Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 42,1

May 2012

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE (SANBI) PRETORIA

Obtainable from SANBI, Private Bag X101, Pretoria 0001,

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

request.

BOTHALIA

Bothalia is named in honour of General Louis Botha, first

Premier and Minister of Agriculture of the Union of South

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

the furtherance of botanical science. The main fields covered

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

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

published annually.

Bothalia Contents: five booklets containing a list of authors

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

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

(1996-2000); d) to vols 31-37 (2001-2007); to vols 38^11

(2008-2011).

FLORA OF SOUTHERN AFRICA (FSA)

A taxonomic treatise on the flora of the Republic of South

Africa, Lesotho, Swaziland, Namibia and Botswana. Contains

descriptions of families, genera, species, infraspecific taxa,

keys to genera and species, synonymy, literature and limited

specimen citations, as well as taxonomic and ecological notes.

Project discontinued.

FSA contributions 1-19 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:

25-40.

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 & P.H. Raven. 1997. Bothalia 27:

149-165.

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

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

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

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

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

249-252.

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

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

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

143-146.

1 8: Salicaceae s. str. M. Jordaan. 2005. Bothalia 35: 7-20.

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

2011. Bothalia 41: 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.

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

• Prodromus of South African Megafloras. Devonian to Lower

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

from: A. A. Balkema Marketing, Box 317, Claremont 7735,

RSA.

• Towards Gondwana Alive. Promoting biodiversity and stem-

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

• Heyday of the gymnosperms: systematics and biodiversity

of the Late Triassic Molteno fructifications, 2003, by J.M.

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 initated by or executed

in partnership with SANBI.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 42,1

Scientific Editor: J.C. Manning

Production Editor: Y. Steenkamp

SANBI

Biodiversity for Life

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2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI 01, Pretoria 0001

ISSN 0006 8241

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May 2012

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

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

Bergh, Dr N. South African National Biodiversity Institute, Cape Town, RSA.

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

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

Coates Palgrave, Mrs M. P.O. Chisipite, Harare, Zimbabwe.

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

Geerinck, Dr D. Universite Libre de Bruxelles, Brussels, Belgium.

Geldenhuys, Mr C. Northern Cape Department of Environment and Nature Conservation, Springbok,

RSA.

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

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

Hind. Dr D.J.N. Royal Botanic Gardens, Kew, UK.

Klak, Ms C. University of Cape Town, Cape Town, RSA.

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

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

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

Martinez-Azorin, Dr M. Instituto Universitario de la Biodiversidad, Alicante, Spain.

Renner, Prof. Dr S. Ludwig-Maximilians Universitat, Miinchen, Germany.

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

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

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

Trias Blasi, Dr A. Royal Botanic Gardens, Kew, UK.

Vogel, Dr J. Natural History Museum, London. England, UK.

Date of publication of Bothalia 41,2

Oct. 2011.

CONTENTS

Bothalia 42,1

1. Revision of the genus Sphenosty’lis (Fabaceae: Phaseoleae) in South Africa and Swaziland. A.N.

MOTEETEE and B.-E. VAN WYK 1

2. Anatomy of myxospermic diaspores of selected species in the Succulent Karoo, Namaqualand,

South Africa. H. FOTOUO MAKOUATE, M.W. VAN ROOYEN and C.F. VAN DER

MERWE 7

3. Recircumscription and distribution of elements of the ‘ Ceterach cordatwn complex ( Aspleniunr.

Aspleniaceae) in southern Africa. R.R. KLOPPER and N.R. CROUCH 15

4. A revision of Tecophilaeaceae subfam. Tecophilaeoideae in Africa. J.C. MANNING and P.

GOLDBLATT 21

5. Notes on African plants:

Aizoaceae. Corpuscularia gracillima is the correct binomial for the illegitimate C. gracilis. J.

NIEDERLE 49

Asteraceae. Berkheva jardineana ( Arctotideae — Gorteriinae), a new dwarf perennial from the

Swartruggens, Western Cape. J.C. MANNING and P. GOLDBLATT 57

Asteraceae. Osteospermum norlindhianum and O. nordenstamii, two new species of Osteo-

spermum sect. Trefenestratae (Calenduleae) from the Greater Cape Floristic Region.

J.C. MANNING, P. GOLDBLATT and N.A. HELME 59

Fabaceae. A new species of Bauhinia from southern Mozambique and the reinstatement of

Bauhinia macrantha. E.J.D. SCHMIDT 44

Hyacinthaceae. Drimia echinostachya , correcting a nomenclatural oversight. U. EGGLI and

N.R. CROUCH 43

Hyacinthaceae. New combinations in Ledebouria. J.C. MANNING and P. GOLDBLATT 47

Iridaceae. Taxonomic notes on Moraea (Irideae) in Congo and adjacent countries. P. GOLD-

BLATT and J.C. MANNING 49

Monimiaceae. Taxonomic and nomenclatural notes on the monotypic genus Xymalos and gen-

eral information on the family Monimiaceae. M. JORDAAN and M. LOTTER .... 51

6. Obituary. Eduardo Jose dos Santos Moreira Mendes (1924-2011). G.F. SMITH, E. FIGUEIRE-

DO and L. CATARINO 67

7. Obituary. Kathleen Dixon Gordon-Gray (nee Huntley) (1918-2012). E.J. MOLL 69

New combinations, names, sections, series, species and statuses in Bothalia 42,1 (2012)

Bauhinia burrowsii E.J.D. Schmidt, 44

Berkheya jardineana J.C. Manning & Goldblatt, sp. nov., 57

Corpuscularia gracillima (L. Bolus) Niederle, comb, nov., 49

Cyanella lutea L.f subsp. rosea (Eckl. ex Baker) J.C. Manning & Goldblatt , stat. nov., 37

Cyanella marlothii J.C. Manning & Goldblatt , sp. nov., 28

Drimia echinostachya (Baker) Eggli & N.R. Crouch, comb, nov., 43

Ledebouria linioseta (A.J.Hankey & P.D.Lebatha) J.C. Manning & Goldblatt , comb, nov., 48

Ledebouria transvaalensis (Van der Merwe ) J.C. Manning & Goldblatt, comb, nov., 48

Moraea schaijesiorum Geerinck, syn. nov., 50

Moraea ventricosa f. boussardiana Geerinck, syn. nov., 50

Moraea ventricosa var. macrantha f. wit tea na Geerinck, syn. nov., 50

Ornithogalum cooperi Baker, syn. nov., 43

Osteospermum nordenstamii J.C. Manning & Goldblatt, sp. nov., 62

Osteospermum norlindhianum J.C. Manning & Goldblatt, sp. nov., 60

Section Drimiopsis (Lindl. & Paxton) J.C. Manning & Goldblatt, stat. nov., 48

Section Resnova (Van der Merwe) J.C. Manning & Goldblatt, stat. nov., 48

Series Hyacinthoides J.C. Manning & Goldblatt, ser. nov., 32

Series Luteae J.C. Manning & Goldblatt, ser. nov., 35

IV

Bothalia 42,1: 1-6(2012)

Revision of the genus Sphenostylis (Fabaceae: Phaseoleae) in South

Africa and Swaziland

A.N. MOTEETEE*f and B.-E. VAN WYK*

Keywords: Fabaceae, Leguminosae, Nesphostylis Verde., Phaseoleae, Phaseolinae, southern Africa, Sphenostylis E.Mey., Vigna Savi

ABSTRACT

A taxonomic revision of the genus Sphenostylis E.Mey. (tribe Phaseoleae) in southern Africa is presented. The genus is

distinguishable by its characteristic wedge-shaped, dorsiventrally flattened style tip. Of the seven known species in the genus,

only two, S. angustifolia and S. marginata, occur in South Africa and Swaziland; and the former is endemic to these countries.

The two species differ in the size, shape, and venation of the leaflets; as well as the length of the petiole, peduncle, and pod. A

key to the two species is provided and the correct nomenclature, typification, and known geographical distributions are given.

INTRODUCTION

Sphenostylis E.Mey. belongs to the subtribe Phaseo-

linae of tribe Phaseoleae (Fabaceae). It is named for

the wedge-shaped, dorsiventrally flattened style (from

the Greek word spheno = wedge). It is a small genus

comprising seven species, all occurring in the tro-

pics and southern parts of the African continent (Gillett

et al. 1971; Potter 1992; Potter & Doyle 1994; Schrire

2005). The Indian species S. bracteata (Baker) Gillett,

transferred from Dolichos L. to Sphenostylis by Gillett

(1966), was moved to Nesphostylis Verde, by Potter &

Doyle (1994) based on a cladistic analysis of morpho-

logical data. Three species of Sphenostylis are used as

food sources in Africa. Sphenostylis stenocarpa (Hochst.

ex A. Rich.) Harms is known as African yam bean or

girigiri (Burkhill 1995), and cultivars of this tropical

African species are grown for their seeds and tubers in

tropical and West Africa. The leaves, flowers, pods, and

seeds of S. schweinfurthii Harms and S. erecta Hutch, ex

Baker f. are wild-harvested. The roots of S. erecta are

also used as fish poison (Potter 1992).

Sphenostylis appears to have a close relationship with

Nesphostylis (Potter & Doyle 1994) with which it shares

the dorsiventrally flattened style tip; but it differs by fea-

tures of the calyx (Lackey 1981), the standard petals,

and the stamens (Potter 1992). In Nesphostylis , the inner

surface of the calyx is pubescent, standard appendages

are present, the base of the vexillary stamen is hooked,

and the stamen apices are dilated. None of these cha-

racters are present in Sphenostylis. Based mainly on the

narrow pods, several botanists including Harvey (1862),

Bentham (1865) and Taubert (1894), relegated Spheno-

stylis into synonymy with the genus Vigna Savi (Gillett

1966). Harms (1899) reinstated the genus and expanded

it to include related species previously placed in Doli-

chos and Vigna. Sphenostylis differs from these two

genera by its distinctive style tip. Furthermore, Vigna

generally has peltate stipules (stipules not peltate in

Sphenostylis). In addition, a phylogeny based on mole-

cular data shows that Vigna and Sphenostylis are not

* Department of Botany and Plant Biotechnology, University of Jo-

hannesburg, P.O. Box 524, Auckland Park, 2006 Johannesburg.

|amoteetee@uj.ac.za.

MS. received: 2011-03-04.

as closely related as previously thought, and that Sphe-

nostylis is rather closer to Dolichos and Macrotyloma

(Wight & Am.) Verde. (Wojciechowski et al. 2004).

Sphenostylis can be distinguished from Dolichos, Ma-

crotyloma, and Vigna by the absence of appendages on

the standard petals (appendages present on the standard

petals in the latter three genera).

Sphenostylis is represented in South Africa and Swa-

ziland by two species, namely S. angustifolia Sond. and

S. marginata E.Mey.; the former being endemic to the

two countries.

MATERIALS AND METHODS

Plant material was studied mainly from herbarium

specimens housed in JRAU, NH, NU, and PRE (acro-

nyms after Holmgren et al. 1990); and also in the field.

Habit affinities are described according to Mucina &

Rutherford (2006).

TAXONOMY

Sphenostylis E.Mey., Commentariorum de Plantis

Africae Australioris: 148 (1836); Baker fi: 670 (1929);

Burtt Davy: 418 (1932); E. Phillips: 427 (1951); Wilc-

zek: 273 (1954); Verde.: 389 (1970); J.B.Gillett et al.:

670 (1971); Compton: 286 (1975); R. A. Dyer: 275

(1975); Potter & Doyle: 389-406 (1994); Germish.: 296

(2000); Verde. & Doygaard: 68 (2001). Type species:

S. marginata E.Mey.

Prostrate, climbing (twining), or erect shrubs or per-

ennial herbs, arising from a thick, woody rootstock.

Leaves pinnately 3-foliolate (Figure ID); stipules per-

sistent, ovate-lanceolate or ovate-acuminate; leaflets

ovate, elliptic or linear, with four linear stipels, one at

the base of each lateral leaflet and two at the base of the

terminal one (Figure IE), appressed-pubescent when

young but becoming glabrescent with age. Inflorescence:

flowers in congested heads on long peduncles, peduncles

much longer than leaves; bracts linear or oblong-lan-

ceolate, small, falling off at a very early stage or absent;

bracteoles 2, ovate-lanceolate or oblong-ovate, caducous

or persistent. Calyx tube campanulate, bilabiate, the two

Bothalia 42,1 (2012)

FIGURE 1. — Sphenostylis marginata (A, B, D) and S. angustifolia (C, E-H): A, flowers showing the twisted standard petals; B, flower showing

the cuneate style tip; C, keel tip and cuneate style tip; D, leaf in adaxial view; E, flowering branch (note the stipules and stipels); F, pod; G,

flower in front view (note the twisted petals); H, seed. Scale bars: A, 8 mm; B, 12 mm; C, E-G: 5 mm; D, H: 2 mm. Photographs: A, B, D

by David Styles; C, E-H by Ben-Erik van Wyk.

lobes of the upper lip partially or entirely connate, lobes

very short and blunt. Corolla purple, purplish pink, vio-

let, whitish pink, or yellow; standard suborbicular, sym-

metrical or twisted (Figure 1A & 1G) with 2 indexed

auricles, but without appendages, with well-developed,

channelled claw; wings obliquely obovate, eared near

base, with short linear claw; keel incurved, ± concave-

convex beaked, with short linear claw. Androecium

diadelphous (with 9 stamens fused into a tube, vexil-

lary stamen free), 5 basifixed anthers alternating with 5

dorsifixed anthers. Ovary narrowly oblong, pubescent,

3 12-ovuled; style penicillate below the stigma; stigma

dorsiventrally flattened, ciliate on the margins (Figures

IB & 1C). Fruit linear, compressed, twisting after dehis-

cence, glabrescent to densely silky, with persistent style

at tip (Figure I F), 2-several-seeded, dehiscent. Seeds

oblong, uniformly black or brown to reddish brown

speckled black, minutely papillose (Figure 1H).

Key to species of Sphenostylis in South Africa and

Swaziland:

la Leaflets 20—45 mm wide; petiole 50-70 mm long; peduncles

220-300 mm long; fruit 95-120 mm long S. marginata

lb Leaflets 10-19 mm wide; petiole 8-16 mm long; peduncles

75-1 15 mm long; fruit 55-85 mm long S. angustifolia

Bothalia 42,1 (2012)

3

1. Sphenostylis angustifolia Sond. in Linnaea 23:

33 (1850); R.A.Dyer: t 1010 (1947); Burtt Davy: 418

(1932); Compton: 287 (1975). Vigna angustifolia (Sond.)

Benth. ex Harv.: 240 (1862). Type: South Africa, Gau-

teng, 2528 (Pretoria): Magaliesberg, (-DC), Zeyher 524

(S, specimen on the left, lecto.!, here designated; BM!,

K!, isolecto.). Syntype: KwaZulu-Natal, 2931 (Stanger):

Port Natal [now Durban], (-CC), Gueinzius 624 ( S ! ).

Note: the Zeyher specimen in the Sonder Herbarium

in S is chosen as lectotype because this is probably the

specimen that Sonder used in his description. The twig

on the left is chosen because it bears a flower and some

immature fruits. Curiously Potter (in sched.) chose the

Zeyher specimen in K as lectotype but this choice was

apparently never published.

Erect suffrutex, rarely climbing and twining, much-

branched, up to 0.5 m tall, spreading. Leaflets oblong or

linear-lanceolate, 30-60 x ( 7—) 1 0—1 9 mm; petiole 8-16

mm long; stipules ovate-acuminate, 3. 5-5.0 x 2-3 mm.

Inflorescences subumbellate axillary racemes; peduncles

(55 — )75— 1 1 5 mm long, with 2-4 flowers; flowers pink

or purple, with a whitish or yellowish centre, 12-25 mm

long; bracts linear, ± 1 .5 mm long, caducous; bracteoles

ovate-lanceolate, 2. 0-2. 5 x ± 1.5 mm, persistent. Calyx

sparsely pubescent, with ± equal lips, upper lip 6-8 mm

long, lower lip 6-8 mm long; lobes rounded, those of the

upper lip joined for almost their entire length. Corolla

pinkish red; standard suborbicular, 15-25 x 15-26 mm,

glabrous; wings obovate, 14-24 * 4-7 mm, without sur-

face sculpturing, apex rounded; keel falcate, 14—24 x

6-10 mm, apex rounded, pocket absent. Ovary 9-13 mm

long, linear-oblong. Fruit linear, 55-85 x 5-6 mm, gla-

brescent, many-seeded. Seeds reniform, ± 7 x ± 4 mm,

brown, reddish brown speckled black, or uniformly

black (Figure 2). Flowering time : Sept.-Feb.

Diagnostic characters'. Based on Potter & Doyle’s

cladistic analysis (1994), Sphenostylis angustifolia

appears to have close affinities with S. zimbabweensis

R.Mithen, which is restricted to the Highlands of Zim-

babwe. The two species share the deciduous bracts and

the persistent bracteoles. Sphenostylis zimbabweensis

is, however, differentiated by the more prostrate growth

form (as opposed to the erect growth form of S. angusti-

folia)1, and the broader, ovate to elliptic leaflets (leaflets

narrower and oblong to linear-lanceolate in S. angus-

tifolia). Sphenostylis angustifolia can be distinguished

from S. marginata by the narrower leaflets and the much

shorter petioles, peduncles, and pods (dimensions are

given in the key).

Distribution and habitat'. Sphenostylis angustifolia

occurs in South Africa (Fimpopo, North-West, Gauteng,

Mpumalanga and KwaZulu-Natal Provinces) and Swazi-

land (Figure 3). It grows scattered in Zeerust Thornveld

(SVcb 3), Central Sandy Bushveld (SVcb 12), Soutpans-

berg Mountain Bushveld (SVcb 21), Polokwane Plateau

Bushveld (SVcb 23), Granite Fowveld (SV1 3), Tza-

FIGURE 2. — Vegetative and reproductive morphology of Sphenostylis angustifolia'. A, flowering branch; B1 & B2, abaxial view of bracteoles; C,

calyx opened out with upper lobes to left; D, standard petal; E, wing petal; F, keel petal; G, stamens; H, pistil; I. lateral view of pod. Vouch-

ers; A from A.O.D. Mogg 35461 (JRAU); B-I from B-E. van Wyk 1438 (JRAU). Scale bar: A, 30 mm; B, 1 mm; C-F, H, 3 mm; G, 2 mm;

I, 10 mm.

4

Bothalia 42,1 (2012)

FIGURE 3. — Distribution of Sphenostylis angustifolia.

neen Sour Bushveld (SVI 8), Legogote Sour Bushveld

(SVI 9), Swaziland Sour Bushveld (SVI 14), Southern

Lebombo Bushveld (SVI 16), Carletonville Dolomite

Grassland (Gh 15), Soweto Highveld Grassland (Gm 8),

Rand Highveld Grassland (Gm 11), KaNgwane Montane

Grassland (Gm 16), Maputaland Coasal Belt (CB 1),

KwaZulu-Natal Coastal Belt (CB 3).

Additional specimens examined

LIMPOPO. — 2230 (Messina): Soutpansberg, Entabeni Forestry

Station at Muchindudi Fall, (-CC), 27 Jan. 1954, L.E. Codd 8392

(PRE). 2329 (Polokwane): Polokwane, Farm Eersteling, on slope of

mtn, (-AB), 1 Jan. 1992, G.J. du Toit 2238 (NH). 2330 (Tzaneen):

Modjadjis Reserve near Duiwelskloof, (-CB), 24 May 1938, J.D.

Krige 164 (PRE). 2428 (Nylstroom): Waterberg, (-AD). 6 Nov. 1978,

G. Germishuizen 909 (PRE); Naboomfontein, (-BD), 13 Dec. 1934,

E.E. Galpin 133130 (PRE); between Warmbaths and Pietersburg

[Polokwane], (-CB), 4 Nov. 1985, B.J. Pienaar 636 (PRE); 18 km

from Nylstroom on road to Warmbaths near Groot Nyl turn-off, (—

CD), 4 Nov. 1985, G. Germishuizen 3347 (PRE). 2429 (Zebediela):

near Daggakraal 50 m NE of Potgietersrus, (-AA), 3 Jan. 1954, B.

Maguire 2569 (PRE); Arabie, Camp 1, (-CD), 7 Jan. 2007, W. Ellery

357 (PRE). 2430 (Pilgrim’s Rest): Lekgalameetse Nature Reserve,

The Downs, SE of Makwens, (-AA), 16 Oct. 1985, M. Stalmans 718

(PRE).

NORTH-WEST.— 2526 (Zeerust): Marico Distr., ± 10 km NE of

Wondergat, (-CD), 6 Feb. 1983, C. Reid 676 (PRE). 2527 (Rusten-

burg): Rustenburg Nature Reserve, (-CA), 27 Feb. 1970, N. Jacobsen

845 (PRE). 2627 (Potchefstroom): on road from Frederikstad to Rys-

mierbult, (-AC), 31 Oct. 1978, B. Ubbink 733 (PRE); Ventersdorp,

Goedgedacht, (-CA), 29 Dec. 1930, J.D. Sutton 511 (PRE).

GAUTENG. — 2528 (Pretoria): La Montagne Rand, N of Cham-

bord W/S, (-CA), 30 Sept. 1978, A.E. van Wyk 2405 (PRE); 28 miles

[45 km] from the National Herbarium on road to Boekenshoutskloof,

(-CB), 15 Apr. 1971, L.A. Coetzer 73 (PRE); Fountains Valley, (-

CC), 3 Oct. 1948, J.M. Watt 4565 (PRE); Doomkloof, Smutskoppie,

(-CD); 24 Nov. 1985, B-E. van Wyk 1438 (JRAU); Renosterkop, NE

of Bronkhorstspruit, (-DB); 6 Dec. 1987, B-E. van Wyk 2730 (JRAU).

2628 (Johannesburg): Melville Koppies Nature Reserve, (-AA), 4

Feb. 1987, B-E. van Wyk 2600-2604 (JRAU); Heidelberg, (-AD), Nov.

1927, A. Thode A1 311 (PRE).

MPUMALANGA. — 2430 (Pilgrim’s Rest): Drie Rondavels look-

out. (-BC), 2 Feb. 1982, A.E. van Wyk, R. Dahlgren & P.D.F. Kok

5487 (PRE). 2529 (Witbank): Loskopdam Game Reserve, (-AD),

13 Dec. 1966, G.K. Theron 728 (JRAU, PRE); between Witbank

and Middelburg along N4 highway, (-CD), 20 Feb. 1991, P. Her-

man 1338 (PRE). 2530 (Lydenburg): 7 km S of Sabie, (-BB), 29

Sept. 2005, J.J. Meyer 4530 (PRE); Wonderkloof Nature Reserve, (—

BC), 17 Nov. 1978, J.P. Kluge 1408 (PRE). 2531 (Komatipoort): Ka-

Ngwane, Songimvelo Game Reserve, (-CC), 8 Dec. 1992, N.L. Meyer

29 (PRE).

SWAZILAND. — 2631 (Mbabane): Hhohho Dist., Nyokane, 15 km

from Piggs Peak-Mbabane turnoff on road to Maphalaleni, (-AB),

23 Oct. 1963, R.H. Compton 31707 (PRE); Dabriach, (-AC), 18 Oct.

1958, R.H. Compton 28097 (PRE); Hlatikulu, (-CD), Oct. 1910, M

Steward 10081 (PRE). 2632 (Bela Vista): Lebombo Mountains, near

fence separating Swaziland and Mozambique, (-CA), 22 Nov. 2002,

M.K. Maserumule 74 (PRE).

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Beacon Hill, off

Panorama Terrace, Wyebank, (-DD), 10 Jan. 2005, D. Styles 2180

(NH).

2. Sphenostylis marginata E.Mey., Commentari-

orum de Plantis Africae Australioris: 148 (Feb. 1836);

Baker f.: 148 (1929); Wilczek 6: 274 (1954); Verdoorn:

t. 1521 (1968); J.B.Gillett et ah: 671 (1971); Compton:

287 (1975); Lock: 438 (1989); Potter & Doyle: 403

(1994); Verde. & Doygaard.: 71 (2001). Vigna margin-

ata (E.Mey.) Benth. ex Harv.: 240 (1862); De Wild.: 98

(1921). Type: KwaZulu-Natal, 3030 (Port Shepstone):

’ in graminosis ad ostia fluvii Omsamculo' [mouth of

Umzimkulu River], (-CB), Drege s.n. (K!, lecto., desig-

nated by Potter & Doyle (1994) [as ‘holo’.], G!, MO!,

P!, isolecto.)

Note: although the P specimen which bears an origi-

nal Drege label with locality details corresponding

exactly to those given in the prognosis as well as Mey-

er’s handwriting (fmihV) would have been more appro-

priately selected as lectotype. Potter & Doyle’s (1994)

designation of the K specimen as the holotype consti-

tutes effective lectotypification (Art. 9.8).

Prostrate suffrutex with twining stems up to 1.5 m

long, arising from a woody rootstock. Leaflets ovate,

elliptic or oblong, 45—1 1 0(— 125) x 20 — 40( — 45) mm; pet-

iole ( 1 5— )50— 70 mm long; stipules oblong-lanceolate or

ovate, 3-7 x 2-4 mm. Inflorescence pseudo-umbellate,

axillary raceme, long-stalked; peduncles 220-300 mm

long, few-flowered; flowers mauve, 14-16 mm long;

bracts oblong-lanceolate, ± 1.5 mm long, caducous,

bracteoles ovate-lanceolate, 2-3 x 1. 5-2.0 mm, persist-

ent. Calyx pubescent, lobes rounded, with equal lips,

upper and lower lips 5-6 mm long, lobes of upper lip

joined for almost their entire length. Corolla pinkish

red; standard broadly obovate, 13-17 x 14-23 mm, gla-

brous; wings obovate, deep purplish pink 10-17 x 4-6

mm, without surface sculpturing, apex rounded; keel ±

equal to wings, 12-18 x 4-6 mm, paler or white, apex

rounded, pocket absent. Ovary 10-15 mm long, linear-

oblong. Fruit linear, 95-120 x 4-8 mm, glabrescent,

5-8-seeded. Seeds reniform, ± 5.0 x ± 3.5 mm, brown

or reddish brown speckled black (Figure 4). Flowering

time : Nov.-Feb.

Diagnostic characters : Sphenostylis marginata dif-

fers from S. angustifolia in having broader leaflets and

longer petioles, peduncles, and pods (see key for dimen-

sions).

Verdcourt (1970) divided what he called the Spheno-

stylis marginata complex, which included the two cen-

tral and East African species S. erecta (Baker f.) Baker f.

and S. obtusifolia Harms, into three subspecies, namely

subsp. marginata (occurring only in South Africa and

Swaziland), subsp. obtusifolia (Harms) Verde., and

subsp erecta (Baker f.) Verde. However, based on mor-

phological and DNA data. Potter & Doyle (1994) pro-

Bothalia 42,1 (2012)

5

FIGURE 4. — Vegetative and reproductive morphology of Sphenostvlis marginata: Al, flowering branch; A2, inflorescence; B1 & B2, abaxial view

of bracteoles; C, calyx opened out with upper lobes to left; D, standard petal; E, wing petal; F, keel petal; G, stamens; H, pistil; I, lateral view

of pod. Vouchers; A from N. Grobbelaar 1648 (PRE); B-F from Commins 855 (PRE); H from Acocks 20928 (PRE); I from N. Grobbelaar

1648 (PRE). Scale bar: Al, A2, 30 mm; B, 1 mm; C, 2 mm; D-F, H, 3 mm; G, 4 mm; I, 15 mm.

posed a narrow view of S. marginata by limiting the

species concept to subsp. marginata (the other two sub-

species were excluded). Sphenostylis marginata subsp.

erecta was restored to species level and subspecies

obtusifolia was transferred to 5. erecta (as subspecies).

Distribution and habitat: Sphenostylis marginata

occurs in South Africa (Limpopo, Mpumalanga, Kwa-

Zulu-Natal and Eastern Cape Provinces), and Swaziland

(Figure 5). It grows in Central Sandy Bushveld (SVcb

12), Ohrigstad Mountain Bushveld (SVcb 26), Gran-

FIGURE 5. — Distribution of Sphenostylis marginata.

ite Lowveld (SVI 3), Legogote Sour Bushveld (SVI

9), Zululand Lowveld (SVI 23), PaulPietersburg Moist

Grassland (Gm 15), KaNgwane Montane Grassland (Gm

16), Income Sandy Grassland (Gs 7), Ngongoni Veld

(SVs 4), Eastern Valley Bushveld (SVs 6), Maputaland

Coasal Belt (CB 1), and KwaZulu-Natal Coastal Belt

(CB 3).

Additional specimens examined

LIMPOPO. — 2428 (Nylstroom): 13 km NE of Wannbaths [Bela-

Bela] on road to Nylstroom, (-CD), 18 Nov. 1981, C. Reid 439 (PRE).

MPUMALANGA. — 2430 (Pilgrim’s Rest): foothills SE of

Magalieskop, Mariepskop Dist., (-DB), 6 Dec. 1990, H.P. van der

Schijf 5878 (PRE). 2531 (Komatipoort): 25 km from White River to

Hazyview, near White Waters Forest Station, (-AA), 3 Jan. 1984, M

Jordaan 298 (PRE); Kruger National Park, Lower Sabie dam, (-BB),

26 Nov. 1990, V.R. Bredenkamp 468 (PRE); Eerste Geluk no. 16,

Uitkyk, (-CA), 26 Mar. 1975, C.H. Stirton 1729 (PRE).

SWAZILAND. — 2631 (Mbabane): Hhohho Dist., Masilela

area, on Maphalaleni Rd., (-AB), 27 Jan. 1994, G. Ger-

mishuizen 7152 (PRE); Little Usutu River, (-AC), 27 Oct.

1956, R.H. Compton 26163 (NH, PRE); Stegi, (-BD), 22

Dec. 1960, R.H. Compton 30388 (NH, PRE); S of Mankai-

ana, (-CA), 6 Nov. 1949, J.L. Sidey 1933 (PRE). 2731 (Louws-

burg): at camp on hilltop before reaching Hluti, (-BA), 8 1931, LB.

Pole-Evans 3364 (PRE).

KWAZULU-NATAL. — 2730 (Vryheid): 0.9 km towards Vryheid

from Natal Spa, Freddie Coetzee’s farm, (-BD), 2 Dec. 1988, P.D.F.

Kok & B.J. Pienaar 1282 (PRE). 2731 (Louwsburg): Nongoma, (-

DC), 20 Nov. 1960, M.J. Wells 2060 (PRE). 2830 (Dundee): hill near

Glencoe, (-AA), 22 Feb. 1993, J. Medley-Wood 4823 (PRE); Dundee

Dist., on banks of Buffalo River, 1 km S of P.O. Vantsdrift, (-AB),

22 Dec. 1946, L.E. Codd 2377 (PRE); Elandslaagte, Blanerne Farm,

6

Bothalia 42,1 (2012)

32 km from Ladysmith on road to Newcastle, (-BA), 15 Nov. 1994,

A.M. Ngwenya 1295 (PRE). 2831 (Nkandla): Babanango Dist., val-

ley N of Izulu Hill, (-CC), 16 Jan. 1946, J.P.H. Acocks 12317 (PRE).

2832 (Mtubatuba): Masundwini, Hluhluwe Game Reserve, (-AA), 7

Nov. 1971, P.M. Hitchins 635 (NU, PRE); Palm Ridge Farm, (-AC),

3 Oct. 1967, E.R. Harrison 128 (PRE). 2931 (Stanger): Twinstreams

Farm, Mtunzini, (-DC), 12 Oct. 1984, I. Garland s.n.. & G. Nichols

805 (PRE). 3030 (Port Shepstone): St Michaels-on-Sea, (-AB), 1 Feb.

1985, B.J. Pienaar 587 (PRE); Port Shepstone (-CD), 24 Mar. 1967,

R.G. Strey 7420 (PRE); Hibberdene, (-DA), 09 Mar. 1970, R.G. Strey

9701 (NU).

ACKNOWLEDGEMENTS

The curator of PRE is gratefully acknowledged for

assistance with herbarium material and the curator of P

is thanked for the image of the Sphenostylis marginata

type specimen. The University of Johannesburg and the

National Research Foundation provided financial sup-

port. We thank David Styles for showing us a population

of S', marginata and for providing photographs of the

species.

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can National Biodiversity Institute, Pretoria.

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ture, South Africa.

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nostylis and Nesphostvlis (Leguminosae: Phaseoleae) based on

morphological and chloroplast DNA data. Systematic Botany 19:

389—406. "

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kinder, & M. Lock, Legumes of the World: 393 — 430. Royal

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33. Berlin.

TAUBERT. P 1 894. Leguminosae. In A. Engler & K. Prantl, Die natiir-

lichen Pflanzenfamilien 3: 70-399. Engelmann, Leipzig.

VERDCOURT. B. 1970. Studies in the Leguminosae-Papilionoideae for

the Flora of tropical east Africa: 111. Kew Bulletin 24: 380—442.

VERDCOURT, B. & DOYGAARD, S. 2001. Sphenostylis. In G.V.

Pope & R.M. Polhill (eds). Flora Zambesiaca 3, 5: 68. Royal

Botanic Gardens, Kew.

VERDOORN, I.C. 1968. Sphenostylis marginata. Flowering plants of

Africa: t. 1521.

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Urundi VI: 274. Institut National pour l’Etude Agronomique du

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WOJCIECHOWSKI, M.F., LAVIN, M. & SANDERSON, M.J. 2004.

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Bothalia 42,1: 7-13 (2012)

Anatomy of myxospermic diaspores of selected species in the

Succulent Karoo, Namaqualand, South Africa

H. FOTOUO MAKOUATE*, M.W. VAN ROOYEN*t and C.F. VAN DER MERWE**

Keywords: arid regions, diaspores, dispersal, myxospermy, Namaqualand, scanning electron microscopy. Succulent Karoo

ABSTRACT

Environmental conditions encountered in arid ecosystems differ vastly from those in more mesic ecosystems. Dispersal

strategies in arid environments reflect these differences and many mechanisms have evolved that restrict or hinder dispersal.

Myxospermy is a trait developed by plant species from arid regions to restrict diaspore dispersal by means of an anchorage

mechanism. Several of the abundant plant species in Namaqualand, within the arid Succulent Karoo Biome, display myxo-

spermy. Diaspores of these species produce copious amounts of mucilage when they are moistened and are anchored to the

soil once the mucilage dries out again. This study investigated the origin of the mucilaginous layer of 12 species anatomically,

using both light and scanning electron microscopy. The mucilage production of the species investigated could best be grouped

into three types: 1, epidermal and sub-epidermal cells of seeds and achenes; 2, specialized tissue in wings or the pappus of

achenes; and 3, mucilage excreting hairs. Previous systems for classifying the different types of mucilage production did not

recognize the mucilaginous nature of wings or a pappus. A short note on the composition of the mucilage is included.

INTRODUCTION

Plants have developed many functional traits that

allow them to adapt and survive in different environ-

ments. Seed dispersal is an important functional trait

that influences the population structure and the spatial

and temporal turnover of species within a plant commu-

nity. Myxospermy is the phenomenon where the epider-

mis of the diaspore (seed/fruit) contains mucilaginous

cells, which swell and become sticky when in contact

with water (Gutterman & Shem-Tov 1997; Van Rheede

van Oudtshoom & Van Rooyen 1999). Myxospermy is

classified as an antitelechoric dispersal mechanism and

is encountered more often in arid than in mesic envi-

ronments. The original explanation for the prevalence

of antitelechory in desert environments was that these

mechanisms were adaptive responses to the particularly

high mortality of dispersed seeds in deserts and that

they had evolved as mechanisms to reclaim the mother

site (Murbeck 1919; Zohary 1937; Stopp 1958; Ell-

ner & Shmida 1981; Van Rooyen et al. 1990). Ellner &

Shmida (1981 ) however, argued that antitelechory was a

side effect of characters whose adaptive value was not

directly related to dispersal. Among the benefits derived

from antitelechory are the spreading of germination over

time and the provision of suitable conditions for germi-

nation and subsequent seedling establishment.

There are many divergent ideas about the ecological

importance of myxospermy, and it probably fulfils vari-

ous functions. When wet, myxospermic diaspores adhere

to the soil; and after drying out, remain glued to soil par-

ticles and thus resist being carried to unfavourable loca-

tions by wind. Flowever, wet diaspores can also adhere to

the feet or fur of animals and be dispersed zoochorically.

The good contact between the mucilage layer and the

soil surface increases water absorption for germination.

* Department of Plant Science, University of Pretoria, 0002 Pretoria,

t greteI.vanrooyen@up.ac.za.

** Laboratory for Microscopy and Microanalysis, University of Preto-

ria, 0002 Pretoria.

MS. received: 2010-01-15.

and as a result, myxospermic species germinate more

successfully on the surface of the soil than non-myxo-

spermic species (Bregman & Graven 1997; Zaady et al.

1997; Van Rheede van Oudtshoom & Van Rooyen 1999).

The water held by the mucilage could also serve as a

water reservoir for seedling establishment (Gutterman

et al. 1967; Gutterman 1993, 1996), although this func-

tion of the mucilage has been challenged (Murbeck 1919;

Grubert 1974). Furthermore, the adherence of the seed to

the soil by the mucilage prevents massive collection by

seed predators (Gutterman 1993), and the repeated wet-

ting of mucilaginous seeds during many nights with dew

may affect the repair mechanisms of the DNA, cell mem-

branes, and organelles; thereby enhancing seed viabil-

ity for many years (Osborne et al. 1980/1981; Leprince

et al. 1993; Huang et al. 2008). The mucilage has also

been reported to have a stimulatory action on germina-

tion (Gat-Tilman 1995) or it may control the germination

process by excluding the passage of oxygen when there

is excess moisture (Gutterman et al. 1967, 1973; Gutter-

man 1 996; Tamara et al. 2000).

Although several studies have drawn the attention to

the ecological importance of myxospermy in the flora

of Namaqualand (Rosch 1977; Van Rooyen et al. 1990;

Van Rheede van Oudtshoom & Van Rooyen 1999), the

germination behaviour of myxospermic species (Fot-

ouo Makouate 2008) and the response of myxospermic

species to grazing pressure (Fotouo Makouate 2008),

little attention has thus far been given to the origin and

chemical composition of the mucilage of these myxo-

spermic diaspores. The objectives of this study were to

investigate the origin and the chemical composition of

the mucilage of a representative sample of myxospermic

species in Namaqualand, South Africa.

STUDY AREA AND SPECIES

Diaspores of the species investigated in this study

were all collected in the Namaqualand Hardeveld Biore-

gion of the Succulent Karoo (Mucina & Rutherford

Bothalia 42,1 (2012)

2006). The Succulent Karoo is an arid winter rainfall

region stretching along the West Coast of South Africa

and Namibia; and is recognized by the IUCN as one of

only two entirely arid global hotspots of biodiversity

(Conservation International 2009). Many of the biologi-

cally unique features of this biome have been attributed

to its climate, i.e. the effective and relatively predict-

able seasonal rainfall, and the relatively moderate win-

ter temperatures (Mucina & Rutherford 2006). A promi-

nent feature of the Namaqualand Hardeveld Bioregion is

the extravagant spring floral display of winter-growing

annuals (Van Rooyen 1999).

Representative species of four families were investi-

gated: within the Lamiaceae, the perennial shrub Salvia

dentata\ within the Acanthaceae, two dwarf peren-

nial shrubs Acanthopsis horrida and Blepharis jure at a\

within the Brassicaceae, the annual Heliophila thunber-

gii var. thunbergii; and within the Asteraceae, the per-

ennial shrubs Pentzia incana and Othonna cylindrica

as well as the annual species Cotida barbata, Monocu-

lus (Tripteris) hyoseroides, Senecio arenarius, Ursinia

cakilefolia, Oncosiphon grandiflorum and Foveolina

dichotoma.

MATERIALS AND METHODS

Scanning electron microscopy (SEM)

Dry seeds/achenes were mounted on a stub and made

conductive with Ru04 vapour (Van der Merwe & Pea-

cock 1999). For wet samples, seeds/achenes of each

species were soaked in water for 24 hours before fixa-

tion. Seeds/achenes were then fixed in 2.5% glutaral-

dehyde in 0.075 M phosphate buffer. pH 7.4, for one

hour, and rinsed three times (five minutes per rinse) in

0.075 M phosphate buffer and dehydrated in an ethanol

series with the respective concentrations of 30%, 50%,

70%, 90%, and 100%. This was followed by critical

point drying in liquid CO,, whereafter the seeds/achenes

were mounted on a stub and made conductive with RuO,

vapour. Samples were viewed with the use of a JSM 840

scanning electron microscope (SEM) (JEOL, Tokyo,

Japan), and photographs were taken with the computer

program Orion 6.60.4.

Light microscopy

Further studies were done on diaspores where the

origin of the mucilage could not be clearly established

with the aid of an SEM. Dry samples were fixed in 4%

(v/v) formaldehyde in 50% (v/v) ethanol (FAA) to pre-

vent mucilage release. The wet samples were immersed

in water for 24 hours. Wet samples were fixed in 2.5%

glutaraldehyde in 0.075 M phosphate buffer, pH 7.4, for

one hour, and rinsed three times (five minutes per rinse)

in 0.075 M phosphate buffer, and dehydrated in an etha-

nol series with the respective concentrations of 30%,

50%, 70%, 90% and 100%. Samples were subsequently

infiltrated over two days with pure LR- White resin and

polymerized at 65°C for 24 to 36 hours. Sections were

cut with a Reichert Ultracut E microtome. Each sec-

tion was stained, mounted in immersion oil, and viewed

with a Nikon Optiphot light microscope (LM) (Tokyo,

Japan). Photographs were taken with a Nikon Digital

camera DXM 1 200 (Tokyo, Japan) and the computer

program Nikon ACT- 1 (Tokyo, Japan).

Diaspore staining was done according to various

manuals of microscopic staining (McClean & Ivimey

Cook 1941; O’Brien & McCully 1981; Lawton &

Ettridge 1986). Seven different stains were used to test

for the presence of various compounds, the details of

which are given in Table 1 .

TABLE 1 . — Stains used for various compounds and expected results

RESULTS AND DISCUSSION

A n atomical investigation

On the basis of the anatomical origin of the mucilage,

Zohary (1937) distinguished seven different types. Gru-

bert (1974) recognized many more types, but his three

main categories were based on whether the origin was

1, restricted to epidermal structures, e.g. epidermis or

hairs; 2, from epidermal and sub-epidermal layers; or 3,

restricted to sub-epidermal layers.

Neither the classification systems of Zohary (1937)

or Grubert (1974) accommodate diaspores where the

mucilage production occurs on wings of achenes, as

was reported in the present study. In general, diaspores

of most myxospermic species are smooth-coated and do

not contain obvious appendages to advance telechory.

Contrary to the belief that none of the myxospermic

Asteraceae possess a pappus (Grubert 1974), many

myxospermic species from Namaqualand are winged or

possess a winged pappus. The presence of wings would

allow anemochoric dispersal during phase I dispersal

(Chambers & MacMahon 1994), but further anemo-

choric dispersal would be prevented once the achenes

were moistened and remained attached to soil particles.

The mucilage production of the species investigated

in the present study could best be grouped into three

main types:

1 Epidermal and sub-epidermal cells of seeds and

achenes:

1.1 Cells of seeds bursting only in the centre

1 .2 Cells of achenes bursting across the entire surface

2 Specialized tissue in wings or pappus of achenes

3 Hairs:

3.1 On achenes

3.2 On seeds

Epidermal mucilage

Heliophila thunbergii var. thunbergii (Brassicaceae)

Bothalia 42,1 (2012)

9

FIGURE 1. — Heliophila thunbergii var. thunbergii, seeds. A-D, SEM: A, entire dry seed; B, high magnification of dry epidermis; C, high magnifi-

cation of wet seed surface; D, section of wet seed. E, F, LM: E, section of the dry seed; F, cross section of wet epidermis, e, epidermal layer,

m, mucilage.

The scanning electron micrograph of dry seeds of

H. thunbergii var. thunbergii reveals a circular seed

(Figure 1A). The epidermal layer (e) consists of circular

cells with a raised margin and a cavity in the centre (Fig-

ure IB). Upon wetting, the mucilage swells and the epi-

dermal cells become incapable of containing the excess

mucilage and therefore rupture. Once wet, the seed is

covered by a gelatinous film (Figure 1C), similar to the

mucilaginous seed coat of two species of the Brassi-

caceae described by Gutterman & Shem-Tov (1997).

The thin section of the dry seed also reveals the epider-

mal cells filled with mucilage (m) (Figure IE). The sec-

tion of the wet seed reveals the epidermis with ruptured

cells from which filaments of mucilage (m) emerge (Fig-

ure ID, F).

Salvia lanceolata (Lamiaceae)

A similar pattern was observed for the seeds of S. lan-

ceolata (Lamiaceae). Once in contact with water, the

outer layer of the epidermis absorbs water quickly, the

epidermis cells swell, the cuticle ruptures and the con-

tents are excreted as long, continuous threads. Hedge

(1970) reported that the contents of the epidermis cells

of some Lamiaceae are secreted in the shape of long,

continuous, helically coiled threads.

FIGURE 2. — Cotula barbata, achenes. A, B, D, E, SEM: A, entire dry achene; B, high magnification of the epidermis; D, entire wet achene; E, high

magnification of wet epidermis covered with thick layer of mucilage. C, F, LM: C, cross section of dry achene coat, with mucilage (arrow)

retained in epidermal layer; F, cross section of wet epidermis of which outer layer has burst, releasing mucilage from epidermal cells, m,

mucilage.

10

Bothalia 42,1 (2012)

FIGURE 3. — Foveolina dichotoma, achenes. A-C, SEM: A, entire achene of F. dichotoma ; B, high magnification of the epidermis of F. dichotoma ;

C, wet achene. Oncosiphon grandiflorum , achenes: D, E, LM: D, thin cross section of dry epidermis; E, thin cross section of wet epidermis

with broken epidermal cells that liberate mucilage, e, epidermal layer, m, mucilage.

Cotula barbata (Asteraceae)

In the dry state, the one side of the achene of C. bar-

bata is undulated (Figure 2A), whereas the other side of

the achene is smooth (Figure 2B). The mucilage struc-

ture observed in C. barbata is similar to that of H. thun-

bergii var. thunbergii with the mucilage contained in the

epidermal layer (Figure 2C). When C. barbata achenes

come into contact with water (Figure 2D), the epidermal

cells burst across the entire outer surface (Figure 2E,

F) and not only the centre of the cell as in the case of

H. thunbergii var. thunbergii.

Foveolina dichotoma. Oncosiphon grandiflorum and

Pentzia incana (Asteraceae)

These three species are taxonomically closely related,

and it appears that they have the same way of producing

mucilage. The SEMs show that the epidermis of the dry

achenes of F. dichotoma consists of parallel longitudinal

cells (Figure 3 A, B), as was also found for O. grandi-

florum and P. incana. Following wetting, fixation, and

critical point drying, the achenes are covered with a

gelatinous film but the pappus remains unchanged (Fig-

ure 3C). Light microscope sections of the dry achene of

O. grandiflorum show that the epidermal cells are filled

FIGURE 4. — Ursinia cakilefolia,

SEM of achenes: A, entire

dry achene with winged pap-

pus; B, high magnification

of dry winged pappus con-

sisting of radially elongated

cells; C, entire wet achene

with uncoiled winged pappus;

D, high magnification of wet

winged pappus showing long

strands of uncoiled mucilage,

wp, winged pappus.

Bothalia 42,1 (2012)

with mucilage (Figure 3D), which is released when the

cells burst open upon swelling in contact with water

(Figure 3E).

Mucilaginous wings

Ursinia cakilefolia (Asteraceae)

Dry achenes of U. cakilefolia have a winged pappus

(wp) (Figure 4A), consisting of a single layer of radi-

ally elongated cells with helically thickened cell walls

(Figure 4B). When the pappus comes into contact with

water, the cells separate at the middle lamellae and the

helical thickenings uncoil to form long strands of muci-

lage (Figure 4C). The different steps through which the

pappus uncoils are clearly visible under the SEM (Fig-

ure 4D). The same process was observed in U. nana

subsp. nana (Rosch 1977).

Monoculus hyoseroides (Asteraceae)

Achenes of M. hyoseroides are three-winged. The

epidermis of the seed-bearing part of the achene is undu-

late and does not become mucilaginous (Figure 5A). The

section through the wings shows hexagonally shaped

cells completely filled with mucilage and separated by

thick cell walls (Figure 5B). Upon wetting, the cells

separate and the wings become completely mucilagi-

FIGURE 5. — Monoculus hyose-

roides, SEM of achenes: A,

entire dry achene; B, high

magnification of a section of

the wing; C, entire wet achene

with disintegrated wing; D,

high magnification of long

strands of mucilage.

nous (Figure 5C) and the long mucilage strands remain

attached to the achene (Figure 5D).

Mucilaginous hairs

Senecio arenarius (Asteraceae)

In the dry condition, the shiny white hairs (h) have

a finger-like shape and lie against the achene surface

(Figure 6A, B). The interior of the hair seems to be

filled with a coiled thread as seen at high magnification

(Figure 6B). In the wet condition, the hairs spread ± at

right angles to the seed. Apparently, when the mucilagi-

nous hairs absorb water the threads uncoil and become

thin and long (Figure 6C). The same phenomenon was

observed for Othonna cylindrica, as well as for O. flori-

bunda (Rosch 1977).

Acanthopsis horrida and Blepharis furcata (Acan-

thaceae)

In the dry condition, the hairs on the seed surface

of A. horrida (Figure 7 A, B) and B. furcata are tightly

appressed to the seed coat and are covered with a thick

deposit (Figure 1C). At high magnification it can be seen

that these hairs contain coiled spirals (Figure 7B). Once

in contact with water the thick deposit expands, the hairs

become swollen and spread ± at right angles to the seed

FIGURE 6. — Senecio arenarius, SEM of achenes: A, entire dry achene; B, high magnification of dry achene with hair lying against it; C, high

magnification of wet hairs, h, hair.

12

Bothalia 42,1 (2012)

FIGURE 7. — Acanthopsis horrida, SEM of seeds: A, entire dry seed; B, hairs; C, dry epidermis covered with thick deposit; D, wet seed; E, swollen

hairs; F, uncoiled strands of mucilage.

coat (Figure 7D). The tips of the hairs break open and

release a multitude of strands of mucilage (Figure 7E,

F).

Chemical composition

Three stains reacted positively with the mucilage of

all species: methylene blue, ruthenium red. and toluidine

blue; thus confirming the presence of polysaccharides

such as cellulose and pectic substances. On the basis

of the intensity of the reaction, the epidermal mucilage

seemed to contain less cellulose than the mucilaginous

wings and mucilaginous hairs. In some species, a slight

reaction was also observed with aniline blue, indicating

the presence of callose in the mucilage.

Cellulose is the most abundant plant polysaccharide

found in the form of microfibrils in cell walls and muci-

lage. Dry amorphous and fibrous cellulose can absorb

considerable amounts of water and becomes soft, flex-

ible, and viscous. Pectin is a heterogeneous grouping

of acidic structural polysaccharides with the main mol-

ecules being the D-galacturonic acid residues. They do

not possess exact structures and are complex. As for

cellulose, pectin is a water binder, is highly viscous,

and can form a firm gel when in contact with water. In

all cases, the mucilage of the diaspores investigated

contained both cellulose and pectic substances. This

could be due to the presence of microfibres of cellulose

embedded in the pectin compounds forming a complex

matrix. Both pectic substances and cellulose were also

reported in the seed mucilage of Arabidopsis thaliana

(Brassicaceae) (Windsor et al. 2000; Willats et al. 2001 ;

Macquet et al. 2007). In a detailed chemical and macro-

molecular study of the composition of A. thaliana seed

mucilage (Brassicaceae), Macquet et al. (2007) found

that the mucilage was made up of two layers: a water-

soluble layer that could be separated from the seed and

an inner layer that remained firmly attached to the seed.

The inner layer was itself constituted of two domains,

the internal one of which contained cellulose. The major

pectin of both the water-soluble and adherent seed muci-

lage was rhamnogalacturonan I.

This study has only provided a rough indication of

the composition of the mucilage of the diaspores investi-

gated. The chemistry of mucilage is complex and needs

a more detailed study to determine the exact type of

polysaccharide-forming mucilage in each diaspore.

CONCLUSIONS

The mucilaginous layer of the diaspores of 12 species

from Namaqualand was investigated anatomically using

both light and scanning electron microscopy. The struc-

tural origin of the mucilage produced by the diaspores

was diverse. Diaspores of myxospermic plant species

might produce mucilage in different ways; but in gen-

eral, this function resides in the epidermal cells and

other external appendages such as hairs and wings. Pre-

vious systems to classify the different types of mucilage

production did not recognize the mucilaginous nature of

wings or pappus. Despite the diversity of the origin and

structure of the mucilage, it performs more-or-less the

same ecological functions.

ACKNOWLEDGEMENT

The authors gratefully acknowledge the support

received from the German Federal Ministry of Educa-

tion and Research (BMBF) through the BIOTA South

Project and the National Research Foundation under

grant no. 61277.

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Bothalia 42,1 : 15-20(2012)

Recircumscription and distribution of elements of the 6 Ceterach

cordatum ’ complex ( Asplenium : Aspleniaceae) in southern Africa

R.R. KLOPPER* and N.R. CROUCH**

Keywords: Aspleniaceae, Asplenium , Ceterach , ferns, pteridophytes, southern Africa, xerophytes

ABSTRACT

Ceterachoid aspleniums in southern Africa have long been treated as a single widespread and variable taxon, Asplenium

cordatum (Thunb.) Sw. (= Ceterach cordatum Thunb.). In addition to A. cordatum, a further two ceterachoid taxa are now rec-

ognized as occurring in the Flora of southern Africa (FSA) region, namely A. capense (Kunze) Bir, Fraser-Jenk. & Lovis and

A. phillipsianum (Kummerle) Bir, Fraser-Jenk. & Lovis. We provide full descriptions and distributions of these three taxa.

INTRODUCTION

Pteridophyte treatments for the Flora of southern

Africa (FSA) (Roux 1986; Schelpe & Anthony 1986;

Burrows 1990) and Flora zambesiaca (FZ) (Schelpe

1970) regions, have considered a single xerophytic

rock fern, Ceterach cordatum Thunb., to occur widely

throughout the region and to show great morphological

variability. Ceterach is currently treated as a subgenus of

Asplenium L. (Crabbe et al. 1975; Bir et al. 1985; Roux

2001). Subgenus Ceterach is distinguished from subge-

nus Asplenium in our region by its lack of indusia and

the presence of densely-set scales (paleae) on the abax-

ial lamina surface, versus indusiate sori and sparsely-

set scales in subgenus Asplenium (Roux 2001). Moore

(1857) early recognized that the strictly African cete-

rachoid species were anatomically and morphologically

distinct from the Eurasian-Macronesian elements, and

therefore excluded them from Asplenium subgen. Ceter-

ach. More recent molecular analyses have demonstrated

the polyphyly of subgenus Ceterach , implicating homo-

plasy in the dense scale cover and in pinnatisect lami-

nae of these asplenioid ferns (Pinter et al. 2002; Van den

Heede et al. 2003). The taxonomic implications are that

southern African taxa referred earlier to subgenus Cete-

rach should either be accommodated in a new subge-

nus, or that distinction at a subgeneric level ( sensu Roux

2001) should be abandoned altogether (Van den Heede

et al. 2003).

The multiple origins in Asplenium of redundant

indusia and dense abaxial scales have been attributed

to independent adaptation to rocky xeroseres across its

range (Van den Heede et al. 2003). We further postulate

that the pinnatisect fronds common to both groups is a

poikilohydric modification that allows for frond integ-

rity to be maintained in the inrolled, dessicated state

(Figure 1), and for even restoration following rehydra-

tion. Ceterachoid taxa in southern Africa are extremely

* Biosystematics Research and Biodiversity Collections Division,

South African National Biodiversity Institute, Private Bag X 1 0 1 , 0001

Pretoria / Department of Plant Science, University of Pretoria, 0002

Pretoria. E-mail: r.klopper@sanbi.org.za.

** Ethnobotany Unit, South African National Biodiversity Insti-

tute, P.O. Box 52099, Berea Road, 4007 Durban / School

of Chemistry, University of KwaZulu-Natal, 4041 Durban.

E-mail: n.crouch@sanbi.org.za.

MS. received: 2010-05-20.

desiccation-tolerant; a member of this group has been

shown by Gaff (1977) to tolerate relative humidities in

the 0-5% range for at least six months, with an initial

water potential (\p) of 18%, expressed in terms of the rel-

ative humidity at 28°C.

We concur with Roux (2009a) in his reinstatement of

Asplenium capense (Kunze) Bir, Fraser-Jenk. & Lovis

as distinct from A. cordatum (Thunb. ) Sw., but we also

identify A. phillipsianum (Kummerle) Bir, Fraser-Jenk.

& Lovis as occurring in the FSA region (Table 1 ). This

taxon extends from the island of Socotra off Soma-

lia through East and Central Africa to the northern

provinces of South Africa, as far south as the northern

regions of Mpumalanga, Gauteng, and North West Prov-

ince. Pappe & Rawson (1858) recognized the wide-

spread European and North African taxon Asplenium

ceterach L. (= Ceterach officinarum Lam. & DC) as

South African, based on a Krebs collection made on the

Baviaans River. This species is not currently considered

to occur south of the Sahara, and has been taken as a

misidentification (Roux 1986, 2009a).

We provide the recircumscription and distribution of

the three ceterachoid elements of Asplenium currently

known from southern Africa.

FIGURE 1 . — Inrolled pinnatisect leaves of a desiccated plant of Asple-

nium cordatum , Nieu Bethesda, Eastern Cape Province. Photo:

N. Crouch.

16

Bothalia 42,1 (2012)

TABLE 1 . — Characters distinguishing ceterachoid members of Asplenium occurring in the FSA region.

MATERIALS AND METHODS

All ceterachoid Asplenium material held in the fol-

lowing herbaria (totalling almost 600 specimens) were

studied: Geo-Potts Herbarium (BLFU), University

of the Free State, Bloemfontein; Buffelskloof Nature

Reserve Herbarium (BNRH), Lydenburg; Bolus Her-

barium (BOL), University of Cape Town, Cape Town;

Selmar Schonland Herbarium (GRA), Albany Museum,

Grahamstown; the Herbarium at the Royal Botanic Gar-

dens, Kew (K), London, United Kingdom; Mpumalanga

Parks Board Herbarium (LYD), Lydenburg; Compton

Herbarium (NBG), SANBI, Cape Town; KwaZulu-Natal

Herbarium (NH), SANBI, Durban; National Museum

Herbarium (NMB), Bloemfontein; Bews Herbarium

(NU), University of KwaZulu-Natal, Pietermaritzburg;

National Herbarium (PRE), SANBI, Pretoria; H.G.W.J.

Schweickerdt Herbarium (PRU), University of Preto-

ria, Pretoria; A.P Goossens Herbarium (PUC), North

West University, Potchefstroom; and the South African

Museum Herbarium (SAM), SANBI, Cape Town.

The JSTOR Plant Science website (http://plants.jstor.

org) was consulted for type material held in other her-

baria. Images of type specimens were directly obtained

from The Museum of Evolution Herbarium (UPS), Upp-

sala University, Uppsala, Sweden. Types seen electroni-

cally are cited as e!

Herbarium acronyms follow Holmgren el ah (1990).

Author citations used follow the standardized author

abbreviations provided by the International Plant Names

Index (http://www.ipni.org).

TAXONOMY

Three taxa are recognized in southern Africa, all chas-

mophytic saxicoles (Jacobsen 1983):

Key to the species

la. Fronds always pinnate, with stronger, rigid rachis; pinnae

free from rachis (petiolulate), with flared to somewhat

auriculate/cordate base, abaxially very densely covered

with overlapping, broadly debate scales; sori small, dis-

crete 1 . A. cordatum

lb. Fronds pinnatisect to pinnate, with weaker, flexible rachis;

pinnae adnate to rachis, with broader decurrent base (at

least in distal half of frond), abaxially sparsely covered

with ovate lanceolate scales; sori large and confluent:

2a. Pinnae adnate to rachis and decurrent apically, becoming free

from rachis (petiolulate) with ± flared to auriculate/cordate

base basiscopically; rachis winged apically 2. A. capense

2b. Pinnae adnate to rachis and decurrent throughout; rachis

almost winged throughout 3. A. phillipsianum

1. Asplenium cordatum (Thunb.) Sw. in Journal

fur die Botanik 1800,2: 54 (1801). Acrostichum corda-

tum Thunb.: 171 (1800). Grammitis cordata (Thunb.)

Sw.: 23, 217 (1806). Cincinalis cordata (Thunb.) Desv.

(1811). Notholaena cordata (Thunb.) Desv.: 92 (1813).

Gymnogramma cordata (Thunb.) Schltdl.: 16 (1825).

Ceterach cordatum (Thunb.) Desv.: 223 (1827). Type:

South Africa, ‘e Cap bonae Spei’, Thunberg s.n. (UPS-

Thunb 24439, holo. el).

Ceterach crenata Kaulf.: 85, 86 (1824), nom. ille-

git. superfl. Type: as for Acrostichum cordatum Thunb.

[McNeill et al. (2006) Art. 7.5].

Gymnogramma namaquensis Pappe & Rawson:

42 (1858). Gymnogramma cordata var. namaquensis

(Pappe & Rawson) Sim: 212 ( 1892). Ceterach cordatum

var. namaquensis (Pappe & Rawson) Sim: 176 (1915).

Type: South Africa, Namaqualand, between rocks near

Modderfontein, 1856, Whitehead s.n. (K, holo.!).

Grammitis cordata [var. and] subvar. nudiuscula

Hook.: t.7 (1860), nom. illegit. superfl. Type: as for

Gymnogramma namaquensis Pappe & Rawson [see

Roux (2009a)].

Bothalia 42,1 (2012)

17

Grammitis cordata var. pinnato-pinnatifida Hook.: t.7

(1860), nom. inval. [McNeill et al. (2006) Art. 26.2],

Gymnogramma cordata var. subbipinnata Hook.: t.7

(1860). Ceterach cordatum var. subbipinnata (Hook.)

Kiimmerle: 289 (1909). Type: South Africa, ‘elevated

mountain of Macaliesberg’, Ecklon & Burke s.n. [miss-

ing, see Schelpe & Anthony (1986: 206) and Roux

(2009:83)].

Gymnogramma cordata var. bipinnata Sim: 212

(1892). Type: South Africa, Namaqualand, without pre-

cise locality, Holland s.n. [NBG, lecto.!, designated by

Schelpe & Anthony (1986: 206)].

Notholaena inaequalis Kunze y eckloniana (Kunze)

Kuntze var. rawsonii (Pappe) Kuntze forma minor

Kuntze: 379 (1898). Type: South Africa, [Eastern Cape],

‘Capland’, Cradock, 940 m, 12 Feb. 1894, Kuntze s.n.

[NY, lecto., designated by Roux (2009b: 228)].

Rhizome to 3-6 mm diam., erect or procumbent;

scales sessile, clathrate, very narrowly lanceolate, 2-3

x 0. 5-0.6 mm, acuminate, frequently with a hair point,

irregularly serrate, bicolorous, chestnut-brown distally,

brown proximally, with paler margin throughout, with

narrowly oblong to ovate cells. Fronds tufted, suberect

to erect; stipe ( 1 0—) 1 7 — 40( — 60 ) mm long, dark chest-

nut-brown, densely scaled when young, becoming sub-

glabrous with age, scales sessile, lanceolate, 3. 0-3. 5

x 0.7-0. 9 mm, acuminate, irregularly serrate, glossy,

sometimes bicolorous, chestnut-brown, sometimes with

narrow paler margin, with oblong to ovate cells; lamina

subcoriaceous, involute and inrolled when dry, elliptic

or narrowly elliptic to oblanceolate in outline, shallowly

to very deeply 2-pinnatifid, sometimes 2-pinnate, (20—

)50— 1 20(— 1 50) x ( 10—) 1 5— 40(— 50) mm, basal pinnae

gradually decrescent; rachis not winged between pinnae,

scales as for stipe but lanceolate, 3.0 x 0.9 mm; pinnae

(4 — )6— 1 9(— 3 0 ) x (2— )3— 7(— 1 3 ) mm, free from rachis

(petiolulate), with flared to ± auriculate-cordate base,

narrowly oblong to oblong-lanceolate, bluntly acute,

margin irregularly scalloped to incised, glabrous above

at maturity, abaxially very densely scaled, scales sessile,

debate, ± 2.0 x | .5 mm, attenuate, serrate, glossy, light

reddish brown, with short roundish to oval cells; sori

linear along (obscure) veins, up to 2 mm long, exindusi-

ate, almost totally obscured by scales. Figure 2A-D.

Etymology : cordatum = heart-shaped, referring to the

basally lobed pinnae.

Distribution and ecology: Asplenium cordatum

is widespread in South Africa, Lesotho, Swaziland,

Namibia, Botswana, and Zimbabwe (Figure 3); also in

Angola, Tanzania, Kenya, Uganda, and Ethiopia.

It occurs in rocky crevasses in exposed, hot, and dry

habitats, often at the base of boulders, well away from

water. A dense scale cover on the abaxial surface of the

fronds serves to protect the sori of this xerophytic spe-

cies.

Asplenium cordatum is very variable in size, with the

largest specimens originating from the dry areas of the

north-western parts of the Western and Northern Cape

provinces. This immense variability is reflected in the

long list of synonyms for this taxon. Burrows (1990)

noted that northwards of the Limpopo River it becomes

progressively rarer, reaching the extreme of its range in

Ethiopia.

Although purported to occur in Madagascar (Roux

2009a), where a ceterachoid element has been reported

from a single locality on Mount Morahahiva (Tardieu-

Blot 1957), this collection best matches Asplenium phil- ■

lipsianum (Tardieu-Blot 1958: Fig. XVII).

Vouchers: H.H. Burrows 3294 (GRA); E. Esterhuysen

25624 (BOL); D. Galpin 4782a (BLFU); H.H. W. Pear-

son 8557 (BOL, K, NBG); L.E. Taylor 2913 (NBG).

2. Asplenium capense (Kunze) Bir, Fraser-Jenk.

& Lovis in Fern Gazette 13,1: 61 (1985). Ceterach ca-

pense Kunze: 496 (1836). Grammitis capensis (Kunze)

T.Moore: Ixiii (1857). Type: South Africa, ‘Port Natal et

Afrique meridionalea’, Drege s.n. [G, lecto.!, isolecto.,

designated by Roux (1986: 352)]; ‘Ceded territory,

bergwaldungen an den Quellen des Katriver, Oberhalb

Philipstown’, Ecklon & Zeyher s.n. (UPS, syn. e!).

Gymnogramma capensis Spreng. ex Kaulf.: 183

(1831), nom. nud. Ceterach cordatum var. capense

(Spreng. ex Kaulf.) Hieron. ex Kiimmerle: 287 (1909).

Type: South Africa, ‘Cap. Bon spei: in einer Felsenritze

am Lowenberg’, Zeyher s.n. FI. Cap. No. 273 (HAL,

holo. e!; BOL, iso.!).

Grammitis cordata var. pinnata Hook.: t.7 (1860),

Type: as for Ceterach capense Kunze [see Roux

(2009a)].

Rhizome to 4 mm diam., erect or procumbent; scales

sessile, clathrate, lanceolate, 3-5 x (.0-1.5 mm, acu-

minate, frequently with a hair point, irregularly serrate,

bicolorous, with dark brown central region and paler

margins, with narrowly oblong cells. Fronds tufted,

erect to suberect; stipe 1 5— 35(— 60) mm long, chestnut-

brown to dark chestnut-brown, densely scaled, scales

sessile, narrowly triangular, 3-5 x 0.5-1 .0 mm, acu-

minate, irregularly finely serrate, glossy, sometimes

bicolorous, with narrow rust coloured central region

and broad straw coloured margin, with oblong to ovate

cells; lamina herbaceous, involute and inrolled when

dry, elliptic to narrowly obovate in outline, pinnatisect

to pinnate, sometimes shallowly 2-pinnatifid, (80 — )90 —

1 20(— 1 90) x (20-)30-45(-52) mm, basal pinnae gradu-

ally decrescent; rachis somewhat winged apically, not

winged between widely spaced pinnae basally, scales

as for stipe but 2-4 x 0. 7-1.0 mm; pinnae ( 1 0—) 1 5—

25(— 28) x (4— )5— 1 0(— 1 1 ) mm, adnate to rachis with

decurrent base apically, becoming free from rachis

(petiolulate) with somewhat flared to auriculate-cordate

base basiscopically, ovate-oblong, obtuse, margin sinu-

ate to scalloped, glabrous above at maturity, abaxially

very sparsely scaled, scales sessile, deflate to lanceo-

late, 1.5-2. 5 x 0.4-0. 7 mm, attenuate, finely serrate,

glossy, bicolorous, with narrow rust coloured central

region and broad straw coloured margin, with narrowly

oblong cells; sori linear along (obscure) veins, 3-6 mm

long, becoming confluent at maturity, exindusiate, not

obscured by scales. Figure 2E-H.

Bothalia 42,1 (2012)

FIGURE 2. — A-D, Asplenium cordatum, J.E. Burrows 1110 (BNRH): A, complete frond; B, cordate pinna base; C, abaxial surface of pinna with

dense scales; D, lamina scale. E-H, Asplenium capense, H.H. Burrows 2891 (BNRH): E, complete frond; F, pinna base; G, abaxial surface

of pinna with sparse scales; H, lamina scale. I -M, Asplenium phillipsianum, J.E. Burrows & S. M. Burrows s.n. (BN RH ): 1, complete frond;

J, adnate pinna base; K, abaxial surface of pinna with sparse scales; L, lamina scale; M, gemmae at frond apex. Scale bar: A, E, I, 20 mm;

B, C, F, G, J, K, M, 5 mm; D, H, L, 1 mm. Artist: Sandra Burrows.

Bothalia 42,1 (2012)

19

FIGURE 3. — Distribution of Asplenium cordatum in the FSA region.

Etymology : capense = pertaining to the Cape.

Distribution and ecology. Asplenium capense occurs

from the Cape Peninsula through the Western and East-

ern Cape, KwaZulu-Natal, the Free State and northern

provinces of South Africa (Figure 4), extending sporadi-

cally to central and tropical East Africa.

This species typically grows in sandy soil on forest

floors, as well as under coastal dune scrub, often fairly

close to streams and under trees on steep damp earth

banks. It is usually associated with riparian or open for-

est, most often in lightly rather than deeply shaded con-

ditions.

Vouchers: Th.C.E. Fries, T. Norlich & H. Weimark

30-8 (BOF, K); D.B. Miiller 890 (NMB; PRE); R. Sch-

lechter 2703 (GRA, K); E.M. van Zinderen Bakker 1135

(BFFU, PRE); C.J. Ward 12400 (NH, NU, PRE).

3. Asplenium phillipsianum (Kiimmerle) Bir, Fra-

ser-Jenk. & Lovis in Fern Gazette 13,1: 62 (1985).

FIGURE 4. — Distribution of Asplenium capense in the FSA region.

FIGURE 5. — Distribution of Asplenium phillipsianum in the FSA

region.

Ceterach phillipsianum Kiimmerle: 287 (1909). Type:

Somalia, ‘Ferns from deep shade Wagga Mountain, anno

1897’, Phillips s.n. (BM, holo.; K, iso.!)

Rhizome to 4 mm diam., erect or procumbent; scales

sessile, clathrate, lanceolate, 3-4 x 0. 6-1.0 mm, acu-

minate, frequently with a hair point, irregularly serrate,

sometimes bicolorous, with dark chestnut-brown central

region, sometimes with darker apex, with paler margins

throughout, with narrowly oblong to ovate cells. Fronds

tufted, suberect to spreading; stipe 5— 20(— 30) mm long,

dark chestnut-brown, densely scaled, scales sessile, lan-

ceolate, 3-4 x 0. 7-1.0 mm, acuminate, irregularly ser-

rate, glossy, bicolorous, distally dark brown and proxi-

mally chestnut-brown with paler margins throughout,

with narrowly oblong to ovate cells; lamina herbaceous,

involute and inrolled when dry, elliptic to obovate

in outline, pinnatisect to very shallowly 2-pinnatifid,

(5— )70— 1 00(— 130) x ( 1 6— )24— 33(— 50) mm, lower pinnae

gradually decrescent, occasionally produces 1-3 gem-

mae situated adaxially in the sinus of the distal pinnules;

rachis almost winged throughout, scales as for stipe

but 2. 5-3.0 x 0. 6-1.0 mm, concolorous reddish brown,

sometimes chestnut-brown towards apex; pinnae ( 6— ) 1 0—

20(-25) x (3-)4-6(-9) mm, adnate to rachis with decur-

rent base throughout, narrowly ovate-oblong, obtuse,

margin entire to weakly sinuate, glabrous above at matu-

rity, abaxially sparsely scaled, scales sessile, lanceolate,

1. 7-2.4 x 0.6-0. 8 mm, attenuate, finely serrate, glossy,

brown, with oblong cells; sori linear along (obscure)

veins, 2-3 mm long, becoming confluent at maturity,

exindusiate, not totally obscured by scales. Figure 2I-M.

Etymology r. phillipsianum = commemorates the Eng-

lish explorer Ethelbert Fort-Phillips ( 1 857-1944) who in

1897 collected the type specimen in Somalia.

Distribution and ecology >: Asplenium phillipsianum

occurs from the northern provinces of South Africa

(Limpopo, North West, Gauteng, and Mpumalanga)

(Figure 5), through central and tropical Africa as far

north as Socotra; also known from Reunion and Mada-

gascar.

20

Bothalia 42,1 (2012)

This species is typically found close to streams,

shaded under trees on steep damp earth banks.

Of the three regional ceterachoid species, A. phillip-

sianum is the only member noted to be gemmiferous (J.

Nel, pers. comm.) (Figure 2M).

Vouchers: J.E. Burrows & S.M. Burrows s.n.

(BNRH); M.F. Glen PRE62127 (PRE); E. Relief & S.E.

Strauss 2154 (PRE); J.P. Roux 3195 (NBG); A. Winter-

boer s.n. (PRU).

Excluded name

Ceterach cordatum var. pinnatifida Sim: 177 (1915).

The syntypes cited by Sim (1915) include representa-

tives of both A. capense and A. phillipsianum.

ACKNOWLEDGEMENTS

Dr Hugh Glen, KwaZulu-Natal Herbarium, SANBI,

Durban, is thanked for translating Latin texts and for

providing nomenclatural advice; Prof. John McNeill,

Royal Botanic Gardens, Edinburgh, for providing

nomenclatural advice; Ms Hester Steyn, National Her-

barium, SANBI, Pretoria, for producing updated dis-

tribution maps. The Curators of BLFU, BNRH, BOL,

GRA, K, LYD, NBG, NH, NMB, NU, PRE, PRU, PUC

and SAM kindly facilitated access to their collections.

REFERENCES

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HAWKSWORTH, D.L., MARHOLD, K„ NICOLSON, D.H.,

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Bothalia 42,1: 21—41 (2012)

A revision of Tecophilaeaceae subfam. Tecophilaeoideae in Africa

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

Keywords: Africa, Cyanella Royen ex L., Eremiolirion J.C. Manning & F.Forest, new species, systematics, Tecophilaeaceae, Walleria J.Kirk

ABSTRACT

Family Tecophilaeaceae subfam. Tecophilaeoideae is revised for the Flora of southern Africa region, with the inclu-

sion of the tropical Walleria mackenzii J.Kirk for completeness. The genera Cyanella Royen ex L. (9 spp.), Eremiolirion

J.C. Manning & F.Forest (1 sp.) and Walleria J.Kirk (3 spp.) are treated, with keys to the genera, species and subspecies; and

full descriptions and distribution maps. A formal infrageneric classification is proposed for Cyanella , in which sect. Trigella

(Salisb.) Pax & K.Hoffm. is revived for the species with a 3 + 3 arrangement of stamens. The new species, C. marlothii

J.C. Manning & Goldblatt, is described from the Richtersveld; and C. pentheri Zahlbr. is resuscitated from the synonomy

of C. hyacinthioides Royen ex L. Pink-flowered plants of normally yellow-flowered C. lutea have a separate geographical

distribution and are recognized as subsp. rosea (Eckl. ex Baker) J.C. Manning & Goldblatt.

INTRODUCTION

Tecophilaeaceae is a small family of seven or eight

genera and ± 25 species from California, Chile, and

southern and tropical mainland Africa (Simpson &

Rudall 1998). The reported occurrence of the fam-

ily in Madagascar (Simpson & Rudall 1998) is based

on Walleria paniculata Fritsch, a synonym of Dianella

ensifolia (L.) DC. (Hemerocallidaceae). The family

is best represented in Africa, where almost two thirds

of the species are found. Cyanastrum Oliv. (3 spp.)

and Kabuyea Brummitt (1 sp.) are strictly tropical, but

Walleria J.Kirk (3 spp.), Eremiolirion J.C. Manning

& F.Forest (1 sp.), and Cyanella Royen ex L. (9 spp.),

are primarily distributed in subtropical and temper-

ate southern Africa. Members of the family are peren-

nial herbs with a cormous, usually tunicated rootstock,

basal (rarely cauline) leaves, and long-lasting flowers,

typically in racemose or paniculate, cymose inflores-

cences; but sometimes solitary and axillary. The flowers

are actinomorphic or zygomorphic, with 3 + 3 petaloid

tepals fused into a short tube adnate to the ovary, and

six stamens, all fertile or some reduced to staminodes,

with ± porose dehiscence. The ovary is inferior or semi-

inferior and 3-carpellate, and matures into a loculicidal

capsule (Simpson & Rudall 1998; Heywood et al. 2007).

The two tropical African genera, Cynastrum and

Kabuyea , have been the subject of a detailed review

(Brummitt et al. 1998), in which they were segregated

as subfam. Cynastroideae, with the remaining genera of

the family retained in subfam. Tecophilaeoideae. The

taxonomy of the southern African species is relatively

well understood, and both Cyanella and Walleria were

revised fairly recently (Carter 1962; Scott 1991; Cowley

& Brummitt 2001), including historical and morpho-

logical details. Since then, however, the genus Eremio-

lirion has been established to accommodate Cyanella

amboensis Schinz, which was excluded from Cyanella

* 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: 2011-02-01.

by Scott (1991), but unplaced. We have also published

additional observations on the distribution and morphol-

ogy of Walleria gracilis (Salisb.) S. Carter (Manning et

al. 2001). It is now clear that there is more variation in

some species of Cyanella than was recognized by Scott

(1991), and three subspecies have since been described

in C. alba L.f. (Manning et al. 2005). Field study and

examination of herbarium material of C. hyacinthoides

Royen ex L. suggest that this species is currently too

broadly circumscribed, and that C. pentheri Zahlbr.

should be resuscitated from synonymy. In addition, the

clear geographical segregation between the typical yel-

low-flowered and the pink-flowered forms of C. lutea

L.f. is appropriately reflected by the recognition of dis-

tinct subspecies for them. A collection from the Rich-

tersveld, until now identified as C. orchidiformis Jacq.,

differs from that species and from all others in the genus

in having all six filaments connate into a staminal tube.

It evidently represents an unnamed species that we

describe here.

Currently, therefore, there is no comprehensive treat-

ment for the family in southern Africa and the available

treatment of Cyanella is inadequate and incomplete in

some respects. We provide here a complete review of the

genera and species occurring in the Flora of southern

Africa region, including also the tropical African Walle-

ria mackenzii J.Kirk for completeness. We also propose

a new infrageneric classification for Cyanella that asso-

ciates morphologically similar species in two sections,

with the larger of the two, sect. Cyanella , subdivided

into two series.

MATERIALS AND METHODS

Type specimens or digital images of types from the

relevant herbaria were examined for all names, as well

as all available herbarium specimens in BOL, NBG,

PRE, and SAM (herbarium acronyms after Holmgren

et al. 1990). Particular use was made of high-resolution

digital images on the Aluka website (www.aluka.org),

and of the Herbarium of the Linnean Society of London

(www.linnean-online.org).

22

Bothalia 42,1 (2012)

TAXONOMY

Key to genera

la Corm not tunicated; leaves cauline; flowers solitary in leaf

axils; seeds verrucose or papillate, with tufts of trichomes,

brown Walleria

lb Corm with fibrous tunics; leaves basal; flowers in racemose

or paniculate cymes; seeds rugose, glabrous, black:

2a Foliage leaves 2; inflorescence a divaricate panicle; pedi-

cels without a bracteole; flowers actinomorphic; stamens

monomorphic, central, and symmetrical Eremiolirion

2b Foliage leaves 3-12; inflorescence a raceme, usually

branched, rarely condensed and flowers apparently soli-

tary; pedicels bracteolate; flowers zygomorphic; stamens

dimorphic, in two groups of 3 + 3 or 5 + 1 Cyanella

Walleria J.Kirk in Transactions of the Linnean Soci-

ety 24: 497 (1864). Type species: Walleria nutans J.Kirk

[lecto., designated by E.P.Phillips: 207 (1951)].

Androsyne Salisb: 61 ( 1866). Type species: A. gracilis

Salisb. = Walleria gracilis (Salisb.) S. Carter.

Deciduous geophytes with deep-seated, non-tuni-

cated corm; subterranean portion of stem developing

paired adventitious roots at each node, aerial portion of

stem erect or straggling, smooth, scabrid or armed with

recurved prickles. Cataphylls numerous, scattered along

subterranean portion of stem, small, tubular, membra-

nous. Foliage leaves numerous, all cauline, alternate,

sessile or amplexicaul, linear to ovate, acute or cir-

rhose and tendrilliferous, midrib sometimes armed with

recurved prickles beneath. Inflorescence of solitary, axil-

lary flowers, or rarely bracteole subtending a second

flower; pedicels erect or cernuous, smooth or prickly,

with solitary bracteole inserted ± midway. Flowers

actinomorphic, erect or nodding, rotate, white to blue;

tepals connate below into short tube, ± similar. Stamens

6, monomorphic, erect-symmetrical, inserted at mouth

of tube; filaments short; anthers basifixed, erect, free,

or connivent around style, narrowly lanceolate, dehisc-

ing by apical pores, outer surface scabridulous in basal

± 1/2. Ovary ± superior, with several ovules per locule;

style terete, erect, filiform. Capsules ovoid to subglo-

bose. Seeds ovoid, brown, surface warty or produced

into finger-like papillae, each with apical tuft of minute

trichomes. Basic chromosome number', x = 12 (Goldblatt

& Manning 1989).

3 spp., southern and southern tropical Africa.

Etymology', the genus is named for Horace Waller,

who made the first collections of both tropical African

species during an expedition to central Africa in 1863.

Ethnobotany. the corms comprise part of the tradi-

tional diet of the San, Tswana, and other indigenous

tribes ( e.g . Leffers 2008; also Lugard 289, Maguire

2194, Snyman & Noailles 231, Story 6117).

Key to species

la Flowers erect, tepals 13-22 mm long; anthers free, not con-

nivent, blue, purple, or black with yellow base and apex,

6-12 mm long; style 8—15 mm long 1 . W. mackenzii

lb Flowers nodding, tepals 6-16 mm long; anthers connate and

connivent, yellow at least in basal 1/2, 4-8 mm long; style

4. 0-8. 5 mm long:

2a Plants erect or sprawling, free-standing, mostly

unbranched; stems and pedicels smooth, scabrid or with

hooked prickles; leaves not cirrhose; tepals plain white,

pink, mauve, or blue 2. W. nutans

2b Plants usually straggling or climbing, well branched; stems

and pedicels always armed with hooked prickles; upper

leaves cirrhose, with tendril-like apex; tepals white with

basal purple blotch 3. W gracilis

I . Walleria mackenzii J.Kirk in Transactions of

the Linnean Society 24: 497, t. 52/2 (1864). Type:

Nyasaland [Malawi], Manganja Hills, near Bishop

Mackenzies Mission, 1863, H. Waller sub J. Kirk s.n.

K256015 (K, holo.!). Illustration: Cowley & Brummitt

(2001).

W. angolensis Baker: 262 (1878). Type: Angola,

Huilla, 18 Dec. 1859, Welwitsch 1749 (BM, holo.!; K,

iso.!).

Deciduous geophyte, 180-900 mm high. Corm sub-

globose or depressed-globose, 20-40 mm diam. Stem

erect, mostly simple or with 1 or 2 branches, smooth

or rarely scabrid or minutely prickly. Leaves ovate to

narrowly lanceolate, 30-110 x (4— )5— 20(— 28) mm,

upper narrower, base cuneate or weakly cordate but

not amplexicaul, apex acute or rarely cirrhose, midrib

smooth, sometimes scabrid or minutely prickly. Flow-

ers solitary in axils in central portion of stem, erect,

sometimes with additional flower developed in axil of

bracteole; pedicels ascending and ± erect at flowering,

straight or flexible, becoming deflexed or pendulous

in fruit, 13-60 mm long, smooth or scabrid, with lan-

ceolate bracteole 10-26 mm long inserted ± halfway,

rarely lacking; tepals white, pink, or mauve to pale

or bright blue, spreading, elliptic-lanceolate, 13-22 x

2. 5-6. 5 mm, inner slightly narrower than outer. Stamens

erect, free and not connivent; filaments 1-3 mm long,

awl-shaped; anthers 6-12 mm long, blue to purple or

black with yellow base and apex, pores circular, apical.

Ovary subglobose-pyramidal, 3-lobed above, ± 3 mm

long; style 8-15 mm long. Capsule subglobose or ovoid,

10-20 mm long, maturing to dark yellow. Seeds ovoid,

± 5 mm long, dark mahogany-brown, papillate, papillae

becoming longer and more finger-like in distal half, each

with apical tuft of minute trichomes. Flowering time :

mainly Nov. -Jan. (-Mar.), shortly after the onset of the

rains.

Distribution and ecology: distributed across southern

tropical Africa, from the higher-lying parts of central

Angola, Zambia, and southern Democratic Republic of

Congo, through Malawi into southern and western Tan-

zania [see Carter (1962) for map]. The species is largely

restricted to higher rainfall areas, where it occurs in

open woodland and savanna, often in rocky outcrops.

Diagnosis and relationships : distinguished from other

species of Walleria by its generally more robust habit,

erect, mostly larger flowers with tepals 13-22 mm long,

and free anthers not cohering at the tips, predominantly

blue to purple or black with only the base and tips yel-

low, and dehiscing through terminal, circular pores.

Walleria mackenzii is likely to be confused only with

W. nutans , which has nodding flowers with tepals 6-16

mm long and connivent anthers, connate at the tips, and

dehiscing through short, subapical, introrse slits.

Bothalia 42,1 (2012)

23

2. Walleria nutans J.Kirk in Transactions of the Lin-

nean Society 24: 497, t. 52/1 (1864). W. mackenzii var.

nutans (J.Kirk) Baker: 498 (1879). Type: Nyasaland

[Malawi], Manganja Hills, near Bishop Mackenzies

Mission, 1863, H. Waller sub J. Kirk s.n. K256018 (K,

holo.!). Illustration: Dyer: 1321 (1960).

W. muricata N.E.Br.: 145 (1909). Type: Bechuana-

land [Botswana], near Palapye, Jan. 1 898, Lugard 289

(K, holo.!).

W. baumii Dammer: 361 (1912). Types: Angola,

Kunene-Kubangoland, Kalolo, 22 Nov. 1899, Baum 448

(BM, syn.); Angola, Habungo, 28 Nov. 1899, Baum 448

(BM, syn.).

W. hockii De Wild.: 8 (1915). Type: Northern Rho-

desia [Zambia], Kafue Valley, 1911, A. Hock s.n.

BR8642639 (BR, holo.!).

Deciduous geophyte (70-) 100-300 mm high. Corm

subglobose or depressed-globose, 20-30 mm diam. Stem

erect or sprawling but never climbing, mostly simple or

with 1 or 2 branches, rarely more, smooth or variously

prickly with delicate, recurved prickles 0.5-1 .5 mm

long. Leaves linear to narrowly lanceolate, (30— )70— 1 50

x ( 2— )5— 7(— 12) mm, upper narrower and attenuate, base

cuneate or weakly cordate but not amplexicaul, midrib

smooth or with recurved prickles beneath. Flowers soli-

tary in axils in central portion of stem, nodding, some-

times with additional flower developed in axil of bracte-

ole; pedicels suberect but sharply decurved distally,

20-50(-80) mm long, smooth or scabrid, with lanceolate

bracteole 10-15 mm long inserted in upper third or quar-

ter; tepals white, pink, or mauve to pale blue, recurved

or reflexed, lanceolate, (6-) 10-1 6 x 2-5 mm. Stamens

connivent, connate at tips; filaments 0. 5-1.0 mm long;

anthers (4— )6— 8 mm long, mostly yellow with narrow

purple band across distal third and with grey tips, slits

short, subapical, introrse. Ovary subglobose-pyramidal,

3-lobed above, ± 3 mm long; style 5. 0-8. 5 mm long.

Capsule ovoid, shortly apiculate and 3-lobed above,

8-17 mm long, green, yellow or orange. Seeds ovoid,

± 5 mm long, dark mahogany-brown, papillate, papil-

lae becoming longer and more finger-like in distal half,

each with apical tuff of minute trichomes. Chromosome

number. 2n = 12 (Goldblatt & Manning 1989). Flower-

ing time: Nov.-Jan.(-Mar.). Figure 1A, B.

Distribution and ecology, widely distributed through

subtropical Africa, from the higher-lying parts of cen-

tral and northern Namibia and southern Angola through

Zambia into eastern Botswana and the northern part

of South Africa, where it has been recorded from the

Soutpansberg into central Limpopo, adjacent Mpuma-

langa and North West Province, and southwest as far as

Taung in Northern Cape (Figure 2). Plants occur in open

savanna, mostly in sandy soils but also on limestone

flats and dolomite rock sheets.

Diagnosis and relationships', closely resembling the

southwestern Cape W. gracilis, with which it shares

nodding flowers with apically connivent anthers dehisc-

ing through introrse, subapical pores and sometimes

prickly stems, pedicels, and abaxial leaf midribs. Walle-

ria nutans is distinguished by its free-standing, mostly

FIGURE 1. — A, B, Walleria nutans : A, flower; B, detached anther.

C-K, W. gracilis'. C, flowering plant; D, flower; E, outer tepal;

F, inner tepal; G, half-flower; H, androecium with style; 1, gyn-

oecium; J, capsule; K, seed. Scale bar: A, C-F, J, 10 mm; B, G-I,

2 mm. Artist: John Manning.

24

Bothalia 42,1 (2012)

FIGURE 2. — Distribution of Walleria nutans.

unbranched stems, leaves without tendril-like tips, and

unmarked, white, pink, or mauve to pale blue tepals.

The stems, pedicels and underside of the leaf mid-

ribs may be smooth or variously armed with recurved

prickles, but these are delicate, almost bristle-like, and

mostly < 1 mm long, and the anthers are mostly yellow,

with the purple and grey banding restricted to the apical

third. The presentation of the flowers is subtly different

in the two species: pedicels in W. nutans are essentially

suberect up to the level of insertion of the bracteole in

the upper third or quarter, at which point the pedicels are

sharply decurved, whereas the bracteoles in W. gracilis

are mostly inserted ± midway along the pedicels, which

are therefore more arcuate.

Vernacular name: bush potato.

Representative specimens

NAMIBIA. — 1723 (Singalamwe): Singalamwe, (-CB), 23 Nov.

1973, Pienaar & Vahnneijer 209 (PRE). 1820 (Tarikora): Gautscha

Pan, E of Karakuwise, (-DD), 27 Dec. 1952, Maguire 2194 (NBG);

Cigarette, NE of Karakuwise, (-DD), 19 Jan. 1953 (fruiting), Maguire

2275 (NBG). 1914 (Kamanjab): Ombutu, (-BC), 25 Feb. 1969, Grob-

belaar 85 (PRE). 1917 (Tsumeb): Tsumeb, (-BA), Dec. 1935, Boss

35483 (PRE). 1920 (Tsumkvve): 157 miles [250 km] E of Grootfon-

tein, Simkue, (-DA), 14 Jan. 1958, Story 6117 (PRE). 2016 (Otji-

warongo): Farm Uitsig, 60 km E-NE of Otjiwarongo, (-BC), 5 Mar.

1984 (ex hort.), Lavranos 21034 (NBG). 2017 (Waterberg): Water-

berg, Farm Okamuru, (-CA), 5 Mar. 1974, Merxmiiller & Giess 30063

(PRE). 2118 (Steinhausen): 15 km along Kapps Farm road from Stein-

hausen to Windhoek, (-CC), 15 Mar. 1988 (fruiting), Gotdblatt &

Manning 8802 (MO, PRE). 2215 (Trekkopje): Aukas, (-AA), 28 Nov.

1980, Dinter 654 (SAM); Farm Neuschwaben, Undasbank, (-DB), 8

Mar. 1953 (fruiting), Kinges 3061 (PRE). 2217 (Windhoek): Wind-

hoek, Farm Lichtenstein, (-CD), 20 Jan. 1923, Dinter 4310 (SAM).

BOTSWANA.— 2225 (Mokatini): N of Lephephe, 100 km W of

Serowe, (-BC), Feb. 1982 (fruiting), Snyman & Noailles 231 (PRE).

2426 (Mochudi): Mochudi, (-AC), without date, Rogers 6739 (BOL).

LIMPOPO. — 2229 (Waterpoort): Soutpansberg, Wylies Poort,

Ingwe Farm, (-DD), 18 Dec. 1960, Hardy 407 (PRE). 2329 (Peters-

burg) [Polokwane]: Buffelsberg near Munnik, (-DB), Dec. 1932,

Schyveickerdt 1036 (PRE); Broederstroom, (-DD), 19 Nov. 1949,

Prosser 1361 (NBG). 2428 (Nylstroom): Vaalwater Poort on Nyl-

stroom road, (-AC), 16 Dec. 1960, Hardy & Bayliss 421 (PRE).

NORTH WEST. — 2526 (Zeerust): Lichtenburg, Grasfontein,

(-CC), Dec. 1929, Sutton 338 (PRE). 2527 (Rustenburg): Broeder-

stroom, (-DD), 19 Nov. 1949, Prosser 1361 (PRE).

MPUMALANGA. — 2430 (Pilgrim’s Rest): Nooitgedacht mtn, near

Branddraai, (-DA), 24 Nov. 1933, Young A688 (BOL," PRE).

NORTHERN CAPE. — -2724 (Taung): Barkly West, Madipelessa,

(-CA), 26 Feb. 1937, Acocks 1822 (PRE).

3. Walleria gracilis (Salisb.) S. Carter in Kew Bul-

letin 16: 189 (1962). Androsyne gracilis Salisb.: 61

(1866). Type: stated as from Nicobar Islands but prob-

ably from South Africa, Western Cape, comm. William

Marsden [BM, holo.l; drawing in Salisbury mss. 8: 818

(BM)]. Illustration: Manning et al.AA-Al (2001).

W. armata Schltr. & K. Krause in Krause: 235 (1921).

Type: South Africa, [Western Cape, near Klawer], [Farm]

Windhoek, 8 July 1 896, R. Schlechter 8074 (B, holo. [not

seen]; BM!, BR!, COI!, GRA!, K, MO!, PRE!, S!, iso.).

[The collection was published as Schlechter 2074 in the

protologue, evidently a misprint].

Deciduous geophyte, 100-700 mm high. Corm sub-

globose or depressed-globose, 20-30 mm diam. Stem

straggling or climbing, well branched, with recurved

prickles 1.0-1. 5 mm long in upper parts. Leaves lanceo-

late to narrowly lanceolate, (30— )70— 1 20 x 5-10 mm,

upper narrower and attenuate-cirrhose, apex coiling and

tendril-like, amplexicaul, midrib with recurved prick-

les beneath. Flowers solitary in axils in central por-

tion of stem, nodding, rose-scented; pedicels arcuate,

20-40 mm long, sparsely prickly, with lanceolate bracte-

ole 6-10 mm long inserted ± midway; tepals white with

purple blotch at base, recurved or reflexed, lanceolate,

10-16 x 2. 5-3. 5 mm. Stamens connivent, connate at

tips; filaments 0. 5-1.0 mm long; anthers 5-6 mm long,

yellow in lower 1/2 and purple above with grey tips, slits

short, subapical, introrse. Ovaty subglobose-pyramidal,

3-lobed above, ± 2 mm long; style ± 4 mm long. Cap-

sule ovoid, ± 15 mm long, shortly apiculate and 3-lobed

above. Seeds ovoid, ± 5 mm long, dark mahogany-

brown, with conspicuous apical cluster of finger-like

papillae, each topped with tuft of minute trichomes, rest

of seed ± smooth but covered with trichome-tufts. Flow-

ering time : June and July. Figure 1C-K.

Distribution and ecology: the species has a limited,

curiously scattered distribution along the west coast of

South Africa. It is best known from the lower reaches of

the Olifants River in Western Cape, where it has been

recorded along the foot of the Gifberg east of Klawer

and on Pakhuis Pass, some 50 km to the south (Figure

3). At these localities, the species occurs in deep sand

among outcrops of Cape sandstone in arid fynbos vege-

tation. There is evidently a large disjunction in the distri-

bution, based on a single enigmatic collection made by

Rudolph Marloth in 1925 from near Kuboes in the Rich-

tersveld. This locality, 60 km upstream from the mouth

of the Orange River, is 350 km north of Klawer, and to

date W. gracilis has not been re-collected there; nor from

the intervening country. Although the identity of the

Kuboes collection is not in doubt, it is unfortunately a

plant that was cultivated to flowering in Cape Town five

years later; and although the label is explicit in identi-

fying the location at which the tuber was originally col-

lected, the possibility that the locality has been confused

must be considered until the species is rediscovered in

the Richtersveld.

Plants may reach up to 600 mm in height when sup-

ported by small shrubs, but are much shorter in the open.

The nodding, Solanum- like flowers are evidently adap-

tated to buzz pollination, probably by solitary bees in the

family Apidae: Anthophorinae (Manning et at. 2001 ).

Bothalia 42,1 (2012)

25

FIGURE 3. — Distribution of WaUeria gracilis.

Diagnosis and relationships : the species closely

resembles W nutans from subtropical Africa and was

treated as conspecific with it by Phillips (1951), but the

two are quite distinct. WaUeria gracilis is recognized

by its straggling or climbing habit, well-branched stem,

more robust prickles ± 1.0-1. 5 mm long, upper leaves

drawn into coiled, tendril-like tips, distinctive white

flowers marked with a large purple blotch at the base of

each tepal, and anthers that are yellow only in the lower

half. The seeds of W. gracilis are also distinctive in

being essentially smooth in the basal half (apart from the

trichome-tufts) with a dense apical cluster of finger-like

papillae. Flowering in W. gracilis takes place during the

winter, whereas W. nutans blooms in summer.

Additional specimens seen

NORTHERN CAPE.— 2817 (Vioolsdrif): Kubus [Kuboes] main

kloof, 29 Aug. 1925 [fl. in cult. June 1930], Marloth 12358 (PRE).

WESTERN CAPE.— 3118 (Vanrhynsdorp): Klawer, Farm Wind-

hoek, NW foothills of Gifberg, (-DA), mid-July 1998, Manning 2180

(NBG), 25 June 2005, Manning 295 IB (NBG), Forest & Manning 542

(NBG). 3219 (Wuppertal): Clanwilliam, Cedarberg [Pakhuisberge],

Farm Alpha, (-AA), 20 July 1941, Bond 7053 (BOL, NBG).

Eremiolirion J.C. Manning & F.Forest in Bothalia

35: 117 (2005). Type species: Eremiolirion amboense

(Schinz) J.C. Manning & C.A.Mannheimer.

Deciduous geophyte with deep-seated, tunicated

corm, tunics decaying into firm-leathery, coarsely net-

ted fibres extending into neck. Cataphyll 1, extend-

ing to ground level and enclosing leaf sheaths. Foliage

leaves 2, basal, narrowly lanceolate-canaliculate, leath-

ery. Inflorescence a divaricately branching, paniculate

cyme with bracts subtending branches and pedicels only;

pedicels ebracteolate, cemuous at tip, elongating slightly

in fruit and straightening. Flowers actinomorphic, nod-

ding, campanulate, white flushed pink or maroon abaxi-

ally; tepals connate below into short tube with minute,

fringed corona present at mouth of tube, dimorphic,

outer oblong, inner pandurate. Stamens 6, monomor-

phic, erect-symmetrical, inserted near mouth of tube;

filaments short; anthers basifixed, erect and connivent

around style, narrowly lanceolate, dehiscing by oblong

apical pores. Ovary half inferior, with several ovules per

locule; style terete, erect, filiform. Capsules ovoid to

globose. Seeds ellipsoid-pyriform, blackish brown, testa

surface rugose.

1 sp., central and northwest Namibia, southwest

Angola.

Etymology, the name is a compound of the Greek ere-

mios (desert or wilderness) and lirion (lily).

Ethnology, the corms are part of the traditional diet of

the local tribes ( Giess , Volk & Bleissner 6039).

Eremiolirion amboense (Schinz) J.C. Manning &

C.A.Mannheimer in Bothalia 35: 117 (2005). Cyanella

amboensis Schinz: 943 (1902). Type: South West

Africa [Namibia], Amboland [Ovamboland], Ondonga,

[Ondongwa], without date, Rautanen 344 (Z, holo. ! ).

Plants (60—) 1 00—250 mm high. Corms deep-seated,

30 mm diam; tunics decaying into firm-leathery,

coarsely netted fibres extending into neck 10-60 mm

long, pale whitish brown. Leaves 2, basal, suberect, nar-

rowly lanceolate, (10-) 15-25 x (8-) 10-20 mm, attenu-

ate, canaliculate with prominent midrib abaxially, leath-

ery. Inflorescence a divaricately branching, paniculate

cyme with ( l-)3-7-branches, up to 30-flowered; pedi-

cels cemuous at tip, 15-25 mm long, elongating slightly

in fruit and straightening, ultimately 20^10 mm long.

Flowers nodding, campanulate, white flushed pink or

maroon abaxially at base of outer tepals, fragrant; peri-

anth tube ± 4 mm long, with fringed corona 0.5-1 .0 mm

high at mouth of tube forming collar extending over

ovary to surround base of style; outer tepals spreading

from base, oblong, 15-20 x 5-7 mm, obtuse, margins

revolute, inner tepals at first suberect but spreading in

upper 1/2, pandurate and short-clawed, claw ± 2 mm

long, blade ovate, 13-18 x 7-10 mm, apex slightly cuc-

ullate, margins crisped. Stamens monomorphic; fila-

ments terete, ± 0.25 mm; anthers narrowly lanceolate,

9-10 mm long, yellow, dehiscing by oblong apical pores

1.5 mm long. Ovary half-inferior; ovules ± 6 per locule;

style 10-12 mm long, extending shortly beyond anthers,

white. Capsules ovoid to globose, 10-12 x 8-12 mm.

Seeds ellipsoid-pyriform, 4. 0-4. 5 x 3. 0-3. 5 mm, black-

ish brown; testa surface rugose. Flowering time : (mid-

Jan.-)Feb.-Mar.(-early Apr.). Figure 4.

Distribution and ecology, locally common through

the higher-lying parts of west-central and northwestern

Namibia, occurring along the better watered, western edge

of the escarpment from west of Mariental in the south

to Kaokoland in the north (Figure 5) and in southwest-

ern Angola near Lake Arco. The species typically occurs

in colonies, often numbering many individuals, in sandy

loam or heavy clay soils, especially in stony or gravelly

situations. Flowering is dependent on rainfall.

The flowers close at night ± 21:00, re-opening in

the morning ± 09:00. They are fragrant during the day,

with a jasmine-like fragrance at first but later smelling

of stale urine, and are visited by bees and the occasional

moth (Ward, Ward & Ward 10518).

Vernacular name : desert snowdrop.

Representative specimens

ANGOLA. — Namibe Prov., Lake Arco, Jan. 2009 (fl. ex cult. Mar.

2012), Harrower 4061 (NBG).

26

Bothalia 42,1 (2012)

FIGURE 4. — Eremiolirion amboense: A, flowering plant; B, flower; C, inner tepal; D, outer tepal; E, half-flower; F, androecium with style; G,

capsule; H, seed. Scale bar: A-D, G, 10 mm; E, F & H, 2 mm. Artist: John Manning.

NAMIBIA. — 1713 (Swartbooisdril): West of Ombazu, (-DD), 9

Apr. 1973, Giess & Van der Walt 12658 (WIND). 1913 (Sesfontein):

Kunene, Barab River, (-DB), 23 Mar. 2001, Burke 1020 (WIND).

1914 (Kamanjab): Etendeka Mountain Camp, (-DD), 28 Feb. 2004,

Mannheimer 2510 (NBG, WIND). 1915 (Okaukuejo): Etosha,

Adamax, (-BB), 16 Jan. 1974, Le Roux 597 (PRE, WIND). 2014

Bothalia 42,1 (2012)

27

(Khorixas): S side of watershed Ugab/Huab Rivers W of Brandberg,

(-CA), 10 Apr. 1989, Ward, Ward & Ward 10518 (PRE, WIND). 2114

(Uis): Omaruru, (-BA), 20 Mar. 1967, Giess 9708 (PRE, WIND).

2315 (Rostock): Swakopmund, W of Kuiseb Canyon, (-BD), 10 Feb.

1966, Giess 9131 (PRE, WIND); Farm Greylingshof SW 107, (-BD),

16 Feb. 1963, Giess, Volk & Bleissner 5158 (PRE, WIND).

Cyanella Royen ex L ., Genera plantarum, edn. 5: 149

(1754). Type species: Cyanella hyacinthoides Royen ex

L.

Pharetrella Salisb.: 47 (1866). Cyanella sect. Phare-

trella (Salisb.) Pax & K.Hoffm.: 427 (1930). Type spe-

cies: P. alba (L.f.) Salisb. = Cyanella alba L.f.

Trigella Salisb.: 46 (1866). Cyanella sect. Trigella

(Salisb.) Pax & K.Hoffm.: 427 (1930). Type species:

T. orchidiformis (Jacq.) Salisb. = Cyanella orchidiformis

Jacq.

Note\ Pax & Hoffmann (1930) inadvertently trans-

posed the species and diagnoses of their sections Phare-

trella and Trigella , assigning Cyanella alba to sect.

Trigella and C. orchidiformis to sect. Pharetrella , thus

precisely opposed to Salisbury’s (1866) original place-

ment. As Pax & Hoffmann were explicitly making com-

binations based on Salisbury’s genera, however, the

types are fixed according to Salisbury's designations,

which are followed here.

Deciduous geophytes with deep-seated, tunicated

corm, tunics decaying into fibrous or firm-leathery,

coarsely netted fibres, sometimes extending into neck.

Cataphyll 1, extending to ground level and enclos-

ing leaf sheaths, entirely sheathing or with short leafy

blade. Stem simple or branched, smooth or minutely and

sparsely scabridulous. Foliage leaves 3-12, basal, lan-

ceolate to linear-lanceolate and canaliculate or filiform-

terete, firm-textured or softer, margins plane, undulate

or crispulate, smooth or scabridulous or ciliate, sur-

face mostly glabrous, rarely puberulous. Inflorescence

a raceme, usually branched, rarely highly condensed

and flowers apparently solitary, with bracts subtend-

ing branches and pedicels; pedicels suberect or spread-

ing, with solitary bracteole inserted ± midway. Flowers

zygomorphic (perianth only weakly so through tepal ori-

entation) or asymmetric (enantiomorphic) through stylar

flexure, spreading-rotate, white, yellow, orange, pink,

or mauve to blue, sometimes distinctly veined or pat-

terned, scented; tepals free, spreading or reflexed, ± sim-

ilar or weakly dimorphic with inner broader, ovate to

oblanceolate, lower concave or ± cucullate. Stamens 6,

dimorphic, either with 3 smaller posterior stamens plus

3 larger anterior stamens, or 5 smaller posterior stamens

plus I larger anterior stamen and then lowermost either

median or flexed laterally to left or right, suberect, upper

stamens arcuate, lower stamen(s) declinate; filaments

stout; anthers basifixed, upper sometimes adherent, nar-

rowly lanceolate, dehiscing by apical pores or short,

introrse slits. Ovary half-inferior, with several ovules

per locule; style terete, declinate, filiform, median or

flexed opposite lower stamen in enantiomorphic species.

Capsules ovoid to globose. Seeds ovoid, black, or dark

brown, testa surface rugose or scalariform. Basic chro-

mosome number, x = 12 (Omduff 1979).

9 spp., southern Namibia and southwestern South

Africa, mainly winter rainfall parts.

Etymology >: the name is a compound of the Greek

kyanus (blue) and -ella (diminutive), alluding to the

small blue flowers of Cyanella hyacinthoides , the first

species to be described.

Ethnobotany: the corms comprise part of the tradi-

tional diet of the Nama tribes ( Archer 410).

I. Section Trigella (Salisb.) Pax & K.Hoffm. in Die

natiirlichen Pflanzenfanrilien 15a: 427 (1930). Trigella

Key to species

la Stamens 3 + 3; flowers pink or mauve (sect. Trigella)'.

2a Leaves linear, occasionally narrowly lanceolate, 2-8 mm wide; perianth not patterned; capsules subglobose-ovoid, 6-10 mm long;

plants from southern Namibia and Richtersveld:

3a Tepals 10-12 mm long; filaments connate < halfway into short tube ± 1 mm long; anthers yellow throughout; style 10-15 mm long,

± twice as long as lower stamens

1 . C. ramosissima

3b Tepals 13-20 mm long; filaments connate halfway or more into tube 1-2 mm long; anthers greyish or mauve distally; style ± 6 mm

long, only slightly longer than lower stamens 2. C. marlothii

2b Leaves lanceolate, 10-30 mm wide; perianth sometimes patterned; capsules ovoid-ellipsoid to oblong, 10-25 mm long; plants from

Richtersveld to Western Cape:

4a Tepals (8— )1 0 — 1 5(— 20) mm long; posterior (upper) filaments arcuate or geniculate-sigmoid, ± evenly thick throughout, not flexuous

distally; anterior (lower) anthers 5-6 mm long 3 . C. orchidiformis

4b Tepals 8-10 mm long; posterior (upper) filaments swollen basally, geniculate-sigmoid and filiform in distal half and strongly flexu-

ous; anterior (lower) anthers 2. 5-3.0 mm long 4. C. cygnea

lb Stamens 5 +1; flowers white, yellow, orange, pink, or mauve to blue (sect. Cyanella):

5a Pedicels suberect; filaments connate at base only; style laterally deflexed to left or right opposite lower stamen and flowers enantiomorphic:

6a Raceme not congested; pedicels 15-30 mm long 8. C. lutea

6b Raceme congested, flowers apparently solitary among leaves; pedicels 80-120 mm long 9. C. alba

5b Pedicels ± geniculate, spreading horizontally at first then sharply flexed upwards, rarely suberect or arcuate; filaments connate for half

or more; style median and flowers not enantiomorphic:

7a Raceme lax, lower flowers 1. 5-3.0 x their length apart; bracteoles sub-basal; perianth orange 7. C. aquatica

7b Raceme dense, lower flowers 0.5-0. 6 x their length apart; bracteoles usually inserted in distal half of pedicel, rarely sub-basal; peri-

anth white, pink, or mauve to blue:

8a Upper cataphyll purple-reticulate; leaves linear, mostly 1-4 mm wide, margins conspicuously ciliate in basal half with long,

shaggy cilia 2-3 mm long but ± smooth distally

6. C. pen t her i

8b Upper cataphyll usually pale, rarely purple-reticulate; leaves linear or lanceolate, mostly 4-15 mm wide, margins smooth or ciliol-

ate along entire length with short hairs up to 1 mm long 5. C. hyacinthoides

28

Bothalia 42,1 (2012)

Salisb. : 46 (1866). Type species: Cyanella orchidiformis

Jacq.

Flowers never enantiostylous; perianth pink to

mauve, sometimes patterned. Stamens 3 + 3, lower

anthers tapering, upper anthers ± sagittate. Ovary, style

median.

1. Cyanella ramosissima (Engl. & Krause) Engl. &

Krause in Krause, Botanische Jahrbticher fur Systematik

57: 239 (1921). Iphigenia ramosissima Engl. & Krause:

124 (1910). Type: Namibia, Aus, Kubub, Oct. 1906, P.

Range 139 (Z, holo.; SAM, iso.!).

C. krauseana Dinter & G.M. Schulze: 525 (1941).

Type: Namibia, Klinghardtsgebirge, 23 Sept. 1922, M.K.

Dinter 3955 (B, holo.f; PRE!, SAM!, iso.).

Plants 80-200 mm high. Corms moderately or very

deep-seated, 1 5-30 mm diam., tunics of coarsely net-

ted, wiry fibres, extending shortly into a fibrous neck

to 20 mm long, pale brown or grey. Basal leaves 4-6,

spreading or suberect, linear to narrowly lanceolate,

50—1 50(— 200) x 2-8 mm, acute to attenuate, plane,

canaliculate or rarely involute, with prominent mid-

rib and ribbed veins abaxially, firm-textured, glabrous,

margins often ± undulate, usually ciliolate. Inflores-

cence a dense raceme up to 1 5(-20)-flowered, simple

or 1- or 2-branched, lower flowers 0.2-0. 5 x pedicel

length apart; pedicels suberect but deflexed at bracteole,

mostly 15-30 mm long; bracteoles mostly inserted in

upper third or quarter. Flowers facing outwards, pale to

deep pink or mauve with darker veins, fragrant; tepals

spreading, outer elliptic, 13-20 x 3-4 mm, apiculate,

inner oblanceolate, 13-20 x 4-7 mm, narrowed below.

Stamens dimorphic, 3 + 3; filaments of posterior cluster

sometimes almost geniculate, 2.5-3.0(-4.0) mm long,

swollen basal ly and connate into short tube up to 1 mm

long, yellow, anthers ± sagittate, outer smaller, ± 1 .5 mm

long, median ± 2 mm long, yellow; filaments of anterior

cluster deflexed, 2. 0-2. 5 mm long, shortly connate for

up to 1 mm, anthers 4-5 mm long, yellow. Ovary half-

inferior, style medially deflexed, 10-15 mm long, almost

twice as long as lower stamens. Capsules erect, sub-

globose-ovoid, 7-10 x 7 mm, 3-lobed. Seeds unknown.

Flowering time : mainly Jul. and Aug. (-early Oct.).

Distribution and ecology, restricted to the win-

ter rainfall part of southern Nambia, where it has been

FIGURE 5.- — Distribution of Eremiolirion amboense , o; Cyanella ram-

osissima, • .

recorded on the higher ground, 350-1 050 m, from Aus

and the Klinghardt Mtns along the Huib Hoch Plateau,

extending into the central Richtersveld in South Africa

as far south as Eksteenfontein (Figure 5). The species

occurs on open stony flats, alluvial ridges, rocky terraces

or sometimes on sandy or calcareous flats, in arid succu-

lent karoo shrubland or sparse desert vegetation.

Diagnosis and relationships: readily recognized by

the linear leaves, 2-8 mm wide, and dense raceme of

large, pink to mauve flowers with 3 + 3 arrangement of

stamens with plain yellow anthers, and a consistently

long style, 10-15 mm long, thus almost twice as long as

the lower stamens. Cyanella ramosissima may be con-

fused with vegetatively similar C. marlothii, which has

smaller flowers with the filaments of all six stamens con-

nate for half or more of their length into a tube 1-2 mm

long, bicoloured anthers, and a short style, ± 6 mm long.

The distinctive combination of narrow leaves and a

long style separates C. ramosissima from the forms of

C. orchidiformis with unpattemed tepals. The two spe-

cies share smaller upper lateral anthers and otherwise

resemble one another very closely although they are

readily distinguished in fruit, as C. ramosissima has

much smaller, subglobose or ovoid capsule, 7-10 mm

long vs. the large, oblong or ellipsoidal capsules,

12-15 mm long of C. orchidiformis. Although C. orchid-

iformis mostly has the style shorter than the lower

anthers, occasional collections (see below) have elon-

gated styles like those of C. ramosissima. In the absence

of fruits, such aberrant plants can be identified by their

broader, soft-textured leaves and slightly larger anthers,

tinged greyish distally. The two species are essentially

allopatric, overlapping in their distribution only in the

Richtersveld, where C. ramosissima is restricted to the

mountainous central region whilst C. orchidiformis

extends around the fringes.

Representative specimens

NAMIBIA. — 2616 (Aus): Farm Klein Aus, (-CB), 11 Aug. 1959,

Giess & Van Vuuren 756 (BOL, PRE); 200 m N of T-junction, (-CB),

21 Oct. 1983, Van Berkel 538 (NBG); Luderitz District, Farm Aub, (—

CB), without date, Lavranos & Pehlemann 21700 (MO); Aus Town-

lands, (-CB), Sept. 1983, Lavranos & Pehlemann 21592 (MO). 2715

(Bogenfels): Klinghardtberge, (-BD), 17 Aug. 1986, Van Berkel 571

(NBG, PRE); W Hockster Mtns, 2 km NW of Hockster, (-BD), 21 July

1986, Van Berkel 558 (NBG). 2716 (Witputz): Namuskluft, (-DD), 1 1

July 1988, Bruyns 3191 (NBG); Farm Spitskop, Rosh Pinah, (-DD),

Aug. 1981, Lavranos 19935 (MO). 2817 (Vioolsdrif): Orange River

just east of confluence with Fish River, (-AA), 1 July 1989, Oliver

9177 (NBG).

NORTHERN CAPE. — 2817 (Vioolsdrif): crest of ridge near Hot-

tentotsparadys, (-AC), 9 Sept. 1996, Bayer & Puttock SAF96157

(NBG); Stinkfontein Mtns, near foot of Cornellsberg, (-CA), 22 Aug.

1994, Goldblatt & Manning 9952 (NBG); E of Eksteenfontein, (-CD).

July 1989, Williamson 4264 (NBG).

2. Cyanella marlothii J.C. Manning & Goldblatt, sp.

nov.

TYPE. — Northern Cape, 2817 (Vioolsdrif): sandy

flats between Jasper’s werf and Doompoort [Doring-

poort Farm at W foot of Ploegberg], (-CA), 26 Aug.

1925, R. Marloth 1211 (PRE, holo.).

Plants 200-350 mm high. Corms moderately deep-

seated, 15-30 mm diam., tunics of coarsely netted, wiry

Bothalia 42,1 (2012)

29

FIGURE 6. — Cyanella floral details.

A, B, C. orchidiformis : A,

flower; B, stamens and style.

C, D, C. cygnea : C, flower; D,

stamens and style. E, C. mar-

lothii, stamens and style. F-H,

C. hyacinthoides : F, flower;

G, stamens and style; H, dor-

sal view of upper stamens. I,

J, C. aquatica : I, flower; J, sta-

mens and style. K., L, C. lutea:

K, flower; L, stamens and

style. M, N, C. alba : M, flow-

er; N, stamens and style. Scale

bar: A, C, F, I, K, M, 10 mm;

B, D, E, G, H, J, L, N, 2 mm.

Artist: John Manning.

fibres, extending shortly into a fibrous neck to 20 mm

long, pale brown or grey. Basal leaves 4-6, suberect,

linear or linear-lanceolate, 50-100 x 2-6 mm, acute to

attenuate, canaliculate or involute, with prominent mid-

rib and ribbed veins abaxially, firm -textured, glabrous,

margins ± undulate, sparsely scabridulous-ciliolate.

Inflorescence a moderately dense raceme up to 20-11 ow-

ered, with up to 2 branches, lower flowers 0.5-0. 6 x

pedicel length apart; pedicels suberect, deflexed at

bracteole, mostly 20-30 mm long; bracteoles inserted

in upper third. Flowers facing outwards, pale mauve

(‘blue’) with darker veins, presumably fragrant; tepals

spreading, outer elliptic, 10-12 x 2-3 mm, apiculate,

inner oblanceolate, 10-12 x 2-3 mm, narrowed below.

Stamens dimorphic, 3 + 3; filaments erect but deflexed

apically, 2-3 mm long, connate halfway or more into

cylindrical tube 1-2 mm long; posterior anthers ± sag-

ittate, outer smaller, 1. 5-2.0 mm long, median 2.0-

2.5 mm long, yellow but greyish or mauve distally, ante-

rior anthers 3^1 mm long, yellow basally but greyish

or mauve in distal 2/3. Ovary half-inferior; style medi-

ally deflexed, ± 6 mm long, extending shortly beyond

anthers. Capsules subglobose, 6-7 mm diam., 3-lobed.

Seeds unknown. Flowering time'. Aug.-Sept. Figure 6E.

Distribution and ecology, thus far known from a sin-

gle collection from sandy flats near the Ploegberg, south

of Kuboes in the Richtersveld (Figure 7).

Diagnosis and relationships', this distinctive species

has the 3 + 3 arrangement of stamens that characterizes

sect. Trigel/a , but is distinguished from other members

in the section by having the filaments of all six sta-

mens connate for half to two-thirds of their length into

a cylindrical or conical tube that completely encloses

30

Bothalia42,l (2012)

FIGURE 7. — Distribution of Cyanella marlothii, o; C. orchidi-

formis, •.

the ovary. Although the species is based on just a sin-

gle collection, this comprises three essentially identical

individuals. These plants were included in C. orchidi-

formis until now, despite their unique androecium. In

this context it is significant that Marloth, who also col-

lected true C. orchidiformis from Steinkopf on the same

trip as C. marlothii , correctly identified the former but

treated the latter as C. capensis (now C. hyacinthoides),

a clear indication that he considered the Ploegberg col-

lection to be distinct from C. orchidiformis , although he

was misled by the connate filaments into misidentifying

it as C. hyacinthoides. The latter does not occur in the

Richtersveld, and is in any event immediately distin-

guished by its 5 + 1 arrangement of anthers and by the

spreading-geniculate pedicels.

Among the members of sect. Trigella , C. marlothii

resembles C. ramosissima in its narrow leaves, 2-6 mm

wide, pale mauve or blue flowers with darker veins, and

apparently ± globose capsules, but is separated from it

by its smaller flowers with tepals 10-12 vs. 13-20 mm

long, bicoloured vs. plain yellow anthers, and shorter

style, ± 6 mm long and only slightly longer than the

anthers vs. 10-15 mm long and ± twice as long as the

anthers.

We have considered the possibility that the collec-

tion may be hybrid between a member of sect. Tri-

gella ( C . cygnea is recorded from the Kuboes area) and

C. hyacinthoides , but discount this in view of the con-

sistent appearance of the plants and the lack of other

intermediate characters. This possibility did not suggest

itself to Marloth, who did not record any potential parent

species at the site. The absence of additional collections

of the taxon is unfortunate but not unique — no further

plants of W. gracilis have been recorded from the Rich-

tersveld since Marloth’s collection on 29 August 1925,

just three days after his collection of C. marlothii (but

see this species for further comment).

3. Cyanella orchidiformis Jacq., Collectanea 4: 211

(1791). Trigella orchidiformis (Jacq.) Salisb. : 46 (1866).

Type: South Africa, without locality or collector, illus-

tration in Jacquin, leones plantarum rariorum 2: t. 447

(1786-1793).

Plants 150-500 mm high. Conns moderately or very

deep-seated, 15-30 mm diam., tunics of coarsely net-

ted, woody fibres, sometimes connate below into flat

claws, extending shortly into a fibrous neck up to 20 mm

long, chestnut-brown. Basal leaves 4-6, suberect or

spreading, lanceolate, 70-250 x 1 0— 25(— 30) mm, acute

to attenuate, plane or canaliculate, with prominent mid-

rib abaxially, soft-textured, glabrous, margins plane or

undulate, smooth or ciliolate-scabridulous. Inflores-

cence a moderately dense raceme up to 35-flowered,

with 1 or 2 branches, lower flowers 0.5-0. 8 x pedicel

length apart; pedicels suberect and deflexed at bracteole,

mostly 15-30 mm long; bracteoles mostly inserted in

upper third or quarter, sometimes in lower half or quar-

ter. Flowers facing outwards, pink or mauve with darker

veins, sometimes with darker centre, or with paler cen-

tre variously speckled with dark pink, the whole outlined

with darker shading, fragrant; tepals spreading, outer

elliptic, (8—) 1 0— 1 5(— 20) x 4-5 mm, apiculate, inner

oblanceolate, (8 — ) 1 0—1 5( — 20) x 5-6 mm, narrowed

below. Stamens dimorphic, 3 + 3; filaments of poste-

rior cluster arcuate to geniculate-sigmoid, 2-5 mm long,

connate at extreme base only, ± evenly thick through-

out, yellow with white base, anthers ± sagittate, outer

smaller, 1-2 mm long, median 2-3 mm, yellow, but grey

to purple distally; filaments of anterior cluster deflexed,

1.5-2. 5 mm long, connate at extreme base, anthers

5-6 mm long, pale yellow at base, greyish or purple dis-

tally. Ovary half-inferior; style medially deflexed, (4-)5-

13 mm long, ± as long as or extending well beyond

anthers. Capsules erect, ovoid-ellipsoid to oblong,

14-25(-30) x 8-10 mm, pale with purplish reticulation.

Seeds ovoid-ellipsoid, 3^1 x 1. 5-2.0 mm, glossy black,

rugose. Chromosome number : 2n = 24 (Ornduff 1979).

Flowering time : (mid-)late Jul.-late Sept. Figure 6A, B.

Distribution and ecology, occurring along the west-

ern escarpment, from just north of Steinkopf in northern

Namaqualand to Citrusdal in the Olifants River Valley

(Figure 7). Collections from the Richtersveld cited under

this species by Scott (1991) are referable to C. cygnea ,

evident from their filiform, sigmoid upper filaments and

smaller anthers. Plants grow mostly in clay or loamy

soils, often in rock crevices in granite or sandstone,

where they benefit from extra moisture through run-

off among rocks along the courses of seasonal streams,

especially in Namaqualand.

Diagnosis and relationships : the most common

and widespread of the three species of sect. Trigella ,

C. orchidiformis, is recognized by its lanceolate leaves,

10-25 mm wide, and racemes of pink to mauve flowers,

mostly darker or patterned toward the centre, with the

anthers partially or almost wholly greyish or purple, and

large, ovoid-ellipsoid fruits, 14-25 mm long. The three

species are essentially parapatric or allopatric, although

both C. cygnea and C. orchidiformis have been collected

near Steinkopf ( Marloth 6761. 6761 A). An exceptionally

large-flowered variant with tepals 20 x 6-7 mm has been

collected on the Gifberg Pass, growing in sandstone soil

after fire ( Goldblatt & Porter 13190), and may be poly-

ploid.

Cyanella orchidiformis is closely allied to C. cygnea,

with which it shares the distinctive large fruits, patterned

perianth, and coloured anthers, but from which it is dis-

tinguished by its generally larger flowers, with tepals

mostly 10-13 mm long vs. 8-10 mm long, and its unex-

Bothalia 42,1 (2012)

31

ceptional stamens. The upper filaments in C. orchidi-

formis are arcuate or weakly geniculate, without a bul-

bous base and not evidently filiform in the distal half,

and the lower anthers are relatively large, 5-6 mm long.

The style is very variable in length, mostly 5-10 mm

long, but occasionally up to 15 mm long. In contrast,

C. cygnea has mostly smaller flowers, with tepals

8-10 mm long and very distinctive stamens, with the

upper filaments geniculately sigmoid and sharply nar-

rowed and filiform in the distal half, with much smaller

lower anthers, 2. 5-3.0 mm long, and a short style

3-4 mm long. The range of C. orchidiformis is largely

to the south and east of C. cygnea but both species have

been collected near Steinkopf.

The relatively broad leaves, 10-30 mm wide, and

large capsules, readily distinguish C. orchidiformis

from C. marlothii and C. ramosissima , which have nar-

row leaves 2-8 mm wide and smaller, subglobose-ovoid

fruits 7-10 mm long.

Vernacular name : waterraap.

Representative specimens

NORTHERN CAPE.— 2917 (Springbok): Steinkopf, ( -BA),

Aug. 1925, Marloth 6761 (NBG); 6.5 km W of Steinkopf, (-BA), 29

Sept. 1986, Perry <& Snijman 3560 (NBG); between Springbok and

Steinkopf beyond Bulletrap, (-BC), 29 Sept. 1986, Peny & Snij-

man 3555 (NBG); Spektakel, (-DA), 25 Aug. 1941, Compton 11398

(NBG); Eselsfontein, (-DA), 8 Sept. 1950, Barker 519 (NBG). 3017

(Hondeklipbaai): Spoegivier, (-AD), 12 Sept. 1982, Archer 295

(NBG). 3018 (Kamiesberg): 6 miles [9.6 km] north of Garies, (-CA),

3 Sept. 1945, Leighton 1398 (PRE); Kamiesberg, 41.5 km from turn-

off to Kliprand, (-DC), 15 Sept. 2006, Goldblatt & Porter 12759A

(MO, NBG). 3117 (Lepelfontein): Towerberg Pass between Komkans

and Kotzesrust, (-BB), 3 Sept. 1976, Boucher 3160 (NBG). 3 1 19 (Cal-

vinia): Lokenburg, (-AC), 23 Aug. 1980, Van Berkel 204 (MO).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Meerhofkasteel, (-

AA) , 8 Aug. 1984, Snijman 805 (NBG); Farm Quaggaskop 125, (-

AB) , 11 Aug. 1977, Le Roux 2282 (NBG); 15 miles [24 km] NW of

Koekenaap, (-AD), 19 Aug. 1970, Hall 3766 (NBG); between Trawal

and Olifants River bridge, shale bank, (-DC), 27 Aug. 1991, Goldblatt

& Manning 9121 (MO); Gifberg Pass, Keurlandshoek, (-DD), 25 Sept.

2008, Goldblatt & Porter 13190 (MO, NBG). 3218 (Clanwilliam):

Clanwilliam, (-BB), 5 Aug. 1896, Schlechter 8417 (MO, NBG); 29

July 1943, Lewis NBG1814/32 (NBG); Olifants Dam, (-BB), 14 Sept.

1847, Barker 4768 (NBG). 3219 (Wuppertal): Biedouw [Bidouw]

Valley, (-AA), 23 Sept. 1952, Barker 1748 (NBG); Cedarberg Forest

Reserve, Langrug, (-AC), 21 Aug. 1983, Viviers 496 (NBG); Rondegat

River Valley 16 km NW of Algeria, (-BC), 8 Sept. 1976, Thompson

2812 (NBG); near Citrusdal, (-CC), 6 Sept. 1949, Steyn 390 (NBG).

Long-styled morphs

3017 (Hondeklipbaai): Grootvlei, (-BB), Sept. 1945, Lewis 1380

(SAM); 7 Sept. 1945, Barker 3716 (SAM). 31 18 (Vanrhynsdorp): Hol-

bak Farm, near Doombaai [Doringbaai], (-CD), 5 Sept. 1964, Hal 1 164

(NBG). 3218 (Clanwilliam): S of Clanwilliam, (-BB), 20 Sept. 1954,

De Vos 1719 (NBG); 10 miles [18 km] S of Clanwilliam, (-BB), July

1948, Lewis 2999 (SAM).

4. Cyanella cygnea G. Scott in South African Journal

of Botany 57: 50 (1991). Type: South Africa, [Northern

Cape], 51.4 km from Springbok along road to Komag-

gas, 16 Sept. 1988 [cult, at Karoo Botanic Garden,

Worcester from material collected ± 1978], P.L. Perry

1119 (NBG, holo.!; K, MO, PRE!, iso.).

Plants (150-)200-500 mm high. Corms moder-

ately or very deep-seated, 15-30 mm diam., tunics

of coarsely netted, woody fibres, sometimes connate

below into flat claws, extending shortly into a fibrous

neck up to 20 mm long, chestnut-brown. Basal leaves

4-6, suberect, lanceolate, 80-200 x 1 0— 20(— 25 ) mm,

acute to attenuate, plane or canaliculate, with promi-

nent midrib and ribbed veins abaxially, soft-textured,

glabrous, margins smooth or ciliolate-scabridulous.

Inflorescence a dense or moderately dense raceme up

to 35-flowered, with 1-4 branches, lower flowers 0.2—

0.5 x pedicel length apart; pedicels suberect, deflexed

at bracteole, mostly 1 5-30 mm long; bracteoles mostly

inserted in upper third or quarter. Flowers facing out-

wards, pink with paler centre variously speckled with

dark pink, the whole outlined with darker shading, fra-

grant; tepals spreading, outer elliptic, 8-10 x 4-5 mm,

apiculate, inner obovate, 8-10 x 5-6 mm, narrowed

and short-clawed below. Stamens dimorphic, 3 + 3; fila-

ments of posterior cluster geniculate-sigmoid, 2-5 mm

long, distally filiform and strongly flexuous, swollen

basally, connate at extreme base only, yellow with white

base, anthers ± sagittate, 1. 5-2.0 mm long; filaments of

anterior cluster deflexed, 1.0-1. 5 mm long, connate at

extreme base, anthers 2. 5-3.0 mm long, pale yellow but

greyish in distal half or third. Ovary half-inferior; style

medially deflexed, 3-4 mm long, not extending beyond

anthers. Capsules erect, ovoid-ellipsoid, ( 1 2—) 1 5—20

x 8-10 mm. Seeds ovoid-ellipsoid, 3-4 x 1. 5-2.0 mm,

glossy black, rugose. Flowering time : late Aug.-early

Oct. (-early Nov.). Figure 6C, D.

Distribution and ecology >: restricted to the higher-

lying parts of northern Namaqualand, where it has been

collected in the Richtersveld along the Ploegberg and

Stinkfontein Mtns, from Kuboes to Eksteenfontein, near

Steinkopf, and along the edge of the escarpment around

Komaggas, some 60 km to the south (Figure 8). Plants

grow in rocky situations in open succulent karoo shrub-

land, typically where there is additional moisture such as

along watercourses or in gorges.

Diagnosis and relationships', closely allied to

C. orchidiformis, with which it shares characteristi-

cally mottled flowers and large, ovoid-ellipsoid cap-

sules ± 15 mm long, and greyish or purple markings

or speckling on the anthers. Cyanella cygnea typically

has smaller flowers, with tepals 8-10 vs. (8 — )1 0—1 5( —

20) mm long, but is best identified by its stamens. The

FIGURE 8. — Distribution of Cyanella cygnea.

32

Bothalia 42,1 (2012)

strongly geniculate-sigmoid filaments of the posterior

(upper) stamens are bulbous at the base and filiform

in the distal half, giving them a characteristic flexuous

form, the outer pair slightly longer than the median.

All three anthers in the posterior cluster are subequal in

size, 1 .5-2.0 mm long, and the lower anthers are equally

larger, 2. 5-3.0 mm long. In contrast, the upper stamens

in C. orchidiformis are ± uniformly thick except at the

extreme apex and not evidently flexuous, the outer

anthers are slightly smaller than the median, and the

lower anthers are larger, 5-6 mm long, sometimes with

the median larger than the laterals. The distributions of

the two species are largely complementary, with C. cyg-

nea occurring to the north and west of C. orchidiformis ,

but they overlap around Steinkopf.

Vernacular name : wildebeet (wild beet) (Scott 1991 ).

Representative specimens

NORTHERN CAPE.— 2816 (Oranjemund): mtns SW of Kuboos

[Khubus], (-BD), 11 Sept. 1973, Lavranos 10834 (MO, PRE). 2817

(Vioolsdrif): Richtersveld, Kodaspiek, (-AA), 2 Sept. 1977, Oliver,

Tolken & Venter 492 (MO); Armmanshoek, (-AC), Aug. 1995, G. & F.

Williamson 5654 (NBG); Richtersveld, near Kubus [Khubus], (-CA),

13 Aug. 1983, Archer 391 (NBG, PRE); Ploegwater at S portion of

Ploegberg, (-CA), 7 Sept. 1991, Germishuizen 5483 (PRE); Stinkfon-

teinberg SW of Vanzylsrus, (-CA), 4 Sept. 1977, Oliver, Tolken & Ven-

ter 626 (NBG); stony flats 4 km N of Eksteenfontein, (-CD), 23 Aug.

2001, Goldblatt & Porter 11751 (MO); 8 km N of Eksteenfontein,

22 Aug. 1994, Goldblatt & Manning 9940 (MO). 2917 (Springbok):

Steinkopf, (-BA), Aug. 1925, Marloth 6761A (NBG); Steinkopf, (—

BC), 9 Aug. 1898, M. Schlechter 119 (MO, PRE); Klipfontein, (-BA),

Sept. 1929, Grant 4840B (MO); Komaggas, Van Reenen se Water, (-

DC), 26 Aug. 1983, Van Wyk 6501 (PRE).

II. Section Cyanella

Flowers sometimes enantiostylous; perianth white,

yellow, orange, pink to mauve, or blue, never patterned.

Stamens 5 + 1; anthers ± oblong. Ovary, style some-

times flexed to left or right.

Series Hyacinthoides J.C. Manning & Goldlbatt, ser.

nov.

Flowers not enantiostylous; pedicels ± geniculate

(horizontally spreading then flexed sharply upwards) or

arcuate; perianth white, orange, pink, or mauve to blue.

Stamens: filaments connate halfway or more. Ovary:

style not flexed sideways. Type species: Cyanella hya-

cinthoides Royen. ex L.

5. Cyanella hyacinthoides Royen ex L., Genera

plantation, edn 5: addendum [522] (1754). C. capen-

sis L.: 985 (1759), norm illegit. superfl. C. pulchella

Salisb.: 249 (1796), nom. illegit. superfl. [Note: Scott’s

(1991) lectotypification of C. pulchella against Jac-

quin’s (1776-1777) illustration of C. capensis L. is

unwarranted and incorrect. There is no indication that

Salisbury had any intention other than of replacing Lin-

naeus’s name with his own]. Type: South Africa, without

precise locality, date or collector, ex herb. Royen Herb.

Linn. 430.2 (LINN, holo.!).

Plants 1 50-400(-500) mm high. Conns deep-seated,

25-30 mm diam., tunics of coarsely netted, wiry or

woody fibres, not or extending shortly into a fibrous

neck to 20 mm long, pale brown or grey. Basal leaves

4 — 9( 1 2), suberect or spreading, linear to narrowly lan-

ceolate, 60-200(-250) x (2— )4— 1 5( — 25) mm, acute to

attenuate, plane, canaliculate or rarely involute, mid-

rib and veins prominent beneath (abaxially), firm-tex-

tured, usually glabrous but veins sometimes scabridu-

lous or puberulous to villous abaxially with hairs up

to 1 mm long, rarely both surfaces densely puberulous

throughout, margins ± undulate or crispulate, usually

ciliolate-scabridulous, sometimes flushed purple basally;

upper cataphyll usually pale, rarely purple-reticulate or

fenestrate. Inflorescence a moderately dense raceme

up to 25-flowered, with 2-4 branches, rarely with sec-

ond order branchlets and thus paniculate, lower flow-

ers 0.3-0. 6 x pedicel length apart; pedicels usually

geniculate, horizontal in basal 1/2 or 2/3 then abruptly

flexed upwards at ± right angles, rarely suberect or arcu-

ate, mostly 20-30 mm long; bracteoles mostly inserted

between lower and upper third, rarely sub-basal. Flow-

ers facing outwards, pale to deep mauve or blue, rarely

white or pink, fragrant; tepals spreading, ovate to obo-

vate, 8-10 x 3 4 mm, apiculate. Stamens dimorphic, 5 +

1; filaments of posterior cluster 1.0-2. 5 mm long, outer

sometimes slightly longer than inner, connate ± halfway

or almost completely into tube 1 .0-2.0 mm long, yellow,

sometimes with small intrastaminal lobules between

bases of filaments, anthers 1 .5— 2.0(— 2.8) mm long,

yellow; anterior stamen with filament ± 1 mm long,

connate to upper cluster for ± half length, anther 2.5-

4.0 mm long, yellow. Ovary half-inferior; style medially

deflexed, 3^4 mm long, not extending beyond anthers.

Capsules erect on geniculate pedicels, subglobose,

5-6 mm diam., 3-lobed and retuse. Seeds ovoid, ± 2 mm

diam., rugulose. Chromosome numbers: 2n = 24, 28 &

48 (Ornduff 1979). Flowering time: mainly mid-Sept-

mid-Dec. but mid-Aug.-mid Oct. in Namaqualand. Fig-

ure 6F-H.

Distribution and ecology: Cyanella hyacinthoides is

widely distributed through the southern African winter-

rainfall region, from just north of Steinkopf southwards

through the higher-lying parts of Namaqualand into

the southwestern Cape as far east as the Gouritz River

(Figure 9), from near sea level to over 1 200 m. It has

been recorded along the Roggeveld Escarpment south

to Matjiesfontein but is absent from the arid Tanqua

River basin and Little Karoo, apart from a single col-

lection south of Oudtshoorn at the foot of the Outeniqua

Mtns. The species has a wide edaphic amplitude and has

FIGURE 9. — Distribution of Cyanella hyacinthoides (pubescent

forms, o).

Bothalia 42,1 (2012)

33

been collected on granite, sandstone, and limestone sub-

strates, although it favours loamy or clay soils, where it

is most often found as a component of renosterveld or

succulent karooid communities. It is tolerant of distur-

bance and thrives in old lands and along road verges.

Cyanella hyacinthoides is extremely variable in its

foliage. Plants typically produce 4-6 lanceolate leaves

but some forms may produce up to a dozen linear- in vo-

lute leaves. These narrow-leaved plants are scattered

throughout the range of the species. A more circum-

scribed ecotype occurs along the higher parts of cen-

tral Namaqualand, between Kotzesrus and Springbok.

Plants there tend to have the lower leaf surface vari-

ously scabridulous or villous, with the hairs restricted

to the leaf margins and the adaxial veins and midrib.

In extreme forms, the hairs are shaggy and up to I mm

long but there is a significant variation in the density

and length of the vestiture, even within a single local-

ity, from scarcely puberulous to densely villous leaves.

A collection from north of Komaggas (Barker 7412)

shows a second type of vestiture, with both leaf sur-

faces closely and evenly puberulous. Populations from

elsewhere in the range generally have the leaf surfaces

glabrous, but some plants may have the lower surface

sparsely and minutely scabridulous along the veins.

There is no association between vestiture and other veg-

etative features, such as leaf width or shape. The devel-

opment of leaf pubescence in populations from this part

of Namaqualand has also been recorded in species of

Trachyandra (Asphodelaceae) (Manning & Goldblatt

2007) and Haemanthus (Amaryllidaceae) (Snijman

1984), and appears to represent a widespread ecological

strategy.

Tetraploids have been detected among several wild

populations of Cyanella hyacinthoides (Ornduff 1979),

and it is thus possible that unusually robust specimens

that have been remarked on by various collectors are

polyploids.

Diagnosis and relationships : Cyanella hyacinthoides

is distinguished by the moderately dense, branched

racemes of mauve to blue (rarely white or pink) flow-

ers with 5 + 1 arrangement of stamens with the filaments

connate for ± half their length or more. The connate

filaments and generally horizontally spreading pedi-

cels serve to distinguish the species from pink-flowered

forms of C. lutea , in which the stamens are ± free and

the pedicels mostly suberect.

The species is closely allied to C. pentheri , with

which it has been much confused, and the two were

treated as conspecific by Scott (1991). They are essen-

tially alike in their inflorescence, although the flowers

in C. pentheri are typically paler, mostly white to pale

mauve, but they differ strikingly in their foliage. The

leaves of C. pentheri are linear-aristate and canalicu-

late-involute with margins that are often crispulate and

conspicuously ciliate only towards the base with shaggy

hairs 2. 0-3.0 mm long. Similar long cilia also fringe

the upper cataphyll, which is funnel-shaped, and boldly

pigmented with deep purple along the edges and veins,

giving it a characteristic fenestrate appearance. Although

C. hyacinthoides is highly variable in its foliage, the spe-

cies only rarely produces similarly narrow, crispulate

leaves and in such cases they are either glabrous or are

ciliolate-pubescent along their entire length, with much

shorter hairs 0. 2-1.0 mm long, and the upper cataphyll

is usually unmarked, very rarely ( Goldblatt & Porter

11896) purple-fenestrate. Although the two taxa have

been recorded growing in close proximity in several

localities (see discussion under C. pentheri ), no interme-

diates between them have been found.

Vernacular names : raap, hotnotsraap, klipraap.

Representative specimens

Typical form

NORTHERN CAPE.— 2917 (Springbok): E of Rosies, (-BA), 15

Oct. 1988, Williamson 3978 (NBG); Springbok, 15 miles [24 km] E

of town, (-CB), 12 Oct. 1947, Rodin 2193 (PRE); between Spektakel-

berg and Komaggas, (-DA), 21 Aug. 1982, Le Roux 2957 (NBG).

3017 (Hondeklipbaai): Spoegrivier, (-AD), 20 Sept. 1983, Archer

410 (NBG); Kamieskroon, Skilpad Nature Reserve, (-BB), 18 Sept.

1995, Cruz 92 (MO, NBG); sandy flats E of Kamieskroon at foot of

pass, ( BB ), 3 Nov. 1982, Goldblatt 6651 (MO). 3118 (Vanrhynsdorp):

5 km S of Bitterfontein, (-AB), 9 Sept. 1985, Duncan 184 (NBG);

Vanrhynsdorp, Zandkraal Farm, (-DB), 7 Sept. 1949, Barker 5662

(NBG); 13 km from Vanrhysdorp on road to Nieuwoudtville, (-DB),

31 Aug. 1986, Fellingham 1116 (PRE). 3119 (Calvinia); Oorlogskloof

Nature Reserve, Farm Driefontein, (-AC), 1 Nov. 1996, Pretorius 398

(NBG); along Nieuwoudtville-Loeriesfontein road, (-AB), 11 Sept.

1986, Steiner 1360 (NBG); Doringbos Valley, (-CC), 27 Sept. 1970,

Barker 10725 (NBG). 3217 (Vredenburg): Witteklip Rocks, (-DD), 19

Sept., Perry 3197 (MO). 3218 (Clanwilliam): Lamberts Bay, Nortier

Experimental Farm, (-AB), 6 Nov. 1974, Boucher 2569 (NBG); irri-

gation dam near Clanwilliam, (-BB), Sept. 1935, Smuts PRE59124

(PRE); Clanwilliam, 6.2 km S of Ramskop, (-BB), 26 Sept. 1986,

Peny 3523 (NBG); Piketberg, approaching Moravian Mission at

Goedverwag, (-DC), 3 Oct. 1984, Peny 3214 (MO, NBG). 3219

(Wuppertal): Bidouw, Welbedacht Farm, (-AA), 22 Sept. 1952, John-

son 537 (NBG). Koue Bokkeveld, Ondertuin, (-CC), 28 Dec. 1978,

Hanekom 2519 (MO). 3220 (Sutherland): Roggeveld, Soekop Farm,

(-AA), 11 Sept. 2006, Rosch 660 (NBG). 3219 (Wuppertal): Citrus-

dal, (-CC), 30 Sept. 1944, Barker 3075 (NBG). 3318 (Cape Town):

Langebaan, (-AA), 5 Oct. 1969, Axelson 80 (NBG); Yzerfontein, De

la Rey Farm, (-AC), 15 Oct. 1995, Boucher 2557 (NBG); Groenekloof

[Mamre], (-AC), 1850, Zeyher 1718 (NBG); Buck Bay Farm, (-CA),

29 Nov. 1978, Boucher 4156 (PRE); Robben Island, (-CD), 14 Nov.

1985, Lloyd 574 (NBG); Cape Peninsula, Kamps [Camps] Bay, (—

CD), Dec. 1897, Thode s.n. (NBG); Malmesbury, Burgers Post Farm,

(-DA), 17 Oct. 1979, Boucher <£ Shepherd 4839 (NBG); Langver-

wacht above Kuils River, (-DC), 22 Nov. 1973, Oliver 4806 (NBG);

Paarl Mountains Nature Reserve, (-DD), 26 Oct. 1994, Swanepoel 50

(NBG); Jonkershoek, (-DD), 27 Nov. 1973, Smith 141 (NBG). 3319

(Worcester): Ceres, Lakenvlei Farm, (-BC), 19 Oct. 1941, Barker

2004 (NBG); Rawsonville, (-CA), 18 Oct. 1980, Walters 2322 (NBG);

Worcester, (-CB), 17 Oct. 1980, Walters 2310 (NBG); E approach to

Franschhoek Pass, (-CC), 8 Nov. 1987, Goldblatt & Manning 8583

(MO, PRE); Madeba Farm, W of Robertson, (-DD), 8 Oct. 1986,

Hilton-Taylor 1765 (NBG). 3320 (Montagu): Matjiesfontein, (-BA),

24 Oct. 1921, Foley 120 (PRE). 3322 (Oudtshoom): lower N slopes

of Outeniqua Mtns, near Sebrafontein Farm, (-CC), 23 Oct. 1985,

Vlok 1216 (NBG). 3418 (Simonstown): Simonstown, Redhill Plateau,

(-AB), 19 Nov. 1970, Goldblatt 5168 (MO); Cape Peninsula, Noord

Hoek, (-AB), 30 Nov. 1943, Wasserfall 674 (NBG); Muizenberg, (—

AB), Feb. 1907 (mostly in fruit), Rogers TM25828 (PRE); Betty’s

Bay, 2 Dec. 1970, Ebersohn s.n. (NBG). 3419 (Caledon): Hermanus,

Vogelgat, (-AD), 30 Oct. 1986, Williams 3719 (MO, NBG); Genaden-

dal, (-BA), 1854, Roser PRE15439 (PRE); Gansbaai, Grootbos Nature

Reserve, (-CB), 8 Dec. 2007, Lutzeyer s.n. (NBG). 3420 (Bredasdorp):

Swellendam, Bontebok National Park, (-AB), Dec. 1962, Liebenberg

6779 (NBG. PRE); De Hoop, Potberg Nature Reserve, (-AD), 28 Nov.

1978, Burgers 1598 (NBG); Riversdale, Reisiesbaan Siding, (-AB), 31

Oct. 1979, Bohnen 7043 (NBG). 3421 (Riversdale): near Still Bay on

Rietvlei Road, (-AD), 13 Nov. 1982, Bohnen 8152 (NBG); limestone

hills S of Albertinia, (-AD), 4 Dec. 1985, Goldblatt 7421 (MO); Farm

Platbos, 2 km S of Aasvogelberg to Gouritz River, (-BC), 10 Dec.

1981, Stirton 10261 (NBG).

34

Bothalia 42,1 (2012)

Hairy forms

NORTHERN CAPE. — 2917 (Springbok): Spektakel Pass, (-DA),

4 Sept. 1951, Martin 835 (NBG), 11 Sept. 1993, Goldblatt & Man-

ning 9715 (MO); Ezelsfontein, (-DA), 8 Sept. 1950, Barker 6656

(NBG); 5 miles [8 km] N of Komaggas, (-DB), 4 Sept. 1951, Barker

7412 (NBG); between Brakwater and Komaggas, (-DB), 9 Sept. 1950,

Barker 6679 (NBG); 64.5 km W of Okiep towards Nababiep, (-DB),

26 Sept. 1986, Perry 3550 (NBG). 3017 (Hondeklipbaai): 7 miles

[11 km] NW of Kamieskroon, (-BB), 25 Sept. 1952, Acocks 16477

(PRE); Kamieskroon, (— BB), 22 Aug. 1959, Barker 9001 (NBG);

Garies Hill, (-BD), 2 Sept. 1951, Barker 7403 (NBG); 19 km S of

Kotzesrus, (-DD), 16 Sept. 2001, Goldblatt & Porter 11896 (MO,

NBG). 3018 (Kamiesberg): 26 km S of Garies on road to Bitterfontein,

Farm Mostertsvlei, (-CA), 30 Sept. 1987, Reid 1310 (PRE).

6. Cyanella pentheri Zahlbr. in Annalen des kaiser-

lichen naturhistorischen Museums 15: 26 (1900). Type:

South Africa, [Western Cape], Olifantrivier [Olifants

River], Aug. [without year], Penther 400 (W, holo.f).

Neotype: South Africa, [Western Cape], Clanwillam,

Biedouw [Bidouw], Welbedacht Farm, 22 Sept. 1952,

A.J. Middebnost 1741 (NBG, neo., designated here;

SAM, iso.).

Note'. The type of Cyanella pentheri is presumed

lost (Scott 1991) but Zahlbruckner’s (1900) description

is quite clear and we designated an extant specimen to

serve as a neotype.

Plants 100-400 mm high. Conns deep-seated,

25-30 mm diam., tunics of coarsely netted, wiry or

woody fibres, extending in a short or very long fibrous

or papery neck to 100 mm long, pale brown. Basal

leaves (5— )9— 1 7, suberect, often ± twisted or coiled api-

cally, linear, 60-150 * 1 — 4(— 5 ) mm, attenuate, canalic-

ulate-involute, with prominent midrib and ribbed veins

abaxially, firm-textured, glabrous or veins puberulous

abaxially, margins straight or ± undulate or crispulate,

conspicuously ciliate in basal parts only with shaggy

hairs 2. 0-3.0 mm long but glabrous distally; upper

cataphyll prominent, with crispulate margins villous

as in leaves, strongly flushed purple towards edge and

along veins, thus fenestrate, sometimes also villous on

veins. Inflorescence a moderately dense raceme up to

25-flowered, simple or up to 4-branched, lower flow-

ers 0.3-0. 6 x pedicel length apart; pedicels genicu-

late, horizontal in basal half or 2/3 then abruptly flexed

upwards at ± right angles, mostly 20-30 mm long;

bracteoles mostly inserted between lower and upper

third, rarely sub-basal. Flowers facing outwards, white

to pale mauve or blue, fragrant; tepals spreading, ovate,

7 11 x 3-4 mm, apiculate. Stamens dimorphic, 5+1;

filaments of posterior cluster 2. 0-3. 5 mm long, connate

± 1/3 to 2/3 into tube 1.0-1. 5 mm long, yellow, anthers

1.5-2. 5 mm long, yellow; anterior stamen with filament

± 1 mm long, connate to upper cluster for ± half length,

anther 2. 5-3. 5 mm long, yellow. Ovary half-inferior;

style medially defiexed, 3-4 mm long, not extending

beyond anthers. Capsules erect on geniculate pedicels,

subglobose, 5-6 mm diam., 3-lobed and retuse. Seeds

unknown. Flowering time', late Aug.-early Oct.

Distribution and ecology'. Cyanella pentheri has a

restricted distribution through the middle reaches of

the Olifants River Valley from north of Citrusdal to

Klawer, extending along the foot of the Gifberg onto the

Bokkeveld Escarpment, and inland to the Bidouw and

Doring River Valleys (Figure 10). Plants favour rocky

FIGURE 10. — Distribution of Cyanella pentheri.

places, often sandstone, mainly in arid fynbos.

Diagnosis and relationships : Cyanella pentheri

has mostly been treated as conspecific with C. hva-

cinthoides, essentially because of the confusion between

true C. pentheri and what we regard as pubescent forms

of C. hvacinthoides. Florally, the two species are cer-

tainly alike in their moderately dense, branched racemes

of spreading, white or mauve to blue flowers with 5 + 1

arrangement of stamens with the filaments connate for ±

half their length or more, but they differ significantly in

their foliage. The leaves of C. pentheri are consistently

linear and canaliculate-involute, mostly l^t mm wide,

with margins that are conspicuously ciliate only towards

the base with long, shaggy hairs 2. 0-3.0 mm long. Simi-

lar, long cilia also fringe the upper cataphyll, which is

funnel-shaped, and strikingly pigmented with deep pur-

ple along the edges and veins, giving it a characteristic

fenestrate appearance. The leaves of C. hvacinthoides,

in contrast, are mostly lanceolate and 4-15 mm wide,

rarely narrower, with margins either smooth or ciliolate-

pubescent along their entire length, with much shorter

hairs 0. 2-1.0 mm long, and the upper cataphyll is usu-

ally unmarked. Pubescent forms of C. hvacinthoides

from central Namaqualand have leaves that are variously

puberulous to villous, but never with the long cilia char-

acteristic of C. pentheri.

The variation in vestiture in C. hvacinthoides is not

correlated with leaf shape, unlike the situation in C. pen-

theri. This is compelling evidence that C. pentheri repre-

sents a distinct genotype, which is further corroborated

by the fact that the vegetative differences between the

two species are maintained wherever the two have been

collected together, notably north of Klawer at Zandkraal

Farm (Barker 5648 vs Barker 5662), Welbedacht Farm

in the Bidouw Valley (Middelmost 1741 vs Johnson

537) and Clanwilliam (Perry 3526 vs Barker 4771). We

have examined both taxa growing together just outside

Clanwilliam ourselves and at none of these localities

have we found intermediates between them.

Vernacular name', klipraap.

Representative specimens

NORTHERN CAPE.— 3119 (Calvinia): Nieuwoudtville, Wil-

lems River Farm, (-AC), Sept, [without year], Leipoldt 789 (NBG);

Nieuwoudtville, hills near Groenrivier, (-AC), Sept, [without year],

Leipoldt 790 (NBG).

Bothalia 42,1 (2012)

35

WESTERN CAPE. — 3118 (Vanrhynsdorp): Zandkraal, (-DA),

7 Sept. 1949, Barker 5648 (NBG). 3119 (Calvinia): foot of Van

Rliyn’s Pass, (-AC), 22 Aug. 1950, Barker 6447 (NBG, SAM). 3218

(Clanwilliam): intersection of Citrusdal road with Klawer-Clanwil-

liam road, (-BB), 14 Sept. 1985, Scott 25 (NBG); Olifants Dam, (—

BB), 14 Sept. 1947, Barker 4771 (NBG, SAM); Clanwilliam, near

dam, (-BB), Sept. 1947, Lewis 2400 (SAM); Botterkloof Pass SE of

Kameelberg, (-CD), 9 Sept. 1983, Oliver 8052 (NBG); Kanolvlei, (—

DD), 6 Sept. 1951, Barker 7448 (NBG). 3219 (Wuppertal): Diamond

Drift, Biedouw River between Pakhuis and Wuppertal, (-AA), Aug.

1939, Leipoldt 3114 (PRE); Biedouw Valley, 2 km along road to Doom

River, (-AA), 22 Aug. 1993, Goldblatt & Manning 9632 (MO); road

to Algeria, (-AC), 6 Sept. 1980, Le Roux 2813 (NBG). Without precise

locality'. Olifantsrivier, Dec. [without year], Zeyher s.n. SAM20551

(SAM).

7. Cyanella aquatica Oberm. ex G. Scott in South

African Journal of Botany 57: 40 (1991). Type: South

Africa, [Northern Cape], Nieuwoudtville, Kdipkoppies,

21 Sept. 1986, G. Scott 66 (NBG, holo.!; PRE, iso.!).

Plants up to 500 mm high. Corms shallow or mod-

erately deep-seated, 20 mm diam., tunics of papery or

leathery layers, not extending into neck, pale whitish

brown. Basal leaves ± 5 or 6, suberect, linear-lanceolate

or narrowly lanceolate, 200-350 x 10-15 mm, attenu-

ate, canaliculate with prominent midrib abaxially, soft-

textured, bright green, glabrous. Inflorescence a lax

raceme, up to 15-flowered, simple or with 1-3 branches

from near base, lower flowers 1. 5-3.0 x pedicel length

apart; pedicels geniculate, horizontal in basal 1/2 to 2/3

then abruptly flexed upwards at ± right angles, mostly

15-20 mm long but lowermost up to 30 mm long;

bracteoles basal or sub-basal. Flowers facing outwards,

bright orange, veined green on reverse, fragrant; tepals

spreading, outer ovate, 9-12 x 3^4 mm, recurved-apicu-

late, inner short-clawed, claw ± 1 mm long, blade ovate,

9-11 x 4-5 mm. Stamens dimorphic, 5 + 1; filaments

of posterior cluster 2. 0-2. 5 mm long, outer slightly

longer than inner, connate ± halfway or more into tube

1. 5-2.0 mm long, yellow, anthers 1. 5-2.0 mm long, yel-

low; anterior stamen with filament ± 1 mm long, connate

to upper cluster for most of length, anther ± 3 mm long,

yellow. Ovary half-inferior; style medially deflexed.

± 3 mm long, not extending beyond anthers. Capsules

erect on geniculate pedicels, subglobose, ±8x6 mm,

3-lobed and retuse. Seeds ovoid, 3.0 x 2.5 mm, rugulose.

Chromosome number. 2n = 24 (Ornduff 1979: as ‘Klip-

koppies’ population of C. hyacinthoides). Flowering

time : mid-Sept.-early Nov. Figure 61, J.

Distribution and ecology, known originally only from

the rocky outcrops immediately east of Nieuwoudtville,

inland of the edge of the Bokkeveld Escarpment,

C. aquatica has recently been collected significantly

further inland just south of Calvinia, but is still the most

local one of species in the genus (Figure 1 1). Plants are

restricted to dolerite dykes, along watercourses or drain-

age lines where the soil becomes seasonally waterlogged

during the winter months.

Diagnosis and relationships', distinguished by the

lax, sparsely branched racemes, ± basal bracteoles on

sharply sigmoid pedicels, and bright orange flowers.

Cyanella aquatica is superficially similar to C. hya-

cinthoides, which also has a 5 + 1 arrangement of sta-

mens with the filaments connate for ± half their length

or more, but which differs in its fibrous corm tunics and

dense racemes of white or pink to blue flowers with

the bracteoles usually inserted near the middle of the

pedicels or above, only rarely near the base. The two

taxa are ecologically separated, with C. hyacinthoides

favouring better drained, sandy or gritty soils. In peri-

anth colour, C. aquatica might be confused with yellow-

flowered C. lutea, but that species has suberect pedicels

with the bracteoles inserted ± midway along, filaments

that are ± free to the base, and a laterally deflexed style.

Cyanella lutea is also ecologically separated, favouring

fine-grained clay soils in renosterveld or drier karroid

vegetation.

Representative specimens

NORTHERN CAPE. — 3119 (Calvinia): Niewoudtville, Klipkop-

pies, (-AC), 15 Sept. 1961, Barker 9531 (BOL, NBG, PRE); 5 Nov.

1962, Barker 9764 (NBG); trek path E of Nieuwoudtville near Cal-

vinia road, (-AC), 29 Oct. 1996, Goldblatt dt Manning 10581 A (MO);

Farm Driefontein. SW of Calvinia, SW slopes of Driefontein-se-Berg,

in watercourse among dolerite rocks, (-DA), 23 Sept. 2009, Goldblatt

& Manning 13419 (NBG, MO).

Series Luteae J.C. Manning & Goldblatt, ser. nov.

Flowers ± enantiomorphic; pedicels suberect; peri-

anth white, yellow, or pink. Stamens', filaments free,

anthers sometimes spotted or maculate. Ovaiy : style

and lower anther weakly or strongly flexed sideways in

opposite directions. Type species: Cyanella lutea L.f.

8. Cyanella lutea L.f., Supplementum plantarum: 201

(1782). Type: South Africa, without precise locality or

date, Sparrman s.n. Herb. Linn. 430.1 (LINN, holo.!).

Plants (120-) 150-350 mm high. Corms moderately

to deep-seated, 20-25 mm diam., tunics of coarsely net-

ted, fibrous, leathery or woody fibres, sometimes con-

nate below into claws, extending shortly in a neck to

30 mm long, rarely into a fibrous neck up to 100 mm

long, brown. Basal leaves 4-15 mm, suberect or spread-

ing, linear-hemiterete to lanceolate, 30-200(-250) x

2 — 1 5( — 20) mm, acute to attenuate, leathery, plane or

canaliculate, glabrous, margin smooth or ciliolate-

scabridulous. Inflorescence a moderate or dense raceme

up to 15-flowered, with 1-3 branches congested near

base, thus emerging from among leaves, rarely with

accessory branchlets and thus paniculate, lower pedicels

0.2-0. 8 x their length apart; pedicels suberect, rarely

arcuate or almost geniculate, 1 5— 30(— 50) mm long;

bracteoles mostly inserted between ± halfway and upper

third, sometimes in basal third or sub-basal. Flowers ±

36

Bothalia 42,1 (2012)

enantiomorphic, facing outwards, yellow or pink to pur-

ple, usually flushed darker on reverse, with dark veins,

fragrant; tepals spreading, outer oblong-elliptic, 1 0—1 5(—

18) x 2—4 mm, apiculate, inner elliptic-ovate, 1 0—1 5( —

1 8) x 3-7 mm, acute, narrowed basally or very short-

clawed. Stamens dimorphic, 5 + 1 ; filaments of posterior

cluster 2.5 — 4.0 mm long, connate only at extreme base,

± linear, yellow, anthers 2—4 mm long, yellow, usually

finely spotted black or maroon; anterior stamen with fil-

ament deflexed ± laterally, 4-5 mm long, linear, connate

to upper cluster at extreme base only, anther 4-7 mm

long, thus ± twice as large as upper, yellow, brown, or

mauve. Ovary half inferior; style ± laterally deflexed to

left or right opposite lower stamen, 6-10 mm long, not

FIGURE 12. — Cyanella lutea: A, flowering plant; B, half-flower; C,

capsule; D, seed. Scale bar: A-C, 10 mm; D, 2 mm. Artist: John

Manning.

extending beyond lower anther. Capsules erect, sub-

globose-retuse, 6-8 mm diam., 3-lobed. Seeds ovoid,

± 2 mm diam.. rugulose. Chromosome number. 2n = 24

(subsp. lutea : Omduff 1979). Flowering time : mainly

Aug.-Nov. Figures 6K, L; 12.

Distribution and ecologyn the most widely distributed

species in the genus, C. lutea , extends through the win-

ter rainfall region of southern Namibia and South Africa

and around the interior margin of the central plateau but

is absent from the central and Great Karoo (Figure 13).

Pink-flowered plants, often with narrower leaves,

have been distinguished taxonomically several times,

but differ consistently from the typical yellow-flowered

form only in perianth colour. Baker (1871) initially rec-

ognized var. rosea from the Eastern Cape but subse-

quently (Baker 1880) changed his mind. This decision

was followed by Scott (1991). However, the two colour

morphs are geographically segregated: pink-flowered

plants are recorded from the edges of the winter rainfall

region into interior southern Africa, typically in sandy

soils; and yellow-flowered plants are restricted to the

southwestern Cape and nearby, on clay soils. We accord-

ingly treat them here as distinct subspecies.

Diagnosis and relationships : distinguished from other

members of sect. Cyanella by its racemes of pink or yel-

low, ± enantiostylous flowers with almost free filaments,

connate only at the extreme base, and the lower anther

± twice as large as the upper anthers. Yellow-flowered

plants are readily recognized by their colour but pink-

flowered plants could be confused with C. hyacinthoides

around Springbok in Namaqualand, where both occur.

Cyanella hyacinthoides is recognized by its partially

connate upper filament cluster, with the lower anther

mostly less than twice as long as the upper, and by its

spreading-geniculate pedicels. Subspecies rosea has also

been confused with C. ramosissima (sect. Trigella ), but

the arrangement of the stamens is quite different in the

two species.

Key to subspecies

la Leaves mostly lanceolate, (2— )5— 1 5(— 20) mm wide; perianth

pale to golden yellow, rarely orange, often flushed reddish

on reverse; plants from southwestern Cape, from Nieu-

woudtville to Uitenhage 8a. subsp. lutea

lb Leaves linear to linear-lanceolate, 2—1 0(— 1 2 ) mm wide; peri-

160 18° 20" 22° 24° 26° 28°

FIGURE 13. — Distribution of Cyanella lutea subsp. lutea, •; C. lutea

subsp. rosea, o.

Bothalia 42,1 (2012)

37

antli pink to purple; plants from southern Namibia and

northern Namaqualand across interior of South Africa into

Eastern Cape as far as Humansdorp 8b. subsp. rosea

8a. subsp. lutea

C. racemosa Schinz: 394 (1895). Type: South Africa,

[Western Cape], in arenosis [sandy] Camp Ground pro-

pre [proper]. Cape Town, 12 June 1892, Schlechter 839

(Z, holo.; PRE, iso.!).

C. lutea forma angustior Zahlbr.: 27 (1900). Type:

South Africa, [Western Cape], Caledon, Oct. [without

year], Penther 494 (W, holo.f).

Leaves 4-10, mostly lanceolate, rarely linear, (2-)5—

1 5(— 20) mm wide. Flowers pale to golden yellow, often

flushed reddish on reverse or tinged orange.

Distribution: endemic to winter rainfall South Africa,

where it has been recorded from the Bokkeveld Escarp-

ment and southern Roggeveld to the Cape Flats and

Bredasdorp in the south and eastwards through the Lit-

tle Karoo to Uitenhage. The subspecies is essentially

restricted to renosterveld shrubland on fine-grained clay

or laterite soils, rarely on stony limestone flats.

Diagnosis: recognized by the yellow perianth, often

flushed reddish on the reverse and thus with an orange

tinge, and the typically lanceolate leaves, mostly 5 — 1 5( —

20) mm wide, rarely narrower and grass-like. Flower-

ing time: mainly Sept.-Oct. but to Nov. in the southern

Cape.

Representative specimens

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville Reserve,

(-AC), 12 Oct. 1983, Perry & Snijman 2372 (NBG, PRE); Oor-

logskloof Nature Reserve, 15 km SW of Nieuwoudtville, (-AC), 14

Oct. 1996, Pretorius 388 (NBG); Lokenburg, (-AC), 23 Aug. 1980,

Van Berkel 207 (MO). 3220 (Sutherland): Roggeveld Escarpment,

Ouberg Pass, (-AD), 6 Sept. 2006, Rosch HR538 (NBG); Sutherland,

Houthoek, (-CA), 13 Sept. 1971, Hanekom 1575 (PRE); Koedoesberg,

(-CC), 1 Sept. 1973, Oliver 4378 (NBG).

WESTERN CAPE-3218 (Clanwilliam): Farm Nurust, about 8

miles [13 km] N of Porterville, (-DD), 22 Sept. 1966, Loubser 2107

(NBG). 3318 (Cape Town): Bobbejaanberg above Groene Kloof

[near Mamre], (-AD), Oct., Ecklon & Zeyher 269 (MO); N of Tiger-

berg [Tygerberg], (-CC), 20 Sept. 1947, Barker 4808 (NBG); Stel-

lenbosch, Elsenburg, (DD), 5 Oct. 1938, Penfold 153 (NBG). 3319

(Worcester): Saron, (-AA), Oct. 1896, Schlechter 10633 (MO); Tul-

bagh, (-AC), Oct. 1920, Marloth 9939 (NBG); 5 miles [8 km] from

Ceres at bottom of Theron’s Pass, (-AD), 11 Nov. 1974, Snijman 9

(NBG); Karoopoort, (-BA), 26 Sept. 1944, Compton 16054 (NBG);

Tanqua Karoo, near Bloukop, (-BD), 22 Sept. 1975, Thompson 2549

(NBG); Karoo Garden, Worcester, (-CB), 11 Sept. 1969, Tarr s.n.

(NBG); Worcester, Langerug Koppie, (-CB), 23 Sept. 1974, Walters

1207 (NBG); Rooihoogte Pass, (-DB), 28 Oct. 1980, Mauve, Reid &

Wikner 197 (NBG). 3320 (Montagu): Laingsburg, Cabidu, (-AB), 28

Sept. 1951, Compton 22890 (NBG); Whitehill, (-BA), 20 Sept. 1943,

Compton 14874 (NBG); S of Ashton, (-CC), 21 Sept. 1941, Barker

2032 (NBG); 14 km E of Montagu, Klipheuwel Farm. (-CC), 16 Oct.

1998, Manning 2195 (NBG). 3321 (Ladismith): Vleiland, N of Klein

Swartberge, (-AC), 10 Oct. 1976, Thompson 3183 (NBG); road to

Waterkloof NW of Ladismith, (-AD), 23 Oct. 1980, Mauve, Reid &

Wikner 105 (NBG); S side of Rooiberg, (-CB), 22 Nov. 1983, Mauve,

Van Wyk & Pare 40 (NBG); Van Wyksdorp, (-DA), 12 Sept. 1983,

Bohnen 8297 (NBG). 3322 (Oudtshoom): Prince Albert route 407

to Klaarstroom, Farm Welgelegen, (-AC), 1 Sept. 2006, Roux 4199

(NBG); George Forest, (-CD), 25 Nov. 1950, Martin 638 (NBG); De

Rust, Ostekloof Farm, (-DA), 28 Sept. 1971, Dahlstrand 2088 (MO,

PRE); Knysna, Barrington, (-DD), 14 Nov. 1949, Barker 6068 (NBG).

3419 (Caledon): Kogelberg State Forest, Remhoogte, (-AA), 25 Oct.

1984, Brits 23 (NBG); Greyton, (-BA), 21 Oct. 1967, Bayliss 4019

(MO, NBG); 5 miles [8 km] NW of Riviersonderend, (-BB), 17 Sept.

1949, Heginbotham 83 (NBG); Swellendam to Stormsvlei, (- BB), 3

Oct. 1974, Goldblatt 2924 (MO); slopes of Kleinberg, ± 3 km NW of

Napier, (-BD), 19 Oct. 1976, Thompson 3206 (NBG, PRE); ± 15 km

NW of Napier, Fairfield Farm, (-BD), 3 Oct. 1994, Kemper IPC644

(NBG); Bredasdorp, Bosheuwel, (-BD), 6 Oct. 1982, Cowling 1882

(NBG). 3420 (Bredasdorp): Kathoek Farm, 30 km E of Bredasdorp,

(-AD), 11 Oct. 1981, Mauve & Hugo 140 (NBG); De Hoop, Potberg

Nature Reserve, (-AD), 12 Oct. 1978, Burgers 1276 (NBG); Swellen-

dam, Bontebok Park, (-AB), 20 Sept. 1965, Grobler 490 (NBG);

Struisbaai, ± 5 km on Bredasdorp road, (-CC), 26 Oct. 1 987, Felling-

ham 1366 (NBG). 3421 (Riversdale): Blombos Road, 8-10 km S of

Riversdale, (-AA), 11 Oct. 1993, Goldblatt & Manning 9792 (NBG);

Reisiesbaan siding, (-AB), 31 Oct. 1979, Bohnen 7051 (NBG); Still

Bay, (-AD), 16 Oct. 1978, Bohnen 4463 (NBG). 3422 (Mossel Bay):

Great Brak, (-AA), 21 Sept. 1959, Lewis 5601 (NBG). 3423 (Knysna):

Plettenberg Bay, (-AB), 21 Nov. 1953, Taylor 4320 (NBG).

EASTERN CAPE. — 3323 (Willowmore): flats between Hotsprings

and Toorwater, (-AC), 5 Oct. 1971, Oliver 3646 (NBG, PRE); Vled-

ermuis area between Fullerton & Heuningklip, (-BA), 14 Sept. 1973,

Oliver 4582 (NBG); Baviaanskloof, Adamskraal, (-BC), 22 Oct.

1999, Desmet 2095 (NBG); Bellvue, ± 4 km from Avontuur, (-CC),

11 Nov. 1978, Botha 2188 (PRE); Suuranysberge, Voelkraal Farm, (—

CC), 1 Oct. 1984, Stirton 10903 (NBG). 3324 (Steytlerville): Kruis-

rivier-Hankey Dist., (-CB), [without date], Manson 297 (NBG); poort

between Patensie and Cambria, (-DA), 11 Sept. 1973, Thompson 1885

(NBG).

8b. subsp. rosea (Eckl. ex Baker) J.C. Manning &

Goldblatt , stat. nov. Cyanella lutea var. rosea Eckl.

ex Baker: t. 259 (1871). Type: South Africa, [Eastern

Cape], Queenstown, 1860, T. Cooper 270 (K, holo.!).

Note: Scott ( 1991 ) was of the opinion that no material

of Cooper’s collection had been preserved and thus lec-

totypified the name against the illustration in Refugium

Botanicum , which was drawn from plants collected and

cultivated by Thomas Cooper. There exists, however, a

specimen at Kew, collected by Cooper in 1 860 at Queen-

stown in the Eastern Cape where this form has since

been re-collected, and labelled with the name Cyanella

rosea. There seems no reason to doubt that it represents

the original collection from which the cultivated plants

were derived. This material, as the holotype, takes prec-

edence over the illustration (McNeil et al. 2006: Art.

9.10 & 9.17). Baker’s (1871 ) citation of the Ecklon man-

uscript name, Cyanella rosea Eckl., which appeared as

a printed label on some herbarium collections, including

Ecklon 255 (NBG), is a clear indication that the correct

author citation for the epithet is Eckl. ex Baker.

C. lineata Burch.: 589 (1812). Type: South Africa,

Bechuanaland [Northern Cape], near Moshowa [Mosha-

weng] River, without exact date [1811-1812], Burchell

2256-2 ( K, holo.!).

C. odoratissima Ker Gawk: t. 1111 (1827). Type:

South Africa, Cape of Good Hope, without precise local-

ity, date or collector, cultivated in Tate’s nursery, Lon-

don, apparently not preserved, illustration in Ker Gawk,

The Botanical Register 13: t. 1111 (1827). [Note: Scott’s

(1991) attribution of the name to Lindley is incorrect, as

John Bellenden Ker [-Gawler] wrote the text for the first

14 volumes (Stafleu & Cowan 1976), and John Lindley

only assumed authorship from vol. 15].

C. lutea var. angustifolia Schinz: 48 (1896). Type:

Namibia, Oas [Huib-Hoch Plateau], Oct. 1891, Fleck

232 (Z, holo.!).

Leaves 6-12, linear-hemiterete to linear-lanceolate.

38

Bothalia 42,1 (2012)

2—1 0(— 1 2) mm wide. Flowers pale to deep pink or pur-

ple. Flowering time : mainly Aug.-Sept. in Namaqualand

and Bushmanland; Oct.-Dec. in the interior and Eastern

Cape.

Distribution', recorded from central Namaqualand

around Springbok and the Huib-Hoch Plateau in south-

ern Namibia, inland through Bushmanland along the

Orange and Vaal Rivers as far as Kuruman in North-

ern Cape and Smithfield in the southern Free State,

thence southwards through the eastern Upper Karoo to

Humansdorp (Figure 13). Plants have been recorded

mainly from sandy, sometimes calcareous, flats in

Nama-Karoo shrubland or drier grassland, in the Kuru-

man area typically beneath small bushes. The subspecies

is relatively poorly documented for such a large range.

Diagnosis : distinguished by its generally narrower,

often grass-like leaves 2-12 mm wide, and its pink

perianth. Plants from Namaqualand-Bushmanland and

southern Namibia are especially distinctive in their very

small stature, numerous, semi-terete leaves, and ± con-

gested inflorescence branching near the base, giving

them a characteristic caespitose appearance.

Representative specimens

FREE STATE. — 3026 (Aliwal North): Smithfield, (-BA), Oct.

[without year or collector], STE12787 (NBG).

NORTHERN CAPE.— 2623 (Morokweng): Vryburg, (-DB), Sept.

1924, Henrici 160 (PRE). 2723 (Kuruman): 36 miles [57.6 km] E-NE

of Van Zylsrus, 2 miles [3 km] N of Kuruman River on Tsabong road,

(-AD), 17 Oct. 1961. Leistner 2886 (PRE). 2818 (Warmbad): 2 miles

[3 km] S of Goodhouse, (-DD), 27 July 1950, Lewis 3003 (SAM),

63739 (PRE); Goodhouse, (-BD), 27 July 1950, Barker 6262 (NBG).

2819 (Ariamsvlei): Augrabies, (-DB), 21 Aug. 1954, Compton 24474

(NBG); Augrabies Falls National Park, (-DB), 22 Aug. 2005, Steyn

759 (NBG, PRE). 2820 (Kakamas); 12 miles [19 km] E of Kakamas,

(-DB), 28 Aug. 1963, Hardy & Rauh 1560 (PRE). 2823 (Griekwa-

stad): Brakfontein, (-CD), 20 Sept. 1988, Saaiman 227 (PRE). 2824

(Kimberley): Kuruman River 16 miles [25.6 km] W of Kuruman-

Gordonia boundary, (-BA), 18 Oct. 1961, Leistner 2893 (PRE). 2917

(Springbok): along Goodhouse road, (-BD), 20 Sept. 1980, Van Ber-

kel 260 (NBG); near Springbok, (-DD), Sept. 1939, Lewis 750 (SAM);

Droedap [SE of Springbok], (-DD), 27 Aug. 1941, Barker 2029

(NBG). 2918 (Gamoep): Aggenys, (-BD), 13 Oct. 1971, Wisura 2264

(NBG). 2922 (Prieska): Prieska, (-DA), [without date], Biyant s.n.

PRE38351 (PRE). 3017 (Hondeklipbaai): Theunis se Dam, 36 km S

of Little Rock Caravan Park on Droedap road, (-BB), 25 Aug. 1977,

Thompson & le Roux 37 (NBG); Droedap, (-BB), 27 Aug. 1941,

Esterhuysen 5894 (PRE). 3023 (Britstown): De Aar. (-DB), 30 Aug.

1895, Solly s.n. PRE38315 (PRE). 3024 (De Aar), Rolfontein Nature

Reserve, Springbok Flats, (-BB), 9 Sept. 1982, Coetzee s.n. PRE61030

(PRE).

WESTERN CAPE. — 3223 (Rietbron): 20 km from Farm Riet-

bron on road to Murraysburg, (-BA), 13 Oct. 1983, Relief & Reid 521

(PRE).

EASTERN CAPE. — 3126 (Queenstown): lower slopes, (-DD),

1893, Galpin 1568 (PRE). 3127 (Lady Frere): Little Bushy near

Cala, (-DA), Dec. 1910, Royffe s.n. TM25721 (PRE). 3225 (Somer-

set East): Mountain Zebra National Park, (-AD), 4 Oct. 1 979, Du Toil

155 (PRE); Addo National Park, (-BC), Nov. 1962, Liebenberg 6620

(PRE). 3226 (Fort Beaufort); Bushman’s River Mouth, (-DB), 2 Dec.

1941, Barker 2034 (NBG). 3227 (Stutterheim): Queenstown, Bram

Neck, (-AA), 28 Oct. 1946, Thorns s.n. (NBG); between Fish River

and Governor’s Kop, (-BD), 16 Oct. 1961, Batten 1-P1.83 (NBG).

3325 (Port Elizabeth): Kommadagga, (-BB), 27 Nov. 1973, Bayliss

6199 (MO); Vanstadensberg, (-CC), Dec. [without year], MacOwan

1086 (SAM); near Zwartkop River, (-DC), Nov. [without year], Eck-

lon 255 (NBG, SAM). 3424 (Humansdorp): Humansdorp, (-BB), 14

Oct. 1928, Gillen 2397 (NBG).

9. Cyanella alba L.f, Supplementum plantarum: 201

(1782). Pharetrella alba (L.f.) Salisb.: 47 (1866). Type:

South Africa, without precise locality or date, Thunberg

s.n. Herb. Linn. 430.4 [LINN, lecto.!, designated by

Scott: 46 ( 1991 )].

Plants 80-200 mm high. Conns deep-seated, 15—

25 mm diam., tunics of coarsely netted fibres, extend-

ing into neck up to 50 mm long, pale brown. Basal

leaves ± 10-20, erect, filiform to linear, (40-)50-100 x

0. 5-3.0 mm, attenuate, leathery, bright green, glabrous.

Inflorescence a highly congested, simple raceme such

that flowers apparently solitary among leaves; pedicels

suberect, (80-) 100-200 mm long; bracteoles either sub-

basal or inserted in upper half. Flowers enantiomorphic,

facing outwards, white or pale pink or pale yellow, fra-

grant; tepals spreading, cucculate, outer elliptic, 12-20

x 5-7 mm, recurved-apiculate, inner ovate, 12-20 x

7 — 12(— 1 5) mm, acute, narrowed basally or short-clawed,

claw up to 1 mm long. Stamens weakly dimorphic or

submonomorphic, 5 + 1; filaments of posterior clus-

ter 3-5 mm long, connate only at extreme base, awl-

shaped, white, anthers 3. 5-5. 5 mm long, yellow, some-

times marked with black spot on upper surface near

base, sometimes cohering; anterior stamen with filament

deflexed laterally, 3-4 mm long, awl-shaped, connate

to upper cluster at extreme base, anther 4-6 mm long,

yellow. Ovary half-inferior; style laterally deflexed

opposite lower stamen, 7-9 mm long, not extending

beyond lower anther. Capsules erect, ellipsoid, 13-15 x

7-8 mm, 3-lobed. Seeds ovoid, ± 2 mm diam., rugulose.

Chromosome number. 2n = 24 (subsp. flavescens: Orn-

duff 1979). Flowering time : (late Aug.-)mid-Sept.-mid-

Oct.(Nov.). Figures 6M, N; 14.

Distribution and ecology, the species has a scattered

distribution along the western mountains in Western

Cape, where it is known from the Bokkeveld Escarp-

ment, the Cedarberg and Olifants River Mtns, and the

base of the Swartruggens (Figure 15). These three areas

of occurrence correspond to the distribution of the three

subspecies that we recognize. Cyanella alba is restricted

to clay soils in renosterveld shrubland.

Diagnosis and relationships', one of the easiest spe-

cies to identify on account of its highly congested inflo-

rescence axis with extremely elongate pedicels, the

flowers thus apparently borne on 1 -flowered peduncles

rather than in a raceme. The raceme is never branched,

and up to a maximum of nine flowers are produced,

thus very much fewer than in other species. The flow-

ers are strongly enantiostylous, and either white to pale

pink with uniformly yellow anthers, or pale yellow with

maculate anthers. These colour morphs, which are geo-

graphically segregated, correlate with the position of

the bracteole on the pedicels, and we recognize them as

three subspecies. The large, ellipsoid capsule, 13-15 mm

long, is unique in sect. Cyanella, resembling those of

C. cygnea and C. orchidiformis in sect. Trigel/a.

Key to subspecies

la Leaves filiform, 0.5-1. 5 mm diam.; flowers white; bracteoles

subbasal, not readily visible among leaves and thus appar-

ently absent 9c. subsp. minor

lb Leaves linear-filiform, 1-3 mm wide; flowers white or yel-

low; bracteoles inserted in distal half of pedicel, thus

Bothalia 42,1 (2012)

FIGURE 14. — Cyanella alba : A, subsp. alba , showing distal bracteole;

B. subsp. minor, showing subbasal bracteole. Scale bar: 10 mm.

Artist: John Manning.

clearly present:

2a Flowers 3-9 per plant, white or pale pink; anthers uni-

formly yellow 9a. subsp. alba

2b Flowers 1-4 per plant, pale yellow or outer tepals white;

upper anthers marked with black blotch adaxially near

base 9b. subsp. flavescens

9a. subsp. alba

Plants (80-) 100-200 mm high. Leaves linear,

1-3 mm wide. Inflorescence 3-9-flowered; pedicels with

bracteole in distal half. Flowers white to pale pink. Sta-

mens: anthers uniformly yellow. Figure 14A.

Distribution : endemic to the Bokkeveld Escarpment,

from just north of Nieuwoudtville southward to Men-

zieskraal near Botterkloof (Figure 15).

Diagnosis: characterized by the long pedicels, (80-

) 100-200 mm long, with the bracteole inserted between

one third and three-quarters along, and white or pale

39

pink flowers flushed darker pink on the reverse. The

anthers are uniformly yellow, with the upper cluster free

or coherent. Plants are often well grown, producing 3-9

flowers. The position of the bracteoles in the distal half

of the pedicels distinguishes subsp. alba from subsp.

minor from the Tanqua Basin to the south, which has

similar flowers but subbasal bracteoles.

Representative specimens

NORTHERN CAPE.— 3119 (Calvinia): N of Nieu-

woudtville, Grasberg Farm, (-AC), 16 Sept. 1961,

Barker 9457 (NBG); Nieuwoudtville Reserve, (-AC), 8

Sept. 1983, Perry & Snijman 2351 (NBG); ± 15 km S of

Nieuwoudtville, Matjiesfontein Farm, (-AC), 13 Sept.

1976, Thompson 2902 (NBG); Lokenberg Farm, (-CA),

26 Sept. 1933, Acocks 17263 (PRE); 4 Sept. 1985, Snij-

man 905 (NBG); Menzieskraal Farm, (-CA), 29 Sept.

1933, Markotter s.n. (NBG).

9b. subsp. flavescens J.C. Manning in Manning et al.

in Bothalia 35: 119 (2005). Type: South Africa, West-

ern Cape, Biedouwberg, 26 Aug. 1 896, Schlechter 8686

(SAM, holo.!, BOL!, PRE!, iso.).

Plants 120-200 mm high. Leaves linear-filiform,

1-2 mm wide. Inflorescence I -4-flowered; pedicels

with bracteole in distal 1/2. Flowers pale yellow or outer

tepals white. Stamens: anthers yellow, upper five coher-

ent and maculate with dark blotch on upper side near

base.

Distribution: restricted to the northern Cedarberg and

Olifants River Valley, between Clanwilliam and Wup-

pertal, and especially common in the Biedouw River

Valley (Figure 1 5).

Diagnosis: a very distinctive taxon recognized by its

pale yellow flowers (sometimes the outer tepals white)

with the upper anthers coherent and marked on the upper

side with a black blotch near the base. Up to four flow-

ers are produced per plant.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): Clanwilliam, (-BB), 4

Aug. 1896, Schlechter 8405 (BOL, PRE); 10 km S of Clanwilliam, (—

BB), 12 Sept. 1997, Goldblatt & Manning 10741 (MO, NBG). 3219

(Wuppertal): Biedouw Mtn, (-AA), 20 Sept. 1937, Lewis s.n. (NBG);

bottom of hill to Biedouw Valley, (-AA), 9 Aug. 1984, Perry 3145

FIGURE 15. — Distribution of Cyanella alba subsp. alba, •; subsp. fla-

vescens, o; subsp. minor, A .

40

Bothalia 42,1 (2012)

(NBG); Farm Welbedacht, (-AA), 20 Sept. 1937, Barker 283 (NBG);

Koudeberg near Wuppertal, (-AA), 4 Oct. 1897, Bolus 9095 (NBG);

Citadel Kop, (-AA), 7 Sept. 1953, Compton 24237 (NBG); near Wup-

pertal, (-AA), 28 Aug. 1951, Martin 811 (NBG).

9c. subsp. minor J.C. Manning in Manning el al. in

Bothalia 35: 119 (2005). Type: South Africa, Western

Cape, Karoopoort, 27 Sept. 1944, Barker 3024 (NBG,

holo.!).

Plants 80-150 mm high. Leaves filiform, 0.5-1. 5 mm

wide. Inflorescence 1-3-flowered; pedicels with bracte-

ole subbasal. Flowers white to pale pink with darker

pink on reverse. Stamens: anthers uniformly yellow. Fig-

ure 14B.

Distribution : highly localized and known only from

just north of Karoopoort in the southern Tanqua Karoo

basin (Figure 15).

Diagnosis : distinguished from the typical subspecies,

which has similar white or pale pink flowers and uni-

formly yellow anthers, by the shorter pedicels, mostly

< 100 mm (rarely up to 150 mm long) with the bracte-

oles sub-basal and thus difficult to distinguish from the

leaves. This led Manning et al. (2005) to conclude that

bracteoles were absent, and we were only able to estab-

lish the true situation after having the opportunity of dis-

secting live plants. The plants are typically small in stat-

ure, with only 1-3 flowers per plant.

Representative specimens

WESTERN CAPE. — 3319 (Worcester): Karoopoort, (-BA), 19

Sept. 1938, Levyns 6236 (BOL); Tanqua Karoo N of Karoopoort, (—

BA), 9 Sept. 2007, Goldblatt <6 Porter 12970 (NBG); 13 Sept. 2009,

Goldblatt, Manning & Porter 12970 (MO, NBG).

EXCLUDED SPECIES

Walleria paniculata Fritsch: 493 ( 1896). Type: Mada-

gascar, Ins. St Marie, without date, Paulay s.n. (GZU,

holo.) = Dianella ensifolia (L.) DC. (Hemerocalli-

daceae) (Perrier de la Bathie 1938).

ACKNOWLEDGEMENTS

Our sincere thanks to Anne-Lise Fourie, Flead Librar-

ian at the South African National Biodiversity Institute,

and Mary Stiffler, Research Librarian at the Missouri

Botanical Garden, for their assistance in locating nec-

essary references. This is the last of our contributions

edited by Beverley Momberg before her retirement, and

we are deeply grateful to her for her years of sterling

service.

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LINNAEUS, C. 1754. Genera plantarum, edn 5. Salvius, Stockholm.

LINNAEUS, C. 1759. Svstema naturae , edn 10. Salvius, Stockholm.

LINNAEUS, C. (fil.). 1782 [‘1781’]. Supplementum plantarum. Braun-

schweig, Uppsala.

MANNING, J.C. & GOLDBLATT, P. 2007. Trachyandra arenicola

and T. montana (Asphodelaceae), two new species from South

Africa. Bothalia 37: 26-31.

MANNING, J.C., GOLDBLATT, P. & BATTEN, A. 2001. Walleria

gracilis. Flowering Plants of Africa 57: 44-47.

MANNING, J.C., FOREST. F. & MANNHE1MER, C.A. 2005. Eremio-

lirion, a new genus of southern African Tecophilaeaceae, and

taxonomic notes on Cyanella alba. Bothalia 35: 115-120.

McNEILL, J„ BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,

HAWKSWORTH, D.L., MARHOLD, K„ NICOLSON, D.H..

PRADO, J„ SILVA, PC., SKOG, J.E., WIERSMA, J.H. & TUR-

LAND, N.J. 2006. International Code of Botanical Nomencla-

ture (Vienna Code) adopted by the seventeenth International

Botanical Congress, Vienna, Austria, July 2005. Gantner, Liech-

tenstein. [Regnum Vegetabile 146],

ORNDUFF, R. 1979. Chromosome numbers in Cyanella (Tecophilae-

aceae). Annals of the Missouri Botanical Garden 66: 581-583.

PAX, F. & HOFFMANN, K. 1930. Amaryllidaceae. In A. Engler & K.

Prantl, Die naturlichen Pflanzenfamilien , edn 2, 15a: 39 1 — 430.

Engelman, Leipzig.

PERRIER DE LA BATHIE, H. 1938. Liliacees. In H. Humbert, Flore

de Madagascar. Gouvemement General de Madagascar, Tanan-

arive.

PHILLIPS, E.P. 1951. The genera of South African flowering plants.

Memoirs of the Botanical Survey of South Africa No. 25. Depart-

ment of Agriculture, Pretoria.

SALISBURY, R.A. 1 796. Prodromus stirpium in horto ad Chapel Aller-

ton vigentium. Hooker, London.

SALISBURY, R.A. 1866. The genera of plants. Van Voorst, London.

Bothalia 42,1 (2012)

41

SCHINZ, H. 1895. Beitrage zur K.enntnis der alrikanischen Flora 111.

Amaryllidaceae. Bulletin de l Herbier Boissier 3, ser. 1 : 394, 395.

SCHINZ, H. 1896. Die Pflanzenwelt Deutsch-Sudwest-Afrikas. Bulle-

tin de I 'Herbier Boissier 4, ser. l,App. Ill: 1-57.

SCHINZ, FI. 1902. Beitrage zur Kenntnis der alrikanischen Flora XIV.

Haemodoraceae. Bulletin de I 'Herbier Boissier 2, ser. 2: 943, 944.

SCOTT, G. 1991. A revision of Cyanella (Tecophilaeaeceae) excluding

C. amboensis. South African Journal of Botany 57: 34-54.

SIMPSON, M.G. & RUDALL, P.J. 1998. Tecophilaeaceae. In K.

Kubitzki, The families and genera of vascular plants III. Flow-

ering plants — monocotyledons: 429^436.

SNIJMAN, D. 1984. A revision of the genus Haemanthus L. (Ama-

ryllidaceae). Journal of South African Botany , Suppl. vol. 12.

National Botanical Gardens, Cape Town.

STAFLEU, F.A. & COWAN, R.S. 1976. Taxonomic literature 1: A-G.

Bonn, Scheltema & Holkema, Utrecht.

ZAHLBRUCKNER, A. 1902. Plantae pentherianae. Annalen des kai-

serlichen naturhistorischen Museums 15: 1-73, t. I-IV.

Bothalia 42,1:43-66 (2012)

Notes on African plants

VARIOUS AUTHORS

HYACINTHACEAE

DRIM1A ECHINOSTACHYA, CORRECTING A NOMENCLATURAL OVERSIGHT

Crouch et al. (2010) reassessed the distribution and

correct identification of a bulb species recently noted in

the ethnomedicinal plant trade in KwaZulu-Natal. They

used the name Drimia cooperi (Baker) Baker, following

Jessop (1977) and subsequent checklist citations (Reid

1993; Manning & Goldblatt 2003, 2006).

Subsequently, it was discovered that this name is an

illegitimate later homonym (McNeill et al. 2006: Art.

53.1) of Drimia cooperi Baker (1868), which is regarded

as a synonym of Ledebouria concolor (Baker) Jessop

by Jessop (1970). The name Drimia cooperi (Baker)

Bentham ex Baker was actually the combination for

Ornithogalum cooperi Baker (1873) made by Baker

(1897). Transfer of the epithet cooperi to the genus

Drimia is illegitimate because the resulting combination

is preoccupied by Drimia cooperi Baker ( 1868). Accord-

ing to the rules of Botanical Nomenclature (McNeill

et al. 2006), the next earliest available epithet must be

used. Crouch et al. (2010) determined that the holotype

of Urginea echinostachya Baker (1897) is conspecific

with Ornithogalum cooperi Baker (1873), and Urginea

echinostachya Baker is therefore the next earliest avail-

able name for the species. The necessary transfer to the

genus Drimia is published below. In the past, D. echi-

nostachya was regarded as a synonym of D. macrocen-

tra (Baker) Jessop (Jessop 1977), despite the conspicu-

ous morphological differences (Crouch et al. 2010).

An argument could be raised concerning the correct

place of transfer of Ornithogalum cooperi Baker (1873)

to Drimia : Baker (1897) attributed the new combination

for this taxon to Bentham (in Bentham & Hooker 1883).

However, consideration of Bentham & Hooker (1883)

reveals that Bentham did not actually use the name

Drimia cooperi , but simply remarked that Ornithoga-

lum cooperi Baker appeared to belong to Drimia. This

cannot be interpreted as a valid combination under the

present nomenclatural rules (McNeill et al. 2006: Art.

33.1, Ex. 2). Herbarium acronyms follow Holmgren et.

al. (1990).

Drimia echinostachya (Baker) Eggli & N.R. Crouch,

comb. nov. Urginea echinostachya Baker, Flora capen-

sis 6: 468 (1897). Type: Natal [KwaZulu-Natal], Inanda,

J.M. Wood 276 (K, holo.l; NH, iso!).

Ornithogalum cooperi Baker: 284 (1873), syn. nov.;

Bentham: 808 (1883). Drimia cooperi (Baker) Bentham

ex Baker: 443 (1897), nom. inval. (Art. 33.1) et illegit.

(Art. 53.1), non D. cooperi Baker: t. 18 (1868) [= Lede-

bouria concolor (Baker) Jessop]. Type: Cape [Eastern

Cape], ‘ad oram orientalis’, Barber s.n. (TCD, lecto.,

designated by Jessop: 287 (1977);-K, photo.!).

ACKNOWLEDGEMENTS

The staff of the Mary Gunn Library (South African

National Botanical Institute (SANBI), Pretoria) facili-

tated access to literature, and Dr H. Glen (SANBI) is

thanked for helpful discussions and Latin translations.

REFERENCES

BAKER, J.G. 1 868. Drimia cooperi. Saunders Refugium Bolanicum 1 :

t. 18.

BAKER, J.G. 1873. Revision of the genera and species of Scilleae

and Chlorogaleae. Journal of the Linnean Society, Botany 13:

209-292.

BAKER, J.G. 1897. Order CXXXVII. Liliaceae. Flora capensis 6:

253-528. Reeve, London.

BENTHAM, G. & HOOKER, J.D. 1 883. Genera plantarum, vol. 3, part

2. Reeve, London.

CROUCH, N.R., WILLIAMS, V.L., EDWARDS, T.J. & BRUETON,

V.J. 2010. Hyacinthaceae. Drimia cooperi in KwaZulu-Natal,

and the ethnomedicinal trade. Bothalia 40: 75-78.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index herbariorum, part 1: The herbaria of the world , edn 8.

New York Botanical Garden. New York.

JESSOR J.P. 1970. Studies in the bulbous Liliaceae: 1. Scilla. Schizo-

carphus and Ledebouria. Journal of South African Botany 36:

233-266.

JESSOP, J.P. 1977. Studies in the bulbous Liliaceae in South Africa: 7.

The taxonomy of Drimia and certain allied genera. Journal of

South African Botany 43: 265-3 1 9.

MANNING, J.C. & GOLDBLATT, P. 2003. Hyacinthaceae. In G. Ger-

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

checklist. Strelitzia 14: 1054—1071.

MANNING, J.C. & GOLDBLATT, P. 2006. Hyacinthaceae. In G. Ger-

mishuizen, N.L. Meyer, Y. Steenkamp & M. Keith, A checklist

of South African plants. Southern African Botanical Diversity

Network Report No. 41: 952-971. SABONET, Pretoria.

MCNEILL, J„ BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,

HAWKSWORTH, D.L., MARHOLD, K„ NICOLSON, D.H.,

PRADO, J., SILVA, PC., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. 2006. International Code of Botanical Nomen-

clature (Vienna Code) adopted by the Seventeenth International

Botanical Congress Vienna, Austria, July 2005. Regnum Vegeta-

ble 146: 1-568. Gantner Verlag, Liechtenstein.

REID, C. 1993. Hyacinthaceae (Part B). In T.H. Arnold & B.C. De Wet,

Plants of southern Africa: names and distribution. Memoirs of

the Botanical Survey of South Africa Flo. 62: 146-1 55. National

Botanical Institute, Pretoria.

*U. EGGLI & * **N.R. CROUCH

* Sukkulenten-Sammlung, Zurich, Mythenquai 88, CH-8002 Zurich,

Switzerland. E-mail: urs.eggli@zuerich.ch.

** Ethnobotany Unit, South African National Biodiversity Insti-

tute, PO. Box 52099, 4007 Berea Road, Durban / School

of Chemistry, University of KwaZulu-Natal, 4041 Durban.

E-mail: n. crouch@sanbi.org. za.

MS. received: 2010-1 1-30.

44

Bothalia 42,1 (2012)

FABACEAE

ANEW SPECIES OF BAUHINIA FROM SOUTHERN MOZAMBIQUE AND THE REINSTATEMENT OF BAUHINIA MACR.4NTHA

Bauhinia burrowsii E.J.D. Schmidt, sp. nov.

TYPE. — Mozambique: Inhambane Province, near

Mapinhane, 22° 13.385' S and 34° 54.347’ E, 4 April

2010, E.J.D. Schmidt 5022 (PRE, holo.; BNRH, Ernst

Schmidt Herbarium, K, LMA, PRU, iso.).

Shrub up to 5 m high; bark smooth and grey, some-

times covered with small lenticels; branches terete,

reddish brown, sparsely puberulous. hairs reddish and

soon lost, becoming grey with age; remnants of older

branchlets form stout, branched thom-like structures.

Leaves alternate, 2-lobed to ± two-thirds of way, lobe

apices rounded, base truncate to rounded; leaf lamina

wider than long, 20-50 x 30-70 mm; venation palmate

with 9 primary veins, veins darker than lamina in dried

specimens, reticulation not prominent; dark green and

hairless above, slightly paler below with white minutely

appressed-puberulous indumentum, virtually invisible

to the naked eye; margin entire; petiole 8-20 mm long.

Inflorescence a terminal minutely appressed-puberulous

raceme, up to 55 mm long, with up to 18 flowers; pedi-

cels 3-10 mm long. Flowers bisexual and pentamerous,

buds ± 45 mm long, upper part (sepals) ± 25 mm long

and linear-lanceolate in outline; hypanthium ± 20 mm

long, finely puberulous. Petals white, very narrowly

elliptic to obovate, 20-33 x 2-4 mm, margin crisped,

outside covered in small basifixed glands (± 2 mm long),

also present in young vegetative parts. Stamens 5, very

slender, 30—40 mm long, white; anthers 2. 5-3. 5 mm

long, dorsifixed close to middle of anther. Stigma 1 .0-

1 .5 mm in diam., peltate. Style 3-6 mm long. Pod dehis-

cent, narrowly oblanceolate to linear-oblong, 60-220

x 17-28 mm, dark brown. Seeds up to 14 per pod,

brown to olive, obovate to subcircular, 7-12 x 5-10 mm.

Figures 1-5.

Diagnostic characters and affinities : the limited dis-

tribution and small flowers with narrow petals are diag-

nostic for B. burrowsii. The closest affinity would be

Bauhinia petersiana Bolle. In flower, the new species

is easily distinguished by the lack of prominent pink

stamens and the generally smaller flowers. The width

of petals in B. petersiana is at least 6 mm, whereas in

B. burrowsii , the maximum is 4 mm. Furthermore, the

anthers in the latter species are dorsifixed close to the

middle of the anther; whereas in B. petersiana , they are

situated more towards one side and are much larger — a

minimum of ± 6 mm (up to 14 mm) long — as opposed

to a maximum of 3.5 mm long in B. burrowsii (Brenan

1967; Coates Palgrave 2002; Schmidt et al. 2007). In

short, the flowers of B. burrowsii are about half the size

of B. petersiana with a different colour and a morpho-

logically different anther attachment (Figure 5). In her-

barium specimens, the leaves of B. burrowsii are gloss-

ier above than those of B. petersiana.

Distribution and habitat', currently only known from

the type locality to the east of Mapinhane and a col-

lection made by Barbosa & Balsinhas on 28 March

1952 [G. Barbosa & A. Balsinhas 5042 (K, photo.!)].

Although collected the same time of the year, the latter

specimen does not have flowers and is therefore easily

mistaken. There is some doubt as to where exactly this

specimen was collected, since it refers to a Vilanculos-

Mabote-Mambone-Ynrrawenfo’ [crossing]. If this refers

to the turn-off at Mapinhane (which seems to be the only

logical conclusion, although the name is not used), this

would be close to the type locality for B. burrowsii. It

can be regarded as endemic to the area. It occurs in open

woodland. This is part of the southern Zanzibar-Inham-

bane regional mosaic (White 1983). Another endemic,

Croton inhambanensis Radcl.-Sm. was described from

the same area. This species was subsequently collected

as far south as Paindane on the coast and B. burrowsii

may also occur as far south as the C. inhambanensis.

Another endemic plant described from the vicinity is

Croton aceroides Radcl.-Sm.

Status'. Bauhinia burrowsii can be described as

locally common where the type specimen was collected.

The area is sparsely populated and the main economic

activity is non-commercial existence agriculture, i.e.

the planting of maize and cassava on a few scattered

small plots. A commercial cattle farm has recently been

established in the area. Thus far, this agriculture has had

a minimal effect on the survival of the plant. Further

studies are needed to establish if plants of the new spe-

cies are indeed threatened. If it does occur further south

in the Paindane area it would be under threat locally

because of the threat of human expansion.

Etymolog)’: the epithet honours John Burrows, for

his unselfish contribution to the knowledge of the plants

of southern Africa. The proposed common name for the

species is the Manyikeni Bauhinia , after Zimbabwe de

Manyikeni , a historical site close to the type locality. A

visit to the site by the author led to the discovery of the

plant.

Other specimens examined

MOZAMBIQUE. — 2234 Mabote: ‘entre o cruzamento de estradas

Vilanculos-Mambone-Maboti e o Maboti, a 8.7 km de cruzamento'

[between the cross-roads Vilanculos-Mambone-Maboti and the 8.7 km

crossing], 28 Mar. 1952, G. Barbosa & A. Balsinhas 5042 (K, photo.!);

near Mapinhane, (-BA) 22° 13.388’S and 34° 54.395'E, 27 Mar. 2009,

E.J.D. Schmidt 4670 (Ernst Schmidt Herbarium, LMA. PRE); ibid., (—

BA) 22° 1 3.37 1'S and 34° 54.370'E, 4 Apr. 2010, E.J.D.Schmidt 5020

(Ernst Schmidt Herbarium, LMA).

THE REINSTATEMENT OF BAUHINIA MACRANTHA

During the research into the species of Bauhinia ,

it became apparent that the current classification of

Bauhinia petersiana Bolle is unsatisfactory. Brummitt &

Ross (1975) treated the differences between the taxa pre-

viously known as B. macrantha Oliv. and B. petersiana ,

and came to the conclusion that they should be treated

as subspecies, namely B. petersiana Bolle subsp. serpae

(Ficalho & Hiern) Brummitt & J.H.Ross and B. petersi-

Bothalia 42,1 (2012)

45

FIGURE 1. — Bauhinia burrowsii, E.J.D. Schmidt 4670 (PRE): A, flowering branchlet; B, fruit. Scale bar: 20 mm. Artist: Carla Grinstad.

46

Bothalia 42,1 (2012)

FIGURE 2. — Bauhinia burrowsii , habit in type locality, April 2010.

Photographer: Ernst Schmidt.

FIGURE 3. — Bauhinia burrowsii, flowers at type locality, March 2009.

Photographer: Ernst Schmidt.

FIGURE 4. — Bauhinia burrowsii, flowers and fruit at type locality,

March 2009. Photographer: Ernst Schmidt.

ana Bolle subsp. petersiana. The name serpae referred

to the previously entity known as B. macrantha and was

a result of an unfortunate decision in Seattle in 1 969 at

the International Code of Botanical Nomenclature meet-

ing (Coetzer & Ross 1977). The subspecies name was

later changed to B. petersiana Bolle subsp. macrantha

(Oliv.) Brummitt & J.H.Ross (Brummitt & Ross 1982).

The main reasons for regarding the taxa as subspecies

and not full species are that their distributions are almost

entirely allopatric and that there is, according to the

authors, sufficient overlap of characteristics to justify it.

An allopatric distribution may be an indication that

two entities are geographical different races of the same

species, and therefore subspecies. However, it is also a

well-known source of full species known as allopatric

speciation. It is interesting, however, that in Zimbabwe

and Zambia the two entities do occur together. They are

therefore strictly not allopatric. There are no reports or

collections of intermediates from these areas in which

they occur together. If they were indeed of the same spe-

cies one would have expected intermediates. There are

almost no variations within the entities throughout their

respective ranges. Therefore the evidence on distribution

supports the recognition of these entities as full species.

Furthermore a combination of overlapping features is

not conclusive proof of entities being mere subspecies

and not full species. It may also be used to differentiate

between species. There are many keys that use a com-

bination of overlapping keys to differentiate between

species. Field observation and field botanists support the

full species status of these entities.

FIGURE 5. — Photographic com-

parison between flowers of

Bauhinia petersiana subsp.

petersiana ( left) and B. bur-

rowsii (right). Photographer:

Emst Schmidt.

1

Bothalia 42,1 (2012)

47

FIGURE 6. — Known distribution of Bauhinia burrowsii.

The author therefore respectfully reinstates the enti-

ties as full species as it was before 1975 and the correct

names are as follows (there are no subspecies for these

species):

Bauhinia macrantha Oliv. in Flora of Tropical East

Africa 2: 289 (1871 ). Type: Angola, Ninda, Serpa Pinto

9 (LISU, holo.).

Bauhinia petersiana Bolle in Peters, Naturwissen-

schaftliche Reise nach Mossambique: Botanik 1 : 24

(1861). Type: Mozambique, ‘rios de Sena’, Peters s.n.

(B, holo. f, K).

ACKNOWLEDGEMENTS

The author gratefully acknowledges Carla Grins-

tad for the line drawings, Mrs Beverley Momberg and

the anonymous referees for their help with the text, and

the Instituto de Investigaqao Agraria de Mozambique

(HAM) for permission to collect plants in Mozambique.

REFERENCES

BRENAN, J.P.M. 1967. Flora of tropica / East Africa: Leguminosae

subfamily Caesalpinioideae. Crown Agents, London.

BRUMM1TT, R.K. & ROSS, J.H. 1975. The relationship of Bauhinia

petersiana and B. micrantha (Leguminosae-Caesalpinioideae).

Kew Bulletin 30: 593-595.

BRUMMITT, R.K. & ROSS, J.H. 1982. Anew combination for an Afri-

can Bauhinia. Kew Bulletin 37: 236.

COATES PALGRAVE, M. 2002. Trees of southern Afi-ica, edn 3.

Struik, Cape Town.

COETZER, L.A. & ROSS, J.H. 1977. Caesalpinioideae. Bauhinia.

Flora of southern Africa 1 6,2: 47 — 55.

SCHMIDT, E„ LOTTER,' M. & McCLELAND, W. 2007. Trees and

shrubs of Mpumalanga and Kruger National Park, edn 2. Jaca-

na, Johannesburg.

WHITE, F. 1983. The vegetation of Africa: a descriptive memoir to

accompany the Unesco/AETFAT/UNSO vegetation map of Afii-

ca. Unesco, Paris.

E.J.D. SCHMIDT*

* Ernst Schmidt Herbarium, P.O. Box 28, 1241 Kiepersol, South Africa.

MS. received: 2011-02-09.

HYAC1NTHACEAE

NEW COMBINATIONS IN LEDEBOURIA

The close relationship among Drimiopsis Lindl. &

Paxton, Ledebouria Roth and Resnova Van der Merwe

(Hyacinthoideae: Massonieae) that was highlighted by

Jessop (1972) has subsequently resulted in their segrega-

tion as the subtribe Ledebouriinae within tribe Masson-

ieae (Miiller-Doblies & Miiller-Doblies 1997). Charac-

ters that define the group include the mostly spotted or

streaked leaves, commonly multiple inflorescences per

annual growth cycle, reduced or obsolete bracts, and

paired ovules per locule; often with just a single seed

developing in only one or two of the locules. The leath-

ery capsule walls and the complete dehiscence, with

the locules reflexing completely, are diagnostic for the

group.

The generic status of the three taxa is less clear,

however, and Resnova has usually been treated as syn-

onymous with either Drimiopsis (Jessop 1972) or

Ledebouria [as Scilla] (Phillips 1951). More recently,

Miiller-Doblies & Miiller-Doblies (1997) proposed their

reinstatement at generic level, a treatment that was fol-

lowed by Lebatha et al. (2006). The development of

molecular techniques has prompted a renewed inter-

est in the systematic status of the three genera, but pre-

liminary phylogenetic analyses of plastid sequence data

have either failed to provide compelling evidence sup-

porting the recognition of three genera (Manning et al.

2004), or have actually demonstrated that Ledebouria

is paraphyletic unless it includes Drimiopsis and Res-

nova., although the latter two are evidently monophyletic

(Pfosser & Speta 1999; Pfosser et al. 2003; Wetschnig

& Pfosser 2003; Lebatha et al. 2006; Wetschnig et al.

2007). A morphological cladistic analysis of the group

provided only weak support (jacknife support value 55)

for a monophyletic Ledebouria, based on a limited sam-

pling of just eight species in the genus (Lebatha et al.

48

Bothalia 42,1 (2012)

2006). Proposed synapomorphies for Ledebonria s.str.

were a conical and severally lobed ovary, a style that

is longer than the ovary, and punctate-reticulate pol-

len (Lebatha et al. 2006). Of these states, ovary shape

and lobing are not consistent across the genus, with both

ovoid and turbinate and 3- or 6-lobed ovaries present in

the genus (Venter 2008). Ledebouria s.str. thus remains

very weakly diagnosable.

The lack of convincing molecular evidence support-

ing the recognition of the three genera as monophyletic

lineages led Manning et al. (2004) to include both Drim-

iopsis and Resnova in a broadly circumscribed Ledebou-

ria. They suggested that the various floral differences

between the three taxa that are generally used to jus-

tify their separation at generic level probably represent

linked pollination syndromes, and are thus not indepen-

dent. In any event, there is no doubt that the three taxa

are closely allied, and their taxonomic status at this stage

is primarily a matter of choice, although molecular data

favours their treatment as a single genus.

The broader circumscription of Ledebouria pro-

posed by Manning et al. (2004) has been implemented

in the latest checklist of southern African plants (Man-

ning & Goldblatt 2006), but a new species of Drimiopsis

described since then (Hankey et al. 2008) lacks a nomen-

clature! combination in Ledebouria. We provide this com-

bination here to facilitate its integration into checklists

and floras that adopt the broader generic circumscrip-

tion. A combination is also provided for Resnova trans-

vaalensis Van der Merwe, which was treated by Manning

et al. (2004) as conspecific with Resnova humifusa Van

der Merwe, following Muller-Doblies & Miiller-Doblies

(1997), but which we now consider to be distinct.

We also take this opportunity to formalize the sta-

tus of Drimiopsis and Resnova at sectional level within

Ledebouria. Current morphological and molecular ana-

lyses suggest that these two groups are monophyletic

and it is useful to have a formal way of referring to

them as infrageneric clusters within Ledebouria (Le-

batha et al. 2006). It is likely that additional sections

will be required in order to render sect. Ledebouria

monophyletic, but this will require a more complete

analysis of the group.

Ledebouria Roth, Novae plantarum species prae-

sertim Indiae orientalis: 194 (1821). Type: Ledebouria

hyacinthina Roth

Section Drimiopsis (Lindl. & Paxton) J.C. Manning

& Goldblatt, stat. nov.

Drimiopsis Lindl. & Paxton: 73 (1851). Type:

Drimiopsis maculata Lindl. = Ledebouria petiolata

J.C. Manning & Goldblatt

Ledebouria linioseta (A.J. Hankey & P.D. Lebatha)

J.C. Manning & Goldblatt, comb. nov. Drimiopsis lini-

oseta A.J. Hankey & P.D. Lebatha, in Hankey et al.: 72

(2008). Type: South Africa, Mpumalanga, between

Roossenekal and Lydenburg, 25 Nov. 1999, Hankey &

Turner 900 (PRE, holo.; K, iso.).

Section Resnova (Van der Merwe) J.C. Manning &

Goldblatt, stat. nov.

Resnova Van der Merwe: 46 (1946). Type: Resnova

humifusa (Baker) U.Miill.-Doblies & D.Mtill.-Doblies,

designated by Muller-Doblies & Muller-Doblies (1997:

59).

Ledebouria transvaalensis (Van der Merwe)

J.C. Manning & Goldblatt, comb. nov. Resnova trans-

vaalensis Van der Merwe: 46 (1946). Type: South

Africa, [Mpumalanga], Piet Retief Distr., Amsterdam,

Van der Merwe s.n. PRE26432 (PRE, holo.).

REFERENCES

HANKEY, A.J., BUYS, M.H. & LEBATHA, P.D. 2008. Drimiopsis

linioseta (Hyacinthaceae), a new species from the Sekhukhune-

land centre of endemism. South Africa. Bothalia 38: 12-1 A.

JESSOP, J.P. 1972. Studies in the bulbous Liliaceae in South Africa: 2.

Drimiopsis and Resnova. Journal of South African Botany 38:

151-162.

LEBATHA, P.D., BUYS, M.H. & STEDJE, B. 2006. Ledebouria , Res-

nova and Drimiopsis'. a tale of three genera. Taxon 55: 643-652.

LINDLEY, J & PAXTON, J. 1 85 1 . Paxton 's Flower Garden vol. 2. Lon-

don.

MANNING, J.C. & GOLDBLATT, P. 2006. Hyacinthaceae. In G.

Germishuizen, N.L. Meyer, Y. Steenkamp & M. Keith (eds), A

checklist of South African Plants : 952-971. Southern African

Biodiversity Network Report No. 41. SABONET, Pretoria.

MANNING, J.C., GOLDBLATT, P. & FAY, M.F. 2004. A revised

generic synopsis of Hyacinthaceae in sub-Saharan Africa, based

on molecular evidence, including new combinations and the

new tribe Pseudoprospereae. Edinburgh Journal of Botany 60:

533-568.

MULLER-DOBLIES, U. & MULLER-DOBLIES, D. 1997. A partial

revision of the tribe Massonieae (Hyacinthaceae). Feddes Rep-

ertorium 32: 49-96.

PHILLIPS, E.P. 1951. The genera of South African flowering plants Ed.

2. Botanical Survey Memoir No. 25. Department of Agriculture,

Division of Botany and Plant Pathology, Pretoria.

PFOSSER, M. & SPETA, F. 1999. Phylogenetics of Hyacinthaceae

based on plastid DNA sequences. Annals of the Missouri Botani-

cal Garden 86,4: 852-857.

PFOSSER, M„ WETSCHNIG, W„ UNGAR, S. & PRENNER, G.

2003. Phylogenetic relationships among genera of Massonieae

(Hyacinthaceae) inferred from plastid DNA and seed morphol-

ogy. Journal of Plant Research 116: 115-132.

ROTH, A.W. 1821. Novae plantarum species praesertim Indiae orien-

talis. Halberstad.

VAN DER MERWE, F.Z. 1946. Aantekeninge vir die hersiening van die

genus Scilla L. in Suid-Afrika. ‘n Nuwe genus: Resnova. Tvd-

skrif vir Wetenskap en Kuns 6: 41 — 46.

VENTER. S. 2008. Synopsis of the genus Ledebouria Roth (Hya-

cinthaceae) in South Africa. Herbertia 62: 85-1 55.

WETSCHNIG, W. & PFOSSER, M. 2003. The Scilla plumbea puz-

zle— present status of the genus Scilla sensu lato in southern

Africa and description of Spetaea lachenaliiflora, a new genus

and species of Massonieae (Hyacinthaceae). Taxon 52: 75-92.

WETSCHNIG, W„ KNIRSCH, W„ ALL S.S. & PFOSSER, M. 2007.

Systematic position of three little known and frequently mis-

placed species of Hyacinthaceae from Madagascar. Phyton Al :

321-337.

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, 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: 2010-11-08.

Bothalia 42,1 (2012)

49

AIZOACEAE

CORPUSCULAR] A GRACILLIMA IS THE CORRECT BINOMIAL FOR THE ILLEGITIMATE C. GRACILIS

L. Bolus validly published Delosperma gracile

L. Bolus in 1958 and later, in 1961, but based on a dif-

ferent type. She rectified this error (Bolus 1964) by pro-

viding the new name D. gracillimum L. Bolus for the

illegitimate later homonym. Hartmann (2001), when

transferring D. gracillimum from Delosperma to the

genus Corpuscularia erred in using the epithet gracilis

(Hartmann 2001). According to the International Code

of Botanical Nomenclature, for any taxon below the

rank of genus, the correct name is the combination of the

correct genus name and the final epithet of the earliest

legitimate name of the taxon in the same rank (McNeill

et al. (2006): Article 1 1 .4), which in this case is D. gra-

cillimum. Therefore, the correct name within the genus

Corpuscularia is C. gracillima. If no available epithet

existed, Hartmann could have re-used the epithet gra-

cilis as new, without reference to L. Bolus (Art. 58.1),

but this was not the case. Furthermore, because she

cited Delosperma gracillimum L. Bolus as a synonym,

the binomial Corpuscularia gracilis H.E.K. Hartmann

is superfluous and illegitimate (McNeill et al. (2006):

Art. 52.1). Herbarium acronyms follow Holmgren et al.

(1990).

Corpuscularia gracillima (L. Bolus) Niederle comb,

nov. Delosperma gracillimum L. Bolus in Journal of

South African Botany 30: 243 (1964), as nom. nov. pro

D. gracile L. Bolus (1961 ) non L. Bolus (1958). D. grac-

ile L. Bolus: 180 (1961), horn, illegit. Corpuscularia

gracilis (L. Bolus) H.E.K. Hartmann: 177 (2002 ‘2001’),

nom. illegit. superfl. Type: South Africa, Cape [Eastern

Cape], Seven Fountains near Grahamstown, 1959, Hall,

H. NBG 102/59 (BOL 131214— central branch, lecto.!,

here designated).

L. Bolus treated the whole gathering Hall, H. NBG

102/59 as the type. According to McNeill et al. (2006),

Art. 8.2, Ex. 1., the whole sheet BOL131214 cannot

serve as type because it contains four branches, which

1 interpret as belonging to the original collection, and

further flowers dated 28 March 1961. The problem con-

cerns nearly all sheets prepared by L. Bolus who later

added flowers to specimens. The names published by

her on or after 1 January 1958 would be problematic if

the collections indicated as types were of living plants

or seeds in reality. My lectotypification overcomes the

problem, whether the branches belong to the original

gathering or to plants cultivated by L. Bolus.

ACKNOWLEDGEMENTS

Helpful comments by two anonymous referees and

literature references provided by Chris Schroder are

gratefully acknowledged.

REFERENCES

BOLUS, H.M.L. 1958. Notes on Mesembryanthemum and some allied

genera 3, pp 289 — 4 1 7. University of Cape Town, Cape Town.

BOLUS, H.M.L. 1961. Notes on Mesembryanthemum and allied gen-

era. Journal of South African Botany 27 : 1 69—1 80.

BOLUS, H.M.L. 1964. Notes on Mesembryanthemum and allied gen-

era. Journal of South African Botany 30: 231-243.

HARTMANN, H.E.K. 2001 . Illustrated handbook of succulent plants:

Aizoaceae A-E. Springer, Berlin.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index herbariorum, part 1: The herbaria of the world , edn 8.

New York Botanical Garden. New York.

MCNEILL, J„ BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,

HAWKSWORTH, D.L., MARHOLD, K„ NICOLSON, D.H.,

PRADO, J„ SILVA, PC., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. (eds.) 2006. International Code of Botanical

Nomenclature ( Vienna Code) adopted by the Seventeenth Inter-

national Botanical Congress Vienna, Austria. July 2005. A.R.G.

Gantner Verlag, Ruggell, Liechtenstein. [Regnum Veg. 146].

J. NIEDERLE

PfF MU, Brno, Kotlarska 2, E-mail: niederle@inail.muni.cz.

MS. received: 2011-06-16.

IRIDACEAE

TAXONOMIC NOTES ON MORAEA (IRIDEAE) IN CONGO AND ADJACENT COUNTRIES

The genus Moraea Miller, now with some 210 spe-

cies (including several in press or manuscript), is centred

in southern Africa (Goldblatt 1986; Goldblatt & Manning

2010), but extends through tropical Africa to Ethiopia and

Eritrea in the north and to Nigeria and Cameroon in the

west. Outside the continent, two species of sect. Gynan-

driris (Parlatore) Goldblatt (Goldblatt 1998) occur in the

Mediterranean basin and Middle East. Goldblatt (1977)

recognised 24 species in tropical Africa, but later sepa-

rated M. stricta Baker from M. thomsonii Baker (Goldblatt

1986). Of these 25 tropical African species, six are shared

with eastern southern Africa, leaving 1 9 species restricted

to tropical Africa. Geerinck (2001), focusing on the genus

in Shaba Province of Congo (Kinshasa), has recently

described one new species, M. schaijesiorum Geerinck,

and reduced two species recognised by Goldblatt (1977)

to synonymy in M. ventricosa Baker: M. macrantha Baker

to varietal rank as var. macrantha (Baker) Geerinck and

M. verdickii De Wild, to the rank of forma in var. macran-

tha. In addition, Geerinck described f. boussardiana Geer-

inck of var. ventricosa and f. witteana Geerinck of var.

macrantha. These infraspecific taxa were distinguished by

minor details of perianth size and colour, with var. ventri-

cosa and var. macrantha based on flower size, specifically

50

Bothalia 42,1 (2012)

outer tepals and anthers, and f. boussardiana and f. wit-

teana by flower colour alone.

Morciea ventricosa is particularly distinctive among

the several large-flowered species of subgen. Gran-

diflora Goldblatt in its yellow, white, or blue flowers

with outer tepals 40-55 mm long, inner tepals 37-45

mm long, and anthers 8-11 mm long. Both the larger

outer and smaller inner tepal limbs are characteristically

spathulate, obtuse and ultimately fully reflexed, with the

broad style branches and crests arching to lie nearly hor-

izontally, and the ovary is always included in the spathes

(Goldblatt 1977, 1993, 1996). The species favours wet

habitats such as stream banks, seeps, and dambos; and

occurs across central Africa from Angola to Tanza-

nia. Moraea macrantha , with much the same range, is

a plant of well-drained, open woodlands or grasslands,

and has larger, consistently blue flowers with outer

tepals 57-80 mm long, inner tepals 55-75 mm long, and

anthers 12-15 mm long. Both species flower late in the

season, mainly March to May or June.

Moraea verdickii is less well known to us but it

blooms early in the season, mostly November to Feb-

ruary, usually has an exserted ovary, and has relatively

large yellow, or rarely white flowers, with outer tepals

mostly 60-80 mm long (rarely to 100 mm) and the

inner tepals 45-70 mm long, acute and evidently erect

(as far as we are able to determine) (Goldblatt 1977;

see also Geerinck 2005: photo 4). In comparison, plants

of M. ventricosa that we have examined always have

smaller flowers with an included ovary and the diagnos-

tic, markedly obtuse inner and outer tepals with fully

reflexed limbs. These features are also evident in a pho-

tograph accompanying the type of f. boussardiana.

In view of the correlated morphological and ecological

characteristics, we feel justified in maintaining Moraea

verdickii and M. macrantha as separate species distinct

from M. ventricosa. Within M. ventricosa as thus circum-

scribed, however, we consider the rank of forma to be triv-

ial and not worthy of recognition, and we therefore reduce

the fonns to synonymy. The forms in this case appear to

be local colour variants and a full review of the tropical

African species over their entire ranges should be under-

taken before any infraspecific taxa are identified. Those

wishing to recognise Geerinck’s forms remain free to do

so as the names exist and constitute an alternative treat-

ment of the group, at least in Congo.

Finally, we have examined the type of Moraea scha-

ijesiorum and conclude that it differs in no significant

way from M. clavata R.C. Foster, until now recorded in

the literature only from Angola and Zambia, to the south

and west of Congo. However, a collection of M. clavata

from Shaba [Schaijes & Michiel 2083 , 6 Sept. 1983,

17 km WSW of Kolwezi (K)], not cited by Geerinck,

is the first record of the species in the Congo. The type

of M. schaijesiorum was collected in 1988, and both it

and the earlier Shaba collection match typical M. cla-

vata in their relatively small, yellow flowers with outer

tepals 20-35 mm long, short foliage leaf inserted in the

middle of the stem, single sheathing leaf, and spathes

45-70 mm long. The early flowering time (November

in the type), is also consistent with M. clavata , the main

blooming time of which is from October to Decem-

ber (Goldblatt 1977). The occurrence of the species in

Shaba, not far from the Zambian frontier is not surpris-

ing. We have no hesitation in reducing M. schaijesiorum

to synonymy in M. clavata.

Moraea clavata R.C. Foster in Contributions from

the Gray Herbarium 114: 49 (1936), as nom. nov. pro

Moraea gracilis Baker: 272 (1878), horn, illegit. non

A.Dietr. (1833). Type: Angola, Huila, near Lopollo

River, Welwitsch 1545 (BM, lecto.!, designated by Gold-

blatt: 287 (1977); K!, LISUI, isolecto.).

M. schaijesiorum Geerinck: 1 1 (2003), syn. nov.

Type: Congo, [Shaba], plateau de Manika, road from

Kolwezi to Musokatanda, 8 Nov. 1988, Schaijes 4144

(BR, holo.!).

Moraea macrantha Baker , Flora of tropical Africa 7:

340 (1898). M. ventricosa var. macrantha (Baker) Geer-

inck: 4 (2001 ). Type: Malawi, Northern Province, Whyte

s.n. (K, holo.!).

Moraea arnoldiana De Wild.: 16 (1902). Type.

Congo, [Shaba], Kasenga, Verdick 606 (BR, holo.!).

Moraea ventricosa Baker in Bulletin of Miscellane-

ous Information, Royal Botanic Gardens, Kew 1895:

73 (1895). Type: Zambia, Northern Province, Fwambo,

Carson 37/1984 (K, holo.!).

M. bequaertii De Wild.: 540 (1913). M. ventricosa f.

bequaertii (De Wild.) Geerinck: 4 (2001). Type: Congo,

[Shaba], Lubumbashi, 5 Apr. 1912, Bequaert 316 [BR,

lecto.!, designated by Goldblatt: 280 (1977)].

M. ventricosa f. boussardiana Geerinck: 4 (2001),

syn. nov. Type: Burundi, Kigamba, 3 Apr. 1977, Reek-

mans 5982 (BR, holo.!).

Moraea verdickii De Wild, in Annales du Musee du

Congo Beige, Bot. 4: 17 (1902). M. ventricosa var. ma-

crantha f. verdickii (De Wild.) Geerinck: 5 (2001 ). Type:

Congo, [Shaba], Lukafu, Verdick 281 (BR, holo.!).

M. ventricosa var. macrantha f. witteana Geerinck: 5

(2001), syn. nov. Type: Congo, Parc National Upemba,

Riv. Kenia, 28 Mar. 1947, De Witte 2470 (BR, holo.!).

ACKNOWLEDGEMENTS

We thank Odile Weber, Royal Botanic Gardens, Kew,

for assistance with exsiccatae details and the Director,

Brussels Herbarium, for loan of type material.

REFERENCES

BAKER, J.G. 1878. Report on the Liliaceae, Iridaceae, Hypoxida-

ceae, and Haemodoraceae of Welwitsch’s Angolan Herbarium.

Transactions of the Linnean Society. London. Botany, ser. 2, 1 :

245-273.

BAKER, J.G. 1895. CCCCLI-Diagnoses Africanae, IV. Bulletin of

Miscellaneous Information. Roval Botanic Gardens. Kew. 1895:

63-75.

BAKER, J.G. 1898. Irideae. In W.T. Thiselton-Dyer, Flora of tropica!

Africa 1: 337-376. Reeve, London.

DE WILDEMAN, E. 1902. Etudes sur la flore du Katanga. Enumeration

systematique. Annales du Musee du Congo Beige, Botanique 4:

1-241.

Bothalia 42,1 (2012)

51

DE WILDEMAN, E. 1913. Decades novarum specierum florae katan-

gensis. VII I— X I . Repertorium specierum novarum regni vegeta-

bilis 1 1 : 535-547.

DIETRICH, A.G. 1833. Species plantarum. Nauck, Berlin.

FOSTER, R.C. 1936. Notes on nomenclature in Iridaceae. Contributions

from the Gray Herbarium of Harvard University 1 14: 37-50.

GEERINCK, D. 2001. Les variations infraspecifiques de Moraea ven-

tricosa Baker (Iridaceae d’Afrique centrale). Taxonomania 4: 4,

5.

GEERINCK, D. 2003. Une nouvelle espece pour la famille des Iri-

daceae au Congo-Kinshasa: Moraea schaijesiorum Geerinck.

Taxonomania 10: 11.

GEERINCK, D. 2005. Iridaceae. In D. Geerinck, C. Schaijes & M.

Schaijes, Flore d'Afrique Centrale. Jardin Botanique National

de Belgique. Me'ise.

GOLDBLATT, R 1977. Systematics of Moraea (Iridaceae) in tropical

Africa. Annals of the Missouri Botanical Garden 64: 243-295.

GOLDBLATT, R 1986. The moraeas of southern Africa. Annals of Kirs-

tenbosch Botanic Gardens 14: 1—224.

GOLDBLATT, R 1993. Iridaceae. In G.V.Pope, Flora zambesiaca. Flora

Zambesiaca Managing Committee, London.

GOLDBLATT, R 1996. Iridaceae. In R.M. Polhill, Flora of tropical East

Africa. Balkema, Rotterdam.

GOLDBLATT, P. 1998. Reduction of Barnardiella, Galaxia, Gynan-

driris, Hexaglottis, Homeria and Roggeveldia in Moraea. Novon

8: 371-377.

GOLDBLATT, P. & MANNING, J.C. 2010. Moraea intermedia and M.

vuvuzela (Iridaceae: Iridoideae), two new species from western

South Africa, and some nomenclatural changes and range exten-

sions in the genus. Bothalia 40: 146-153.

P. GOLDBLATT* & J.C. MANNING** ***

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

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

blatt@mobot.org

** 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;

E-mail: j.manning@sanbi.org.za

MS. received: 2011-03-16.

MONIMIACEAE

TAXONOMIC AND NOMENCLATURAL NOTES ON THE MONOTYP1C GENUS XYMALOS AND GENERAL INFORMATION ON THE

FAMILY MONIMIACEAE

INTRODUCTION

Harvey (1863) originally described Xylosma mono-

spora Harv., based on specimens from KwaZulu-Natal

but without precise localities ( Cooper 1251 and Cooper

1204), as belonging to the Flacourtiaceae. In 1878,

Bentham described Toxicodendrum acutifolium Benth.

from the Eastern Cape for the same taxonomic entity

but placed it in the Euphorbiaceae. Later Engler (1895,

1900) described the species yet again, this time from

Tanzania, under the name Paxiodendron usambarense

Engl, but placing it in the Lauraceae.

Baillon (1887) compared Harvey’s X. monospora

(which has a monocarpellate ovary with a solitary ovule,

sessile stigma, and drupaceous, non-dehiscent fruit)

with Hyaenanche Lamb., then in the Euphorbiaceae

but now in Picrodendraceae (APG III 2009). Species

of Hyaenanche have 4-locular ovaries with two ovules

per locule, 3-5 styles connate in the lower half, and

dehiscent capsules. Baillon (1887) concluded that the

two entities placed respectively by Bentham (1878) in

Xylosma G.Forst. and by Harvey (1863) in Toxicoden-

drum were in fact conspecific but did not belong in

either Toxicodendrum or Xylosma. He therefore placed it

in a new genus Xymalos Baill., which is an anagram of

Xylosma, which means fragrant wood — referring to the

leaves and wood that are lemon-scented when crushed.

He placed Xymalos in the Bixaceae.

De Jussieu founded the family Monimiaceae in 1809

based on three genera, Monimia Thou., Atherosperma

Labill. (a monotypic genus from Australia and Tasma-

nia), and Siparuna Aubl. (a large neo-tropical genus

with over 70 species). Oliver (1896) was the first to

place Xymalos in the Monimiaceae and also emended

the generic description. This familial transfer was fol-

lowed by amongst others Baker & Wright (1909), Per-

kins & Gilg ( 191 1 ), Wright (1912), Engler (1915), Verd-

court (1968), and Stannard (1997). Warburg (1893)

recognized Xymalos as closely related to the Flacour-

tiaceae, but Gibbs (1917) and Hutchinson (1973) both

placed Xymalos in the Trimeniaceae. Currently Xymalos

is placed in Monimiaceae subfamily Mollinedioideae

(Thome 1974; Philipson 1987, 1993). Molecular data

support this classification (Renner 1998, 1999). Moni-

miaceae is a pantropical family in the order Laurales and

in a broad sense comprises 440 species and 34 genera

(Philipson 1987, 1993). The small families Atherosper-

mataceae and Siparunaceae have since been excluded

from the Monimiaceae. Trimeniaceae is excluded from

the order Laurales — a decision that is well supported

by phylogenetic and morphological studies (Mooney

el al. 1950; Renner 1999; Renner & Chanderbali 2000;

Romanov et al. 2007; APG III 2009). Mabberley (2008)

recorded 24 genera in the family but the new mono-

typic genus, Grazielanthus, from the Brazilian Atlantic

Coastal Forest has since been described by Peixoto &

Pereira-Moura (2008). The family currently comprises

150-220 species of shrubs and small trees in 25 genera

(Renner et al. 2010). It has a highly disjunct, presum-

ably Gondwanan distribution, mainly in the southern

hemisphere (Renner et al. 2010). Xymalos monospora

occurs in montane forests in east tropical and southern

Africa, West Cameroon, and on Bioko, an island once

connected to Africa (Figure 1 ).

Monimiaceae, together with the closely related fami-

lies Lauraceae and Hemandiaceae ( Gyrocarpus Jacq.),

are characterized by having secretory cells containing

so-called ‘ethereal oil’ (ethereal oil cells) and a unila-

cunar nodal anatomy (Mooney et al. 1950). Monim-

iaceae can be recognized by decussate to subopposite,

exstipulate leaves; twigs that are often flattened below

the somewhat swollen nodes; leaves that are pellucid-

52

Bothalia 42, 1 (2012)

FIGURE 1. — Distribution of Xymalos monospora based on specimens

housed in the National Herbarium, Pretoria, and on the literature

(Verdcourt 1968; Stannard 1997).

dotted due to the presence of numerous glands or ‘ethe-

real oil cells’ (Lorence 1985), and are lemon-scented

when crushed. The lack of stipules is considered by

Hickey & Wolfe (1975) to be derived within the Lau-

rales. Xymalos has leaves with margins that possess a

‘monimioid tooth’ with an undivided median vein and

an indurated cap (Figure 4), another derived character

in the Laurales (Lorence 1985), or the leaves are some-

times entire. Leaves are pinnately veined with prominent

venation on the lower surface and the secondary veins

originate at uniform angles to the costa (Lorence 1985),

looping and joining well within the margin, sometimes

leaving a bullate or puckered surface above. Plants are

usually dioecious or monoecious and the yellow flowers

are very small and mostly unisexual, except for Horto-

nia Wight ex Am., which have bisexual flowers (Endress

& Igersheim 1997). The latter is a Sri Lankan genus,

sister to Xymalos (Renner et al. 2010), and possesses

flowers with an elaborate perianth comprising spirally

arranged petaloid and sepaloid series, 6-14 free, stalked

carpels borne on a discoid receptacle, appendaged sta-

mens and with staminodes present (Lorence 1985). The

receptacle in Monimiaceae is deeply concave, cupuli-

fortn or urceolate (Mooney et al. 1950). The family is

characterized by apocarpous gynoecia with uniovulate

carpels (Endress & Igersheim 1997), surrounded by a

flat or concave floral cup (Lorence 1985). The fruit(lets)

are presented on the opened floral cup. Male flowers

have a 4-8-lobed perianth with few to many stamens,

with or without appendages and arranged in whorls or

sometimes spirally or irregularly disposed, either with-

out a pistillode or with remnants of one. The anthers

are 2—- 1-sporangiate. Staminodes are absent or present

in female flowers, carpels are few to many (only one in

Xymalos) and sessile or stipitate, with a solitary, anatro-

pous ovule. In the genera Kibara Endl. (Malaysia) and

Xymalos (Africa) the discoid stigma is sessile on the

turbinate ovary, but a style is present in all other genera,

typically short and included, but exserted in Monimia

(Philipson 1986, 1993). Ovules in Xymalos are crassinu-

cellate and bitegmic (Leinfellner 1966) and the carpels

are strongly ascidiate in the Mollinedioideae (Endress

& Igersheim 1997). The fruit are either separate drupe-

lets with a ± strongly developed putamen formed by the

lignified endocarp, or apocarpous berrylets frequently

enclosed in a persistent, well-developed fleshy hypan-

thium and/or receptacle (Philipson 1986; Romanov et

al. 2007). Romanov et al. (2007) described these two

fruit forms in the family and distinguished four different

types of drupe(lets) differing in their endocarp structure.

(Copper et al. (2006) record Xymalos mossambi-

censis Cavaco as occurring in tropical Africa. Cavaco

( 1949) described this species from Mozambique but the

name was not taken up by Verdcourt in his account of

the Flora of tropical East Afiica (1968), nor by Stan-

nard in his account for Flora zambesiaca (1997), both

of which only mention Xymalos monospora. Both of

these authors and subsequent publications by Philipson

(1993), Jordaan (2000, 2003), Da Silva et al. (2004) and

Mabberley (2008) consider Xymalos a monotypic genus

occurring in tropical and southern Africa. In Mozam-

bique, Xymalos is recorded north of the Zambezi River

at Ribaue, Gurue (Stannard 1997) and Nampula (Da

Silva et al. 2004) and south of the Zambezi at Tsetssera

(Stannard 1997). Cavaco collected a specimen south of

the Save River (southern Mozambique) and somewhere

between Mapai and Pafuri (Kruger National Park area)

(type specimen mentioned in the protologue). In this

area between Save River and Mapai there is certainly no

suitable habitat for Xymalos monospora , which usually

grows in Afromontane forest, and the closest locality,

Tsetssera, is roughly 400 km to the north of the Mapai-

Pafuri record. Therefore, the locality is very doubtful

as given in the protologue, namely: ‘Sul do Save, entre

Mapai et Pafuri, in Mozambique’.

Cavaco compared the holotype of X. mossambi-

censis with specimens of X. monospora from southern

and tropical Africa housed in the Paris Herbarium (e.g.

Rudatis 1418, Stolz 2063, Holst 4249) and concluded

that his specimen constituted a new species. We have

not seen the type but Cavaco’s (1949) protologue and

accompanying illustration indicate that X. mossambi-

censis differs essentially from X. monospora in having

larger leaves with entire margins and fasciculate rather

than racemose inflorescences. Throughout the distribu-

tion range of X. monospora , the leaf margins vary from

entire (Figure 2) to coarsely serrate with widely spaced

gland-tipped teeth (Figure 3). Inflorescences are always

axillary and vary from cymose to racemose. There are

no significant differences between X. mossambicensis

and X. monospora, and they are therefore considered as

conspecific.

TAXONOMY

Xymalos monospora (Harv.) Bail I. in Bulletin Men-

suel de la Societe Linneenne de Paris 1: 650 (1887);

Warb.: 53 (1893); Oliv.: t. 2444 (1896); Sim: 288, t.

121 (1907); Baker & C.H. Wright: 170 (1909); Per-

kins & Gilg: 10, t. 5 (1911); C.H. Wright: 493 (1912);

F.W. Andrews: 7 (1950); J.Leonard: 402, t. 39 (1951);

Keay: 55 (1954); Verde.: I, t. 1 (1968); Fouilloy: 109

(1974); R.B.Drumm.: 237 (1975); Pooley: 96 (1993);

A.E.van Wyk & P.van Wyk: 302 (1997); Stannard: 43

(1997); Lotter: 114 (2002); M. Coates Palgrave: (2002);

Bothalia 42,1 (2012)

53

FIGURE 2. — Xymalos monospora showing entire leaves. Photogra-

pher: M. Lotter.

FIGURE 3. — Xymalos monospora showing toothed leaves and some

leaf damage. Photographer: M. Lotter.

Boon: 104 (2010). Xylosma monospora Harv.: 52, t. 181

(1863). Type: South Africa, Natal [KwaZulu-Natal],

without precise locality, Cooper 1251 (TCD — dig-

ital image!, lecto., designated here; K — digital image!,

PRE!, isolecto.). [Note: Of the two collections cited in

Harvey’s protologue of Xylosma monospora , Cooper

1251 is chosen as the lectotype because there are more

duplicates of this collection than of Cooper 1204.]

Toxicodendrum acutifolium Benth.: 214 (1878). Type:

South Africa, Eastern Cape, ‘British Caffraria’, Barber

& Barber 10 (K — digital image!, holo.).

Paxiodendron usambarense Engl.: 182 (1895). Xyma-

los usambarensis (Engl.) Engl.: 310 (1901). Type: Tan-

ganyika [Tanzania], E Usambara Mountains, Gonja,

Bulwa, Holst 4249 (B, holo.t; P — digital image!, lecto.,

designated here; COI — digital image!, JE — digital

image!, K — digital image!, isolecto.).

P. usambarense var. serratifolia Engl.: 182 (1895).

Type: Tanganyika [Tanzania], Kilimanjaro, Marangu,

Volkens 2264 (B, holo.t; JE — digital image!, lecto., des-

ignated here).

P ulugurense Engl.: 389 (1900). Xymalos ulugurense

(Engl.) Engl.: 310 (1901); Baker & C.H. Wright: 170

(1909). Type: Tanganyika [Tanzania], Uluguru Moun-

tains, Lukwangule Plateau, Goetze 274 (B, holo.t; K —

digital image!, lecto., designated here).

Xymalos mossambicensis Cavaco: 45 (1949). Type:

‘Moqambique [Mozambique], Sul do Save, entre Mapai

et Pafuri’, Cavaco 89 (P, holo.).

Note : Most type specimens were seen as digital

images on websites. Where the holotypes have been

destroyed during World War II in the Berlin Herbarium,

as in the case of Holst and Goetze specimens, lectotypi-

fication is covered by article 9.15 of the Code (McNeill

et al. 2006), which provides for narrowing the lectotype

to a single specimen. There are records of duplicates of

Holst 4249 verified by Engler, isotypes of Paxiodendron

usambarense , in K, COI, JE and P, all of them flower-

ing material, and all specimens show entire leaves. The

Paris specimen is selected because it is the only one

which gives the exact type locality in Tanzania. All the

syntypes of Paxiodendron usambarense Engl. var. ser-

ratifolia Engl, (three Volkens specimens) were destroyed

in Berlin, but one extant isosyntype, Volkens 2264. is

seen on ALUKA [http://plants.jstor.org/], housed at JE,

and is therefore selected as the lectotype. This speci-

men has leaves with widely-spaced teeth in contrast

with all the isotypes of the typical P. usambarense with

entire leaves. There is only a fragment of the duplicate

of Goetze 274 at K, which is selected as the lectotype for

Xymalos ulugurense.

Evergreen tree up to 8(-20) m, mostly single-

stemmed, with dense, rounded, spreading crown. Bark

silvery grey or grey-brown, rough, slightly fissured,

with prominent lenticels, flaking in large scales to leave

concentric, ridged markings; inner wood lemon-yellow;

sap reddish. Leaves simple, opposite, glabrous, lemon-

scented when crushed, pellucid-dotted; lamina narrowly

elliptic to obovate, 40-200 * 15-100 mm, apex acute,

rounded or shortly acuminate, base cuneate, margin usu-

ally irregularly and coarsely glandular-serrate, some-

times entire; midrib sunken above, prominent beneath,

lateral veins 6-9, looping well within margin, reticu-

late venation prominent on both surfaces; petiole up to

30 mm long, glabrous. Inflorescences racemose or pa-

niculate, solitary or paired in leaf axils, 1 0— 35(— 70) mm

long; peduncles 5-10 mm long; bracts triangular,

1.0-2. 5 mm long. Flowers unisexual, small, yellowish

or greenish; pedicels 1-3 mm long, velvety. Perianth

1-2 mm long, hairy. Petals absent. Male flowers: peri-

anth 4-6-lobed; lobes rounded to ovate; stamens 6-15 or

more; anthers subsessile. Female flowers: perianth 3-5-

lobed; lobes ovate, rounded to triangular, velvety, mar-

gins ciliate; ovary obovoid, cylindrical or turbinate, gla-

brous; stigmas sessile, discoid, thick, glabrous. Fruit an

ovoid to ellipsoid drupe, (5-) 10-25 x 3-15 mm, slightly

asymmetrical, glabrous, orange or reddish, fleshy,

crowned with persistent stigma. Seed solitary, ellipsoid,

compressed, 10 « 8 mm, white (Figure 4). Flowering

time: June-Oct.

Diagnostic characters: leaves of Xymalos monospora

are opposite and translucent gland-dotted, with a lemon

smell when crushed. The bark is very distinctive, flak-

ing to reveal circular or worm-like markings, and when

cut has a lemon-like smell (Lotter 2002). Small, fragrant

yellow flowers, the sexes on separate trees, appear from

June to October (Lotter 2002), arranged in racemes and

54

Bothalia 42,1 (2012)

FIGURE 4. — Xymalos monospora, A, C & E, Mohle 389 (PRE); B, Van der Schijff 5630 (PRE); D, Compton s.n. (PRE53940)\ F, De Winter <£

Killick 8954 (PRE). A, flowering tw ig x 1;B, leaf with entire margin x 1 ; C, tip of tooth on leaf margin * 2; D, female flower x 8; E, male

flower x 4; F, fruiting branch x 1. Artist: Daleen Roodt.

Bothalia 42,1 (2012)

55

FIGURE 5. — Xymalos monospora showing leaf damage. Photographer: M. Jordaan.

often produced in clusters from old leafless wood (Sim

1907). The fruits are ovoid, smooth, 1 -seeded drupes,

often enclosed by a perianth or a fleshy receptacle, with

a persistent stigma.

Distribution and habitat : Xymalos monospora grows

in Afromontane, scarp or coastal forests, or in grass-

lands associated with forests, in moist areas near water-

courses, from near sea level to above 2 000 m. It occurs

in eastern tropical and southern Africa, from southern

Sudan as far south as the Eastern Cape. Tadesse (2000)

suggests that the species could also occur in Ethiopia

but has not yet been recorded there. Distinctive disjunct

populations occur in the highland forests of Nigeria on

the Gotel Mountains (Aluka Library [http://plants.jstor.

org/]), Cameroon Mountains (Cable & Cheek 1998) and

on the Island of Bioko, formerly Fernando Po (Keay

1954) (Figure 1 ). The range of Xymalos monospora falls

well within the archipelago-like Afromontane Region of

Endemism (White 1978, 1983). This African phytocho-

rion is of special biogeographical and evolutionary sig-

nificance because of its putatively ancient status.

Pollination: Xymalos monospora is wind- or insect

pollinated, but this needs further investigation (Philip-

son 1993). Quantities of pollen are produced by the

male flowers, and most specimens collected possess

male flowers, with very few of the ± 50 specimens in the

National Herbarium, Pretoria, with female flowers.

Leaf damage', there is some damage to the leaves on

± 100 specimens examined, usually in the form of black

patches or holes in the leaf blade (Figure 5). Xymalos

monospora is the host for an unusually large number of

fungal pathogens (Doidge 1950), which are probably the

cause of the damage to the leaves. The leaf edges are

also sometimes eaten away (Figure 3) and the species

is a host for the larva of the butterfly Papilio dardanus

Brown (mocker swallowtail) (Papilionidae) (Picker et al.

2004; butterflycorner website [en.butteiflycomer.net]).

ACKNOWLEDGEMENTS

We would like to thank Hester Steyn for preparing the

distribution map, and the artist Daleen Roodt for the line

drawing.

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Bothalia 42,1 (2012)

57

ASTERACEAE

BERKHEYA JARDINEANA (ARCTOT1DEAE — GORTERIINAE), A NEW DWARF PERENNIAL FROM THE SWARTRUGGENS,

WESTERN CAPE

Berkheva Ehrh., a genus of ± 80 species centred in

southern Africa, was last revised by Roessler (1959)

but many species remain poorly known; to some extent

at least because of the challenges posed to collectors by

these often viciously spinescent plants. Berkheva is dis-

tinguished by its shrubby or perennial habit, spinescent

involucral bracts, mostly radiate (rarely discoid) capit-

ula, ± deeply alveolate receptacle with uniformly thin-

walled cavities, and pappus of ± 20 denticulate scales

in one or two rows (Roessler 1959; Bremer 1994). The

genus is polyphyletic as currently defined (Funk & Chan

2008; Funk et al. 2008; Karis et al. 2009), with a prelim-

inary molecular analysis of the subtribe nesting the gen-

era Culliimia R.Br. ex Aiton, Cuspidia Gaertn., Didelta

L’Her. and Heterorachis Sch.Bip. ex Walp. in various

parts of Berkheva. Substantial revision of generic cir-

cumscription is required to render it monophyletic.

Of 23 species of Berkheya recorded from the core

Cape Floristic Region, 16 are endemic (Goldblatt &

Manning 2000; Manning et al. 2010), including the

recently described B. chrysanthemoides J.C. Manning &

Goldblatt from the Bokkeveld Mountains (Manning et

al. 2010). Here we describe another novelty discovered

in 2009 on the Swartruggens northeast of Ceres.

Berkheya jardineana J.C. Manning & Goldblatt , sp.

nov.

TYPE. — Western Cape, 3219 (Wuppertal); Knolfon-

tein, Swartruggens, 60 km NE of Ceres, (-DC), 26 Oct.

201 1, 7. Jardine 1738 (NBG, holo.; MO, S, iso.).

Dwarf, tufted perennial to 10 cm high at flowering,

forming small clumps from branching, woody rhizome;

stems erect, cobwebby or thinly felted, flushed purple.

Leaves crowded at base of stems, basal foliage leaves

spathulate in outline, softly leathery, blade suborbicu-

lar, 15-35 mm diam., weakly folded along midline and

coarsely 7- or 9-dentate, primary teeth excurrent in pale,

relatively soft spine ± 3 mm long, with smaller second-

ary and tertiary antrorse spines 1-2 mm long between

primary spines, cobwebbed or thinly felted on both sur-

faces and with scattered minute, gland-tipped hairs, gla-

brescent above, veins raised beneath, narrowed into pet-

iole-like base 15-25 mm long, more densely cobwebbed

than blade; cauline leaves smaller, lanceolate, 10-20 *

3- 6 mm, sessile and weakly amplexicaul, weakly 3- or

4- jugate, teeth excurrent into soft spines ± 3 mm long

and with 1 or 2 smaller antrorse spines along margins

between primary spines, thinly felted on both surfaces

and with scattered minute, gland-tipped hairs. Capitula

1-3 in shortly pedunculate racemes, discoid, 25^-0 mm

across involucre and 1 5-20 mm across disc; involu-

cral bracts 4- or 5-seriate, bracts basally connate for ±

5 mm, patent-reflexed, narrowly lanceolate, flat, apex

excurrent in a yellowish spine 2-3 mm long, outer and

median series with 3-5 pairs of patent spines similar to

apical spine, outer surface thinly cobwebbed and with

minute, gland-tipped hairs, innermost bracts with 1-3

pairs of spines in distal half grading into several antrorse

spinules in basal half, glabrous except towards apex,

outer one or two series mostly 6-8 x 2-3 mm, median

two series 10-12 x 3 mm, inner series oblong-lanceo-

late, 5-6 x 1 .5 mm; receptacle deeply alveolate, margins

irregularly fimbriate with straw-like spinules 0. 5-1.0

mm long. Corolla funnel-shaped, yellow, densely glan-

dular-pubescent on tube and more sparsely along lobe

margins, ± 7 mm long, tube ±3.5 mm long, lobes erect,

narrowly lanceolate, ± 3.5 mm long, penicillate. Sta-

mens: anthers tailed, with lanceolate apical appendage,

± 4 mm long; endothecial cells with inner periclinal wall

thickenings not evidently displaced. Achenes turbinate,

± 2.5 mm long, 10-ribbed, those of outer florets densely

antrorsely pubescent with short, twin hairs but inner

achenes progressively more glabrous. Pappus scales

sub-biseriate, ± 20, oblong-obovate, denticulate, outer

series slightly shorter than inner, 1.0-1. 5 x 0.5 mm. Pol-

len lophate. Flowering time: Oct.-Dec. Figure 1.

Distribution and ecology: known so far only from

the Farm Knolfontein on the Swartruggens, a semi-arid

mountain range 60 km east of Ceres at the eastern mar-

gin of the Cape Floristic Region (Figure 2). The spe-

cies is locally common on sandy flats in areas of dry,

rocky sandstone soils, mostly in full sun. It appears to

be highly localized; although several subpopulations are

known, they are all within approximately 1 km of one

another.

Diagnosis and relationships: this petite species is dis-

tinguished in the genus by its diminutive size and curi-

ous growth form, developing into small, multi-stemmed

clumps or low cushions through branching of the woody

underground stem. The leaves are clustered at the base

of the aerial stems, which reach only to 10 cm high at

flowering, later elongating to 15 cm in fruit. The basal

foliage leaves are highly distinctive, being softly leath-

ery and spathulate in outline with a coarsely toothed,

suborbicular blade only weakly armed with pale, rela-

tively soft-textured spines. Both surfaces are thinly cob-

webby-felted. The species is not pyrophytic and the low

growth form is likely an adaptation to its exposed habi-

tat.

The rosulate habit, relatively small, discoid capitula,

and sub-biseriate pappus of small obtuse scales place

Berkheya jardineana in ser. Rigidae (Roessler 1959).

Other members of the series are larger perennial herbs or

shrubs with pinnatifid or lobed leaves.

Etymology*: the species is named for Ivor and Cora

Jardine, who first collected specimens in 2009. They

have devoted many years to documenting the flora of the

Swartruggens (Jardine & Jardine 2010), in the course of

which they have discovered several novelties, including

Hesperantha lithicola Goldblatt & J.C. Manning (Iri-

daceae) and Trieenea occulta J.C. Manning & Goldblatt

(Scrophulariaceae).

58

Bothalia 42,1 (2012)

FIGURE 1 .- Berkheya jardineana, Jardine 1738 (NBG). A, Plant; B, foliage leaf; C, involucral bracts (outermost on left, innermost on right); D,

outer floret showing puberulous ovary and pappus of small, obtuse scales, plus portion of alveole margin; E, achenes (outer on left, inner on

right) ( Jardine 1747). Scale bar: A, B, 10 mm; C, 8 mm; D, E, 2 mm. Artist: John Manning.

Bothalia 42,1 (2012)

59

Additional specimens seen

WESTERN CAPE. — 3219 (Wuppertal): Knolfontein, Swartrug-

gens, 60 km NE of Ceres, 1 201 m, (-DC), 26 Oct. 2009, Jardine &

Jardine 570 (NBG); 21 Nov. 2011, Jardine 1747 (NBG); 12 Dec.

201 1, Jardine 1761 (NBG); 14 Dec. 2011 , Jardine 1766 (NBG).

REFERENCES

BREMER, K. 1994. Asteraceae, cladistics and classification. Timber

Press, Portland, Oregon.

FUNK, V.A. & R. CHAN. 2008. Phylogeny of the spiny African daisies

(Compositae, tribe Arctotideae, subtribe Gorteriinae) based on

trnL-F , ndhF , and ITS sequence data. Molecular Phylogenetics

and Evolution 48: 47-60.

FUNK, V.A., CHAN, R. & KEELEY, S.C. 2008. Insights into the evolu-

tion of the tribe Arctoteae (Compositae: subfamily Chicorioideae

s.s.) using trnL-F, ndhF, and ITS. Taxon 53: 637-655.

GOLDBLATT, P. & J.C. MANNING. 2000. Cape Plants: A Conspectus

of the Cape Flora. Strelitzia 9. National Botanical Institute &

Missouri Botanical Garden.

JARDINE, 1. & JARDINE, C. 2010. Four seasons on the Swartruggens.

Veld & Flora 96: 179-181.

KARIS, P.O., FUNK, V.A., MCKENZIE, R.J., BARKER, N.P. & R.

CHAN. 2009. Arctotideae. In: Systematics, Evolution and Bioge-

ography of Compositae , Funk, V.A., A. Susanna, T. Stuessy, arid

R. Bayer (Eds.), I APT, Vienna, Austria, pp. 285-3 10.

MANNING, J.C., KARIS, P.O. GOLDBLATT, P. & HELME, N.A.

2010. Berkheya chrysanthemoides and Heterorachis hystrix , two

new species of Arctotideae-Gorteriinae (Asteraceae) from the

southwestern Cape. Bothalia 40: 219-224.

ROESSLER, H. 1959. Revision der Arctotideae-Gorteriinae (Compo-

sitae). Mitteilungen der Botanischen Staatssammlung Mimchen

3: 71-500.

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 Life

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: 2012-01-03.

ASTERACEAE

OSTEOSPERMUM NORLINDHIANUM AND O. NORDENSTA MIL TWO NEW SPECIES OF OSTEOSPERMUM SECT. TREFENESTRATAE

(CALENDULEAE) FROM THE GREATER CAPE FLORISTIC REGION

Calendulae is a small tribe of ± 120 spp. with a

marked centre of diversity in southern Africa, where

± 80% of the species occur (Nordenstam & Kallersjo

2009). Available phylogenetic analyses (Nordenstam

1994, 2006; Nordenstam & Kallersjo 2009) retrieve

Garuleum Cass, and Dimorphotheca Vaill. ex Moench

as two early-branching lineages, with no question about

their taxonomic status. Generic delimitation within the

remainder of the tribe, however, remains unsatisfac-

tory (Manning & Goldblatt 2008). The species are cur-

rently segregated among ten genera: Calendula L. (± 15

spp.), Chrysanthemoides Fabr. (2 spp.), Gibbaria Cass.

(2 spp.), Inuloides B.Nord. (1 sp.), Monoculus B.Nord.

(2 spp.), Nephrotheca B.Nord. & Kallersjo (1 sp.), Nor-

lindhia B.Nord. (3 spp.), Oligocarpus Less. (2 spp.),

Osteospennum L. (± 45 spp.) and Tripteris Less.

(20 spp.) (Nordenstam 2007; Nordenstam & Kallersjo

2009). In spite of recent generic fragmentation both

Osteospermum and Tripteris remain polyphyletic (Nor-

denstam & Kallersjo 2009), necessitating the recognition

of additional genera if this treatment is to be carried to

its logical conclusion.

In practical terms, few synapomorphies are avail-

able for the recognition of these new segregates, most

of which are mono- or oligotypic, and it is unfortunate

that dismemberment of the group has been implemented

piecemeal and in advance of a well-sampled and well-

supported analysis of the tribe. Even with the very nar-

row circumscription of Tripteris proposed by Norden-

stam (2007) to include just those species with winged,

apically3-fenestrate achenes, the molecular analysis in

Nordenstam & Kallersjo (2009) still locates taxa with

other kinds of achenes (among them the genus Monocu-

lus) among typical Tripteris species, rendering the latter

polyphyletic.

The alternative, and in our opinion more useful treat-

ment from both a practical and theoretical perspective

(Manning & Goldblatt 2008), is to adopt a synthetic

view of Osteospermum as constituting the monophyletic

clade sister to Dimorphotheca as resolved in the phy-

logenetic analyses presented by Nordenstam (2006) and

Nordenstam & Kallersjo (2009), thus including the gen-

era Calendula, Chrysanthemoides, Gibbaria, Inuloides,

Monoculus, Nephrotheca, Norlindhia, Oligocarpus and

Tripteris. Segregate lineages within Osteospermum sens,

lat. are then usefully treated at sectional level, as was

initiated by Norlindh (1943) in the last comprehensive

revision of the tribe in sub-Saharan Africa. Although

the current sectional boundaries clearly require sub-

stantial revision in the light of modem phylogenetic

analyses, shuffling of species among them will have no

impact on the nomenclature. This is the treatment that

60

Bothalia 42,1 (2012)

has been adopted in the two floras treating the species

of the Greater Cape Floristic Region (Manning & Gold-

blatt, in press; Snijman in press.) and we follow it here

in describing two new species of Osteospermum subg.

Tripteris (Less.) T.Norl. from the semi-arid parts of the

Greater Cape Floristic Region.

Osteospermum sect. Trifenestratae T.Norl. [as Trife-

nestrata\, which coincides with the narrow definition of

the genus Tripteris (sensu Nordenstam 2007), comprises

± 18 species of perennial herbs, subshrubs or shrubs

widely distributed through Africa, with some 7 species

endemic to the winter rainfall parts of South Africa and

southern Namibia. It is diagnosed by 3-winged achenes

with all three sides of the apical airchamber thin-walled

and translucent (‘fenestrate') (Norlindh 1943). Here we

describe two new species in the section. Osteospermum

norlindhianum from the Cold Bokkeveld resembles

O. dentation Burnt. f. from sandy flats along the West

Coast of Western Cape in its herbaceous, perennial habit

and its heteromorphic achenes but differs in details of

the foliage and fruit. It had been recognized as distinct

by Norlindh during the preparation of his monograph of

the tribe (Norlindh 1943) but it is only now that fruit-

ing material has become available. O. nordenstamii from

quartz fields in southern Namaqualand, is a recently dis-

covered, gnarled dwarf shrublet with consistently oppo-

site, basally connate leaves, suggesting a relationship

with the common Namaqualand shrub, O. oppositifolium

(Aiton) T.Norl.

Osteospermum norlindhianum J.C. Manning &

Goldblatt , sp. nov.

TYPE.— WESTERN CAPE, 3219 (Wupper-

tal): Swartruggens, Knolfontein 60 km NE of Ceres,

32°50'49.7"S, 1 9°37'20.8"E, l 260 m, (-DC), 26 Oct.

2011 , /. Jardine 1729 (NBG, holo.; MO, S, iso.).

Tufted perennial from woody crown, 300-450 mm

high, caudex producing a cluster of annual flowering

stems; flowering stems suberect, laxly branched, 2-3

mm diam. at base, pubescent with mix of straggling,

eglandular hairs and short, gland-tipped hairs, flushed

purple at base. Leaves congested basally, lowermost sub-

opposite with bases imbricate but not connate, becom-

ing alternate distal ly as internodes lengthen, decreasing

in size acropetally and ultimately grading into linear-

subulate bracts, lower leaves oblanceolate, mostly 40-90

x 6-9 mm, leathery, sparsely to densely pubescent with

mix of acute, eglandular hairs and gland-tipped hairs,

adaxial surface ± asperulous, abaxial surface ± glan-

dular-pubescent intermixed with scattered, straggling,

eglandular hairs, these longer and denser along margins,

blade tapering below into long, petiole-like base, mid-

rib prominent abaxial ly, margins sparsely and weakly

dentate, apiculate. Capitula heterogamous, radiate, ± 20

mm diam., on slender, branched peduncles forming lax,

compound corymbs, nodding in fruit; involucre shal-

lowly campanulate, 7-8 mm diam.; involucral bracts

12-15, sub-uniseriate, lanceolate, 3. 5-4.0 x 1.0-1. 5

mm, acute, densely glandular-pubescent, with scarious

margins 0.25-0.50 mm wide; receptacle flat, glabrous.

Ray florets female-fertile, 8-10; tube sparsely glandular-

pubescent, ± 1 mm long; lamina spreading but recoil-

ing in afternoon, narrowly elliptic, 4-veined, ± 3 x as

long as involucre, 9-10 x 3 mm, bright yellow, some-

times flushed reddish abaxially. Anthers vestigial, free,

reduced to 4 subulate staminodes ± 0.8 mm long, yellow.

Ovary obovoid, 3-angled, ± 2 mm long, densely glandu-

lar-puberulous; style terete, branching ± 0.5 mm above

mouth of tube, branches narrowly elliptic-lanceolate,

obtuse, ± 1 mm long, yellow, lateral margins stigmatic.

Achenes heteromorphic in some capitula, 3-winged or

unwinged: winged achenes 10-12 x 7-8 mm, body tur-

binate, ± 7 mm long, transversely ribbed and glandular-

scabrid, with cylindrical, apical, trifen-estrate aircham-

ber, windows ovate, ± 2.5 x 2.0 mm, wings translucent,

2-3 mm wide but sometimes one or two only partially

developed; unwinged achenes 0-3, fusiform-rostrate,

10-12 x 1.5 mm, glandular-pubescent, body purplish,

rostrum and stipe greenish. Disc florets functionally

male, numerous; corolla narrowly funnel-shaped, ± 3

mm long, yellow; tube glandular-hairy, ± 2 mm long;

lobes suberect, triangular, ± 1 mm long. Anthers 2 mm

long, yellow; base tailed, tails equalling filament collar;

apical appendage ovate. Ovary ± terete, ± 1.5 mm long,

glandular-pubescent; style terete, on short stylopodium,

shortly bifid, lobes deltoid, acutely papillate with basal

fringe of longer trichomes. Flowering time : Sept.-May.

Figures 1; 3A-C.

Distribution and ecology, restricted to the drier, east-

ern edge of the Cedarberg and Cold Bokkeveld, where

it has been recorded from Matjiesrivier in the south-

ern Cedarberg and the Skurweberg and Swartruggens

ranges in the Cold Bokkeveld (Figure 2) between 1 000

and 1 260 m. The species is a component of arid fynbos

communities on the drier, hotter, northern slopes and the

plants, like many species in the genus, are highly aro-

matic.

Diagnosis and relationhips: Osteospermum norlindh-

ianum belongs to the small group of species in sect.

Trifenestratae that includes O. dentatum and O. conna-

tum DC. and that is characterized by a tufted, perennial

habit with the lower leaves congested and opposite or

subopposite, and annual flowering stems developing lax,

corymbose synflorescences of moderately-sized capitula

with the tendency to produce dimorphic achenes. Most

fruits develop three conspicuous wings but in some

heads up to half of the achenes are unwinged and fusi-

form-rostrate, with reduced or obsolete apical windows.

In this group, O. connatum from the Cedarberg is dis-

tinctive in its mostly subsessile, semi-amplexicaul or

conspicuously auriculate leaves, and relatively small

winged achenes, 5-6 mm long. Both O. norlindhianum

and O. dentatum (Figure 4) from sandy coastal flats

between Saldanha and Hermanus (Figure 2), in contrast,

have leaves narrowed to a petiole-like base and larger

achenes, 9-12 mm long, and the two are superficially

very similar.

Apart from their eco-geographical separation, the

two species are distinguished by details of their foliage

and fruits. The leaves of O. norlindhianum are oblan-

ceolate without basal auricles, mostly 6-9 mm wide and

obscurely toothed, with scattered, relatively long, strag-

gling eglandular hairs on the underside and also along

the margins, rendering them ± ciliate; the involucral

bracts are glandular-puberulous with scarious margins

0.25-0.50 mm wide (Figure 3A); and the seed body in

Bothalia 42,1 (2012)

61

COMPTON HERBARIUM

'NBG02695S7 -0'

COMPTON HERBARIUM

NATIONAL BOTANICAL INSTITUTE, KIRSTEN0OSCH, CAPE TOWN

FIGURE 1. — Osteospermum norlindhianum, Swartruggens, Jardine 1729 (NBG).

62

Bothalia 42,1 (2012)

FIGURE 2. — Distribution of Osteospermum dentatum , o; O. nor-

lindhianum, • ; O. nordenstamii, A.

the winged achenes is conspicuously glandular-scabrid

and ± turbinate, with a broadly cylindrical distal third

containing the air chamber, a broadly ovoid and trans-

versely ribbed central section with a median longitudi-

nal sulcus, and an abruptly narrowed slender basal third

(Figure 3B). The leaves of O. dentatum are oblong to

obovate and generally broader, 10-20 mm wide, mostly

with basal auricles, and usually more coarsely toothed

or lacerate, with both surfaces and the margins scabrid-

ulous with short, stiff eglandular hairs; the involucral

bracts are scabridulous with narrow scarious margins

0.10-0.25 mm wide (Figure 3D); and the fruit body is

smoothly ellipsoid and subglabrous, with a median lon-

gitudinal sulcus through the entire distal two thirds (Fig-

ure 3E).

Hilliard & Burtt (1985) commented on the spatial/

altitudinal separation within species pairs in Calenduleae

in O. attenuation Hilliard & Burtt (± 1 675 to 2 300 m)

and O. grandidentatum DC. (sea level to ± 1 500 m);

and in Dimorphotheca fruticosa (L.) B.Nord. (near sea

level) and D. caulescens (Harv.) Harv. (± 1 650 m). The

new O. norlindhianum (± 1 000 to 1 260 m) and O. den-

tatum (near sea level) constitute a third example of this.

Etymology, named for Tycho Norlindh, whose mon-

ograph on the sub-Saharan members of the tribe (Nor-

lindh 1943) remains current. Norlindh also correctly

annotated the flowering collection Esterhuysen 12709

(BOL) as representing an unnamed species related to

O. dentatum but was prevented from formally describ-

ing and naming it by the lack of fruits. The species was

recently re-collected, in flower and in fruit, by Ivor and

Cora Jardine during their survey of their property in the

Swartruggens.

Additional specimens seen

WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, Matjiesrivier

Nature Reserve, above Easter Cave, 950 m, 32°28'38"S, 19°22'43"E,

(-AD), 24 Apr. 1996, Lechmere-Oerte l 15 (NBG); Matjiesrivier

Nature reserve, W of Zuurfontein west gate, 1 000 m, 32°27'S, 19°24'E

(-AD), 5 Oct. 1997, Lechmer-Oertel 873 (NBG); rocky ridge above

Winkelhaaks River, E of Bokkeveld Sneeukop, (-CD), 20 Apr. 1946,

Esterhuysen 12709 (BOL); Swartruggens, Knolfontein, 60 km NE of

Ceres, 1 259 m, 32°50'58.2"S, 19°37'25.4"E, (-DC), 11 Feb. 2009,

Jardine & Jardine 1093 (NBG).

Osteospermum nordenstamii J.C. Manning & Gold-

blatt, sp. nov.

TYPE. — Northern Cape, 3017 (Hondeklipbaai):

Riethuis quartz fields, 100 m E of road to Kommagas

from Riethuis, 30°04'52.5"S, 17°26'08.7"E, 170 m, (-

AB), 24 Aug. 2011, Helme 7342 (NBG, holo.).

Gnarled, dwarf shrublet to 100 mm high, flowering

on densely leafy short-shoots; vegetative shoots pros-

trate and laxly leafy with intemodes 10-15 mm long,

stems smooth and weakly compressed when young and

flushed reddish, ± 2 mm diam., glabrescent with thinly

scattered, eglandular hairs, developing a pale brown-

ish, corky bark, greyish and rectangularly fissured when

old, 3^1 mm diam. Leaves opposite, decussate, suberect,

oblanceolate, (9—) 1 5—3 0(— 3 5 ) x (3-)4-8 mm, sessile,

connate for up to 3 mm and shortly sheathing at base

of short shoots, obtuse or rarely apiculate, leathery or

sub-succulent, pubescent when young with mix of strag-

gling, eglandular hairs and short, gland-tipped hairs,

especially along margins but subglabrous when fully

expanded, densely woolly in axils, adaxial surface flat

or shallowly concave, margins yellowish-translucent and

homy, prominent when dry, abaxial midrib prominent

basally and decurrent on stem. Capitula heterogamous,

radiate, ± 25 mm diam., shortly pedunculate, solitary

and terminal on short shoots, rarely a second capitulum

developing from an axillary shoot near apex, nodding in

fruit; peduncles 15-25 mm long and 0.5-0. 8 mm diam.

but up to 1.0 mm diam. in fruit, leafless but with 1 or

2 scattered, linear-subulate bracts 3-5 mm long, thinly

pubescent with a mix of straggling, eglandular hairs

and short, gland-tipped hairs, flushed purple; involu-

cre shallowly campanulate, ± 10 mm diam.; involucral

bracts 15-17, sub-uniseriate or biseriate, lanceolate, 5-6

x 1. 5-2.0 mm, acute, ± glabrous or thinly puberulous

with scattered gland-tipped hairs, with scarious margins

0.3-0. 5 mm wide; receptacle flat, glabrous. Ray florets

female-fertile, 20 to 22, tube glandular-pubescent, ± 1

mm long, lamina spreading but recoiling in afternoon,

narrowly elliptic, 4-veined, ± twice as long as involu-

cre, 11-12 x 4 mm, dull yellow; anthers vestigial, free,

reduced to 4 subulate staminodes, yellow; style terete,

branching ± I mm above mouth of tube, branches nar-

rowly elliptic-lanceolate, obtuse, ± 1.5 mm long, yellow,

lateral margins stigmatic; ovary obovoid, 3-angled, ± 2

mm long, glandular-pubescent; achenes homomorphic,

3-winged, 10-12 x 8-10 mm, body narrowly turbinate,

± 7 mm long, subglabrous or sparsely glandular-pubes-

cent, with apical, trifenestrate airchamber, windows sub-

rotund, ± 1 mm diam., wings translucent, flushed purple,

3^1 mm wide. Disc florets functionally male, numerous;

corolla narrowly funnel-shaped, ± 4 mm long, yellow

tipped blackish; tube glandular-pubescent, ± 3 mm long;

lobes suberect, triangular, ± 1 mm long; anthers 2 mm

long, dark purple with yellow filaments; base tailed, tails

equalling filament collar; apical appendage ovate. Ovaty

compressed-ovoid with lateral ribs, ± I mm long, gla-

brous; style terete, bifid, lobes deltoid, acutely papillate

with basal fringe of longer trichomes. Flowering time :

July-Sept. Figures 3F, G; 5; 6.

Distribution and ecology, restricted to the coastal

plain of southern Namaqualand, where it has been col-

lected northeast and southeast of Riethuis in Northern

Cape, and near Koekenaap and Vredendal in West-

ern Cape (Figure 2). Osteospermum nordenstamii is

Bothalia 42,1 (2012)

63

[

F

FIGURE 3. — Achenes and involu-

cral bracts. A-C, Osteosper-

mum norlindhianum , Jardine

1729 (NBG); A, involucral

bracts; B, winged achene;

C, wingless achene; D, E,

O. dentatum', D. involucral

bracts [Langebaan, Bosenberg

& Rutherford 89 (NBG)]; E,

winged achene [Wildevoel-

vlei, O 'Callaghan 704

(NBG)]; F, G, O. nordens-

tamii, Helme 7342 (NBG)];

F, involucral bract; G, winged

achene. Scale bar: A, D. F,

1 mm; B, C, E, G, 2 mm. Art-

ist: J.C. Manning.

locally common among white quartz pebbles, mainly (Schmiedel & Jurgens 1999). The two known areas

on north-facing slopes, and is yet another highly local- of occurrence at Koekenaap and Riethuis are some

ized endemic of the Namqualand lowland quartz fields 175 km apart with very little suitable quartz patch habi-

64

Bothalia 42,1 (2012)

, ’ j. A

UNIVERSITY OF CAPE TOWN.

HERBARIUM AFAICANUM 60LUSIANUM.

hlo«« fax i ,-iv

. , MiOJ

- cx^4*n.i4-nv Oji-x.ta.CjM. *)$Uui y

Life V'iivt;

Fi"' W a.m. y,» u*in^ tffLee,

FIGURE 4. — Qpeospermum dentatum, Kommetjie, Bolus 23203 (NBG).

Bothalia 42,1 (2012)

65

FIGURE 5. — Flowering plant of Osteospermum nordenstamii at the

type locality near Riethuis. Photographer: N.A. Helme.

FIGURE 6. — Fruiting plant of Osteospermum nordenstamii at the type

locality near Riethuis Photographer: N.A. Helme.

tat between them. Low rainfall at both places is sup-

plemented by substantial precipitation from coastal fog,

especially in autumn and winter.

At the type locality near Riethuis the species grows

with the locally endemic succulents Drosanthemopsis

vaginata, Meyerophytum meveri, Conophytum conca-

vum and Monilaria globosa (Aizoaceae), and with the

more widespread Hirpicium alienatum (Asteraceae) and

Cheiridopsis robusta (Aizoaceae). An undescribed spe-

cies each of Othonna (Asteraceae) and of Eriospermum

(Ruscaceae) were also recorded in the area. The two

southern localities fall within the Knersvlakte centre of

endemism (Van Wyk & Smith 2001).

Diagnosis and relationships'. Osteospermum norden-

stamii is readily distinguished from all other species by

its dwarf habit (Figures 5 & 6), forming small, gnarled,

± creeping shrublets less than 10 cm high, with decus-

sate, oblanceolate leaves with entire margins, mostly

obtuse apices, and distinctly woolly axils. Plants pro-

duce prostrate or spreading, relatively distantly leafy

vegetative shoots but flower only from closely leafy

short-shoots, which produce solitary, dull-yellow capit-

ula on short, terminal peduncles. The leaves have dis-

tinctive translucent, homy margins especially evident in

dried herbarium material.

The opposite, entire, subglabrous leaves of Oste-

ospermum nordenstamii are otherwise found only in

O. oppositifolium among the species of sect. Trifenes-

tratae, and suggest that its relationships lie here. The

two species are readily separated by their habit and also

by bark, foliage and inflorescence. O. oppositifolium is

common thoughout western Namaqualand as far south

as Clanwilliam, forming colonies on stony and rocky

slopes. It is a rounded shrub, mostly 0. 5-1.0 m high but

occasionally as low as 30 cm, with linear to oblanceo-

late, mostly acute leaves, 20-60 x 2—1 0(— 1 2) mm, with

the margin only weakly thickened and without woolly

axils (rarely a few sparse hairs are present). The stems

develop relatively thin, longitudinally striate bark quite

unlike the corky, quadrately fissured bark of O. nor-

denstamii, and the capitula are typically borne in lax

corymbs at the ends of the branches, not on short shoots.

They are mostly larger than in O. nordenstamii, 25-35

mm diam., and the ray florets vary in colour from rich

golden yellow in Namaqualand to pale yellow in the

Olifants River Valley. Both species have been recorded

at the same locality north of Koekenaap ( Nordenstam

907).

Etymology’: named for Bertil Nordenstam, who has

published extensively on Asteraceae in general and on

the tribe Calendulae in particular, and who first collected

the species in 1962.

Other specimens seen

NORTHERN CAPE. — 3017 (Hondeklipbaai): Riethuis/Oubees,

30°7'49"S, 17°25'20"E, quartz patch, 150 m, (-AB), 24 Aug. 1999,

Desmet 244 (NBG); S of Riethuis on track to Taaibosduin, quartzite

koppie, (-AB), 13 Aug. 2006, Koekemoer 3342 (PRE); along road to

Soebatsfontein from Wallekraal/Hondeklipbaai road, 161 m, (-BC), 16

Sept. 2000, Koekemoer & Funk 1957 (PRE).

WESTERN CAPE.— 31 18 (Vanrhynsdorp): 3 miles [4.8 km] NE of

Koekenaap, (-AD), 15 July 1962, Nordenstam 581 (S); Moedverloor,

4 miles [6.4 km] N of Holrivier Bridge, quartzite fields and patches,

(-AD), 10 Aug. 1962, Nordenstam 907 (S); 4 miles [6.4 km] N of

Koekenaap, quartzite kopje W of road, (-AD), 25 Aug. 1962, Nor-

denstam 1022 (S); 4 km N of Koekenaap, quartzite outrcrop, (-AD),

6 Sept. 1974, Nordenstam & Lundgren 1686 (S); Koekenaap, 31.46°S,

18.31°E, (-AD), without date, Desmet 3636 (NBG); Koekenaap, por-

tion of old Farm 630, 50 m, (-CB), 16 Aug. 1997, Desmet & Ellis

1243 (NBG).

ACKNOWLEDGMENTS

Material was collected under permits from the con-

servation authorities of Northern and Western Cape.

BOL, NBG, PRE and SAM, the main herbaria with

good representation of collections of Cape species, were

consulted for records of the two new species (herbarium

acronyms after Holmgren et al. 1990). We thank cura-

tors of these herbaria for allowing access to their collec-

tions, and the referees for their constructive comments.

REFERENCES

HILLIARD, O.M. & BURTT, B.L. 1985. Notes on some plants of

southern Africa chiefly from Natal XI. Notes from the Royal

Botanic Garden Edinburgh 42: 227-260.

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.

66

Bothalia 42,1 (2012)

MANNING, J.C. & GOLDBLATT, P. 2008. Tripteris calcicola (Aster-

aceae), a new calciphilous species from Western Cape, South

Africa. Bothalia 38: 85-88.

MANNING, J.C. & GOLDBLATT, P. in press. Plants of the Greater

Cape Floristic Region 1 : The Core Cape Flora. Strelitzia #. South

African National Biodivsersity Institute, Pretoria.

NORDENSTAM, B. 1994. New combinations in the Calenduleae.

Compositae Newsletter 25: 46-M9.

NORDENSTAM, B. 2006. Generic revisions in the tribe Calenduleae

(Compositae). Compositae Newsletter 44: 38-49.

NORDENSTAM, B. 2007. Tribe Calenduleae Cass. In J. Kadereit &

C. Jeffrey (vol. eds), K. Kubitzki (series ed.). The families and

genera of vascular plants. VIII Flowering plants-Eudicots-Aste-

rales, pp. 241-245. Springer, Berlin, Heidelburg, New York.

NORDENSTAM, B. & KALLERSJO, M. 2009. Calendulae. In V.A.

Funk, A. Susanna, T.F. Stuessy & R.J. Bayer (eds). Systematics,

Evolution and Biogeography of Compositae'. 293-314. Interna-

tional association for Plant Taxonomy, Vienna.

NORLINDH, T. 1943. Studies in the Calenduleae. I. Monograph of the

genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and

Chrysanthemoides. Bloms, Lund.

SCHMIEDEL, U. & JURGENS, N. 1 999. Community structure on unu-

sual habitat islands: quartz-fields in the Succulent Karoo, South

Africa. Plant Ecology 142: 57-69.

SN1JMAN, D. in press. Plants of the Greater Cape Floristic Region 2:

The Extra-Cape Region. Strelitzia #. South African National

Biodiversity Institute, Pretoria.

VAN WYK, A.E. & SMITH, G.F. 2001. Regions of floristic endemism

in southern Africa. Umdaus, Hatfield.

J.C. MANNING*,** P. GOLDBLATT*** & N.A. HELME****

* Compton Herbarium, South African National Biodiversity Institute,

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

sanbi.org.za.

** Research Centre for Plant Growth and Development, School of Life

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

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.

**** P.O. Box 22652, Scarborough 7975, South Africa.

MS. received: 2012-01-24.

Bothalia 42,1: 67-68 (2012)

OBITUARY

EDUARDO JOSE DOS SANTOS MORE1RA MENDES (1924-2011 )

On 24 September 2011, Eduardo Mendes (Figure

1 ) passed away at his home in Lisbon, Portugal, at the

grand old age of 86 — just 2 months short of his 87th

birthday — following a stroke. He was one of the last of a

generation of colonial plant taxonomists who had exper-

tise on the flora of Angola and some of the other former

African colonies of Portugal, and his passing signifies

the end of an era.

Eduardo Jose dos Santos Moreira Mendes was born

in Lisbon on 26 November 1924. He studied in Lisbon

and obtained his first degree in biological sciences in

1946 at the University of Lisbon where he studied under

the tutelage of, among others, the renowned cytogeneti-

cist, Prof. Flavio Resende, who had a special interest

in southern African petaloid monocots, including aloes

and their kin (Smith & Figueiredo 201 la). Mendes was

a botany lecturer at the University of Lisbon from 1 946

to 1954, where he taught courses in the Faculties of Sci-

ences, Pharmaceutical Sciences, and Medicine. During

that period he developed a research programme on cryp-

togams, and published several papers on this group. In

1954 he was employed as assistant to the project ‘Mis-

sao Botanica de Angola e Mozambique’ and in 1956

became a researcher at the Junta de Investigaqoes do

Ultramar, later superseded by the Instituto de Investi-

gaqao Cientifica Tropical (IICT). From 1974 to 1986,

Mendes was Director of the Centro de Botanica of the

IICT, in Lisbon, Portugal. While Director of the Centro,

he used an interesting way of summoning staff to his

office: a switch on his desk was connected to a bell that

FIGURE 1. — Eduardo Mendes (26 November 1924-24 September

2011) at his home in Lisbon, Portugal. Photograph: Gideon F.

Smith, 23 September 2010.

rang in the Institute’s corridors and each staff member

had a Morse code-style code that, when sounded, meant

that he or she must immediately go to the Director’s

office (Martins 1994).

Throughout his career Mendes maintained an interest

in the floras of Angola and Mozambique, and did con-

siderable work on these. He revised several families for

the Conspectus florae angolensis ; publishing a series of

papers under the title ‘Additiones et Adnotationes Florae

Angolensis’ that continued until 1973. He further revised

several families for the Flora zambesiaca in a set of

papers under the series entitled ‘Additiones et Adno-

tationes Florae Mozambicanae’. During his career he

published over 70 papers and was editor or co-editor of

Conspectus florae angolensis , Flora zambesiaca , Flora

de Mozambique, and Garcia de Orta, Serie Botanica.

Mendes participated in two botanical expeditions

to Angola, during which he collected several thousand

herbarium accessions (Figure 2). The first expedition

that Mendes undertook to Angola took place from Sep-

tember 1955 to March 1956 and focused mostly on the

southwestern parts of the country, where he collected

1 750 numbers (Figure 3). The second expedition took

place between December 1959 and May 1960, mostly

in the southeastern region, between Cuvango and Cuito-

Cuanavale, and resulted in the numbers 1 800 to 4 064

in Mendes’s African collection (Mendes 1980; Martins

1994).

FIGURE 2. — A young, mustachioed Eduardo Mendes photographed in

Angola during the first of the two expeditions he undertook to

the country. Photographer unknown. Reproduced courtesy of the

Mendes family, Lisbon, Portugal.

68

Bothalia 42,1 (2012)

FIGURE 3. — A series of tents pitched around the field collecting

vehicle served as shelter during the first of the expeditions that

Mendes undertook to Angola. Photograph: Eduardo Mendes.

Reproduced with the permission of the Instituto de Investiga^ao

Cientifica Tropical, Lisbon, Portugal.

After his retirement from the Centro de Botanica,

Mendes disappeared from the botanical landscape. Two

of the authors (GFS and EF) visited him at his home on

23 September 2010 to obtain information on the prov-

enance of the material on which the name Aloe mendesii

Reynolds (Asphodelaceae), a cliff-dwelling species from

Angola, was based. He was very helpful and vividly

remembered the material, including when and where it

was collected and what happened to it — quite remarka-

bly— over 55 years ago! However, Mendes sadly passed

away one week before the resulting paper appeared in

print (Smith & Figueiredo 201 lb).

The work and collections of Mendes remain lit-

tle known in South Africa as he never collected in the

country as far as we could ascertain, and was therefore

not included in Gunn & Codd’s (1981) benchmark work

on the botanical exploration of southern Africa, and was

only briefly mentioned in Figueiredo & Smith (2008).

However, his collecting activities in Angola contributed

significantly to the primary botanical knowledge of that

country, and some of the specimens that he collected

decades ago are still being used as the types of new

plant names. Sousa et al. (2010), for example, recently

described Cyphostemma mendesii F. Sousa in the Vita-

ceae based on some of Mendes’s Angolan material. Thir-

teen further taxa were described from, and named for,

Eduardo Mendes from collections he made in Angola.

Eleven are currently accepted, including the distinctive

succulent tree Euphorbia eduardoi L.C. Leach, Uvaria

mendesii Paiva, Ceropegia mendesii Stopp, Phyllanthus

mendesii Brunei ex Radcl.-Sm., Adenodolichos mendesii

Torre, Crotalaria mendesii Torre, Indigofera mendesii

Torre, Vigna mendesii Torre, Rotheca mendesii (R.Fem.)

R.Fern., and Aloe mendesii Reynolds.

REFERENCES

FIGUEIREDO, E. & SMITH, G.F. 2008. Plants of Angola / Plantas de

Angola. Strelitzia 22: 1-282. South African National Biodiver-

sity Institute, Pretoria.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. A. A. Balkema, Cape Town.

MARTINS, E.S. 1994. Homenagem a botanicos com colaborafao rele-

vante no Centro de Botanica do IICT. I. Eduardo Mendes. Garcia

de Orta. Serie Botanica, 12: 11-14.

MENDES, E.J. 1980. A Junta de Investigates Cientlficas do Ultramar

e a flora de Africa: Missoes e Centro de Botanica. Boletim da

Sociedade Broteriana, Serie 2 54: 201-215.

SMITH, G.F. & FIGUEIREDO, E. 2011a. Obituary. Franz Sebastian

Muller (1913-2010). Bothalia 41 : 365-368.

SMITH, G.F. & FIGUEIREDO, E. 2011b. Provenance of the material

on which the name Aloe mendesii Reynolds (Asphodelaceae),

a cliff-dwelling species from Angola, is based. Bradleya 29:

61-66.

SOUSA, F„ FIGUEIREDO, E. & SMITH, G.F. 2010. Cyphostemma

mendesii (Vitaceae), a new species from Angola. Phytotaxa 7:

35-39.

G.F. SMITH*, E. FIGUEIREDO** and

L. CATARINO***

* Office of the Chief Director: Biosystematics Research & Biodiversity

Collections, South African National Biodiversity Institute, Private Bag

XI 01, Pretoria, 0001 South Africa / H.G.W.J. Schweickerdt Herbarium,

Department of Plant Science, University of Pretoria, Pretoria, 0002

South Africa / Centre for Functional Ecology, Departamento de Cien-

cias da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal.

Email: g.smith@sanbi.org. za.

** Department of Botany, P.O.Box 77000, Nelson Mandela Metropo-

litan University, Port Elizabeth, 6031 South Africa / Centre for Func-

tional Ecology, Departamento de Ciencias da Vida, Universidade de

Coimbra, 3001-455 Coimbra, Portugal Email: estrelafigueiredo@hot-

mail.com.

*** Jardim Botanico Tropical, Instituto de Investigate Cientifica Tro-

pical, Trav. Conde da Ribeira 9, 1300-142 Lisboa, Portugal. Email:

lmfcatarino@gmail.com.

Bothalia 42,1 : 69-70 (2012)

OBITUARY

KATHLEEN DIXON GORDON-GRAY (NEE HUNTLEY) (1918-2012)

Kathleen Gordon-Gray passed away peacefully on 13

January 2012, eleven days before her 94th birthday. She

is survived by her only daughter Celia.

Gordon-Gray was no ordinary person; she was in fact

a most extraordinary and generous woman. Not only

was she the South African expert in Cyperaceae and

several other plant families occurring in KwaZulu-Natal

(KZN) (see scholar.google.co.za for a list of some of her

publications), but she was a wife, mother, and an excel-

lent lecturer in the then Botany Department at the Uni-

versity of Natal (now UKZN) on the Pietermaritzburg

campus. Gordon-Gray was also a much sought after

confidant and friend to students and colleagues alike.

It was her gentle and caring manner, ability to listen

and her almost infinite patience and desire for perfec-

tion that were her enduring qualities. I never heard her

speak ill of anyone and, if she had a fault, it was that

she accepted her lowly academic status in the Depart-

ment— as women often did in those days; foregoing

promotion to realize her professional calling. Even to

the last, wracked with arthritis, she was alert and work-

ing on Cyperaceae with Jane Browning and another of

her close co-workers C.J. (Roddy) Ward, a well-known

KZN plant collector and field ecologist (and ex-student

from her earliest days as a Junior Lecturer).

Kath Gordon-Gray was born in Pietermaritzburg and

graduated with a B.Sc. and M.Sc. (1939). Her Ph.D.

was awarded in 1959 (all degrees from Natal University,

now UKZN). After teaching at Girls Collegiate School

(1940-1945), she was appointed in the Botany Depart-

ment as a Junior Lecturer to teach mainly ex-service

men; later as Lecturer (1951-1966 (Gunn & Codd

1981)) under Professor A.W.J. Bayer who was Head of

Department at the time; as Senior Lecturer ( 1967-1976,

after Professor Bayer retired); and finally as Associate

Professor (1977-1978). She was, sadly, required by the

regulations of the time, to retire at the age of 60 in 1978

(Glen & Germishuizen, 2010); but continued to work on

her beloved KZN plants and the Cyperaceae in particu-

lar.

Gordon-Gray was essentially a ‘home’ girl, collecting

some 4 000 specimens mainly from KZN. My fondest

memories of her date back to the 1960s when the then

Botany Department in Pietermaritzburg was arguably

in its heyday. From the late 1950s to the early 1970s,

a remarkable number of botanists passed through the

Department (Moll 1987) and many undergraduates and

postgraduates came under the spell of Doc G-G, as she

was fondly known. Personally, if it had not been for

Gordon-Gray, I would have become a zoologist and I

know of many others who pursued a career in botany

because of her influence. Thus, almost single-handedly,

she recruited many people into botany who later became

FIGURE 1. — Three keen botanists sorting through live material of Aca-

cia robusta in the herbarium that had been collected for leaf and

pod measurements as part of a study to quantify the difference

between the two subspecies of ,4. robusta subspp. clavigera and

robusta. Left to right Eugene Moll, Kath Gordon-Gray, and Ken

Tinley (Photographer: J.H. Ross).

well known in their particular fields, and who then went

on themselves to recruit other botanists and ecologists.

Some noteworthy students and colleagues until the

1980s were Don Killick, Roddy Ward, Denzil Edwards,

Mike Wells, Clare Archer (Reid), Jim Ross, Fiona

Getliffe, Roger Ellis, Charles Stirton, Esme Hennessy,

Ken Tinley, Brian Downing, Brian Huntley, Ed Granger,

and Trevor Arnold.

One thing that she is well known for is her annota-

tions on herbarium sheets, that are often accompanied by

her meticulous drawings and notes of what she believed

to be diagnostic features (for example see Goetghebeur

& Coudijzer, 1985: 245), and on JSTOR some sheets

such as the Type of Fimbristylis bequaertii De Willd.

[Family Cyperaceae]; BR0000008639400, have illus-

trations by Gordon-Gray attached (J. Browning pers.

comm. January 2012).

When she died, Gordon-Gray was still working on

Cyperaceae with Jane Browning who compiled a com-

prehensive list of notes on her co-worker that she kindly

shared with me. ft was typical of Gordon-Gray to shun

the lime-light. She was however belatedly awarded the

South African Association of Botanists’ Silver Medal in

1998, though she certainly deserved greater recognition

earlier in her career. As she disliked being photographed,

pictures of her are rare and she shunned National and

International Meetings; preferring to work alone and in

close collaboration with those she knew and trusted. Yet.

through her teaching and supervision, she was inspira-

tional and provided sound theoretical and practical train-

ing in the fundamentals of taxonomy, anatomy, breed-

70

Bothalia 42,1 (2012)

ing systems and the ecology of her beloved sedges and

grasses.

ACKNOWLEDGEMENTS

Thanks to Celia Gordon-Gray, Jane Browning, and

Roddy Ward for reading and commenting on this obituary.

REFERENCES

GLEN, H.F. & GERMISHUIZEN, G. (compilers) 2010. Botanical

exploration of southern Africa, edition 2. Strelitzia 26. South

African National Biodiversity Institute, Pretoria.

GOETGHEBEUR, P. & COUDIJZER, J. 1985. studies in Cyperaceae

5 — the genus Bulbostylis in Central Africa. Bulletin du Jardin

Botanique National de Belgique 55: 207-259.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. A. A. Balkema, Cape Town.

MOLL, E.J. 1987. What future the ecological profession? Bulletin of the

South African Institute of Ecologists 6: 30, 31

E.J. Moll*

* Extraordinary Professor, Department of Biodiversity and Conserva-

tion Biology, University of the Western Cape, Bellville. E-mail: emoll@

telkomsa.net.

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BOTHALIA

Volume 42,1 May 2012

CONTENTS

1. Revision of the genus Sphenostylis (Fabaceae: Phaseoleae) in South Africa and Swaziland. A.N.

MOTEETEE and B.-E. VAN WYK 1

2. Anatomy of myxospermic diaspores of selected species in the Succulent Karoo, Namaqualand,

South Africa. H. FOTOUO MAKOUATE, M.W. VAN ROOYEN and C.F. VAN DER

MERWE 7

3. Recircumscription and distribution of elements of the ‘ Ceterach cordatum' complex ( Asplenium :

Aspleniaceae) in southern Africa. R.R. KLOPPER and N.R. CROUCH 15

4. A revision of Tecophilaeaceae subfam. Tecophilaeoideae in Africa. J.C. MANNING and P.

GOLDBLATT 21

5. Notes on African plants:

Aizoaceae. Corpuscularia gracillima is the correct binomial for the illegitimate C. gracilis. J.

NIEDERLE 49

Asteraceae. Berkheya jardineana (Arctotideae — Gorteriinae), a new dwarf perennial from the

Swartruggens, Western Cape. J.C. MANNING and P. GOLDBLATT 57

Asteraceae. Osteospermum norlindhianum and O. nordenstamii, two new species of

Osteospermum sect. Trefenestratae (Calenduleae) from the Greater Cape Floristic

Region. J.C. MANNING, P. GOLDBLATT and N.A. HELME 59

Fabaceae. A new species of Banhinia from southern Mozambique and the reinstatement of

Bauhinia macrantha. E.J.D. SCHMIDT 44

Hyacinthaceae. Drimia echinostachya , correcting a nomenclatural oversight. U. EGGLI and

N.R. CROUCH 43

Hyacinthaceae. New combinations in Ledebouria. J.C. MANNING and P. GOLDBLATT 47

Iridaceae. Taxonomic notes on Moraea (Irideae) in Congo and adjacent countries. P.

GOLDBLATT and J.C. MANNING 49

Monimiaceae. Taxonomic and nomenclatural notes on the monotypic genus Xymalos and

general information on the family Monimiaceae. M. JORDAAN and M. LOTTER 5 1

6. Obituary. Eduardo Jose dos Santos Moreira Mendes (1924-201 1). G.F. SMITH, E. FIGUEIREDO

and L. CATARINO 67

7. Obituary. Kathleen Dixon Gordon-Gray (nee Huntley) (1918-2012). E.J. MOLL 69

Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • AGRIS • BIOSIS: Biological Abstracts /RRM • CABS • CABACCESS • CAB

ABSTRACTS • ISI: Current Contents, Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon: reviews and notices.

ISSN 006 8241

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