagh Waterfall, possibly in

1831 when they visited this site (Glen & Germishuizen

2010), and almost contemporaneously by the naturalist

and traveller Carl Drege, who found it in the Du Toitsk-

loof Mtns sometime between 1826 and 1834. The date

1840 marked on the type collection at Kew is not the

date of collection as Drege returned to Europe in 1834

(Glen & Germishuizen 2010), and is probably the date

that the specimen was offered for sale and acquired by

the herbarium. These two collections formed the basis

of both Watsonia minuta Klatt (1882) and W. lapey-

rousioides Baker (1892). At the time of her revision of

Thereianthus, Lewis (1941) had not been able to iden-

tify Klatt’s W. minuta and although she transferred the

name to Thereianthus she treated the taxon as a doubt-

ful species, possibly near T. racemosus, based on Klatt’s

curious and quite mistaken description of the perianth

tube as a mere 4 mm long. She accordingly recognized

T. lapeyrousioides as a separate species. Ironically, and

quite inexplicably, she overlooked the significance of

the Ecklon & Zeyher 189 syntype of W. minuta at SAM,

which she cited under T. lapeyrousioides and which

would have established the identity of Klatt’s species. It

was left to Goldblatt (1989) to recognize that this was

the earliest name for the species.

The Ecklon & Zeyher collections were distributed

under the unpublished name Beilia triticea Eckl. (now

applied to Thereianthus bracteolatus), while the Drege

material was misidentified as Anomatheca juncea (Ker

Gawl.) Ker Gawl. (= Freesia verrucosa). Plants from the

Cold Bokkeveld, treated by Lewis (1941) as T. lapey-

252

Bothalia 41,2 (201 1)

rousioides var. elatior G.J. Lewis, are recognized here as

a distinct species, T. intermedins.

Additional specimens examined

WESTERN CAPE. — 3318 (Cape Town): Jonkershoek Mtns, (—

DD). 1826, Marloth s.n. (PRE); Jan. 1888, 'Mar loth s.n. (NBG); Jonk-

ershoek State Forest, Stellenboschberg, (-DD), 8 Dec. 1975, Haynes

1180 (NBG); Stellenbosch. Swartboskloof, (-DD), 25 Oct. 1960,

Van der Merwe 23-94 (NBG, PRE). 3319 (Worcester): Winterhoeks-

berg, (-AA), Nov., Ecklon & Zevher Irid. 188 (72.11) (MO); Tulbagh,

Watervalsberge, east of Suurvlakte Falls, (-AC), 29 Dec. 2000, Hans-

ford 22 (NBG); lower Wellington Sneeukop, (-CA), 23 Dec. 1945,

Esterhuysen 1254 (BOL); Bain’s Kloof [Bainskloof], (-CA), 12 Nov.

1896, Schlechter 9126 (BOL, MO, PRE); 28 Nov. 1926, Grant 2648

(BOL, MO); 27 Nov. 1939, Bond 5 (NBG); 17 Jan. 1945, Compton

16922 (NBG); 9 Nov. 1946, Compton 18682 (NBG); 6 Nov. 1948,

Acocks 15238 PRE); Nov. 1949, Stokoe s.n. SAM60115 (SAM); Dec.

1957, Loubser 866 (NBG); 24 Nov. 1979, Goldblatt 5221 (PRE); 27

Nov. 1990, Steiner 2265 (MO, NBG); Baviaanskloof, above pools in

middle of valley, (-CA), 12 Jan. 1993, Goldblatt & Manning 9505

(MO); (-CA), 15 Jan. 2011 [fruiting]. Manning 3327 (NBG); Wolwek-

loof Forest Reserve, (-CA), 20 Oct. 1946, Barker 4249 (NBG); Lim-

ietberg, (-CA), 28 Nov. 1982, Esterhuysen 35830 (MO, PRE), Jackson

s.n. (NBG); Slanghoek Mtns, Witte River Valley, (-CA), Nov. 1922,

Thorne SAM46563 (SAM), 21 Nov. 1943, Wasserfall 614 (NBG), 28

Dec. 1975, Esterhuysen s.n. (MO); Du Toit's Kloof, (-CA), Jan. 1880,

Bolus 5497 (BOL); 10 Dec. 1945, Esterhuysen 12369 (BOL); Dec.

1949, Stokoe s.n. (NBG, SAM60114); Rawsonville, veld opposite

Farm Gevonden, (-CA), 3 Nov. 1962, Walters 912 (NBG); Du Toit-

skloof Mtns, Elandsberg catchment, (-CC), 29 Dec. 2000, Low 6770

(NBG); Slanghoek Valley, The Cossacks, (-CC), 16 Dec. 1997, Oliver

11004 (NBG); Paarl, Haalhoek Spitzkop, (-CC), 2 Jan. 1947, Ester-

huysen 13528 (BOL, NBG), 19 Dec. 1981 Esterhuysen 35731 (BOL,

MO); Wemmershoek area, Klein Drakenstein Mtns above Zachari-

ashoek, (-CC), 1 Jan. 1983, Oliver 7928 (NBG, PRE); French Hoek

[Franschhoek], (-CC), 23 Dec. 1895, Bolus 4010 (MO, NBG, PRE),

Jan. 1949, Stokoe s.n. SAM60178 (SAM); Franschhoek Pass, Adolph's

Kop, (-CC), 15 Dec. 1944, Esterhuysen 11047 (BOL, PRE); western

Riviersonderend Mtns, 26.4 km west of Olifantsberg, (-CD), 18 Dec.

2007, Helme 5172 (NBG). 3418 (Simonstown): Jonkershoek Valley,

near top of Kurktrekker, (-BB), 1 Dec. 1989, Oliver 9352 (NBG);

Kogelberg Forest Reserve, near Rooiels, (-BD), 1 1 Nov. 1952, Rycroft

1354 (NBG). 3419 (Caledon): Villiersdorp, (-AB), 23 Nov. 1930, Nel

s.n. (NBG).

5. Thereianthus juncifolius (Baker) G.J. Lewis

in Journal of South African Botany 7; 37 (1941). Mor-

phixia juncifolia Baker: 238 (1876a). Watsonia jun-

cifolia (Baker) Baker: 178 (1892). Freesia juncifolia

(Baker) Klatt in T.A. Durand & Schinz 5: 188 (1894).

Lapeirousia juncifolia (Baker) N.E.Br.: 194 (1931).

Type: South Africa, [Western Cape], without precise

locality or date, [Piquetberg Div. fide Baker (1896)],

Zeyher 1619 (K [K320516 ex herb. Wallich ], lecto.!,

here designated; BOL!, K (3 sheets)!, S!, SAM! (2

sheets), isolecto.) [The Herb. Wallich duplicate is

selected as lectotype from among four duplicates at

Kew as having been available to Baker at the time and

as bearing the inscriptions ‘ Morph ixia juncifolia Baker’

and ‘Type’].

Anomatheca calamifolia Klatt: 355 (1882), nom. superfl. pro Mor-

phixia juncifolia Baker (1876b). Type: South Africa, [Western Cape],

Twenty Four Rivers, Zeyher [as Drege] 1619 (S, 'Herb. Klatt', holo.l;

BOL!, K (3 sheets)!, S!, SAM (2 sheets)!, iso.).

Ixia zeyheri Baker: 166 (1892), nom. superfl. pro Morphixia jun-

cifolia Baker (1876a) [based on the same type]. Type: South Africa,

[Western Cape], without precise locality or date, Zeyher 1619 (K

[K320516 ex herb. Wallich ], lecto.!, here designated; BOL!, K (3

sheets)!, S!, SAM (2 sheets)!, isolecto.).

Slender plants 1 20-200(-300) mm high. Corm glo-

bose, 5-10 mm diam.; tunics of fine-textured, netted

fibres, accumulating with age and forming thick neck up

to 60 mm long around base of stem. Stem flexed outward

above sheath of second leaf and then erect, unbranched

or rarely with short branch from axil of uppermost

scale-leaf, 0. 5-1.0 mm diam. below spike. Cataphylls

dry and papery, reddish brown, uppermost long and

reaching to lower leaf blade. Leaves (3)4, lowermost

inserted below ground, blade mostly reaching to spike

or exceeding it, filiform-terete, 150-500 x 0.5-1. 5 mm,

thick-textured, striate with slightly raised veins when

dry, upper leaves cauline with lower 1 or 2 inserted on

lower V3 of stem and mostly imbricate with lowermost

sheathing for < V2 length with blades (20-)40-100 mm

long, uppermost 1 or 2 leaves scale-like and entirely

sheathing, inserted ± midway between foliage leaves and

spike and well separated from spike. Spike erect, elon-

gate, laxly (2)3-1 0-flowered, lower internodes 3— 5(— 7)

x longer than bracts but progressively shorter acro-

petally and uppermost internodes ± as long as bracts;

bracts not imbricate, outer 3-5 mm long, acute to trun-

cate, soft-textured and green with papery brown tips,

inner slightly longer than outer and similar in texture,

acute or truncate. Flowers suberect, purple to lilac with

white throat; perianth actinomorphic, salver-shaped;

tube straight, 11-18 mm long, widening gradually and

evenly from base, 1.0-1. 5 mm diam. at mouth; tepals

spreading and slightly cupped, ovate or obovate, sub-

equal, 10-15 x 3. 5-6.0 mm. Stamens central; filaments

erect, 4-6 mm long, exserted 3-4 mm from tube; anthers

3-4 mm long, lilac or pale yellow; pollen lilac or cream-

coloured. Ovaty ovoid, ± 1 .5 mm long; style central,

dividing between base and middle of anthers, branches

± 3 mm long, divided for > V2 their length, spreading or

recurved. Capsules flask-shaped, 9-15 mm long. Seeds

fusiform, 6-7 mm long, with chalazal thread ± 13 mm

long, thus ± twice as long as seed body, sharply bent

back ± midway and hooked or crook-like, sometimes

twisting around adjacent seeds, reddish brown, longitu-

dinally rugulose. Flowering time : Dec.-Feb., rarely to

Mar. Figure 9.

Distribution and ecology, a mainly montane spe-

cies occurring from the southern Cedarberg through the

Olifants River Mtns to the Skurweberg north of Ceres,

with a disjunction to the Hottentots Holland, Kogelberg

and Klein River Mtns in the south, and also recorded on

the lower southern slopes of the Riviersonderend Mtns

(Figure 10). The species is thus apparently absent from

the mountains between Ceres and Stellenbosch. Plants

are restricted to seeps, streamsides, drainage lines and

marshy ground, and may be locally abundant, mostly

between 500 and 1 000 m but to near sea level along

coastal ranges. The leaves are still green and fresh at

flowering. As in most species in the genus, flowering is

stimulated by fire.

The long-tubed, salver-shaped, purple flowers sug-

gest pollination by long-proboscid flies, possibly Bom-

byliidae or Tabanidae, and butterflies but we have not

observed any visits.

Diagnosis and relationships', one of the most distinc-

tive members of the genus, Thereianthus juncifolius is

characterized by its slender habit, filiform-terete leaves,

0.5- 1.5 mm diam., and very lax spike of salver-shaped,

purple flowers with slender tube 11-18 mm long and

ovate-obovate tepals. The lower flowers are well sepa-

Bothalia 41,2 (201 1 )

253

FIGURE 9. — Thereianthus juncifo-

lius. Manning 3329. A, whole

plant; B, outer bract; C, inner

bract; D, capsules; E, seed.

Scale bar: A, 10 mm; B-D, 2

mm; E, 0.5 mm. Artist: John

Manning.

254

Bothalia 41,2 (2011)

FIGURE 10. — Known distribution of Thereianthus juncifolius, •.

rated from one another by internodes three to five times

longer than the bracts. The species is also distinctive in

its bracts, the inner as long as or slightly longer than

and similar in texture to the outer, both soft-textured

and green throughout except for the brown, papery tips.

In all other species the inner bracts are shorter than the

outer and membranous or papery except along the veins.

The capsules are markedly flask-shaped with a narrow,

tapering neck and fusiform seeds with a long, filiform

chalazal extension ± twice as long as the seed body. The

placenta is restricted to the lower portion of the capsule

and the tapered neck or beak accommodates the chalazal

extensions of the seeds, which extend up the neck of the

capsule to the apex and then flex sharply back towards

the seed body. This combination of unusual characteris-

tics sets the species apart from others in the genus but

the flask-shaped capsules and the seeds with long, fili-

form chalazal extensions indicate a close alliance with T.

minutus.

History, the unusual morphology of the species

caused a great deal of uncertainty about its generic posi-

tion among early workers. The lax spike and salver-

shaped flowers determined its initial description in the

genus Morphixia (Baker 1876a) but the subsequent

realization of the taxonomic importance of divided style

branches encouraged various authors to place it in all

possible genera with this character, namely Watsonia

(Baker 1892), Freesia (Klatt 1894) and Lapeirousia

(Brown 1931). Although Baker (1892) initially misun-

derstood the affinities of Thereianthus juncifolius , he

rapidly appreciated its relationship to other species in

the genus by including it with them in subgen. Beilia

of Watsonia. His simultaneous redescription of the spe-

cies as Ixia zeyheri , based on the same collection, is best

viewed as an aberration since none of the herbarium

material seen by him bears this name. Klatt’s (1882)

independent description of the species in the genus Ano-

matheca Ker Gawk, again based on the same collection

by Zeyher, anticipates his later inclusion of that genus in

Freesia.

Additional specimens examined

WESTERN CAPE.— 3219 (Wuppertal): S Cedarberg, Sneeuberg,

(-AC), I Mar. 1947, Esterhuysen 13842 (BOL); Algeria Forest Station,

Sneeuberg, 4200-4800' [1 280-1 460 m], (-AC), 5 Feb. 1982, Viviers

121 (NBG); Apollo Mtn., Kromrivier Valley, 1 340 m, (-AC), 24 Jan.

1986, Taylor 11482 (NBG); Elandskloof, Hexberg slopes, marshy site,

(-CA), 12 Dec. 1982, Esterhuysen 35847 A (MO); Keerom, E foot of

Twenty Four River Mtns [Olifantsrivierberge], (-CC), 3 Dec. 1950,

Esterhuysen 17863 (BOL, NBG); Grootfontein, next to Ratelrivier, (-

CC), 12 Dec. 1982, Van Zyl 2418 (NBG); Koue Bokkeveld, Skoonge-

sig, Fonteinbult, (-CC), 29 Dec. 1978, Hanekom 25 73 (MO, PRE);

Grootwinterhoek Forest Station, rocky flats, along stream, (-CC), 27

Dec. 1995, Goldblatt 10455 (MO); 3319 (Worcester): mtns near Por-

terville, (-AA), Nov. 1912, Edwards s.n. BOL16150 (BOL); Groot

Winterhoek Forestry Station, 650 m, (-AA), 2 Dec. 1983, De Kock 52

(NBG); Groot Winterhoek, road to De Tronk at Groot Kliphuis river

crossing, (-AA), 10 Dec. 2007, Helme 5091 (NBG); eastern slopes of

Witsenbergen, (-AA), Dec. 1919, Andreae 178 (NBG); Skurfdeber-

gen near Wagensboom River, (-AB), 27 Jan. 1897, Schlechter 10169

(BOL); Schurfteberg Pass, (-AB), 20 Dec. 1944, Compton 16799

(NBG). ‘Worcester, beim Waterfall,’ (-AC), Dec. Ecklon & Zeyher

hid. 297 (1.12) (MO). 3418 (Simonstown): margins of streams flow-

ing into Buffels River near Pringle Bay, (-BD), 27 Jan. 1936, Pillans

8185 (BOL); Betty's Bay, (-BD), 10 Feb. 1956, Levyns 10451 (BOL);

marsh on mtns behind Rooiels-Stalberg-Pringle E Peak, (-BD), 10

Jan. 1971, Esterhuysen 32561 (BOL, MO); Kogelberg, (-BD), 16

Jan. 1944, Esterhuysen 10010 (BOL); mtns north of Palmiet River

Mouth, (-BD), 26 Jan. 1947, Esterhuysen 13667 (BOL, NBG); mtns

above Betty’s Bay, marshy slopes in valley at base of Platberg, (-BD)

27 Jan. 1980, Esterhuysen 35387 (BOL, MO); Kogelberg State Forest,

Oudebosch, 80 m, (-BD), 20 Jan. 1992, Kruger 302 (MO, NBG). 3419

(Caledon): Hottentots Holland Mtns, east of Landdroskop Hut, (-AA),

6 Jan. 2000, Oliver & Oliver 11470 (NBG); Grabouw, (-AA), Jan.

1943, Stokoe s.n. SAM55716 (SAM); Nuweberg Forest Reserve, slopes

of Landdrost kop, damp gully above path to Somerset Sneeukop, (-

AA), 11 Jan. 1987 Goldblatt 8351 (MO); Aries Kraal, (-AA), 1 Jan.

1945, Compton 16851 (NBG); Lebanon, (-AA), Jan. 1968, Kruger

640 (NBG); Highlands Forest Reserve, hiking trail, marshy places, (—

AC), 29 Jan. 1987 Goldblatt 8469 (MO); summit of Klein River Mtns,

Bellevue, (-AD), 2 Feb. 2008, Helme 5322 (NBG); Vogelgat, N of

Mt Frustration, (-AD), 1 Mar. 1987, Williams 3785 (NBG); Vogelgat,

Rock-in-the-Path, (-AD), 1 Feb. 1986, Williams 3643 (NBG); Her-

manus, Femkloof Nature Reserve, 400’ [122 m] (-AD), 21 Jan. 1981,

Orchard 583 (MO); Greyton, foot of Riviersonderend Mtns, (-CA), 16

Jan. 1988, Oakes s.n. (NBG).

Sect. Thereianthus

Leaves linear or falcate, plane and either leathery

without raised veins or strongly ribbed or corrugate with

prominently raised veins, or ellipsoid in section; bracts

(7— )8— 1 5 mm long, closely and prominently veined with

raised veins (sclerenchymatous phloem caps prominent).

Pollen operculum 1 -banded, exine perforate. Seeds

(where known) ellipsoid-ovoid and angled, without

chalazal extension, brown to blackish.

Ser. Bracteolatus J.C. Manning & Goldblatt , ser.

nov.

Lamina foliorum non costata nec corrugata sine

nervis prominentibus vel incrassatis, floribus actinomor-

phis tepalis patentibus et tubo curvato, staminibus ± dec-

linatis.

Type species: Thereianthus bracteolatus (Lam.)

G.J. Lewis.

6. Thereianthus bulbiferus Goldblatt &

J.C. Manning, sp. nov.

Plantae 140-200 mm altae, cormo globoso 10-15 mm

diam., tunicis fibrosis crassis interdum infra unguiscen-

tibus, caule usitate eramoso cormelos 5-7 in axilla folii

infimi facienti, foliis 4, infimo lineari-falcato, 70-120

x ( 1 .5— )2— 3 mm, coriaceis, spica erecta dense 7- ad

1 5-flora, bracteis breviter imbricatis, bractea externa

7— 9(— 1 0) mm longa interna quam Ji ad 3A plo breviore,

floribus coeruleis tepalis inferioribus prope basin nota

alba mediana munitis, perianthio actinomorpho, tubo

Bothalia 41,2 (2011)

255

7-9 mm longo infundibuliformi, tepalis subaequalibus

vel interioribus pauce latioribus oblanceolatis patentibus

parum cupulatis (10-) 13-20 x 3-6 mm, staminibus ±

declinatis, filamentis 6-8 mm longis ± 5-6 mm ex tubo

exsertis, antheris 4 mm longis purpureis, ramis styli pro-

funde furcatis 3-5 mm longis.

TYPE. — Western Cape, 3319 (Worcester): Bo-Her-

mon, Elandsberg Nature Reserve, alluvial flats near

river, after fire, (-AC), 7 Nov. 2010, J.C. Manning 3312

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

Plants 140-200 mm high. Corm globose, 10-15 mm

diam.; tunics of coarsely netted fibres, sometimes form-

ing claws below, accumulating with age and forming

short neck around base of stem. Stem erect, flexuous or

flexed outward only above sheath of second leaf, usu-

ally unbranched, rarely with short branch from upper

leaf axil, 2. 0-2. 5 mm diam. below spike, developing

conspicuous cluster of 5-7 cormels, each 3^1 mm diam.,

in axil of lowest leaf. Cataphylls dry and papery, red-

dish brown. Leaves 4, lowermost inserted below ground,

blade reaching base of spike or not, ± dry at flower-

ing, linear-falcate, often sinuous-twisted, 70-120 x

( 1 .5— )2— 3 mm, leathery, with 3 ± equally distinct veins

or with main vein slightly more distinct, not thickened

or raised when fresh but prominent when dry, upper 3

leaves cauline with lower 2 inserted on lower V3 of stem

and sheathing for ± V2 length, sheaths shortly imbricate,

blades 50-70 x 3^) mm, uppermost leaf a little distant

from lower leaves and well separated from spike, usu-

ally with short blade 20-50 mm long, rarely almost

entirely sheathing. Spike erect, densely 7-15-flowered;

bracts imbricate for ± V2 their length, outer 7— 9(— 1 0)

mm long and up to two intemodes long, obtuse or trun-

cate, either green below or entirely brown and leathery,

conspicuously and closely veined, with narrow membra-

nous margins, inner bract V2 to 3A as long as outer, bifid,

± membranous with narrow brown zones along veins.

Flowers patent, pale to mid-blue, lower three tepals

with white median mark near base; perianth actinomor-

phic; tube slightly curved, 7-9 mm long, funnel-shaped,

exserted 2-5 mm beyond bracts; tepals oblanceolate,

subequal or inner slightly broader, spreading and slightly

cupped, ( 1 0 — ) 1 3 — 20 x 3-6 mm. Stamens ± declinate;

filaments 6-8 mm long, exserted ± 5-6 mm from tube;

anthers all facing upwards or median (lower) facing

downwards or outwards, 4 mm long, purple; pollen lilac.

Ovary obovoid, 2. 0-2. 5 mm long; style ± deflexed to lie

beneath stamens, dividing at middle of anthers, branches

3-5 mm long, divided to near base. Capsules ovoid, 6 x

3 mm long, slightly shorter to slightly longer than bract.

Seeds subglobose-angled, 1. 5-2.0 mm long, chalaza ±

excavated, yellowish brown, obscurely scalariform-rug-

ulose. Flowering time : Nov. (early Dec.). Figure 1 1.

Distribution and ecology n a local endemic between

Piketberg and Wellington, Thereianthus bulbiferus is

most common along the western foot of the Elandskloof

Mtns near Hermon, where it is known from several pop-

ulations on Elandsberg Nature Reserve (Figure 12). The

southernmost population near Agter Paarl, known only

from a single specimen collected over forty years ago. is

now likely extinct through agriculture. Plants at Elands-

berg Nature Reserve grow on seasonally moist or wet

alluvial flats, primarily in Swartland Alluvial Fynbos

(Mucina & Rutherford 2006) communities but extending

into adjacent Swartland Alluvial Renosterveld. Flower-

ing is most prolific after fire but occasional individuals

will flower in open or cleared spaces in the intervening

years. The species co-occurs throughout its range with

T. spicatus , which favours drier soils. At Elandsberg,

T. bulbiferus occurs as single-species communities on

the lower, moister flats but on the slopes, where it is

restricted to drainage lines, it interdigitates with T. spi-

catus growing on adjacent drier sites. The single collec-

tion from near Agter Paarl ( Thompson 622) is a mixed

gathering comprising a single stem of T. bulbiferus and

two stems of T. spicatus , suggesting that the two species

grew in close proximity there too, and both species have

also been collected on the Piketberg, although in differ-

ent localities.

Visits by monkey beetles (Coleoptera: Hopliini) have

been observed at Elandsberg Nature Reserve (unpub-

lished obs.) but the relatively short-tubed flowers pro-

duce nectar and are probably also visited by antho-

phorine bees.

Diagnosis and relationships'. Thereianthus bulb-

iferus is diagnosed by its leathery leaves without raised

central and secondary veins when fresh, the lower leaf

mostly characteristically falcate or arcuate and often

sinuous-twisted, and by the moderately dense spikes of

blue flowers with relatively short, funnel-shaped peri-

anth tube, 7-9 mm long exserted up to 5 mm beyond the

bracts, actinomorphic perianth with white markings at

the base of all the tepals, and ± declinate stamens and

style. Flowers with declinate stamens are found else-

where in sect. Thereianthus only in T. bracteolatus , and

in both species the perianth tube is curved so that the

flowers face sideways rather than upwards. Both spe-

cies also have inconspicuously veined leaves. The seeds,

with an unusual chalazal depression or small excavation

rather than crest, are distinctive for T. bulbiferus.

Thereianthus bulbiferus is unique in the genus in pro-

ducing a cluster of cormels from the axil of the lower-

most leaf. These cormels, 3-4 mm diam., are present

at flowering time, and although they may be lost from

some pressed specimens, their presence can be read-

ily inferred by the split in the leaf sheath through which

they protruded. Florally, the species is distinguished

from T. bracteolatus by the light to mid-blue perianth

with white markings on all tepals, which are incon-

spicuously veined, the outer longitudinal veins ± evenly

branched along their entire length. The flowers of T.

bracteolatus , in contrast, are dark blue to purple without

evident white markings but with the longitudinal veins

splashed or spotted with purple. The outer longitudinal

veins in this species are typically only weakly branched

in the distal half so that the tepals appear to be parallel-

veined from the base.

All other members of sect. Thereianthus have

strongly and closely ribbed leaves, and mostly arcuate

stamens (except T. ixioides , which has symmetrical sta-

mens). Fresh leaves of the members of the T. spicatus

alliance have the veins separated from one another but

as the leaves dry and shrink, the veins become closely

appressed, leaving no gaps between them. The leaves of

T. bracteolatus and T. elandsmontanus, in contrast, are ±

256

Bothalia 41,2 (2011)

FIGURE 1 1 . — Thereianlhus bulb-

iferus, Manning 3312. A,

whole plant; B, flower, front

view; C, half flower; D, outer

(left) and inner (right) bract;

E, details of stamens and

stigma; F, capsule; G, seeds.

Scale bar: A-C, 10 mm; D, E,

3 mm; F, 2 mm; G, 0.5 mm.

Artist: John Manning.

smooth and leathery when fresh and although the veins

become evident as the leaf dries, they remain separated

from one another by intervening blade tissue and by

lower order veins.

History, although collected on two separate occa-

sions several decades ago, Thereianthus bulbiferus was

confused with both T. bracteolatus and T. spicatus,

and it was only in 2010 when we were able to properly

Bothalia 41,2 (2011)

257

16° 18" 20° 22° 24° 26° 28°

FIGURE 12. — Known distribution of Thereianthus bracteolatus, •; T.

bulbiferus, O.

examine living plants in the field at Elandsberg Nature

Reserve that it became clear that they represent a distinct

species.

Etymology, the specific epithet alludes to the charac-

teristic cormels on the lower part of the stem.

Additional specimens examined

WESTERN CAPE.— 3218 (Clanwilliam): Piketberg, Sandleegte

[Buglerspost, at top of Versveld Pass], 2400' [730 m], (-DC), 3 Dec.

1973, Linder 130 (BOL). 3318 (Cape Town): Pause’s plot on Protea

F.U.C. road off Agter-Paarl road, (-CB), 2 Dec. 1968, Thompson 622A

(NBG). 3319 (Worcester): Bo-Hermon, Elandsberg Nature Reserve,

Vervlak, sandstone flats burned in Mar., (-AC), Nov. 1999, Manning

s.n. (NBG); 7 Nov. 1999, Parker 390 (NBG).

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

in Journal of South African Botany 7: 42 (1941). Gladi-

olus bracteolatus Lam.: 725 (1788). Type: South Africa,

without precise locality, said to have been collected by

Sonnerat s.n. (P: Herb. Lamarck, holo. ! ).

Ixia punctata Andrews: t. 177 (1801). Watsonia punctata

(Andrews) Ker Gawl.: 229 (1804). Gladiolus punctatus (Andrews)

Roem. & Schult.: 425 (1817). Type: South Africa. 'Cape of Good

Hope’, said to have been collected and sent by G. Hibbert, illustration

in Andrews, The botanist's repository 3: t. 177 (1801).

Gladiolus subulatus Vahl: 97 (1805), nom. illegit. superfl. pro G.

bracteolatus Lam. (1788). Watsonia subulata Klatt, nom. nov. pro G.

subulatus Vahl, nom. illegit. et superfl. pro G. bractolatus Lam. Type:

South Africa, without precise locality or date, Bulow s.n. C1000844

(C, holo.!).

Gladiolus triticeus Thunb.: 194 (1807). Watsonia triticea (Thunb.)

Spreng.: 150 (1824). Beilia triticea (Thunb.) Eckl.: 43 (1827).

Micranthus triticeus (Thunb.) Heynh.: 405 (1847). Watsonia punc-

tata var. triticea (Thunb.) Baker: 177 (1892). Beilia spicata var. triti-

cea (Thunb.) Kuntze: 305 (1898). Type: South Africa, without precise

locality, Thunbergs.n. UPS-THUNB1084 (UPS-THUNB, holo.!).

Plants (80-)150-200(-300) mm high. Corm globose,

10-15 mm diam., developing several large cormels at

base; tunics of coarsely netted fibres, thickened and

forming woody claws below, accumulating with age and

forming collar up to 20 mm long around base of stem.

Stem erect, flexuous or flexed outward only above sheath

of second leaf, unbranched, ( 1 .5— )2.0— 2.5 mm diam.

below spike. Cataphylls dry and papery, reddish brown.

Leaves 4(5), lowermost inserted below ground, blade

reaching base of spike or not, ± dry at flowering, lin-

ear, elliptic or terete in section, 70-170 * 0.5—1 ,5(— 2.0)

mm, leathery, with 3 ± equally distinct veins or with

main vein slightly more distinct, not thickened or raised

when fresh but prominent when dry, upper 3(4) leaves

cauline with lower 2 inserted on lower 'A of stem and

sheathing for ± 'A length, sheaths usually shortly imbri-

cate, blades 50-70 mm long, uppermost leaf a little dis-

tant from lower leaves and well separated from spike,

entirely sheathing or with short blade 20-50 mm long.

Spike erect, densely 7-15(-30)-flowered; bracts imbri-

cate for ± 'A their length, 8—1 0( — 1 2) mm long and ±

two internodes long, outer entirely brown and leathery,

obtuse or truncate (rarely apiculate), conspicuously and

closely veined, with narrow membranous margins, inner

'A to ’A as long as outer, bifid, ± membranous with nar-

row brown zones along veins. Flowers ± patent, purple

to violet with dark blotches and streaks on longitudinal

veins and especially midline; perianth actinomorphic;

tube slightly curved, (7— )8— 1 0(— 1 5 ) mm long, funnel-

shaped. usually exserted 2 — 6( — 1 0) mm beyond bracts,

rarely just included; tepals oblanceolate, subequal or

inner slightly broader, spreading and slightly cupped.

( 10 — )1 1—17 x 3-6 mm. Stamens unilateral, ± declinate,

sometimes weakly so and then ± central; filaments 8-10

mm long, exserted ± 6-8 mm from tube, usually flexed

sharply upwards apically; anthers all facing upwards,

usually white (rarely purple), 4-6 mm long; pollen white

(rarely brown when anthers purple). Ovary’ obovoid,

2. 0-2. 5 mm long; style ± deflexed to lie beneath sta-

mens, dividing opposite middle of anthers, branches 3-5

mm long, divided to near base. Capsules ovoid, ± 6 mm

long. Seeds subglobose- or ellipsoid-angled, ± 2 mm

long, chalaza truncate, brown, testa colliculate. Flower-

ing time : late Nov. -early Jan. figure 13A-G.

Distribution and ecology >\ the only species of There-

ianthus on the Cape Peninsula, T. bracteolatus occurs

from the Peninsula and Stellenbosch along the coastal

ranges of the Hottentots Holland, Kogelberg and Klein

River/Babilonstoring Mtns to Bredasdorp in the east,

with isolated records from near Caledon (Drayton Sid-

ing) and the lower slopes of the Riviersonderend Mtns

above Greyton (figure 12). Somewhat surprisingly, the

species has not been recorded from the Caledon Swart-

berg. Although there is an early record, Zeyher 1608

(SAM), from the Twenty four Rivers Mtns near Porter-

ville. the species has never been re-collected here and as

this collection is a mixed one that includes a stem of T.

longicollis, which is known from this location, the asso-

ciation of T. bracteolatus with the locality is probably an

error.

The species is locally common after fire on sandy

flats and lower slopes below 1 000 m, although plants

will also flower in cleared places along roads in the

absence of fire. The distribution of Thereianthus bracte-

olatus largely overlaps with that of T. spicatus, with the

notable exception of the Cape Peninsula where only T.

bracteolatus is found, but T. spicatus favours more stony

sites and often heavier soils. We have encountered both

species growing adjacent to each other at Kogelberg,

with T. bracteolatus on damper soils and flowering in

December and T. spicatus immediately adjacent on

rocky slopes and flowering in January.

The purple or violet flowers with a relatively short

perianth tube are evidently adapted to pollination by var-

ious nectar-feeding bees.

258

Bothalia 41,2 (2011)

FIGURE 13. — Thereianthus bracteolalus. A-E, Goldblatt 9032: A, whole plant; B, flower, side view; C, outer tepal; D, inner tepal; E, details of

stamens and stigma. F, G, Rodgers 2829: F, capsule; G, seeds. T. spicatus. H— J, Goldblatt 6837: H, whole plant; I, flower, front view; J,

flower, side view. K, L, Goldblatt & Manning 9567: K, capsules; L, seed. Scale bars: A, B, H-J, 10 mm; C, D, F, K, 2 mm; E, 3 mm; G, L,

0.5 mm. Artist: John Manning.

Diagnosis and relationships'. Thereianthus bracteo-

latus is unmistakeable when seen alive, with very nar-

row leaves, 0.5—1 .5(— 2) mm diam., and terete or elliptic

in section, and dense spikes of sideways-facing, deep

purple or violet flowers with curved perianth tube (7— )8—

1 0(— 1 5 ) mm long, and relatively long, unilateral, dec-

imate stamens. The flowers, which lack nectar guides,

have darkly pigmented veins, giving them a character-

istic blotched-striate appearance. Although Lewis (1941)

noted that the branching pattern of the veins was dis-

tinctive for the species, this difference is not evident in

all plants. Like other species in the genus, each tepal

has three primary longitudinal veins but unlike other

species, the lateral veins are often only branched in

the basal half so that the tepals appear to be ± parallel-

veined for most of their length. In the other species the

veins are not darkly pigmented and thus not evident,

and the lateral longitudinal veins are ± evenly branched

throughout their length.

Perianth tube length is somewhat variable, usually

8-10 mm but 10-15 mm long in an isolated population

at Drayton Siding east of Caledon ( Goldblatt 394). The

anthers and pollen are usually white but plants from

Bothalia 4 1 .2 (2011)

259

the Silvermine area of the Cape Peninsula have purple

anthers with brown pollen. Another isolated popula-

tion, from the lower slopes of the Riviersonderend Mtns

above Greyton ( Esterhuysen 20763), has stamens that

are less obviously declinate than usual but matches the

species in vegetative and other floral characters, includ-

ing the distinctive tepal venation. This population may

repay further consideration.

The smooth leaves and especially the declinate sta-

mens (but not the characteristic tepal venation) are both

shared with Thereianthus bulbiferas from gravelly flats

between Malmesbury and Paarl. This species has pale

to mid-blue flowers with distinct white blotches at the

base of each tepal, and shorter filaments, 6-8 mm long.

Most characteristically, the plants develop a cluster of

cormels in the axil of the lower leaf, whereas cormels

in T. bracteolatus are developed at the base of the par-

ent corm. Although T. bracteolatus is sometimes con-

fused with T. spicatus, especially in the dried state when

details of the perianth coloration and stamen orientation

are obscured, the two are readily distinguished by the

difference in leaf texture: leathery and terete or elliptic

in section without raised veins in T. bracteolatus versus

plane and ribbed or corrugate with raised veins in T. spi-

catus. Living plants of T. spicatus are readily separated

by their blue (rarely white or pink) flowers with white

markings on the lower three tepals outlined with a pur-

ple chevron and lacking darker venation, the ± straight

or distally curved perianth tube, 12-25 mm long, and the

much shorter, arcuate filaments, 3^4 mm long.

History, the second species of Thereianthus to be

described after T. spicatus, T. bracteolatus was named in

the genus Gladiolus by French biologist J.P.B. Lamarck

(1788) from specimens said to have been collected by

naturalist and draughtsman Pierre Sonnerat, who could

only have collected this material in 1773, on the out-

going leg of his voyage to Isle de France (now Mauri-

tius). This name was overlooked by the English botani-

cal artist and engraver, Henry Andrews (1801), who

described Ixia punctata from plants cultivated in Lon-

don that had been sent to collector and botanical patron

G. Hibbert from the Cape of Good Hope. In continen-

tal Europe, the species was described under the name

Gladiolus triticeus for a third time a few years later by

Thunberg (1807), based on his own collections from

the Cape. At the time, Thunberg expressed doubt as to

whether his species was truly different from G. spica-

tus L. (i.e. T. spicatus) and must have appreciated the

similarity between these species, ultimately to be treated

in a separate genus. J.G. Baker (1892) subsequently

treated Thunberg’s name as a variety of Andrew’s spe-

cies, which had by now been transferred to Watsonia as

W. punctata (Andrews) Ker Gawl. The nomenclature

was clarified by Lewis (1941), who correctly identified

Lamarck’s name as the earliest for the species, and also

established the genus Thereianthus for plants until then

treated as subgen. Beilia of Watsonia.

Additional specimens examined

WESTERN CAPE. — 3318 (Cape Town): Table Mountain, (-CD),

Jan. 1898, Thode 6017 (NBG), Dec. 1950, Pillars 10276 (MO);

Cape Peninsula, Camps Bay, (-CD), Dec. without year, Zeyher 5038

(SAM); 5 July 1943 [fr.]. Barker 2416 (NBG); above S end of Kirst-

enbosch, (-CD), 28 Nov. 1953, Esterhuysen 22404 (BOL); Kenil-

worth, (-CD), Dec. without year, Bolus 3305 (NBG); Kenilworth Race

Course, (-CD), 11 Dec. 1969, Esterhuysen 32359 (BOL, MO, NBG);

near Stellenbosch, (-DD), Dec. 1924, Rogers 28929 (BOL, NBG).

3319 (Worcester): Franschhoek Pass, east side, (-CC), 26 Nov. 1939,

Compton 8185 (NBG). 3418 (Simonstown): Vlaggenberg [Vlakken-

berg], (-AB), 1 Jan. 1896, Wolley Dod 345 (BOL); Bergvliet Farm. (—

AB), 16 Dec. 1976, Purcell 152 (SAM); Constantia Nek, (-AB), 24

Jan. 1929, Gillett 3365 (BOL, NBG); Steenberg Plateau, (-AB), Dec.

1944, Lewis 56067 (SAM); Silvermine Valley, (-AB), Dec. 1944,

Lewis 56066 (SAM); mntns above Muizenberg, (-AB), Jan. 1881,

Bolus 3305 (BOL, NBG); Red Hill, (-AB), 21 Dec. 1963, Taylor

5616 (NBG); Farm Buffelsfontein W of Redhill, (-AB), 17 Dec. 1984,

Snijman 833 (NBG); Patreis Vlei [Patrys Vlei], (-AD), 3 Jan. 1941,

Salter 8586 (BOL, NBG); Cape Point Reserve, sandy plain on Circu-

lar Drive, (-AD), 31 Dec. 1989, Goldblatt 9032 (MO); Cape Point, (-

AD), 1 Jan. 1968, Taylor 726 7 (NBG); Steenbras, (-BB), Dec. 1945,

Lewis 58735 (SAM); Palmiet River Valley, (-BD), Nov. 1944, Lewis

56408 (SAM); between Kogel Baai and Rooi Els, (-BD), Nov. 1945,

Lewis 58736 (SAM); Kogelberg State Forest, Palmiet River Valley, (—

BD), 29 Nov. 1991, Kruger 97 (NBG); Palmiet River Mouth, (-BD),

7 Dec. 1962, Grobler 71 (NBG). 3419 (Caledon): Aries Kraal, (-AA),

18 Nov. 1944, Leighton 808 (BOL); 18 Nov. 1944, Leighton 3361

(NBG); Lebanon Forest Reserve, (-AA), 14 Jan. 1971, Haynes 510

(NBG); Grabouw near Palmiet River, (-AA), Dec. 1899, Bolus 3305

(BOL); Bot River, (-AC), 9 Dec. 1951, Taylor 3810 (NBG); Honing-

klip. (-AC), without date, Taylor 5119 (NBG); Hermanus Mtn, (-AD),

26 Dec. 1943, Leighton 357 (BOL); Hermanus, Rotary Way, (-AC), 8

Jan. 1975, Walters 1452 ( NBG); Hermanus, Vogelgat Nature Reserve,

(-AD). 7 Jan. 1984, Burman 1273 (BOL); Femkloof Nature Reserve,

500 m, (-AD), 2 Jan. 1976, Orchard 359 (MO, NBG); top of Shaw’s

Pass, (-AD), Jan. 1957, Lewis 63514 (BOL); foothills of River Sonder

End [Riviersonderend] Mtns, (-BA), Apr.-May 1950 [fr.], Lewis 2280

(SAM); Zonder End [Riviersonderend], (-BA), without date, Barnard

448 (SAM); Riviersonderend Mtns near Greyton, (-BA), 30 Nov.

1952, Esterhuysen 20763 (BOL, MO, NBG); Caledon, Drayton Sid-

ing, (-BA), 16 Dec. 1968, Goldblatt (BOL); north end of Akkedisberg

Pass, fynbos slopes, burned last summer, (-BC), 19 Nov. 1979, Gold-

blatt 5186 (MO); Elandskloof, (-BD), 17 Dec. 1896, Schlechter 9765

(BOL, MO); between Gansbaai and Baardscheerdersbos, (-DA), 1 1

Nov. 1980, Snijman 380 (NBG); between Wolvengat and Elim, Farm

239, (-DA), 25 Nov. 2000, Mucina 251100/5 (NBG); Bredasdorp,

plateau on mountain top, (-DB), 19 Dec. 1930, Galpin 22442 (BOL).

Doubtful locality : 3319 (Worcester): Twenty Four Rivers, (-AA), with-

out date, Zeyher 1608 [excluding single specimen of T. longicollis as

indicated] (SAM).

Ser. Thereianthus

Leaves ribbed or corrugate, with thickened and raised

veins. Flowers suberect with ± straight tube; perianth

actinomorphic or more usually zygomorphic. Stamens

arcuate, rarely central.

8. Thereianthus spicatus (L.) G.J. Lewis in Jour-

nal of South African Botany 7: 39 (1941). Gladiolus

spicatus L.: 37 (1753). Ixia spicata (L.) Willd.: 200

(1797), horn, illegit. non Burm.f. (1768). Gladiolus

alopecuroides Pers.: 46 (1805), nom. illegit. pro Gladi-

olus spicatus L. et horn, illegit. non L. (1756). Micran-

thus spicatus (L.) Heynh.: 405 (1847). Paulomagnu-

sia spicata (L.) Kuntze: 702 (1891). Beilia spicata (L.)

ex Kuntze: 305 (1898) [Beilia spicata (L.) Eckl. nom.

inval.]. Type: from South Africa without any details ex

Herb, van Royen (?L, not found). Neotype, designated

here: South Africa, [Western Cape], Jonkershoek Val-

ley, contour path below Berg River Neck, 29 Dec. 1989,

Goldblatt & Manning 9028 (NBG. neo.; K, MO, PRE,

iso.).

Thereianthus spicatus var. linearifolius G.J. Lewis: 40 (1941).

Type: South Africa, [Western Cape], mountains near Franschhoek,

Salter 2973 (BOL, holo.!).

Gladiolus rubens Vahl: 98 (1805). Watsonia rubens (Vahl) Ker

Gawl.: sub t. 1072 (1807). Type: South Africa, not cited, probably at C.

260

Bothalia41,2 (2011)

Plants (150-)200-500(-600) mm high. Conn glo-

bose, 10-20 mm diam.; tunics of coarsely netted fibres,

thickened and claw-like at base, accumulating with age

and forming neck up to 20 mm long around base of

stem. Stem erect or flexed outward above sheath of sec-

ond leaf, unbranched or rarely with short branch from

axil of uppermost leaf, 1. 5-3.0 mm diam. below spike.

Cataphylls dry and papery, reddish brown. Leaves

4(5), lowermost inserted below ground, blade reach-

ing or exceeding spike (rarely shorter than spike), ±

dry at flowering, linear, ( 100-)200-250 x 1— 2(— 4) mm,

closely 3-5(-7)-ribbed, glabrous or minutely scabridu-

lous along ribs and margins, upper 3 leaves cauline,

second and third leaves usually ± imbricate and envel-

oping stem but uppermost leaf distant, blades succes-

sively shorter, uppermost leaf only rarely reaching base

of spike, entirely sheathing or with short blade 20-60

mm long. Spike erect, densely (3-)6-20(-30)-flowered;

bracts imbricate for V: to 2A length, (7— )8— 1 3 (— 15) mm

long and two or three internodes long, outer acute, either

green below or entirely brown and leathery, conspicu-

ously and closely veined, with narrow membranous mar-

gins, inner slightly shorter, bifid, ± membranous with

narrow brown zones along veins. Flowers suberect,

mostly pale to mid-blue, rarely white or pink, lower

three tepals each with white marking near base outlined

distally with purple chevron; perianth subactinomor-

phic or weakly zygomorphic; tube ± straight or slightly

curved outwards in distal 3 mm, ( 1 2—) 1 5— 1 8(— 25 ) mm

long, 1.0-1. 5 mm diam. at mouth, ± dilated gradually

and evenly from base to tip, exserted 6-15 mm beyond

bracts, 0.8-1. 8 x as long as tepals; tepals elliptical-

oblanceolate, inner three slightly smaller, spreading and

slightly cupped, 10-18 x 3-5 mm. Stamens arcuate; fila-

ments 3—4 mm long, exserted 2 mm from tube; anthers

4-5 mm long, purple; pollen purple. Ovary obovoid,

2. 0-2. 5 mm long; style arching above anthers, dividing

at ± middle of anthers, branches 4-5 mm long, divided

to near base. Capsules narrowly ovoid, 6-7 x 3 mm.

Seeds ellipsoid-angled, ± 2 mm long, black or brown,

testa striate-colliculate. Flowering time : (Nov.) Dec.-

Jan., rarely into early Feb. Figure 13H-L.

Ecology and distribution', the most common species

in the genus, Thereianthus spicatus is distributed along

the coastal mountains and foothills in the southwestern

part of Western Cape, from Piketberg and Porterville

southwards to the Palmiet River Mouth and onto the

lower slopes of the Riviersonderend Mtns but is absent

from the Cape Peninsula (Figure 14). The species is

mostly found on sandstone slopes but also grows on

shale bands and on granite, from near sea level to over

1 000 m. Flowering is strongly stimulated by fire but

plants will bloom in older, open veld and in fire-breaks.

Flower colour is somewhat variable in the species,

usually mid- to deep blue but sometimes white or pink.

Although the different forms are often mixed, there is an

altitudinal component to perianth colour in populations

in the Jonkershoek Valley, where the proportion of white

flowers increases with elevation. The relatively great

variation in perianth tube length suggests adaptation

to the local pollinator fauna. Mostly 15-20 mm long,

the perianth may be as long as 25 mm in plants from

Grootwinterhoek ( Helme 5141 ) and in Idas Valley, Stel-

lenbosch (Levyns 1458), and as short as 12 mm around

Somerset West ( Martley s.n. BOL22440, Schlechter

7245). Leaf width in the species is mostly 1-2 mm but

plants with broader leaves, 3—4 mm wide, are known

from Bainskloof, Stellenbosch, Kuils River and Somer-

set West. A collection of unusually delicate plants, with

small tepals 8 mm long (Esterhuvsen 22748) from Bai-

ley’s Peak matches the species in other respects, notably

the ribbed leaves, and perianth tube 1 5-20 mm long, and

we regard it as a local, montane form.

The moderately long-tubed flowers are evidently

adapted to pollination by correspondingly long-tongued

insects, and we have seen the species being visited by

the butterfly Cynthia cardui (Nymphalidae) at Jonker-

shoek. The perianth tube is narrowed towards the base,

and nectar wells up for some distance into the tube,

where it is accessible to long-tongued anthophorine bees

as well, which probably also play a role in pollination.

Diagnosis and relationships', the central species in

series Thereianthus, T. spicatus is distinguished from the

other members of the genus with strongly ribbed leaves

(71 elandsmontanus, T. ixioides and T. longicollis) by

flowers with a moderately long perianth tube, ( 1 2 — ) 1 5 —

1 8( — 25) mm long, usually curved outwards slightly near

the tip and dilating gradually and evenly from base to

apex. The tepals are relatively large, elliptical-oblanceo-

late, 10-18 x 3-5 mm, and the stamens are arcuate, with

filaments 3-4 mm long. Although usually glabrous, the

leaves in some plants are scabridulous along the ribs and

margins. This is especially evident in the collection from

Piketberg but occurs sporadically elsewhere as well.

Perianth length is more variable in this species than in

others, and longer-tubed forms are sometimes misidenti-

fied as Thereianthus longicollis — both species have been

collected in the mountains above Tulbagh Waterfall and

above Saron. We have seen both in the field and there is

no doubt that they are distinct. Longer-tubed plants of T.

spicatus, such as those from the Grootwinterhoek Mtns

above Porterville with tubes reaching to 25 mm long, are

distinguished from T. longicollis by their proportionally

larger tepals, always less than twice as long as the tube.

The critical differences separating T. spicatus from the

closely allied T. ixioides and also from T. bracteolatus

(ser. Bracteolatus), with which it has been confused in

the past, are discussed under those species.

Bothalia 41,2 (2011)

261

History, the earliest known species in the genus,

Thereianthus spicatus, was first named in the genus

Gladiolus (Royen 1740; Linnaeus 1753) but subse-

quently transferred to various other genera, including

Ixia L., Watsonia , Micranthus and the invalidly pub-

lished Beilia (Ecklon 1827). This reflects the uncertainty

about generic circumscriptions that prevailed for many

years. The original collector and date of collection are

unknown but the species was evidently cultivated in

the botanical gardens at Leiden. The Netherlands, in the

1740s. Forms with narrower leaves were distinguished

by Lewis (1941) as var. linearifolius but this distinction

has no taxonomic merit (Manning & Goldblatt 2004),

with individual collections showing a range of leaf

widths from 1-3 mm (e.g. Esterhuysen 13673).

Nomenclatural note : the species was based by Lin-

naeus (1753) on the polynomial Gladiolus foliis lineari-

bus, cattle simplicissimo, fioribus spicatus (Royen 1740)

but no specimen under this inscription could be located

at Leiden (G. Thijsse pers. comm. Nov. 2010) and must

be presumed lost — types of the other three Gladiolus

species listed by Royen (1740) are all at Leiden. Pos-

sibly, therefore, no authentic material was available to

Linnaeus. A specimen with the binomial G. spicatus

in the Linnean Herbarium (LINN 59.3) and inferred to

have come from Traugott Gerber (fl. 1739-1741) by

Jackson (1912). was regarded by Jarvis (2007) as origi-

nal material. Critically, however, this specimen does

not match the protologue, having just a single sheathing

leaf, and it cannot therefore be considered to be origi-

nal material. Although it was identified as the holotype

by Manning & Goldblatt (2004), their designation is

not nomenclaturally effective in lacking a definitive

phrase designating it as a lectotype (Jarvis 2007; Art.

9.21 : McNeill el al. 2006). Significantly, the specimen is

not in fact T. spicatus as currently understood, being a

species of Gladiolus, evidently G. communis. The cur-

rent taxonomic association of the name can be traced to

Thunberg (1807) (UPS-THUNB 1 072) but the basis for

this remains unknown. We designate a neotype here to

fix its application in this sense.

Additional specimens examined

WESTERN CAPE. — 3218 (Clanwilliam): Piketberg, suurvlakte

[west of Avontuur near top of Versveld Pass], 780 m, (-DC), 23 Dec.

1973, Linder 187 (BOL). 3318 (Cape Town): Paarl Mtn, near top, (—

DB) , 2 Dec. 1929, Grant 5027 (BOL, MO); Penhill, Eersterivier, (—

DC) , 10 Dec. 1977, Raitt 288 (NBG); Kuils River, (-DC), Dec. 1931,

Basson s.n. BOL2430 (BOL); Joostenberg, (-DD), 14 Nov. 1965, Lewis

6222 (NBG); Jonkershoek, (-DD), 11 Dec. 1975, Kruger 119 (NBG);

Jonkershoek Valley, below Berg River Nek, (-DD), 29 Dec. 1989,

Goldblatt 9028 (MO. NBG); Jonkershoek, Swartboskloof, (-DD), 5

Dec. 1960, Van der Merwe 24-43 (NBG); 15 Jan. 1988, Forsyth 504

(NBG); Jonkershoek, Dwarsberg, (-DD), 9 Feb. 1959, Rycroft 2147

(NBG); Stellenboschberg above Coetzenberg, (-DD), Nov. 1992, De

Vos 2739 (NBG); Stellenbosch, Idas Valley, (-DD), 5 Dec. 1925, Lev-

yns 1458 (BOL). 3319 (Worcester): Groot Winterhoek, road to De

Tronk at Groot Kliphuis river crossing, (-AA), 10 Dec. 2007, Helme

5141 (NBG); Tulbagh, Waterv alberge, east of Suurvlak Falls, (-AC),

29 Dec. 2000, Hansford 21 (NBG); Elandsberg Nature Reserve,

Bosplaas trek. (-AC), 18 Nov. 1996, Parker 114 (NBG); Bains-

kloof Mtns, Bailey’s Peak, (-CA), 21 Feb. 1954, Esterhuysen 22748

(BOL); Bainskloof, (-CA), 1 Jan. 1941, Kies s.n. NBG62273 (NBG);

Limietberg, (-CA), 6 Jan. 1983, Goldblatt 6837 (MO, NBG); Wem-

mershoek Mtns, April Peak, (-CC), 15 Dec. 1940, Esterhuysen 4098

(BOL); Klein Drakenstein Mtns, upper Kasteelkloof, (-CC), 9 Jan.

1969, Kruger 879 (NBG); Du Toitskloof, (-CC). Jan. 1882, Tyson 905

(SAM); 1 Jan. 1950, Hall 285 (MO. NBG); Elandspad Catchment in

Agtertafelberg Kloof, (-CC), 28 Dec. 2000, Low 6769 (NBG); Zach-

ariaskloof Catchment, (-CC), 28 Dec. 1978, Kruger 1777 (NBG);

Franschhoek Pass, (-CC), Dec. 1929, Thode A2223 (NBG); 17 Jan.

1939, Gillett s.n. NBG62371 (BOL, NBG); 25 Jan. 1933, Pillans 7624

(BOL). 3418 (Simonstown): Somerset West, (-BA), Dec. 1930, Marl-

ley s.n. BOL22440 (BOL); Somerset West, Schaapenberg, (-BA), 14

Nov. 2006, Helme 4097 (NBG); Helderberg Nature Reserve, Porcu-

pine Buttress, (-BB), 3 Dec. 1993, Runnalls 639 (NBG); Sir Lowry’s

Pass, (-BB), 15 Jan. 1896, Schlechter 7245 (BOL); Dec. 1899, Bolus

4210 (BOL); 19 Dec. 1942, Compton 14222 (NBG); near foot of Sir

Lowry’s Pass, (-BB), 27 Jan. 1993, Goldblatt & Manning 9567 (MO,

NBG); Steenbrasdam, (-BB), 22 Dec. 1983, De Kock 73 (NBG); Steen-

bras Forest Reserve, (-BB), 24 Jan. 1951, Barker 7211 (BOL, NBG);

Elgin Forest Reserve, (-BB), Jan. 1951, Lewis 2906 (SAM); Kogel-

berg Forest Reserve, Blousteenberg, (-BD), 22 Dec. 1969. Boucher

972 (NBG); Betty's Bay, (-BD), 13 Dec. 1973, Vogts s.n. NBG173047

(NBG); Harold Porter Reserve, (-BD), 22 Nov. 1963, Topper s.n.

NBG70755 (NBG); Kogelberg, Oudebosch, (-BD), Dec. 1952, Stokoe

s.n. NBG173067 (NBG); slopes above Palmiet River Mouth, (-AC), 20

Dec. 1934, Salter 5180 (BOL); 26 Jan. 1947, Esterhuysen 136 73 (BOL,

NBG). 3419 (Caledon): Aries Kraal, (-AA), 30 Dec. 1944, Compton

16823 (NBG); sandy plateau above Viljoens pass, (-AA), 31 Jan. 1986,

Goldblatt 7627 (MO); Palmiet River Mtns, (-AC), Jan. 1947, Stokoe

s.n. (MO); Genadendal, (-BA), 28 Dec. 1894, Guthrie 3584 (NBG).

9. Thereianthus ixioides G.J. Lewis in Journal of

South African Botany 7: 41 (1941). Type; South Africa,

[Western Cape], Bain's Kloof [Bainskloof] at summit, 8

Dec. 1934. T.M. Salter 511 7 (BOL. holo.l).

Watsonia filifolia E.Mey. in Drege: 82 (1843), nom. nud. [South

Africa, Du Toitskloof, without date, Drege s.n. (MO)]. Watsonia punc-

tata var. filifolia E.Mey. ex Baker: 16: 159 (1877), nom. nud.

Plants 250-500 mm high. Conn globose, 12-20 mm

diam.; tunics of coarsely netted fibres, thickened and

claw-like at base, accumulating with age and forming

short neck around base of stem. Stem erect or flexed

outward above second sheath, unbranched, 1.0-1. 5 mm

diam. below spike. Cataphylls dry and papery, reddish

brown. Leaves 4(5), lowermost inserted below ground,

blade reaching or exceeding spike, green or dry at flow-

ering, linear, 200^100 x 1. 0-2.0 mm, closely 3-ribbed,

upper 3(4) leaves cauline, sheaths of all except upper-

most ± imbricate, blades successively shorter, up to 1 50

mm long, uppermost leaf entirely sheathing, distant from

next lower and separated from base of spike. Spike erect,

compact, densely 6-12-flowered; bracts imbricate for

V2 to 2L length, 8-13 mm long and two or three inter-

nodes long, outer acute or obtuse, either green below or

entirely brown and leathery, conspicuously and closely

veined, with narrow membranous margins, inner slightly

shorter, bifid. ± membranous with narrow brown zones

along veins. Flowers suberect, white to pale mauve

(rarely mid-blue), unmarked or lower three tepals

each with purple marking near base (rarely ± all tepals

marked); perianth actinomorphic; tube straight, 7 — 1 2( —

15) mm long, 0. 5-1.0 mm diam. at mouth, filiform and

tightly clasping style throughout, abruptly expanded in

distal 1 mm, exserted 1-9 mm beyond bracts, 0. 5-1.0 *

as long as tepals; tepals elliptical-oblanceolate, spread-

ing and slightly cupped or flat, 12-15 x 4-6 mm. Sta-

mens central, erect or diverging apically with anthers

spreading; filaments 3^4 mm long, swollen slightly at

base and occluding mouth of perianth tube; anthers 5-7

mm long, lilac or pale yellow; pollen cream-coloured.

Ovary obovoid. 2. 0-2. 5 mm long; style central, divid-

ing between base and middle of anther, branches ± 3-4

mm long, divided to near base. Capsules ovoid. 7-1 0 x

3-5 mm. Seeds ellipsoid-angled, 2-3 mm long, yellow-

ish brown testa striate-colliculate. Flowering time : late

Nov.-Dee. or early Jan. Figure 15.

262

Bothalia 41,2 (201 1 )

FIGURE 15. — Thereianthus ixioides. A-C, Manning 2198'. A, whole

plant; B, flower, front view; C, details of stamens and stigma. D,

E, Manning 3326: D, capsules; E, seeds. Scale bar: A, B, 1 0 mm;

C, 3 mm; D, 2 mm; E, 0.5 mm. Artist: John Manning.

Distribution and ecology. Thereianthus ixioides is a

narrow endemic of the mountains between Wellington

and Franschhoek, with most collections predictably from

the Bainskloof and Du Toitskloof Passes that traverse the

Limietberg and Klein Drakenstein Mtns respectively, and

from the Zachariaskloof catchment in the Wemmershoek

Mtns (Figure 16). Plants are locally common on cooler,

south-facing sandstone slopes at mid altitudes, mainly

around I 000 m, often in seasonally damp places, along

drainage lines or water courses. Flowering is stimulated by

fire but plants also bloom in open veld several years old.

FIGURE 16. — Known distribution of Thereianthus ixioides. •; T.

elandsmontanus , O.

The compact spikes of white to lilac (rarely blue),

actinomorphic flowers are probably adapted to a gen-

eralist pollination system and visited by a variety of

insects. The filiform perianth tube clasps the style tightly

for most of its length, with the mouth occluded by the

swollen bases of the filaments. Each flower produces

only trace amounts of nectar, which rises to the mouth of

the filiform perianth tube through capillary action, where

it can be accessed by insects with short mouthparts. The

flowers have a remarkable similarity in shape, colour

and markings to those of Ixia metelerkampiae, another

local endemic of the mountains near Wellington and vis-

its by the hopbine beetle (Scarabaeidae: Hopliini), Peri-

trichia sp., to both species were reported by Goldblatt et

al. (2000). Visits by hopbine beetles (Heterochelus sp.)

have also been observed at Franschhoek Pass (R. Saun-

ders, pers. comm. Jan. 2011), and pollination by these

beetles may be the primary strategy.

Diagnosis and relationships'. Thereianthus ixioides

is recognized by its compact, subcapitate spike of actin-

omorphic, white or pale lilac (rarely mid-blue) flow-

ers with filiform, straight perianth tube 7 — 1 2( — 15 ) mm

long and up to 1 mm diam., tightly clasping the style for

most of its length and abruptly dilated only in the dis-

tal 1 mm, with the filaments thus completely exserted.

The stamens are symmetrically arranged, with ± hori-

zontally spreading anthers, and the filaments occlude the

mouth of the perianth tube. It has been confused with T.

spicatus but that species has a mostly longer, ± cylindri-

cal perianth tube ( 1 2-) 15-25 mm long that is evenly and

gradually dilated from the base and not tightly clasping

the style, and unilateral, arcuate stamens, with the base

of the filaments shortly included within the top of the

perianth tube and not occluding the mouth.

History. Thereianthus ixioides appears to have been

first collected in Du Toitskloof by botanical collector

Johann Franz Drege, who was active in the southwest-

ern Cape from 1826-1834. This material was distributed

under the name Watsonia filifolia E.Mey., and although

published by Meyer (Drege 1843), the name was not

accompanied by a description and is thus nomenclatu-

rally illegitimate. It was later treated as a variety of T.

spicatus by Baker (1877) under the name W. punctata

v ar. filifolia. Our association of the name W. filifolia with

T. ixioides is based on a duplicate of the Drege collec-

tion bearing this name in the herbarium of the Missouri

Bothalia 41,2 (2011)

263

Botanical Garden (MO). The species was re-collected

from the same locality half a century later in 1882 by

the botanist William Tyson. His material, also identified

as Watsonia punctata (now T. bracteolcitus), raised no

interest. The next collection, that of Adele Grant made

in Bainskloof in 1926 was, however, recognized as rep-

resenting a new species of Watsonia (the genus to which

species of Thereianthus were then customarily referred),

an opinion that was confirmed when T.M. Salter, spe-

cialist on the genus Oxalis, also came across it in Bains-

kloof in December 1934. The species was subsequently

described by Lewis (1941) in her newly established

genus Thereianthus , based largely on Salter’s ample

material, and remained the most recent species to be rec-

ognized in the genus for more than half a century (Man-

ning & Goldblatt 2004).

Additional specimens examined

WESTERN CAPE. — 3319 (Worcester): Bain’s Kloof [Bainskloof),

(-AC), 26 Nov. 1926, Grant 2644 (BOL, MO); Dec. 1957, Loubser

865 (NBG); Limietberg, north of Bain’s Kloof [Bainskloof], 3000'

[914 m], (-CA), 6 Jan. 1983, Goldblatt 6837 (MO, NBG); Bains-

kloof, just before summit. (-AC), 26 Nov. 1998, Manning 2198

(NBG); Baviaanskloof, above pools in middle of valley, (-CA), 15

Jan. 2011 [fruiting]. Manning 3326 (NBG); Klein Drakenstein. Haal-

hoek Spitzkop, (-CC), 2 Jan. 1947, Esterhuysen 13541 (BOL); Klein

Drakenstein Mtns, 1.5 km SE Hugenootskop. (-CC), 14 Jan. 2007,

Helme 4510 (NBG); Du Toitskloof, (-CC), Oct.-Jan. without year

[1826-1834], Dr'ege s.n. (MO); 1882 [without month], Tyson 904

(SAM); Dec. 1950, Lewis 2292 (SAM); 10 Dec. 1950, Barker 7166

(NBG); Dec. 1951, Stokoe s.n. SAM63513 (SAM); Wemmershoek

Valley, Dec. 1944, Lewis 849 (SAM); Wemmershoek. (-CC), 14 Dec.

1940, Esterhuysen 4039 (BOL); Haalsneeuwkop, 4000' [1 220 m], (—

CC), Dec. 1944, Stokoe s.n. SAM56189 (SAM); near Pofadder Pool

on Elandspad River, (-CC), 24 Nov. 1985, Fischer s.n. NBG133017

(NBG); Du Toitskloof Mtns, Elandspad catchment in Agtertafelberg

Kloof, 1 020 m, (-CC), 28 Dec. 2000, Low 6773 (NBG); Zachari-

ashoek, Kasteelkloof catchment, (-CC), 2100' [640 m], 14 Dec. 1972,

Smith 69 (NBG); 30 Dec. 1982, Viviers 1122 (NBG); Zachariashoek

catchment, (-CC), 18 Dec. 1969, Haynes 260 (NBG).

10. Thereianthus elandsmontanus Goldblatt &

J.C. Manning, sp. nov.

Plantae 140-240 mm altae, cormo globoso 10-15 mm

diam.. tunicis fibrosis crassis, caule erecto eramoso, cat-

aphyllis papyraceis rubro-brunneis, foliis 4, infimum sub

terra insertum laminis linearibus 3- vel 4-nervosis, spica

erecta dense 7- ad 15-flora, bracteis imbricatis 8— 1 0(—

13) mm longis, bractea externa infra viridi vel omnino

brunnea coriacea, interna pauciter breviore, floribus

cremeis ad pallide cameis tepalis inferioribus in medio

marroninis striatis, perianthio leviter zygomorpho, tubo

(17-) 20-23 mm longo cylindrico, tepalis lineari-oblan-

ceolatis patentibus pauciter cupulatis 12-15 x 2-3 mm,

staminibus unilateralibus arcuatis, filamentis 4-5 mm

longis ex tubo 2-3 mm exsertis, antheris 4 mm longis

atropurpureis, styli ramis fere ad basem furcatis ± 3 mm

longis.

TYPE. — Western Cape, 3319 (Worcester); Bo-Her-

mon, Elandsberg Nature Reserve, Slangkop. low sand-

stone ridge, two years after fire, (-AC), 7 Nov. 2010,

J.C. Manning 3313 (NBG, holo.; MO, iso.).

Plants 140-240 mm high. Corm globose, 10-15 mm

diam.; tunics of coarse, matted fibres, accumulating with

age and forming short neck around base of stem. Stem

erect, unbranched, ± 1.5 mm diam. below spike. Cata-

phylls dry and papery, reddish brown. Leaves 4, lower-

most inserted below ground, blade exceeding spike, ±

dry at flowering, linear, 200-250 * 2-3 mm, closely 3-

or 4-ribbed, upper 3 leaves cauline, sheaths ± imbricate

and enveloping entire stem, blades successively shorter,

up to 100 mm long, uppermost leaf sheath ± reaching to

base of spike, with short blade 20-30 mm long. Spike

erect, densely 7-15-flowered; bracts imbricate for Ji to

2h length, 8—1 0(— 1 3) mm long and two or three inter-

nodes long, outer acute, either green below or entirely

brown and leathery, conspicuously and closely veined,

with narrow membranous margins, inner bract slightly

shorter, bifid, ± membranous with narrow brown zones

along veins. Flowers suberect, cream-coloured to pale

flesh-pink, flushed purple on tube, lower three tepals

with median linear maroon streak extending halfway

along blade; perianth weakly zygomorphic; tube slightly

curved outwards in distal 3 mm, ( 1 7— )20— 23 mm long,

± 1.5 mm diam. at mouth, ± cylindrical but widening

slightly distally, exserted 13-15 mm beyond bracts;

tepals linear-oblanceolate, inner three slightly smaller,

spreading and slightly cupped, 12-15 x 1. 5-3.0 mm.

Stamens arcuate; filaments 4-5 mm long, exserted 2-3

mm from tube; anthers 4 mm long, blackish purple; pol-

len purple. Ovary obovoid, 2. 0-2. 5 mm long; style arch-

ing above anthers, dividing between base and middle of

anthers, branches, ± 3 mm long, divided to near base.

Capsules narrowly ovoid, 5-7 x 3 mm. Seeds ellipsoid-

angled, ± 2 mm long, yellowish brown, testa striate-col-

liculate. Flowering time'. Nov. Figure 17.

Distribution and ecology’'. Thereianthus elands-

montanus is known from a single small population on

Slangkop, a low sandstone ridge on Elandsberg Nature

Reserve near the foot of the Elandskloof Mtns near

Hermon (Figure 16). Just a couple of dozen individuals

were found. Although only discovered two years after a

fire had cleared the site, it is clear that the species also

flowered in the first year after the bum since one of the

individuals had the remains of a previous flowering stem

still associated with the corm.

The moderately long-tubed, cream-coloured to flesh-

pink flowers with conspicuous narrow, maroon markings

on the lower three tepals bear a striking resemblance to

those of several co-occurring species of Iridaceae, nota-

bly Lapeirousia anceps (L.f.) Ker Gawl. and Tritonia

undulata (Burm.f.) Baker, that are adapted to pollination

by long-proboscid tabanid flies in the genus Philoliche,

and it is likely that Thereianthus elandsmontanus is also

a member of this pollination guild.

Diagnosis and relationships'. Thereianthus elands-

montanus is distinguished from the other ribbed-leaved

species in the genus (ser. Thereianthus ), by its cream-

coloured to pale flesh-pink flowers with moderately

long tube, 17-23 mm long and 1 .5-1.8 * as long as the

narrow, ± linear tepals which are 1. 5-3.0 mm wide, the

lower three each with a conspicuous maroon median

streak extending over the lower half. Unfortunately

the flowers do not retain their colour in the herbarium,

drying bluish lilac. T. elandsmontanus is most likely to

be confused with T. spicatus, which has a tube of simi-

lar length and which also occurs at Elandsberg Nature

Reserve but this species has blue (rarely white or pink

flowers) with broader, lanceolate tepals, 3-5 mm wide,

the lower three marked with a white flash at the base

264

FIGURE 17. — Thereianthus elandsmontanus, Manning 3313. A, whole

plant; B, flower, front view; C, flower, side view; D, capsules;

E, seed. Scale bar: A-C, 10 mm; D, 2 mm; E, 0.5 mm. Artist:

John Manning.

bordered distally by a small blue chevron, and slightly

shorter filaments, 3^4 mm long vs 4-5 mm long.

Although these differences may seem minor, we are

familiar with both species on the reserve and are con-

vinced that they are distinct. Another related species,

T. longicollis from the nearby Saronsberg and Eland-

skloof Mtns, has a much longer tube, 25-40 mm long

and 2. 0-2. 5 x longer than the tepals, the lower three of

which are marked with a dark, transverse chevron. The

Bothalia 41,2 (2011)

leaves in T. elandsmontanus are often slightly broader

than in T. longicollis, 2-3 mm vs 1-2 mm, with 3 or 4

vs consistently 3 raised veins. We considered the pos-

sibility that the Elandsberg plants might be an isolated,

short-tubed form of this species, possibly recognizable

at subspecific rank but the consistent morphological and

ecological differences between them suggest that rec-

ognition at species level is more appropriate. Typical T.

longicollis is a montane species and the clear separation

in tube length between it and T. elandsmontanus argues

for complete genetic isolation between them.

History: the most recently discovered species in the

genus, Thereianthus elandsmontanus was collected for

the first time in November 2010 on a low sandstone

ridge on Elandsberg Nature Reserve. This ridge has been

explored botanically since 1996 by the owners of the

property. Dale and Elizabeth Parker, but the species was

only noticed when it flowered in response to a controlled

bum of the ridge in early 2008.

Etymology: the name derives from the Elands-

berg Nature Reserve, where the species appears to be

endemic.

11. Thereianthus longicollis (Schltr.) G.J. Lewis

in Journal of South African Botany 7: 39 (1941). Watso-

nia longicollis Schltr.: 106 (1900). Type: South Africa,

[Western Cape], Tulbagh Waterfall, in saxosis, ± 1200'

[365 m], 10 Nov. 1896, R. Schlechter 9055 (B, holo.;

BOL!, GRA1, MO!, PRE! (4 sheets), iso.).

Watsonia punctata var. longicollis Baker: 177 (1892). Type: South

Africa, [Western Cape], Worcester [District], Zeyher s.n. K400455 (K,

holo.!).

Plants 150-350 mm high. Conn globose, 10-15 mm

diam.; tunics of coarsely netted fibres, thickened and

claw-like at base, accumulating with age and forming

short neck around base of stem. Stem erect, unbranched,

± 1.5 mm diam. below spike. Cataphylls dry and papery,

reddish brown. Leaves 4, lowermost inserted below

ground, blade reaching or exceeding spike, ± dry at

flowering, linear, 200-250 x 1-2 mm, closely 3-ribbed,

upper 3 leaves cauline, sheaths ± imbricate and envel-

oping entire stem or slightly distant, blades succes-

sively shorter, up to 150 mm long, uppermost leaf

sheath reaching to base of spike or separated from it,

with short blade 20-60 mm long. Spike erect, densely

7- 1 5(-20)-flowered; bracts imbricate for 'A to 2A length,

8— 1 2(— 1 7 ) mm long and two or three internodes long,

outer acute, either green below or entirely brown and

leathery, conspicuously and closely veined, with nar-

row membranous margins, inner slightly shorter, bifid,

± membranous with narrow brown zones along veins.

Flowers suberect, white or pale pink to purple-pink with

purple tube, lower three tepals each with purple chev-

ron or diamond-shaped markings near base; perianth

weakly zygomorphic; tube slightly curved outwards in

distal 3 mm, 25-40 mm long, ± 2 mm diam. at mouth,

± cylindrical but dilated slightly distally, exserted 20-30

mm beyond bracts, 2. 0-2. 5 x longer than tepals; tepals

oblong-oblanceolate, inner three slightly smaller, spread-

ing and slightly cupped, 12-17 x 2. 5-5. 5 mm. Stamens

arcuate; filaments 6 mm long, exserted 3 mm from tube;

anthers 4-6 mm long, purple; pollen purple. Ovaty

obovoid, 2. 0-2. 5 mm long; style arching above anthers.

Bothalia 41,2 (2011)

265

FIGURE 18. — Thereianthus longicollis, Manning 3325. A, whole plant;

B, flower, front view; C, flower, side view; D, dehisced capsules;

E, seed. Scale bar: A-C, 10 mm; D, 2 mm; E, 0.5 mm. Artist:

John Manning.

slopes of the Elandskloof Mtns. Plants grow in sandy

gravel or loamy ground on rocky sandstone slopes, and

flower regularly in open proteoid or restioid fynbos.

The very long-tubed, pink or purple-pink flowers with

purple markings on the lower three tepals have the stereo-

typical adaptations for pollination by long-proboscid flies.

At or near the type locality above Tulbagh Waterfall we

observed pollination visits by the tabanid fly, Philoliche

rostrata, the proboscis of which measured ± 40 mm, a

close match to the tube length in Thereianthus longicollis.

At this site other representatives of this pollination guild

included T. minutus, Geissorhiza confusa (Iridaceae) and a

long-tubed and large-flowered race of Lobelia coronopifo-

lia (Lobeliaceae) with a pink corolla.

Diagnosis and relationships ; Thereianthus longicol-

lis is separated from all other species in the genus by

its unusually long perianth tube, 25^-0 mm long and

2.0-2. 5 times longer than the tepals. The species could

only be confused with longer-tubed forms of T. spica-

tus , which have also been recorded from the mountains

above Porterville and which we have seen ourselves

above Tulbagh Waterfall but the perianth tube in T. spi-

catus is shorter, mostly less than 20 mm long (rarely

up to 25 mm) and always less than twice as long as the

tepals. and also narrower, at most 1.5 mm diam. at the

mouth. A second species in ser. Thereianthus with long-

tubed flowers, T. elandsmontanus from the foothills of

the Elandskloof Mtns, has a consistently shorter perianth

tube, 17-23 mm long, narrower tepals 2-3 mm wide,

with the lower half largely covered by linear purple

markings, and often broader leaves with 3 or 4 ribs (see

that species for further discussion).

History : Thereianthus longicollis was first encoun-

tered by botanical collectors Christian Frederick Ecklon

and Carl Ludwig Zeyher, who were active in the Cape

Colony through the middle of the nineteenth century

(Glen & Germishuizen 2010). Their material from Tul-

bagh Waterfall was distributed under the name Watso-

nia (Beilici) triticea (now T. bracteolatus) and the taxon

was first formally recognized as a long-tubed variant of

T. bracteolatus under the name Watsonia punctata var.

longicollis by Baker (1892, 1896). Although no material

was initially listed under this variety. Baker (1896) later

cited Zeyher s.n. under this name, thereby typifying it

unambiguously against a specimen bearing that inscrip-

dividing at ± middle of anthers, branches ± 4-5 mm

long, divided to near base. Capsules narrowly ovoid,

6-7 x 2. 5-3.0 mm. Seeds ellipsoid-angled. ± 2.5 mm

long, yellowish brown, testa striate-colliculate. Flower-

ing time ; mostly mid-Nov.-late Dec. Figure 18.

Distribution and ecology. Thereianthus longicollis is

a narrow endemic of the mountains between Porterville

and Wolseley (Figure 19), thus including the southern

end of the Grootwinterhoek Mtns, Saronsberg and the

Elandskloof Mtns. The species has been collected sev-

eral times at the type locality near the Tulbagh Waterfall

but is otherwise known from just a few records to the

north above Saron and Porterville, and most recently

from Elandsberg Nature Reserve on the lower western

16° 18° 20° 22° 24° 26° 28°

FIGURE 19. — Known distribution of Thereianthus longicollis, •.

266

Bothalia 41,2 (2011)

tion at the Kew Herbarium, which had been acquired

on 1 April 1865 from H.G. Reichenbach. The species

was subsequently re-collected at Tulbagh Waterfall on

10 November 1896 by botanist Rudolph Schlechter,

who realized that Baker was incorrect in associating the

plants with T. bracteolatus and described it as a distinct

species under the name W. longicollis, citing only his

collection. Although the labels on his specimens indicate

that Schlechter had initially intended to recombine Bak-

er's variety at specific rank, the name as later published

is presented as a new species and we treat it as such.

Additional specimens examined

WESTERN CAPE. — 3319 (Worcester): Twenty Four Rivers, (—

AA), without date, Zeyher 1608 [only one of' the specimens as indi-

cated on sheet] (SAM); De Hoek Estate, near Saron, (-AA), Nov.

1941, Stokoe s.n. SAM55617 (SAM); slopes of Great Winterhoek Mtns

near Saron, (-AA), Nov. 1941, Stokoe s.n. (BOL); Porterville, plateau

above [Farm] Gelukwaards, (-AA), Dec. 1910, Edwards 80 (BOL);

Bo-Hermon, Elandsberg Nature Reserve, lower slopes of Elandskloof

Mtns at southern boundary of farm at end of fence-road, (-AC), 5 Dec.

2010, Manning 3325 (NBG); Tulbagh Waterfall, (-AC), Nov. without

year, Ecklon cfi Zeyher s.n. SAM70719 (SAM); 16 Nov. 1941, Comp-

ton 12413 (NBG); [Elandskloof Mtns], Ontongsberg [Ontongskop],

lower slopes, (-AC), 16 Nov. 1941, Isaac s.n. (BOL); Voelvleiberge

[Elandskloofberge], ascent to Protea plain, (-AC), 29 Dec. 2000,

Hansford 20 (NBG).

EXCLUDED SPECIES

Beilia spicata var. brevifolia Kuntze: 305 (1898).

Type: South Africa, [Western Cape], Mowbray bei Caps-

tadt, collector and date unknown (specimen not located).

The brief description of this taxon \ folds brevibus 1-2

cm lads acuminads ’ and its collection at Mowbray,

a suburb of Cape Town, make it certain that it is not

Thereianthus spicatus , which has much narrower leaves

and is absent from the Cape Peninsula. The only pos-

sibility in Thereianthus is thus T. bracteolatus but the

broad leaves are inconsistent with this species as well

and suggest that it might be a species of Watsonia but its

identity remains unclear.

ACKNOWLEDGEMENTS

We thank Elizabeth Parker for her assistance and com-

panionship in the field; Roy Gereau for advice on nomen-

clatural questions and for revising our Latin descriptions;

Otto Leistner for his assistance with our Latin; Mary

Stififler, Research Librarian, Missouri Botanical Gar-

den, for providing copies of needed literature; Stephen

Boatwright for very kindly preparing the micrographic

plates; Michelle Smith for the electronic maps; Jonathan

Colville for beetle identifications; Amida Johns for assist-

ing us in the field; and the curator of the Bolus Herbar-

ium (BOL) for the loan of specimens. Collecting permits

were provided by CapeNature, South Africa.

REFERENCES

ANDREWS, H. 1801 . Ixia punctata. The botanist’s repository 3 : 1. 177.

BAKER, J.G. I 876a. New species of Ixieae. Journal of Botany, London

14: 236-239.

BAKER, J.G. 1876b. New Gladioleae. Journal of Botany, London 14:

333-339.

BAKER, J.G. 1877 [as 1878]. Systema iridearum. Journal of the Lin-

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Bothalia 41,2: 269-276 (2011)

Romulea pilosa and R. quartzicola (Iridaceae: Crocoideae), two new

species from the southern African winter rainfall region, with nomen-

clatural corrections including new names for R. amoena , R. neglecta

and R. rosea var. reflexa

J.C. MANNING*. P. GOLDBLATT** and A.D. HARROWER***

Keywords: Iridaceae, new species, nomenclature, Romulea Maratti, southern Africa, taxonomy

ABSTRACT

Romulea pilosa J.C. Manning & Goldblatt and R. quartzicola J.C. Manning & Goldblatt are two narrow endemics from

the southern African winter rainfall region. An early, fragmentary collection of R. pilosa from Riviersonderend lacked the

diagnostic corm and was thus mistakenly associated with R. tetragona (sect. Ciliatae) as \sx.flavandra MP.de Vos because of

the highly distinctive pilose, H-shaped leaf. The rediscovery of the taxon in the wild shows it to be a previously unrecognized

member of sect. Aggregatae, distinguished by its unusual foliage and bright orange flowers. R. quartzicola was grown to

flowering from seeds collected from quartz patches in southern Namaqualand and proved to be a new species of sect. Ciliatae,

distinguished by its early flowering, short, subclavate leaves with reduced sclerenchyma strands, and bright yellow flowers

with short bracts. R. neglecta M.P.de Vos, a rare endemic from the Kamiesberg in Northern Cape, is a later homonym for the

Mediterranean R. neglecta Jord. & Fount, and the earliest name for this plant is shown to be R. speciosa (Ker Gawl.) Baker,

typified by an illustration in Andrews’ The botanist's repository. An epitype is designated to fix the application of the name.

We have also examined the type illustration of R. pudica (Sol. ex Ker Gawl.) Baker, hitherto treated as an uncertain species,

and are confident that it represents the species currently known as R. amoena Schltr. ex Beg., and takes priority over it as

being the earlier name. The type of R. reflexa Eckl., a new name for the later homonym I. reflexa Thunb. and the basionym

of R. rosea var. reflexa (Eckl.) Beg., has been mistakenly identified as an Ecklon collection but is in fact the collection that

formed the basis of Thunberg’s I. reflexa. This collection is actually a form of R.flava Lam., and the name R. rosea var. re-

flexa is thus moved to the synonomy of that species. The variety currently known under this name should now be known as R

rosea var. muirii (N.E.Br. ) Goldblatt & J.C. Manning. Finally, the protologue of R. parviflora Eckl., until now treated under

the synonomy of R. obscura Klatt var. obscura, is in fact consistent with R. rosea var. australis (Ewart) M.P.de Vos, and we

therefore include the name in the synonomy of the latter.

INTRODUCTION

Romulea Maratti, one of the larger genera of Iri-

daceae (Goldblatt & Manning 2008), is widely distrib-

uted through eastern sub-Saharan Africa, the Medi-

terranean and the Near East. It is centred in the winter

rainfall region of southern Africa, where 80 % of the

species occur ( De Vos 1 972, 1 983; Manning & Goldblatt

2001,2004,2006).

Flower structure in the genus is conservative (Gold-

blatt et al. 2002), with a few notable exceptions, and

characters of the corm are essential in the primary subdi-

vision of the genus into subgenera and sections (Manning

& Goldblatt 2001, 2004). Flower colour and markings, as

well as the texture and details of the membranous mar-

gins of the bracts provide the primary specific characters.

Additional valuable anatomical characters include the

distribution of vascular bundles in the leaf and the devel-

opment of sclerenchyma caps and strands.

Five new species have been described since the

last review of the genus (Manning & Goldblatt 2001,

* Compton Fterbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town / Research Centre for

Plant Growth and Development, School of Biological and Conservation

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

X01, Scottsville 3209, South Africa.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

den, P.O. Box 299, St. Louis, Missouri 63166, USA.

*** Kirstenbosch National Botanical Garden, South African National

Biodiversity Institute, Private Bag X7, 7735 Claremont, Cape Town.

MS. received: 2011-04-14.

2004, 2006), and a further two highly local endemics

are described here, bringing to 83 the number of spe-

cies recorded from southern Africa and raising the total

number in the genus to ± 97 spp.

Romulea pilosa , a spring-flowering species endemic

to stony renosterveld in the Overberg south of Rivier-

sonderend, was collected at least as long ago as the

early 1930s but the fragmentary material was misidenti-

fied until recently. It is named for the distinctive, pilose

leaves. R. quartzicola , a recently discovered, winter-

flowering species from quartz patches in the southwest-

ern comer of the Knersvlakte, is named for its unusual

habitat. We also deal with the nomenclature of Romulea

neglecta M.P.de Vos ( 1983), a later homonym. The earli-

est legitimate name for the species is R. speciosa (Ker

Gawl.) Baker (1877), which coincidentally has the same

type as R. neglecta.

Collections at BOF, NBG, PRE and SAM, the main

herbaria with good representation of Cape species, were

consulted (herbarium acronyms after Holmgren et al.

1990).

TAXONOMY

1. Romulea pilosa J.C. Manning & Goldblatt, sp.

nov.

Ad sectionem Aggregatae M.P.de Vos pertinens et

recognita caule subterraneo vel breviter aerio, foliis rela-

tive brevibus falcatis H-formibus 4-6 mm latis, margin-

270

Bothalia 41,2 (201 1 )

bus late alatis ciliatisque, floribus salmoneis vel auran-

tiacis cupulo flavo, tubo perianthii infundibuliformi ± 5

mm longo parte inferiori ± 1 mm longo, tepalis oblan-

ceolatis 20-30 x 7-10 mm, filamentis 5-6 mm longis

minute pilosis in dimidio inferiori, antheris ± 4 mm

longis.

TYPE. — Western Cape, 3420 (Bredasdorp): Farm

Fonteinkloof, between Riviersonderend and Protein, hill

W of homestead, (-AA), 15 Aug. 2009, Manning 3220

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

Plants ± 60 mm high; stem subterranean or shortly

aerial, simple or with up to 2 branches. Conn subglo-

bose, asymmetric, obliquely flattened towards base

with crescent-shaped basal ridge; tunics hard, smooth,

dark brown, split into clusters of convergent fibrils

on basal ridge and splitting along clusters into nar-

row vertical strips, drawn into short fibrils up to 2 mm

long above. Cataphylls 3, membranous, flushed green-

ish above ground. Foliage leaves 2, lowermost 1 basal,

blades falcate, H-shaped in section, lowermost 50-70

x 4-6 mm, upper shorter, lateral ribs reduced, median

(apparently marginal) ribs extended laterally to form

flange-like wings 1-2 mm wide along each side of

blade, densely villous along margins of wings with hairs

1-2 mm long in two diverging rows. Inflorescence of

solitary flowers; peduncles angled in section and vil-

lous along angles in distal part; outer bracts subobtuse,

13-15 mm long, green with narrow, translucent mem-

branous margins, strongly keeled along median and

submarginal veins, longitudinally indexed along sub-

marginal veins, median keel sometimes double, keel

and submarginal veins densely pubescent, inner bracts

green with broad translucent white margins, ± as long

as outer, bicarinate with keels densely pubescent. Flow-

ers deeply cup-shaped, cup ± 10 mm deep, salmon-pink

to dull orange but golden yellow in lower half to two

thirds, yellow extending beyond cup margin onto lower

third of limb, outer tepals flushed coppery on reverse,

unscented, 30-40 mm diam.; perianth tube funnel-

shaped, 5 mm long with lower narrow portion ± 1 mm

long, tepals oblanceolate, 20-30 x 7-10 mm. Stamens

yellow; filaments inserted at base of cup, free, 5-6 mm

long, minutely pilose in lower half; anthers parallel, 4

mm long. Style dividing opposite lower half of anthers,

branches ± 1 .5 mm long, divided for ± half their length.

Capsules oblong-ellipsoid, 7-9 x 5 mm long, fruiting

peduncles diverging, straight. Seeds subglobose, ± 2 mm

diam. Flowering time : Aug. Figure 1.

Distribution and ecology, known with certainty only

from a single locality south of Riviersonderend on the

Farm Fonteinkloof (Figure 2), where a small population,

numbering less than fifty plants, occurs in a remnant

patch of renosterveld vegetation, too stony to plough.

The plants are wedged among small vertical riffles of

shale on the crest of a hill as part of a Ruens Silcrete

Renosterveld community, a highly endangered veg-

etation type found as scattered pockets in the Overberg

region (Mucina & Rutherford 2006).

Diagnosis and relationships : the oblique conn with

tunics split along the basal ridge into clusters of con-

verging fibrils place Romulea pilosa in sect. Aggrega-

tae M.P.de Vos (1972), making it the ninth member of

this small group. Its relatively short, falcate, H-shaped

leaves 4-6 mm wide, with ciliate, winged margins, and

its salmon to orange flowers with a yellow cup, make

it instantly recognizable in the genus. Similar winged,

ciliate leaves are known only in two other species in

the genus, R. hirsuta Schltr. and R. tetragona M.P.de

Vos. Although associated in sect. Hirtae Beg. by De

Vos (1972) on the basis of their similar leaf morphol-

ogy, these two species have fundamentally different

conn tunic morphologies, leading Manning & Gold-

blatt (2001) to segregate them respectively to subgen.

Romulea sect. Cilatae (M.Pde Vos) J.C. Manning &

Goldblatt and subgen. Spatalanthus (Sweet) Baker. The

discovery of a third, evidently unrelated species with

similar, H-shaped and winged leaves now suggests that

this unusual leaf morphology has evolved independently

at least three times within the genus. The three species

concerned occupy quite different geographical regions

with differing ecology, and the adaptive value of this

leaf type is not immediately clear.

Within sect. Aggregatae , Romulea pilosa is evidently

most closely allied to R. dichotoma (Thunb.) Baker and

R. jugicola M.Pde Vos, both of which typically have

aerial stems, ciliate peduncles, and a solitary basal leaf

with ciliate rib margins. The leaves of R. dichotoma in

particular tend to have wider longitudinal grooves, with

the marginal ribs prominently expanded and almost

wing-like. Both species are found on clay soils in renos-

terveld communities in the southern Cape. R. jugicola ,

known only from the foothills of the Kammanassie Mtns

in the Fittle Karoo, is distinguished by its well-exserted

aerial stems, up to 30 mm tall, with a well-developed,

fibrous neck around the base, and orange-yellow peri-

anth; R. dichotoma is more widespread, extending from

Stanford to Humansdorp, and has mostly magenta or

rarely salmon-pink flowers with a yellow cup. It is most

evidently separated from R. pilosa by its generally taller

stem, up to 350 mm high, usually branched dichoto-

mously near the top, and narrower, distinctly 4-ribbed

leaves, 1-2 mm in diameter.

The leaves of Romulea pilosa are anatomically con-

sistent with sect. Aggregatae in their well-developed

sclerenchyma girders joining the primary vascular bun-

dles to the epidermis (Manning & Goldblatt 2001), an

evidently apomorphic character state that is only inter-

mittently developed in subgen. Romulea. Although gird-

ers are also developed in the secondary bundles in other

members of the series, they are almost confined to the

primary bundles in R. pilosa (occurring only rarely in

one or two secondary bundles near the wing margins

in some leaves). Stomata are present on the blade and

‘inner’ wing surfaces but are lacking on the ‘outer’ wing

surface, i.e. those making the acute angle along the leaf

margins.

Morphologically similar, H-shaped leaves have

evolved repeatedly in several genera of the Cape Iri-

daceae. Notable examples in Gladiolus sect. Homoglos-

sum are the Gladiolus gracilis-G. caeruleus-G. recur-

vus group in ser. Carinatus as well as most members

of ser. Homoglossum and Tristis; and some members of

Geissorhiza sect. Engysiphon. We have observed that the

wings in these species curve inward over the stomatif-

erous grooves when the plants are water-stressed, and

Bothalia 41,2 (2011)

271

FIGURE 1. — Romulea pilosa , Manning 3220. A, whole plant; B, outer bract; C, inner bract; D, stamens and style; E, detail of leaf

margin; F, t/s leaf; G, detail of corm fibril clusters. Scale bars: A-C, 10 mm; D, 2.5 mm; E, 1 mm; F, 0.5 mm; G, 0.2 mm. Art-

ist: John Manning.

speculate that this mechanism facilitated the develop-

ment of a relatively broader photosynthetic leaf surface

while still controlling water loss.

History, described here from a population discovered

in August 2009 by Napier residents and dedicated con-

servationists, Cameron and Rhoda McMaster, the spe-

cies appears to have been first collected in 1932 or 1933

by Grace Neethling of Riviersonderend. This discovery

is represented by a fragmentary specimen in the Bolus

herbarium, comprising a solitary detached leaf and a sin-

gle dissected flower. The plant from which these parts

were picked was cultivated at Kirstenbosch but subse-

quently disappeared and thus no corm was preserved.

The fragments are accompanied by a coloured painting

of the floral parts showing the distinctive orange tepals

w'ith their yellow cup. Although initially intending to

describe the collection as a new species. De Vos (1972)

subsequently treated it as Romulea tetragona war. flavan-

dra M.P.de Vos (otherwise endemic to the southwest-

ern Karoo), although noting the anomalous colour of

the flowers of the Riviersonderend collection. With the

discovery of a wild population, it is now evident that

this plant is quite unrelated to R. tetragona and does

indeed represent a distinct species. The locality where

Miss Neethling found her plant is unknown, although

the Neethling family owned the Farm Bon Esperance at

the foot of the Riviersonderend Mtns at the time (Lewis

1947). Most of the natural vegetation below these moun-

tains has since disappeared under cultivation and we

will never know if her collection came from the present

locality or from another population in the vicinity.

272

Bothalia 4 1 .2 (2011)

FIGURE 2. — Known distribution of Romulea pilosa, O; R. quartzi-

cola , •.

Additional material seen

WESTERN CAPE. — 3419 (Caledon): Riviersonderend, (-BB), fid.

[flowered] at Kirstenbosch, 24 Aug. 1933, G. Neethling s.n. BOL24789

(BOL).

2. Romulea quartzicola J.C. Manning & Gold-

blatt, sp. nov.

Ad sectionem Ciliatae (M.P.de Vos) J.C. Manning

& Goldblatt pertinens et recognita florentia praecoci,

foliis relative brevibus obtusis subclavatis, floribus gran-

dis canarinis, tubo perianthii profundo 7-8 mm longo,

tepalis 20-22 x 6-10 mm, filamentis, 7-9 mm longis

quern antheris longioribus, bracteis ovatis vel subor-

bicularibus 8-13 mm longis, marginibus membranaceis

angustis ecoloratis vel purpureo-suffusis.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Kners-

vlakte. Farm Moedverloor, ± 15 km NE of Lutzville,

(-AD), 21 June 2010, A.D. Harrower & J.C. Manning

4395 (NBG, holo.; MO, iso.).

Plants 50-80 mm high; stem aerial, up to 35 mm

long, simple or with 1 branch. Conn subglobose, ± 10

mm diam., asymmetric, obliquely flattened towards

base with crescent-shaped basal ridge; tunics hard,

smooth, dark brown, split into numerous fine parallel

fibrils on basal ridge and into narrow teeth up to 5 mm

long above. Cataphylls 3, membranous, flushed green-

ish above ground. Foliage leaves (1)2, all basal, blades

mostly ± as long as flowering stem, cylindrical-subcla-

vate, obtuse, narrowly 4-grooved, 30-80 x (1.0-) 1.5—

2.5 mm; cauline leaf present only when lateral inflor-

escence developed, inserted midway or two thirds up

stem, much shorter than basal leaves, entirely sheathing

or with blade up to 30 mm long. Inflorescence of soli-

tary flowers; outer bracts broadly elliptical-suborbicular,

concave, obtuse, green with narrow, translucent (some-

times purplish tinted) membranous margins, 8-13 mm

long, inner bracts similar but with broader membranous

margins, ± as long as outer, notched apically. Flowers

deeply cup-shaped, 25-30 mm diam., cup 12-15 mm

deep, bright chrome-yellow (rarely peach-coloured),

with 3-5 inconspicuous dark veins at base of cup, rarely

Hushed copper or pale green on reverse, weakly clove-

scented; perianth tube funnel-shaped, 7-8 mm long

with lower narrow portion ± 1 mm long, tepals oblan-

ceolate, 20-22 x 6-10 mm. Stamens yellow; filaments

inserted at base of cup, lower 2^1 mm included within

perianth tube, free, pubescent in lower two thirds, espe-

cially towards base, 7-9 mm long; anthers parallel, 4-5

mm long, thecae well separated by broad connective

0.50-0.75 mm wide. Style dividing opposite upper half

of anthers, branches ± 3 mm long, divided for entire

length into six filiform branchlets. Capsules and seeds

unknown. Flowering time: mid-June to July. Figure 3.

Distribution and ecology’-, known so far from several

small populations on the Farm Moedverloor, northeast

of Futzville in the extreme southwestern Knersvlakte

(Figure 2). Plants are restricted to the drainage basin

of the Moedverloor River, where they are localized on

southern or southwest-trending slopes, typically just

below the crest of the quartzite outcrops that fringe the

eastern edge of the basin, but occasionally on the lower

slopes. The species is relatively common where it occurs

but populations never extend over a large area. Plants

grow among quartz pebbles only in relatively dense

stands of Knersvlakte Dwarf Vygieveld (Mucina &

Rutherford 2006), a dwarf succulent shrubland domi-

nated by various small Aizoaceae in the genera Argvro-

derma , Cheiridopsis, Conophytum and Monilaria. This

highly specific ecology appears to be determined largely

by the cooler aspect and by the associated higher pre-

cipitation. The rocky crests on which the species mostly

occurs face the ocean, and the localized presence of

lichens on the summit rocks suggests that coastal fogs,

which are a feature of the region in the autumn and win-

ter, play an important role in providing additional mois-

ture to these sites, as attested also by the relatively dense

associated succulent shrublets compared with adjacent

slopes with a more northerly aspect. The Moedverloor

basin has exposed an extensive and rather localized area

of these quarzite slopes, the Moedverloorberg to the

north being granitic and the hills to the south primarily

loamy sand. A localized distribution and low population

size is a characteristic of other quartz-loving species in

the region (Snijman & Harrower 2009). Focal changes

in soil chemistry, depth, particle size, aspect, insolation

and available moisture appear to be responsible for this

very patchy ecology (Schmiedel & Jurgens 1999).

Other locally endemic geophytes from the southwest-

ern Knersvlakte include Bulbine dactylopsoides and

B. haworthioides (Asphodelaceae), Lachenalia patula ,

Ornithogalum naviculum , and the recently described

Drimia fimbrimarginata (all Hyacinthaceae) (Snijman &

Harrower 2009).

The flowers open between 13:00-14:00 in the after-

noon and close in the evening near sunset. They remain

closed in cold and overcast weather.

Diagnosis and relationships : the conn with a fimbri-

ate basal ridge places Romulea quartzicola in sect. Cili-

atae (M.P.de Vos) J.C. Manning & Goldblatt of subgen.

Romulea , where it is distinguished by its early flower-

ing, relatively short, obtuse, subclavate leaves, large,

deeply funnel-shaped, canary-yellow flowers with tepals

20-22 x 6-10 mm and perianth tube 7-8 mm long, and

the relatively long filaments, 7-9 mm long, much longer

than the anthers. The bracts are ovate or suborbicular.

273

Bothaha 41,2 (2011)

8-13 mm long, and both the inner and outer bracts have

narrow, colourless or purple-flushed membranous bor-

ders. The leaves of R. quartzicola are morphologically

distinctive in broadening appreciably in the distal half,

thus somewhat clavate in appearance, but they are ana-

tomically simple in lacking vascular girders as well as

rib marginal bundles or strands. All four primary bun-

dles have incomplete sclerenchyma sheaths restricted to

phloem caps, and even these are absent in the second-

ary bundles. This is a relatively uncommon condition, as

most species in the section have complete sclerenchyma

bundle sheaths around the lateral primary bundles (De

Vos 1972; Manning & Goldblatt 2001). The leaves of

FIGURE 3. — Romulea quartzicola,

Harrower 2756. A, whole

plants; B, outer bract; C, inner

bract; D, stamens and style; E,

anther, outer face; F, details

of style branches; G, t/s leaf.

Scale bars: A-C, 10 mm; D,

E, 2.5 mm; F, 0.5 mm. Artist:

John Manning.

R. quartzicola are thus among the least sclerified in the

entire genus, matched in this regard, as far as is known,

only by R. namaquensis M.P.de Vos.

The combination of yellow flowers and green bracts

without the conspicuous, brown-streaked membranous

margins that characterize the Romulea pearsonii group

of species, places R. quartzicola in a small group of

sect. Ciliatae that comprises R. citrina Baker, R. ellip-

tica M.P.de Vos, R. lutea J.C. Manning & Goldblatt, R.

montana Beg. and R. sulphurea Beg. Among this group

it is readily distinguished from all except R. sulphurea

by the relatively long filaments, ± twice as long as the

274

Bothalia 41,2 (2011)

anthers. The rarely collected R. sulphured, which was

represented only by the type (collected in 1897) until

we relocated it in 1999 (Manning & Goldblatt 2001),

is still known only from the Pakhuis Pass in the north-

ern Cedarberg, where it forms dense populations on

seasonally wet sandstone pavement, flowering in early

spring, in July and August. It is distinguished from R.

quartzicola not only by its ecology and distribution but

by its filiform leaves, ± 0.5 mm diam., smaller, sweetly

scented flowers with elliptical tepals 12-20 mm long,

mostly smaller bracts, 7-10 mm long, the inner with

broad, brown-speckled membranous margins, and by the

shorter anthers, 2. 5-3.0 mm long.

Perianth colour is an important character in Romulea

to the extent that species are either pink- or yellow-flow-

ered but never both. Plain yellow flowers without dark

blotches on the inner surface are uncommon in Romulea

and restricted to a handful of species of sect. Ciliatae,

only one of which, R. lutea J.C. Manning & Goldblatt,

has been recorded from southern Namaqualand. This

species, a recent discovery from sandy washes west of

Koekenaap (Manning & Goldblatt 2008), has plain,

golden yellow flowers superficially similar to those of R.

quartzicola but it differs not only in its later flowering,

August to September, but in its narrower leaves, ± 1 mm

in diameter, with only the lowermost leaf basal, shorter

perianth tube, ± 3 mm deep, and shorter filaments, 4 mm

long and ± as long as the anthers. The two species dif-

fer also in leaf anatomy. In R. lutea the lateral primary

bundles have complete sclerenchyma sheaths and the

secondary bundles have well-developed phloem caps

extending to the epidermis as girders. Both of these con-

ditions are lacking in R. quartzicola.

History, the species was originally collected as a

single fruit in July 2005. Seed was germinated in April

2006 in the nursery at Kirstenbosch Botanical Garden,

and four plants were grown to maturity. These flowered

for the first time on 6 June 2009, when it became clear

that they represented an undescribed species. Type mate-

rial was collected the following year.

Additional material seen

WESTERN CAPE. — 3118 (Vanrhynsdorp): Knersvlakte, Farm

Moedverloor, ± 15 km NE of Lutzville, (-AD), 15 May 2010 [flow-

ered in cultivation, plants originally collected 22 July 2005], A.D. Har-

rower 2756 (NBG); 21 June 2010, A.D. Harrower & J.C. Manning

4397 (NBG), 4399 (NBG).

NAME CHANGES AND NOMENCLATURAL CORRECTIONS

1 . The protologue of Trichonema pudica (Ker Gawler

1810), the basionym for Romulea pudica (Ker Gawl.)

Baker ( 1 877), includes a painting of a pink-flowered spe-

cies of Romulea with dark longitudinal streaks in a pale

cup. This plant was brought from the Cape by a Miss

Symonds and given to Messrs Whitley and Borme of

Brompton, London, in whose greenhouse it was raised.

They later provided the flowering specimen to the artist

who executed the painting reproduced in Curtis’s Botani-

cal Magazine. Depicting a plant with two basal leaves

and a symmetrical, bell-shaped conn with a neatly scal-

loped basal rim, this illustration clearly represents the

species currently known under the name R. amoena, a

narrow endemic of the Bokkeveld Mtns south of Nieu-

woudtville in Northern Cape. Ker Gawler (1810) cited

one specimen in the protologue, Ixia pudica Sol. ms in

Herb. Banks (now the herbarium of the British Museum

of Natural History). The plant so annotated by Solan-

der was collected by Oldenburg and has no date, precise

locality or number. De Vos (1972, 1983) identified it as

R. flava Lam. and we agree with her determination after

having examined a digital image of the specimen.

Ker Gawler first listed the name Ixia pudica when

he described the genus Trichonema (Ker Gawler 1802),

including it among one of several species that he

believed correctly belonged in his new genus. He later

(Ker Gawler 1804, 1809) listed ‘ Trichonema pudica

Gawl.’ as one of several species of Trichonema but

it was only in 1810 that the validating protologue was

published (Ker Gawler 1810). De Vos (1983) cited the

painting as the type (‘iconotype’) of T. pudica and we

endorse her action. The painting, showing a plant with

red perianth, pale, darkly striped cup, and two basal

leaves, differs sharply from the alternative choice of

type, the Oldenburg specimen, which has a single basal

leaf and apparently had a white perianth with a plain

yellow cup. Significantly, the trivial name, rose-flow-

ered trichonema, that accompanies the protologue, pro-

vides a direct and clear reference to the painting. Only

the painting, therefore, accords with the words and title

of the protologue and we formally designate it the lec-

totype. Although De Vos (1983) was not able to associ-

ate the painting (and thus the name) with any wild spe-

cies, we are confident that it is a good representation of

R. amoena (Beguinot 1907), the only known species of

this genus with a dark pink to red perianth with pale,

streaked cup, two basal leaves, and a bell-shaped conn

scalloped along the basal rim. We accordingly reduce R.

amoena to synonymy under R. pudica, a combination

provided by Baker (1877). Daniel Solander evidently

provided the inspiration for the name of the species but

the specimen bearing his annotation is not the type.

Romulea pudica (Sol. ex Ker Gawl.) Baker in Jour-

nal of the Linnean Society, Botany 16: 89 (1877). Tricho-

nema pudicum Sol. ex Ker Gawl: t. 1244 (1810) [T. pudi-

cum Sol. ex Ker Gawl.: 223 (1804) et sub t. 1225 (1809),

nom. nud.]. Ixia pudica (Sol. ex Ker Gawl.) Roem. &

Schult.: 377 (1817). Geissorhiza pudica (Sol. ex Ker

Gawl.) Klatt: 655 (1866). Bulbocodium pudicum (Sol.

ex Ker Gawl.) Kuntze: 700 (1891). Romulea rosea var.

pudica (Sol. ex Ker Gawl.) Baker: 42 (1896). Lectotype

here designated: South Africa, without precise locality,

illustration in Ker Gawl. in Curtis’s Botanical Magazine

31: t. 1244 (1810). [The syntype, without precise locality,

Oldenland s.n. annotated Ixia pudica by Solander (BM) =

R. flava Lam. is in serious conflict with the protologue.]

Romulea amoena Schltr. ex Beg.: 334 (1907), syn. nov. Type:

South Africa, [Northern Cape], Bokkeveld Mtns, Papelfontein [Pap-

kuilsfontein], Schlechter 10896 (G, holo.; BM, BOL, GRA, K, MO,

PRE, S, Z, iso.).

2. The phrase ‘ Blumen rosenroth klein (flowers

rose-red, small) constitutes the entire diagnosis for

C.F.Ecklon’s Romulea parviflora, which he collected on

the Cape Peninsula near Green Point. The species, typi-

fied by a specimen in the Stockholm Herbarium so anno-

tated by Ecklon (Nordenstam 1972), was determined as

Bothalia 41,2 (2011)

275

R. obscura Klatt by De Vos ( 1972). Nordenstam ( 1972)

felt that this diagnosis was sufficient to validate Eck-

lon’s species, and that, as the earlier name it should thus

replace R. obscura. De Vos (1973) disagreed, regarded

Ecklon’s species as inadequately diagnosed and there-

fore invalidly published, and accordingly listed the name

in synonymy both under R. obscura var. obscura and

under R. rosea var. australis Ewert (De Vos 1972). Later,

in the Flora of southern Africa (De Vos 1983) she cited

it solely as a synonym of R. obscura var. obscura. This

species usually has yellow to apricot, occasionally pale

pink flowers (De Vos 1972), whereas R. rosea , includ-

ing its varieties, has pale to deep pink or red flowers.

We are of the opinion that the species is therefore better

assigned to R. rosea on the basis of flower colour. The

small flower size indicates that it should be assigned to

synonymy under R. rosea var. australis.

Romulea rosea var. australis (Ewart) M.P.de

Vos in Journal of South African Botany, Supplement 9:

254 (1972). Romulea cruciata var. australis Ewart: 43

(1907). Type: Australia, Victoria, near Melbourne, Tovey

s.n. [MEL, lecto., designated by De Vos: 252 (1972);

BOL, iso.].

Romulea parviflora Eckl.: 19 (1827). Trichonema parviflorum

(Eckl.) Steud.: 702 (1841), syn. nov. Type: South Africa, [Western

Cape], Cape Peninsula, Green Point, Sept. 19-24, Ecklon s.n. (S,

holo.).

3. First described as Ixia reflexa by Thunberg (1811),

the epithet was transferred to Romulea by Ecklon

(1827), who explicitly cited Thunberg’s species as the

basionym. Ixia reflexa of Thunberg is, however, a homo-

nym for a species of that name described by Andrews

(1797) [now, incidentally, Ixia scillaris L.]. Ecklon's use

of the name is to be treated nomenclaturally as a new

name dating from 1827 (McNeil et al. 2006: Art. 58.1)

but it remains typified by the type of Thunberg’s homo-

nym. The taxon is conspecific with R. flava var. flava

(De Vos 1972; 1983). Beguinot (1908), and later De Vos,

however, mistakenly regarded Ecklon’s name as typi-

fied by his (Ecklon’s) own collections bearing that name,

which are a form of R. rosea , and accordingly treated

them as var. reflexa of that species. Based on the actual

type, however, R. rosea var. reflexa is a nomenclatu-

ral synonym of R. flava. The variety var. reflexa sensu

Beguinot, a local endemic of the Riversdale District,

then requires a new name and its only synonym, at spe-

cies rank, is R. muirii , which is available for the taxon.

We accordingly provide the combination below.

Romulea rosea var. muirii (N.E.Br. ) J.C. Manning

& Goldblatt, comb, et stat. nov. Romulea muirii N.E.Br.:

467 (1932). Type: South Africa, [Western Cape], Rivers-

dale, Muir 4848 (K, holo.).

R. rosea var. reflexa sensu Beg. (1908) et De Vos (1972), non

(Eckl.) Beg. (1908) [= R. flava (Lam.) M.P.de Vos].

Romulea flava (Lam.) M.P.de Vos in Journal of

South African Botany, Supplement 9: 98 (1972). Ixia

flava Lam.: 109 (1791). Type: without precise locality or

collector [P: Herb. Lamarck, lecto., designated by M.P.

deVos: 99(1972)].

R. rosea var. reflexa (Eckl.) Beg.: 158 (1908). R. reflexa Eckl.: 18

(1827), as nom. nov. pro Ixia reflexa Thunb.: 55 (1811), hom. illegit.,

non 1. reflexa Andr. (1797), syn. nov. Type: South Africa, without pre-

cise locality, Thunberg s.n. UPS-THUNB984 [UPS-THUNB, lecto.,

designated by De Vos: 27 (1983)].

4. In her Flora of southern Africa account of Rom-

ulea, De Vos (1983) recognized Ixia neglecta Schult.

(1822) as an earlier name for what was until then R.

oliveri M.P.de Vos, a narrow endemic of the Kamies-

berg in Namaqualand, Northern Cape. While true, the

combination that she proposed in Romulea is techni-

cally a new name because she deliberately excluded

the type of both I. neglecta and its synonym Tricho-

nema speciosum Ker Gawl. Her new name is, however,

a homonym for the Mediterranean R. neglecta Jord. &

Fourr. (1868). Ixia neglecta of Schultes (1822) was itself

a new name in Ixia for Trichonema speciosum (Ker

Gawler 1804) — not the later account of the species in

1812 cited by De Vos — because I. speciosa Andrews

bars its transfer to Ixia. The type of T. speciosum (and

of/, neglecta Schult.) is a plate in Andrews (1801 ) titled

Ixia bulbocodium var. flore speciosissimo , a polynomial.

Ker Gawler ’s Trichonema speciosa is based solely on

the Andrews illustration and not, as De Vos and others

believed, on the illustration in Curtis s Botanical Maga-

zine-. t. 1476 (Ker Gawler 1812), which is the type that

De Vos cited for R. neglecta. This plate is, nevertheless,

evidently the same species (and probably the same plant

or its descendants) as was illustrated by Andrews. Baker

(1877) provided the combination R. speciosa , citing Ixia

bulbocodium var. speciosa Andrews (in error for the

polynomial listed above), and Ker Gawler ’s Trichonema

speciosa. In addition. Baker listed in synonymy R. tabu-

laris Eckl., at the time a manuscript name on specimens

distributed by Ecklon, later R. tabularis Eckl. ex Beg.

in 1907. Baker also correctly listed in synonymy, Ixia

neglecta Schult.

The type of Trichonema speciosa and Ixia neglecta

is a painting of a red-flowered Romulea , the dark tepal

cup pale at the base and the reverse of the outer tepals

boldly striped with pale and dark longitudinal lines. The

poorly rendered conn seems consistent with R. neglecta

but the plant is illustrated with one basal and one sub-

basal leaf. The bracts are consistent with R. neglecta

as understood by De Vos, the inner with membranous,

darkly streaked margins. Andrews’s plant was provided

in 1801 by George Hibbert, Member of Parliament and

patron of botany, who we suspect received conns from

James Niven. Hibbert had commissioned Niven in 1798

to collect plants for him at the then Cape Colony and

Niven is known to have visited the Kamiesberg in 1799,

where he collected specimens of the endemic Moraea

longiflora Ker Gawk, which was illustrated in Curtis’s

Botanical Magazine', t. 712 (Ker Gawler 1804; Goldblatt

1986: 104). Thus this early illustration of a then remote

Namaqualand endemic is not unprecedented.

The presence of two, rather than a single basal leaf,

is not consistent with Romulea neglecta , which usually

has a single basal leaf, but the anomaly may be the result

of its cultivation in England or of a misrepresentation by

the artist. The plant illustrated later in Curtis’s Botani-

cal Magazine however, also from stock derived from

Hibbert's introduction, has a single basal leaf. Because

of this inconsistency, we designate an epitype for R. spe-

ciosa, the name that must now be applied to the species.

As the type collection of R. oliveri still represents the

276

Bothalia 41,2 (2011)

only actual specimen of R. speciosa that is known, we

designate this collection as the epitype.

Romulea speciosa (Ker Gawl.) Baker in Journal

of the Linnean Society, Botany 16: 89 (1877). Tricho-

nema speciosum Ker Gawl.: 223 (1804). Ixia neglecta

Schult.: 279 (1822), nom. nov. pro Trichonema spe-

ciosum Ker Gawl. [non Ixia speciosa Andrews (1801)].

Bulbocodium speciosum (Ker Gawl.) Kuntze: 700

(1891). Type. South Africa, without precise locality,

possibly collected by J. Niven, illustration in Andrews

(1801), The botanist’s repository 3: t. 170 ‘ Ixia bulboc-

odium var . flore speciosissimo (1801). Epitype: South

Africa, [Northern Cape], Kamiesberg, Farm Welkom, 14

Sept. 1970, Oliver 3169 (NBG, epi. ! ; PRE, iso.).

R. oliveri M.P.de Vos: 116 (1972). Type: South Africa, [Northern

Cape], Kamiesberg, Farm Welkom, 14 Sept. 1970, Oliver 3169 (NBG,

holo.!; PRE, iso.).

R. neglecta M.P.de Vos: 29 (1983), horn, illegit., non R. neglecta

Jord. & Fourr. (1868). Type: South Africa, without precise locality,

possibly collected by J. Niven, illustration in Curtis’s Botanical Maga-

zine 14: t. 1476 ' Ixia speciosa' ( Ker Gawler 1812).

ACKNOWLEDGEMENTS

We thank Roy Gereau, Missouri Botanical Garden,

and Kanchi Gandhi, Harvard University, for invaluable

assistance with nomenclatural questions and Mary Stif-

fler, research librarian, Missouri Botanical Garden, for

cheerfully providing references not available electroni-

cally. Type material was collected under a permit from

CapeNature.

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Bothalia 41,2: 277-282 (2011)

FSA contributions 19: Asteraceae: Anthemideae: Eumorphia

N. SWELANKOMO*

INTRODUCTION

The genus Eumorphia DC. belongs to the tribe

Anthemideae in the subfamily Asteroideae of the family

Asteraceae. It consists of six species and two subspecies

(Hilliard & Burtt 1982), which are endemic to southern

Africa. It occurs widely in southern Africa in Limpopo,

Mpumalanga, Swaziland, Free State, KwaZulu-Natal.

Lesotho, Western and Eastern Cape (Herman et al.

2003).

MATERIAL AND METHODS

A total of seven Eumorphia taxa were studied from

herbarium specimens housed in PRE. In each species,

morphological characters were investigated. The length

of the corolla, length and width of the involucre, length

of the style and ovary, number of ray florets, where pos-

sible, and length of cypselae were measured with the aid

of a Bausch & Lomb StereoZoom® Series microscope.

Two cypselae in each taxon, except in the case of E.

swaziensis and E. corymbosa, were examined by means

of a scanning electron microscope (JSM-840, JEOL,

Tokyo, Japan). Anthers were studied under a dissecting

and light microscope and photographs were taken with

a digital camera (Canon Powershot S80 Digital Camera)

mounted on the light microscope (Olympus Vanox-S).

Type specimens seen on the website are distinguished by

the code e! in citations below. Acronyms for herbaria are

listed in Holmgren et al. (1990) and in Smith & Willis

(1999).

Eumorphia DC., Prodromus 6: 2 (1838); Harvey:

153 (1865); Kallersjo: 538 (1985); Bremer & Hum-

phries: 94 (1993); Bremer: 448 (1994); Herman et al:

137 (2000); Oberprieler et al.: 355 (2007); Oberprieler et

al.: 643 (2009). Type species: E. dregeana DC.

Shrubs or shrublets, glabrous or pubescent. Leaves

opposite, alternate in E. davyi and E. swaziensis, eri-

coid, sometimes clustered, entire or lobed. Capitula

heterogamous, radiate, terminal, solitary or corymbose

in E. corymbosa and E. swaziensis, shortly peduncu-

late or subsessile. Involucre campanulate to subglo-

bose or cylindrical in E. corymbosa. Involucral bracts

in 3-5 rows, outer sometimes with 2-5 glands on back

near apex, inner ciliate near apex, sometimes all with

broad, membranous apices. Receptacle flat or con-

vex, rarely conical, paleate or sometimes epaleate in E.

davyi ; paleae longer than disc florets, linear, sometimes

with bent, apical appendage. Ray florets female, fer-

tile; corolla white sometimes flushed rose-pink, lamina

* Biosystematics Research and Biodiversity Collections Division,

South African National Biodiversity Institute, Private Bag XI 01, Pre-

toria, 0001. E-mail: N.Swelankomo@sanbi.org.za.

MS. received: 2011-02-23.

oblong to linear, 3-toothed, 2-4 times as long as tube,

glandular. Style slightly branched to branched with stig-

matic areas along edges. Disc florets bisexual, fertile,

yellow; corolla tube sometimes campanulate above, with

5 ovate lobes. Anthers minutely caudate with linear, api-

cal appendages; endothecial tissue polarized. Style lin-

ear, branched, truncate, branches with stigmatic areas

along edges. Cypsela ribbed, non-myxogenic, cylindri-

cal, minutely papillose especially on ribs, with dentate

or entire apical rim. Pappus absent. Figure 1 A-C.

Etymology r. the generic name Eumorphia is derived

from the Greek words, eu- meaning well and morphe

meaning form; referring to the neatness of the foliage

(Jackson 1990: 91 ).

Conservation status: according to the latest Red List

of South African plants (Raimondo et al. 2009), all spe-

cies are listed as LC (Least Concern).

Key to species

la Capitula corymbosely arranged, < 5 mm diam.:

2a Leaves opposite, narrowly linear, entire 1 . E. corymbosa

2b Leaves alternate, mostly trifid just above middle of stem

2. E. swaziensis

lb Capitula solitary, > 5 mm diam.:

3a Leaves alternate 3. E. davyi

3b Leaves opposite:

4a Subshrub up to 0.6 m tall, receptacle epaleate or with few

marginal paleae 4. E. prostrata

4b Erect shrubs to 1m tall, receptacle paleate:

5a Leaves linear-terete, closely imbricate, up to 3.5 mm

long 5. E. dregeana

5b Leaves entire or 3-lobed, up to 8 mm long 6. E. sericea

1. Eumorphia corymbosa E. Phillips in Journal

of South African Botany 16: 20 (1950). Type: Western

Cape, 3222 (Beaufort West): on top of Molteno Pass,

5500' [1804 m], (-BA), May, J.P.H. Acocks 14340,

(PRE, holo.!).

Woody subshrub, 0. 1-1.0 m high, single-stemmed

at base becoming branched, glabrous; stem very rough.

Leaves opposite, narrowly linear, entire, upper 4-5 x 1 .2

mm, lower 10-18 x 1.2 mm, some clustered at nodes,

acuminate. Capitula heterogamous, radiate, small, 5 x

2.5 mm when pressed, corymbose at ends of branches,

shortly pedunculate. Involucre cylindrical; involucral

bracts in 5 rows, imbricate, glabrous; outer ovate. 1 x

0.5 mm, membranous along edges; middle ovate-lanceo-

late, l.lxl mm, membranous along edges; inner 5 x 1

mm, lacerate, tips membranous. Receptacle flat, paleate;

paleae linear-lanceolate, 2 mm long, almost as long as

disc florets. Ray florets female, white; corolla limb glan-

dular. 3 mm long, 3-toothed; corolla tube shorter than

limb, sparsely glandular. Style terete, 1.5 mm long,

branches linear, obtuse, 0.4 mm long. Ovary ellipsoid,

ribbed, glabrous. Disc florets bisexual, fertile, yellow;

corolla tube 1.2 mm long, glandular; corolla limb 1

mm long, glandular, 5-lobed; lobes ovate, 0.2 mm long.

278

Bothalia 41,2 (201 1 )

FIGURE 1, — A-C, scanning electron micrographs of cypsela of E.

davyi, Brusse 5010: A, surface with papillae especially on ribs;

B, surface at higher magnification; C, showing apical rim.

Anthers linear, minutely caudate, with lanceolate api-

cal appendage. Style terete, branches linear, truncate, 2

mm long. Ovary’ 1.2 mm long, ± ellipsoid, slightly stri-

ate, glabrous. Cypsela 2 mm long, glabrous, cylindri-

cal, minutely papillose especially on ribs, crowned with

thickened apical rim. Pappus absent. Flowering time :

Feb.-Mar.

Distribution and habitat'. Eumorphia corymbosa is

endemic to the Western Cape in the Beaufort West Divi-

sion, from the National Park, Mountain View, down to

the foot of the Nuweveld Mtns as far as Molteno Pass

(Figure 2), at I 000-1 828 m. It grows in seasonal wet

soils along rivers and in marshy habitats.

Notes'. Eumorphia cotymbosa can be distinguished

from other species in the genus by the small corym-

bosely arranged capitula at the branch endings; cylindri-

cal involucre and opposite leaves.

Other specimens examined

WESTERN CAPE. — 3222 (Beaufort West): Karoo National Park,

Mountain View, (-BA), 2 Jan. 1985, Shearing 828 (PRE); near Abra-

hamspoort Farm, on top of Nuweveldsberg Plateau, (-BA), 9 Mar. 1990,

Viok 2276 (PRE); foot of Nuweveld Mtns, on bank of small river flow-

ing at foot of mountain, (-BA), 15 Feb. 1978, Gibbs Russell. Robinson.

Herman & Downing 235 (PRE); Mountain View Farm, among dolerite

boulders at top of hill, (-BA), 14 Feb. 1978, Gibbs Russell, Robinson.

Herman & Downing 124 (PRE); Matjieskloof, 24 miles [38.4 km] north

of Beaufort West, (-BA), 16 May 1950, Acocks 15893 (PRE).

2. Eumorphia swaziensis Compton in Journal of

South African Botany 33: 300 (1967). Type: Swaziland,

2631 (Mbabane): hill northeast of Mbabane, (-AC),

4000' [I 312 m], 15 Mar. I960, R.H. Compton 29963,

(SDNH, holo. e!; PRE, two sheets!, NBG e!, iso.).

Small, mainly glabrous shrub, 0.3-0. 6 m high, with

erect woody stems, densely branched in upper florif-

erous parts. Leaves alternate, very numerous, erecto-

patent, mostly trifid just above the middle; rachis and

lobes filiform, fle.xuous, apiculate, lobes subopposite,

up to 20 mm long; many leaves in upper parts of stems

undivided, filiform. Inflorescence irregularly corymbose,

capitula numerous, erect, each borne on short pedun-

cle, subtended by short linear bract; peduncle and bract

finely and sparsely hairy. Involucre ±5x2 mm, com-

pact, campanulate, glabrous. Involucral bracts'. 3 rows,

imbricate; outer ovate, 1.2 x 0.5 mm; middle 2.2 x 1

mm; inner 3.5 x 1.1 mm, with narrowly membranous-

fimbriate margins. Receptacle paleate; paleae mem-

branous, linear, slightly shorter than disc florets. Ray

florets 5, female, white; corolla limb broadly oblong to

ovate, ±3x2 mm, shortly 3-lobed, glabrous; corolla

tube sparsely glandular, 1 mm long. Style linear, slightly

curved, 0.1 mm long with stigmatic areas along edges.

12 14 16 18 20 22 24 26 28 30 32

FIGURE 2. — Known distribution of Eumorphia corymbosa , ■; E.

davvi , •; E. dregeana , *; E. prostrata , A; and E. swaziensis,

+ .

Bothalia 41,2 (2011)

279

Ovary 1.8 mm long, slightly striate. Disc florets bisex-

ual, 5-lobed, yellow; lobes ovate, 0.1 mm long; corolla

glandular; corolla limb 1 mm long; corolla tube 1

mm long. Anthers minutely caudate, with short api-

cal appendages; filaments thickened. Style 2 mm long,

branched, style branches linear, truncate, 0.15 mm long,

with stigmatic areas along edges. Ovary linear, 1.1 mm

long, slightly striate. Cypsela: mature ones not seen.

Pappus absent. Flowering time : Mar.-Apr.

Distribution and habitat : Eumorphia swaziensis, as

the name suggests, is endemic to Swaziland, where it

occurs in grassland on the hilltops, at 1 219-1 372 m

around Mbabane (Figure 2).

Notes : Eumorphia swaziensis can be distinguished

from other species in the genus by having alternate

leaves; irregularly corymbose inflorescences and cam-

panulate involucres.

Specimen examined

SWAZILAND. — 2631 (Mbabane): Black Umbuluzi Valley, (-AC),

24 Mar. 1978, Kemp 1304 (PRE).

3. Eumorphia davyi Bolus in Transactions of the

South African Philosophical Society 16: 387 (1906);

Retief & Herman: 306 (1997). Type: Eastern Transvaal

[Mpumalanga], 2430 (Pilgrim’s Rest): Graskop, near

Pilgrim’s Rest, (-DD), 29 Jan. 1906, J. Burn Davy 1474,

(BOL, holo. e!; PRE!, K e! , iso.).

Ericoid shrub, 0.18-2.0 m high, single-stemmed at

base becoming branched only in upper third, forming

a compact, rounded bush, glabrous. Stem very rough.

Leaves alternate, imbricate, up to 1 1 x 0.5 mm, glabrous,

acute, mucronulate. Capitula shortly pedunculate or sub-

sessile, radiate. Involucre campanulate, 5-6 x 6 mm,

finely hairy. Involucral bracts in 3 rows, imbricate; outer

narrow, 3^1 x 0.5 mm, slightly hairy; middle lanceolate,

4- 6 x 1 mm, acuminate, lacerate membranous; inner

with ovate apex, 6-7 x 1 mm, membranous, discolor-

ous. Receptacle conical, epaleate. Ray florets female,

fertile, white; corolla limb oblong, 3-toothed, 7 mm

long; corolla tube 2 mm long. Style up to 1.8 mm long,

linear, branched, style branches 0.2 mm long, round

with stigmatic areas along edges. Ovaiy 3 mm long,

slightly striate. Disc florets many, bisexual, fertile, yel-

low; corolla limb 1 mm long; corolla tube 0.8 mm long,

5- lobed; lobes ovate, 0.15 mm long. Anthers minutely

caudate, apex round; filaments thickened. Style linear,

1.8 mm long, branched, style branches 0.2 mm long,

truncate with stigmatic areas along edges. Ovary 1.8

mm long. Cypsela closely ribbed, slightly curved, 2.2

mm long, glabrous but minutely papillose especially on

ribs, crowned with thickened apical rim. Pappus absent.

Flowering time : Sept.-April.

Distribution and liabitaf. Eumorphia davyi is found

in Mpumalanga at Graskop, Pilgrim's Rest and as far

north as Mariepskop, and in Limpopo Province in

Polokwane [Pietersburg] and The Downs (Figure 2) at

1 200-2 286 m. E. davyi grows in grassveld.

Notes : Eumorphia davyi is named in honour of

Joseph Burtt Davy (1870-1940), a British botanist,

founder of the Pretoria National Herbarium, a student

of the flora of California, agriculturist, sylviculturist and

ecologist (Stafleu & Richard 1976). E. davyi can be dis-

tinguished from other species in the genus by alternate

leaves and a conical, epaleate receptacle.

Other specimens examined

LIMPOPO. — 2430 (Pilgrim’s Rest): The Downs, (-AA), Nov.

1918, Rogers 21557, 22003 (PRE).

MPUMALANGA. — 2430 (Pilgrim’s Rest): at top of Mariepskop,

(-DB), 15 Jan. 1960, Van der Schijff 4835 (PRE); Mount Sheba, road up

mountain from Mount Sheba Hotel to main Lydenburg-Pilgrims’s Rest tar

road, (-DC), 17 Apr. 1987, Brusse 5010 (PRE); Graskop, (-DD), 28 Mar.

\919, Kluge 1845 (PRE).

4. Eumorphia prostrata Bolus in Transactions of

the South African Philosophical Society 16: 388 (1906);

Hilliard: 332 (1977). Type: Eastern Cape, 3028 (Matatiele):

Barkly East, summit Doodman’s Krans Mtn, (-CA), 8 Mar.

1904, E.E. Galpin, 6700. (BOL, holo. e!; K e! , PRE, two

sheets!, SAM e!, iso.).

Low, erect, spreading subshrub, up to 0.6 m high,

often prostrate over rock sheets or hanging down cliff

faces; branches sericeous, soon glabrous, old ones rough

with leaf scars, often rooting, twigs closely leafy. Leaves

opposite, sericeous, silvery, hairs appressed, primary

leaves linear or 2- or 3-forked at tips, up to 1 0 x 1.5 mm,

appearing fascicled from closely leafy axillary shoots.

Capitula solitary, terminal, shortly pedunculate, radiate.

Involucre campanulate, up to 7 x 9 mm when pressed,

villous. Involucral bracts in 3 rows, imbricate; outer

narrowly ovate, 2. 5-5. 5 x 0.5 mm; middle obtuse, 5.5-

6.0 x 1.1 mm, tips membranous; inner oblong, 6. 5-7.0 x

1.1 mm, discolorous, membranous. Receptacle epaleate

or with a few marginal paleae. Ray florets female,

3-toothed, white tinged pink at base or sometimes at

back; corolla limb 5-7 mm long; corolla tube 1.8 long.

Style up to 2.2 mm long, branched, style branches 0.8

mm long, with stigmatic areas along edges. Ovary 1.2

mm long, slightly striate. Disc florets bisexual, 5-lobed,

yellow; lobes ovate, 0.1 mm long; corolla limb 2 mm

long; corolla tube 1 mm long. Anthers minutely caudate,

apical appendage elongated, obtuse, filaments thicken-

ed. Style up to 2.8 mm long, slightly branched, style

branches 0.2 mm long with stigmatic areas along edges

and papillate appendages. Ovary 1.1 mm long. Cypsela

narrowly cylindric, 2 mm long, closely ribbed, minutely

papillose especially on ribs, crowned with thickened api-

cal rim. Pappus absent. Flowering time'. Dec.-Apr. Fig-

ure 3.

Distribution and habitat'. Eumorphia prostrata is only

known from a small area of the high Drakensberg, from

Tina Head (Doodman's Krans and Naudesnek) where

the boundaries of Lesotho, the Barkly East and Maclear

Districts meet, to Thamathu Pass (Bushman’s Nek area)

and the Drakensberg Garden area in Underberg District,

KwaZulu-Natal (Hilliard 1977), at 1 830-2 970 m (Fig-

ure 2). In its natural environment, it appears to tolerate

extreme cold (-10°C) in winter, strong buffeting winds

and fairly high daytime temperatures in summer (Oliver

1989). E. prostrata grows in mountain grassveld in sea-

sonally wet areas.

Notes', although it seems close to Eumorphia seri-

cea, this species has a prostrate habit and a receptacle

epaleate or with a few marginal paleae.

280

Bothalia 41,2 (2011)

FIGURE 3. — Eumorphia prostrata. A, habit; B, leaf surface; C, ray floret; D, disc floret, Hoener 1722. Scale bar; A, 10 mm; B-D, 0.33 mm. Artist:

Gillian Condy.

Specimens examined

KWAZULU-NATAL. — 2929 (Underberg): Bushman’s Nek, Tha-

mathu Pass, (-CC), 4 Nov.1976, Hilliard & Burtt 8930 (PRE); vicin-

ity of Tarn Cave above Bushman’s Nek, (-CC), 20 Jan. 1984, Hilliard

& Burtt 17400 (PRE); Thamatuwe (= Thamathu) Pass, (-CC), 17

Jan. 1 969, Killick & Vahrmeijer 3996 ( PRE).

LESOTHO. — 2929 (Underberg): Sehlabathebe National Park,

Matsa a Mafikeng, (-CC), 16 Dec. 1967, Hoener 1722 (PRE).

EASTERN CAPE. — 3028 (Matatiele): Rhodes gravel road/jeep

track between Naudesnek and Ben Macdhui, (-CA), 12 Dec. 1999,

Koekemoer 1632 (PRE).

5. Eumorphia dregeana DC., Prodromus 6: 3

(1838); Harv.: 153 (1865). Type: Eastern Cape, 3124

(Hanover): Middelburg, Sneeuweberg between Kompas-

berg and Rhenosterberg, 5000-6000' [1 640-1 964 m],

(- DC), Sept., Drege s.n., (G-DC, holo. e! ; MO e!, M el,

PRE!, SAM e!,Ee!,Pe!,K e!, NBG e!, iso.).

Multi-stemmed, erect subshrub, 0. 3-1.0 m high.

Leaves linear-terete, closely imbricate, opposite, slightly

pubescent, up to 3.5 mm long, ± 1 mm broad, entire.

Capitula shortly pedunculate, radiate, solitary. Involu-

cre campanulate, 4-7 mm wide when pressed. Involu-

cral bracts in 3 rows, imbricate; outer ovate, glabrous

or slightly pubescent, 2 x 1 mm; middle 2.2 x 1.2 mm;

inner 2.5 x 1.25 mm, obtuse with membranous tips.

Receptacle paleate; palea membranous, linear, 4 mm

long. Ray florets female, 3-toothed, white; corolla limb

5 mm long; corolla tube 1 mm long. Style up to 1.2 mm

long, slightly branched, style branches 0.8 mm long,

with stigmatic areas along edges. Ovary 2 mm long,

slightly striate. Disc florets bisexual, fertile, 5-lobed, yel-

low; lobes ovate, 0.2 mm long; corolla limb glandular,

1.2 mm long; corolla tube glandular, I mm long. Anthers

minutely caudate, apical appendage elongated, obtuse.

Style up to 1 .2 mm long, branched, style branches 0.25

mm long with stigmatic areas along edges and pap-

illate appendages. Ovaty 1.5 mm long, slightly stri-

ate. Cypsela closely ribbed, 1.5-1. 8 mm long, slightly

curved, glabrous but minutely papillose especially on

ribs, crowned with thickened apical rim. Pappus absent.

Flowering time : Feb.-Apr.

Distribution and habitat'. Eumorphia dregeana occurs

in the southeastern Free State and the northwestern

Eastern Cape, at 1 311-1 980 m (Figure 2). According

to specimens in the National Herbarium, E. dregeana

grows in False Mountain Renosterveld (Karoo Escarp-

ment Grassland), on sandy slopes and in vleis.

Notes'. Eumorphia dregeana is named in honour of

Jean Franyois (Johann Franz) Drege (1794-1881), a Ger-

Bothalia 41,2 (2011)

281

man plant collector in South Africa (1826-1834) (Stafleu

& Richard 1976). E. dregeana can be distinguished from

other species in the genus by having closely imbricate,

opposite leaves and paleate receptacles.

Specimens examined

FREE STATE. — 2927 (Maseru): Thaba Phatshwa Mtn, south of

Tweespruit, high up on eastern slope of mountain, (-AC), 3 Apr. 1973,

Muller 1163 (PRE).

EASTERN CAPE. — 3124 (Hanover): Cradock, Wapadsberg, pla-

teau, (-DD), 28 Mar. 1947, Acocks 13552 (PRE). 3125 (Steynsburg):

Middelburg, Leeukop Farm, (-DB), 12 Mar. 1950, Theron 746 (PRE).

3224 (Graaff-Reinet): Oudeberg, (-DD), Apr. 1890, Bolus 1063 (PRE).

NORTHERN CAPE. — 3124 (Hanover): Noupoort, 11 km from

Noupoort on road to Colesberg via Oorlogspoort, (-BB), 3 Apr. 1981,

Herman 533 (PRE).

6. Eumorphia sericea J.M.Wood & M.S. Evans

in Journal of Botany 35: 488 (1897); Grierson: t. 608

(1972); Hilliard: 332 (1977). Type: Natal, summit of

Drakensberg, near Bushman's River Pass, 8000-10 000'

[2 624-3 280], Apr. 1896, M.S. Evans 751, (NH, holo.;

PRE!, K, iso.).

Key to subspecies of Eumorphia sericea

6a Plant with thick, silky, woolly indumentum, cypsela crowned by

a thickened slightly lobulate rim 6a. E. sericea subsp. sericea

6b Plant more robust with thinner indumentum, pappus scales

forming a corona on cypsela .... 6b. E. sericea subsp. robustior

6a. subsp. sericea

Much-branched, dwarf shrub, 0.2-1. 3 m high; outer

branches sometimes decumbent, nude below, rough

with leaf scars, grey (because of its thick indumentum)

silky-woolly, glabrescent, twigs closely leafy. Leaves

opposite, grey silky-woolly, hairs fluffed up from leaf

surface, primary leaves linear or digitately 3-lobed, ± 8

x 1.5 mm, appearing fascicled from closely leafy, dwarf

axillary shoots. Capitula solitary, shortly pedunculate,

radiate. Involucre campanulate, ± 7 x 10 mm when

pressed, villous. Involucral bracts : 3 rows, imbricate;

outer 3x1.5 mm, ovate; middle 4x1.5 mm; inner 6.5 x

1.5 mm, obtuse with scarious and lacerate tips. Recepta-

cle paleate; paleae membranous, narrowly lanceolate, ±

5 x 0.5 mm. Ray florets female, 3-toothed usually white,

but sometimes rose-pink, or white flushed with pink;

corolla glandular; corolla limb 10.5 mm long; corolla

tube up to 1.8 mm long. Style 3 mm long, branched,

style branches 0.3 mm long with stigmatic areas along

edges. Ovary 2.2 mm long, slightly striate. Disc florets

bisexual, glandular, 5-lobed, yellow; lobes ovate, 0.1

mm long; corolla limb 2 mm long; corolla tube 1 mm

long. Anthers minutely caudate, apical appendages elon-

gated, obtuse; filaments thickened. Style up to 2.5 mm

long, branched, style branches 0.18 mm long with stig-

matic areas along edges. Ovary 2.2 mm long, slightly

striate. Cypsela narrowly cylindric, ± 2.5 mm long,

closely ribbed, glabrous but minutely papillose espe-

cially on ribs, crowned with thickened apical rim. Pap-

pus absent. Flowering time : Nov.-Apr.

Distribution and habitat'. Eumorphia sericea subsp.

sericea occurs in Free State, KwaZulu-Natal, Lesotho

and Eastern Cape Province, at 2 296-3 414 m (Figure

4). E. sericea subsp. sericea grows in seasonally wet

areas.

Note', although close to E. prostrata , it has paleate

receptacles.

Specimens examined

FREE STATE. — 2828 (Bethlehem): Mont-aux-Sources, summit

above chain ladder, (-DB), 14 Jan. 1982, Roux 1083 (PRE).

KWAZULU-NATAL. — 2929 (Underberg): top of Sani Pass, (-CB),

23 Mar. 1962, Acocks 22159 (PRE).

LESOTHO. — 2928 (Marakabeis): Tsienyane Valley, over Menia-

meng Pass, (-AA), 8 Jan. 1955, Coetzee 484 (PRE). 2929 (Under-

berg); Castle Buttress, on summit of Drakensberg, (-AA), 18 Dec.

1952, Killick 1873 (PRE).

EASTERN CAPE. — 3028 (Matatiele): Barkly East, Drakensberg,

Doodman’s Krans Mtn, (-CA), 7 Mar. 1904, Galpin 6699 (PRE).

6b. subsp. robustior Hilliard & B.L.Burtt in Notes

from the Royal Botanic Garden Edinburgh 40: 248

(1982). Type: Eastern Cape, 3127 (Lady Frere): Elliot,

Fetcani Pass, common bush up to 5' [1.64 m], on slopes

and especially in rock-fall scrub, (-BB), 7500' [2 460

m], 22 Jan. 1979, O.M. Hilliard & B.L. Burtt 12331

(NU, holo.; E, PRE!, K, M, S, NBG, iso.).

Green bush because of its thinner indumentum on

leaves, more robust, hence its name, pappus scales form-

ing a corona on cypsela. Flowering time : Dec-May.

Distribution and habitat'. Eumorphia sericea subsp.

robustior is found in Naudesnek down to Spitskop Farm,

at 2 073-2 755 m (Figure 4). This subspecies grows in

damp areas.

Specimens examined

EASTERN CAPE. — 3027 (Lady Grey): Barkly East, Rhodes to

Naudesnek, at foot of cliff, (-DD), 22 Feb. 1971, Hilliard & Burtt

6732 (PRE); Rhodes, roadside between Ugie and Barkly East, (-DD).

10 May 1985, Hutchings 1706 (PRE). 3028 (Matatiele): Barkly East,

Naudesnek, (-CA), 28 Nov. 1971, Hilliard 5213 (PRE); Naudesnek,

between Maclear and Rhodes, (-CC), 7 Dec. 1983, Van Wvk 6713

(PRE). 3127 (Lady Frere): Elliot, bastervoetpad, rocky ground on hill-

side and along stream, (-BB), 15 Feb. 1983, Hilliard & Burtt 16657

(PRE).

28 30 32

FIGURE 4. — Known distribution of Eumorphia sericea subsp. sericea ,

3; and subsp. robustior , •.

282

Bothalia 41,2 (201 1 )

ACKNOWLEDGEMENTS

The Microscopy Unit of the University of Pretoria is

thanked for the use of their Scanning Electron Micro-

scope, and my fellow SANBI colleagues, namely P.

Herman for guidance and discussions, Gillian Condy

for kindly preparing the line drawing, Elizma Fouche

for her assistance with SEM plates and Hester Steyn for

producing the distribution maps.

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Bothalia 41,2: 283-288 (2011)

Review of the genus Xenoscapa (Iridaceae: Crocoideae), including X,

grandiflora , a new species from southern Namibia

J.C. MANNING* and P. GOLDBLATT**

Keywords: Iridaceae, new species, southern Africa, taxonomy, Xenoscapa (Goldblatt) Goldblatt & J.C. Manning

ABSTRACT

The small genus Xenoscapa (Goldblatt) Goldblatt & J.C.Manning, endemic to the southern African winter rainfall region,

is reviewed. The new species X. grandiflora is described from the deeply dissected southern part of the Huib Hoch Plateau

in southern Namibia. It differs from the two known species in the genus in its significantly larger, pale lilac flowers. Full

descriptions and accounts of all three known species are provided, with distribution maps and illustrations.

INTRODUCTION

Xenoscapa (Goldblatt) Goldblatt & J.C.Manning, one of

the smallest genera in Iridaceae, currently comprises two

species from the winter rainfall region of southern Namibia

and southwestern South Africa. Both are small, deciduous

geophytes with two or three, soft-textured, prostrate foli-

age leaves and unusual, single-flowered, mostly shortly

branched spikes (Goldblatt & Manning 1995, 2008). The

vegetative similarity between them extends to the flowers,

which are long-tubed with relatively small, narrow tepals

and shortly exserted. arcuate stamens.

The distinctive combination of single-flowered spikes,

tubular flowers, and deeply divided style branches caused

a great deal of uncertainty about the taxonomic position

of the genus. The slender perianth tube and divided style

branches caused the first known species, Xenoscapa fis-

tulosa (Spreng. ex Klatt) Goldblatt & J.C.Manning, to be

described in the genus Ovieda Spreng. (an illegitimate later

synonym of Lapeirousia Pourr.), from which it was sub-

sequently transferred to Lapeirousia (Baker 1877). The

rounded conns with fibrous tunics are anomalous there,

however, and it was accordingly moved to Anomatheca

as the monotypic section Xenoscapa (Goldblatt 1972).

A later morphological cladistic analysis indicated that

the species was also misplaced here, combining a unique

mix of mainly apomorphic character states, notably the

unusual inflorescence, reduced leaf number and cylin-

drical capsules, containing plesiomorphic, angled seeds.

As a result, Xenoscapa was recognized as an independ-

ent genus, and increased to two species with the addition

of a new species from central Namaqualand, X. uliginosa

Goldblatt & J.C.Manning. The precise relationships of

Xenoscapa remained uncertain until recent analysis of

plastid sequence data placed it as sister to the clade com-

prising Devia-Crocosmia-Freesia (Goldblatt et al. 2006).

The four genera are currently treated as the tribe Freesieae

(Goldblatt & Manning 2008).

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town / Research Centre for

Plant Growth and Development, School of Biological and Conservation

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

X01, Scottsville 3209, South Africa.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

den, P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2011-02-01.

The two known species of Xenoscapa are distin-

guished by small differences in perianth size and colour,

height of the flowering stems in fruit, and the presence

or absence of floral fragrance (Table 1 ). X. fistulosa is

relatively widespread, occurring throughout the range

of the genus, from the Huib Hoch Plateau in southern

Nambia southwards along the Namaqualand escarpment

and the interior mountains of the southwestern Cape,

with two outlying populations along the West Coast

(Goldblatt & Manning 2000a). X. uliginosa , in contrast,

is a highly local endemic restricted to middle and upper

elevations of the Kamiesberg, where it may co-occur

with X. fistulosa (Goldblatt & Manning 1995). A recent

collection from the summit of Hohenzollem Peak in

the Hunsberge in southern Namibia, slightly east of the

recorded range of X. fistulosa , is distinctive in its sig-

nificantly larger, pale lilac flowers with relatively larger

anthers, and represents a third species, described here

as X. grandiflora. We take this opportunity to provide a

review of the genus Xenoscapa.

MATERIALS AND METHODS

This study is based on an examination of living plants

in the field and of specimens in the following herbaria:

BOL, MO, NBG, PRE and SAM (acronyms according

TABLE 1. — Distinguishing morphological characters mXenoscapa

284

Bothalia 41,2 (2011)

to Holmgren et al. 1990). The abbreviation of author

names follows Brummitt & Powell (1992).

TAXONOMY

Xenoscapa (Goldblatt) Goldblatt & J.C. Manning

in Systematic Botany 20: 172 (1995). Anomatheca sect.

Xenoscapa Goldblatt: 88 (1972). Type species: X. fistu-

losa (Spreng. ex Klatt) Goldblatt & J.C. Manning.

Seasonal perennials with small, globose conn, rooting

from the base and axial in origin; tunics of finely retic-

ulate fibres. Cataphylls 2 or 3, pale and membranous.

Leaves 2 or 3, unifacial, prostrate, soft-textured, with

definite midrib; margins with columnar epidermal cells.

Stem short or long, erect, terete, often with 1-3 short

branches. Inflorescence of solitary, sessile flowers termi-

nal on axes; bracts green, leathery, inner bracts slightly

shorter or more often slightly longer than outer, often

notched apically. Flowers zygomorphic, tubular or sal-

ver-shaped, either cream-coloured and sweetly scented

or pink with contrasting markings on lower tepals and

unscented, with nectar from septal nectaries; perianth

tube cylindric and elongate; tepals subequal, spreading

or dorsal erect, slightly larger, and spoon-shaped. Sta-

mens unilateral and arcuate, filaments shortly exserted.

Ovary globose, sessile; style filiform, with short, deeply

divided, recurved branches. Capsules oblong to cylin-

dric, cartilaginous. Seeds strongly angled, with promi-

nent chalazal crest, slightly wrinkled, matte, surface

colliculate-rugulose. Basic chromosome number x =11.

Three species, in the southern African winter rainfall

zone of southern Namibia and western South Africa.

Etymology, from the Greek, xenos, for strange and

scapa, for flowering stem, because of the inflorescence,

unusual for subfamily Crocoideae in bearing solitary

flowers on the main and lateral branches instead of

spikes of multiple flowers.

Key to species

la Flowers white or cream-coloured (rarely very pale pink),

without contrasting nectar guides but perianth tube often

flushed purple, fragrant; perianth tube 1.0-1 .5 mm diam.

at mouth; tepals 4—7 x 1-3 mm; anthers 1.0-2. 5 mm long;

capsules (8—) 1 2-1 8 mm long 3 .X.fistutosa

lb Flowers lilac to dark pink or purple with contrasting nectar

guides on lower tepals, unscented; perianth tube 1. 5-2.0

mm diam. at mouth; tepals 6-14 x 2. 0-5. 5 mm; anthers

2—4 mm long; capsules 10-13 mm long:

2a Flowers dark pink; perianth tube 22-28 mm long; tepals

6-7 x 2-3 mm; anthers ± 2 mm long; plants from

Kamiesberg in central Namaqualand 2. X. uliginosa

2b Flowers pale lilac; perianth tube 33-35 mm long; tepals

12-14 x 4. 0-5. 5 mm; anthers 3.5— 4.0 mm long; plants

from Hunsberge in southern Namibia \.X. grcmdiflora

1 . Xenoscapa grandiflora Goldblatt & J.C. Manning,

sp. nov.

Ilaec species quoad folia prostrata oblonga et capsu-

las oblongo-cylindricas Xenoscapae fistulosae etX. ulig-

inosae similis, sed ab eis floribus pallide lilacinis ino-

doris, tepalis iuferioribus dilute malvino-suffusis, tubo

perianthii 33-35 mm longo, et tepalis 12-14 x 4. 0-5. 5

mm differt.

TYPE. — Namibia, 2717 (Chamaites): Hunsberge, Ho-

henzollem Peak, (-CC), 1 July 2007 [fi. in cult. 13 July

2010], E. van Jaarsveld sub J.C. Manning 3303 (NBG,

holo.; MO, iso.).

Deciduous geophyte, 40-100 mm high, including

flowers. Corm subglobose, 6-8 mm diam.; tunics of

coarse, netted fibres. Leaves 3, basal, prostrate, oblong,

lower two largest, 45-60 x 1 5-20 mm, upper leaf ± half

as large. Stem terete, inclined at base but then stiffly

erect, up to 45 mm long, with up to 3 short branches

10-15 mm long, each subtended by a short cauline bract.

Inflorescence of solitary flowers on main and lateral

axes; bracts green, 8-9 mm long, inner slightly longer

than outer and often shortly forked apically. Flow-

ers zygomorphic with angle widest between upper and

lower lateral tepals, tinged pale lilac, darker on tube,

lower 3 tepals each with pale creamy yellow patch at

base outlined in purple, unscented; perianth tube erect,

cylindric, 33-35 mm long, wider and curved just below

apex, 2 mm diam. at mouth; tepals subequal but inner

slightly wider than outer, 12-14 x 4. 0-5. 5 mm, weakly

cucullate apically, dorsal erect, others spreading hori-

zontally at right angles to tube. Stamens : filaments 3-5

mm long, exserted 2-4 mm from tube; anthers 3. 5-4.0

mm long, dark purple; pollen grey-blue. Ovary ellip-

soid; style branching opposite anther bases, branches 4

mm long, deeply forked and apically recurved. Capsule

oblong-cylindric, 10-13 * 3^1 mm. Seeds ± 1 mm diam.

Flowering time\ June-July. Figure 1A-D.

Distribution and ecology r. known only from the type

collection, made on the summit of Hohenzollern Peak at

the southern end of the Hunsberge in southern Namibia

(Figure 2) on 1 July 2007 by horticulturist Ernst van

Jaarsveld. The plants were tightly wedged in crevices in

the shelter of rock outcrops immediately below the east-

ern edge of the summit plateau, and were just past flow-

ering. Corms that were collected then have subsequently

flowered in cultivation. Vegetatively, these plants match

photographs of wild plants taken at the time and there

is no reason to expect that the dimensions derived from

the cultivated specimens will differ in any meaning-

ful way from specimens in the wild. The Hunsberge,

in the deeply dissected southern part of the Huib Hoch

Plateau, are a recognized site of ecological importance

in Namibia, known for a high level of endemism among

vertebrates (Barnard et al. 1998).

Diagnosis and relationships'. Xenoscapa grandiflora

is distinguished by its large, unscented, pale lilac flow-

ers marked with purple chevrons on the lower tepals,

a perianth tube 33-35 mm long and the tepals 12-14 x

4. 0-5. 5 mm.

Xenoscapa grandiflora is most likely to be confused

with X. uliginosa from the Kamiesberg, both of which

have relatively larger, unscented, generally pink flowers

with darker chevrons on the lower tepals but the latter

has bright pink or purple flowers with a shorter perianth

tube, 22-28 mm long, and smaller tepals, 6-7 mm long,

with the dorsal tepal typically suberect or spreading to

expose the stamens and style rather than remaining erect

Bothalia 41,2 (201 1 )

285

FIGURE 1 . — Xenoscapa species: A-D,

X. grandiflora. Van Jaarsveld

sub Manning 3303 : A, flower-

ing plant; B, flower, front and

side views; C, capsules; D,

seed. E, F, X. uliginosa, Gold-

blatt & Manning 9244: E, flow-

ering plant; F, capsule. G, H, X.

fistulosa, Goldblatt & Manning

9374: G, flower and capsule; H,

capsule. Scale bar: 10 mm. Art-

ist: John Manning.

after anthesis (Figure IE). The common and widespread

X. fistulosa has smaller flowers with uniformly white or

cream-coloured (rarely pale pink) tepals (although the

tube is usually flushed maroon) and a distinct and notice-

able floral fragrance (described as sweetly clove-like).

Both Xenoscapa fistulosa and X. grandiflora are

self-fertile and autogamous, with all flowers setting full

capsules of seed that appear well formed (reproductive

biology is unknown in X. uliginosa). The unscented,

lilac flowers with elongate perianth tube and dark pur-

ple markings on the lower tepals of X. grandiflora are

consistent with pollination by long-proboscid flies, the

pollination system inX. uliginosa (Manning & Goldblatt

1996; Goldblatt & Manning 2000b).

2. Xenoscapa uliginosa Goldblatt & J.C. Manning

in Systematic Botany 20: 173 (1995). Type: South

Africa, [Northern Cape], Namaqualand, Farm Modder-

fontein. lower east-facing slopes of Sneeukop, ± 1 400

m, 18 Sept. 1981, Goldblatt [& Manning \ 9244 (NBG,

holo.!;K!,MO!,PRE!,iso.).

Deciduous geophyte, 30-50 mm high, including flow-

ers. Conn subglobose, 5-8 mm diam.; tunics of fine, net-

ted fibres. Leaves 3, basal, prostrate, oblong, lower two

largest, 20 — 40 x 5-18 mm, upper leaf ± half as large.

Stem terete, inclined at base but then stiffly erect, up

to 40 mm long, with up to 3 short branches 5-10 mm

long, each subtended by a short cauline bract. Inflores-

cence of solitary flowers on main and lateral axes; bracts

286

Bothalia 41,2 (2011)

16° 18° 20° 22° 24° 26° 28°.

FIGURE 2. — Known distribution of Xenoscapafistulosa, •; X. grandi-

flora , A;X. uliginosa, O.

green, 6-7 mm long, inner slightly longer than outer

and often shortly forked apically. Flowers zygomorphic,

dark pink or purple, darker on tube, lower 3 tepals each

with pale creamy yellow patch at base outlined in pur-

ple, unscented; perianth tube erect, cylindric, (20— )22 —

28 mm long, wider and curved just below apex, 1. 5-2.0

mm diam. at mouth; tepals subequal but inner slightly

wider than outer, 6-7 x 2-3 mm, weakly cucullate api-

cally, dorsal erect or suberect and exposing anthers. Sta-

mens'. filaments 3-5 mm long, exserted ± 2 mm from

tube; anthers ± 2 mm long, dark purple; pollen grey-

blue. Ovary ellipsoid; style branching opposite anther

bases, branches 2 mm long, deeply forked and apically

recurved. Capsule oblong, ± 10 x 3—4 mm. Seeds ± 1

mm diam. Flowering time : mid Sept.-Oct., rarely into

early Nov. Figure IE, F.

Distribution and ecology: restricted to the Kamies-

berg in central Namaqualand, and known originally only

from the Sneeuberg but recently collected in the Lang-

kloof east of Stalberg, at the foot of Rooiberg (Figure 2).

Xenoscapa uliginosa has a specialized habit along the

margins of seasonal seeps and in rock flushes. During

the growing season the roots are constantly irrigated by

water percolating through the mossy sward in which the

conns are anchored.

The long-tubed, bright pink flowers are pollinated by

the long proboscid fly Prosoeca peringueyi (Nemestrini-

dae) (Manning & Goldblatt 1996).

Diagnosis and relationships: Xenoscapa uliginosa is

recognized by its moderately large, dark pink or purple,

unscented flowers marked with purple chevrons on the

lower tepals, perianth tube 22-28 mm long, and tepals

6-7 x 2-3 mm. Plants from the Sneeuberg have the

dorsal tepal ± suberect to expose the anthers and style

but a recent collection from the Langkoof on the south-

ern flank of the Rooiberg massif (Goldblatt & Porter

13575), representing the second known station for the

species, has the dorsal tepal erect over the stamens and

style, and the length of the perianth tube at the lower end

of the range recorded for the species. In other respects,

however, including the short, oblong capsides, the Rooi-

berg plants match those from the type locality and there

is at present no reason to suspect that they are not con-

specific.

The species might be confused with Xenoscapa gran-

diflora from southern Namibia but that species has larger,

pale lilac flowers with a much longer perianth tube,

33-35 mm long, and the dorsal tepal 12-14 mm long.

Flowering in X. uliginosa is also generally later, in Sep-

tember and October (rarely into November) rather than in

mid-winter.

Additional material examined

NORTHERN CAPE. — 3017 (Hondeklipbaai): Modderfonteins-

berg, (-BB), 4 Nov. 1830, Drege 2635 (P); Sneeukop, above Modder-

fontein Farm, (-BB), 15 Sept. 1990, Rourke & Nelson 1936 (NBG);

Sneeukop, Bovlei Farm, (-BB), 28 Sept. 1991, Bean & Trinder-Smith

2634 (BOL). 3018 (Kamiesberg): Langkloof, Farm Naartjiesdam, (-

AC), 23 Sept. 2010, Goldblatt & Porter 13575 (NBG, MO).

3. Xenoscapa fistulosa (Spreng. exKlatt) Goldblatt

& J.C. Manning in Systematic Botany 20: 172 (1995).

Ovieda fistulosa Spreng. ex Klatt: 781 (1863). Lapei-

rousia fistulosa (Spreng. ex Klatt) Baker: 155 (1877).

Anomatheca fistulosa (Spreng. ex Klatt) Goldblatt: 88

(1972). Type: South Africa, [Western Cape], Olifants

River, Clanwilliam, without date, Ecklon & Zevher Irid.

254 (B, holo.!; MO!, PRE!, iso.).

Deciduous geophyte, (50-)70-200 mm high, includ-

ing flowers. Conn subglobose, 5-8 mm diam.; tunics

of fine to medium-textured, netted fibres. Leaves 3,

basal, prostrate, oblong, lower two largest, 20-50(-

70) x 5—1 8(— 20) mm, upper leaf ± half as large. Stem

terete, inclined at base but then stiffly erect, up to 180

mm long, with up to 4 short or longer branches 5-30

mm long, each subtended by a short cauline bract.

Inflorescence of solitary flowers on main and lateral

axes, sometimes a second inflorescence developing

in axil of upper leaf; bracts green, 5-7 mm long, inner

slightly longer than outer and often shortly forked api-

cally. Flowers zygomorphic, white or cream-coloured,

rarely pale pink, flushed purplish on tube, unmarked,

with sweet-spicy fragrance; perianth tube erect, cylin-

dric, 1 8— 25(— 33 ) mm long, wider and curved just below

apex, 1.0-1. 5 mm diam. at mouth; tepals subequal, 4-7

x 1-3 mm, weakly cucullate apically, dorsal erect but

others spreading ± horizontally at right angles to tube.

Stamens: filaments 3-5 mm long, exserted ± 2 mm from

tube; anthers 1.0-2. 5 mm long, dark purple; pollen grey-

blue. Ovary ellipsoid; style branching between base and

middle of anthers, branches 2 mm long, deeply forked

and apically recurved. Capsule oblong to cylindric,

( 8—) 1 2—20 x 2. 5-4.0 mm. Seeds ± 1 mm diam. Flower-

ing time: mainly Aug.-Sept., rarely in late July. Figure

IG, H.

Distribution and ecology: widely distributed along

the near interior of the southern African Atlantic coast

and the western Karoo, Xenoscapa fistulosa extends

from Aurusberg and the southern edge of the Huib Hoch

Plateau in southern Namibia through the higher-lying

parts of the Richtersveld and Namaqualand into West-

ern Cape, where it occurs in the northern Cedarberg and

inland onto the Roggeveld Escarpment, thence south-

wards into the Worcester Valley as far east as Montagu

and thence to Faingsburg and the northern foothills of

the Klein Swartberg, with two isolated stations along

the coast, one at Vredenburg and the other on the lower

slopes of Fion’s Head on the Cape Peninsula (Figure 2).

Bothalia 41,2 (2011)

287

The species is mostly restricted to shale or gneiss/gran-

ite substrates, rarely on other rock types in the Rich-

tersveld and southern Namibia, from near sea level to

almost 1 300 m. It does not occur on sandstone-derived

soils of the Cape System and is thus virtually absent

from the southwestern Cape mountain systems — the two

coastal stations in the southwestern Cape are on granite

outcrops.

Plants are invariably restricted to cool, seasonally

moist or wet and largely shaded situations in the lee

of rocks or boulders, with the corms often anchored in

moss pads. The flowering season is substantially length-

ened in years of favourable rainfall by the development

of a secondary inflorescence from the upper leaf axil.

This has not been observed in the other two species.

The pale, mostly white or cream-coloured flowers

of Xenoscapa fistulosa with their sweet-spicy fragrance

suggest that the species is adapted to moth pollination.

It co-occurs with long-proboscid fly-pollinated X. uligi-

nosa in the Kamiesberg ( Bean & Trinder-Smith 2642 &

2634 respectively), and hybrids between the two (Gold-

blatt 9244 A MO) are known (Goldblatt & Manning

1995).

Diagnosis and relationhips : Xenoscapa fistulosa is

recognized by relatively long-tubed, white or cream-

coloured (rarely pale pink) flowers with a strong sweet-

spicy fragrance. The slender perianth tube is mostly

1 8-25 mm long, exceptionally 30-33 mm long in plants

from Pakhuis Pass and the northern Cedarberg (Leipoldt

s.n., Goldblatt 544), and only 1.0-1. 5 mm in diameter at

the mouth. The tepals are mostly smaller and narrower,

4 — 6(— 7) x 1-3 mm. than in pink-flowered X. grandiflora

and X. uliginosa (Table 1 ). Most collectors remark on

the strong floral fragrance, which immediately distin-

guishes the species from its unscented congeners.

Although the species is typically white-flowered,

populations from the Richtersveld and southern Namibia

often have pale pink flowers [see Williamson (2000): 127

Xenoscapa fastuosa (sic.) for illustration]. The smallest-

flowered species in the genus, X. fistulosa may grow

much taller than the other species, the stem reaching

up to 180 mm long, with the lateral branches up to 30

mm long. The capsules are similarly often significantly

longer, up to 18 mm long and strongly cylindric. The two

outlying coastal populations, from Witteklip at Vreden-

burg and Lion’s Head on the Cape Peninsula, although

well separated geographically from the main range of

the species, do not evidently differ morphologically from

inland populations.

Additional material examined

NAMIBIA. — 2716 (Witputz): Karas, Aurusberge, saddle overlook-

ing Roter Kamm, (-CB), 28 Sept. 1996, Marmheimer & Burke 393

(WIND); Rosh Pinah. Spitskop, (-DC), 25 Sept. 1981, Muller & Horn

1599 [sic] (WIND); 9 Aug. 2000, Bruy ns 8314 (NBG); Rosh Pinah,

Sonberg, (-DD), 3 Sept. 2000, Bruyns 8855 (NBG); Witputz, Zebra-

fontein, (-DD), 22 Sept. 1981, Miiller & Horn 1599 [sic] (PRE). 2816

(Oranjemund): Diamond Area No. 1, Obib Mtn Peak, (-BA), 3 Sept.

1989, Van Wyk 9028 (PRE).

NORTHERN CAPE.— 2817 (Vioolsdrif): Langermanskop, (-AA),

29 July 1993, Van Jaarsveld & Bezuidenhout 13430 (NBG); Rosyn-

tjiesberg, neck north of Lelieshoek, (-AC), 30 Aug. 1977, Oliver,

Tolken & Venter 294 (PRE); Stinkfontein Mtns, Comellsberg, (-CA),

6 Sept. 1977, Oliver, Tolken & Venter 710 (PRE); summit of Ploeg-

berg complex, (-C'D), 5 Aug. 1979, Van Berkel 104 (NBG). 2917

(Springbok): Steinkopf, (-AA), Aug. 1925, Marloth 6769 (NBG,

PRE); Spektakelberg, Farm Naries, (-AA), 26 Aug. 1983, Van Wyk

6452 (PRE); Bulletrap-Nigramoep road, 1 km east of Doringrivier,

(-BC), 26 Aug. 1981, Van Berkel 426 (NBG); Springbok, NE of

Platjiesfontein Farm, (-DA), Sept. 1995, G & F Williamson 5703

(NBG); Concordia, (-DB), Sept. 1883, Bolus 695 (BOL, SAM). 3017

(Hondeklipbaai): Sneeukop, Bovlei Farm, (-BB), 28 Sept. 1991, Bean

& Trinder-Smith 2642 (BOL); 16 miles [25.6 km] SW of Garies, (-

DB), without date, Hal! 3763 (NBG). 3018 (Kamiesberg): granite

dome NE of Farm Outuin, (-AA), 21 Aug. 2001, Goldblatt & Porter

11734 (MO, NBG); Studer’s Pass, (-AC), 19 Aug. 1986, Bean 1717

(BOL); Grasberg, (-CB), 22 Aug. 1999, Desmet 166 (NBG); Klip-

poort-se-Berg, (-CB), 21 Aug. 1999, Desmet 140 (NBG); Langberg,

W of Loeriesfontein, (-DB), 5 Sept. 2006, Goldblatt & Porter 12766

(MO, NBG). 3119 (Calvinia): Menzieskraal Farm, (-AC), Sept. 1899,

Leipoldt 825 (SAM); lower slopes of Hantam Mtns, Akkerendam, (—

BD), 22 Jul. 1961, Lewis 5807 (NBG); Menzieskraal Farm, (-CB), 22

Aug. 1986, Snijman 1077 (NBG); Keiskie-se-poort, (-DB), 17 Sept.

2008, Goldblatt & Porter 13134 (MO, NBG). 3220 (Sutherland):

Tankwa Karoo National Park, Farm Klipfontein, (-AA), 14 Sept.

2007, Sachse 554 (PRE); Tankwa Karoo National Park, Farm Klein-

fontein, (-AA), 6 Aug. 2006, Klopper 335 (PRE); Agterkop Farm, (—

AA), 5 Aug. 2006, Steyn 901 (PRE); Soekop Farm, Bohoek, (-AA),

8 Aug. 2006, Rosclt 462 (NBG); Tankwa Karoo National Park, near

Paulshoek, (-AC), 27 Aug. 2004, Steyn 589 (PRE); Kuduberg, Muis-

hondhoogte, (-CA), Sept. 1921, Marloth 10382 (PRE); Houthoek, (—

CA), 15 Aug. 1968, Hanekom 1092 (PRE); Verlatekloof Pass, (-DA),

25 Aug. 2004, Steyn 553 (PRE).

WESTERN CAPE. — 3118 ( Vanrhynsdorp); Mauwerskop, (-DB),

20 Aug. 1986. Snijman 1058 (NBG). 3217 (Vredenburg): Witte-

klip, near Vredenburg, (-DD), 1 Sept. 1944, Lewis 890 (SAM). 3218

(Clanwilliam): Olifants River, (-BB), 25 Aug. 1894, Schlechter 5024

(BOL); east bank of lower Clanwilliam Dam. (-BB), 8 Sept. 1976,

Thompson 2819 (NBG). 3219 (Wuppertal): Clanwilliam, Alpha Farm,

(-AA), 20 July 1941, Bond 1054 (NBG); Pakhuis Pass, (-AA), 3 Sept.

1933, Leipoldt s.n. BOL45054 (BOL); near top of Bidouw Pass, (-

AA), 14 Sept, [without year], Goldblatt 544 (BOL). 3220 (Sutherland):

Farm Thyskraal, (-CC), 7 Sept. 1986, Cloete & Haselau 292 (NBG).

3318 (Cape Town): Lion’s Head. (-CD), Aug. 1924, Rennie & Giffen

s.n. SAM27416 (SAM); slopes of Lion’s Head, (-CD), Aug. 1940,

Lewis 4293 (SAM). 3319 (Worcester): Karoopoort, (-BA), 18 Sept.

1938, Hafstrom & Acocks 315 (PRE); 16 Sept. 1971, Thompson 1261

(NBG); Hex River Pass, near top, (-BD), 2 Sept. 1992, Goldblatt &

Manning 9374 (PRE); Worcester. Karoo Garden, (-CB), 28 July 1975,

Dobay 21 (NBG); 13 Aug. 1970, Bayer 2 (NBG); Stettyn area. Farm

Kleindoom, (-CD), 1971 [without precise date], Oliver 3280 (NBG,

PRE); N end of Farm Alfalfa, near Moordkuil, (-DA), 26 Sept. 1983,

Goldblatt & Snijman 6971A (MO, NBG). 3320 (Montagu): Whitehill

Ridge, (-BA), 17 Aug. 1942, Compton 13377 (NBG); Oudeberg, NE

of Montagu, (-CB), 27 July 1959, Acocks 20533 (PRE); 8 km SE of

Montagu, Farm Amanzi, (-CD), 15 Aug. 1993, Manning 2028 (NBG);

S foothills of Voetpadsberg, Farm Doringkloof, (-DB), 23 Aug. 1985,

Van der Kooy 30 (NBG). 3321 (Ladismith): Kleinswartberg foothills,

road to Seweweekspoort. Farm Modderfontein, (-AC), 21 Sept. 2003,

Goldblatt & Porter 12327 (MO, NBG).

ACKNOWLEDGMENTS

We are grateful to Ernst van Jaarsveld, horticulturist

at Kirstenbosch National Botanical Gardens, for bring-

ing Xenoscapa grandiflora to our attention; the curators

of BOL, MO and PRE for providing access to their col-

lections; and to Colleen Mannheimer for providing addi-

tional records from southern Namibia.

REFERENCES

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nean Society, Botany 16: 61-180.

BARNARD, P„ BETHUNE, S. & KOLBERG, H. 1998. Biodiversity

of terrestrial and freshwater habitats. In P. Barnard, Biologi-

cal diversity in Namibia: a country study: 57-187. Namibian

National Biodiversity Taskforce, Windhoek.

288

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BRUMMITT, R.K. & POWELL, C.E. 1992. Authors of plant names.

Royal Botanic Gardens, Kew.

GOLDBLATT, P. 1972. Revision of Lapeirousia and Anomatheca in the

winter rainfall area of southern Africa. Contributions from the

Bolus Herbarium 4: 1—111.

GOLDBLATT. P. DAVIES. T.J., MANNING, J.C., VAN DER BANK,

M. & SAVOLAINEN, V. 2006. Phylogeny of Iridaceae subfami-

ly Crocoideae based on a combined multigene plastid analysis.

Aliso 22: 399-411.

GOLDBLATT, P. & MANNING, J.C. 1995. Phylogeny of the African

genera Anomatheca and Freesia (Iridaceae: Ixioideae), and a

new genus Xenoscapa. Systematic Botany 20: 161-178.

GOLDBLATT, P. & MANNING, J.C. 2000a. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanical

Institute, Cape Town & Missouri Botanical Garden, St Louis.

GOLDBLATT, P. & MANNING, J.C. 2000b. The long-proboscid fly

pollination system in southern Africa. Annals of the Missouri

Botanical Garden 87: 146-170.

GOLDBLATT, P. & MANNING, J.C. 2008. The iris family: natural

history and classification. Timber Press, Oregon.

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.

KLATT, F.W. 1863. Revisio iridearum. Linnaea 32: 689-784.

MANNING, J.C. & GOLDBLATT, P. 1996. The Prosoeca peringueyi

(Diptera: Nemestrinidae) pollination guild in southern Africa:

long-tongued flies and their tubular flowers. Annals of the Mis-

souri Botanical Garden 83: 67-86.

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Umdaus, Hatfield, Pretoria.

Bothalia 41,2: 289-326 (2011)

Notes on African plants

VARIOUS AUTHORS

NYCTAGINACEAE

NOTES ON COMM1CARPUS IN SOUTHERN AFRICA, INCLUDING A NEW RECORD FOR NAMIBIA

Commicarpus Standi, is a genus in the Nyctaginaceae

with its main distribution range in Africa (Meikle 1978).

In southern Africa, nine species occur which are widely

distributed in the region, with its centre of diversity in

Namibia (Germishuizen & Meyer 2003). No formal

taxonomic research has specifically been conducted on

Commicarpus in southern Africa. Most of the existing

knowledge of the group is based on floras from else-

where in Africa (Baker & Wright 1909; Hutchinson &

Dalziel 1927; Meikle 1954; Stannard 1988; Whitehouse

1996).

Anthocarp morphology is considered to provide the

most differential characters of the family (Smith 1976;

Willson & Spellenberg 1977; Bohlin 1988; Douglas

& Manos 2007). An anthocarp consists of a fruit (an

achene or utricle) enclosed in a persistent, accrescent,

hard, leathery or fleshy base of the calyx tube (Bogle

1974). The anthocarp of the southern African Com-

micarpus species is 5-10 x 2-3 mm, cylindrical, fusi-

form. clavate or elliptic-clavate with ten obscure ribs

and large, dark, mucilaginous, wart-like glands scattered

over the surface. The apex is surrounded by ten glands

which are either stalked or sessile.

An investigation of the anthocarp morphology of the

southern African Commicarpus species has revealed

numerous misidentifications in southern African her-

baria. These misidentifications were probably based on

leaf morphology and growth form which do not vary

considerably between Commicarpus species. Finer mor-

phological differences of the anthocarps facilitate the

construction of a key to accurately differentiate between

the taxa below genus level.

Key to species

la Anthocarp cylindrical with sessile glands around apex

1 . C. chinensis subsp. natalensis

lb Anthocarp fusiform, clavate or elliptic-clavate with stalked

and sessile glands around apex:

2a Anthocarp elliptic-clavate 2 . C. pilosus

2b Anthocarp fusiform or clavate:

3a Anthocarp fusiform:

4a Ring of five stalked glands alternating with five sessile

glands around apex; anthocarp 4—6 mm long

3. C. squarrosus or 4. C. friiticosus

4b Ring of ten stalked glands around apex; anthocarp 7-9

mm long:

5a Surface of anthocarp below apex covered sparsely

with short-stalked glands (< 1 mm long) grouped

together 5. C. plumbagineus

5b Surface of anthocarp below apex covered with scat-

tered sessile glands 6 . C.fallacissimus

3b Anthocarp clavate:

6a Anthocarp broadly clavate; surface covered with pro-

minent glandular hairs; apex surrounded by ten

short-stalked glands (< 1 mm) 7. C. decipiens

6b Anthocarp clavate; surface glabrous to puberulent;

apex surrounded by five stalked glands alternating

with five sessile glands:

7a Apex surrounded by five thick-stalked glands, stalks

< 1 mm long 8. C. pentandrus

7b Apex surrounded by five thin-stalked glands, stalks

1-2 mm long 9. C. helenae var. helenae

The key has facilitated the re-identification of large-

fruited Commicarpus specimens collected from Namibia.

We have identified these specimens as Commicarpus

helenae (Roem. & Schult.) Meikle var. helenae , a vari-

ety with a wide distribution from the Middle East into

Africa, which also occurs on the islands of the Canar-

ies, Cape Verde and the type locality, St Helena (Meikle

1978). In the Flora of southern African ( FSA ) region, it

was hitherto only known from Botswana and the Lim-

popo Province of South Africa and was not considered

to occur in Namibia (Germishuizen & Meyer 2003). In

southern Africa, the taxon has been regarded as of lim-

ited occurrence, as the circumscription of Commicarpus

helenae var. helenae has generally been misunderstood.

Based on anthocarp structure, we can confirm that many

southern African specimens of C. helenae var. helenae

have been misidentified and erroneously placed under C.

fallacissimus (Heimerl) Heimerl ex Oberm., Schweick. &

I.Verd.. C. pentandrus (Burch.) Heimerl, C. pilosus (Hei-

merl) Meikle, C. plumbagineus (Cav.) Standi, var. plum-

bagineus and C. squarrosus (Heimerl) Standi. Especially

the misidentifications as C. fallacissimus, C. pilosus or C.

squarrosus can be ascribed to the semisucculent leaves

and growth form that these taxa share with C. helenae

but the anthocarps differ considerably in shape and gland

characters (Figure 1). As a result, the distribution patterns

of these taxa in the FSA region have also been misinter-

preted.

We can therefore confirm the presence of Commi-

carpus helenae var. helenae in Namibia based on the

anthocarp structure of the specimens being most simi-

lar to what is generally regarded as typical for the spe-

cies (Struwig et al. 2010). The anthocarp is clavate and

tapers markedly from the apex to the base with a ring

of five, long, stalked glands around the apex and two

rings of sessile glands below. However, a specimen from

Khowarib ( Struwig 44) in the Kaokoveld, differs from

other specimens in that the five stalked glands around

the apex are longer (2 mm) than those recorded for other

specimens from southern Africa (1 mm) (Figure 2). This

taxon is being investigated further.

290

Bothalia 41,2 (2011 )

FIGURE I . — Anthocarps of Commicarpus: A, C. helenae var. helenae', B, C.fal/acissimus ; C, C. pilosus: and D, C. squarrosus. Scale bars: 5 mm.

Photographs: S.J. Siebert.

Commicarpus helenae ( Roem . & Schult .) Meikle

in Hooker’s leones Plantarum 7, 4: t. 3694 (1971).

Boerhavia helenae Roem. & Schult.: 73 (1822). Type:

St Helena, Ladder Hill, Burchell l [K, neo., designated

by Whitehouse ( 1 996).-Aluka image, website accessed

08-02-2010],

Boerhavia slellala Wight: 6, t. 875 (1843) non B. stellata Bojer:

188 (1842) nom. i I legit. C. stellatus (Wight) Berhaut: 51 (1953). Type:

India.

[C. verlicillalus sensu Baker & Wright: 6 (1909); sensu Hutchinson

& Dalziel: 153 (1927); sensu Heimerl: 1 17 (1934); sensu Balle: 86 t. 7

(1951); sensu Meikle: I ( 1954); non Standi.: 101 (1916)].

Commicarpus helenae {Roem. & Schult.) Meikle

var. helenae

Herb, from a woody rootstock. Stems 1 .0-1.5 m long,

slender, erect, decumbent or scrambling, branching.

pubescent or glabrous. Leaves ovate, 15-50 * 10-40

mm, ± fleshy, pubescent to glabrous, apex rounded

to acute, apiculate, base cordate, rounded or more or

less truncate, slightly attenuate along petiole; margins

entire to somewhat repand; petioles 4-22 mm long,

sparsely pubescent. Inflorescence narrow, verticillate,

long-pedunculate, peduncles sparsely pubescent; bracts

linear-lanceolate, 1-2 mm long, pubescent. Flowers ses-

sile to very shortly pedicellate. Perianth 4.0-5.0(-6.5)

mm long, lower portion sulcate with 5 prominent vis-

cid glands around apex, upper portion 2. 0-3. 0(— 4.5)

mm long, widely infundibuliform with a short (1 mm)

inconspicuous tube, pubescent; sepals purple, mauve,

magenta, pink, white or yellow. Stamens 2 or 3; fila-

ments 4— 5(— 6) mm long; anthers transverse-elliptic to

rounded, 0.3-0. 5 mm long. Ovary ellipsoid, shortly

stipitate, 0. 5-0.6 mm long, glabrous; style 3.0— 3.5(—

5.5) mm long. Anthocarp clavate, tapering markedly

Bothalia 41,2 (2011)

291

FIGURE 2. — Light microscope

micrographs of anthocarps

of Commicarpus helenae var.

helenae. A, specimen collect-

ed from Kruger National Park,

South Africa, Zambatis 1954;

B, specimen collected from

Khowarib, Namibia, Stniwig

44. Scale bars: 1 mm.

from apex to base, 3. 5-7.0 x 1.0-2. 5 mm, sessile viscid

glands in rings, 5 stalked viscid glands, 1-2 mm long,

alternating with 5 sessile glands around apex Figures

1A; 2 A, B.

Commicarpus helenae has two varieties based on char-

acters of a morphological and biogeographical nature.

Commicarpus helenae (Roem. & Schult.) Meikle var.

barbatus Meikle differs from the typical variety in having

remarkably hairy stems and leaves (Meikle 1979, 1983).

Its distribution is restricted to tropical Africa (Klopper et

al. 2006; African Plant Database 2010). In Africa, the typi-

cal variety occurs as far south as N’watinwambu in the

Kruger National Park, South Africa (Maurin & Van der

Bank OM348).

Distribution and habitat : previously only 1 1 localities

of Commicarpus helenae var. helenae were known for

southern Africa, but our redeterminations have increased

this figure to 25, suggesting that the taxon is much more

12 14 16 18 20 22 24 26 28 30 32

FIGURE 3. — Distribution of Commicarpus helenae var. helenae in

southern Africa.

part of the southern African flora than was previously

thought (Figure 3). It has been recorded from similar

habitat types in Namibia and South Africa. Labels gen-

erally describe the habitat as rocky, in sandy, loamy or

alluvial soil in calcrete or sandstone areas.

Specimens examined

NAMIBIA. — 1713 (Swartbooisdrif): Omitengundi, 500 m north

of road junction, (-AB), 09-04-1998, Schulte 108530 (WIND). 1913

(Sesfontein): Khowarib Rest Camp, underneath Acacia and Mopani

trees, (-BD), 09-02-2009, Stniwig 44 (PUC, WIND). 1916 (Gobaub):

Etosha National Park, Charitsaubplain, near pan with Acacia reficiens ,

(-AA), 18-04-1973, Le Roux 520 (WIND, PRE). 2115 (Karibib):

Spitzkoppe, (-DC), 03-03-1985, Craven 2022 (WIND). 2216 (Otjim-

bingwe): Okomitundu Farm, underneath Acacia trees in front of gate

to house, (-AB), 12-04-2010, Stniwig 183 (PUC, WIND).

BOTSWANA. — 2022 (Lake Ngami): Lake Ngami, Mwaku Pan

near Sehitwa, slopes around Mwaku Pan, (-BD), 12-1969, Van der

Spuy 30 (PRE, UCBG); north of Ngwanalekau Hills, open areas (often

termitaria) in Terminalia prunioides woodland, (-DB), 14-03-1969,

Buerger 1149 (PRE, UCBG). 2122 (Kobe): ± 10 km past Kuke Vil-

lage on road to Maun, (-AB), 06-03-1996, Burgoyne & Snow 5283

(PRE, UCBG). 2123 (Pink Pan): central Kalahari, (-AA), 02-02-1991,

Barnard 586 (PRE, UCBG). 2125 (Lothlekane): Orapa, (-AD), 16-03-

1975, Kerfoot 7748 (PRE. UCBG).

LIMPOPO. — 2229 (Waterpoort): Breslau Game Farm, base of

koppie, (-AC), 05-05-2000, Straub s.n. (PUC); Breslau Game Farm.

(-AC), 30-05-2000, Straub 831 (PRE); Langjan Nature Reserve, (-

CC), 12-1974, Zwanziger 496 (PRE); Soutpan 193, thinly scattered

in outer circle of salt pan, (-CD), 19-11-1932, Obermeyei; Schweick-

erdt & Verdoorn 19 (PRE); Soutpan, west side of pan, (-CD), 21-01-

1931, Bremekamp & Schweickerdt 249 (PRE, PRU). 2230 (Messina):

Tshipiza work station (Greater Kuduland Safaris), (-CB), 06-12-1984,

Van Wyk 6887 (PRE, PRU). 2331 (Phalaborwa): 50 m west of Shing-

wedzi River, over cement drift on bank, (-AA), 11-03-2010, Stniwig

141 (KNP. PUC); 150 m west of cement drift over Shingwedzi River,

upstream of Red Rocks, (-AA), 14-01-1994, Zambatis 1954 (KNP,

PRE); Letaba River, western border Mhlangene, (-CA), 01-12-1992,

Van Rooyen & Bredenkamp 582 (PRU). 2431 (Acomhoek): Kingfish-

erspruit, Timbavati, (-AD), 21-12-1962, Biologiese Afdeling 4592

(KNP, PRE); Klaserie Nature Reserve, Farm Ross 55KU, (-AD),

16-12-1981, Zambatis 1306 (PRE); Timbavati Private Nature Reserve,

Hans Hoheisen Wildlife Reserve Station, Kempiana 90KU, (-AD),

18-12-1982, Zambatis 1526 (PRE); Kruger National Park, Acomhoek,

in grassland, (-BC), 03-1975, Gertenbach 5042 (PRE).

MPUMALANGA. — 2431 (Acomhoek): Kruger National Park,

Nkuhlu Partial Exclosure, Sabie River, (-DD), 08-12-2010, Van Coller.

292

Bothalia 4 1 ,2 (2011)

Siebert & Siebert, 4339 (PUC). 2531 (Komatipoort): N'watinwambu

firebreak about 1 km from SI 14 road, (-BA), 05-12-2005, Maurin &

Van der Bank OM348 (KNP).

ACKNOWLEDGEMENTS

Ms Hester Steyn is thanked for the production of the

distribution map and the curators of the following her-

baria for providing access to study material: KNP, PRE,

PRU, PUC and WIND (acronyms according to Holmgren

et al. 1990). The North-West University and National

Research Foundation provided financial support.

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BOHLIN, J-E. 1988. A monograph of the genus Colignonia (Nyctagi-

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BOJER, M.W. 1842. Descriptions de diverses plantes nouvelles de

Madagascar, des ties Comores et de File Maurice. Boerhaavia

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Naturelles, Botanique 2,18: 188, 189. Fortin, Masson & C.

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DOUGLAS, N.A. & MANOS, P.S. 2007. Molecular phylogeny of

Nyctaginaceae: taxonomy, biogeography and characters asso-

ciated with a radiation of xerophytic genera in North America.

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GERMISHUIZEN, G. & MEYER, N.L. 2003. Plants of southern

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KLOPPER, R.R., CHATELAIN, C„ BANNINGER, V„ HABASHI. C„

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M. STRUWIG*!, S.J. SIEBERT* and E.S. KLAASSEN**

* A.P. Goossens Herbarium, School of Environmental Sciences and De-

velopment, North-West University, Private Bag X6001, 2520 Potchef-

stroom. South Africa.

j Author for correspondence: madeleen.struwig@nwu.ac.za.

** National Botanical Research Institute, Private Bag X13184, Wind-

hoek, Namibia.

MS. received: 2010-07-05.

AIZOACEAE

NEW COMBINATIONS IN ANTI MIMA AND OCTOPOMA (RUSCHIOIDEAE)

1. A new combination in Antimima for Ruschia pari-

petcila (L. Bolus) L. Bolus

Antimima is a large and still unrevised genus of ±

100 species (Dehn 1989; Hartmann 1998b). Many spe-

cies, most of which were described by L. Bolus, were

not compared with similar species, making it unclear to

which species they were most similar.

Features by which species of Antimima may be re-

cognized include 5(or 6)-locular fruits with large clos-

ing bodies and shallow locules, heterophyllous leaves,

and 1( 3)-fiowered inflorescences (rarely aggregated in

well-developed cymes) (Dehn 1989; Hartmann 1998b).

In addition, the leaves in species of Antimima often have

a papillate epidermal surface, whereas they are typically

smooth in Ruschia. Notably, 6-locular fruits are rare in

both Ruschia and Antimima.

A six-locular species of Ruschia from Namaqua-

land, R. hexamera L. Bolus, was recently found to

be conspecific with Antimima crassifolia (L. Bolus)

H.E.K. Hartmann and to represent the earlier name for

the taxon (Klak 2010). A further collection by Pillans

from Wallekraal near Hondeklip Bay in Namaqualand,

described as R. paripetala by L. Bolus in 1927, is also

very similar to A. hexamera (L. Bolus) Klak. The type

specimens of both A. hexamera and R. paripetala pos-

Bothalia 41,2 (201 1 )

293

sess six-locular fruits with closing bodies and both

lack valve wings; the epidermis of the leaves is papil-

late; the inflorescence is 1-3-flowered; and the petaloid

and filamentous staminodes are pink to white. In addi-

tion, L. Bolus described a further collection by Pillans,

collected between Sendelings Drift and Doornpoort,

as R. paripetala var. occultcms L. Bolus (1929: 125).

This variety was considered a synonym of A. perforata

(L. Bolus) H.E.K. Hartmann (Hartmann 1998b). Both

var. occultans and A. perforata have six-locular fruits

and a somewhat similar habit, but differ from R. pari-

petala and A. hexamera by having smaller flowers, up to

14 mm diam. in A. perforata and var. occultans , com-

pared to 27 mm diam. in R. paripetala and A. hexam-

era. Hartmann (2001) described the closing body of R.

paripetala as small (as opposed to large in A. perforata ),

which is generally a characteristic of species of Ruschia.

The somewhat smaller than usual closing body in R.

paripetala may have been the main reason for retain-

ing R. paripetala in Ruschia. However, a reinvestigation

of the closing bodies in the types of R. paripetala , R.

paripetala var. occultans , A. hexamera and A. perforata

found that in none of them were the bodies sufficiently

large to block the entire exit of the locule. The somewhat

smaller size of the closing bodies in A. crassifolia and A.

hexamera was previously discussed by Klak (2010). In

other characteristics, such as the presence of expanding

keels which extend into awns, the shallow locules, the

papillate epidermis and the overall similar habit, the type

specimens are in agreement. The morphological evi-

dence thus suggests that R. paripetala is best placed in

Antimima, where it is the earliest available name for this

group of taxa. The new combination and full synonymy

is given below.

The most northerly collection of this species was

made in the Richtersveld and collections have been

made as far south as Hondeklip Bay in Namaqualand.

Antimima paripetala (L. Bolus) Klak , comb. nov.

Mesembiyanthemum paripetalum L. Bolus, Annals of

the Bolus Herbarium 4: 87 (1927). Ruschia paripetala

(L. Bolus) L. Bolus: 221 (1950). Type: South Africa, [North-

ern Cape], from the coast at Hondeklip Bay, Oct. 1924, Pil-

lans 17759 (BOL, sheet I. lecto.!, here designated).

R. hexamera L. Bolus: 144 (1928). Antimima hexamera (L. Bolus)

Klak: 307 (2010). Type: South Africa, [Northern Cape], hills near

Brakfontein, between Sept, and Oct. 1926, Pillans 5703 (BOL, holo.!).

R. hexamera L.Bolus var. longipetala L. Bolus: 237 (1931), syn.

nov. Type: South Africa, [Northern Cape], Port Nolloth, without date,

M. Schlechter sub SUG8367 (BOL, holo.!).

R. crassifolia L.Bolus: 338 (1958), syn. nov. Antimima crassifolia

(L.Bolus) H.E.K. Hartmann: 71 (1998b). Type: South Africa. [Northern

Cape], Lekkersing, June 1954, H. Hall sub BOL25758 (BOL. holo.!).

Additional material examined

NORTHERN CAPE. — 2816 (Oranjemund): Alexander Bay, along

road from Port Nolloth to Lekkersing, (-BD), 3 Sept. 2001. Klak 776

(BOL); Boegoeberg Suid, (-DC), 26 Oct. 1985, Van Jaarsveld 8229

(NBG). 2916 (Port Nolloth): between Port Nolloth and Holgat, (-

BB). May 1929, Pillans 5766 (BOL). 2917 (Springbok): Vioolsdrif,

between Port Nolloth and Lekkersing, (-AA), 4 Sep. 2001, Klak 780

(BOL); Karrachab Poort. (-AA), 18 July 1970, Wisitra 1625 (NBG).

2. A new combination in Octopoma for Ruschia nana

L.Bolus

Octopoma is a genus of nine species, which can be

subdivided into two groups, one occurring in Namaqua-

land and the other in the Little Karoo (Klak 2010). The

main characteristics of Octopoma , which distinguish

it from Ruschia and Leipolcltia, are the 6- to 8-locular

fruits, with no or only narrow valve wings (Hartmann

1998a). In Ruschia , the fruits are 5(rarely 6)-locular,

with no valve wings or rarely with narrow valve wings,

whereas in Leipoldtia the fruits are ± 10-locular with

broad valve wings. Recently, a new species, Octopoma

tanquanum Klak, was described. This is found in the

Tanqua and Little Karoo and differs from the other spe-

cies of Octopoma by its 6-locular fruits and broad valve

wings (Klak 2010).

Examination of herbarium specimens currently pla-

ced in Ruschia indicates that the little-known species,

R. nana L.Bolus, is conspecific with O. tanquanum.

The type of R. nana was collected at Matjiesfontein by

H. Bolus in September 1908. Unfortunately, this speci-

men lacks fruits, so that the internal morphology of its

capsule remains uncertain. However, the presence of six

calyx lobes and six styles indicate that the fruits would

be 6-locular. The presence of six-locular fruits is very

rare in Ruschia and is an indication that the species may

not belong in Ruschia. A comparison of the type of R.

nana with the type of O. tanquanum shows that both

have extremely similar leaves, which are trigonous,

fused towards the bases and finely serrated along the

keel towards the apex. Also, both type collections have

solitary, cream-coloured flowers and bracteoles which

embrace the base of the flower. Both O. tanquanum

and R. nana flower in late spring to early summer (end

of October to November and December respectively).

Since both type collections are from the Matjiesfontein

area, there is little doubt that the two species are conspe-

cific. The absence of mature fruits on the type of R. nana

could have been one reason why this species was not

previously transferred to Octopoma (Hartmann 1998a).

The following new combination and synonym are pro-

vided.

Octopoma nanum (L.Bolus) Klak , comb. nov.

Ruschia nana L.Bolus, Notes on Mesembryanthemum

and allied genera 2: 75 (1929). Mesembryanthemum

reduction N.E.Br.: 32 non 33 (1930). Type: South Africa.

[Western Cape], Matjiesfontein, Sept. 1908, fl. Cape

Town in Dec. 1908, H. Bolus 15417 (BOL, holo.!).

O. tanquanum Klak: 302 (2010), syn. nov. Type: South Africa,

Western Cape, Tanqua Karoo, 7 km west of Matjiesfontein, Farm Aas-

voelbos, 26 May 2007, Bruyns 10797 (BOL, holo.!).

Additional specimens examined

NORTHERN CAPE. — 3220 (Sutherland): between Hottentots

Kloof and Sutherland, (-CC), Leipoldt s.n. (BOL).

WESTERN CAPE. — 3319 (Worcester): Karoopoort, (-BA), Little-

wood sub KG534/59 (BOL); east of Karoopoort, (-BB), 17 Aug. 2002,

Bruyns 9158 (BOL); 5.5 miles [8.8 km] from Touws River on Worces-

ter road (-BD), Acocks 15289 (BOL). 3320 (Montagu): Touws River

Dist., Tanqua Karoo, Melkboskraal Farm, (-AA), 12 Sept. 2006, Klak

1373 , (BOL); 7 km west of Matjiesfontein, Aasvoelbos Farm, (-BA),

28 May 2007, Klak 1441 (BOL); Matjiesfontein, (-BA), Pillans 2057

(BOL); Ceres Karoo. Herre SUG10508 (BOL); Bloutoring station, (-

CB), 22 June 2009, Bruyns 11381 (BOL); Sewefontein Farm, (-DA),

Bruyns 11389 (BOL); Tilney Farm, (-DC), 26 May 2002, Bruyns 9025

(BOL).

294

Bothalia 41,2 (201 1 )

ACKNOWLEDGEMENTS

The National Research Foundation (NRF) is thanked

for financial support towards this study. The Curator of

the Compton Herbarium (NBG) is thanked for permis-

sion to examine herbarium material.

REFERENCES

BOLUS, H.M.L. 1927. Novitates Africanae. Annals of the Bolus Her-

barium 4: 87.

BOLUS. H.M.L. 1928. Notes on Mesembryanthemum and allied gen-

era, part 1 : 144.

BOLUS, H.M.L. 1929. Notes on Mesembryanthemum and allied gen-

era. part 2: 75, 125.

BOLUS, H.M.L. 1931. Notes on Mesembryanthemum and allied gen-

era, part 2: 237.

BOLUS, H.M.L. 1950. Notes on Mesembryanthemum and allied gen-

era, part 3: 221 .

BOLUS, H.M.L. 1958. Notes on Mesembryanthemum and allied gen-

era, part 3: 338.

BROWN, N.E. 1930. Mesembryanthemum. Gardeners' Chronicle ser.

3, 87: 32.

DEHN, M. 1989. Untersuchungen zum Merkmalsbestand und zur Stel-

lungderGattungHnfrm/waN.E.Br. emend. Dehn (Mesembryan-

themaceae Fenzl). MitteiJungen aus dem Institut fur Allgemeine

Botanik Hamburg 22: 189-215.

HARTMANN, H.E.K. 1998a. New combinations in Ruschioideae,

based on studies in Ruschia (Aizoaceae). Bradleya 16: 44-91.

HARTMANN, H.E.K. 1998b. New combinations in Antimima (Rus-

chioideae, Aizoaceae) from southern Africa. Bothalia 28: 67-82.

HARTMANN, H.E.K. 2001. Illustrated handbook of succulent plants:

Aizoaceae F-Z. Springer, Berlin, Germany.

KLAK, C. 2010. Three new species and two new combinations, in the

Aizoaceae from the Western and Northern Cape of South Africa.

South African Journal of Botany 76: 299-307.

C. KLAK*

* Bolus Herbarium, University of Cape Town, 7701 Rondebosch, Cape

Town. E-mail: Cornelia. Klak@uct.ac.za.

MS. received: 2010-10-29.

VITACEAE

A NEW AND AN OVERLOOKED RECORD OF CYPHOSTEMMA IN ANGOLA

The family Vitaceae was recently catalogued for

Plants of Angola (Figueiredo & Smith 2008) by Retief

(2008), who recognised 20 species in the genus. During

ongoing work on the flora of Angola, two additional

species of Cyphostemma (Planch.) Alston were recently

newly recorded for the country. These are Cyphostemma

wittei (Staner) Wild & R.B.Drumm. and C. congestion

(Baker) Desc. ex Wild & R.B.Drumm. C. congestion had

already been recorded for Angola in an obscure publica-

tion that was overlooked by previous reviewers of the

family in Angola.

Cyphostemma congestum (Baker) Desc. ex Wild

& R.B.Drumm. in Flora zambesiaca 2,2: 473 (1966). Vitis

congesta Baker in Oliv.: 412 (1868). Cissits congesta

(Baker) Planch, in A. DC. & C.DC.: 590 (1887). Cypho-

stemma congestion (Baker) Desc.: 120 (1960), comb,

inval. [ICBN Art. 33.4, no clear indication of basionym].

Type: Angola, Chibiza, Meller s.n. (K, holo.).

Cissus fleckii Schinz: 640 (1908a). Cyphostemma fleckii (Schinz)

Desc.: 121 (1960), comb, inval. [ICBN Art. 33.4, no clear indication

of basionym], Cyphostemma fleckii (Schinz) Desc.: 221 (1967). Type:

South West Africa [Namibia], Hereroland, Fleck 762 (Z, holo.).

Cissus amboensis Schinz: 699 (1908b). Type: South West Africa

[Namibia], Hereroland, Dinter 186 (syn.), Amboland, Wulfhorst 176 (syn. ).

Cyphostemma congestum has been recorded from

Namibia, Botswana, South Africa, Mozambique, Zam-

bia and Zimbabwe (African Plants Database 2010). This

species shows variation in the density of stem glands:

in Namibia in the west, the stems can be moderately to

sparsely glandular, while they are more so towards the

eastern parts of its distribution range. The PRE speci-

mens from the Angolan side of the Ruacana Falls, cited

below, have stems that are puberulous with multicellular

hairs, without capitate glandular hairs.

Merxmuller & Schreiber (1969) synonymized Cis-

sus amboensis with Cyphostemma congestum and men-

tioned the occurrence of the taxon in Angola. This was

overlooked by Retief (2008). The specimens listed here

confirm the occurrence of C. congestum in Angola.

Specimens examined

ANGOLA. — Cunene, Ruacana Falls, 30 April 1962, Kotze 57

(PRE!); Cunene, top of Ruacana Falls on Angolan side, 30 April 1962,

Ry croft 2423 (PRE!). Figure 4.

Cyphostemma wittei (Staner) Wild & R.B.Drumm.

in Kirkia 2: 141 (1961). Cissus wittei Staner in Wildeman

& Staner: 49 (1932). Type: from Congo (Katanga).

Cyphostemma wittei was previously only known from

the Democratic Republic of Congo, Tanzania and Zam-

bia. It is a variable species with the stems sometimes

glandular, whereas in other cases they are eglandular.

FIGURE 4. — Distribution of Cyphostemma congestum. •; and C. wit-

tei, A. in Angola.

Bothalia 41,2 (201 1 )

295

This variation has even been encountered on the same

plant (Milne-Redhead 2839, K, cited by Wild & Drum-

mond 1966). The stems on the LISC specimen from

Angola cited below are eglandular.

Specimen examined

ANGOLA. — Moxico, Cameia, Barros Machado ANG.XII. 54-51

(LISC!). Figure 4.

The two new records bring the total number of

Cyphostemma species known for the country, to 22.

Nine of these known Angolan species of Cyphostemma

are endemic.

ACKNOWLEDGEMENTS

We thank Ms Joana Abreu for providing additional

information from LISC herbarium. Ms Ronell Klop-

per of SANBI’s Biosystematics Division, Pretoria, is

thanked for kindly preparing the distribution map. An

anonymous referee is thanked for collegial contributions.

REFERENCES

AFRICAN PLANTS DATABASE (version 3.3). 2010. Conservatoire

et Jardin botaniques de la Ville de Geneve and South African

National Biodiversity Institute, Pretoria, http://www.ville-ge.ch/

musinfo/bd/cjb/africa/ (accessed August 2010).

CANDOLLE, A.L.P.P. DE & CANDOLLE, A.C.P. DE. 1887. Mono-

graphiae Phanerogamarum. Prodromi nunc continuatio, nunc

revisio auctoribus Alphonso et Casimir de Candolle aliisque

botanicis ultra memoratis. Part 5,2: 305-654. Masson, Paris.

DESCOINGS, B. 1960. Un genre meconnu de Vitacees: comprehension

et distinction des genres Cissus L. et Cyphostemma (Planch.)

Alston. Notulae Systematicae 16: 113-125.

DESCOINGS, B. 1967. Note rectificative a propos de la nomenclature

des Cyphostemma (Vitacees). Naturalia Monspeliensia , Serie

Botanique, 18: 226.

FIGUEIREDO, E. & SMITH, G.F. 2008. Plants of Angola/Plantas de

Angola. Strelitzia 22. South African National Biodiversity Ins-

titute, Pretoria.

MERXMULLER, H. & SCHREIBER, A. 1969. 80.Vitaceae. Prodro-

mus einer Flora von Siidwestafrika : 1-8.

OLIVER, D. 1868. Flora of tropical Africa, vol. 1. Reeve, Ashford,

Kent.

RETIEF, E. 2008. Vitaceae. In E. Figueiredo & G.F. Smith, Plants of

Angola/Plantas de Angola, Strelitzia 22: 1 68, 1 69. South African

National Biodiversity Institute, Pretoria.

SCHINZ, H. 1908a. Beitrage zur Kenntnis der Afrikanischen Flora

(suite). Bulletin de I’Herbier Boissier, ser. 2, 8: 625-640.

SCHINZ, H. 1908b. Beitrage zur Kenntnis der Afrikanischen Flora

(suite). Bulletin de I'Herbier Boissier, ser. 2, 8: 699-703.

WILD, H. & DRUMMOND, R.B. 1961. Vitaceae from the Flora zam-

besiaca area: 2. Kirkia 2: 132-143.

WILD, H. & DRUMMOND, R.B. 1966. Vitaceae. In E. Launert, Flora

zambesiaca 2, 2: 439^192. Flora Zambesiaca Managing Com-

mittee, London.

WILDEMAN, E. DE & STANER. P. 1932. Contribution a l ’etude de

la Flore du Katanga, supplement 4: 1-1 16. D. van Keerberghen,

Bruxelles.

F. DE SOUSA*, E. FIGUEIREDO** and G.F. SMITH***

* Department of Plant and Environmental Sciences, University of Gothenburg,

Box 461, SE-40530 Goteborg, Sweden. E-mail: filipedeportugal@gmail.com.

** Department of Botany, P.O.Box 77000, Nelson Mandela Metropolitan Uni-

versity, Port Elizabeth, 6031 South Africa / Centre for Functional Ecology,

Departamento de Ciencias da Vida, Universidade de Coimbra, 3001-455 Coim-

bra, Portugal. E-mail: estrelafigueiredo@hotmail.com (corresponding author).

*** Biosystematics Research & Biodiversity Collections, South Afri-

can National Biodiversity Institute, Private Bag X101, 0001 Pretoria

/ H.G.W.J. Schweickerdt Herbarium, Department of Plant Science,

University of Pretoria, 0002 Pretoria / Centre for Functional Ecology,

Departamento de Ciencias da Vida, Universidade de Coimbra, 3001-

455 Coimbra, Portugal. E-mail: g.smith@sanbi.org.za.

MS. received: 2010-08-22.

APOCYNACEAE (ASCLEPIADOIDEAE-CEROPEGIEAE)

FIRST RECORDS OF ORBEA COOPERI IN GAUTENG AND MPUMALANGA PROVINCES, FSA REGION

The last full revision of the genus Orbea (N.E.Br.)

L.C. Leach by Bruyns (2002), consisted of 56 species

distributed throughout Africa and southwestern Arabia

with ± 3 1 species south of the equator and ± 24 in South

Africa (Bruyns 2002, 2005). As a result of some taxo-

nomic changes, more species were moved to the genus

(Plowes 2004, 2007; Bruyns 2005; Raffaelli et al. 2008;

Meve 2009) bringing the total in Orbea to the current 61

species (sensu Bruyns).

The current known distribution records show that

Orbea cooperi (N.E.Br.) L.C. Leach occurs in the drier

western and central interior of South Africa. Charles

Craib and Gillian Condy, Warren McCleland, Tony de

Castro and eventually the second author reported the

existence of an Orbea (presumably O. cooperi) from

Mpumalanga during 2008-2009. New distribution

records were also collected from the Devon area (Leeu-

kop, Gauteng) and near Greylingstad (Platkop, Mpuma-

langa). The new records constitute a northeastern exten-

sion of the distribution range of O. cooperi (Figure 5).

The habitat at the new sites is stony ground characterized

by dolerite outcrops and sheets, surrounded by grassland

with predominantly black turf soil. O. cooperi grows

amongst the rocks in these dolerite outcrops. This habi-

tat becomes waterlogged and muddy in spring and sum-

mer after rain. As the season progresses, the soil dries

out and becomes hard and cracked in winter. Associated

species found at these localities include succulents such

as Crassula setnlosa and Euphorbia clavarioides, the

karroid dwarf shrub, Eriocephalus karooicus and herbs,

including Lessertia cf. depressa and Jamesbrittenia

stricta, intermixed with other herbs and grasses.

At first, the new records were thought to represent a

new species or subspecies based on the flowers that were

produced by various cuttings grown in the nursery of the

Pretoria National Botanical Garden (PNBG). The flow-

ers of these plants were devoid of vibratile hairs (mar-

ginal cilia) and were much smaller than the reported size

for Orbea cooperi (Figure 6). The exact same plants

grown in the PNBG’s nursery did, however, in the third

296

Bothalia 41,2 (2011)

TABLE 1. — Comparison of main characters to distinguish between

Orbea cooperi and O. tapscottii

FIGURE 5. — Known distribution adapted from Bruyns 2005 of Orbea

tapscottii (shaded light grey) and O. cooperi (shaded dark grey).

Point records for material housed at PRE as follows: Orbea

tapscottii, •; O. cooperi, ▲ . New distributions of O. cooperi

reported here, ■ (both sight records and herbarium specimens

at PRE).

year of flowering produce normal-sized flowers. In addi-

tion, the second author also collected flowering mate-

rial from a site in the vicinity of Greylingstad, Mpuma-

langa, that was furnished with vibratile hairs. Therefore,

the first author could confirm in 2009 that all these new

records were Orbea cooperi.

The first sterile cutting presented by Craib was

thought to be Orbea tapscottii (I.Verd.) L.C. Leach.

Orbea cooperi and O. tapscottii are closely related.

The main distinguishing characters are listed in Table

1 for comparison. It lias been proposed that because of

the absence of marginal cilia in the Devon population,

it may warrant assignment of subspecific rank (Darrel

Plowes pers. comm.). The authors are, however, uncer-

tain whether this would be wise, as specimens from the

Leeukop population do possess marginal cilia. Although

no other populations between Devon and Leeukop have

been located, the habitats are similar to each other and

markedly dissimilar to other habitats of O. cooperi.

Proper investigation of a large quantity of flowers at each

population needs to be done in order to resolve this issue.

This new distribution is, however, disjunct from any

of the other known collections made of this species.

This may indicate (as for many other species) that vast

areas still need to be surveyed in order to ascertain the

complete distribution range of this taxon (and many oth-

ers). In addition, the vegetation at the newly discovered

locations in Mpumalanga is open grassland with pre-

dominantly black turf interspersed by dolerite outcrops

and intrusions, which makes that habitat quite unusual

for this taxon. The distribution of Orbea cooperi and its

nearest relative O. tapscottii is given in Figure 5. The

listed new records all come from an area between the

distribution ranges of these two sister species. Orbea

tapscottii is usually associated with acacia savanna and

O. cooperi is mainly found in both karoo and acacia

savanna vegetation.

Voucher specimens of new collections of Orbea cooperi

GAUTENG. — 2628 (Johannesburg): Highveld Bridge, Farm Leeuwkop,

±2.15 km directly N of Devon, 26°20'06.7" S, 28°47'01.0" E, (-BD), fl.

13-11-2008, Bester 8539 (PRE!).

MPUMALANGA. — 2628 (Johannesburg): Balfour Dist., Farm

Platkop just outside Greylingstad, 26°4T00" S, 28°5T41" E, (-DB),

fl. 29-11-2009, Berruti 53A (PRE!); 26°4T02.58" S, 28°51'41.46" E,

FIGURE 6.— Flower and stems of Orbea cooperi from Leeukop locality, Bester 8539: A, vertical view; B, angled side view. Scale bars: A, B, 6

mm.

Bothalia 41,2 (201 1 )

297

Berruti 53B (PRE!); 26°40'59.76" S, 28°51'42.6" E, fl. 3-10-2009, Bei-

ruti 52 (PRE!); Lucas Kloppers' farm, 26°39’49.92" S, 28°51'57.78" E

(sight record, S. Berruti); Farm Platkop, 543 1R, rocky area adjacent

to district road. 26°39'46" S, 28°51'54" E, fl. 22-02-2009, McCleland

PRE850944 (PRE! in spirits).

ACKNOWLEDGEMENTS

Tony de Castro, Warren McCleland, Charles Craib

and Gillian Condy are thanked for bringing the location

of this species to our attention. We are grateful to Mpu-

malanga Parks Board for granting a collection permit for

obtaining the material.

REFERENCES

BRUYNS, P.V. 2002. Monograph of Orbea and Ballvanthus ( Apocynace-

ae-Asclepiadoideae-Ceropegieae). Systematic Botany Mono-

graphs 63: 1-195.

BRUYNS, P.V. 2005. Stapeliads of southern Africa and Madagascar ,

vol. 1. Umdaus Press, Hatfield, Pretoria.

MEVE, U. 2009. Taxonomic changes in Arabian taxa of Orbea Haw.

(Ceropegieae). Asklepios 105: 19-22.

PLOWES, D.C.H. 2004. Orbea elegans Plowes sp. nov. , an attractive

new stapeliad from northern South Africa (Apocynaceae-Ascle-

piadoideae-Ceropegieae). Asklepios 90: 14—17.

PLOWES, D.C.H. 2007. Two new stapeliads from Yemen. Excelsa 21:

11-14.

PLOWES, D.C.H. 2008. Angolluma taitica (Bruyns) Plowes comb. nov.

Asklepios 103: 4.

PLOWES. D.C.H. 2009. The Transkei Orbea saga. Asklepios 105: 9-18.

PLOWES. D.C.H. 2010. When is a Cara/luma not a CarallumaP. Ask-

lepios 107: 3-22.

RAFFAELLI, M„ MOST1, S. & TRADELLI, M. 2008. Apocynaceae of

Oman: Orbea nardii, sp. nov. and Pentatropis bentii, first find-

ing. Webbia 63: 161-167.

S.P. BESTER* and S.M. BERRUTI**

* Author for correspondence: National Herbarium (PRE), South Afri-

can National Biodiversity Institute, Private Bag XI 01, 0001 Pretoria.

E-mail: S.Bester@sanbi.org.za.

** School of Environmental Sciences and Development, Private Bag

X6001, North-West University, 2520 Potchefstroom / P.O. Box 426,

2415 Greylingstad.

MS. received: 2010-08-18.

HYACINTHACEAE

ORNITHOGALUM LEBAENSE TRANSFERRED TO ALBUCA

Extensive phylogenetic analysis of nuclear and plas-

tid DNA sequence data has unequivocally demonstrated

that Ornithogalum L. is paraphyletic as traditionally

circumscribed (e.g. Obermeyer 1978). Two alternative,

generic treatments for subfamily Omithogaloideae have

recently been proposed, both derived essentially from

the same data set and representing alternative hierar-

chical treatments of comparable monophyletic groups.

The more conservative (Manning et al. 2009) recog-

nizes four genera: Dipcadi Medik. and Pseudogaltonia

(Kuntze) Engl, remain as conventionally circumscribed;

Albuca L. is enlarged to include Ornithogalum subgen.

Osmyne and subgen. Urophyllotr, and Ornithogalum

includes the remaining species in the genus plus Galto-

nia Decne. and Neopatersonia Schonland. In this sys-

tem. the major lineages within Albuca and Ornithoga-

lum are treated as subgenera and sections. The second

option ( Martinez- Azorin et al. 2011) is a much more

radical departure from the traditional classification,

treating all major lineages at generic level, resulting in

the recognition of 19 genera in the subfamily, many of

them new. Although theoretically defensible, the practi-

cal value of this option in sub-Saharan Africa is severely

limited by the low level of morphological support for

several of the genera. One consequence of this is that

the South African species O. toxicarium C. Archer &

R.H. Archer remains unplaced in this classification until

molecular data are forthcoming.

The recently described Ornithogalum lebaense Van

Jaarsv. (2010) from Angola is currently unplaced in

either classification but is closely allied to the taxon

previously treated as O. longibracteatum Jacq. (Ober-

meyer 1978) and now treated either as Albuca bracte-

ata (Thunb.) J.C. Manning & Goldblatt (Manning et al.

2009) or as Stellarioides canaliculata Medik. (Martinez-

Azorin et al. 2011). As we have already argued (Man-

ning et al. 2009), we favour a more conservative treat-

ment of the genera and have already implemented this

treatment in various local floras. We therefore advo-

cate treating this species in Albuca subgen. Urophvllon

(Salisb.) J.C. Manning & Goldblatt and provide the req-

uisite new combination to facilitate this.

Albuca lebaensis (Van Jaarsv.) J.C. Manning

& Goldblatt , comb. nov. Ornithogalum lebaense Van

Jaarsv. in Herbertia 64: 92 (2010).

REFERENCES

MANNING. J.C., FOREST, F., DEVEY, D.S., FAY, M.F. & GOLD-

BLATT. P. 2009. A molecular phylogeny and a revised classifica-

tion of Omithogaloideae (Hyacinthaceae) based on an analysis

of four plastid DNA regions. Taxon 58: 77-107.

MARTINEZ-AZORIN, M„ CRESPO, M.B., JUAN, A. & FAY, M.

2011. Molecular phylogenetics of subfamily Omithogaloideae

(Hyacinthaceae) based on nuclear and plastid DNA regions,

including a new taxonomic arrangement. Annals of Botany 107:

1-37.

OBERMEYER, A. A. 1978. Ornithogalum: a revision of the southern

African species. Bothalia 12: 323-376.

VAN JAARSVELD, E.J. 2010. Ornithogalum lebaense, a new cliff-

dwelling Ornithogalum (Hyacinthaceae) from southwest Angola

(Benguella Province). Herbertia 64: 91-103.

J.C. MANNING* and P. GOLDBLATT**

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town / Research Centre for

Plant Growth and Development, School of Biological and Conservation

Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag

X01, 3209 Scottsville, South Africa.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

den. P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2011-03-31.

298

FABACEAE

A NEW SPECIES OF XIPHOTHECA FROM THE WESTERN CAPE, SOUTH AFRICA

Bothalia 41,2 (201 1 )

Xiphotheca Eckl. & Zeyh. is a papilionaceous genus

which belongs to the tribe Podalyrieae, together with

Amphithalea Eckl. & Zeyh., Liparia L., Podalyria Lam.,

Stirtonanthus B.-E.van Wyk & A.L.Schutte, Cyclopia

Vent., Virgilia Poir. and Calpurnia E.Mey. (Schutte &

Van Wyk 1 998).

In a taxonomic revision of the genus, Schutte (1997)

recognized nine distinct species, all of which are

endemic to the Cape Floristic Region of South Africa.

Based on a cladistic analysis of morphological data, two

sections were established within the genus: Xiphotheca

sect. Congestae A.L.Schutte, comprising four species

(all with congested, non-pedunculate inflorescence units

and the lower calyx lobe longer than the lateral lobes)

and X. sect. Xiphotheca, comprising five species (all

with extended and pedunculate flowering units and the

lower calyx lobe as long as the lateral lobes).

While doing field work during 2000 and 2001 as part

of the Cape Lowland Project of the Botanical Society of

South Africa, Nick Flelme collected an odd specimen in

the Overberg area. Follow-up field work confirmed that

it is indeed an unknown species of Xiphotheca. This spe-

cies is described below.

Xiphotheca rosmarinifolia A.L.Schutte , sp. nov.,

X. phylicoidi A.L.Schutte & B.-E.van Wyk similis, sed

foliis angustioribus, floribus minoribus, pedunculis,

pedicellis bracteisque brevioribus differt.

TYPE. — Western Cape, 3420 (Bredasdorp): 33 km

NE of Bredasdorp on Farm Plaatjieskraal 54; 0.6 km

NW of farmhouse on road to Sonderkoskop, 240 m, (-

AD), 1 Aug. 2001, N.A. Helme 2086 (NBG, holo.).

Erect, multi-stemmed shrub, up to 0.5 m tall, sprout-

ing from woody rootstock after fire. Branches seri-

ceous, glabrescent. Leaves alternate, simple, 10-16 x

1-2 mm, lamina linear, with strongly revolute margins,

pubescent on adaxial surface, glabrescent, densely seri-

ceous on abaxial surface, apex acute; petiole ± 1 mm

long, sericeous; stipules inconspicuous. Inflorescences

axillary, with paired (geminate) flowers, grouped into

flowering units of up to 20 flowers; peduncle short, < 1

mm long, sericeous. Bracts linear, ± 1 mm long, fused

at base with pedicel for 0.5 mm. Pedicel 3 mm long,

sericeous. Bracteoles minute, situated at thickening on

pedicel. Corolla yellow, fading brown with age, longer

than calyx, glabrous. Calyx narrowed to base, not intru-

sive; lobes triangular, acuminate; upper two lobes fused

higher up than lower three lobes; lower lobe as long as

upper four lobes, pubescent. Standard petal suborbicu-

lar, apex emarginate, base cuneate. Wing petals oblong,

longer than keel; auricle weakly developed, pocket

developed as a thickened lobe towards inside, incon-

spicuous on outer surface. Keel petals oblong, pocket

weakly developed, apex obtuse. Stamens diadelphous,

vexillary filament free; anthers ± uniform in shape and

size, alternately dorsifixed and sub-basifixed. Ovary

with 2 or 3 ovules, densely sericeous; pistil sessile; style

slender, curved upwards, glabrous. Pods coriaceous.

obliquely oblong, laterally compressed, constricted

between seeds, 2- or 3-seeded, densely pubescent, gla-

brescent. Seeds oblong-reniform, brown to deep dark

brown; hilum elliptic, surrounded by a fleshy collar-like

aril. Flowering time : August (flowering after fire). Fig-

ure 7.

Diagnostic features and affinities : Xiphotheca ros-

marinifolia falls within X. sect. Xiphotheca, owing to

its characteristic pedunculate inflorescences, extended

flowering units and the lower lobe of the calyx being as

long as the lateral lobes. Within this section, it is closely

related to X. phylicoides — both species are sprouters and

both have leaves with strongly revolute margins. How-

ever, X. rosmarinifolia differs from X. phylicoides in the

narrower and linear leaves, 1-2 mm wide, the shorter

peduncle (< 1 mm long), bracts (± 1 mm long) and pedi-

cel (3 mm long). In X. phylicoides the leaves are 2. 5-5.0

mm wide and elliptic, the peduncle is 1.5-2. 5 mm long,

the bracts are 3^1 mm long and the pedicel is 3. 5-5.0

mm long. The two species are allopatric, with the latter

recorded from three localities on the Outeniqua Mtns

near Mossel Bay, whereas X. rosmarinifolia is known

only from a single locality near Bredasdorp.

The specific epithet refers to the leaves that resemble

those of the well-known herb, rosemary.

Distribution and habitat, the species is known only

from the type locality on the Farm Plaatjieskraal near

Bredasdorp (Figure 8), where it was discovered by Nick

Helme in 2001. It grows in Renosterbos-Fynbos transi-

tional vegetation in a very dry habitat on the northwest-

ern slope of a silcrete hill in deep, pink, clayey soil with

white quartz pebbles.

The type material was collected in flower during

August 2001, following a fire in summer earlier that

year. This species appears to flower only within the

first year after a fire. Follow-up visits to the site during

spring in subsequent years confirmed this observation.

Conservation : Xiphotheca rosmarinifolia appears to

be a very restricted species, known from only a single

location in the Lower Breede River Valley. Despite addi-

tional field work in the area, no more populations could

be found. This species is seriously threatened because it

grows in an area of high agricultural potential that could

be ploughed in future. In addition, the area is subject to

inappropriate fire management and grazing by livestock.

Less than 200 mature individuals are known. Hence the

species is listed as Critically Endangered (CR) in Rai-

mondo et al. (2009).

According to Mucina & Rutherford (2006) the vegetation

type in the area is classified as Eastern Ruens Shale Renos-

terveld, which is classified as Critically Endangered, since

over 80 % of the original extent of this vegetation has been

lost, mainly for cropland. The Farm Plaatjieskraal harbours

one of the largest remaining patches of this vegetation type

in the Overberg area. Several other threatened species have

also been recorded from this site: Acmadenia macropetala

Bothalia 41,2 (201 1 )

299

FIGURE 7. — Xiphotheca rosmarinifolia. A— I, Helme 2086, NBG: A, flowering branch, showing arrangement of flowers; B, leaf abaxial view, show-

ing revolute margins; C, flower in lateral view; D, calyx (upper lobes to left); E, standard petal, adaxial view; F, wing petal; G, keel petal; H,

anthers; I, pistil. J, K, Vlok & Schulte 445, NBG: J, pod in lateral view; K, seed. Scale bars: A, J (at A), 5 mm; B-G, I, K (at B), 5 mm; H, 1

mm. Artist: A.L.Schutte-Vlok.

FIGURE 8. — Known distribution of

Xiphotheca rosmarinifolia.

(VU = Vulnerable), Acrodon deminutus (VU), Aspalathus

grobleri (EN = Endangered), Brownanthus fratermis (EN),

Drosanthemum quadratum (EN), Erica venustiflora subsp.

glandulosa (VU), Gibbaeum haaglenii (EN), Leucadendron

coriaceum (EN) and Relhania gamotii (VU) (pers. obs.;

N.A. Helme pers. comm. ). The site is therefore of very high

conservation significance and every effort should be made to

secure this site under the Stewardship programme of Cape-

Nature (Western Cape Nature Conservation Board).

Other material examined

WESTERN CAPE. — 3420 (Bredasdorp): Farm Plaatjieskraal, NE

of Bredasdorp, on silcrete hill NW of fannhouse on road to Sonderkos-

kop, (-AD), 09-09-2001, J.H.J. Vlok & A.L. Schutte 445 (NBG).

ACKNOWLEDGEMENTS

I am very grateful to Nick Helme for bringing the col-

lection to my attention. CapeNature is thanked for gran-

300

Bothalia 41,2 (2011)

ting me the opportunity to take sabbatical leave. This

publication stems from the revision of Xiphotheca which

formed part of a Ph.D. thesis in Botany at the Rand Afri-

kaans University (now University of Johannesburg). I

am indebted to Prof. B.-E. van Wyk for supervising the

study. My husband, Jan Vlok, kindly accompanied and

assisted me on field trips. Dr O. Leistner kindly assisted

with the translation of the diagnosis into Latin.

REFERENCES

MUCINA. L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of

South Africa, Lesotho and Swaziland. Strelitzia 19. South Afri-

can National Biodiversity Institute, Pretoria.

RAIMONDO, D„ VON STADEN, L„ FODEN, W„ VICTOR, J.E.,

HELME, N.A., TURNER, R.C., KAMUNDI, D.A. & MAN-

YAMA, P.A. (eds). 2009. Red List of South African plants 2009.

Strelitzia 25. South African National Biodiversity Institute, Pre-

toria.

SCHUTTE, A.L. 1997. A revision of the genus Xiphotheca (Fabaceae).

Annals of the Missouri Botanical Garden 84: 90-102.

SCHUTTE, A.L. & VAN WYK, B-E. 1998. Evolutionary relationships

in the Podalyrieae and Liparieae (Fabaceae) based on morpho-

logical, cytological and chemical evidence. Plant Systematics &

Evolution 209: 1-31.

A.L. SCHUTTE-VLOK*

* Department of Botany and Plant Biotechnology, University of Johan-

nesburg, P.O. Box 524, 2006 Auckland Park, Johannesburg. Present ad-

dress: CapeNature, Private Bag X658, 6620 Oudtshoom, South Africa.

MS. received: 2010-03-17.

CYPERACEAE

NEW NAMES AND NEW COMBINATIONS IN CYPERUS FOR SOUTHERN AFRICA

Since the genus Mariscus Vahl is no longer upheld

(Archer 2000) and the genus Monandrus Vorster was

never published, several taxa treated by Vorster (1978)

for southern Africa require new combinations or new

names in Cyperus L. in order to reflect current taxo-

nomic treatments incorporating molecular data (Muasya

et al. 2009; Huygh eta!. 2010).

1. Cyperus atriceps (Kiik.) C. Archer & Goetgh.,

comb. & stat. nov. Types: South West Africa [Namibia],

Grootfontein, Dinter 73 7 7 (B*, lecto., here designated-

PRE, photo.!; B** (second sheet), K, PRE!).

C. aristatus Rottb. var. atriceps Kiik. in Mitteilungen des Tluiring-

ischen Botanischen Vereins 2, 50: 8 ( 1943). Mariscus aristatus (Rottb.)

Cherm. var. atriceps (Kiik.) Podlech: 523 (1960).

Monandrus atriceps (Kiik.) Vorster, comb. nov. ined.: 333 (1978).

Vorster (1978) found this to be a distinctive species,

widely distributed in Namibia, Botswana and central

South Africa. In his key, based on fruit characters, he

distinguished it from Cyperus squarrosus L. (syn. C.

aristatus Rottb.) by ‘Fruits obovoid, covered with coni-

cal sparsely-spaced papillae’ vs ‘Fruits obpyramidal,

covered with closely-spaced flat-topped papillae’.

2. Cyperus austro-africanus C. Archer & Goetgh.,

nom. nov. Type: South Africa, Eastern Cape, Basche,

Drege 4383 (B, lecto., designated by Vorster (1981); B

(second sheet), P (three sheets), S).

Mariscus dregeamts Kunth, Enumeratio plantarum 2: 120 (1837);

Vorster: 245 (1978), non Cyperus dregeanus Kunth [= C. pulcher

Thunb.].

M. dregeanus var. buchananii C.B. Clarke: 188 (1897). Type: Natal

[KwaZulu-Natal], Durban Flat, Buchanan s.n. (K, holo.).

M. dubius sens. lat. : auctt. plur.

Due to the pre-existing Cyperus dregeanus Kunth, the

species requires a new name upon transfer to Cyperus

[cf. International Code of Botanical Nomenclature Art.

* Sheet bearing Kiikenthal’s determinavit label dated 1 2.vi. 1 937.

** Sheet bearing Kiikenthal’s determinavit label dated 2.i. 1 940.

53.1 (McNeill et al. 2006)]; the chosen epithet describes

the almost exclusively southern African distribution of

this taxon.

3. Cyperus decurvatus (C.B. Clarke) C. Archer

& Goetgh., comb. & stat. nov. Type: Mocambique

[Mozambique], Ressano Garcia, Schlechter 11952 (K,

holo.-PRE, photo.!; B, BOL, BR, G, GRA, PRE!, Z).

Mariscus vestitus (Hochst.) C.B. Clarke var. ? decurvata C.B. Clarke

in Botanischer Jahrbucher 38: 134 (1906). Cyperus indecorus Kunth

var. decun’atus (C.B. Clarke) Kiik.: 545 (1936). Mariscus indecorus

(Kunth) Podlech var. decurvatus (C.B. Clarke) Podlech: 524 (1960).

C. indecorus Kunth var. dinteri Kiik.: 545 (1936). Mariscus inde-

corus (Kunth) Podlech var. dinteri (Kiik.) Podlech: 524 (1960). Type:

South West Africa [Namibia], Gneisberge bei Farm Hoffnung, 1 700

m, Dinter 2724 (B, SAM).

Mariscus rehmannianus C.B. Clarke: 196 (1898); Vorster: 280

(1978), non Cyperus rehmannianus (C.B. Clarke) Kuntze [= Pycreus

rehmannianus C.B. Clarke], Type: South Africa, Transvaal [Gauteng],

lagoon near Pienaars River, Nelson 13 (K-PRE, photo.!); Pretoria,

Rehmann 4479 (K-PRE, photo.!, Z-PRE, photo.!); Transvaal, without

precise locality, Holub s.n. (K-PRE, photo.!).

According to Vorster (1978: 280), the sheet of Sch-

lechter 11952 at B has the name Mariscus rehman-

nianus inscribed in Clarke's handwriting. There is no

inscription on any of the sheets relating to M. vestitus

var. decurvatus ; however, the K sheet, which has ample

material, is presumed to be the holotype.

4. Cyperus palmatus (Lye) C. Archer & Goetgh.,

comb. & stat. nov. Type: Tanganyika [Tanzania], Ufipa

Dist., Milepa-Zimba, Bullock 3625 (K, holo.).

C. usitatus Burch, subsp. palmatus Lye in Nordic Journal of Bot-

any 3: 228, fig. 22 (1983).

C. fulgens C.B. Clarke var. contractus Kiik.: 122 (1935), non Cype-

rus contractus Steud. Type: South West Africa [Namibia], Amboland

[Ovamboland], Otjituo, auf dem sandige Boden des Omuramba-Oma-

lako, Dinter 894 (B, holo.-PRE, photo.!; K-PRE, photo.!, SAM-PRE,

photo.!).

C. fulgens sens, lat.: Podlech: 14 (1967).

C. palmatus Vorster, sp. nov. ined.: 347 (1978).

Bothalia 41,2 (2011)

301

In addition to Tanzania, this species is recorded from

northern South Africa, Zambia and Kenya. The first

author has not seen the type or cited specimens of Cype-

rus usitatus subsp. palmatus, but is convinced from the

description and illustration provided that the synonymy

listed above all relates to this taxon. It appears to be

coincidental that Vorster (1978) used the same epithet

for his new species as Lye (1983) did for his new sub-

species.

The species undoubtedly belongs to the group includ-

ing Cyperus fulgens and C. usitatus (plus several more

in tropical Africa), all of which produce small, tunicated

corms (not often collected) distant from the plant base at

the tips of very thin, easily broken rhizomes. This group

does require revision, but C. palmatus has such a dis-

tinctive inflorescence form (aptly described by the spe-

cific epithet) that in the interim it is preferable to treat it

at species level.

5. Cyperus uitenhagensis (Steud.) C.Archer &

Goetgh ., comb. nov. Type: South Africa, Eastern Cape,

Uitenhagen [Uitenhage], Zeyher 91 (B, holo.-PRE,

photo.!).

Mariscus uitenhagensis Steud., Synopsis plantarum glumacearum.

2. Cyperaceae: 317 (1855); Vorster: 253 (1978).

Mariscus marlothii (Boeck.) C.B. Clarke var. globospica

C.B. Clarke: 187 (1897). Cyperus capensis (Steud.) Endl. var. pseu-

domarlothii forma globospica (C.B.Clarke) Kuk.: 540 (1936). Type:

South Africa, Western Cape Province, Uniondale Div., rocky hill near

Groot River, Burchett 5020 (K, holo.: PRE [fragment]!).

C. capensis (Steud.) Engl. var. pseudomarlothii K.iik.: 540 (1936).

Types: South Africa, Transvaal [Gauteng], Pretoria, Rehmann 4035 (B,

K, Z), 4728 (not seen); Mogg 578 (K. PRE!); [Pretoria], Hiigel liber

Aapies Rivier, Rehmann 4324 (B. Z); Johannesburg, Moss 5924 (J, K);

Transvaal [Mpumalanga], Witbank. Rogers 2569 (J, GRA, Z); Natal

[KwaZulu-Natal], Pietermaritzburg. Wilms 2359 (K); Mozambique,

Ressano Garcia bei Lourenco Marques, Schlechter 11926 [= C. austro-

africanus\, 11950 (B, Z). Note: C. capensis var. polyanthemus Kuk.:

540 (1936) was treated as a synonym of Mariscus chersinus N.E.Br.

[Cyperus chersinus (N.E.Br.) Kuk.].

Mariscus marlothii sensu C.B.Clarke: 590 (1894) non (Boeck.)

C.B.Clarke.

M. capensis sensu Schonland: 29 (1922), p.p., non (Steud.) Schrad.

REFERENCES

ARCHER, C. 2000. Cyperaceae. In O.A. Leistner, Seed plants of south-

ern Africa: families and genera. Strelitzia 10: 594-605.

CLARKE, C.B. 1894. Cyperaceae. In T. Durand & H. Schinz, Conspec-

tus florae Africae 5: 526-692. Charles vande Weghe, Brussels.

CLARKE, C.B. 1897. Cyperaceae. In W.T. Thiselton-Dyer, Flora cap-

ensis 7: 149-192. Reeve, Kent.

CLARKE, C.B. 1898. Cyperaceae. In W.T. Thiselton-Dyer, Flora cap-

ensis 7: 193-310. Reeve, Kent.

CLARKE, C.B. 1906. Cyperaceae africanae. In A. Engler, Beitrage zur

Flora von Afrika 29. Botanische Jahrbiicher 38: 131-136.

HUYGH. W„ LARRIDON, I„ REYNDERS, M„ MUASYA, A.M.,

GOVAERTS, R„ SIMPSON, D.A. & GOETGHEBEUR, P.

2010. Nomenclature and typification of names of genera and

subdivisions of genera in Cypereae (Cyperaceae): 1. Names of

genera in the Cyperus clade. Taxon 59: 1883-1890.

KUNTH, C.S. 1837. Enumeratio plantarum 2. Cotta, Stuttgart.

KUKENTHAL, G. 1935. Cyperaceae-Scirpoideae-Cypereae. In A.

Engler, Das Pflanzenreich Fam. IV, 20, Heft 101 : 1-160. Engel-

mann, Berlin.

KUKENTHAL, G. 1936. Cyperaceae-Scirpoideae-Cypereae. In A.

Engler, Das Pflanzenreich Fam. IV, 20, Heft 101: 161-671.

Engelmann, Berlin.

KUKENTHAL, G. 1943. Neue oder nicht geniigend bekannte Cypera-

ceen. Mitteilungen des Thuringischen Botanischen Vereins N.F.

50: 1-13.

LYE, K.A. 1983. Studies in African Cyperaceae 25. New taxa and com-

binations in Cyperus L. Nordic Journal of Botany 3: 213-232.

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., W1ERSEMA, J.H. &

TURLAND, N.J. 2006. International Code of Botanical Nomen-

clature (Vienna Code). Regnum Vegetabile 146.

MUASYA, A.M., VRIJDAGHS, A.. SIMPSON, D.A., CHASE, M.W.,

GOETGHEBEUR, P. & SMETS, E. 2009. What is a genus in

Cypereae: phytogeny, character homology assessment and gener-

ic circumscription in Cypereae. Botanical Review 75: 52-66.

PODLECH, D. 1960. Ubereinige Cyperaceen Sudafrikas. Mitteilungen

der Botanischen Staatssammlung Miinchen 3: 521-530.

PODLECH, D. 1967. Cyperaceae. In H. Merxmuller, Prodromus einer

Flora von Siidwestafrika 165. Cramer, Lehre.

SCHONLAND, S. 1922. Introduction to South African Cyperaceae.

Memoirs of the Botanical Survey of South Africa No. 3. Govern-

ment Printer, Pretoria.

STEUDEL, E.G. 1855. Synopsis plantarum glumacearum 2. Cyper-

aceae. Metzler, Stuttgart.

VORSTER, P.J. 1978. Revision of the taxonomy o/Mariscus Vahl and

related genera in southern Africa. D.Sc. thesis, University of

Pretoria.

VORSTER, P.J. 1981. The identity and typification of Mariscus dregea-

nus. Bothalia 13: 444—446.

C. ARCHER* and P. GOETGHEBEUR**

* South African National Biodiversity Institute, Private Bag X 1 0 1 , 000 1

Pretoria. E-mail: c.archer@sanbi.org.za.

** Department of Biology, Research Group Spermatophytes, Ghent

University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium.

MS. received: 2011-04-20.

SCROPHULARIACEAE

ANTICHARIS JUNCEA, AN OVERLOOKED RECORD FOR SOUTH AFRICA, WITH NOTES ON ITS TYPE LOCALITIES AND

FLOWER MORPHOLOGY

The genus Anticharis Endl. consists of± 10 species in

Africa, Arabia and India, six of which are known from

the Flora of southern Africa region (Smithies 2000).

The plants are small, erect or suberect perennials (rarely

annuals), sometimes suffruticose herbs or dwarf shrubs,

glabrous to glandular-pubescent with alternate, entire

leaves. The purple corolla tube is elongated and broad

with a wide open, usually white throat. Anticharis is eas-

ily distinguished from the genera Aptosimum Burch, ex

Benth. and Peliostomum E.Mey. ex Benth (all members

of tribe Aptosimeae) by having only two fertile stamens

and not four (Hiem 1904; Smithies 2000).

In recent floristic inventories only two species of

Anticharis , namely A. scoparia (E.Mey. ex Benth.)

Hiern ex Benth. & Hook.f. and A. senegalensis (Walp.)

Bhandari, are listed as occurring in South Africa (in

Northern Cape and in Northern Cape and Limpopo

respectively) (Smithies & Ready 2003; Smithies 2006).

302

Bothalia 41,2 (2011)

A specimen of A.juncea L. Bolus (Steyn 1696), collected

during a field trip to the Northern Cape in August 2009,

was initially thought to be the first record of the species

for South Africa. Up to now A. juncea (= A. genistoides

Schinz) was considered in regional inventories as being

confined to Namibia (Smithies & Ready 2003). The

purpose of this contribution is to report on the floristic

significance of this find, to provide clarity on the origi-

nal collecting localities of the species and to designate a

lectotype. The availability of live material also permits

a more complete morphological description of the flow-

ers, and we provide the first close-up images of the fresh

flowers to be published for the species.

An investigation into the localities of the specimens

of Anticharis juncea housed in PRE and NBG proved

informative. These collections are incorporated in the

National Herbarium’s Pretoria (PRE) Computerised

Information System (PRECIS), a mega-database used,

amongst others, to produce checklists of the South Afri-

can flora and to plan itineraries of collecting trips. Most

collections of A. juncea are from the Warmbad District

(Namibia) but the protologue of A. juncea cites three

syntypes, Pearson 3609 and 3600 from Bushmanland

and Pearson 4002 from Great Namaqualand, south of

Warmbad, Namibia (Brown et al. 1915). The specimens

from Bushmanland were collected near a place called

Wortel in January 1909. There are a number of refer-

ences to a place with the name of Wortel. These range

from the Rehoboth area (Bethanie District) of Namibia

in the north to Gamoep further south in the Namaqua-

land part of the Northern Cape of South Africa. A close

look at Pearson’s itinerary reveals that he was collecting

in Bushmanland (South Africa) during December 1908

and January 1909. Only in late January 1909 did he start

collecting in the Bethanie District, Namibia. It is very

likely that Pearson’s specimens from Bushmanland were

collected near the Farm Wortel (2918BB), northwest of

Pella, South Africa. Pearson was therefore not only the

first botanist to have discovered A. juncea , but also the

first to have collected herbarium material of this species

in South Africa, a record which has hitherto not been

reflected by floristic inventories.

FIGURE 9. — Anticharis juncea ,

close-up of corolla, A, side

view; B. view from below.

Scale bars: 5 mm. Photo-

graphs: M. Koekemoer.

Bothalia 41,2 (2011)

303

Anticharis juncea is a glabrous, perennial shrublet

up to 300 mm high, with striate stems that are almost

leafless to distantly leafy, with alternate, narrow, linear

leaves, 2-5 * 1 mm. The flowers are axillary, solitary,

tubular and carried on a bi-bracteolate pedicel, 2-4 mm

long. The corolla tube is 18-21 mm long, purple, nar-

row at the base and dilated distally, with two shallow,

lateral pouches (or pockets) near the base, a prominent,

dorsal pouch towards the distal end, a marked diagonal

constriction, best seen from the side, and a longitudi-

nal. ventral groove in the proximal half (Figure 9A, B).

These floral features are clearly visible in live mate-

rial but are obscured when the flowers are pressed and

dried, although the presence of pouches can be vaguely

discerned. The two fertile anterior stamens are inserted

above the base of the corolla and extend a little beyond

the middle of the corolla tube. The capsule is ovate,

bisulcate with an acute apex and dehisces mainly sep-

ticidally but also loculicidally. Although the specific

pollinator(s) of A. juncea is unknown, there is a marked

and widespread association between pollen wasps

(Vespidae: Masarinae) and the members of the Scrophu-

lariaceae tribe Aptosimeae (Benth.) Benth. & Hook.f.,

namely Aptosimum Burch, ex Benth.. Peliostomum

Benth. and Anticharis (Gess & Gess 2004. 2010).

Although it most closely resembles the mainly annual

Anticharis senegalensis, A. juncea differs from the

former in being perennial, glabrous, with fewer, smaller

leaves and shorter pedicels (Brown et al. 1915). The

three South African species may be distinguished using

the following key:

1 a Plants perennial or annual, glandular-pubescent:

2a Plants usually annual; leaves linear, 10-40 mm long; pedi-

cels 8-12 mm long A. senegalensis

2b Plants perennial; leaves linear-lanceolate, 4-13 mm long;

pedicel 3-8 mm long A. scoparia

lb Plants perennial, glabrous; distantly leaved, leaves linear,

2-5 mm long; pedicel 2-A mm long A. juncea

Anticharis senegalensis [= A. linearis (Benth.)

Hochst. ex Asch.] has a wide and markedly disjunct

transequatorial range, occurring in Limpopo and North-

12 14 16 18 20 22 24 26 28 30 32

FIGURE 10. — Known distribution of Anticharis juncea , •; A. scopa-

ria, I; and A. senegalensis , A.

em Cape in South Africa, Botswana, Namibia (Figure

10), also in Angola and Zimbabwe in the south, then fur-

ther north in arid parts of North Africa through Arabia to

India. The distribution of A. juncea, on the other hand, is

much more restricted, with all known records confined

to a small area in the extreme south of Namibia and

adjacent parts of the Northern Cape. It is usually found

on stony or rocky soils within the Succulent Karoo

Biome (Rutherford & Westfall 1968), or the Desert

Biome in South Africa and the bordering Nama-Karoo

Biome in Namibia (Mucina & Rutherford 2006). Bio-

geographically, A. juncea is endemic to the Gariep Cen-

tre of Endemism (Van Wyk & Smith 2001), in particu-

lar the very hot and arid, deeply dissected region along

the Lower Orange River Valley, downstream from the

Augrabies waterfalls. The distribution of A. juncea over-

laps with that of A. scoparia, also a perennial and Gariep

Centre endemic, but a species with a more northwesterly

range (Figure 10). A. scoparia is glandular-pubescent

with linear-oblong leaves (Hiem 1904).

Anticharis juncea is an example of a species that was

overlooked and left out of South African floristic check-

lists for more than 100 years since its first discovery,

because of the incorrect interpretation of the localities

in the protologue. Computerized herbarium specimens

offer improved access to large quantities of data as well

as expanded analytical potential (Rhoads & Thomp-

son 1992). However, to be analysed spatially, botanical

data must have locality information that can be related

to a point on a map. It is very important that the locality

information should be related to the correct point on a

map.

Anticharis juncea L. Bolus in N.E.Br., L. Bolus &

E. Phillips in Annals of the South African Museum 9,4:

263, 264 (1915). Type: Northern Cape, 2918 (Gamoep):

Bushmanland, Wortel, pass near base of kopje (-BB),

10 Jan. 1909, Pearson 3600 (BOL, scan!, Iecto., desig-

nated here).

Note : all three syntypes mentioned in the protologue

are mounted together on the same sheet in BOL where

Louisa Bolus worked. The three collections are, however,

clearly delineated by pencil lines and each has its own

label. The syntype Pearson 3600 is considered the bet-

ter specimen and is chosen here as lectotype. There are

two sheets of the syntype Pearson 3609 in NBG [marked

I and II]. Sheet II represents Anticharis juncea and

although it carries the number 3609 , is rather confusingly

numbered and apparently contains material from two dif-

ferent collections. On one of the two specimens mounted

on this sheet is a field note with the number 5413. At a

later stage, someone has written 4002 on it. this number

being that of the third Pearson syntype.

Other specimens seen

NAMIBIA. — 2818 (Warmbad): Great Namaqualand [southern

Namibia], kopje south of Warmbad, (-CD), 27 Jan. 1909, Pearson

4002 (BOL, scan!); Goodhouse Poort, (-CD), 22 June 1989, Van Wyk

8639 (PRE). 2819 (Ariamsvlei): Warmbad Dist., 16 miles [25.6 km]

SSW of Charly’s Puts, Geinab Gorge, (-BA), 17 May 1955, Acocks

18173 (PRE); Warmbad Dist., Farm Vellorsdrift: WAR 93, Granitberge

am Oranje bei Brucke, (-CB), 17 May 1963, Giess, Volk & Bleiss-

ner 7051 (PRE); Warmbad, ± 1 km NW of Orange River along road

between Onseepkans and Karasburg, (-CB), 1 Feb. 1974, Davidse de

Loxton 6197 (PRE).

304

Bothalia 41,2 (2011)

NORTHERN CAPE. — 2918 (Gamoep): ridge E of Goodhouse

turn-off en route to Klein Pella, (-BA), 23 Aug. 2009, Steyn 1696

(PRE); Bushmanland, near Wortel, (-BB), 10 Jan. 1909, Pearson 3609

(BOL, scan!; NBG Sheet I, scan!).

ACKNOWLEDGEMENTS

The author would like to thank colleagues for helpful

comments and suggestions. My sincere thanks to Prof.

Braam van Wyk who greatly improved the manuscript.

Grateful thanks to Terry Trinder-Smith and Edwina

Marais for help in locating herbarium specimens in BOL

and NBG, respectively. A collecting permit was provided

by Northern Cape Nature Conservation.

REFERENCES

BROWN, N.E., BOLUS, L. & PHILLIPS. E.P. 1915. Scrophulariaceae.

Annals of the South African Museum 9,4: 263, 264.

GESS, S.K. & GESS, F.W. 2004. Distributions of flower associations of

pollen wasps (Vespidae: Masarinae) in southern Africa. Journal

of Arid Environments 57: 17-44.

GESS, S.K. & GESS, F.W. 2010. Pollen wasps and flowers in southern

Africa. SANBI Biodiversity Series 18. South African National

Biodiversity Institute, Pretoria.

HIERN. W.P. 1904. Scrophulariaceae. In W.T. Thiselton-Dyer, Flora

capensis 4,2: 121—420. Reeve, London.

MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South

Africa, Lesotho and Swaziland. Strelitzia 19. South African

National Biodiversity Institute, Pretoria.

RHOADS, A.F. & THOMPSON, L. 1992. Integrating herbarium data

into a geographic information system: requirements for spatial

analysis. Taxon 41 : 43—4-9.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Botanical Research Insti-

tute, Pretoria.

SMITHIES, S.J. 2000. Scrophulariaceae. In O.A. Leistner, Seed plants

of southern Africa: families and genera. Strelitzia 10: 508-537.

SMITHIES, S..I. 2006. Scrophulariaceae. In G. Germishuizen, N.L.

Meyer, Y. Steenkamp & M. Keith, A checklist of South African

plants. Southern African Botanical Diversity Network Report

No. 41. SABONET, Pretoria.

SMITHIES, S.J. & READY, J.A. 2003. Anticharis. In G. Germishuizen

& N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 863. National Botanical Institute, Pretoria.

VAN WYK, A.E. & SMITH, G.F. 200 1 . Regions offloristic endemism in

southern Africa. A review with emphasis on succulents. Untdaus

Press, Hatfield, Pretoria.

H.M. STEYN*

* National Herbarium, South African National Botanical Institute, Pri-

vate Bag XI 01, 0001 Pretoria.

MS. received: 2010-09-07.

ASTERACEAE

LACHNOSPERMUM NEGLECTUM (ASTEROIDEAE: GNAPHALIEAE), ANEW AND OVERLOOKED SPECIES FROM THE

WORCESTER VALLEY, WESTERN CAPE

Lachnospermum Willd., a small genus in the Meta-

lasia group of tribe Gnaphalieae, is distinguished by its

solitary or few, relatively large, campanulate capitula;

multicellular hairs on the outside (abaxial) surface of the

corolla lobes; long, unicellular hairs on the cypselas; and

a rim or shallow cup at the top of the cypsela surround-

ing the pappus (Hilliard & Burtt 1981; Ward et al. 2009).

It currently comprises three named species (Koekemoer

2002), although a fourth unnamed species was included

by Goldblatt & Manning (2000) in their enumeration of

the genus as L. sp. 1 . Vegetatively similar to L. fascicula-

tum (Thunb.) Baill., this taxon had been identified by the

German botanist Rudolf Schlechter as a distinct species

under the manuscript name L. neglectum , based on a col-

lection of his from Brandvlei near Worcester made in the

late nineteenth century. Schlechter was familiar with both

species, having collected true L. fasciculatum a short dis-

tance further up the Breede River Valley just four years

earlier, but the superficial similarity between the taxa led

most botanists to identify both as L. fasciculatum , with

the signal exception of British botanist N.E. Brown. The

earliest collection of the Brandvlei taxon appears to have

been made by W. Tyson in the Hex River Valley in 1881,

and an indication that N.E. Brown concurred with Sch-

lechter in considering it to represent a novelty, is borne

out by an annotation on the duplicate at the Bolus Her-

barium to this effect by Cape Town botanist Harry Bolus,

dated lOJanuary 1903.

The Brandvlei taxon has now been collected from

multiple different populations in the Worcester and Hex

River Valleys. Although vegetatively similar to Lachno-

spermum fasciculatum , these populations differ consist-

ently from that species in several features, notably the

size and shape of the capitula, the colour and disposition

of the involucral bracts, and the size of the florets. We

follow Schlechter in treating them as a distinct species

under his epithet neglectum , which is now even more

apposite than it was at the time that he suggested it.

Specimens from BOL, NBG, PRE and SAM, the main

herbaria with good representation of Cape species, were

consulted for records of the two new species (herbarium

acronyms after Holmgren et al. 1990). The Aluka website

(www.aluka.org) was searched for duplicates of the Sch-

lechter collection of Lachnospermum neglectum.

Lachnospermum neglectum Schltr. ex J.C. Manning

& Goldblatt, sp. nov.

Lachnospermum fasciculato (Thunb.) Baill. habitu

suffruticoso intricate ramoso similis sed capitulis late

cylindricis, 7-9 x 6-8 mm, paleis ± 10 subulato-pilosis,

phyllariis ± 100 in seriebus ± 12 imbricatis extends 1-2

mm longis intends 6. 0-7.0 x 1.5 mm oblongis margini-

bus membranaceis distaliter dense coactis in laminam

attenuatam subulatam deflexam stramineam 0. 5-2.0 mm

longam contractis, flosculis flavis ±5.5 mm longis, et

cupula apical i ovarii cypselarumque symmetrica differt.

TYPE. — Western Cape, 3118 (Worcester): ‘in colli-

bus aridis prope Brand Vlei’ [Brandvlei], (-CB), 9 Jan.

1896, Schlechter 9934 [K, holo.; BOL, HBG, PRE (2

sheets), S, Z, iso.].

Intricately branched shrublet, 400-800 mm high;

branches often spreading horizontally, densely whitish

Bothalia 41,2 (2011)

305

or yellowish tomentose with matted hairs, ± sparsely

leafy. Leaves ericoid, spreading or declinate, with

rosette-like axillary fascicles ± half as long, to as long

as subtending leaf, involute, narrowly lanceolate-eri-

coid, 2. 0-2. 5 x 1 mm. acute or apiculate, mostly ± half

twisted, adaxial surface felted, abaxial surface arach-

nose. Capitula discoid, 7-9 x 6-8 mm, terminal on pri-

mary and secondary branches, solitary or up to three in

umbellate clusters, ± 20-flowered. Involucre broadly

campanulate, 7-9 x 6-8 mm; bracts ± 100 in ± 12 imbri-

cating series, progressively longer acropetally, outermost

1-2 mm long, innermost 6. 0-7.0 x 1.5 mm, oblong with

cartilaginous stereome and membranous margins, gla-

brous in basal half but densely felted in distal half with

cohering, white hairs, tapering into an attenuate, sharply

deflexed. brown-papery, awl-like lamina 0. 5-2.0 mm

long. Receptacle paleate; paleae ± 10, setaceous, ± 6 mm

long, sparsely villous distally. Florets yellow, ± 5.5 mm

long; lower part of tube cylindrical. 3 mm long, limb

narrowly campanulate, 5-lobed. lobes lanceolate, ± erect

and not spreading, ± 1 mm long with thickened margins,

ciliate along inner margin and penicillate apically with

sparse gland-tipped hairs on abaxial surface. Anthers

2.5 mm long including ovate apical appendage, base

tailed. Ovary cylindrical with shallow, fimbriate crown,

6-winged, wing ribs densely pubescent; style terete with

swollen base on distinct stylopodium, branching just

below mouth of tube, branches ± 1 mm long, lateral

margins stigmatic, apices truncate, penicillate. Cypsela

cylindric with six longitudinal ribs or wings and shallow

8-10-lobed crown, 3. 0-3. 5 x 1.5 mm. ribs asymmetri-

cally disposed, with one median rib on inner face and

five ribs congested on outer face, ribs densely villous

along ridges with long, eglandular hairs in several rows

plus scattered gland-tipped hairs, crown fimbriate and

adpressed scabridulous, pale straw-coloured. Pappus

bristles ± 80, connate basally and also shortly fused into

fascicles, ± 4.5 mm long, barbellate with apical barbs

longer. Flowering time'. Jan.-Feb. Figures 1 1 A-I; 12.

Distribution and ecology'. Lachnospermum neglec-

tum is confined to the upper Breede River Valley and its

tributaries, where it has been collected from around De

Dooms in the Hex River Valley and between Worcester

and Robertson in the Breede River Valley itself, extend-

ing onto the lower northern slopes of Jonaskop at the

western end of the Riviersonderend Mtns (Figure 13).

Plants are recorded as locally common in transitional

renosterveld-fynbos on sandstone alluvium and on sandy

flats, from 200-600 m, or on lower sandstone slopes.

Several other ericoid shrubs are narrow endemics of

the Worcester Valley, notably Euchaetis pungens (Ruta-

ceae), Polhillia obsoleta (Fabaceae) and Leucadendron

flexuosum (Proteaceae).

Diagnosis and relationships'. Lachnospermum neglec-

tum. like L. fasciculatum, is a well-branched, moderately

densely leafy shrublet, with small, ericoid leaves devel-

oping rosette-like axillary brachyblasts, and bearing

1-3 capitula per branch. The brown, scale-like limbs to

the involucral bracts set them apart from the remaining

two species, L. imbricatum (P.J.Bergius) Hilliard and L.

umbellatum (L.f.) Pillans, both of which have conspicu-

ously petaloid, white or pink bracts.

Lachnospermum neglectum is distinguished from L.

fasciculatum by its mostly markedly horizontal branches

and generally smaller involucres, 7-9 x 6-8 mm, obtuse

at the base, and by the sharply deflexed. pale straw-col-

oured, attenuate and awl-like, papery limbs to the bracts.

In L. fasciculatum the branches are mostly ± ascend-

ing and the capitula are larger, 10-15 x 8-11 mm, and

obconic at the base (Figure 1 1 J ), and the involucral

bracts have smaller, erect or suberect, scale-like, acute

or acuminate limbs that are chestnut-brown or plum-red.

The florets in L. neglectum are also substantially smaller

than in L. fasciculatum, ±5.5 mm long vs 7-9 mm long

(Figure 1 IK), and the apical cup on the ovary and cypse-

las is symmetrically developed on all sides vs asymmet-

rical and reduced to a rim on one side in L. fasciculatum

(Figure 11L; see also Koekemoer 2002: plate 31.1, fig.

14).

The two species are geographic vicariants, with Lach-

nospermum fasciculatum more widely spread through

the southwestern Cape. Considered until recently to be

endemic to the Cape Floristic Region, with its northern-

most station in the Pakhuis Mtns (Goldblatt & Manning

2000; Koekemoer 2002), L. fasciculatum has now been

collected north of the Olifants River at Brand se Baai on

the southern Namaqualand coast. From here it ranges

through the Cedarberg and Cold Bokkeveld to Vil-

liersdorp and Greyton at the southern foot of the Rivier-

sonderend Mtns (Figure 13), extending onto the West

Coast and the Cape Flats, from Mamre to Kuils River.

It is largely absent inland of the coastal ranges, except

at the head of the Breede River Valley, where it has been

collected from the foot of the Waaihoek Mtns. This sta-

tion is just 30 km northwest of the most northerly of the

known populations of L. neglectum.

Additional specimens examined

Lachnospermum neglectum

WESTERN CAPE. — 3319 (Worcester): in clivis montis Hex River

Valley, 1500' [450 m], (-BC), Jan. 1881, Tyson 786 (SAM); in con-

valle Hex River, prope De Dooms, (-BC), Jan. 1908, H. Bolus 13125

(BOL); Sandberg Hills on Skerpenheuvel Road, 300-600 m, (-DA),

28 Oct. 1982, Forrester & Bayer 237 (NBG); Jonaskop, roadside to

FM tower, (-DC), 14 Jan. 1979, Boucher 4247 (PRE); Jonaskop.

second gate from bottom, 600 m, (-DC), 9 Feb. 1981, Boucher 5006

(NBG); base of Sencor road, N side of Jonaskop, 30 Jan. 2004, Bergh

1105 (NBG); 7 km SW of Robertson, S side of Sandberg, (-DD), 31

Jan. 2007, Helme 4562 (NBG).

Lachnospermum fasciculatum

WESTERN CAPE. — 3117 (Lepelfontein): Brand se Baai, Harte-

beeste Kom, 74 m, (-BD), 26 Mar. 2003 [fr.], Desmet & Helme 3434

(NBG). 3118 (Vanrhynsdorp): 11.2 km S of Redelinghuys, (-DA), 7

June 1970, Acocks 24271 (PRE); between turn-off to Papkuilsfontein and

Brakvlei, (-DD), 20 Mar. 1996, Koekemoer 1217 (PRE). 3218 (Clanwil-

liam): rocky slopes west of Clanwilliam on road to Graafwater, (-BB),

6 Oct. 2004 [fr.], Goldblatt & Porter 12633 (NBG). 3219 (Wuppertal):

northern Cedarberg, Ribboksvlei Farm, N of Kliphuis Peak, ± 920 m,

(-AA), 14 Feb. 1984, Taylor 10913 (NBG, PRE); Pakhuisvlakte at top

of pass, ± 900 m, (-AA), 14 Jan. 1986, Taylor 11468 (NBG, PRE); 17

km from Clanwilliam on road to Pakhuis, (-AA), 25 Aug. 1995, Rod-

riguez-Oubina & Cruces 2076 (PRE); foot of Pakhuis Pass on E side,

(-AA), 21 Mar. 1996, Koekemoer 1220 (PRE); near Algeria Forest Sta-

tion, Mar. 1940, Stokoe 56987 (SAM); 1 km N of Algeria Forest Sta-

tion on Nieuwoudt’s Pass, ± 517 m, (-AC), 27 Apr. 1981, Le Maitre

196 (NBG. PRE); Algeria, (-AC), 14 Feb. 1936, Compton 6186 (NBG);

Op-de-Berg, Tuiskloof Farm, 756 m, (-CA), 8 Jan. 2003, Botha (3)012

(NBG); along R303 between Op-de-Berg and Citrusdal, (-CA), 11 Feb.

2007, Koekemoer 3483 (PRE); Citrusdal, Allendal Farm, Grysboknek,

306

Bothalia 41,2 (2011 )

FIGURE 1 1. — Lachnospermum negleclum: A-H, Boucher 5006. A, capitulum; B, involucral bracts (outermost on left and innermost on right); C,

palea; D, floret, side view; E, anther; F, stigma; G, cypsela (inner face, pappus removed); H, cypsela (outer face). 1, primary leaves (left Tyson

5877, right Helme 4562). L. fasciculalum, .l-L, Hanekom 3381 : J, capitulum; K, floret, side view; L, cypsela (side view, pappus removed).

Scale bar: A-C, J, 2 mm; D, G, H, 1, K, L, 1 mm; E, F. 0.5 mm. Artist: J.C. Manning.

800 m, (-CA), 23 Mar. 2002, Hanekom 3381 (NBG, PRE); 23 miles [37

km] N of Grootriver turn-off on road from Ceres to Citrusdal, (-CA),

8 Mar. 1966, Taylor 6755 (NBG); Kromrivier, on path to Disa Pools,

(-CB), 12 Mar. 2003, Koekemoer 2686 (PRE); La Fontein, (-CC), 10

Feb. 1978, Boucher 3655 (NBG, PRE). 3318 (Cape Town): sand flats

NE of Blaauberg, (-CD), Jan. 1927, Pillans 6822 (BOL); Malmesbury

Road, (-DA), 9 Mar. 1973, Montgomery 381 (NBG); Malmesbury Dist.,

Burgers Post Farm near Pella, 200 m, (-DA), 15 Mar. 1979, Boucher &

Shepherd 4269 (NBG, PRE), 17 Jan. 1980, Boucher & Shepherd 4929

(NBG, PRE); Kuils River, (-DC), II Mar. 1933, Levyns 4251 (BOL);

Klapmuts, (-DD), 9 Mar. 1959, Barker 8863 (NBG); ± 3 km E of Pniel,

Farm Normandy, 300 m, (-DD), 5 Apr. 2004, Helme 2986 (NBG); Bol-

Bothalia 4 1 ,2 (2011)

307

FIGURE 12. — Lachnospermum neg-

lectum, Schlechter9934 (holo-

type). Digital image courtesy

of Aluka website (www.aluka.

org).

telary, (-DD), 18 Jan. 1941, Compton 10380 (NBG). 3319 (Worcester):

Ceres, slopes of Elandskloof, (-AC), 5 Apr. 1952, Levyns 9834 (BOL);

25 km N of Wellington, Elandsberg Nature Reserve, (-AC), 12 Nov.

1986, De Villiers 40 (NBG, PRE); ‘in planitie ad pedem montis Mosterts-

berg’, (-AC), 30 Jan. 1 892, Schlechter 857 (BOL); Romans River Nature

16” IF 20“ 22“ 24” 26' 2»

FIGURE 13. — Known distribution of Lachnospermum neglectum, O;

and L. fasciculatnm, •.

Reserve, (-AC), 4 Feb. 1981, Nilsson 133 (PRE); Zachariaskloof Catch-

ment, ± 1100' [366 m], (-CC), 20 Feb. 1970, Haynes 293 (NBG, PRE);

Franschhoek Pass on Villiersdorp side, (-CC), 26 Feb. 1948, Compton

20472 (NBG). 3419 (Caledon): near bridge over River Zonder Einde

[Riviersonderend], between Viljoen’s Pass and Villiersdorp, (-AB), Mar.

1933, L. Bolus 21492 (BOL); near Genadendal, (-BA), 3 Mar. 1935, Lev-

yns 4864 (BOL); E of Greyton west of Gobos River, 220 m, (-BA), 23

Feb. 2006, Helme 3899 (NBG).

ACKNOWLEDGEMENTS

We thank the curators of the relevant herbaria for

allowing access to their collections, H.P. Linder for con-

firming the existence of isotype material of L. neglectum

in the Zurich Herbarium, and B. Nordenstam for oblig-

ing us in the same way for Stockholm.

REFERENCES

GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanical

Institute, Cape Town & Missouri Botanical Garden, St. Louis.

308

Bothalia 41,2 (201 1 )

HILLIARD, O.M. & BURTT, B.L. 1981. Some generic concepts in

Compositae-Gnaphaliinae. Botanical Journal of the Linnean

Society 82: 181-232.

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.

KOEKEMOER, M. 2002. Systematics of the Metalasia group in the

Relhaniinae (Asteraceae-Gnaphalieae). Unpublished Ph.D. the-

sis, Faculty of Science, Rand Afrikaans University, Johannes-

burg.

WARD, J„ BAYER, R.J., BRE1TWEISER, I., SM1SSEN, R„ GALBA-

NY-CASALS, M. & UNWIN, M. 2009. Gnaphalieae. In V.A.

Funk, A. Susanna, T.F. Stuessy & R.J. Bayer, Systematics, evolu-

tion, and biogeographv of Compositae: 539-588. International

Association for Plant Taxonomy, Vienna.

J.C. MANNING* and P. GOLDBLATT**

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

den, P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2010-06-03.

AMARYLLIDACEAE

AMMOCHAR1S DESERTICOLA (AMARYLLIDEAE), A NEW SPECIES FROM NAMIBIA AND A KEY TO SPECIES OF THE GENUS

Ammocharis Herb, is a small, sub-Saharan genus in

the family Amaryllidaceae. When last revised in 1939

the genus included five species (Milne-Redhead & Sch-

weickerdt 1939). More recently, phylogenetic analy-

ses using nuclear and limited plastid non-coding DNA

sequences (Meerow et al. 2003; Kwembeya et al. 2007)

have supported the taxonomic reassessments of two

anomalous species from genera closely related to Ammo-

charis (Lehmiller 1992; Snijman & Williamson 1994):

Cybistetes longifolia (L.) Milne-Redh. & Schweick. was

reinstated in Ammocharis and Crinum nerinoides Baker

was transferred to Ammocharis (Snijman & Archer

2003). In addition, Nordal (1982) reduced A. heterostyla

(Bullock) Milne-Redh. & Schweick. to synonymy in A.

angolensis (Baker) Milne-Redh. & Schweick.

The genus is most closely allied to Crinum L. from

which it is distinguished by leaves that, unlike those

of Crinum , are not obviously sheathing at the base and

have an intercalary meristem allowing the base of a leaf

to continue developing intermittently over several grow-

ing seasons. Due to this pattern of regrowth after a leaf

has died back, the mature leaves characteristically have

truncate tips. In addition, the leaves are biflabellately

arranged and most often have flat blades. Ammocharis ,

like all taxa in the tribe Amaryllideae, has fleshy, recal-

citrant seeds which fail to become dry and dormant

before germinating. In contrast to the other genera,

however, Ammocharis has two modes of seed dispersal.

Most commonly the seeds are dispersed by means of the

fleshy fruits being released individually from the fruiting

head in close proximity to the mother plant. A. longifo-

lia, which occupies the extreme western parts of winter

rainfall southern Africa, is exceptional in having a wind-

dispersed infructescence. The entire fruiting head rapidly

dries off and is shed as a single unit that is dispersed by

rolling away in the wind.

The genus comprises two widespread species, Ammo-

charis coranica (Ker Gawl.) Herb, which occurs in sea-

sonally damp areas throughout the summer rainfall area

of southern Africa (Milne-Redhead & Schweickerdt

1939), and A. tinneana (Kotschy & Peyr.) Milne-Redh.

& Schweick., which inhabits grassland or wooded grass-

land, extends from Ethiopia southwards to Botswana and

northern Namibia (Nordal 1982). Other species, which

have relatively restricted distribution ranges, occupy

more specific habitats within the landscape: flood plains

of rivers [A. baumii (Harms) Milne-Redh. & Schweick.]

and limestone pans (A. nerinoides). Namibia has all but

one species of Ammocharis within its borders, whereas

other countries have two species at most.

While on a botanical expedition in the extremely arid

Namib Naukluft Park in late February 2006, Namibian

botanist Dr Patricia Craven photographed flowering

plants of an unknown species that has leaves like Ammo-

charis coranica and A. longifolia but exceptionally long

flowers. Four years later, flowering and fruiting plants

were collected in the same region. The infructescences

were remarkable in being dry, with widely radiating,

straight pedicels, a condition previously known only

in A. longifolia. Following exceptionally good summer

rains in 2011, additional collections of this taxon were

made, allowing a more thorough study of the plants. The

strap-shaped leaves, with truncate tips characteristic of

an intercalary meristem, the unusually long flowers and

the dry infructescences indicate clearly that these newly

discovered Namibian plants are an undescribed spe-

cies of Ammocharis , closely related to A. longifolia. We

describe this new species here as A. deserticola and pro-

vide a key to identify the seven species currently recog-

nized in Ammocharis.

Ammocharis deserticola Snijman & Kolberg, sp.

nov.

Bulbus ± ovatus, tunicatus, 80-110 mm latus,

supeme conspicue attenuatus, collum ± 80 x 25-35

mm. Folia 6-8, disticha, biflabellatim disposita, paten-

tia vel adpressa, ligulata 7-250 x 8-30 mm, vetustiora

apice truncata. Scapus compressus, ad 210 * 12-18 mm.

Umbella 11 -25-flora. Pedicelli 30-60 x 3—4 mm, trans-

verse ± trigoni, sub fructu valde elongati, rigidi et radi-

antes. Flores actinomorphi, tubaeformes, rosei; perigonii

tubus cylindricus, rectus, 50-90 mm longus; segmenta

oblanceolata, 70- 80 x 8-12 mm, patentes, recurvi ver-

sus apicem; stamina e perigonii tubo valde exserta; filam-

enta actinomorpha, 53-60 mm longa; antherae oblongae,

curvatae, 5><1.5 mm, pallidae; ovarium obtuse 3-angu-

latum, 10-15 mm longum; stylus filiformis, exsertus;

stigma integrum. Capsula indehiscens, obovoidea, 20-40

x 15-45 mm, valde 6-costata, rostrata; rostrum 14-50

mm longum. Semina camosa, subglobosa, ± 15 mm lata,

pallida.

TYPE. — Namibia, 2215 (Trekkopje): Erongo region,

Namib Naukluft Park, 5 km N of Langer Heinrich mine

Bothalia 41,2 (2011)

309

on track towards Swakop River, 565 m, (-CD), 17 Feb.

2011, H. Kolberg & T. Tholkes HK2998 (WIND, holo.,

NBG, iso.).

Deciduous bulbous herb, 200-350 mm tall when

flowering. Bulb solitary, hypogeal, ± ovate, 80-110 mm

diam., extended into a neck, ± 80 x 25-35 mm, outer

tunics brown to tan-coloured, with darker brown longi-

tudinal veins, leathery to somewhat brittle, inner tunics

cream-coloured, with tan longitudinal veins. Leaves 6-8,

green at flowering, in 2 opposite spreading fans, laxly

spreading, later becoming prostrate, some curved side-

ways at base, strap-shaped, 7-250 x 8-30 mm, plane,

glaucous-green, glabrous; margin pale, smooth; apex of

mature leaves dying back as if cut across. Inflorescence

1 or rarely 2 mature at once, laxly 1 1-25-flowered,

umbel-like, ± 250 mm diam.; scape erect, longitudinally

compressed, up to 210 x 12-18 mm, green sometimes

flushed pink near base or apex; spathe valves 2, oblong-

lanceolate, up to 50-80 x ± 20 mm, soon reflexed and

dry; bracteoles filiform, ± 35 mm long. Flowers erect in

bud, soon spreading; pedicels erect at first, soon spread-

ing, 30-60 x 3^1 mm, trigonous in cross section, green;

perigone trumpet-shaped, actinomorphic, dark pink,

with paler pink tube, backed with narrow, brownish pink

median stripes on segments, faintly scented; tube cylin-

drical, straight, 50-90 mm long, widening slightly up to

5 mm at throat, firm; segments oblanceolate, 70- 80 x

8-12 mm, recurved distally, mucronate on tip. Stamens

inserted in throat, ± evenly spreading; filaments 53-60

mm long, 3 inner ± 2/3 as long as tepals, 3 outer ± as

long as tepals, ± inwardly curved, pink; anthers dorsi-

fixed, oblong, ± curved, 5.0 x ].5 mm, cream-coloured;

pollen cream-coloured. Ovary ± trigonous in cross sec-

tion, 10-15 mm long, scarcely wider than pedicels and

perigone tube, green, style filiform, reaching 10-15 mm

beyond outer stamens, pink, stigma small, undivided,

minutely penicillate. Infructescence dry; pedicels widely

radiating, straight, thickened, 50-130 x 5-8 mm; cap-

sule indehiscent, obovoid, 20^40 x 15-45 mm, strongly

6-ribbed, with thin, papery walls disintegrating irregu-

larly, with stout, apical beak, 14-50 mm long. Seeds few,

subglobose, ± 1 5 mm diam., fleshy, with a thin, pale,

corky covering. Flowering time : mid-February to early

March. Figures 14, 15.

Distribution and ecology. Ammocharis deserticola is

currently recorded only from the Namib Naukluft Park,

in the central Namib, Namibia (Figure 16). The species

occurs along well-drained, sandy rivulets and in grav-

elly soil on the northern slopes of undulating hills, often

benefiting from fog that drifts inland from the coast. The

surrounding vegetation is sparse and comprises spe-

cies of Stipagrostis (Poaceae), Adenolobus pechuelii

(Fabaceae) and Tetraena stapjfli (Zygophyllaceae).

Rainfall occurs in the summer months with an annual

FIGURE 14. — Habitat and habit of Ammocharis deserticola. A, typical, open habitat in the Namib Naukluft Park, Namibia, between low hills in

gravelly soil; B, plant in bud showing leaves with truncate leaf tips; C, flowering plant showing distinctive, long perigone tube; D, fruiting

plant showing dry infructescence. Photographs by H. Kolberg.

310

Bothalia 41,2 (201 1 )

FIGURE 15. — Ammocharis deserticola. A, dried bulb cut in half lengthwise with pressed leaves and pressed inflorescence; B, pressed infructes-

cence cut in half lengthwise. Voucher specimen: H. Kolberg & T. Tholkes HK2998 (NBG). Scale bars: A, B = 100 mm.

average of 50-100 mm, but it is highly erratic and com-

pletely rainless years are not uncommon. Minimum tem-

peratures on average are 9-1 0°C, whereas they reach an

average of 32-33°C in the summer months (Mendelsohn

et al. 2002). At the type locality, between 300 and 400

plants were visible in February 2011, whereas at two

other localities in the same quarter-degree grid, an esti-

mated 100 to 150 plants were seen. Two populations,

including the one at the type locality, occur near the

Langer Heinrich uranium mine, but are afforded protec-

tion by falling within a national park. The third popula-

tion of up to 100 plants, however, falls within the mine’s

operational area and the plants there will be endangered.

Plans are under way to rescue these and use them in the

restoration of mine sites. There are unconfirmed reports

of this species occurring also in the quarter-degree grids

23 1 5CA and 25 1 5DA.

Diagnostic features and relationships : Ammocharis

deserticola is easily identified by a combination of floral

and fruiting features. The flowers have a characteristi-

cally long perigone tube (50-90 mm) with broad, lan-

ceolate, perigone segments. The infructescence is dry at

maturity and has stout pedicels that radiate ‘spoke-wise’

from their attachment with the scape. Each pedicel car-

ries a dry, heavily 6-ribbed capsule which is topped with

a stout, 14-50 mm long, apical beak. The thin, papery

walls of the capsules break open irregularly, often allow-

ing the extremely large seeds to drop to the ground

before the infructescence is rolled away in the wind.

The relatively broad perigone segments and specialized

infructescence which are shared with A. longifolia sug-

gest a close alliance with that species. In A. deserticola ,

however, the perigone tube is more than twice as long as

that of A. longifolia (50-90 vs 8-15 mm long).

FIGURE 16. — Known distribution of Ammocharis deserticola , ▲; and

A. longifolia , •.

Bothalia 41,2 (2011)

311

More than 600 km separate the populations of Ammo-

charis deserticola from the closest known populations of

A. longifolia in the Sperrgebiet, southern Namibia. Else-

where, A. longifolia is found in South Africa, extending

southwards from the sand plains of the Richtersveld.

through the Namaqualand lowlands to the sand flats of

the Cape Peninsula and Breede River Valley, Western

Cape (Figure 16). It is remarkable that such a showy

species as A. deserticola has been overlooked in the

past. This is probably the result of the bulbs not flow-

ering for long intervals until sufficient rain falls and

because their area of occupancy was largely inaccessible

to botanists prior to the mining activities in the central

Namib.

Other specimens examined

NAMIBIA. — 2215 (Trekkopje): Erongo region. Namib Naukluft

Park, track west of Langer Heinrich mine towards Swakop River, 5 km

N of main mine road, 560 m, (-CD), 17 Mar. 2010, H. Kolberg & T.

Tholkes HK2915 (NBG, PRE, WIND); Erongo region, Namib Nauk-

luft Park, just W of entrance to Langer Heinrich mine, 625 m, (-CD),

17 Feb. 2011 . H. Kolberg & T. Tholkes HK2997 (NBG).

Key to species

la Leaves linear, straight, ± erect A. baumii

lb Leaves narrowly to broadly lorate, at least some falcate,

spreading on ground:

2a Flowers with style included in perigone tube A. angolensis

2b Flowers with style well-exserted from perigone throat:

3a Perigone segments with apex spirally recurved at anthe-

sis A. tinneana

3b Perigone segments recurved at anthesis but never spirally:

4a Pedicels not elongating after anthesis; infructescence

drooping on ground; fruit walls smooth, membranous:

5a Bulb 20-51 mm diam. at maturity; leaves 2 — 4(— 1 0)

mm wide; scape 2. 5-5.0 mm wide; flower head

60-130 mm diam A. nerinoides

5b Bulb 90-160 mm diam. at maturity; leaves (5— )25— 75

mm wide; scape 15-25 mm wide; flower head 1 50—

200 mm diam A. coranica

4b Pedicels lengthening after anthesis; infructescence

detaching at ground level; fruit walls papery, heavily

6-ribbed:

6a Perigone tube 8-15 mm long A. longifolia

6b Perigone tube 50-90 mm long A. deserticola

ACKNOWLEDGMENTS

We are grateful to Tyrone Tholkes who assisted in the

field, Patricia Craven for her images and Michelle Smith

for preparing the figures. The second author acknowl-

edges funding from the Millennium Seed Bank Partner-

ship, Kew and permission from the Namibian Ministry

of Environment and Tourism to collect plants.

REFERENCES

KWEMBEYA, E.G., BJORA, C.S., STEDJE, B. & NORDAL, I. 2007.

Phylogenetic relationships in the genus Crinum (Amaryllidace-

ae) with emphasis on tropical African species: evidence from

trnL-F and nuclear ITS DNA sequence data. Taxon 56: 801-810.

LEHMILLER, D.J. 1992. Transfer of Crinum nerinoides to Ammo-

charis (Amaryllidaceae). Novon 2: 33-35.

MEEROW, A.W., LEHMILLER, D.J. & CLAYTON, J.R. 2003. Phy-

togeny and biogeography of Crinum L. (Amaryllidaceae) inferred

from nuclear and limited plastid non-coding DNA sequences.

Botanical Journal of the Linnean Society 141 : 349-363.

MENDELSOHN. J„ JARVIS, A., ROBERTS, C. & ROBERTSON, T.

2002. Atlas of Namibia: a portrait of the land and its people.

Ministry of Environment and David Phillip Publishers, Cape

Town.

MILNE-REDHEAD. E. & SCH WEICKERDT, H.G. 1 939. A new con-

ception of the genus Ammocharis Herb. Journal of the Linnean

Society, Botany 52: 159-197.

NORDAL, I. 1982. Amaryllidaceae. In R.M. Polhill, Flora of tropical

East Africa. Balkema, Rotterdam.

SNIJMAN, D.A. & ARCHER, R.H. 2003. Amaryllidaceae. In G. Ger-

mishuizen & N.L. Meyer, Plants of southern Africa: an annotated

checklist. Strelitzia 14: 957-967 .

SNIJMAN, D.A. & WILLIAMSON, G. 1994. A taxonomic re-assess-

ment of Ammocharis herrei and Cybistetes longifolia (Amaryl-

lideae: Amaryllidaceae). Bothalia 24: 127-132.

D.A. SNIJMAN* and H. KOLBERG**

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Cape Town. Email: d.snijman@sanbi.org.za.

** Millennium Seed Bank Partnership, Namibia, c/o National Botani-

cal Research Institute, Private Bag 13184, Windhoek, Namibia. Email:

hertak@nbri.org.na.

MS. received: 2011-04-04.

ALLIACEAE

MICROMORPHOLOGY AND CYTOLOGY OF PROTOTULBAGHIA SIE BERTH. WITH NOTES ON ITS TAXONOMIC SIGNIFICANCE

Four genera of the Alliaceae are known to occur in

sub-Saharan Africa, in total comprising 35 indigenous

and two naturalized alien species (Table 2) (Archer 2003;

Klopper et al. 2006). Tnlbaghia L. is the most diverse

and widespread genus of the family in southern Africa,

but the other two indigenous sub-Saharan genera. Allium

L. and Prototulbaghia Vosa, are each represented by a

single restricted-range species, of which the latter has

only been described recently (Vosa 2007). This note sup-

plements the existing description of P. siebertii Vosa,

which is endemic to the Sekhukhuneland Centre of Ende-

mism (Van Wyk & Smith 2001 ), South Africa (Vosa et al.

2011), where it is restricted to the summit of the Leolo

Mountain Range, Limpopo Province. Although macro-

morphological characters of Prototulbaghia have been

documented by Siebert et al. (2008), little is known about

the micromorphology and cytology of P. siebertii.

Live material of Prototulbaghia siebertii was col-

lected from the wild in November 2005 and culti-

vated in clay pots in a mixture of compost, peat, sand

and garden soil. Plants were housed in a greenhouse at

TABLE 2. — Sub-Saharan genera of Alliaceae and no. species per genus

recorded for Flora of southern Afi-ica ( FSA ) (Archer 2003) and

tropical African flowering plants (EPFAT*) respectively (Klop-

per et al. 2006)

talien taxa; * Enumeration des plantes a fleurs d’Afrique tropicale.

312

Bothalia 41,2 (2011)

A

FIGURE 1 7. — Prototulbaghia siebertii. A, micrograph of floral scape showing very small glandular hairs; B, photograph of peculiar hooked tips of

leaves. Plant grown from seed originally collected near Schoonoord, Leolo Mountains, Sekhukhuneland. Scale bars: A. 1 mm; B. 0.75 mm.

18 to 22°C. For mitotic preparations, actively growing

root tips were collected at about 1 1 : 00, pretreated in an

aqueous solution of 0.05 % colchicine for four hours at

greenhouse temperature and fixed overnight in 1 :3 gla-

cial acetic acid/absolute alcohol (Vosa 2000). After a

short hydrolysis in IN hydrochloric acid at 60°C, the

meristematic part of the tip was squashed in a drop of

1.5 % orcein in 45 % acetic acid and stained in Feulgen

for one hour (Vosa 2000). For an assessment of meio-

sis in pollen mother cells, anthers were removed from

young flower buds at an appropriate developmental

stage, before colouring of the perianth, and squashed in

1.5 % aceto-orcein (Vosa 1961) without pretreatment or

fixation on a microscope slide. The slide was left for six

hours and pressed between filter paper. The slide was

heated slightly over a flame, covered with a cover-slip

and sealed with paraffin wax (Vosa 1961). Micrographs

of the living material and of the cytological preparations

were taken with a digital camera.

Observations of the finer morphological features of

Prototulbaghia , both in living plants and herbarium

specimens have revealed four further diagnostic char-

acters of the inflorescence, leaves and anatomy. Most

conspicuous is the scape (or peduncle) which is red-

dish brown in the lower portion during its early stages

of development and is covered by short, whitish, glan-

dular hairs with somewhat recurved tips (Figure 17A).

The glandular hairs on the scape are easily rubbed off,

revealing underneath a single ridge of small translucent

teeth with their tips directed proximally (Figure 18A).

Under higher magnification (x 10), the surface of the

leaves is somewhat pruinose, especially distally, and

near the apex is a single row of tiny translucent, almost

glass-like teeth directed toward the apex. The row of

teeth forms a type of low central keel between two indis-

tinct keels on the abaxial surface as described by Vosa

(2007) (Figure 18B). The teeth on the distal part of the

leaves are easily rubbed off.

The minute teeth on the leaves and scape of Proto-

tulbaghia siebertii have not been recorded in any of the

species of Tulbagliia so far studied. In the allied genus

Allium, at least in section Molium, glandular hairs are

present both on the floral scape and on the leaves (Mann

1959). Flowever, many of the over 250 species of Allium

have not been examined for this character.

The spreading, shiny, dark green leaves of Prototul-

baghia siebertii have peculiar hooked apices (Figure

17B), which turn black in dried material. The apices

of emerging young leaves are slightly coiled outwards.

FIGURE 18. -Prototulbaghia siebertii. A, micrograph of floral scape showing ridge of tiny translucent teeth; B, micrograph of distal part of leaf

showing liny teeth directed towards apex of leaf. Scale bars: A, B. I mm.

Bothalia 41,2 (201 1 )

313

the coils relaxing as the leaf develops, but remaining

thickened and hook-like in mature leaves. Such hook-

like leaf apices have not previously been recorded for

the Alliaceae. Plants are easily uprooted when pass-

ing a hand or fingers through a clump, because of the

entangled hooked apices. We speculate that the hooked

leaves function as an additional mechanism of short

range vegetative dispersal (Vosa 2007). So far the only

other recorded instance of hooked leaf apices assisting

in dispersal of a plant is known for Junaea spectabilis

Kunth. (Arecaceae), the Chilean wine palm (C.G. Vosa

pers. obs. in the wild and in cultivation). In the case of

the palm, this unusual phenomenon has been observed in

the leaflets (pinnae) of the fronds in seedlings and fairly

young plants. This palm grows preferentially in soft for-

est soil, rich in peaty humus, and the hooked or coiled

leaflet apices aid in seedling dispersal when dragged

around by passing animals. This may also be the case in

Prototulbaghia , considering the muddy, wet, humus-rich

soils of its natural habitat.

The fourth micromorphological (anatomical) observa-

tion is a family characteristic. We can confirm the presence

of laticifers which on damage exude a clear watery liquid

in the leaf tissue of Prototulbaghia. Similar laticifers are

also present in Tulbaghia and Allium (Mahlberg 1993).

Prototulbaghia has a diploid chromosome comple-

ment of 2 n = 12. The karyotype is very similar to that

of Group 5 in Tulbaghia (Vosa 1975, 2000, 2009), sup-

porting the suggested close relationship between the two

genera. This poses the question how such an isolated

species, which is not polyploid, could have persisted

with such a presumably limited genetic endowment con-

sidering that polyploids are generally believed to display

increased genetic variability and evolutionary potential

(Hunziker& Schaal 1983).

A preliminary assessment of meiosis in Prototul-

baghia, based on 18 pollen mother cells at metaphase

1, has revealed six regular bivalents with an average

of about 20 chiasmata. Apart from the long arms of

the longest chromosome of the complement which are

unpaired (Figure 19), crossing over appears to be non-

localized. This is very unlike the pattem/situation in

those species of Tulbaghia so far investigated where

crossing over is almost exclusively proximal to the cen-

tromere (Vosa 1975, 2000). This pattern may indicate a

relatively higher degree of recombination than in Tul-

baghia where the average chiasmata, based on the anal-

yses of six diploid species, is much lower (Vosa 1966).

Gender is one of the most important genetic factors

affecting recombination frequency (Burt et al. 1991).

Hence the existence of a two-track heredity in Tulbaghia

(Vosa 1972 ) with a higher degree of genetic recombi-

nation in ovules than in microspores, may possibly be

somewhat enhanced in Prototulbaghia as a compensat-

ing recombination mechanism. Overall such hypotheti-

cal modified genetic inheritance in Prototulbaghia may

mean that the macromorphological uniformity observed

in nature in its populations hides a complex and spe-

cialized genomic arrangement. However, these ideas on

recombination frequency in Prototulbaghia still require

testing.

FIGURE 19. — Prototulbaghia siebertii. Micrograph of metaphase 1 of

meiosis (six bivalents have been numbered in order of decreas-

ing size). Note: in bivalent 1 , unpaired segments corresponding

to long arms of two involved chromosomes (small arrows). In

this metaphase, there are 24 chiasmata, 23 non-localized and one

proximal to centromere indicated by larger arrow. Scale bar: 10

pm.

The lack of recombination in the long arms of the

longest chromosome of Prototulbaghia (Figure 19) may

be a parallel evolutionary trend toward the state found

in all the species of the closely related Tulbaghia. The

existing cytological differences between the two genera

may be due to the hypothetical need of high recombi-

nation in Prototulbaghia for the genome as a whole to

maintain genetic variability as a narrow endemic with a

single population. The paradox is that although it may

seem tempting to regard Prototulbaghia as an entity

with some plesiomorphic features and most probably

sister to Tulbaghia and intermediate to Allium, the exist-

ence of the taxon-specific micromorphological and cyto-

logical features noted above may also be an indication

of its specialized and geographically isolated evolution

(‘trapped’ on its specialized mountain summit habit).

Observations of the cytology discussed above provide

evidence of a diagnostic character that provides support

for the hypothesis that Prototulbaghia may possess fea-

tures that are ancestral to Tulbaghia. Note, however, that

in Tulbaghia the grouping of species based on karyotype

(Vosa 1975, 2000) corresponds only in part to the sub-

divisions of the genus into sections based on floral mor-

phology (Vosa 2009). Such discrepancies demonstrate

that in Tulbaghia floral morphology and the karyotype

in many cases seem to follow independent evolutionary

pathways (Vosa 2009). Despite the similarity in karyo-

type, differences in floral morphology between Proto-

tulbaghia and Tulbaghia have also been documented by

Vosa (2007) and Siebert et al. (2008).

Available molecular phylogenetic analyses of

Alliaceae support a close relationship between Allium

and Tulbaghia, although hitherto only two (77 simmleri

Beauv. and T. violacea Harv.) of the 31 species of Tul-

baghia were considered (Fay & Chase 1996; Friesen et

al. 2006; Nguyen et al. 2008). In such restricted molecu-

lar analyses, the potentially misleading effects of poor

taxon sampling on phylogenetic analyses and their appli-

314

Bothalia 41,2 (201 1 )

cations should be kept in mind (e.g. Heath et al. 2008).

There is an obvious need for a well-sampled molecular

phylogenetic analysis that includes Prototulbaghia and

more Tulbaghia species, together with other members of

the Alliaceae, to test proposed generic demarcations and

evolutionary trends in the family.

For the description of Prototulbaghia siebertii , see

Vosa et al. 2011. Newly observed features are:

Leaves have a row of tiny translucent teeth adpressed

toward the distal abaxial part below the apex; laticifers

present. Spathe valves subtend the inflorescence when the

flower scape emerges, gradually drying off. Flower scape

is covered by short, whitish, glandular hairs; keel present

towards the base, single, persistent, ridge-like, with trans-

lucent teeth. Flowers are fugacious. Perianth segments

are light green in the lower half; inner segments are 4.3

mm wide in the middle. Gynoecium has 2—4 ovules per

locule. Capsule has a withered style on top.

ACKNOWLEDGEMENTS

The North-West University and University of Pretoria

provided financial support.

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SIEBERT, S.J., VOSA, C.G., VAN WYK, A.E. & MULLER, H. 2008.

Prototulbaghia (Alliaceae), a new monotypic genus from

Sekhukhuneland, South Africa. Herbertia 62: 76-84.

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southern Africa. A review with emphasis on succulents. Umdaus

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female meiosis in Tulbaghia. Catyologia 25: 275-281.

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Botanica (Roma) 34: 47-121 .

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2011. Prototulbaghia siebertii. Flowering Plants of Africa 62:

22-28, t. 2264.

C.G. VOSA*, S.J. SIEBERT** and A.E. VAN WYK|

* Author for correspondence: Linacre College, Oxford, United King-

dom & Department of Biology, University of Pisa, 56126 Pisa, Italy.

E-mail: caniovosa@tin.it.

** A.P. Goossens Herbarium, School of Environmental Sciences and

Development, North-West University, Private Bag X6001, 2520 Potch-

efstroom. South Africa. E-mail: stefan. siebert@nwu.ac.za.

t H.G.W.J. Schweickerdt Herbarium, Department of Plant Science,

University of Pretoria, 0002 Pretoria. E-mail: braam.vanwyk@up.ac.

za.

MS. received: 2010-05-28.

HYACINTHACEAE

ALBUCA GARIEPENSIS (ORNITHOGALOIDEAE), ANEW SPECIES OF A. SUBGEN. NAMIBIOGALUM FROM GORDONIA, SOUTH

AFRICA, AND A. PRASINA TRANSFERRED TO ORNITHOGALUM

INTRODUCTION

Generic limits in Hyacinthaceae: Ornitliogaloideae

have been intensively reviewed in recent years using

molecular techniques (Stedje 1998; Pfosser & Speta

1999; Manning et al. 2004, 2009; Martinez-Azorin et

at. 2011). The broad patterns of phylogenetic relation-

ships in the subfamily are now clear but various options

exist for the translation of these patterns into a generic

taxonomy (Manning el ah 2009; Martinez-Azorin et al.

201 1). A recent solution, followed here, provides for the

expansion of Albuca L. to include Ornithogalum sub-

gen. Osmytae (Salisb.) Baker and O. subgen. Urophyllon

(Salisb.) Baker (Manning et al. 2009). Ornithogalum is

consolidated to include the genera Galtonia Decne. and

Neopatersonia Schonland, leaving Dipcadi Medik. and

Pseudogaltonia (Kuntze) Engl, as the remaining gen-

era in the subfamily. In this circumscription, Albuca

is distinguished from Ornithogalum by flowers with

thick-textured, whitish or yellowish tepals with a broad,

green or brown longitudinal band on the adaxial surface

associated with 3-5 mostly simple, medially aggre-

gated veins (Manning et ah 2009). In Ornithogalum , the

tepals are thin- or thick-textured with a narrow or indis-

tinct median band and veins not tightly aggregated in

the middle of the tepals, and the outer veins are often ±

branched.

Several collections of narrow-leaved plants from the

middle reaches of the Orange River in Northern Cape,

Bothalia 41,2 (2011)

315

with relatively short, pyramidal racemes of green-

banded flowers and relatively large, depressed-globose

or cordate capsules, have remained unnamed until now

or have been identified as Ornithogalum tenuifolium

subsp. aridum Oberrn. [now A/buca virens subsp. arida

(Oberm.) J.C. Manning & Goldblatt], Their large cap-

sules, however, preclude any association with A. virens ,

which has smaller, ovoid capsules, and it is now clear

that they represent an undescribed species, which we

describe here as A. gariepensis after the Khoisan name

for the Orange River, known to earlier inhabitants of the

region as the gariep or Great River.

Collections at KMG, NBG, PRE and SAM. the main

herbaria with good representation of collections of

Northern Cape species, were consulted for records of the

new species (herbarium acronyms after Holmgren et al.

1990).

In addition, examination of herbarium specimens

of Ornithogalum prasinum Ker Gawl., which was

transferred to Albuca by Manning et al. (2009) on the

strength of its supposedly close relationship to O. sein-

eri (Engl. & K. Krause) Oberm. (now A. seineri (Engl. &

K. Krause) J.C. Manning & Goldblatt, reveals that it lacks

the perianth features that are diagnostic of Albuca. This,

combined with its karyology with basic chromosome

number x = 8, dictates its transfer back to the genus

Ornithogalum , where its large, retuse capsules suggest a

relationship with O. xanthochlorum Baker.

TAXONOMY

Albuca gariepensis J.C. Manning & Goldblatt sp.

nov.

Geophyta decidua 0.03-0.20 m alta, foliis (4)5— 8(9)

racemum duplo ad sexies longior linearibus pauciter

rotatis vel ad apicem tortis canaliculatis 70-200 x 1-4

mm, racemo subpyramidalo dense 8-25-florum, 30-80

mm longo. bracteis lanceolato-aristatis, inferioribus ad

20 x 3 mm submembranaceis ad papyraceis, pedicelis

± patentibus (5— )8— 1 0(— 1 5) mm longis, floribus rotatis

tepalis liberibus albis carinis viridibus oblongo-ellip-

ticis 7-10 x 2. 5-3. 5 mm, nervis centralibus tribus con-

gestis, staminibus liberibus erectis, filamentis 4-5 mm

longis inferioribus 1.5 mm expansis quadratiformis,

ovario ovoideo breve stipitato ± 3 mm longo profunde

3-angulato, stylo erecto ± 3 mm longo, capsula ambita

depresso-cordata profunde 3-lobata 10-12 x 15-20 mm

diam., seminibus discoideis 5-8 mm diam.

TYPE. — Northern Cape: 2921 (Kenhardt):10-12 km

NE of Putsonderwater along roadside. Farm Kareelaagte,

(-BB), 6 Mar. 2010, Manning 3294 (NBG, holo.; MO,

PRE. iso.).

Deciduous geophyte, 0.03-0.20 m high. Bulb solitary,

pyriform, 10-25 mm diam.; outer tunics dry and thinly

leathery, brownish, sometimes forming a loose neck

and finely transversely barred, inner tunics tightly over-

lapping, white. Leaves overtopping raceme and two to

six times as long, (4)5— 8(9), suberect or spreading, not

clasping below, linear, slightly coiled or twisted apically,

canaliculate and rounded abaxially, 70-200 x \-4 mm,

acute, fleshy, dull bluish green. Inflorescence usually

solitary but up to 3 from a bulb, subpyramidal, densely

8-25-flowered, 30-80 mm long at flowering, elongating

slightly and ± cylindrical in fruit, scape green and 1 .5-

3.0 mm diam. at ground level but subterranean portion

tapering strongly and white; bracts lanceolate-aristate,

lowermost up to 20 x 3 mm and uppermost ±7x2 mm,

apex often slightly coiled, whitish and submembranous

when young but becoming papery; pedicels ± spread-

ing, (5— )8— 1 0(— 1 5) mm long. Flowers held horizontally,

rotate, 14-20 mm diam., faintly scented of lemon-coco-

nut; tepals biseriate with outer series overlapping inner,

free, suberect in basal 1. 5-2.0 mm then spreading, white

with green keels adaxially, outer tepals oblong-elliptical,

7-10 x 2. 5-3.0 mm, slightly keeled apically and peni-

cillate, with 3 centrally congested veins, inner tepals

elliptical, 7-10 x 3. 0-3. 5 mm, concave apically and

penicillate, with 3 centrally congested veins. Stamens

free, erect; filaments 4-5 mm long, two thirds as long as

tepals, verruculose, both series with quadrate expansion

in basal 1.5 mm, outer expansion ± 1 mm wide, inner

1.5 mm wide; anthers versatile, ± 2 mm long at anthe-

sis, pale creamy yellow. Ovary ovoid and shortly stipi-

tate, ± 3 mm long, strongly 3-angled, green; style erect,

tapering, 3-angled, ± 3 mm long, acute with trigonous,

papillate stigma, green in lower half and white distally.

Capsule depressed-cordate in outline, deeply 3-lobed,

10-12 x 15-20 mm. Seeds discoid, 5-8 mm diam.. col-

liculate, glossy black. Flowering time ; January to early

April. Figures 20; 21.

Distribution and ecology. Albuca gariepensis seems

to be restricted to the central portion of the Orange River

drainage basin in northeastern Bushmanland, where it

has been collected in a region bounded by the towns of

Kakamas, Upington and Kenhardt (Figure 22). The spe-

cies has been recorded from calcrete plains and sandy

flats, often in slight washes or drainage lines where

seasonal soil moisture levels are higher. A. gariepensis

appears to be endemic to the Bushmanland Bioregion

of the Nama-Karoo Biome, primarily in Bushmanland

Arid Grassland (Mucina & Rutherford 2006). Flower-

ing occurs in late summer, the exact timing evidently

depending on the seasonal rains. The collection Snij-

man 244 includes a plant bearing both mature fruiting

and flowering stems, indicating two growth flushes some

time apart, presumably in reponse to two distinct rainfall

events.

Diagnosis and relationships : the species of Albuca

with ± monomorphic tepal whorls are segregated among

four subgenera that are defined by a combination of

vegetative, floral, and fruiting characters (Manning

et al. 2009). The several, canaliculate leaves and the

pyramidal raceme of white flowers with shortly stipi-

tate, 3-lobed ovary developing into a depressed, apically

retuse capsule containing large, discoid seeds place A.

gariepensis in subgen. Namibiogalum (U.Mtill.-Doblies

& D.Miill.-Doblies) J.C. Manning & Goldlbatt (Manning

et al. 2009). Here, it is characterized by its several, lin-

ear-canaliculate leaves that are loosely coiled or twisted

apically and much longer (three to six times) than the

short inflorescence; its moderate-sized flowers, 14-20

mm in diameter, borne on pedicels mostly 8-10 mm

long; and relatively large, deeply 3-lobed capsules. The

316

Bothalia 41,2 (2011)

FIGURE 20. — Albuca gariepensis, Manning 3294. A, flowering plants; B, bract; C, flower; D, outer tepal; E, inner tepal; F, outer stamen, adaxial

(upper) and abaxial (lower) views; G, inner stamen; H. ovary, lateral and dorsal views; I, capsule; J, seed. Scale bar: A-C, I, J, 10 mm; B,

C, 3 mm; D-H, 2 mm. Artist: John Manning.

Bothalia 41,2 (201 1 )

317

FIGURE 21. — Albuca gariepensis.

A, flowering plant, Man-

ning 3294. Photograph by

John Manning. B, fruiting

plant, Magee & Boatwright

229. Photograph by Anthony

Magee.

species is most similar to A. seineri (Engl. & K. Krause)

J.C. Manning & Goldblatt in foliage and inflorescence

but the latter is an altogether larger species, with ±

straight or arching leaves up to 400 * 30 mm (vs 70-200

x 1-4 mm), and larger flowers 24-40 mm in diameter

borne on longer pedicels, up to 50 mm long. Its area of

occurrence is north of that of A. gariepensis, ranging

from northern Namibia through Botswana and the north-

ern parts of Northern Cape (Prieska and Hopetown Dis-

tricts) into the western Free State, northern Mpumalanga

and Limpopo (Obermeyer 1978).

Smaller plants of Albuca gariepensis may be con-

fused with A. toxicaria (C. Archer & R.H. Archer)

J.C. Manning & Goldblatt from southern Namibia and

central South Africa, which has with similar, almost fili-

form leaves but which has smaller flowers, ± 10 mm in

diameter, very short pedicels 1.5-2. 2 mm long, minute,

deltoid bracts ± 1 .2 mm long, and much smaller, ovoid

capsules up to 10 x 5 mm containing elliptical, peripher-

ally winged seeds 8 * 5-6 mm (Archer & Archer 1999).

The filaments in Albuca gariepensis are abruptly

expanded at the base into quadrate wings, unlike the

smoothly tapering filaments in A. seineri and A. toxi-

caria. Similar, basally expanded filaments are charac-

teristic of some other species in the subgenus, notably

A. rautanenii (Schinz) J.C. Manning & Goldblatt and

A. stapfii (Schinz) J.C. Manning & Goldblatt. but the

ovary in both these species is abruptly expanded above

the stipe into a 6-lobed disc. Although the ovary in A.

gariepensis is slightly widened above the stipe, it does

not develop the characteristic lobes that are present in

these two species.

Additional specimens seen

NORTHERN CAPE. — 2820 (Kakamas): 31 km W of Kakamas

along road to Pofadder and Onseepkans, 800 m, (-CC), 31 Jan. 1974,

Davidse 6169 (PRE); 20 km S of Kakamas on road to Loeriesfontein,

789 m, (-DC), 14 Mar. 2010, Magee & Boatwright 229 (MO, NBG).

2821 (Upington): Upington, industrial commonage, 2700 ft [1 680 m],

1 Apr. 1980, Snijman 244 (NBG). 2921(Kenhardt): near Steenkop, SE

of Kenhardt, 900 m, (-AD), 7 Jan. 1989, Bntyns 3461 (NBG).

Omithogalum prasinum

Ornithogalum prasinum Ker Gawl. is relatively w ide-

spread through the drier parts of central South Africa,

extending into Namibia and Botswana (Obermeyer

1978). The species was misattributed to John Lindley

by Baker (1897) and subsequent authors, but John Bel-

lenden Ker[-Gawler] provided the text of the first 14

volumes of The botanical register , whereas Lindley was

responsible for volumes 15-33 (Stafleau & Cowan 1976:

724). Placed by Baker (1897) in O. subgen. Osmyne

(Salisb.) Baker, O. prasinum was subsequently trans-

ferred to subgen. Urophyllon (Salisb.) Baker by Ober-

meyer (1978), who considered it to be closely related

to O. seineri. This treatment was followed by Miiller-

Doblies & Muller-Doblies (1996), who included both

species in O. series Prasinocorymbosa Mtill.-Doblies &

Miill.-Doblies, and later by us when we transferred the

members of O. subgen. Urophyllon to Albuca (Manning

et al. 2009). At this time we had DNA sequence data for

O. seineri but not for O. prasinum and our association of

the two species was determined by the opinions of Ober-

meyer (1978) and Muller-Doblies & Muller-Doblies

(1996).

A review of the cytology of subfamily Ornithogaloi-

deae (Goldblatt & Manning 2011) shows A. prasina to

have a diploid number of 2 n = 16 (Pienaar 1963), thus

x = 8. This makes the species cytologically anomalous

in Albuca , in which the most common base number is x

= 9, and led us to re-examine its floral morphology, par-

ticularly the tepal venation, in relation to that of A. sein-

eri. We confirm that A. seineri (Figure 23 A, B), with 2 n

= 24 (x = 6) (Vosa 1980), has the medially aggregated,

mostly simple veins diagnostic of Albuca but that A.

FIGURE 22. — Known distribution of Albuca gariepensis.

318

Bothalia 41,2 (201 1 )

FIGURE 23. — Albuca seineri , D’Ewes s.n. (NBG). A, outer tepal; B,

inner tepal. Ornithogalum prasinum , Barker 7116 (NBG). C,

outer tepal; D, inner tepal. Scale bar: 2 mm. Artist: John Man-

ning.

Cape], Colesberg Div., Hondeblafs River [Hondeblafri-

vier], fl. ex hort. 1816/18, Burchell bulb 49 (K, holo.!,

iso.!). Albuca prasina (Ker Gawl.) J.C. Manning & Gold-

blatt: 92 (2009).

ACKNOWLEDGEMENTS

Type material was collected with a permit from

Northern Cape Nature Conservation and the necessary