MARY

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Volume

BOTHALI A

9 Jaargang 9

Published by the

Botanical Research Institute

Department of Agricultural Technical Services

Gepubliseer deur die

Navorsingsinstituut vir Plantkunde

Departement van Landbou-Tegniese Dienste

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\ MMN- hitllTIlUT VIR PLANTKUNDE

Edited bv — Onder redaksie van

L. E. Codd. D.Sc.,

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prRpE-T CRIA-

REo/ INSTITUTE

Chief : Botanical Research Institute, Pretoria

Hoof : Navorsingsinstituut vir Plantkunde, Pretoria

THE GOVERNMENT PRINTER-DIE STAATSDRUKKER, PRETORIA

G.P.-S. 36919-1968-69-1,000

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CONTENTS - INHOUD

Page / Bladsy Date/ Datum

Part 1 1-250 30 November 1966

Part 2 251-362 4 August/Augustus 1967

Parts 3 & 4 363-513 7 May/Mei 1969

Page

Bladsy

Anderson, J. G. The genus Andropogon in Southern Africa 5

A new combination in Hyparrhenia 130

A new species of Panicum 341

A new variety of Festuca costata 341

Badenhorst, H. Melastomaceae: a note on Memecylon australe Gilg & Schltr.

and M. grandiflorum R. & A. Fernandes 345

Codd, L. E. The Cassine complex 123

A new species of Canthium 345

New species of Kniphofia 139

Notes on Boerhavia in Southern Africa 113

The South African species of Kniphofia 363

The status of the genus Lydenburgia 124

De Winter, B. A new species of Eragrostis 137

Polexansia De Winter, gen. nov 130

Styppeiochloa De Winter, gen. nov 134

Gunn, M. D. Ethel Mary Doidge (1887-1965) 251

Hilliard, O. M. A new species of Streptocarpus ( Gesneriaceae ) 335

Jessop, J. P. The genus Asparagus in Southern Africa 31

Killick, D. J. B. Two new Erica species from the Drakensberg 126

Klesser, Patricia J. A virus disease of Datura spp 173

Leistner, O. A. The typification of Podocarpus elongatus 142

Marais W. Notes on South African Cruciferae 97

Marasas, W. F. O. New species of Ascomycetes and a new genus of Sphaeropsi-

daceae from Transvaal 203

Marasas, W. F. O. & Rabie, C. J. New and interesting records of South African

Fungi. Part IV 217

Marasas, W. F. O. & Schumann, I. H. Two species of Erysiphaceae from Pre-

toria 245

Page

Blaclsy

Marasas, W. F. O., Van der Westhuizen, G. C. A., Van Warmelo, K. T. &

Papendorf, M. C. New and interesting records of South African Fungi.

Part V ~. 229

Marsh, J. A. Notes on Widdringtonia 124

Obermeyer, A. A. The identity of Bidbinella carnosa 345

The identity of Frit Maria nana 345

Mrs. Reino Pott-Leendertz (1869-1965) and the Transvaal Museum Herbarium 1

A new combination in Ornithogalum 344

A new combination in Trachyandra 344

Two new species of Bulbine 342

Raven, P. H. A revision of African species of Epilobium ( Onagraceae ) 309

Simons, R. H. The genus Ceramium in South Africa 153

The genus Microcladia (red algae) in South Africa 351

A new species of brown alga from South Africa 357

A new species of Dictyotales from South Africa 169

Tolken, H. R. The species of Arthrocnemum and Salicornia ( Chenopodiaceae )

in Southern Africa 255

Van Warmelo, K. T. The cytology of Mycosphaerel/a pinodes 195

The identity of a pea blight fungus in South Africa 183

Verdoorn, I. C. A new species of Dombeya and a new variety of Dombeya

rotundifolia 143

Plate 1. — Mrs. Reino Pott-Leendertz (1869-1965); photograph taken about 1910

7230691

Botha Ha 9, 1 : 1-3.

Mrs. Reino Pott-Leendertz (1869-1965)

and the

Transvaal Museum Herbarium

by

A. A. Obermeyer

Reino Leendertz, the founder of the Transvaal Museum Herbarium, was born

in Medemblik, Holland, on the 2nd January, 1869. She qualified as a pharmacist

in 1891 but then turned her attention to Botany. At Amsterdam she studied under

Professor Hugo de Vries and afterwards became one of his assistants. In July, 1898,

she arrived in Pretoria and was appointed Botanist to the State Museum on the 1st

September, 1898. In her letter of appointment the Director, Dr. J. W. B. Gunning,

mentioned that, although it was contrary to the policy of the Committee to appoint

women on the scientific staff, her exceptional qualifications decided them to waive

this objection. Thus she became the first official botanist of the Transvaal as well as

its first woman civil servant. A large collection of plants had been purchased from

Rudolf Schlechter and these formed the nucleus of the Transvaal Museum Herbarium.

The State Museum was then housed in a building on the Market Square in Pretoria,

the part that is presently used as the covered market. This building, which was rented

for £10 per month, was officially opened by President Paul Kruger in May, 1894. It

soon proved to be too small and plans were made to put up a new building in Boom

Street, where the Zoological Garden was being constructed. A photograph of the

stone-laying ceremony for this new building, on the 22nd July, 1899. shows Miss

Leendertz as the only woman among the museum officials.

War broke out and Pretoria came under the English in 1901. The British com-

mander asked her to stay, but Miss Leendertz decided to return to Holland. At a

girls’ high school in Amsterdam she taught Natural History for a year but she did not

like teaching (in after years she confided to a niece that she had difficulty in keeping

order). So she re-applied for her former post at the museum, now called the Pretoria

Museum, and was accepted immediately, since the Director had a high opinion of

her capabilities. She returned to Pretoria in 1904.

She threw herself with heart and soul into the work and started a special Transvaal

herbarium. She collected extensively in the Transvaal, her numbers eventually reaching

nearly 6,000. Her first excursions were done by bicycle. In the beginning a Mr.

B. H. Rademacher and his sister accompanied her; later Mr. A. T. J. Janse (now

Professor Janse), who was on the staff of the Normal College, became her enthusiastic

companion during week-end excursions. At one stage they decided to make an exhibit

of the Transvaal timber trees. The trees were photographed; wood, bark, flowering

branches and fruits were all loaded onto the bicycles and taken home. Mr. Haak Snr.

who was the owner of the bicycle shop (which afterwards became Haak’s Garage),

once happened to see such a fully loaded bicycle and remarked that they were not

meant to be used as wagons.

Quite often she used her holidays to go on collecting expeditions further afield.

Gatherings from Ermelo (with a beautiful kloof, Mavieriestad, nearby), Zeerust, Bethal,

Potchefstroom, Standerton, Vereeniging, Moorddrift (near Potgietersrus), Heidelberg

and some spots nearer Pretoria (Daspoort. Kameeldrift, Roodeplaat. Irene) are localities

regularly met with in the herbarium.

2

The collection now expanded by leaps and bounds as can be seen from the

following extract from the Annual Report of the Transvaal Museum, 1905-1906:

“ I collected in the neighbourhood of Pretoria 84 species of plants in many duplicates,

nine species of wood, many fruits and seeds to complete the collections. On a botanical

excursion to the Zoutpansberg District, where I stayed at Marabastad, Waterval,

Rietfontein and Eersteling — only a fortnight — I collected 105 species of plants in many

duplicates. Most of them were new to the collection “ Transvaal Plants ”. On a five

days’ excursion to Crocodile River and Skeerpoort, I collected 68 species of plants

of which 50 species were new for the Transvaal collection: and on one to Rooiplaat

(Roodeplaat), Derdepoort, also of five days, 62 species of plants of which 40 species

were new to the collection ”.

In about 1906 the Reverend F. A. Rogers of the South African Church Railway

Mission is also mentioned by her as an assiduous collector, who regularly sent her

large parcels of plants from different localities. The plants were just put between

sheets of paper by him and Miss Leendertz did the rest, the pressing, naming, etc.

She also made duplicate sets for the collector. Other collections, which were purchased

by the Museum, were a collection of Ferns and Fern Allies from Mr. T. R. Sim. Miss

Leendertz pointed out that it was a standard collection, Mr. Sim having based his

book “ The Ferns of South Africa ” on it. Further additions were collections purchased

from Mr. G. N. Thorncroft, of Barberton, and the missionary, Henri Junod of Shilouvane

(near Tzaneen).

No wonder that soon the universal complaint was heard; the boxes housing the

collections “ were filled to the top ” and that she had no assistant to help her with the

pressing, mounting and labelling. The neat sheets with her copy-book handwriting

are a joy to behold. She was an extremely good taxonomist and hardly ever made

mistakes in her identifications although the literature at her disposal in those days

was limited.

In her next report she informs us that a junior assistant was appointed in September,

1907, and that new boxes had arrived. Incorporated in the herbarium that year were

2,436 species of plants and she remarked that “ many families are now fairly well

represented and form useful material for study and reference ”. She then again refers

to the Ferns and Fern Allies collection which through additions from Junod, Burtt

Davy and others had become one of the best in South Africa. She also thanks Dr.

Harry Bolus for his co-operation. In conclusion she mentions that the “ Preliminary

List of Transvaal Vascular Plants which Mr. J. Burtt Davy and myself have prepared ”,

had been read as a paper at the Grahamstown Meeting of the South African Asso-

ciation for Advancement of Science.

She also spent a good deal of time on the preparation of a botanical exhibition,

filling 8 show cases with examples of flowering timber trees of the Transvaal with their

Latin and vernacular names, and 16 other cases “ illustrating the biology of plants,

e.g. insectivorous plants, parasites, poisonous plants, etc., all provided with descriptive

labels in both languages ”,

In April, 1908, the Director, in a report presented in the first issue of the “Annals

of the Transvaal Museum ” informed us that the Herbarium contained 4,000 specimens.

In 1915 the number of sheets catalogued had risen to 14,400 and when she left to go on

pension on the 2nd January, 1925, the last number entered by her was 23,255.

Before that, in 1912, the Natural History and Geological collections of the Transvaal

Museum had been moved to a new building in Paul Kruger Street.

Staff members of the Museum, when on field trips, added many plants to the

herbarium. We frequently meet the names Dr. H. G. Breyer (at one time a Director

of the Museum) and his son, when the latter was stationed at Fort Namutoni in South

3

West Africa; C. Wilde, T. J. Jenkins, C. J. Swierstra, V. Fitzsimons, G. van Dam.

J. Hewitt, M. Collins, Austin Roberts, were industrious collectors. The Maputaland

Expedition, which visited Mozambique in 1914, also yielded a number of plants.

Names of well known South African collectors are also represented in the herbarium,

such as Wilms (apparently his early gatherings were not numbered), F. A. Rogers,

G. Thorncroft, H. Junod, W. Nelson, W. A. H. Harbor, M. Stewart, R. D. Bradfield,

R. E. Symons, J. Medley Wood, Alice Pegler, Florence Paterson, Daly & Cherry,

H. G. Flanagan, P. Krook, A. Penther, W. Tyson, Barbara Smart, Theo Kassner,

A. Stolz, etc.

Together with other well known scientists such as Sir Arnold Theiler, Prof. A. T. J.

Janse and Dr. I. B. Pole Evans, Miss Leendertz was a foundation member of the

Biological Society which was started in Pretoria in 1907. In 1909 she married Mr.

N. W. Pott.

From the time of her retirement in 1925, her interest in botany appears to have

ceased but to the last she remained active and alert and a real lady, much loved by all

around her. After a short illness she passed away on the 14th June, 1965, at the age

of 96.

A few plants were named after her, the best known being Stapelia leendertziae.

There is also a Blepharis leendertziae Oberm., Delosperma leendertziae N. E. Br.,

Indigofera leendertziae N. E. Br., Lotononis pottiae Burtt Davy and Thesium pottiae

N. E. Br. A list of her publications is appended.

Upon her retirement. Miss D. Kaplan succeeded her in 1925. In 1927 Mr. R. G. N.

Young took over and in 1929 Miss A. A. Obermeyer, who remained there until her

marriage in 1938. In 1953 the Board of Trustees of the Transvaal Museum donated

the herbarium, now consisting of about 40,000 numbers, to the National Herbarium,

Pretoria.

Acknowledgements

It was a pleasure to hear Professor A. T. J. Janse reminisce on the time that he

and Miss Leendertz went out collecting on bicycles and I thank him heartily for the

information so freely given. Grateful thanks are also due to Mrs. E. Patterson, a

niece, who lent me the few but important letters and documents Mrs. Pott had

preserved. These were very helpful indeed in building up a picture of her early years

at the Museum. Dr. V. Fitzsimons, the present Director, kindly lent us the photograph

reproduced here.

List of Publications by Mrs. R. Pott-Leendertz

1. The Amaryllidaceae of the Transvaal in Ann. Transv. Mus. 1: 46 (1911). Enumerating 11 genera

and 36 species.

2. A first checklist of the flowering plants and ferns of the Transvaal and Swaziland. J. Burtt Davy

and R. Pott-Leendertz in Ann. Transv. Mus. 3 : 119-182 (May, 1912). 3,240 species are enumerated.

3. A new species of Stapelia from the Transvaal, Stapelia gettleffii Pott in Ann. Transv. Mus. 3: 226,

1. 1 3 (1912).

4. A new record of plant species for the Transvaal, Duvalia polita N. E. Br. in Ann. Transv. Mus.

3: 226 (1912).

5. New species of Alepidea in Ann. Transv. Mus. 4: 206 (1914). A. jenkinsii Pott ; A. basinuda Pott.

6. A new species of Warburgia from the Transvaal, in Ann. Transv. Mus. 6: 60, fig. 1-2 (August 1918).

7. Addendum to the first checklist of the flowering plants and ferns of the Transvaal and Swaziland,

in Ann. Transv. Mus. 6: 119-135 (1918). Another 700 species were added.

'

5

Botlialia 9, 1 : 5-30.

The Genus Andropogon in Southern Africa

by

J. G. Anderson

Abstract

This study is limited to an investigation of the Southern African representatives of the genus

Andropogon, consisting of 14 species, two varieties and one record of doubtful identity. The history,

karyology, leaf anatomy and anatomy of the shoot and embryo are discussed briefly. A key based on

floral and vegetative characters is provided and a detailed description of the genus and of each species

and variety is given. The nomenclature and synonomy for each taxon is included as well as brief notes

on the distribution and habitat. Specimens deposited in the National Herbarium, Pretoria, and several

other South African Institutions are cited in full.

Introduction

No thorough revision of the genus Andropogon, as it occurs in South Africa and

including its nomenclature, has been undertaken since Stapf’s excellent work in

Volume 7 of the Flora Capensis (1898). An unpublished thesis by W. J. Louw (1943)

is available in which special attention is given to the leaf anatomy, but he dealt with

only nine species. A summary of the then known South African species, with short

descriptions and keys, was published by Chippindall in “ The Grasses and Pastures

of South Africa” (1955), but this work did not include nomenclatural information

nor the citation of specimens. Since the publication of this work, three new species

have been described by the present author, and one additional species has been

recorded for South Africa. The genus Hypogynium which included the South African

species H. festucifonnis has been referred back to Andropogon by Clayton in the Kew

Bull. 17, 3: 469 (1964). The present revision was, therefore, undertaken in order to

include the new information, as well as to investigate questions of synonomy and

nomenclature.

Most of the information regarding the anatomy of this genus was obtained from

Louw’s unpublished data but, as he did not deal with A. absyssinicus, A. lacunosus,

A. ravus, A. laxatus and A. festucifonnis, these species were investigated by the present

author.

The material in the National Herbarium, Pretoria, was studied as well as specimens

kindly sent on loan from the following herbaria: Bolus, Durban, Grahamstown,

Kimberley, Natal University and the South African Museum. Type material was

gratefully received on loan from Kew, Lisbon, Paris, Stockholm, Vienna and Zurich.

Special thanks are due to Dr. B. de Winter of the Botanical Research Institute, Pretoria,

for assistance and guidance.

All specimens cited are represented in the National Herbarium, Pretoria (PRE),

unless indicated otherwise.

History

Twelve species of Andropogon were included by Linnaeus in the first edition of

his Species Plantarum (1753) and, of these, A. distachyos L. has generally been accepted

as the type species. Most of the remaining eleven species, owing to their divergent

characteristics, have been assigned to other genera by subsequent authors. After

Linnaeus, the most important contributions to the knowledge of the genus for Southern

Africa came from Nees, Steudel, Hackel and Stapf.

6

In 1841 Nees dealt with 16 species of Andropogon and separated them into two

subgenera viz. Gymnandropogon and Cymbopogon. The subgenus Gymnandropogon

contained six species of which three, A. eucomus, A. amplectens and A. appendiculatus

were newly described from South Africa and are still regarded as true members of

Andropogon. The three remaining species are now placed in the genus Bothriochloa.

Under the subgenus Cymbopogon, Nees included ten species, eight of which were newly

described. These species have since been placed in the genera Monocymbium, Hyparr-

henia and Cymbopogon. Nees also described Heteropogon filifolius in this work, a

species subsequently transferred to Andropogon by Steudel.

Steudel, in 1855, recognised 459 species and divided the genus into five sections

and 29 subsections. Twenty species were cited from South Africa. Whilst several

of these were transferred to other genera by later workers, most of them were retained

as distinct taxa.

Hackel revised the tribe Andropogoneae in A. & C. de Candolle's “ Monographiae

Phanerogamarum ” in 1889 and divided the genus Andropogon into two series viz.

Isozygi and Heterozygi, consisting of thirteen subgenera with 193 species. Nine of

these were mentioned as coming from South Africa. Two new South African species

were described in this work, viz. A. schinzii and A. pilosissimus. During the present

study the holotype of A. pilosissimus was seen and it compared well with specimens

of Hyparrhenia aucta (Stapf) Stent deposited in the National Herbarium, Pretoria.

The necessary combination is made on p. 130 of this issue.

In 1898 the genus as it occurs in South Africa was fully treated by Stapf in the

Flora Capensis, Volume 7. Stapf followed Hackel in separating the genus into the

two series Isozygi and Heterozygi, dividing the South African representatives into

nine subgenera and 32 species. Stapf also states that this genus “ consists of several

heterogeneous elements ” and that “ several of the old genera of which it is made up,

will have to be restored, as for instance Chrysopogon, Cymbopogon and Heteropogon ”.

Only the seven species placed into the subgenus Arthrolophis are at present regarded

as true members of Andropogon.

In the Flora of Tropical Africa, 1919, Stapf treated the species of the genus occur-

ring in tropical Africa. In this work Stapf reinstated a number of genera formerly

placed in synonymy under Andropogon and gave a narrower circumscription to the

latter. The Andropogoneae were separated into sixteen subtribes with the genus

Andropogon placed in the subtribe Andropogonastrae. Stapf recognised 46 species

(and four sections). Of these, four species and one variety were known from South

Africa. Another four species and one variety subsequently proved also to occur in

South Africa.

The most recent work on the genus in this country is by Chippindall in “ The

Grasses and Pastures of South Africa”, (1955), in which ten species and one variety

are included. In this work brief descriptions of the species are supplied but synonymy

and citation of specimens are omitted.

Economic Importance

Economically this genus is not regarded as important and the species are usually

classified as “ sour grasses ”. They are usually grazed only in the young stages becoming

unpalatable when mature. Ecologically, however, certain species are important consti-

tuents in several of the major veld types of South Africa.

Karyology

In their study of A. distachyos, the type species of the genus, Gould (1956) and

Celarier (1956) found that in this species 2n = 36, indicating a basic chromosome

7

number of 9. For A. abyssinicus, Gould reported a count of 16. Subsequent studies

by various authors dealing with several other species of Andropogon, however, have

shown that 10 appears to be the commonest basic number in the genus.

The following table is a summary of the chromosome numbers so far recorded

for the South African species, with the authority and year of publication.

ANDROPOGON

Andropogon L., Sp. PI. 1045 (1753); Gen. PL ed. 5: 468 (1754); Nees. FI. Afr. Austr.

103 (1841), emend.; Steud., Syn. PI. Glum. 1: 363 (1855); Benth. & Flook. f., Gen.

PI. 3, 2: 1133 (1883); Hack, in DC., Monoar. Phan. 6: 359 (1889); Stapf in FI. Cap.

7: 334(1898); FI. Trop. Afr. 9: 208 (1919)f Phill., Gen. ed. 2: 84 (1951); Chippindall

in Gr. & Pastures S. Afr. 1: 495 (1955). Type species: A. distachyos L. (as “ dis-

tachyon ”, 1753).

Spikelets 2-nate, the one sessile, the other pedicelled, those of each pair differing

in sex and also more or less in shape and size or those of the lowest pair of the lowest

raceme male or imperfect or the pedicelled suppressed and the spikelets thus solitary.

Sessile spikelets 2-flowered, dorsally or laterally compressed, rarely pubescent, nearly

always with a bent and twisted awn; callus obtuse, subacute or acute, rarely broad

and obtusely rounded, shortly bearded. Glumes equal to subequal, subcoriaceous or

membranous; lower flat, concave or channelled on the back with sharply indexed

margins at least from the middle upwards, rarely with subcircular or elongated shallow

pits, 2-keeled; upper more or less boat-shaped, keeled upwards, 1-3-nerved, sometimes

aristulate or awned. Lemmas ciliate or ciliolate, rarely glabrous, lemma of lower

floret hyaline, 2-nerved; lemma of upper floret 2-fid or 2-dentate, usually awned from

the sinus, hyaline or firmer and sometimes substipiform below the insertion of the

awn; awn usually 2-8 times the length of the lemma. Paleas hyaline nerveless scales

or absent. Lodicules 2, minute, glabrous. Stamens 3, rarely reduced to staminodes.

Ovary glabrous, styles terminal; stigmas laterally exserted. Grains narrowly lanceolate

to oblong in outline, subterete to plano-convex; embryo about half the length of

the grain; starch grains simple or compound, granules mostly numerous. Pedicelled

spikelets often larger and more conspicuous than the sessile, sometimes small and

suppressed and represented by the pedicel only, more or less dorsally compressed,

rarely hairy. Glumes herbaceous, chaitaceous or membranous; lower usually flat

on the back, rarely grooved or with subcircular or elongated pits, muticous or very

rarely aristulate, usually many-nerved; upper membranous to subcoriaceous, 3-5,

rarely 7-nerved, ciliate or ciliolate on the margins, apex acute, acuminate, mucronate

or shortly awned. Florets: lower reduced to the lemma; upper male or sterile, rarely

bisexual; lemma, if present, hyaline, muticous or very rarely aristulate.

8

Perennials of various habit, rarely annuals, usually compactly caespitose, sometimes

with an oblique or short rhizome, very rarely fascicled or stoloniferous. Culms erect,

rarely geniculate, terete or sometimes compressed below, glabrous, smooth, rarely

hairy upwards, often channelled on the one side, usually simple, rarely branched either

from the base or from all the nodes resulting in a suffrutescent habit, or only from the

upper nodes. Leaves with the sheaths terete or the lower compressed and keeled,

glabrous or sparingly hairy, rarely densely pubescent, smooth, often tinged with purple;

ligules membranous with a ciliate or ciliolate rim, often scarious, truncate, rounded

to oblong or oblong-lanceolate; blades linear, expanded or longitudinally folded

(or revolute in dried specimens), more rarely filiform. Inflorescence of 1-many spike-

like racemes which are either in pairs, with the lower almost sessile, upper shortly

peduncled, or arranged subdigitately, or on a short central axis at the end of the culm

and, if present, its flowering branches ; raceme pairs or corymbs supported by a spathe

or spatheole and frequently collected into a false panicle; racemes glabrous or often

plumose with white or cream-coloured hairs; joints and pedicels linear, filiform,

cuneate or cuneate-clavate, often inflated and hollow or thickened upwards, tips more

or less cupular, auricled or dentate, rarely truncate.

Anatomy

Shoots rounded, compressed or strongly compressed in cross section, rarely cylin-

drical; leaf-blades rolled or folded in the bud; sheaths slightly to prominently keeled.

Leaf-blades flat, V-shaped or rarely terete in cross section, keels acute and prominent

or obtuse, rarely rounded, formed by an adaxial tissue of colourless parenchyma cells,

which are roughly triangular in transverse section. Silicified cells dumb-bell-shaped;

lateral walls smooth, exceptionally with a small central bulge. Suberized cells more or

less rectangular often with undulating lateral walls. Ripple-walled cells more or less

rectangular and arranged with their long axis parallel to the vascular bundles.

Stomata mainly located on the abaxial surface, usually in 1-2 longitudinal rows between

the nerves. Motor cells large and rectangular or hexagonal in shape, occupying £

the thickness of the leaf. Hairs variable and rare, of three kinds: (1) unicellular

usually antrorse sharp-pointed bulbous-based hairs; (2) unicellular tubercle-based

hairs, mostly on the abaxial surface and confined to motor cell and ripple-walled cell

areas; (3) bicellular hairs of which the basal and upper cells are approximately equal

in length; upper cell delicate and with a subobtuse apex. Bundle sheaths: outer sheath

thick-walled, parenchymatous, usually containing chloroplasts; inner sheath absent.

Chlorenchyma consisting of a usually well defined single layer arranged in the form

of an arc and surrounding each vascular bundle, often supported by irregulaily shaped

chlorophyll-bearing cells, mainly adaxially; the two adjacent arcs are either adjoining

or separated by a single row of larger rounded cells. Bulliform cells mainly consisting

of enlarged epidermal cells. Embryo about half the length of the grain; epiblast absent;

deeply cleft between the lower part of the scutellum and the coleorrhiza; the vascular

tissue with a distinct elongation between the point of divergence of the scutellum bundle

and the base of the coleoptile; margins of the primary leaf of the embryo (sheathed

by the coleoptile) overlapping and with several vascular bundles; the coleoptile

sheath with two lateral bundles and the scutellum with one median bundle.

Distribution

A large genus occurring mainly in the tropical and subtropical regions of both

hemispheres. Fourteen species, two varieties and one record of doubtful identity

(from the Natal Drakensberg) have been recorded from South Africa.

Most of the species are widely distributed throughout the eastern grassveld regions

of South Africa. Two species, A. eucomus and A. appendiculatus extend, in addition,

into mainly the coastal districts of the eastern and southern Cape, penetrating into

9

the south-western regions of the Cape. Three species and one variety, viz. A. eucomus,

A. amplectens , A. schinzii and A. schirensis var. angustifolius also occur in the central

regions while A. schinzii, A. eucomus, A. brazzae, A. huillensis, A. schirensis var.

angustifolius and A. gayanus var. squamulatus have been recorded from northern South

West Africa; all occur north of the tropic of Capricorn.

The name “Andropogon” is derived from the Greek words, aner, a man, and

pogon, a beard.

1 Basal parts usually distinctly flattened ; lower and sometimes the upper leaf-sheaths compressed

and keeled, usually strongly so; leaf-blades folded or sharply keeled, rarely expanded;

racemes 1-15:

2 Racemes plumose with silvery-white or cream-coloured hairs; joints and pedicels filiform

and slightly curved with hairs more than 5 mm long:

3 Sessile spikelets 2-3 mm long, the pedicelled supressed or represented by a minute rudiment ;

hairs on the joints and pedicels several times longer than the spikelets. ... LA. eucomus

3a Sessile spikelets 4—6 mm long, the pedicelled usually suppressed or rudimentary, more rarely

well developed; hairs on the joints and pedicels about as long as the sessile spikelets:

4 Racemes 5-12 (rarely 3 or 4), loose and usually flexuous; plant stout, 90-180 cm tall;

leaf-blades up to 30 cm long; lemma of the upper floret of the sessile spikelets

3-4 mm long, usually obscurely 3-nerved, rarely 1-nerved; awn 8-14 mm long

2. A. huillensis

4a Racemes 2-3, straight or slightly curved; plant up to 70 cm tall; leaf-blades up to

15 cm long; lemma of the upper floret of the sessile spikelets 2-3 mm long, usually

1-nerved; awn 15-20 mm long 3. A. laxatus

2a Racemes not plumose; joints and pedicels linear and straight, glabrous, or with hairs up to

5 mm long:

5 Racemes 1-3, 2^4 cm long; plant 20-100 cm tall; leaf-blades up to 30 cm long:

6 Racemes 1; sessile spikelets awnless 4. A. festuciformis

6a Racemes 2-3; sessile spikelets awned with awns up to 12 mm long 5. A. platybasis

5a Racemes 3-20 (if only 2, then racemes hairy), 4—16 cm long; plant 40-250 cm tall; leaf-

blades up to 50 cm long:

7 Sessile spikelets awnless; joints and pedicels glabrous or almost so; leaves expanded

or often revolute, margins scabrid or spinously ciliate 6. A. brazzae

7a Sessile spikelets awned, awns 10-15 mm long; joints and pedicels hairy with hairs up

to 5 mm long; leaves usually folded, rarely expanded, margins scabrid 7. A. appen-

diculatus

la Basal parts not distinctly flattened; lower leaf-sheaths rounded; leaf-blades filiform, convolute

or expanded; racemes 2, exceptionally 3, 4 or 5:

8 Glumes densely or sparingly hairy over the whole surface with hairs up to 4 mm long or,

if glabrous, then with 5-12 subcircular to elongated shallow pits (depressions) on either

side of a deep longitudinal median groove; joints and pedicels linear, straight:

9 Lower glume of the sessile spikelet dorsally flat or only slightly concave; glumes densely

or sparingly hairy with hairs about 4 mm long:

10 Annual; glumes densely hairy; racemes whitish-villous; joints and pedicels densely

hairy on the back and sides 8. A. abyssinicus

10a Perennial; glumes sparingly hairy; racemes appearing glabrous; joints and pedicels

hairy only on the sides 9. A. pilosellus

9a Lower glume of the sessile spikelet dorsally with a deep longitudinal median groove;

glumes glabrous and with 5-12 subcircular to elongated shallow pits on either side

of the groove 10. A. lacunosus

8a Glumes glabrous or ciliate only on the keels and margins, without depressions on either

side of the longitudinal median groove; joints and pedicels cuneate, cuneate-clavate

or cuneate-linear, usually inflated and hollow:

11 Lower glume of the sessile spikelet dorsally with a deep, narrow or wide, longitudinal

groove, groove often so deep that the wingless keels almost meet, especially in spikelets

borne on the upper part of the raceme:

10

12 Leaf-blades broad and rounded to cordate and more or less amplexicaul at the base;

callus acute or subacute 11. A. amplectens

12a Leaf-blades not or only slightly widened, often narrowed towards the base; callus

subacute, obtuse or obtusely rounded (if acute, then leaves filiform or narrowed

towards the base):

13 Upper glume of the sessile spikelet and both glumes of the pedicelled spikelet awned,

awn up to 10 mm long; culms branched, the branches either barren or bearing

inflorescences 12. A. schinzii

1 3a Glumes of both the sessile and pedicelled spikelets awnless, usually acute or mucronate ;

culms simple, rarely with one flowering branch:

14 Sessile spikelets 7-9 mm long; callus broad and obtusely rounded: plants often

with a branched system of knotty rhizomes; inflorescence usually glaucous-

grey or green 13. A. ravus

14a Sessile spikelets up to 7 mm long; callus obtuse, subacute or acute; plants densely

tufted; inflorescence and often the older leaves drying brown or reddish-brown:

15 Sessile and pedicelled spikelets very unequal in length; awn of sessile spikelets

3-5 cm long, stout; callus acute or subacute; leaves usually filiform or

when expanded then up to 3 mm wide, leaf-sheath usually breaking up

into fibres when old 14. A. filifolius

15a Sessile and pedicelled spikelets more or less equal in length; awn of sessile

spikelets up to 3 cm long, slender; callus obtuse to subacute; leaf-blades

usually expanded and 3-7 mm wide; lower leaf-sheaths not breaking up

into fibres 15. A. schirensis var. angustifolius

11a Lower glume of the sessile spikelet dorsally flattened or only very shallowly grooved:

16 Keels of the glumes of the sessile spikelets wingless; culms branched, robust, up to

3-5 m tall, usually glaucous, plants tufted 16. A. gayamts var. squamidatus

16a Keels of the glumes of the sessile spikelets distinctly winged; culms simple, up to

1-2 m tall; plant with a short oblique rhizome 17. A. distachyos

1. Andropogon eucomus Nees, FI. Afr. Austr. 1: 104 (1841); Steud., Syn. PI.

Glum. 1: 390 (1855); Hack, in DC., Monogr. Phan. 6: 421 (1889); Stapf in FI.

Cap. 7: 338 (1898); FI. Trop. Afr. 9: 230 (1919); Chippindall in Gr. & Pastures

S. Afr. 1; 501 (1955). Type: Clanwilliam, Olifantsriver, Eck/on & Zeyher s.n. (S, lecto.!).

Eriopodium krausii Hochst. ex Krauss in Flora 115 (1846). Type: Natal, between

Umlazi River and Durban Bay, Krauss 163 (K!; PRE, fragment!).

Perennial, densely tufted, occasionally with a short creeping rhizome. Culms up

to 90 cm long, erect or slightly geniculate, glabrous, 4-6 noded, compressed below,

simple to about the middle and with 2-6 or more flowering branches from the upper

nodes. Leaves up to 30 cm long and 4 mm broad, folded or rarely expanded, linear,

acute, glabrous or pubscent especially near the ligule, smooth or slightly scaberulous

on the margin, midrib slender, prominent below, lateral nerves 4-5 on each side, very

fine; ligules 0-5 mm long, truncate, membranous, ciliolate; sheaths glabrous or

beaided near the mouth, rarely lower down, firm, basal compressed and keeled, the

remainder shorter than the internodes, terete, distant, the uppermost subtumid.

Inflorescence of 2-5 subdigitate racemes, conspicuously plumose from the silvery-

white hairs surrounding the spikelets and on the rachis and pedicels; spathes linear-

lanceolate, acuminate or acute, usually without a blade, glabrous, pale-green or tinged

with purple or red, at length tightly inrolled. Racemes 3-5 cm long, plumose, often

nodding; joints finely filiform, 2 mm long, silky with soft 1-0-1 -2 cm long hairs;

pedicels similar, 4 mm long, quite barren or occasionally with the minute rudiment

of a spikelet at the apex. Spikelets all sessile, hermaphrodite, often purplish; callus

minute, truncate, long bearded. Glumes equal; lower 2-3 mm long, subcoriaceous,

narrowly lanceolate, concave dorsally, keeled, with 2-3 obscure nerves in each keel;

keels scabrid in the upper half, rarely long bearded near the apex, obtuse and glabrous

in the lower half; apex minutely bifid; upper boat-shaped, 2-3 mm long, membranous,

11

1- nerved, sparingly pilose on the margins in the upper half, dorsally scabrid on the

nerve; apex acuminate or shortly mucronate. Lower floret reduced to a lemma

2- 2-5 mm long, narrowly lanceolate, membranous, hyaline, nerveless, sparingly ciliate.

Upper floret hermaphrodite, lemma 1-5-2 mm long, lanceolate-oblong, 1-nerved,

ciliolate, bifid to about one third, lobes finely subulate, awned from between the lobes;

awn a fine bristle, 12-20 mm long, slender, slightly twisted near the base; palea very

minute, hyaline, nerveless; anthers 0-75 mm long; grain 1 mm long or longer, linear-

lanceolate in outline, subcylindric. Pedicelled spikelets absent or reduced to a minute

rudiment.

This species is widely distributed in southern Africa but has not been recorded

from the drier areas of the central Cape, south-western Orange Free State and southern

South West Africa. It also occurs in Mozambique, Rhodesia, Angola, the Congo

and Uganda. It usually grows in damp places, often in vleis and occasionally in open

grassland or on rocky outcrops.

Cape. — Ceres: Marloth 12643. Clanwilliam: Olifants River at Villa Brakfontein, Zeyher 47. East

London: Convict Station, Galpin 5580. Grahamstown: Hutton 249. Hay: Griquatown, Burchett

2102 (GRA; PRE, fragment); Witsand, Acocks 4474 (KMG); Leistner 1171. Humansdorp: Karee-

douw, Fowcade 2533. Komga: near Kei Mouth, Flanagan 2362. Paarl: Klein Drakenstein Mts.

on Berg River, Drege s.n. (S; PRE, fragment); Dal Josaphat, Tyson 6050 (SAM). Peninsula: Kloof-

neck, Marloth 3717; Steenberg near Lakeside, Bolus 14928 (BOL). Port Elizabeth: Schoemakers

Kop, Paterson 548 (GRA). Riversdale: Kortefontein, N. slopes of Langeberge, Acocks & Hafstrom

2039. Robertson: Omklaar, Stokoe 7196 (BOL). Somerset West: Steenbras River Mouth, Acocks

3936 (KMG). Taung: Thoming, Brueckner 1235; Rodin 3653. Tulbagh: Nieuwkloof, MacOwan

1691 (GRA, SAM). Vryburg: Armoedsvlakte, Mogg 8123. Worcester: Bainskloof Pass, Bolus

11743 (BOL).

Basutoland. — Tejatejaneng: Dieterlen 816.

Orange Free State. — Heilbron: Makouvlei, Brandmuller 44; Viljoensdrift, Rogers 13101.

Natal. — Bergville: Cathedral Peak Forest Research Station, Killick 1435; 1448. Camperdown:

Nagle Dam, Wells 1171 (NU). Dundee: Vants Drift, Codd 185. Durban: Bay End, Strey 4543;

Medley Wood 1654 (NH); 6004; Rehmann 8550 (GRA). Eshowe: Kotze 417. Estcourt: Mountain

hostel, West 253; Champagne Castle, Bayer 1246a. Newcastle: 20 miles S.E., Godfrey S.H. 1887.

New Hanover: Blinkwater, Sim 19361. Nkandla: Melmoth, Tinley 6; Qudeni, Leisegang 57 (NU).

Nongoma: Gerstner 3875 (NH). Pietermaritzburg: Table Mtn., Killick 282 (NU). Signal Hill,

Bews 37 (NU). Port Shepstone: Mogg 13201. Umlazi: Umlaas River, Drege s.n. (S, SAM); Isipingo

Beach, Ward 1275 (NU). Utrecht: Donkerhoek, Devenish 1124.

Swaziland. — Mbabane: Ukutula, Compton 25514; Forbes Reef, Compton 22391.

Transvaal. — Belfast: Schoemanskloof, Smuts 336. Brits: Jacksonstuin, Van Vuuren 400. Ermelo:

Athole Pasture Research Station, Norval 137. Johannesburg: Hutton 249 (GRA). Krugersdorp:

Jack Scott Private Nature Reserve, Wells 2367. Letaba: Duiwelskloof, Westfalia Estate, Scheepers

838. Lichtenburg: Malmanie Oog, Burn Davy 80. Middelburg: Pan Station, Burtt Davy 137. Nel-

spruit: Kaapsehoop, Devils Kantoor, Pole Evans 1008. Petersburg: farm Vaalwater, Skead 31;

Houtbosch, Rehmann 5673 (BOL). Pilgrims Rest: Mariepskop, Van der Schijff 4473. Potchefstroom :

Welverdiend, Louw 870. Potgietersrus : Crecy, Barenbrug M. Pretoria: Rayton vlei, Schweickerdt

1583; Waterkloof, Verdoorn 832; Wonderboom, Louw 864. Vereeniging: Brandmuller 44. Waterberg:

Nooitgedacht 699, Acocks & Naude 42.

Bechuanaland. — Lobatsi: Kanye, Bathosu Dam, Miller B/873.

South West Africa. — Caprivi: Katima Mulilo, Killick & Leistner 3082. Grootfontein : Omuramba,

Hohensee, Boss s.n.; Fontein Omuramba, De Winter & Marais 5041. Okahandja: Wiss & Kinges

994. Otjiwarongo: Waterberg, Liebenberg 4785; Waterberg plateau, Boss s.n.

The Ecklon & Zeyher specimen in Stockholm proved to be the most authentic

material seen. This specimen is written up in Nees’s own handwriting and was for this

reason chosen as the lectotype.

This species is characterized by the absence of pedicelled spikelets. The pedicels,

although present, are either barren or bear only the minute rudiments of a spikelet.

The hairs on the joints and pedicels are dense and several times longer than the spikelets.

These hairs, together with the long slender awns of the lemma of the upper floret, give

the inflorescence an attractive white, plumose appearance.

12

2. A. huillensis Rendle, Cat. Afr. PI. Welw. 2: 146 (1899); Stapf in FI. Trop.

Afr. 9: 231 (1919); Chippindall in Gr. & Pastures S. Afr. 1: 500 (1955). Type:

Angola, Huila, Humpata, Welwitsch 2670 (BM, holo.; PRE, fragment!, photo.!).

Perennial, tufted from a short oblique rhizome. Culms 90-180 cm long, erect,

stout, terete, 5 or more noded, glabrous, very smooth, shiny, simple up to the third

or fourth node. Leaves up to 60 cm long and 4 mm wide, linear, long tapering to a

fine apex, flat or folded, firm and hard, the upper almost reduced to the midrib, smooth

and glabrous or sparingly pilose towards the ligule, scaberulous on the margin; ligules

2 mm long, firm, scarious, ciliolate; sheaths firm, tight, glabrous or bearded near

the ligule, the lower up to 25 cm long, compressed and keeled or sometimes terete,

often shiny or tinged with purple. Inflorescence of 5-12 racemes arranged subdigi-

tately, plumose from the long white hairs on the jonts and pedicels; spathes glabrous,

the lowest resembling the preceding leaves; spatheoles narrow, linear-lanceolate to

linear, tapering to an acute apex, up to 8 cm long, reddish, at length tightly inrolled.

Racemes 4-10 cm long, slender, loose, flexuous and often nodding, plumose, sessile

or with peduncles 1-3 cm long; peduncles glabrous; joints 3-6 mm long, filiform,

often curved, bearded with fine, approximately 7 mm long hairs, apex shallowly cupular;

pedicels similar but approximately 5 mm long. Sessile spikelets 4-6 mm long, linear-

lanceolate, more or less tinged with purple; callus minute, rounded, bearded. Glumes

firmly membranous; lower 4-6 mm long, dorsally deeply or shallowly concave, keeled,

obscurely 3-nerved in each keel; keels scaberulous in the upper half, apex minutely

bifid; upper 5 mm long, narrowly boat-shaped, 1 -nerved, pilose on the margins in

the upper half; apex acuminate or mucronulate. Lower floret reduced to a linear-

oblong lemma, 3-5 mm long, membranous, hyaline, nerveless or obscurely 2-nerved,

pilose on the margins in the upper half, apex acute. Upper floret hermaphrodite;

lemma 3-4 mm long, narrowly linear-lanceolate, membranous, hyaline, obscurely

3-nerved, rarely 1 -nerved, pilose on the margins in the upper half, 2-lobed, lobes acute,

awned from between the lobes, awn a fine bristle 8-14 mm long, usually bent and

twisted below the middle, scaberulous; palea 2-2-5 mm long, oblong, hyaline, nerveless,

bearded near the obtuse apex; anthers 2 mm long; grain 4 mm long, linear-lanceolate.

Pedicelled spikelets variously reduced, sometimes to a glume or entirely suppressed,

occasionally well developed and male and then as long as or slightly longer than the

sessile.

A. huillensis is common in the Transvaal and is also found in Zululand, Swaziland,

South West Africa, Angola, Rhodesia and Mozambique. It usually grows in damp

places, along streambanks and in vleis, very often in sandy soils.

Transvaal. — Barberton: White River, Rogers 22965. Brits: Jacksonstuin, Van Vauren 392.

Johannesburg: Hutton 254. Krugersdorp: Lake, Louw 922. Lydenburg: between Lydenburg and

Dullstroom, Pole Evans 3811. Middelburg: Burtt Davy 13264; Hutchinson 2709. Nelspruit:

Pretoriuskop, Van der Schijff 2648. Pilgrims Rest: Graskop, Driekop Gorge, De Winter & Codd 210.

Potchefstroom: Elandsfontein, Louw 1522. Potgietersrus: Pyramid Estate, Gatpin 8884. Pretoria:

Rayton vlei, Schweickerdt 1593; Premier Mine, Repton 4058. Rustenburg: Rainhill, Story 991;

Codd 1074. Soutpansberg: Phiphiti Waterfalls, Obermeyer s.n. Warmbad: Bell s.n. Waterberg:

Twenty-four Rivers, 13 miles E. of Vaalwater, Codd 994; Rogers 25104.

Natal. — Ubombo: Bazwanu, Strey 5135.

Swaziland. — Hlatikulu: Mhlambanyati Pan, Michelmore 50. Mbabane: Black Mbuluzi Valley,

Compton 31269.

South West Africa. — Caprivi: Andara, Okavango River, Volk 2154.

A. huillensis can be confused with A. eucomus and A. laxatus but is generally a

taller, more robust plant with larger spikelets and with the hairs on the joints, and

pedicels shorter than in A. eucomus. In A. laxatus there are only 2-3 racemes in the

inflorescence whereas there are 5-12 in A. huillensis. In A. huillensis the pedicelled

spikelets are usually suppressed, these being represented by much reduced glumes.

These spikelets are occasionally well developed and male.

13

Welwitsch 2644, collected in Angola, was described by Rendle as A. huillensis

var. minor in Cat. Afr. PI. Welw. 2: 147 (1899). In the Flora of Tropical Africa,

Stapf states that the plant has smaller inflorescences and smaller spikelets and adds

that it is probably only a weaker state of A. huillensis. I have seen an isotype as well

as a fragment and photograph of the holotype and in my opinion this variety should

be included in the synonomy of A. laxatus.

3. A. laxatus Stapf in FI. Trop. Afr. 9: 237 (1919); Chippindall in Gr. & Pastures

S. Afr. 1: 501 (1955). Syntypes: Angola, Welwitsch 2693; Rhodesia, Inyanga , Mundy

s.n. ; Salisbury, Mundy s.n. (K!).

A. ternatus Nees var. africanus Rendle, Cat. Afr. PI. Welw. 2: 147 (1899). Type:

Angola, Huila, dry wooded pastures near Humpata, Welwitsch 2643 (LISU!; PRE,

photo.!). A. huillensis Rendle var. minor Rendle in Cat. Afr. PI. Welw. 2: 147 (1899).

Type: Angola, Huila, rather damp rich pastures on the river Monino, Welwitsch

2644 (LISU!; PRE, photo.!, fragment!).

Perennial, densely tufted. Culms up to 70 cm long; erect, terete, somewhat

compressed below, glabrous, smooth, shiny, sometimes pruinose below the lower

nodes, branched in the upper half, 1-3 noded. Leaves up to 15 cm long and 4 mm

wide, linear, expanded or folded, subacute, firm, flexuous or straight, glabrous or

sparingly hairy near the ligule, margins scaberulous; ligules 1 mm long, scarious,

rounded or truncate, ciliolate; sheaths firm, glabrous, smooth, the lowest up to 6 cm

long, compressed, keeled, the upper looser, much longer and more or less terete.

Inflorescence of 2, rarely 3, racemes on a short common axis, silkily plumose from the

hairs on the joints and pedicels; spathes resembling the preceding leaves and with

short blades; spatheoles very narrow, acute, 6-8 cm long, at length tightly inrolled,

tinged with purple. Racemes 4-6 cm long, slightly flexuous, plumose; peduncles short

or the upper raceme sessile; joints 4-6 mm long, filiform, flattened, bearded with

spreading hairs 5-8 mm long, apex cupular; pedicels similar, 4-5 mm long, apex

obtuse. Sessile spikelets 5-6 mm long, linear-lanceolate; callus small, obtuse, bearded.

Glumes more or less equal, membranous; lower 5-6 mm long, dorsally concave, keeled,

1-2-nerved in each keel; keels scaberulous in the upper half, apex shortly bifid; upper

5-6 mm long, narrowly boat-shaped, 1-3-nerved, pilose on the margins in the upper

half, dorsally scabrid on the nerves, apex mucronate, mucro approximately 1 mm

long. Lower floret reduced to a lemma 4-5 mm long, narrowly oblong, hyaline,

nerveless, pilose on the margins in the upper half, apex acute. Upper floret herma-

phrodite; lemma 2-3 mm long, narrowly oblong, hyaline, usually 1 -nerved, often

pilose on the margins, apex 2-lobed, awned from between the lobes; awn 1- 5-2-0

cm long, slender, kneed and twisted below, scaberulous ; palea 2 • 5-3 mm long, oblong,

hyaline, nerveless, ciliate upwards. Pedicelled spikelets usually reduced, occasionally

well developed and male and then of the same shape and size as the sessile. Glumes

unequal; lower 5-nerved, acuminate; upper 3-nerved, acuminate; lemmas of both

florets, if present, narrowly oblong, acute, 1 -nerved, ciliate.

A. laxatus is a rare species occurring in Angola, Rhodesia, Zambia and the Congo

Republic. It has been recorded only once from South Africa, namely at Graskop

in the Pilgrims Rest district of the Transvaal. It usually grows in wet places, near

streams or in vleis.

Transvaal. — Pilgrim’s Rest: Graskop, Driekop Gorge, De Winter & Codd 218.

Angola. — Huila: Humpata, Texeira 3411. Moxico: Miltie-Redhead 4217.

Zambia. — Mwinilunga: Sinkabolo Dambo, Milne-Redhead 4380. Barotseland: 10 miles N. of

Senanga, Codd 7281.

Congo. — Leopoldville: Kwango, Devred 1495.

14

This species is very closely related to A. huillensis but is a shorter, less robust

plant with 2, rarely 3, racemes to the inflorescence. The lemma of the upper floret

is shorter, 2-3 mm long, but with a much longer awn, 15-20 mm long, than is found

in A. huillensis.

Of the syntypes only Mundy s.n. from Salisbury was seen and consequently no

lectotype was selected.

4. A. festuciformis Rendle in Cat. Afr. PI. Welw. 2: 145 (1899); Clayton in Kew

Bull. 17: 470 (1964). Type: Angola, Huila, Humpata, Welwitsch 7505 (BM, holo. ;

PRE, photo. !).

A. schlechteri Hack, in Bull. Herb. Boiss. ser. 2, 6: 703 (1906). Type: Natal,

Clairmont, Schlechter 3143 (Z, holo.; BOL!).

Hypogynium schlechteri (Hack.) Pilger in Pflanzenfam. ed. 2, 14e: 156 (1940);

Chippindall in Gr. & Pastures S. Afr. 1 : 516 (1955).

Perennial, densely tufted, subflabellate. Culms up to 100 cm long, erect, simple

below but branched upwards, compressed, smooth and often glossy, usually purplish,

2-3-noded. Leaves up to 30 cm long and 3 mm wide, rigid, folded, linear, passing

gradually into the sheath, pubescent on the upper surface near the base, apex acute;

ligules 1 mm long, membranous, acute; sheaths striate, persistent, firm, often tinged

with purple, lower compressed and keeled, glossy. Inflorescence of solitary, peduncled

racemes, crowded to form a narrow panicle 5-15 cm long or scattered along the culm;

spathes scarious, narrow, the lower 2-3 cm long, decreasing in size upwards, acute,

usually tinged with purple. Racemes 2-4 cm long, straight, glabrous; joints and

pedicels unequal, 4-5 mm long, linear, straight, often with a shallow median groove,

scaberulous, apex truncate. Sessile spikelets female, 4-4 • 5 mm long, linear-lanceolate ;

callus short, obtuse, shortly bearded. Glumes subequal; lower 3 5-4 mm long, linear-

lanceolate, bicarinate, subcoriaceous, scaberulous on the keels in the upper half, dorsally

shallowly grooved, 1-nerved in each keel, apex minutely bifid; upper 3-3-5 mm long,

boat-shaped, sharply keeled, subcoriaceous, 1-nerved, scaberulous on the upper half

of the keel, apex acute. Lower floret reduced to the lemma, 2-5-3 mm long, narrowly

oblong, membranous, nerveless, apex obtuse or truncate. Upper floret with the lemma

lanceolate, 2-2-5 mm long, membranous, 1-nerved, apex shortly bilobed or subacute,

awnless; palea absent; anthers reduced to staminodes. Pedicelled spikelets male,

5 • 6-6 mm long, lanceolate. Glumes subequal ; lower 5 • 6-6 mm long, narrowly ovate,

subcoriaceous, 5-7-nerved, glabrous, apex acuminate; upper 4-7-5 mm long, subcori-

aceous, narrowly ovate, 5-7-nerved, apex acuminate. Lower floret with the lemma

3 • 5-4 mm long, lanceolate, membranous, 3-nerved, apex obtuse. Upper floret with

the lemma 3-2-4 mm long, narrowly lanceolate, membranous, 1-nerved, apex acute;

palea absent; anthers 2-5 mm long.

Andropogon festuciformis has been recorded from Natal, Angola, Zambia, the

Congo Republic and the Guinea Republic. It usually grows in damp places, often

in vleis.

Natal. — Durban: Clairmont, Medley Wood 8543. Hlabisa: W. of St. Lucia Estuary, Feely &

Ward9\ Umfolozi flats near Mtubatuba, Liebenberg 591 1. Pietermaritzburg: Table Mtn., Killick 284.

This species differs from all the other South African species in having solitary

racemes. Until recently it was placed in the genus Hypogynium Nees and was separated

from Andropogon mainly on account of the awnless lemmas of the sessile spikelets

and the solitary racemes. W. D. Clayton in the Kew Bulletin (1964) points out that

a few species of Andropogon in the section Piestum (e.g. A. textilis) also have solitary

racemes and that these characters are not sufficient to justify a separate genus. In

A. brazzae (see p. 15) the sessile spikelets are awnless as in A. festuciformis.

15

5. A. platybasis J. G. Anderson in Kirkia 1 : 102 (1961). Type: Transvaal, Belfast,

5^ miles from Dullstroom on road to Lydenburg, De Winter & Codd 180 (PRE, holo.!).

Perennial, tufted with a short rhizome. Culms up to 75 cm long, erect, simple,

2-3-noded, glabrous, striate, compressed; nodes glabrous, internodes glabrous and

smooth except for the upper which are pilose for approximately 3 cm below the

inflorescence. Leaves up to 15 cm long, 3-4 mm wide, folded or expanded in the upper

half, linear with the apex acute, glabrous, occasionally with a few long hairs near the

ligules, margins scaberulous; sheaths strongly compressed and keeled, glabrous,

striate, pallid or suffused with purple, basal persistent; ligules fimbriate, membranous,

1 mm long. Inflorescence of 3, occasionally 2, spike-like racemes arranged digitately

or subdigitately. Racemes 2-3 cm long, straight, greyish-violet; joints and pedicels

2-3 mm long, linear, pilose on the margins. Sessile spikelets female with the anthers

leduced to staminodes, 5-6 mm long, lanceolate. Glumes subequal, membranous,

glabrous; lower 5-6 mm long, narrowly lanceolate, minutely 2-dentate, concave

between the keels with 2 nerves in each keel, scaberulous from the middle upwards;

upper 3-6-4 mm long, narrowly boat-shaped, 3-nerved with the faint lateral nerves

often anastomosing with the distinct midnerve, midnerve excurrent into a 1 mm long

awn. Lower floret reduced to the lemma, 4 mm long, margins inflexed. hyaline, nerveless,

margins ciliate in the upper half. Upper floret with the lemma 2 mm long, lobed to

about the middle, awned from between the lobes; awn 9-12 mm long, lobes ciliate;

palea 0-75 mm long, truncate hyaline, nerveless. Pedicelled spikelets male, 6-7 mm

long, lanceolate. Glumes membranous; lower with a distinct midnerve and 3-4 nerves

on either side, keeled with the keels scaberulous in the upper half, entire at the apex

or with 2 membranous lobes on each side of the slightly excurrent nerve; upper narrowly

boat-shaped, 7 mm long, 3-nerved, membranous, apex mucronulate. Lower floret

reduced to the lemma, 5-5-6 mm long, lanceolate, 3-nerved with a distinct midnerve

and a faint lateral nerve on either side, apex obtuse. Upper floret male; lemma and

palea absent; anthers 3 mm long.

A. platybasis is a rare species recoided from the Belfast, Lydenburg and Wakker-

stroom districts of the eastern Transvaal and from Mbabane in Swaziland. It is a

water-loving grass growing in marshy soils, on streambanks and at high altitudes.

Transvaal. — Belfast: 6 miles N.E. of Dullstroom, Codd & De Winter 3225. Lydenburg: 13-6

miles from Lydenburg on road to Sabie, De Winter & Codd 199. Wakkerstroom: Oshoek, Devenish

910.

Swaziland. — Mbabane, Compton 30984.

A. platybasis is characterised by the upper floret of the sessile spikelet being female

with its anthers reduced to very tiny staminodes. It is also readily distinguished from

other South African species in being generally a smaller plant with 2-3 short, greyish-

violet, racemes.

6. A. brazzae Franch. in Bull. Soc. Hist. Nat. Autun 8; 326 (1895); Stapf in FI.

Trop. Afr. 9: 233 (1919); Robyns, FI. Agrost. Congo Beige 1: 126 (1929). Type:

Congo Republic, between Mokeno and Mongo, Brazza 234 (P, holo., iso.!; PRE,

photo.!).

A. appendiculatus Nees var. genuinus sensu Dur. & Schinz, Etudes FI. Congo 314

(1896), non Hack. (1889), fide Stapf.

Perennial, densely tufted with a short rhizome. Culms up to 250 cm long, erect,

stout, slightly compressed and simple below with a few long, erect, flowering branches

higher up, often channelled, glabrous and smooth, 3-many noded. Leaves up to 50 cm

long and 5 mm wide, linear, long attenuate to a fine apex, expanded or revolute in

dried specimens, puberulous on both surfaces, margins scabrid to spinously ciliate;

ligules membranous, ciliolate, 2 mm long, rounded; sheaths up to 20 cm long and

16

1 cm broad, lowest and intermediate compressed and strongly keeled, firm, striate,

glabrous, often tinged with purple. Inflorescence of 3-15 racemes, simple or divided;

spathes like the preceding leaves but with reduced blades. Racemes 3-10 cm long,

slender, loose and often flexuous, glabrous, pale or purple-brown, joints 4-7 mm long,

linear, slender, slightly thickened upwards, glabrous, scaberulous or shortly ciliate on

the margins, transparent towards the middle line, tips minutely cupular and denticulate;

pedicels similar, 3-5-5 mm long. Sessile spikelets hermaphrodite, 4-6 mm long,

linear-lanceolate, loosely wedged in between joint and pedicel; callus short, obtuse,

glabrous. Glumes subequal; lower 4-6 mm long, subcoriaceous, keeled, shallowly

concave and hyaline between the keels; keels smooth in the lower half, scabrid upwards,

2-nerved in each keel, apex bifid or acuminate; upper 4-5-5 mm long, membranous,

boat-shaped, acuminate, 3-nerved with the midnerve distinct, the lateral nerves shorter

and finer, glabrous or dorsally scabrid in the upper part of the midnerve. Lower floret

reduced to a lemma 3-5-5 mm long, membranous, hyaline, lanceolate, acute or

acuminate, indistinctly 3-nerved, winged in the upper half, glabrous or ciliolate on the

margins. Upper floret with the lemma 4-5-5 mm long, membranous, hyaline, 1 -nerved

or indistinctly 3-nerved, glabrous, acute, mucronate or rarely with rudiments of an

awn up to 1-5 mm long; palea 1-1-5 mm long, ovate-oblong, hyaline, nerveless;

anthers 2 mm long. Pedicelled spikelets male, 4-6 mm long. Glumes subequal; lower

4-6 mm long, linear-lanceolate, subcoriaceous, keeled, 5-nerved with a distinct mid-

nerve and 2 nerves close together in each keel, keels scabrid in the upper half, apex

acute or acuminate; upper 4-5 mm long, lanceolate, often winged in the upper half,

membranous, glabrous or ciliolate on the margins, 3-nerved, acuminate or often

mucronate. Lower floret reduced to a lemma 3-4 mm long, membranous, hyaline,

winged in the upper half, glabrous or ciliolate on the margins, indistinctly 3-nerved,

apex acute. Upper floret with the lemma 3-3 • 5 mm long, lanceolate, glabrous, hyaline,

obscurely 1 -3-nerved, mucronulate; palea 1 mm long, ovate-oblong, hyaline, nerveless;

anthers 2 mm long.

A. brazzae has been recorded from the Okavango Native Territory in South West

Africa, Angola and the Congo Republic. It is a rare species found on streambanks

or in marshy soils.

South West Africa. — Okavango: 16-8 miles W. of Nyangana Mission, De Winter & Marais 4775.

Angola. — Moxico: Lusavo Ferry, Milne-Redhead 4125.

Congo Republic. — Equateur: Conteaux 503; 564; Coquihatville, Germain 4717; Bondo Village,

Germain 1934. Leopoldville: Mongobele, Flamigny 5046. Orientate: Ango, Germain 4311.

This species is allied to A. appendiculatus but differs from that species in that the

spikelets are either awnless or do not produce fully developed awns. Investigation

of the material deposited in the National Herbarium, Pretoria, revealed that in the

sessile spikelets the apex of the lemma of the upper floret is usually acute or mucronate.

In some spikelets the midnerve of this lemma is thickened at the apex and frequently

produced into an awn up to 1-5 mm long. Other interesting features of this species

are that the leaf-blades are frequently revolute in dried specimens and that the leaf

margins are often very distinctly spinously ciliate.

7. A. appendiculatus Nees, FI. Afr. Austr. 105 (1841); Steud., Syn. PI. Glum.

1 : 379 (1855); Hack, in DC., Monogr. Phan. 6: 436 (1899); Stapf in FI. Cap. 7: 339

(1898); Chippindall in Gr. & Pastures S. Afr. 1: 500 (1955). Type: Cape, Klein

Buffelvallei, near Gaatjie, Drege s.n. (S, lecto.!, SAM!).

A. ischaemum Thunb., Prodr. 20 (1794), non Linn. Type: Cape, Thunberg s.n.

(S, iso.!). A. appendiculatus Nees var. genuinus Hack, in DC. Monogr. Phan. 6: 437

(1889). — var. ischaemum Nees, FI. Afr. Austr. 105 (1841). Syntypes: Uitenhage,

Van Stadensriver Mtns., Drege 696!; Zwartkei River, Ecklon s.n.!; Klein Buffelvallei,

near Gaatjie, Drege s.n.! — forma purpurea Nees, l.c., 106 (1841). Type: Winterberg,

17

Ecklon s.n. — var. serrulatus Nees, l.c., 105 (1841), excl. syn. ; Hack, in DC., Monogr.

Phan. 6: 437 (1889). Syntypes: Waterfall near Tulbagh, Ecklon s.n.!; Dutoitskloof,

Drege s.n.!.

Perennial, densely tufted with a creeping, sometimes branched, rhizome. Culms

up to 120 cm long, glabrous, compressed or terete, 4-6 noded, simple below, remotely

branched from the upper 2-3 nodes with 1-3 flowering branches; branches solitary,

long, glabrous. Leaves mostly basal; blades up to 30 cm long and 5 mm broad,

usually folded or sometimes expanded, upper very short or obselete, rigid, glabrous

or bearded near the ligule, acutely keeled, margins scabrid; ligules 1-5-2 mm long,

truncate, membranous, ciliolate; sheaths glabrous or bearded at the mouth, lower very

firm, strongly compressed, keeled, persistent, upper shorter than the internodes, tight,

often tinged with purple. Inflorescence of 2-20 racemes; peduncles generally long

exserted, glabrous. Racemes 4—16 cm long, fascicled, unequal, often slender and loose,

usually strongly flushed or wholly dark purple; joints linear, 4-6 mm long, hairy on the

convex back, hairs up to 5 mm long, white or yellowish, tips subcupular with irregularly

toothed margins; pedicels similar but more slender, produced into a subulate appendage

facing the upper glume. Sessile spikelets hermaphrodite, 6-7 mm long, lanceolate;

callus short, obtuse, scantily bearded. Glumes subequal, subcoriaceous to chartaceous;

lower 6-7 mm long, lanceolate, dorsally concave, keels acute, scabrid or rigidly ciliate

in the upper half, obscurely 3-nerved in each keel, apex bifid, rarely deeply bifid; upper

5-5-6 mm long, boat-shaped, 1-nerved, glabrous or scabrid on the keel. Lower floret

reduced to the lemma, 5-5-6 mm long, oblong-lanceolate, membranous, obscurely

2-nerved or nerveless, pilose on the margins in the upper half, apex acute. Upper

floret with the lemma 4—4-5 mm long, membranous, 1-3-nerved, pilose on the margin

in the upper half, 2-lobed, lobes acuminate, awned from between the lobes; awn

10-15 mm long, slender, geniculate below the middle; palea 1-2-3 mm long, ciliate,

hyaline, nerveless; anthers 4 mm long. Pedicelled spikelets male. Glumes subequal,

subcoriaceous; lower 5-4-6 mm long, oblong-lanceolate, flat or slightly rounded

on the back, scabrid or ciliate on the keels in the upper half, keels acute, midnerve

prominent and with 3 nerves in each keel, nerves often anastomosing near the apex,

minutely awned from a bifid or acute apex; awn 1-7-2 mm long; upper 4-5-5 mm

long, oblong-lanceolate, pilose on the margins in the upper half, 1-3-nerved, apex

acute or shortly mucronate. Lower floret with the lemma 4-5-5 mm long, lanceolate,

membranous, 3-nerved or nerveless, reversedly pilose on the margins in the upper half,

apex acute. Upper floret with the apex of the lemma minutely bifid, otherwise similar

to that of the lower floret; palea 3-7-4 mm long, elliptic, membranous, nerveless,

ciliate on the margins in the upper half; anthers 3-4 mm long.

This is a common and widespread species in South Africa, in which it is endemic,

its distribution ranging from the south-western, eastern and north-eastern Cape to

Natal, Swaziland, Transvaal, Basutoland and the Orange Free State. It is absent

from the drier areas of the central Cape and South West Africa. A. appendiculatus

occurs from sea level up to altitudes of 10,000 ft. above sea level in the Natal Drakens-

berg. It is usually found growing in damp situations, frequently in swampy soils and

very often in the shade.

Cape. — Albany: Grahamstown Nature Reserve, Britten 5961. Aliwal North: Elands Hoek, Bolus

10505. Barkly East: Ben McDhui, Galpin 6896. Caledon: Rivierzondereind, Schlechter 9895 (BOL,

GRA). Ceres: Koude Bokkeveld, Skurfdebergen, Schlechter 10175. Clanwilliam: Sneeuberg, south

Cedarberg, Edwards 162; Juriesberg, Compton 6263 (NBG). Herschel: Majuba Nek, Sterkspruit,

Hepburn 284 (GRA). Humansdorp: Witte Els Bosch Peak, Esterhuysen 6116 (BOL). Keiskamma

Hoek: Cata Forest Reserve, Story 3279. Kentani: Pegler 82. King Williams Town: Amatola Mtns.,

Dyer 268. Knysna: Pirie, Sim 2822. Komga: Flanagan 1020. Lady Grey: 4 miles W., Godfrey

1542. Molteno: Bews 80 (NU). Mount Ayliff: Amalfi, Godfrey & Scott SH. 1503. Paarl: Du

Toits Kloof Mtn., Tyson 6040 (SAM). Port Elizabeth: Liebenberg 5387. Queenstown: Mtaba

Magwele Mtn., Galpin 2634. Stellenbosch: Jonkershoek, Taylor 3253; 4508. Steynsburg: Molteno

Road, Acocks 8668. Stutterheim: Fort Cunynghame, Sim 2731 ; Rogers 12701. Uniondale: Niekerks-

berg Forest Reserve, near Misgund, Taylor 3259. Worcester: Hex River Valley, Tyson s.n. (GRA).

18

Basutoland. — Leribe: Dieterlen 349; 6323 (SAM). Maseru: Maluti Mtns., Staples 158; Roma,

Schutte s.n. (BOL). Quachas Nek: valley of the Legowa River, Pole Evans 84. Quthing: above

Mphaki, Archibald 534 (GRA).

Orange Free State. — Bethlehem: ravine near village, Phillips 3099. Ficksburg: Riverhill Farm,

Potts 3687. Harrismith: Van Reenen, Bews 81 ; 82; 83 (NU). Heilbron: Maccauvlei, Brandmuller 3.

Kroonstad: Pont 152. Senekal: Doornkop, Goossens 885. Winburg: 10 miles N., Godfrey SH. 1561.

Zastron: “ Mooifontein ”, Liebenberg 5247.

Natal. — Bergville: Mont-aux-Sources, Bayer & McClean 317; Cathedral Peak Forest Res. Stn.,

Killick 1153. Dundee: Vants Drift, Codd 1 86. Durban: Wood 6049 (GRA); Umgeni, Rehmann

8652 (GRA). Estcourt: Cathkin Peak Hostel, West 13; Research Stn., Acocks 9958. Klipriver:

Acton Homes, Doidge s.n. Mapumulo: Buchanan 191 (GRA). Mooi River: Meteor Ridge, Mogg

3388. Mpendle: Huntley 456. Nkahdla: Qudeni Forest Reserve, Fisher & Schweickerdt 40 (NH).

Mtunzini: “ Homewith ”, Mogg s.n. Newcastle: road to Utrecht, Edwards 628. New Hanover:

Balgowan, Mogg 3546. Nqutu: Codd 239. Utrecht: Doringkop, Edwards 1037; Boshoffs Vlei,

Edwards 1046.

Swaziland. — Mbabane, Compton 23763; 23840 (NBG); Rogers 11609 (BOL).

Transvaal. — Belfast: Dullstroom, Galpin 13030. Benoni: Bradfield 372. Bethal: Leendertz

10532. Bronkhorstspruit: on road to Pretoria, De Winter & Codd 167. Carolina: 19 miles S.E.

of Machadodorp, Codd 8278. Ermelo: Maviristad, Pott 5208; Athole Pasture Research Stn., Norval

52; Nooitgedacht, Henrici 1284; 1299. Heidelberg: Henley on Klip, Stent H. 21 584. Johannesburg:

Leendertz 6062; Burtt Davy 832; Canada Junction, Cohen 871. Lydenburg: Zwagershoek, Obermeyer

s.n. Petersburg: Blaauwberg, on Mohlakeng Plateau, Codd & Dyer 8989. Pilgrim’s Rest: Mt.

Anderson, Pole Evans 3794. Potchefstroom : Welverdiend, Louw 874 (GRA). Pretoria: Doornkloof,

Irene, Pole Evans 382; Wolwekloof, Mogg 15909: Rayton, Schweickerdt 1698; Wonderboom South,

Louw 863. Schweizer-Reneke : Burtt Davy 1697 . Soutpansberg: Tzaneen, Sampson 141. Standerton:

Beginsel, Burtt Davy 926. Swartruggens: Sutton 933. Vereeniging: Deneysville, Chippindall 343;

Volksrust: Burtt Davy 150. Wakkerstroom: Oshoek, Devenish 1118; Jantjies Hoek pass, Galpin

70217. Wolmaransstad: Oersonskraal 16, Sutton 77.

This species is variable as may be expected when its wide distribution and its

occurrence in a wide range of habitats is taken into consideration yet it is, on the whole,

easily recognizable and distinct.

When describing the species, Nees (1841) divided it into var. serrulatus, var.

ischaemum and the forma purpurea, on grounds of the size of the spikelets and the

difference in pubescence of the lemma of the sessile spikelets. Hackel (1889) retained

the var. serrulatus but placed the var. ischaemum and forma purpurea in synonomy

under his variety genuinus.

When studying the herbarium material, I found that a small form with conspicuously

hairy racemes, as well as forms with more numerous, but less hairy racemes, occur

in the south-western Cape. From this area to the northern provinces the species shows

a gradual decrease in pubescence as well as an increase in size of plants and of the

number of racemes to the inflorescence. A taller plant with several, almost glabrous

racemes, is found in the Transvaal, Natal, and Orange Fiee State. The size of spikelets

and degree of pubescence of the lemma of the sessile spikelets is not correlated with the

different forms.

Study of a large range of material has thus shown that, in spite of the extreme

variability, there is no valid reason for subdividing this species into subordinate taxa.

8. A. abyssinicus R. Br. in Fres. Mus. Senkenb. 2: 146 (1837); Steud., Syn. PI.

Glum. 1: 371 (1855); Hack, in DC., Monogr. Phan. 6: 465 (1889); Stapf in FI. Trop.

Afr. 9: 222 (1919); Chippindall in Gr. & Pastures of S. Afr. 1: 496 (1955). Type;

Abyssinia, Salt s.n. (BM, holo. ; PRE, photo.!).

Annual, loosely tufted. Culms up to 70 cm long, geniculate, 2-4-noded, simple

or sparingly branched from below or at the middle, terete, glabrous, smooth, the

internodes channelled downwards. Leaves up to 25 cm long and 4 mm wide, expanded,

narrowly linear, long tapering to an acute apex, glabrous or sparingly hairy from

minute tubercles, midrib prominent below, broader and whitish above; ligules 0-5 mm

19

long, membranous, fimbriate, truncate; sheaths terete, glabrous or sparingly hairy

from minute tubercles, smooth, shorter than the internodes. Inflorescence of 2 racemes,

one sessile, the other with a short hairy base. Racemes 8-10 cm long, densely whitish-

villous; joints and pedicels 3-5-4 mm long, narrowly cuneate-linear, transparent

between the thickened edges, densely hairy on the back and sides with the hairs

approximately 3 mm long, tips on one side produced into a short, membranous,

crenulate auricle. Sessile spikelets 6-8 mm long, oblong-lanceolate. Glumes subequal,

membranous; lower dorsally flattened, densely hairy, hairs approximately 4 mm

long, keeled, keels with very narrow subhyaline wings, intracarinal nerves 6-7, evenly

distributed, apex minutely bifid; upper narrowly boat-shaped, sparingly hairy, 3-nerved,

apex with a bristle 5-7-6 mm long. Lower floret reduced to a 5 mm long lemma,

linear-oblong, hyaline, nerveless, ciliate. Upper floret hermaphrodite; lemma 4-7-5

mm long, cuneate-oblong, 3-nerved below, 2-lobed to about the middle, awned;

lobes acute, ciliate; awn about 12 mm long, fine, bent at and twisted below the middle;

palea absent; anthers imperfect, 1-2 mm long. Pedicelled spikelets 6-8 mm long,

male or hermaphrodite. Glumes subequal; lower long hairy on the back, keels

narrowly winged, many nerved, nerves very fine and close with a prominent midnerve,

apex 2-mucronate with a fine bristle 4-5 mm long from between the mucros; upper

subhyaline, long hairy on the back, 3-nerved. apex acute or aristulate. Lower floret

with the lemma 4-5-5 mm long, oblong, faintly 2-nerved, ciliate, apex obtuse. Upper

floret with the lemma lanceolate-oblong, bifid, ciliate, often with a mucro from the

sinus or more or less reduced and entire with a male or hermaphrodite flower; anthers

2- 3 mm long.

This species has been recorded from Abyssinia, Uganda, Malawi, the Cape and

Natal. Tn the Republic it is restricted to the Drakensberg, where it grows on mountain

slopes at altitudes ranging from 5,000-9,000 feet.

Natal. — Estcourt: Ntabamhlope Mtn., West 41. Newcastle: Normandien Pass, Edwards 2819.

Cape. — Maclear: 23 miles N., Theron 2203.

Uganda. — Kigezi: Greenway & Eggeling 7144; Kashenji, Johnston 1375.

The holotype of this species was not seen but the South African material was

compared with the description and with a photograph of the holotype in the National

Herbarium, Pretoria. It is also identical with Schimper 223b, collected in Abyssinia

on Mt. Sholoda. The latter specimen is quoted by Stapf (1919) and is deposited

in Kew.

9. A. pilosellus Stapf in FI. Trop. Afr. 9: 221 (1919). Type; Eritrea, Amasen,

At-Zien, Pappi 5285.

Perennial, loosely tufted with a short oblique rhizome. Culms up to 20 cm long,

geniculate, slender, simple, 2-3-noded, glabrous, smooth. Leaves up to 10 cm long

and 3 mm broad, expanded, linear from a narrow base, sparingly hirsute on both

surfaces with bulbous based hairs, margins scabrid, apex acute; ligules 1-5 mm long,

membranous, obtuse; sheaths terete, sparingly hirsute with bulbous based hairs,

persistent. Inflorescence of 2 racemes, one sessile, the other with a short hairy base.

Racemes 3-5 cm long, straight, yellow-green to purplish; joints and pedicels linear,

3- 3 • 5 mm long, subtransparent between the thickened edges, ciliate on both margins

with hairs 1 • 5-2 mm long, tips cupular, truncate. Sessile spikelets hermaphrodite,

7 mm long, lanceolate-oblong; callus short, obtuse, densely bearded. Glumes subequal;

lower 7 mm long, lanceolate-oblong, 7-nerved, subchartaceous, dorsally shallowly

concave, long hirsute, unequally winged from the keels, one wing very narrow, scabrid,

apex unequally 2-dentate, each tooth with a fine mucro; upper 6 mm long, narrowly

boat-shaped, membranous, 3-nerved, shortly pilose on the margins, keel scabrid,

apex minutely bifid, awned, awn 3 mm long. Lower floret reduced to the lemma, 5 mm

20

long, oblong-lanceolate, hyaline, 2-nerved, sparingly pilose on the margins, apex acute.

Upper floret with the lemma 4 mm long, hyaline, 1-nerved, lobed to about the middle,

awned, awn 8 mm long, bent at and twisted below the middle; palea absent. Pedicelled

spikelets neuter, reduced to the glumes; lower 4-5 mm long, lanceolate-oblong, hirsute,

7- nerved, apex minutely bifid, shortly mucronate; upper 2 mm long, oblong, hyaline,

nerveless, apex truncate.

Natal.— Bergville: Drakensberg, low area pass between Saddle and Twins, Edwards 2133.

The plant, from which the above description was drawn up, was collected on the

Drakensberg in Natal in July, 1957, at an altitude of 9,200 feet. To my knowledge

this is the only record for South Africa. It compares very well with Stapf’s description

of A. pilosellus in the Flora of Tropical Africa (1919) and could possibly be a depauperate

specimen of that species. I have not seen the type of A. pilosellus, Pappi 5285, collected

at At-Zien in Eritrea, but Edwards 2133 was sent to Kew for comparison. Mr. W. D.

Clayton kindly examined it and reported that it was probably A. pilosellus but that

a definite identification would not be possible until a detailed study of the East African

montane species of Andropogon had been made.

10. A. lacunosus J. G. Anderson in Bothalia 8: 113 (1962). Type: Transvaal,

Ermelo, Athole Pasture Research Station, Gericke A. 49 (PRE, holo. !).

Perennial, caespitose with a short rhizome. Culms up to 80 cm long, erect, slender,

glabrous, striate, often grooved, terete or the basal internodes slightly compressed,

2-4-noded; nodes glabrous. Leaves up to 20 cm long and 5 mm wide, narrowed

towards the base and tapering to an acute apex, broadest at about the middle, 7-nerved

with the midnerve prominent abaxially, sparsely to densely pilose with bulbous-based

hairs between the nerves, often densely pilose near the ligule; ligules 1 mm long,

membranous, ciliolate, truncate; basal sheaths often tinged with purple. Inflorescence

of 2-3 racemes, 5-5-7 cm long; joints and pedicels linear, 3-4-5 mm long, margins

pubescent. Sessile spikelets hermaphrodite, 6-8 mm long, linear. Glumes unequal;

lower 6-8 mm long, narrowly-lanceolate, deeply grooved on the back, the groove

densey pilose in the lower and with 5-8 variably shaped shallow pits on either side

of the groove, chartaceous, 7-9-nerved, keeled with the keels narrowly winged, ciliate

in the upper half, apex obscurely bifid; upper 5-6 mm long, narrowly boat-shaped,

thinly membranous, 3-nerved with the midnerve distinct and produced into a 1-3 mm

long scabrid bristle, lateral nerves obscure, margins narrowly winged upwards, wings

pilose along the margins, apex obtuse. Lower floret reduced to the approximately

5 mm long lemma, lanceolate, hyaline, nerveless, margins inrolled and pilose in the

upper half, apex obscurely bifid. Upper floret bisexual with the stamens reduced to

staminodes; lemma 4-7-5 mm long, linear, hyaline, nerveless, lobed to about the

middle, lobes acute, pilose along the margins in the upper half, awned from between

the lobes; awn slender, 10-15 mm long; palea absent. Pedicelled spikelets male,

8- 9 mm long. Glumes unequal; lower 8-9 mm long, lanceolate, chartaceous, keeled,

keels and margins pilose, 11 -nerved with the nerves grouped together, one group of

3 in the middle and 2 marginal groups of 4 each, with 2 longitudinal grooves and 5—12

variably shaped, shallow pits on the back, midnerve excurrent into a very short bristle

or apex acute; upper 7-5-8 mm long, lanceolate, membranous, 3-5-nerved, obscurely

winged, pilose on the margins, apex acuminate. Lower floret reduced to the lemma,

6-5-7 mm long, hyaline, obscurely 2-nerved, pilose on the margins; palea absent.

Upper floret male, lemma about 4-5 mm long, hyaline, nerveless, pilose along the

margins, apex emarginate; palea absent; anthers linear, 4 mm long.

This species is rare and has been found only in the Ermelo, Groblersdal, Belfast,

Lydenburg, Pilgrims Rest and Wakkerstroom districts of the eastern and north-eastern

Transvaal. It occurs in vleis and near streams, usually in sandy soils, at altitudes

ranging from 4,700-6,800 feet.

21

Transvaal. — Belfast: Dullstroom, Hector s.n. Ermelo: Athole Pasture Research Station, Letole

River, Malherbe 54; Norval 8; 122. Groblersdal: 28-j miles N.E. by E., Acocks 20908. Lydenburg:

15 miles W. of Sabie, Codd 6441. Pilgrim’s Rest: Mariepskop, Meeuse 9940; near Reitz’s grave,

Van der Schijff 5992; 6 miles N. of Graskop, Codd 6450. Wakkerstroom : Oshoek, Devenish 1090.

A. lacunosus is a distinct species and is unique in the genus Andropogon because

of the shallow, variably shaped pits on the lower glumes of both the sessile and

pedicelled spikelets.

11. A. amplectens Nees, FI. Afr. Austr. 104 (1841); Steud., Syn. PI. Glum. 1 : 372

(1855); Hack, in DC., Monogr. Phan. 6: 453 (1889); Stapf in FI. Cap. 7: 342 (1898);

FI. Trop. Afr. 9: 243 (1919); Chippindall in Gr. & Pastures S. Afr. 1: 498 (1955).

Type: Cape, “ between Omtata and Omgaziana ”, Drege s.n.

Perennial, densely tufted. Culms up to 150 cm long, slender, few-9 noded, erect

or rarely subdecumbent, glabrous, smooth. Leaves glabrous, or rarely hairy to densely

pubescent; blades linear from a wider rounded base or those of the culms cordate

or subcordate and amplexicaul, tapering to a long setaceous apex, up to 30 cm long

and 10 mm wide at the base, those of the innovations up to 15 cm long and 2-5 mm

wide, subglaucous or suffused with purple, usually glabrous, rarely with a few scattered

hairs or densely pubscent, margins somewhat scabrid; ligules 1 mm long, membranous,

fimbriate, truncate or rounded; sheaths terete, tight, glabrous, the lowest and those

of the innovations short, sometimes purplish, subpersistent. Inflorescence of 2, rarely

3 racemes; spathes linear-lanceolate, long tapering upwards, glabrous, pale green, at

length tightly inrolled; spatheoles similar to the spathes, about 8 cm long, tapering

to a setaceous point; peduncles exerted. Racemes 5-8 cm long, straight or somewhat

flexuous; joints and pedicels very similar, 5-6 mm long, clavate or cuneate, distinctly

swollen upwards, densely ciliate on the margins, tips produced into very oblique, slightly

denticulate cupules. Sessile spikelets hermaphrodite, 7-9 mm long, laterally compressed,

wedged in between joint and pedicel, linear-oblong, glabrous, green, often reddish

towards the tips; callus acute to subacute, 2 mm long, shortly bearded. Glumes equal;

lower 7-5-8 mm long, lanceolate, keeled with the keels rounded, broad and smooth

below, acute and scabrid near the tips, narrowly grooved between the keels and with

2 fine lateral furrows in the upper third, apex bifid; upper 7-5-8 mm long, boat-shaped,

subcoriaceous, 1 -nerved, glabrous or the nerve dorsally scabrid, apex mucronate or

aristulate. Lower floret reduced to the lemma, 5-6 mm long, lanceolate, nerveless

or faintly 2-3-nerved, sparingly ciliate, hyaline, apex acute. Upper floret hermaphrodite;

lemma 5-7-6 mm long, linear-lanceolate, sparingly ciliate, 3-nerved, deeply lobed,

lobes acute, awned from between the lobes, awn stout, 3-5-5 cm long, kneed and

pubescent below the middle; palea 1-6-2 mm long, linear-oblong, nerveless, ciliate;

anthers 4 mm long. Pedicelled spikelets male, 9-12 mm long, lanceolate-acuminate,

dorsally compressed. Glumes subequal; lower 9-12 mm long, subherbaceous, linear-

lanceolate, many-nerved between the scabrid keels, usually produced into an awn

2-8 mm long; upper 9-11 mm long, linear-lanceolate, 3-5-nerved, sparingly ciliate

on the keels, apex acuminate. Lower floret reduced to the lemma, 6-9 mm long,

membranous, linear-lanceolate, faintly 3-nerved, apex acute. Upper floret with the

lemma 6-8 mm long, membranous, 1 -nerved, apex bifid; palea 2-3-5 mm long, mem-

branous, nerveless; anthers 4-4-5 mm long.

This species occurs from the East London district of the eastern Cape to Natal

and Swaziland and is widely distributed and common in the Transvaal. From here

it extends into the Orange Free State and northern Cape. It has also been recorded

from Bechuanaland, Mozambique, Rhodesia, Malawi and the Congo Republic. It

usually occurs in dry open veld and on hillsides, but is often found also in marshy

areas and other moist situations.

22

Cape. — Barkly West: Daniels Kuil, Gets 2999 (KMG). East London: Kidds Beach, Comins 1484

(GRA). Hay: Jaspar Hill at Clifton, Acocks 2142. Kentani: near Kei Mouth, Flanagan 981 ; Pegler

1056. Kuruman: Pole Evans 2414; Marloth 1114 (GRA). Mount Currie: Clydesdale, Tyson 931

(SAM). Tsolo: Umtwynkwa Cutting, Galpin 6893. Umtata: “ Morely ”, Drege s.n. (K, S). Vryburg:

Armoedsvlakte, Henrici 44: Bietjiesvlakte, Henrici 73.

Orange Free State. — Heilbron: Maccauvlei, Brandmuller 42. Kroonstad: Vais River, Pont 27;

143. Ventersburg: 12 miles S., Godfrey SH. 1563.

Natal. — Bergville: Ladysmith road, Edwards 661 . Camperdown: Nagle Dam, Wells 1190 (NU);

1865. Estcourt: Acocks 11454. Hlabisa: Hluhluwe Game Reserve, Ward 1977. Lions River: Moll

617. Nkandla; Qudeni, Edwards 1297. Pietermaritzburg; Albert Falls, Comins 265 (NU). Port

Shepstone: Port Edward, Huntley 720; Oribi Flats, McClean 528. Tugela: Bulwer Farm, Edwards

1920. Ubombo: Mpangazi Lake, Strey & Huntley 5017. Umzinto; Dumisa Station, Rudatis 884.

Upumulo: Buchanan 194 (BOL, NH).

Swaziland. — Mbabane: Ngwena Mtn. Range near Forbes Reef, Meeuse 10136; Ukutula, Compton

25787; 26698.

Transvaal. — Brits: Jacksonstuin, Van Vuuren 379. Delmas: Schmc/z 16 (GRA). Ermelo: Experi-

mental Farm, Burtt Davy 7720. Heidelberg: Henley-on-Klip, Stent s.n.; Uitgevallen 197, Burn Davy

13667. Johannesburg: 8 miles S., Lintner 45; Houghton, Cohen 650. Letaba: The Downs, Junod

4096; Duivvelskloof, Scheepers 822. Lichtenburg: Grasfontein, Sutton 392. Marico: Liebenberg

5321. Nelspruit: Nohpe, Van der Schijff 1512; Shabin Kop, De Winter & Codd 458. Pietersburg:

Iron Crown Peak, Swanepoel s.n. Potchefstroom: Burtt Davy 2700. Pretoria: Doornkloof, Pole

Evans 351; Hoorns Nek, Codd 770; Onderstepoort Station, Smith 6169; Premier Mine, Rogers

25048. Ventersdorp: Palmietfontein, Botha 26. Vereeniging: Leeukuil, Story 91. Warmbad: Pole

Evans & Smuts 690. Waterberg: Nooitgedacht 699, Acocks & Naude 31.

Bechuanaland. — Lobatsi: Kanye, Miller B/304.

Rhodesia. — Bubi : Gwampa Forest Reserve, Goldsmith 37/56. Matobo: Matopas, Rattray 300.

Que Que; Gokwe, Bingham 481A.

Mozambique. — Beira: Dondo, Torre 5899; Santaca, Maputo, Gomes <fc Sousa 3955.

I have not seen the type specimen of A. amplectens but the South African material

agrees with the description by Nees (1841) and with a specimen collected by Drege

simultaneously with the type. The latter specimen was recorded from “ Morely ” in

the Umtata district and as this is the type locality there is no doubt as to the identity

of the species.

A. amplectens is a distinct species with the culm leaves characteristically rounded

or subcordate and subamplexicaul at the base, tapering sharply to a long, fine, apex.

The leaves and culms may be dark green or frequently tinged with red, purple or brown.

In its early and flowering stages the plant is usually conspicuously glaucous.

The width of the leaf-bases and the pubescence of the vegetative parts of the plant

vary considerably. Examination of the South African material revealed that the leaf-

bases are from 2-5 mm to 1 - 3 cm wide. This variation occurs in plants from the same

locality and even in leaves on the same plant. For this reason 1 am reluctant to include

any of the broad-leaved South African plants in the var. diversifolius (Rendle) Stapf,

described originally from Angola.

It is of interest to note that plants collected along the coast in the Port Shepstone

district of Natal often have a subdecumbent or sprawling habit, with leaves 5-12 cm

long and 6-10 mm wide.

12. A. schinzii Hack, in DC., Monogr. Phan. 6: 458 (1889); Stapf in FI. Cap.

7: 341 (1898); FI. Trop. Afr. 9; 245 (1919); Chippindall in Gr. & Pastures S. Afr.

1: 499 (1955). Type: South West Africa, Amboland, Oshando, Schinz 670 (Z, holo. ! ;

BOL, iso.!; PRE, photo.!, fragment!).

A. schinzii Hack, ex Schinz in Verh. Bot. Ver. Brandenb. 30: 139 (1888), nom. nud.

Perennial, densely tufted with a slightly bulbous base. Culms up to 120 cm tall,

erect, slender, glaucous, glabrous, terete or often channelled, simple and sometimes

woody below with barren solitary or fascicled branches from the lower nodes and few

23

solitary (rarely 2-3-nate) leafy long and erect flowering branches higher up. Leaves

up to 30 cm long and 8 mm broad, linear from a narrow base, long tapering to a fine

apex, slightly rough or puberulous on both surfaces, rarely with some additional very

long and spreading hairs near the ligule; ligules 1-2 mm long, membranous, truncate,

laterally produced and adnate to the auricles of the sheath; sheaths terete, tight,

glabrous, firm, the lower and intermediate mostly longer than the internodes, shortly

auricled at the mouth, the basal long, persistent. Inflorescence of 2 racemes, conspi-

cuously hairy from the long soft hairs on the joints and pedicels; spathes subherbaceous,

gieen, sometimes faintly tinged with purple, at length tightly inrolled with a more

or less reduced blade. Racemes 5-8 cm long, one sessile, the other with a short base,

straight or flexuous, villous, dull or yellow-green, sometimes purplish; joints and

pedicels equal or subequal, cuneate, stout above, much attenuated towards the base,

4-5 mm long, rigidly ciliate on both margins, cilia 3 -5-4 -5 mm long, tips obliquely

cupular, minutely dentate. Sessile spikelets hermaphrodite (except the lowest 1 or 2

of the sessile raceme), 6-7 mm long, narrowly linear, laterally compressed, loosely

wedged in between joint and pedicel; callus short, obtuse, shortly bearded. Glumes

subequal; lower 6-7 mm long, subcoriaceous, keeled with a deep narrow hyaline

groove between the keels, nerves 4 on each side, one close to the groove, the other

in the keels; keels obtuse and smooth in the lower three-quarters, acute and scaberulous

upwards, apex minutely bifid; upper boat-shaped, 6-7 mm long, membranous, 3-nerved,

pilose on the margins, awned, awn 7-10 mm long, fine, straight. Lower floret reduced

to a lemma, 5-5-7 mm long, linear-oblong, 2-nerved. pilose on the margins, hyaline,

apex acute or obtuse. Upper floret hermaphrodite, lemma 5-6 mm long, oblong,

3-nerved below, pilose on the margins, conspicously 2-lobed, lobes acute, awned from

between the lobes, awn slender, 25-35 mm long, geniculate, scaberulous; palea 3-5 mm

long, linear-oblong, hyaline, nerveless; anthers 3-4 mm long. Pedicelled spikelets

male, dorsally compressed, oblong or lanceolate-oblong, acute, purplish or reddish.

Glumes subequal; lower flat or rounded on the back, 5-6 mm long, membranous,

keeled, keels ciliate, awned from a bifid apex, awn 5-7 mm long, fine, straight; upper

submembranous, 6-7 mm long, 3-nerved, pilose on the margins in the upper half,

awn 1-2 mm long. Lower floret reduced to the lemma, 6-7 mm long, hyaline, 2-nerved,

narrowly keeled, pilose on the margins. Upper floret with the lemma 6-7 mm long,

hyaline, 3-nerved, pilose on the margins, apex mucronulate; palea 4-5-5 mm long,

hyaline, nerveless, apex obtuse or acute; anthers 3-4 mm long.

A. schinzii is found in several districts of the Transvaal, the Hay, Postmasburg

and Kuruman districts of the northern Cape and in the central and northern part of

South West Africa. It has also been recorded from Swaziland. Bechuanaland, Rhodesia

and the Congo Republic. It usually grows on rocky slopes or ledges on hillsides.

Cape. — Hay: Floradale, Ferrar 47 ; 6098 (KMG); Esterhuysen 2368; Bingap, Acucks 5730 (KMG,

NH, BOL). Griqualand West: Klipfontein, Burchett 2164-2 (SAM). Kuruman: Broncote, Ester-

huysen 1152 (BOL, KMG). Postmasburg: Kaapstewel, Cooke s.n. (GRA); 6335 (KMG).

Swaziland. — Mbabane: Komati Bridge, Compton 26818.

Transvaal. — Krugersdorp: Gladysvale, Rodin 3930. Letaba: 174 miles W. of Letaba Camp,

De Winter & Codd 692; Duiwelskloof, Westfalia Estate, Scheepers 666. Messina: Govt. Ranching

Station, Du Plessis s.n.; Dongola, De Winter & Codd 286. Nelspruit: Skipberg, Van der Schijff 3397 .

Potgietersrus : Pyramid Estate, Galpin 8885; Swerwerskraal, Acocks 8869; Doornhoek, Walker s.n.

Pretoria: Klapperkop, Mogg 16231; Wonderboom Reserve, Repton 2070; Rust der Winter Dam,

De Winter & Codd 247. Rustenburg: ! lexkrantz, Codd 1066. Foster River, Bunt Davy 82. Waterberg:

104 miles N.E. of Oslo, Codd 3998.

Bechuanaland. — Francistown: between Totume and Bushman Mine, Pole Evans 3266; Gordon

87; 102.

South West Africa. — Grootfontein : Auros, Volk 594b; Farm Kumkauas, Kinges 2818; 2883;

20 miles W., Schoenfelder 26; 5 miles N., Schweickerdt 2063. Kaokoveld: Omberera, De Winter

& Leistner 5487. Otjiwarongo: Ozondjache, Volk 517. Rehoboth: Paulinenhof, Strey 2575. Wind-

hoek: Sittmann 3; between Haris and Aub, Pearson 9601 (BOL).

24

Rhodesia. — Makoni: Rusape, Davies 1075. Matobo: Matopos, Brain 3874. Shangani: Gwampa

Forest Reserve, Goldsmith 151/55.

A. schinzii is easily distinguished by its slightly bulbous base, the usually branched

culms and the glaucous-grey colour of the culms and leaves. Because of the long

soft hairs on the margins of the joints and pedicels, the racemes are villous in appearance.

A characteristic feature of this species is that the upper glumes of the sessile spikelets

as well as the lower glumes of the pedicelled spikelets are each provided with a straight

awn 5-10 mm long.

13. A. ravus/. G. Anderson in Bothalia 7: 417 (1960). Type: Natal, Bergville,

Cathedral Peak, Killick 1261 (PRE, holo. !).

Perennial, often with a branched system of knotty rhizomes. Culms up to 90 cm

long, usually 45-60 cm, erect, simple, 2-3-noded, glabrous, faintly striate or smooth.

Leaves up to 25 cm long, expanded, rarely almost filiform, 2-6 mm wide, linear, tapering

to a setaceous apex, narrowed towards the base or base slightly rounded, glabrous,

glaucous-grey or green, margins often scabrid; ligule 1-5-2 mm long, membranous,

scarious; sheaths glabrous, striate, pallid or upper suffused with purple, shorter than

the internodes, leaf-blade of uppermost sheath sometimes reduced to a short awn,

mouth often with a few long hairs. Inflorescence of 2 (very occasionally 3) spike-like

racemes, shortly pedicelled or lowermost sessile. Racemes 7-12 cm long, glaucous-grey

or green and usually suffused with dull purple; joints and pedicels 6-7 mm long,

cuneate-clavate, ciliate on the margins, cilia approximately 4 mm long. Sessile spikelets

hermaphrodite, linear-oblong; callus broad and obtusely rounded, bearded. Glumes

subequal; lower 6-9 mm long, dorsally with a deep, narrow or wide median groove,

thinly coriaceous; 3-nerved on each side of the groove, sharply keeled upwards, keels

narrowly winged, scabrid to ciliate in the upper half, apex mucronate; upper boat-

shaped, keel scabrid or ciliate in the upper third, finely ciliate on the margins, 1-3-

nerved, apex subacute. Lower floret with the lemma 6-8 mm long, 2-keeled with

sharply indexed margins, margins ciliate in the upper half, hyaline, 3-nerved, apex

obtuse ; palea a nerveless hyaline scale 2-3 mm long. Upper floret with the lemma

5-6 mm long, deeply lobed, lobes acute, awned from between the lobes, awn 1 • 5-2 cm

long, twisted below the knee; palea absent. Pedicelled spikelets male, dorsally

compressed. Glumes unequal; lower 7-12 mm long, conspicuously broad, keeled,

keels and margins ciliate or with stiff hyaline hairs, apex obtuse or mucronate; upper

shallowly boat-shaped, 6-9 mm long, 3 or faintly 5-nerved, margins ciliate, apex acute.

Lower floret with the lemma 7-8 mm long, margins indexed, sparsely ciliate in the upper

half, hyaline, 3-nerved, apex obtuse; palea a nerveless hyaline scale. Upper floret

with the lemma 5-6 mm long, margins indexed, apex obtuse; palea absent; anthers

linear, 4-5 mm long.

A. ravus is recorded from mountain slopes near Mbabane in Swaziland and in the

foothills of the Drakensberg from Bethlehem in the Orange Free State to Maclear in

the eastern Cape Province. It usually occurs in open grassveld at altitudes ranging

from 4,700-8,000 feet.

Cape. — Maclear: Lower Pitsing, Acocks 12191.

Orange Free State. — Bethlehem: Golden Gate, Story 1957.

Natal. — Bergville: Cathedral Peak, Edwards 1935; 1954 (NU); Mont-aux-Sources, Bayer &

McClean 179; Edwards 639; Cleft Peak, towards Indumeni, Gomel! s.n. Klip River: Van Reenen,

Phillips 8728.

Swaziland. — Mbabane: Ukutula, Compton 25593; Forbes Reef, Codd <£ Midler 305.

25

This species is usually glaucous-grey in colour. It can be confused with A.

amplectens but is distinguished from that species by the shape of the callus. In A.

ravus the callus is conspicuously broad and rounded, whereas it is acute or subacute

in A. amplectens.

A depauperate form of A. ravus has been collected near Cleft Peak in the Cathedral

Peak area. Except for being smaller in stature with narrow reddish leaves, I can

find no difference in spikelet characters. This form is represented by Gomel l s.n. and

Edwards 1935 and 1954.

14. A. filifolius ( Nees ) Steud., Syn. PI. Glum. 1: 374 (1855); Hack, in DC.,

Monogr. Phan. 6: 453 (1889); Stapf in FI. Cap. 7: 342 (1898); Chippindall in Gr.

& Pastures S. Afr. 1: 497 (1955). Type: Uitenhage, Zuurebergen, northern slopes,

Drege s.n. (S, lecto.!).

Heteropogon filifolius Nees, FI. Afr. Austr. 102 (1841). Syntypes: Olifantshoek,

Boschesman River, Ecklon s.n. (SAM!); Uitenhage, Tigerberg, Ecklon s.n.; Zuur-

bergen, Drege s.n. (S!).

Perennial, densely tufted. Culms up to 70 cm tall, simple, erect, glabrous, terete,

3-4-noded. Leaves mostly basal; blades up to 30 cm long, filiform, rarely expanded

and then 2-5 mm wide, firm, flexuous, glabrous or pubescent on the inside near the

ligule, apex acute; ligules coriaceous, 0-5-2 mm long, truncate; sheaths shorter than

the internodes, terete, tight, striate, glabrous, lowest persistent, at length breaking up

into fibres. Inflorescence of 2 spike-like racemes, one sessile, the other with a glabrous

base 7-11 mm long. Racemes 3-10 cm long, straight; joints and pedicels similar,

6-8 mm long, cuneate-linear, attenuated towards the base, rigidly ciliate on the margins,

tips crenulate, cupular. Sessile spikelets hermaphrodite, laterally compressed, wedged

in between joint and pedicel; callus acute to subacute, shortly bearded. Glumes

subequal, subcoriaceous, glabrous; lower 6-8 mm long, linear, keeled, keels rounded,

narrow, almost contiguous, narrowly grooved between the keels, apex obtuse, hyaline;

upper 6-8 mm long, boat-shaped, glabrous, 1-nerved, apex acute with a bristle 1 -3-1 -5

mm long. Lower floret reduced to the lemma, 5-5-6 mm long, lanceolate, membranous,

hyaline, 1-nerved or nerveless, ciliate on the margins, apex acute. Upper floret with

the lemma 4-5-5 mm long, linear, hyaline, 3-nerved, bilobed, lobes oblong, awned

from between the lobes, awn 3-5 cm long, stout, kneed at about the middle, pubescent

below; palea 2-7-3 mm long, ovate, hyaline, glabrous, nerveless, apex acute; anthers

3—4 - 5 mm long. Pedicelled spikelets male, lanceolate, dorsally compressed or subterete,

glabrous. Glumes unequal; lower 9-20 mm long, flat or rounded on the back,

herbaceous, acutely 2-keeled, keels narrowly winged, intercarinal nerves many, apex

lobed with a bristle from between the lobes, bristle about 2 mm long ; upper 8-14 mm long

lanceolate, 3-nerved, ciliate in the upper half or glabrous, apex finely acuminate. Lower

floret reduced to the lemma, 8-13 mm long, linear-oblong, 3-4-nerved, ciliate on the

margins, apex acuminate. Upper floret with the lemma 7-12 mm long, linear, 1-nerved,

glabrous or ciliate on the margins; palea 2-4 mm long, linear, hyaline, nerveless;

anthers 5-7 mm long.

A. filifolius is indigenous to Southern Africa and has been recorded from the

southern and eastern Cape, Basutoland, Natal, Swaziland and the northern and eastern

Transvaal. It grows in sour, open grassland on mountains and hillsides, often in rocky

soil.

Cape. — Albany: Cradock road, Wells 2758 (GRA); Grahamstown, Daly & Sole 109 (GRA).

Komga: near Kei Mouth, Flanagan 2348. Matatiele: Melkfontein, Story 537. Mount Ayliff: Kok-

stad, Goossens 268. Port Elizabeth: Van Stadensberg, Gillett 2384 (BOL).

Basutoland. — Butha Buthe: Moteng, Guillarmod 3842. Leribe: Dieterlen 397a.

26

Natal.- Bergville: Natal National Park, Galpin 10360; Cathedral Peak, Killick 949. Durban:

Clairmont, Schlechter 3092 (GRA). Estcourt: Giants Castle, Wood 10546; Griffins Hill, Acocks

10633; Ntabamhlope, West 470; 917; Mooi River, Mogg 3104. Lions River: Balgowan, Moll 1205.

Mpendle: Tillietudlem, Huntley 455. Nkandla: Codd 235; Qudeni, Fisher & Schweickerdt 30 (NH,

NU). Pietermaritzburg: Town Hill, Fisher 285 (NU); Bews 77 (NU). Utrecht: Edwards 630.

Swaziland. — Mbabane: Ukutula, Compton 24721; 24958.

Transvaal. — Barberton: Burtt Davy 87; Kaapsche Hoop, Pole Evans 1009. Belfast: Burtt Davy

88. Carolina: near Bossis, Burtt Davy 86. Ermelo: Maviristad, Pott 81 . Heidelberg: Vrisgewaagd,

Strey 3068. Letaba: The Downs, Junod 4092; Duiwelskloof, Scheepers 966. Pilgrim’s Rest: 12)-

miles from Graskop on road to Vaalbank, De Winter & Codd 221 ; Graskop, Kowyns Pass, De Winter

<& Codd 153. Waterberg: Krantzberg, Dyer & Verdoorn 4242.

This is a variable species in which several “ forms ” can be distinguished. These

forms differ mainly in size, leaf shape and length of the pedicelled spikelets. The most

common form occurs in the Drakensberg in the Cape, Basutoland and Natal and is

usually a smaller plant of which the leaves are all filiform, the ligules about 0-5 mm long,

the racemes 3-6-5 cm long and the pedicelled spikelets 9-12 mm long. A second

form is represented by plants with the leaves all filiform or filiform mixed with narrowly

expanded leaves, the ligules 0-5-2 mm long, the racemes 6-7 cm long and the pedicelled

spikelets up to about 17 mm long. An extremely robust form, represented by Dyer

& Verdoorn 4242, Pole Evans 1009, and Compton 24758, collected in the Waterberg,

Barberton and Mbabane districts respectively, has leaves which are filiform mixed

with leaves up to 5 mm wide, racemes 7-10 cm long and pedicelled spikelets 17-20

mm long.

Except for the variation in size of the various organs, no other noteworthy differences

could be found. In contrast to the variability in the length of the pedicelled spikelets,

the size of the sessile spikelets remains more or less the same. These forms intergrade

and should probably be regarded as ecotypes.

15. A. schirensis Hochst ex A. Rich. var. angustifolius Stapf in FI. Cap. 7: 340

(1898), sphalm “ angustifolia FI. Trop. Afr. 9: 247 (1919); Chippindall in Gr.

& Pastures S. Afr. 1: 497 (1955). Syntypes: Several. Lectotype: Orange Free State,

Thaba Nchu, Burke 434 (K, lecto.!; SAM!; PRE, fragment).

A. amplectens Nees var. natalensis Hack, in Mem. Herb. Boiss. 20: 9 (1900). Type:

Natal, near De Beers, Drakensberg, Wood 6002 (NH, iso.!).

Perennial, densely tufted. Culms up to 120 cm long, erect, slender, glabrous

and smooth, terete, simple, rarely branched, 3-4-noded. Leaves with the blades up

to 35 cm long and 7 mm wide, expanded, linear from a narrow base, long tapering

to a fine apex, glaucous or very often reddish, glabrous or often pubescent or scabrid

at the base, margins scabrid; ligules 1-5 mm long, membranous, truncate; sheaths

terete, smooth, rarely pubescent near the ligule, shorter than the internodes. Inflo-

rescence of 2 racemes (rarely 2-5), one sessile, the other with a base about 6 mm long.

Racemes 5-8 cm long, straight or flexuous, conspicuously dorsiventral exhibiting

on the one side the sessile spikelets with the joints and pedicels and on the other side

the broad backs of the pedicelled spikelets with the dense cilia of the pedicels between

them; joints and pedicels very similar, clavate-cuneate, 4-5 mm long, densely ciliate

on both margins, cilia usually shorter in the lower portion, 4-5 mm long towards the

apex, tips obliquely cupular or the pedicels produced into a broad tooth. Sessile

spikelets hermaphrodite, narrowly linear, laterally compressed, glabrous, gieen to

reddish-brown, tightly wedged in between joint and pedicel; callus 1-1.5 mm long,

obtuse to subacute, shortly bearded. Glumes equal; lower 5-6 mm long, linear-

lanceolate, thinly coriaceous, keeled with a narrow, deep hyaline groove between the

keels, keels obtuse, smooth below and scaberulous above the middle, 1 -nerved in the

keels, apex obscurely bifid ; upper 5-6 mm long, rounded on the back up to beyond

the middle then slightly compressed and keeled, subcoriaceous, 3-nerved with the side

27

nerves indistinct, margins very narrow, ciliate, apex acute to mucronulate. Lower

floret reduced to the lemma, 5-5-5 mm long, linear-oblong, membranous, hyaline,

2-nerved, winged, shortly ciliate on the wings, apex subobtuse. Upper floret with

the lemma 4-5 mm long, linear-oblong, membranous, hyaline, 3-nerved, deeply 2-lobed,

lobes acute, lanceolate, ciliolate, awned from between the lobes, awn 2-5-3 cm long,

bent and twisted, slender, scaberulous; palea 3-6-4 mm long, ovate-oblong, hyaline,

nerveless, apex obtuse; anthers 3-5 mm long. Pedicelled spikelets male, 7-8 mm

long and 1-2 mm wide, dorsally much compressed, reddish or purplish-brown. Glumes

more or less equal; lower 7-8 mm long, linear-lanceolate, subherbaceous, 7-15-nerved,

keeled, keels acute, scabrid or rigidly ciliolate, apex acute or acuminate; upper 7-8

mm long, membranous, 3-nerved, ciliolate on the margins, apex acuminate. Lower

floret reduced to the lemma, 7-7-5 mm long, narrowly-oblong, membranous, hyaline,

2-nerved, ciliolate on the margins, apex acute. Upper floret with the lemma 6-7 mm

long, linear, hyaline, 1-nerved; anthers 5 mm long.

This variety is common in Natal and the northern, eastern and central Transvaal.

It also occurs in the Orange Free State, Swaziland, the Barkly West, Kuruman,

Mafeking and Vryburg districts of the northern Cape and in South West Africa. Further

north it has been recorded from Angola, Rhodesia, Zambia and Malawi.

It grows mainly in open grassland or on hillsides, usually in stony or sandy soil,

occasionally in loamy soil on the edge of pans or along riverbanks.

Cape. — Barkly West: Daniels Kuil, Esterhuysen 2044; Pillans 2089. Kuruman: 11 miles N.W.,

Leistner 598. Mafeking: near Mosita, Brueckner 371. Vryburg: between Vryburg and Tigerkloof,

Brueckner 1087; Jakkalskop, Scheepers 2757.

Basutoland. — Leribe: Dieterlen 397b.

Orange Free State. — Ficksburg: farm "Aletta ”, Mat thy sen 11. Kroonstad: Vais River, Pont

131. Parys: Cohen 101. Winburg: Pretorius Game Reserve, Van Zinderen Bakker 1076; 1170.

Natal. — Bergville: Sentinel Park to Mont-aux-Sources, Edwards 639 (NU). Dundee: near Wasch-

bank, Pentz & Acocks 10282; near Rorkes Drift, Edwards 673. Durban: Berea, Broad 104

(NU). Estcourt: Giants Castle, West 1618 (NH); Symons 15700. Hlabisa: St. Lucia Estuary,

Feelv, Tinley & Ward 3; W. of Charters Creek, Feeiy, Tinlev & Ward 7. Kliprivier: Van Reenen,

Schlechter 6983 (BOL, GRA). Lions River: Mooi River, Rietvlei, Buchanan 193; 196; Nottingham

Road, McClean 948 (NH). Mpendle: Clarkson 156 (NU). Mtunzini: Ngoya Forest Reserve, Huntley

641; 644 ; 815. Newcastle: Road to Utrecht, Edwards 624; Buchanan 195 (NH). Nkandla: Qudeni

Forest Reserve, Fisher & Schweickerdt 28 (NH, NU). Pietermaritzburg: Table Mtn., McClean 195;

Bews 75 (NU); Inchanga, Most 92 (NU). Utrecht: Amajuba, Burtt Davy 154. Vryheid: Hlobani,

above Mkusi River, Johnstone 387 (NU). Weenen: Farm Schurfdepoort, West 1799 (NH).

Swaziland. — Mbabane: Ukutula, Compton 26696; Forbes Reef road, Compton 25742; Forbes

Reef, Meeuse 10134.

Transvaal. — Barberton: Thorncroft 73a. Belfast: Machadodorp, Williams 6315; Pole Evans

H. 16962. Brits: Jacksonstuin, Mogg 14277; Van Vuuren 457. Carolina: Van der Schijff 2605.

Ermelo: Billys Vlei, Burtt Davy 78. Heidelberg: Henley-on-Klip, Stent s.n. Johannesburg: Hougton,

Cohen 655. Letaba: Westfalia Estate, Scheepers 519; 1236. Lichtenburg: road to Zeerust, Kinges

1415. Lydenburg: Mt. Anderson, Strey 3036. Marico: Liebenberg 5323. Middelburg: Burtt Davy

152. Nelspruit: Shabin, Van der Schijff 1470. Pietersburg: 12 miles E., Van Vuuren 1455. Pilgrim's

Rest: Mariepskop, Van der Schijff 6151; 6 miles N. of Graskop, Codd 6450; Iron Crown Mtn.,

Meeuse 9849. Potgietersrus : Moord Drift, Norlindh & Weimarck 5241. Pretoria: Meintjies Kop,

Pole Evans 364; Hoorns Nek, Codd 769; Rietvlei Reserve, Repton 3227. Vereeniging: Burtt Davy

17194 (BOL). Waterberg: Sandrivierspoort, Codd 921 ; Towoomba, Acocks 8892. Witrivier: Numbi,

Van der Schijff 1605.

South West Africa. — Grootfontein : 30 miles N. of Gautscha Pan, Story 6472; Aha Mts.. Story

6387; between Cigarette and Tsammagaigai, Maguire 2509; Otavi, Dinter 5745. Otjiwarongo: Water-

berg, Volk 1077; Waterberg Plateau, De Winter 2791; 2805. Okavango: 8 miles E. of Tamso, De

Winter & Marais 4682; 4 miles S.W. of Nzinzi, De Winter & Wiss 4008; 5-8 miles E. of Schamvura

Camp, De Winter & Wiss 4435.

Rhodesia. — Bubi: Gwampa Forest Reserve, Goldsmith 19/56. Inyanga: Norlindh & Weimarck

4804. Salisbury: Eyies 1493; Brain 2443.

28

A. schirensis var. angustifolius is characterized by its conspicuously dorsiventral

racemes. On the one side the sessile spikelets with the joints and pedicels can be seen,

whereas the broad backs of the pedicelled spikelets with the dense cilia of the pedicels

between them are evident when viewed from the other side. The whole plant, especially

the racemes, is usually flushed with a reddish or purplish-brown colour.

Typical A. schirensis apparently does not occur in South Africa although several

specimens approach it very closely, in particular those from South West Africa. These

include: De Winter & Marais 4682, De Winter & Wiss 4008, Maguire 2509, Volk 1077,

Scheepers 1236, Van Vuuren 457 and Aeocks 8892. These specimens were compared

with tropical African material and it was found that the plants are smaller and less

robust than in typical A. schirensis and that the leaves, although comparatively wide,

are shorter.

A definite conclusion can be reached only after a study of a more extensive range

of African material has been undertaken.

16. A. gayanus Kunth var. squamulatus ( Hochst . ex A. Rich.) Stapf in Fl.Trop. Afr.

9: 263 (1919); Chippindall in Gr. & Pastures S. Afr. 1: 498 (1955). Type: Abyssinia,

Ferrfera Valley, Schimper 715 (K, iso.!; PRE, photo.!).

A. squamulatus Hochst. ex A. Rich., Tent. FI. Abyss. 2: 454 (1851); Hochst. in

Flora 27: 244 (1844), nom. nud. ; Steud., Syn. PI. Glum. 1: 374 (1855). A. gayanus

Kunth var. cordofanus Hack, in DC., Monogr. Phan. 6: 448 (1889), in part, excl.

Kotschy 54. A. helophilus Schum. in Pflanzenw. Ost-Afr. 5 : 98 (1895). Type: Usam-

bara, Hosigo, Holst 2535 (K, iso.!). A. hylophilus Schum. in Abhandl. Preuss. Akad.

Wiss. Berlin 38 (1894), nom. nud.

Perennial, tufted, glaucous. Culms up to 3-5 m long, stout, terete, glabrous,

smooth, shiny, many-noded with few to many flowering branches. Leaves glabrous

or softly pubescent, rarely villous or tomentose; blades up to 40 cm long and 18 mm

wide, glaucescent or ultimately reddish, expanded, linear to lanceolate-linear, the

lower portion usually much attenuated and often reduced to the midrib, widening

upwards and tapering to a fine apex, midrib prominent, margins scabrid; ligules

2-5-3 mm long, rounded or truncate, scarious, ciliolate; sheaths terete, striate, tight,

if wider than the base of the blade then forming more or less rounded shoulders or

only very shortly produced into auricles adnate to the ligule. Inflorescence of 2 racemes

appearing hairy from the long soft hairs on the joints and pedicels, one sessile, the

other with a glabrous base 7-10 mm long; spathes usually with well developed blades

except in the uppermost, glabrous. Racemes 6-10 cm long, more or less erect and

straight; joints 5-6 mm long, stout, cuneate-clavate, swollen upwards, densely ciliate

on both margins, cilia about 4 mm long, apex cupular, dentate; pedicels similar but

the cupular apex produced into a tooth on each side. Sessile spikelets hermaphrodite,

narrowly-oblong, pale green; callus obtuse, bearded at the base. Glumes equal;

lower 7-9 mm long, chartaceous, narrowly-oblong, scabrid or shortly ciliate on the

upper half of the keels, dorsally only shallowly grooved, 5-7-nerved on each side of

the groove, apex obtuse or minutely bifid; upper 7-9 mm long, boat-shaped, subchar-

taceous, scabrid on the upper half of the keel, ciliate on the margins, 3-nerved with the

midnerve distinct, the lateral shorter and finer, apex obtuse oi minutely 3-dentate.

Lower floret reduced to a linear-oblong lemma, 7-8 mm long, membranous, hyaline,

2-nerved, margins pilose, apex acute or truncate and ciliolate. Upper floret with the

lemma 6-7 mm long, membranous, hyaline, 1 -nerved, oblong, 2-lobed, lobes acute,

awned from between the lobes, awn 2-2-5 cm long, kneed at about the middle,

scaberulous; palea 3-7-4 mm long, oblong, membranous, hyaline, 2-nerved, obtuse

or minutely 2-dentate; anthers 4 mm long. Pedicelled spikelets male, linear-oblong,

29

scaberulous, pale-green. Glumes equal; lower 7-9 mm long, chartaceous, linear-

oblong, dorsally flattened, keeled, scabrid on the keels, many-nerved, the midnerve

distinct and produced into a straight, scaberulous awn 4-6 mm long, apex minutely

bifid or acute; upper 7-9 mm long, narrowly-oblong, membranous, 3-5-nerved, margins

shortly pilose, apex acute, acuminate or mucronulate. Lower floret reduced to the

lemma, 7-8 mm long, membranous, hyaline, 2-3-nerved, linear-oblong, margins shortly

pilose, apex obtuse or acuminate. Upper floret with the lemma 6-7 mm long, mem-

branous, hyaline, 1-nerved, glabrous, apex minutely 2-dentate or mucronate; palea

2-7-3 mm long, more or less oblong, membranous, hyaline, ciliolate on the margins,

apex obtuse; anthers 4 mm long.

A. gayanus var. squamulatus has been recorded from Abyssinia, the Sudan, Nigeria,

Ghana, the Congo Republic, Uganda, Tanzania, Zambia, Rhodesia and Mozambique.

In South Africa it is confined to the northern parts of Natal and Transvaal, Swaziland,

Bechuanaland and northern South West Africa. It usually grows in open grassland

in rocky situations and almost invariably in sandy soil.

Natal. — Hlabisa: W. of Charters Creek, Ward 2859; False Bay Park, Ward 4151. Ubombo:

Strey & Huntley 5011; Mkuzi Game Reserve, Ward 3996.

Swaziland. — Stegi: Malinda Hills, Compton 29790.

Transvaal. — Letaba: Duiwelskloof, Scheepers 639; between Komatipoort and Letaba River,

Rogers 2657. Pietersburg: Leipzig, Schweickerdt 1822. Pilgrim’s Rest: Orpen Dam, Van der Schijff

3473. Sibasa: Punda Milia, Van der Schijff 1826; Acocks 16765; Codd 4212; 5374; Godfrey H. 1715.

Soutpansberg: 5 miles N. of Louis Trichardt, Van Vuuren 1683.

Bechuanaland. — Francistown, Makarikari Pan, Pole Evans 3269; 3303; between Totume and

Bushman Mine, Pole Evans 3267 ; Lower Ngwezimba River, Miller s.n.; Ngamiland, Tsodito Hills,

Banks 45.

South West Africa. — Grootfontein : Aha Mtns., Story 6337; Gemsboklaagte, Schweickerdt 2064;

Otjihenena, Volk V. 68. Otjiwarongo: Bradfield 376; Otjiwarongo-Grootfontein Road, Liebenberg

4859; Waterberg, Volk 423; 999; 1078; Liebenberg 4802. Okavango: Andara Mission Stn., De

Winter & Marais 4827.

Angola. — Huila: Teixaira 55 3; 3459; 3575.

Rhodesia. — Hartley: Makwiro, Mainwaring s.n. (Eyles No. 2624). Marandellas: Eyles 4015.

Matobo: West 2158. Salisbury: Sturgeon 57588. Shangani: Goldsmith 48928.

Mozambique. — Cabo Delgado: 5 km from Mocamboa da Praia, Gomes e Sousa 4693. Lourenco

Marques: Manica, De Lernos & Macuacua 92. Niassa: Maudimba, Hornby 3340.

Zambia. — Chilinga: Sandwith 70. Fort Jameson: Senegallia farm, Grout 124.

Andropogon gayanus var. squamulatus differs from the other South African species

of Andropogon in being more robust with broader leaves. Like A. schinzii the culms

are usually branched, producing few to many flowering branches. Three well defined

varieties of A. gayanus are upheld by Stapf in the Flora of Tropical Africa (1919), of

which only the var. squamulatus is known to occur in South Africa. This variety differs

from typical A. gayanus in that the pedicelled spikelets are scaberulous with the joints

and pedicels ciliate on both margins.

17. A. distachyos L., Sp. PI. 1046 (1753); Steud., Syn. PI. Glum. 1: 372 (1855);

Hack, in DC., Monogr. Phan. 6: 461 (1889); Stapf in FI. Cap. 7: 343 (1898); FI.

Trop. Afr. 9: 218 (1919); Chippindall in Gr. & Pastures S. Afr. 1 : 496 (1955). Type:

Europe, Helvetia, Linn. Herb. 1211.10 (PRE, photo.!).

Perennial, caespitose with a short oblique rhizome. Culms up to 120 cm long,

erect, slender, terete, simple or branched towards the base, glabrous, 3-7-noded, lower

nodes often hairy, basal internodes usually naked. Leaves 5-30 cm long, 1-4 mm,

rarely 5 mm wide, expanded, linear, tapering to a long and sometimes setaceous apex,

rigid to flaccid, hairy or glabrous but usually with pubescent basal shoots, often glabre-

scent, more or less rough above and scabrid on the margins; ligules 2 mm long, mem-

branous with a fimbriate rim, truncate, sometimes pilose from behind; sheaths glabrous

30

or the lower hairy. Inflorescence of 2, rarely 3 racemes, the one sessile, the other shortly

peduncled, rather robust, straight or curved, greenish, often tinged with a dull or livid

purple, 5-10 cm long; joints cuneate-linear, hollow and very slightly inflated, 3-5-4 mm

long, semi-transparent along the middle, glabrous to subpubescent on the back, ciliate

on the outer margin, cilia 1-2 mm long, tips slightly cupular; pedicels similar, slightly

longer and more translucent. Sessile spikelets hermaphrodite, 9-12 mm long, broad-

lanceolate to acuminate, pale-green, often purplish towards the apex; callus short,

obtuse, shortly bearded. Glumes unequal; lower subherbaceous or chartaceous,

dorsally flattened, glabrous or puberulous, keels broadly winged, wings membranous,

whitish or purplish, intracarinal nerves 7-11, partly evancescent below; upper distinctly

shorter than the lower, boat-shaped, rigidly membranous, 3-nerved, softly ciliate,

tips minutely 2-dentate with a bristle 4-6 mm long. Lower floret reduced to the lemma,

5-7 mm long, oblong, acute or subacute, 2-nerved. Upper floret with the lemma 5-7

mm long, slightly gibbous on the back, 2-lobed to below the middle, chartaceous and

3-nerved below, lobes lanceolate, glabrous, awned from between the lobes, awn slender,

about 2-5 cm long, kneed at and twisted below the middle; palea minute or absent;

anthers 3 mm long; grain oblong, 2 mm long. Pedicelled spikelets similar to the

sessile but narrower. Glumes unequal, lower 9-1 1 mm long, subherbaceous to char-

taceous, very narrowly winged and terminated by a bristle 4-6 mm long; upper mem-

branous, 3-nerved, shortly aristulate. Lower floret with the lemma 5-7 mm long,

delicately hyaline, ciliolate, faintly 2-nerved. Upper floret with the lemma 4-5 mm

long, delicately hyaline, glabrous, 1 -nerved, apex slightly notched.

This species occurs in southern Europe, throughout the Mediterranean region,

tropical Africa and southern Arabia. In South Africa it has been recorded from

mountain slopes in the Drakensberg areas of Natal, the Orange Free State and the

northern Transvaal. It usually grows in rocky situations in open grassland, often

preferring wet places.

Orange Free State. — Harrismith: Wittebergen, Buchanan 263 (BOL). Vrede: Quagga Kop,

Acocks 12757.

Natal. — Bergville: Cathedral Peak, Chippindall 355. Estcourt: Cathkin Peak, Edwards 2006;

Mountain hostel, West 145; Ntabamhlope, A cocks 1 1460; 11480. Kliprivier: Codings Pass, Acocks

13487. Underberg: McClean 601.

Transvaal. — Pilgrim's Rest: Mariepskop, Van der Schijff 5583.

Linnaeus originally described this species as A. distachyon. The generic name

Andropogon is, however, masculine and, if Recommendation 75A of the International

Code of Botanical Nomenclature (1961) is taken into consideration, the specific name

should be changed to distachyos.

A. distachyos is readily distinguished from the other South African species by

the broadly winged lower glumes of the sessile spikelets. I have not seen type material

of this species but the South African material was carefully compared with the

description of the species. Furthermore, Chippindall 355 was compared with European

specimens at Kew by Dr. B. de Winter who states that it compares well except that

the South African specimen is slightly larger. I have no doubt that the South African

material is indeed A. distachyos L.

Bews s.n., collected at Himeville in the Underberg district, proved to be of interest.

This specimen, deposited in the herbarium of the Natal University (NU 3539), is

vegetatively similar to A. schirensis var. angustifolius. Even the colouring of the leaves

and racemes is typically reddish-brown. Examination of the spikelets revealed, however,

that the lower glumes of the sessile spikelets are dorsally flattened and broadly winged

as in A. distachyos. Except for minor details the spikelets are typical of this species.

As both A. schirensis var. angustifolius and A. distachyos occur in this area, this specimen

is most probably a hybrid between these two species.

31

Bothalia 9, 1: 31-96.

The Genus Asparagus in Southern Africa*

by

J. P. Jessop

Abstract

A revision of the genus Asparagus in South Africa, South West Africa, Bechuanaland, Basutoland,

and Swaziland has been undertaken.

Notes are given on the value of most of the characters which have been used in the separation

of species, recent literature on the nature of Asparagus assimilatory organs, and a technique for the

examination of chromosomes. Chromosome counts are listed for ten taxa.

There is a key to the forty species and four varieties. In the main part of the work these taxa

are described, and their synonymy, taxonomy, distribution and habitats dealt with. Six species and

one variety are new.

The following are the new species and combinations: A. setaceus (Kunth) (Asparagopsis setacea

Kunth), A. mucronatus, A. macowanii Bak. var. zuluensis (N. E. Br.) (A. zuluensis N. E. Br.), A. rigidus,

A. densiflorus (Kunth) (Asparagopsis densiflora Kunth), A. aethiopicus L. var. angusticladus, A. falcatus

L. var. ternifolius (Bak.) (A. aethiopicus L. var. ternifolius Bak.), A. aspergillus, A. obermeyerae, A.

krebsianus (Kunth) (Asparagopsis krebsiana Kunth), A. acocksii, A. crassicladus.

Several plants of horticultural importance occur in South Africa. The three best known are

A. plumosus, which is reduced here to synonymy under A. setaceus (Kunth) Jessop, and A. sprengeri

and A. myersii. A. sprengeri is being reduced to synonymy under A. densiflorus (Kunth) Jessop,. A.

myersii, which is a nomen nudum, is also regarded as belonging to A. densiflorus.

Introduction

The earliest record of a South African Asparagus, which can be authenticated,

is that of an artist, probably Hendrik Claudius, who accompanied Governor Simon

van der Stel on an expedition to Namaqualand during the second half of 1685 and

the beginning of 1686. One of the illustrations which he made was of A. capensis L.

Subsequently several other species were described, and some of them illustrated, but

by 1753 Linnaeus could include only five South African species of Asparagus in his

Species Plantarum. Thunberg, as a result of his own collecting, brought this number

up to 13 in his Prodromus. The next major contribution was that of Roemer and

Schultes, Systema Vegetabilium (1829), in which nearly thirty species were mentioned.

Many new names, many of them superfluous, were added by Kunth in 1850 when he

divided the genus as it is defined at the present into three genera. He referred seven

of his species to Myrsiphyllum, which consisted of plants with solitary, flattened cladodes

and axillary flowers. The remaining species were divided into Asparagus, with

supposedly dioecious flowers and eight South African species, and Asparagopsis, with

23 South African species.

Baker, whose monograph was published in 1875, provided the most recent complete

work on the genus. In this monograph. Baker recognised 40 species from South Africa

and, 21 years later, in the Flora Capensis, he included an additional four.

Since Baker, several authors have added an occasional name, based especially

on South West African material. The most important work has been that of Schinz

on South West African plants in 1896, Salter, who revised the genus for the Flora

of the Cape Peninsula, and Kies, who revised part of the section Africani in 1951.

Phillips (1951) estimated that the total number of South African species was about 70.

* Based on an M.Sc. thesis, submitted to the University of Cape Town in 1964

7230691-2

32

Partly as a result of these additional species, and in the absence of a revision of

the whole genus, or even the South African species, the available classification was

found to be in considerable need of revision. Salter stressed this need in the Flora

of the Cape Peninsula in 1950, and an examination of such works as Wilman’s Preli-

minary Check List of the Flowering Plants and Ferns of Griqualand West shows a

large number of unidentified specimens.

It was, therefore, decided to undertake a complete revision of the genus in South

Africa, accompanied where possible by field work. Most of this work was undertaken

while the author was a student at Cape Town University. The assistance of many

members of the staff of both the Bolus herbarium and the University of Cape Town

Botany Department is gratefully acknowledged. A special debt is owed to Dr. E. A.

Schelpe who supervised the work and provided much valuable advice. The revision

was completed at the National Flerbarium, Pretoria, where several members of the

staff provided very useful advice. In particular Mrs. A. A. Mauve’s knowledge of

the Transvaal species has been a great help. Acknowledgements are due to the

University of Cape Town Botany Department for the photographs of Asparagus

mucronatus and A. crassicladus. The remaining illustrations were prepared at the

Botanical Research Institute, Pretoria.

Specimens have been loaned, or other assistance has been provided, by the directors

and curators of the following herbaria : The Albany Museum Herbarium; the Herbarium,

Berlin-Dahlem ; The British Museum; the Herbarium of Trinity College, Dublin; the

Royal Botanic Gardens, Kew; the Herbarium of the University of Kiel; the National

Botanic Gardens and South African Museum herbaria at Kirstenbosch ; Rijksherbarium,

Leyden; the Botanische Staatssammlung, Munich; the Natal Herbarium; the National

Museum of Natural History, Paris; the Stellenbosch Herbarium; the Botanic Gardens

and Museum, Uppsala; the Windhoek Herbarium; and the Botanische Garten und

Museum der Universitat, Zurich.

In the distribution lists all specimens cited have been examined by the author.

In the larger species not all specimens seen have been cited, but a specimen from each

magisterial district has been selected for citing. When no herbarium abbreviation

follows the citation of a specimen, a specimen of that collecting is in the National

Herbarium, Pretoria. When there is no specimen in the National Herbarium only

one herbarium is cited for each collecting although duplicates may have been seen

from other herbaria.

Taxonomic Characters

Up to the present, there has been no satisfactory delimitation of the species. This

is partly because no-one has undertaken a major revision of the group since the Flora

Capensis in 1896, and partly because, in the absence of diagnostic floral characters,

most of the species must be defined on vegetative characters. These are often extremely

susceptible to environmental factors, and this is aggravated by the wide distribution

of many of the species.

A list of the characters of use in defining taxa below generic level is given here,

with notes on the general value of the different characters.

(a) Perianth segments

All have six, fairly small (1-6 mm long), white or near white tepals. The members

of the inner whorl are sometimes slightly broader than the outer segments, and the

distal margins are sometimes slightly ciliate. These characters are frequently variable

within a single species. A few species are characterised by having the distal half of

the tepals reflexed.

33

{b) Stamens

The six stamens sometimes have a pair of minute lateral spurs at the bases of their

filaments. This character was first used in Salter’s treatment in the Flora of the Cape

Peninsula, and is probably of value. The anthers vary from greenish through yellow

to orange and black. Work on fresh material suggests that colour is useful, but on

dried specimens the colour can usually not be determined.

(c) Fruit

The fruit is generally a wrinkled, fleshy berry with two or three seeds, and globose

in shape, but dry, smooth fruits as in A. exuvialis, and single-seeded berries as in

A. scandens, occur. One species, A. crispus, has an ovoid berry, and A. glaueus has

a persistent perianth enclosing the fruit. The fruit characters are usually reliable at

specific level.

( d ) Peduncles

All species have flowers borne solitarily on the ultimate flowering stalks. In

A. stipulaceus and A. capensis these stalks are reduced to discs. As all leaves are reduced

to scales, it is not strictly correct to refer to umbels or racemes. By treating the cladodes

as leaves, the use of the conventional terminology will be of value. The term peduncle

will be used for the flower stalk except in the section Racemosi, where the term pedicel

will be used for the final flower stalks and peduncle for the main axis of the inflorescence.

Inflorescences (termed racemes here) consisting of branches bearing only a series of

bracts and pedicels occur. However, in species producing racemes, cladodes some-

times appear at the bases of the pedicels. In two species, A. minutiflorus and A. rigidus,

the inflorescence, which appears to be a reduced raceme, consists of a peduncle, one to

three terminal pedicels, and sometimes one lateral pedicel. In some species both

terminal and axillary flowers occur as in A. africanus, but in others terminal, as in

A. setaceus, or axillary peduncles, as in A. scandens, are diagnostic. The umbels,

referred to by Baker, are clusters of terminal flowers, and are rarely found as the only

type of inflorescence on a plant. However, exceptions to this occur in the sections

Striati and Capenses. The peduncles may be solitary, paired or numerous, and these

characters are sometimes useful at specific level. In all species the peduncles are

articulated and in A. nelsii sometimes twice.

(e) Roots

The roots may be tuberous or fibrous. Classification cannot usually be based

on further subdivision of tuber type as the tubers are particularly susceptible to

environmental conditions. This is discussed further under A. asparagoides. In many

species there is a single root-stock bearing lateral tubers, but in most species of the

section Racemosi there are several roots each bearing lateral tubers. This condition

is preceded in young plants by the direct attachment of the tubers to the root-stock.

(/) Stems

Roots and aerial stems are produced from perennial rhizomes in all species. Annual

and perennial aerial stems occur, but the duration of the life of the aerial stem may

vary under different conditions. In A. exuvialis the aerial stem is perennial, but the

cladodes are deciduous, and after a month or two are replaced. Some species are

predominantly climbers and others erect. In each of these groups exceptions may be

produced by the effect of the environment, particularly the presence or absence of

support and the amount of light reaching the plant. The aerial stem may have a smooth,

wrinkled (Fig. lb), grooved (Fig. la), ridged (Fig. lc) or hispid surface. These characters

are sometimes useful. In A. exuvialis the outer layers of the aerial stems and branches

peel. (Fig. 2d). In several species, including A. compactus and A. retrofractus, the

grooved white outer layers disintegrate, leaving the stem smooth and brown. Another

character of occasional value is aerial stem colour, as in A. denudatus, which has a green

aerial stem.

34

c, A. aethiopicus var. aethiopicus, ridged.

fiteiPR vt.MWwe,

35

( g ) Branchlets

Branchlets, which are ultimate, specialised branches bearing cladodes and flowers,

occur in some species, especially in the section Capenses, where they are fascicled.

(Fig. 2b).

(h) Cladodes

Up to the beginning of the nineteenth century, the photosynthetic organs of

Asparagus were regarded as leaves. By the middle of the twentieth century, however,

it had come to be accepted by most botanists that they are modified branches. Several

conflicting theories exist, and these need to be considered. The three most important

theories propounded this century are those of Arber (1924 & 1925), Schlittler (1953

& 1959) and Kaussmann (1955).

Arber divided the genus Asparagus (sensu lato) into Myrsiphyllum (containing

A. asparagoides and A. undulatus ), in which she considered that the assimilatory organs

were leaves, and Asparagus in which she considered them to be branches. She cut

serial sections through the plumules of A. asparagoides and found what she believed

to be the axillary shoot of which the assimilatory organ was the prophyll. In Asparagus

she, and later Kaussmann, found what they took to be reduced leaves borne on the

organs, which they therefore regarded as cladodes.

While Kaussmann agreed with Arber about the nature of the organs of Asparagus

sensu Arber, and supported the view with developmental investigations, he disagreed

with her about the nature of the organs of A. asparagoides. His ontogenetic work

suggested that these were also cladodes, and not true leaves.

Schlittler took the opposite view to that of Kaussmann and, basing his evidence

on a criticism of other workers’ conclusions, and by comparing the assimilatory organs

of what he believed to be related plants, arrived at the conclusion that they were leaves

in all species.

While no satisfactory conclusion to this controversy has yet been reached, it has

been decided that the term cladode will be used throughout this revision. The present

author feels that the weight of evidence is on the side of this view, and believes this to

be the more generally accepted point of view.

The cladodes may be single, 3-nate or variable. These three characters are

generally reliable. In many species the cladodes are flattened. In most, however,

they are terete or angled, but several species are variable in this character, such as

A. juniperoides. All species have a single vein except the section Myrsiphyllum and

A. striatus. The cladodes are sometimes scabrid, but this is rarely a reliable feature.

(i) Leaves

Leaves are always reduced. Sometimes they are firm, as in A. exuvialis, but usually

with a more or less free, papery and scarious part. (Figs. 2c & d).

(j) Spines

Spines are formed from branches in the section Capenses, one in A. glaucus, three

or more in the others. (Fig. 2b). In all other species where spines are formed they are

formed from reduced leaves. Many species, which usually have spines, have individuals

without them. The spines may be straight or recurved, spreading or reflexed, but

these characters are generally rather variable. In species with spines, the presence

or absence of spines on the ultimate branches is often significant.

36

F.c 2— Diagnostic characters: a, spines formed from modified branches showing aborted nodes;

F,°' 2' “ Sded spines formed ftom modified branches lacking .aborted nodes: “f no!

Hifferentiated and fascicled; c, spine formed from a modified leaf with the non spino

portion chartaceous; d, spines formed from a modified leaf with the non-spinous por 10

hard, deltoid and entire; outer layers of the stem peeling in strips.

37

Subgeneric Groups

Other than Kunth (1850), the only botanists before Baker (1875) who subdivided

the genus, did so on the presence or absence of spines — in many species one of the

most variable characters. Kunth recognised three genera: Asparagus , Asparagopsis

and Myrsiphyllum. Myrsiphyllum included the herbaceous species with flattened

cladodes. Asparagus was separated from Asparagopsis in, supposedly, being dioecious.

Although in A. officinalis L. this is undoubtedly the case, for most of the species he

placed in Asparagus his information was faulty. The species within each genus

recognised by Kunth do not show a particularly large number of characters in common

while, in several cases, undoubtedly similar species were placed in different genera.

As none of the South African species which Kunth placed in his construction of

Asparagus is dioecious, his use of this character to separate Asparagus from Asparagopsis

was incorrectly applied to the South African species. His genus Myrsiphyllum was

retained by Baker (1875 and 1896) as a subgeneric group, and will be treated as a

section in this work.

Baker (1896) included all species in one genus, which he divided into nine sections.

The characters he used for separating these sections were: the presence or absence

of spines; the shape of the cladodes, and the number of these in a cluster; the number

of flowers in a cluster; whether or not the branchlets were fascicled; and the type of

inflorescence.

In each of his sections there are closely similar species, but also many with little

in common. His section Myrsiphyllum has been retained here with a slight change,

and the section Capenses includes all the species he included. While it would be unwise

to propose a revision of the sections of Asparagus with only a knowledge of the genus

in Southern Africa, the following sections appear to reflect the natural grouping of

the species in South Africa. In each, the more important characters are printed in

italics. The species which would be included in each are indicated by their numbers

in this work.

1. Capenses

Roots not tuberous. Aerial stems perennial, woody. Branchlets fascicled.

Cladodes fascicled, terete. Spines axial in origin. Peduncles usually terminal. Anther

filaments not spurred. Berries globose, fleshy or dry.

This section contains the same plants as Baker's section Capenses.

Species: 1-4.

2. African i

Roots not tuberous. Aerial stems usually perennial. Branchlets, when present,

usually solitary. Cladodes fascicled, terete. Spines frequent, foliar in origin. Peduncles

axillary or terminal. Anther filaments not spurred except in A. fasciculatus. Berries

globose, fleshy.

This section has no outstanding characters, but is identifiable by a combination

of root, spine, cladode and inflorescence characters. It contains species from Baker’s

sections Declinati, Umbellati, Thunbergi and Africani.

Species: 5-16.

3. Exuviali

Roots tuberous. Aerial stems perennial , woody. Cladodes fascicled, terete. Spines

foliar in origin, usually small and blunt. Peduncles axillary. Anther filaments not

spurred. Fruit dry and smooth.

A monotypic section containing A. exuvialis, which Baker included in his section

Declinati.

Species: 17.

38

4. Race mo si

Roots tuberous. Aerial stems woody or softly woody. Branchlets often present.

Cladodes fascicled or occasionally solitary, flattened or subulate, with one vein. Spines

often present, foliar in origin. Flowers generally borne in racemes. Anther filaments

not spurred. Fruit a berry.

This section includes Baker’s sections Racemosi, Falcati and A. juniperoides from

his section Myrsiphyllum.

Species: 18-33.

5. Striati

Roots tuberous. Aerial stems perennial. Branchlets absent. Cladodes sometimes

solitary, flattened to terete, firm, with one or several veins. Spines absent or poorly

developed. Peduncles terminal. Anther filaments not spurred.

This section contains part of Baker's sections Striati and Umbellati.

Species: 34 and 35.

6. Sc an dentes

Tubers present. Aerial stems somewhat herbaceous, perennial. Branchlets absent.

Cladodes ternate, flattened, with one vein. Spines absent. Peduncles axillary. Anther

filaments not spurred. Fruit globose, fleshy, with one seed.

This section contains two species which Baker included in his section Striati.

Species: 36 and 37.

7. Crisp i

Tubers present. Aerial stems annual. Branchlets absent. Cladodes ternate,

terete or flattened. Spines absent. Peduncles axillary. Anther filaments spurred.

Fruit ovoid, fleshy, many-seeded.

This section is monotypic, containing A. crispus, which Baker included in his

section Declinati.

Species: 38.

8. Myrsiphyllum

Tubers present. Aerial stems annual. Cladodes flattened, with more than one

vein, solitary. Spines absent. Peduncles axillary. Anther filaments spurred. Fruit

fleshy, globose.

This section is the same as Baker’s, except that A. juniper oides has been removed.

Species: 39 and 40.

Cytological Investigations

While a thorough investigation of chromosome numbers could not be undertaken,

a technique has been used which has provided counts for ten plants.

It was found that young shoots were as satisfactory as either germinating seed

radicles or roots, and that they are more readily obtainable. Pollen Mother Cells

are extremely difficult to get at the right stage as they develop when the buds cannot

easily be recognised.

The following procedure was developed:

39

Shoot tips were placed in 0-01 per cent aqueous colchicine solution for three to

four hours, before being fixed in Acetic Alcohol (1:3) for twenty to twenty four hours.

They were then hydrolised in 20 per cent HC1 at 63 degrees centigrade for three to

eight minutes, depending on the thickness of the material. Two changes of distilled

water for at least five minutes each were used before the shoot tips were squashed

in 2 per cent Aceto-Orcein.

The following numbers were obtained: —

Species No.

A . capensis 40

A . stipulaceus 80 ?

A. fasciculatus 20

A . thunbergianus 20

A . retrofr actus 20

A. densiflorus (sprengeri form) 60

A . krebsianus 20

A. asparagoides (broad cladode form) 20

A. asparagoides (narrow cladode form) 20

A. asparagoides (pubescent form) 20

ASPARAGUS

Asparagus L. , Sp. PI. 313 (1753); Thunb., Prodr. 65 (1794): FI. Cap. ed. Schult. 332

(1823); Bresler, Diss. (1823); R. & S., Syst. Veg. 7: 317 (1829); Kunth, Enum. PI.

5: 57 (1850); Bak. in Journ. Linn. Soc. Bot. 14: 594 (1875); Benth. & Hook, f.,

Gen. PI. 3: 765 (1883); Bak. in FI. Cap. 6: 256(1896); FI. Trop. Afr. 7: 425 (1898);

Marloth, FI. S. Afr. 4: 83 (1915); Krause in Pflanzenfam. Auf. 2, 15a: 362 (1930);

Salter in FI. Cape Penins. 172 (1950); Hutch., Fam. Flow. PI. 2: 608 (1959); Solch,

Beitr. FI. Siidwest-Afr. 31 (1961). Type species: A. officinalis L.

Medeola L., Sp. PI. 339 (1753), partly, as to M. asparagoides L.

Myrsiphyllum Willd. in Ges. Naturf. Fr. Berl. Mag. 2: 25 (1808); Kunth, Enum.

PI. 5: 105 (1850). Type species: M. asparagoides (L.) Willd.

Asparagopsis Kunth in Abh. Akad. Berl. 35 (1842); Enum. PI. 5: 76 (1850). Type

species: A. albus (L.) Kunth.

Hecatris Salisb., Gen. PI. Fragm. 66 (1866). Type species: H. asparagoides (L.)

Salisb.

Roots often tuberous, but frequently fibrous. Stems erect or climbing, annual

or more often perennial and woody, often much-branched; final branches sometimes

fascicled and morphologically distinct from the main branches. Cladodes solitary or

fascicled, with one or more veins. Leaves greatly reduced. Flowers borne singly on

the final flowering stalks, which may be arranged terminally, in the axils of cladodes,

or on specialised branches lacking cladodes; the final flowering stalk always articulated.

Perianth segments six, free or nearly so, more or less similar, ascending or spreading,

up to 6 mm long, white or nearly so. Stamens six, fused to the perianth segments

towards their base. Ovary of three fused carpels containing three chambers; ovules

several, axile; style with three short branches or divided nearly to the base. Berry

1 — several-seeded, globose, or rarely ovoid, usually red, less often black or brown.

Distributed throughout Africa and in most of Europe, Asia and Australia. 40

species recognised in Southern Africa.

40

1 Cladodes conspicuously flattened:

2 Cladodes more than one-veined:

3 Flowers axillary:

4 Stem smooth, usually climbing; tubers sessile or distant 39. A. asparagoides

4a Stem seabird and often with longitudinal wings, erect; tubers distant ... . 40. A. undulatus

3a Flowers terminal 34. A. striatus

2a Cladodes one-veined:

5 Plants herbaceous, twining; flowers axillary or terminal:

6 Cladodes all in one plane 36. A. scandens

6a Cladodes not all in one plane 37. A. ramosissimus

5a Plants woody or, if rather softly woody, then erect; flowers usually racemose:

7 Cladodes solitary:

8 Stems pubescent 18. A. wcyacanthus

8a Stems glabrous:

9 Spur of the leaves spine-shaped; tepals and cladodes without a ciliate margin;

stems usually well lignified; from the southern Cape and the eastern parts

of South Africa 22. A. densiflorus

9a Spur of the leaves not spine-shaped; tepals and usually the cladodes with a ciliate

margin; stems softly woody ; from the western Cape and S.W. A. 19. A. juniperoides

7a Cladodes several at a node:

10 Stems erect 22. A. densiflorus

10a Stems climbing:

1 1 Cladodes more than 35 mm long 26. A. falcatus var. falcatus

11a Cladodes less than 35 mm long:

12 Racemes branched; cladodes usually very dark after drying 26. A. falcatus var.

ternifolius

12a Racemes simple; cladodes not usually darkening on drying:

13 Branches grooved; from the southern and south-west Cape 25. A. aethiopicus

var. aethiopicus

13a Branches smooth; from the eastern Cape and the eastern and northern parts

of South Africa 25. A. aethiopicus var. angusticladus

la Cladodes terete, angled, or absent during the flowering period:

14 Flowers in racemes:

15 Cladodes solitary:

16 Spur of the leaves spine-shaped; tepals and cladodes without a ciliate margin; stems

usually well-lignified; from the southern Cape and the eastern parts of South

Africa 22. A. densiflorus

1 6a Spur of the leaves not spine-shaped ; tepals and usually the cladodes with a ciliate margin ;

stems softly woody; from the southern Cape and S.W. A 19. A. juniperoides

15a Cladodes several at a node, or absent during the flowering period:

17 Cladodes terete or oval in section, 0-8 mm or more thick 32. A. crassicladus

17a Cladodes angled or, if terete, then less than 0-8 mm thick, or absent during the flowering

period:

18 Branches, at least the younger, densely pubescent, scabrid or tubercled; peduncles

grey to brown; flowers white:

19 Branches ridged or grooved; cladodes often absent during the flowering period

27. A. asperigillus

31. A. acock sii

19a Branches smooth

41

18a Branches glabrous or, if slightly pubescent, then the stems erect and the flowers

yellowish or greenish yellow:

20 Stems erect :

21 Final branches grooved 22. A. densiflorus

21a Final branches smooth:

22 Flowers white; from the eastern parts of South Africa.... 23. A. saundersiae

22a Flowers yellowish or greenish yellow; mainly from the western and northern

Cape, Bechuanaland and S.W.A., but recorded from the northern Trans-

vaal 33. A. nelsii

20a Stems twining:

23 Spines present on all branches; branchlets never well differentiated 30. A. krebsianus

23a Spines absent from ultimate branches; branchlets sometimes well differentiated:

24 Stems usually grooved; branchlets differentiated; cladodes more than \ mm

thick, angled; perianth segments 2-5-3 -0 mm long 28. A. racemosus

24a Stems smooth: branchlets not generally differentiated; cladodes very narrow,

less than \ mm thick; perianth segments usually 1 -5-2-0 mm long

24. A. buchananii

14a Flowers not in racemes:

25 Branches spine-tipped; spines formed from modified branches, usually at least 3-nate:

26 Spines solitary; perianth enclosing the fruit 4. A. glaucus

26a Spines rarely solitary; perianth not enclosing the fruit:

27 Flowers sessile; spines completely smooth:

28 Average cladode length less than 5-5 mm; plant widely branched:

29 Cladodes at least 1-5 mm long; branches and stems generally fairly slender;

from the southern and western Cape, and S.W.A. away from the coast

1. A. capensis var. capensis

29a Cladodes less than 1 mm long; branches and stems more robust; from the west

coast 1 . A. capensis var. lit oralis

28a Average cladode length more than 5-5 mm; plants of a “fox-tail” form

2. A. stipulaceus

27a Flowers pedunculate; spines bearing nodes, which are usually reduced to minute

protuberances (Fig. 2a) 3. A. suaveolens

25a Branches not spine-tipped; spines formed from modified leaves, solitary, or absent:

30 Stem rigid; flowers mainly terminally 1-3-nate on peduncles, but sometimes also laterally

solitary on the peduncles:

31 Cladodes more than 0-4 mm broad; spines 4 mm or less long; stems glabrous

21. A. rigidus

31a Cladodes less than 0-3 nm broad; the longest spines at least 5 nm long; stems

usually more or less scabrid or pubescent 20. A. minutiflorus

30a Stems variable; flowers axillary or terminal on normal branches:

32 Cladodes three at a node except at nodes where there is a branch and then only two:

33 Cladodes 1 mm or more broad; fruit globose.... 37. A. ramosissimus

33a Cladodes less than 1 mm broad; fruit ovoid 38. A. crispus

32a Cladodes solitary, paired or numerous at a node, or absent during the flowering

period:

34 Flowers all terminal:

35 Stems smooth, climbing, repeatedly branched; cladodes and ultimate branches

all in one plane 6. A. setaceus

35a Stems grooved, erect or climbing to 2 m or rarely more; final branches and

cladodes not all in one plane.

36 Branches usually reflexed; cladodes laterally usually several; plant more or

less herbaceous, and often climbing or scrambling 35. A. subulatus

42

36a Branches usually ascending; cladodes laterally usually solitary; plant more

woody and robust, always erect 34. A. striatus

34a Flowers all, or at least some, axillary:

37 Cladodes solitary 19. A. juniperoides

37a Cladodes not solitary:

38 Young branches never both white and grooved:

39 Usually climbers or scramblers; stems tortuous:

40 Aerial stems annual, softly woody; peduncles articulated at the base of

the flower 9. A. fascicidatus

40a Aerial stems perennial, rarely annual; peduncle not articulated at the

base of the flowers:

41 Stems smooth or regularly grooved, sometimes lacking cladodes towards

the ends of the branches; spines often well-developed; colour of

stems variable 5. A. africanus

41a Stems irregularly, longitudinally wrinkled; cladodes always evenly

distributed along the entire length of the branches, or completely

absent; spines absent or poorly developed; stems green to greyish

green 7. A. denudatus

39a Stems erect, straight or zigzagging with straight internodes:

42 Outer layers of stems and branches peeling; leaf firm and entire (Fig. 2d)

17. A. exuvialis.

42a Outer layers of stems and branches not peeling; leaf usually with a char-

taceous free portion (Fig. 2c):

43 Plant completely glabrous:

44 Spines absent or poorly developed:

45 Branches green :

46 Stem zigzagging: branches more or less spreading. . 7 . A. denudatus

46a Stem straight; branches virgate 8. A. virgatus

45a Branches not green:

47 Plant up to 1 m high; from the southern Cape 12. A. macowanii

var. macowanii

47a Plant over 1^ nr high: widespread, but not in the southern Cape

12. A. macowanii var. zuluensis

44a Spines present and pungent, at least on the stems and main branches

10. A. thunbergianus

43a Ultimate branches pubescent 11. A. mucronatus

38a Young branches and often all the aerial stems and branches white and grooved:

48 Stems bearing fascicled, simple branches or less often secondary branches

present 29. A. obermeyerae

48a Axes repeatedly branched; branches solitary or less often 2- or 3-nate:

49 Spines present on all ultimate branches:

50 Cladodes ascending, straight; plant over 1 nr high, bushy 15. A. laricinus

50a Cladodes spreading, often arcuate; stems more or less erect, less than

1 m high 14. A. compactus

49a Spines absent from ultimate branches:

51 Cladodes 4 mm or less long 16. A. microrhaphis

51a Cladodes 5 mm or more long:

52 Branches zigzagging; stems often scrambling or climbing; from the

western half of South Africa 13. A. retrofractus

52a Branches straight; stems e ect; from the wetter parts of the southern

Cape and the eastern half of South Africa:

53 Stems up to 1 m high; from the Cape 12. A. macowanii var. macowanii

53a Stems over 1J2 m high; from Natal and the Transvaal 12. A. macowanii

var. ziduensis

43

1. A. capensis L., Sp. PL 314 (1753). Type: Cape of Good Hope, without precise

locality, collector unknown, in Herb. Plukenet (BM, holo.; PRE, photo.!).

Roots fibrous. Stems erect, usually with several wide-spreading branches, up to

1 m high, brown or green, zigzagging, terete, pubescent when young, terminating in

a spine. Branches similar to the stems. Branchlets fascicled, not replacing spines,

spineless except occasionally for a terminal spine, pubescent. Lateral spines formed

from modified branches, usually 3-nate, rarely more numerous, smooth, spreading at

right angles to one another, straight or recurved, the central one longest, brown or

green with a brown tip; some of the central spines on the older parts of the plant

replaced by normal branches. Cladodes terete, usually slightly recurved, 1-5-6 mm

or more long, glabrous or very minutely pubescent. Leaves scarious, with marginal

hairs, sometimes very greatly reduced. Flowers sessile, 1- or 2-nate, terminal. Perianth

segments oblong-obovate, 2-4-5 mm long, white with a green or purple median

streak; margin serrate. Stamens nearly as long as the perianth segments; anthers

about 0-5 mm long, yellow. Ovary about 1 mm long; style about 1 mm or less long.

Berry globose, 4 mm diameter, 1-seeded, red; perianth persistent.

South-west Cape, northwards to the southern parts of South West Africa, and

eastwards to Port Elizabeth.

For key to varieties, see key to species.

(a) var. capensis.

A. capensis L., Sp. PI. 314 (1753); Jacq., Hort. Schoen. t.266 (1798); Thunb., Prodr.

66 (1794); Bresler, Diss. 21 (1826); Bak. in Journ. Linn. Soc. Bot. 14; 617 (1875);

FI. Cap. 6: 263 (1896); Salter in FI, Cape Penins. 173 (1950); Solch, Beitr. FI. Siidwest-

Afr. 36 (1961). A. triacanthus Burm. f., Prod. FI. Cap. 10 (1768), nom. illegit. Type:

as for A. capensis. A. nelsonii Bak. in Journ. Linn. Soc. Bot. 14: 617 (1875); FI.

Cap. 6: 264 (1896). Type: South Africa, Nelson s.n. (BM, holo.; PRE, photo.!).

Asparagopsis passerinoides Kunth, Enum. PI. 5: 90 (1850), nom. illegit. Type: as

for A. capensis.

The typical variety is characterised by the generally smaller plants, narrower

stems and branches, and by the larger cladoles, which are always linear and more than

1 mm long.

This variety occurs over the entire range of the species except the coast of Nama-

kwaland and South West Africa. It occurs from sea-level to about 1,000 feet, especially

in sandy soils. Among shore vegetation it may form dense bushes about 1 m high,

with thick branches and short, thicker cladodes, while at higher altitudes it is generally

more sparsely branched and smaller, with narrower branches and cladodes. It is one

of the commoner species of Asparagus in the winter rainfall area. Flowering is affected

by, but not dependant on, fires. Flowers have been recorded in February and from

April to August. The flowers are sweetly scented.

Cape. — Bellville: Melkbosch, Wasserfall 159 (NBG). Bredasdorp: near Elim, Schlechter 10476

(BOL). Caledon: Grietjesgat, between Sir Lowrys Pass and Palmiet River, Ecklon & Zeyher s.n.

Calvinia: Calvinia. Compton 19484 (NBG). Cape: Lion’s Head, MacOwan 1977 (SAM). Carnarvon:

10 miles N.E. of Carnarvon, Hafstrom & Acocks 194. Clanwilliam: Langekloof, Schlechter 8029

(BOL). Hay: 16 miles E. of Griquatown, Acocks 2191. Hopefield: Hopefield, Jessop 455 (BOL).

Knysna: Keurboom’s River at Postpad, Fourcade 207 (BOL). Laingsburg: Ngaapkop, Compton

14591 (NBG). Malmesbury: Malmesbury, Marloth s.n. (STE). Middelburg: Witkransnek, Acocks

21415. Montagu: Fonteinkloof, Lewis s.n. (NBG). Murraysburg: Murraysburg, Tyson 38 (BOL).

Namakwaland: Garies, Thorne s.n. (SAM). Paarl: Tigerberg, Pillans 8656 (BOL). Piketberg: The

Rest, Barker 3590 (NBG). Port Elizabeth: Red House, Paterson 1109 (BOL). Somerset West:

Strand, Parker 3805 (BOL). Stellenbosch: Stellenbosch, Bond s.n. (NBG). Steynsburg: Steynsburg,

Thode 5636 (STE). Tulbagh: Saron, Schlechter 7886. Uitenhage: Uitenhage, Cooper 1573 (BOL).

Uniondale: Joubertina, Horn s.n. Vanrhynsdorp: near Vanrhynsdorp, Marloth 7794. Victoria

West: above Victoria West reservoir, Smith 2404. Worcester: Nature Reserve, Olivier 8. Wynberg:

Rondebosch, Wolley Dod 1143 (BOL).

44

South West Africa. — Luderitz: Zebrafontein, Merxmuller & Giess 3325 (M); 6 miles N. of Witputz

Police Station, Merxmuller & Giess 3446 (M).

Linnaeus, in his Species Plantarum, wrote: “Asparagus aculeatus, triplici spina,

surrectus. Pink. aim. 54. t. 15./. 4 But this figure does not show a plant with ternate

spines as described. On the page of text, page 54, to which Linnaeus referred, Plukenet

wrote: “Asparagus aculeatus, triplici spina surrectus, Prom. Bon. Spei. Phytogr. Tab.

78. fig. 3 While the polynomial is the same as that used by Linnaeus, the figure

is therefore different. This figure does show a plant with ternate spines. It would,

therefore, s;cm likely that Tab. 78, fig. 3 should accompany Linnaeus’ citation of

“ Pluk. aim. 54 ”, and the reference to Tab. 15, fig. 4 was an error. As the specimens,

labelled A. capensis , in the Linnaean Herbarium do not fit Linnaeus’ description,

they cannot be regarded as types. Both the Plukenet figure and the specimen on which

it was based in the Plukenet Herbarium match the polynomial in having ternate spines.

The specimen is therefore regarded as the holotype.

The binomial Asparagus triacanthus was accompanied, in Burmann’s work,

only by a reference to Plukenet’s Phytographia t. 78, f. 3, which is the figure regarded

by the present author as the illustration on which Linnaeus based his A. capensis.

These names are therefore synonymous.

Asparagopsis passerinoides Kunth is also a superfluous name for A. capensis of

Linnaeus. Kunth cited both Linnaeus’ description and the correct Plukenet figure.

Occasionally an atypical form occurs under trees. This form has longer

cladodes — up to 8 mm long — and a more lax habit. Also, it often has more numerous

spines at a node than is usual in this species. A specimen of this form, collected by

Nelson and now in the British Museum, was named A. nelsonii by Baker. He separated

A. nelsonii from A. capensis on the grounds that its branches were terminated by a

spine. There are few, if any, specimens of A. capensis in which the branches are not

terminated by a spine, however. No difference was found between these forms after

they had been cultivated near one another for two years. Nor is it possible to separate

these forms in the field because of specimens showing intermediate characters.

(b) var. litoralis Suess. & J. Karl, in Mitt. Bot. Staatssam. Miinchen 2: 50 (1950).

Type: South West Africa, Pomona, Dinter 6367 (B, holo.!; BOL!, PRE!, SAM!, Z!).

This variety is characterised by the larger plant, more robust stems and branches,

and small size of the cladodes. The cladodes are so reduced as to give a granular

appearance to the branchlets.

Retricted to the coast from Lambert's Bay in the Clanwilliam district to Luderitz

Bay in South West Africa. Flowers have been recorded in May.

Cape. — Clanwilliam: Lambert's Bay, Muir s.n. (SAM); 5 miles N. of Lambert's Bay, Taylor 3951;

Elandsbaai, Strey 427 (M). Namakwaland: south of Hondeklip Bay, Pi/lans s.n. (BOL).

South West Africa. — Luderitz: Luderitz Bay, Peyer 5 (Z); Pomona, Dinter 6367; Oranjemund,

Merxmuller 2335; Sturmvogelbucht, Merxmuller 2241; Buchubergen, Dinter 6463a (B).

There are specimens which show several intermediate characters between the two

varieties of this species. These occur in littoral vegetation but, although possessing

the more robust form of the second variety, do not have the extremely reduced cladodes.

A specimen of this form was grown near a specimen of the typical form of variety

capensis for about five years. During this time the differences were maintained. This

may indicate that a genetic difference is involved, but this is not of sufficient magnitude

either to create a third variety or to raise variety litoralis to specific status.

2. A. stipulaceus Lam., Encyc. 1: 297 (1783); Bresler, Diss. 5 (1826); R. & S.,

Syst. Veg. 7: 333 (1829); Salter in FI. Cape Penins. 173 (1950). Type: Cap e, Sonnerat

s.n. (P, holo.; PRE, photo.!).

45

Asparagopsis stipulacea (Lam.) Kunth, Enum. PI. 5: 91 (1850).

Asparagus densus Soland. ex Bak. in Journ. Linn. Soc. Bot. 14: 616 (1875). Type:

Cape, without precise locality, Auge s.n. (BM, holo.; PRE, photo.!).

Roots fibrous. Stems erect to 50 cm high, rarely more, not usually producing

branches, pubescent, grey-green to brown, ending in spines, angled or terete, ridged

when young, slightly zigzagging. Branches short, few or absent, not forming secondary

branches, spine-tipped. Lateral spines ternate, the longest occasionally replaced by

a branch, spreading at right angles to one another, straight or slightly curved, green

with a brown-red tip, up to 2 cm long. Branchlets fascicled, spineless or sometimes

terminating in a spine, grooved. Cladodes firm, terete or angled, slightly mucronate,

sometimes scabrid, 5-10 mm long, almost always 3-nate. Leaf scarious, fringed with

hairs. Peduncles represented by a minute disc, terminal, 1- or 2-nate. Perianth segments

oblong-obovate, white with a green or purple median streak, 5-6 mm long, spreading,

the margin serrate. Stamens almost as long as the perianth; anthers 0-7-0 -8 mm long,

yellow-green. Ovary 1-1-5 mm long; style 1 mm long. Berry fleshy, globose, red,

1- or 2-seeded.

This species grows in sandy soils, often in vegetation with a high proportion of

Restionaceae, and usually within sight of the sea, from the west coast of the Cape

Peninsula to the eastern parts of the Bredasdorp district. It is particularly affected

by fires and rarely flowers except after fires. Flowers have been recorded in April,

May and July. Fruits are rarely produced. Even in areas where large numbers of

plants have flowered, it is usually impossible to find fruits. This is possibly associated

with polypoidy.

Cape. — Bredasdorp: Mierkraal, Schlechter 10536; De Hoop, Barker 8731 (NBG); Baardskeerders

bos, Oliver s.n. (BOL); near Gansbaai, Oliver s.n. (BOL). Caledon: Rooi Els, Jessop s.n. (BOL);

Betty’s Bay, Oliver s.n. (BOL). Wynberg: Karbonkelberg, Oliver s.n. (BOL). Simonstown: Vasco

da Gama, Salter 8046A (BOL); .Jessop 135 (BOL); Buffels Bay, Guthrie s.n. (BOL); Oliver s.n. (BOL);

Miller’s Point, Pillans 8532 (BOL); Clovelly, Sailer 8210 (BOL); Kalk Bay, Jessop 47 (BOL); Oliver

s.n. (BOL); St. James, Michell s.n. (BOL); hills near Fish Hoek, Peers s.n.

The type specimen was collected by Sonnerat at the Cape. Two specimens have

been mounted on the same sheet. That on the left is of an Asparagus, and matches

Lamarck’s description. The other is of a Psoralea. The specimen on the left has,

therefore, been selected as the type.

A. capensis and A. stipulaceus are very similar, and their separation is sometimes

difficult. A. capensis is larger and considerably more branched, while A. stipulaceus

is usually more pubescent and has larger claodes and flowers. At Rooi Els where the

two species grow together on a slope above the river, they retain their distinguishing

characters.

Chromosome counts were made on specimens of A. capensis from about four

miles inland from Ysterfontein, in the Malmesbury district, and of A. stipulaceus from

about half way up the mountain behind Kalk Bay in the Simonstown district. The

diploid number was found to be 40 for the former species and very near to — probably

exactly — 80 for the latter. This difference in chromosome number suggests that the

small qualitative differences between the two species is not only the result of inadequate

sampling of the populations of each form.

3. A. suaveolens Burch., Trav. 2: 226 (1822); Pole Evans in Flow. PI. S. Afr.

11: t. 409 (1931). Type: Griquatown, Burchell 1956 (K, holo.!; PRE, photo.!).

A. spinescens Steud. ex R. & S., Syst. Veg. 7: 334 (1829). Type: Cape, without

precise locality, Ludwig s.n. A. triacanthus Willd. ex R. & S., Syst. Veg. 7: 334 (1829).

Type: Cape, Lichtenstein in Herb. Willd. no. 6693. A. burchellii Bak. in Journ. Linn.

Soc. Bot. 14: 618 (1875); FI. Cap. 6: 264 (1896). Syntypes: “Cap. Bon. Spei in

46

aridis ”, Burchell 2962 (K ! ) ; Zeyher ( Asparagus 10); Cooper 1574. A. stipulaceus

sensu Bak. in Journ. Linn. Soc. Bot. 14: 617 (1875); FI. Cap. 6: 264 (1896). A.

spinosissimus Kuntze, Rev. Gen. PI. 3: 315 (1893); Solch, Beitr. FI. Siidwest-Afr. 39

(1961). Type: Cathcart, Kuntze s.n. (NY, holo.; BOL, photo.!). A. omahekensis

Krause in Bot. Jahrb. 51 : 447 (1914); Solch, Beitr. FI. Siidwest-Afr. 38 (1961). Type:

“ Zwischen Gobabis und Oas ”, Dinter 2711 (B, holo.!; SAM !). A. intangibilis Dinter

in Fedde, Rep. 29: 269 (1931); Solch, Beitr. FI. SUdwest-Afr. 37 (1961). Type:

“ Grosse Karasberge ”, Dinter 5168 (B, holo.!).

Asparcgopsis triacantha (Willd.) Kunth, Enum. PI. 5: 91 (1850). A. zeyheri Kunth,

Enum. PI. 5; 92 (1850). Type: Without locality, Zeyher s.n. A. spinescens (R. & S.)

Kunth, Enum. PI. 5: 93 (1850).

Roots fibrous. Stems erect, to 1 m, or occasionally climbing or scrambling to

0- 5 m, zigzagging, spine-tipped, glabrous or pubescent, pale cream coloured to brown.

Branches similar to the stems, usually spreading or ascending, but occasionally reflexed.

Lateral spines 3-, 5- or 7-nate, at least the lateral ones of each group usually bearing

sterile nodes (Fig. 2a), pungent, well-developed, straight or arcuate, spreading or

reflexed. Branchlets fascicled, paired to numerous, herbaceous, arcuate or straight,

up to 3 cm long. Cladocles borne on branchlets, fascicled, subulate, spreading or

ascending, 3-6-nate, 1-10 mm or more long. Peduncles solitary, terminal on branchlets,

1- 4 mm long, articulated near the base. Perianth segments obovate, 1-5-3 -5 mm

long, the outer whorl frequently, perhaps always, with a purple streak. Stamens nearly

as long as the perianth segments, narrow; anthers about 1/3 mm long, orange to yellow.

Style divided for less than half its length; ovary sessile, 12-14-ovuled. Berry one-seeded,

globose, 3—3 - 5 mm diameter, red or black; perianth persistent.

This is one of the commoner species of Asparagus in the summer rainfall area,

occurring in Rhodesia, large parts of South West Africa, Bechuanaland and most of

the areas to the south, and extending into the winter rainfall area as far as the Clan-

william district in the north and Caledon in the east. It has not been recorded from

Basutoland and there are few records from the south-eastern parts of South Africa.

It occurs in coastal vegetation, low bush, open forest and Karroid vegetation. Flowers

have been recorded in most months, but particularly in autumn and early summer.

There does not seem to be any geographical variation in flowering period.

Cape. — Albany: Botha's Hill, Rogers 3417 (BOL). Alexandria: Debega Valley, Galpin s.n. Aliwal

North: Doctor’s Drift, Gerstner 245. Barkly West: near the Vaal River, Nelson 1 76. Bathurst: Port

Alfred, Salisbury s.n. (GRA). Bedford: Witmos, Galpin 3081. Bredasdorp: Mierkraal, Schlechter

10527. Caledon: Caledon, Esterhuysen s.n. (BOL). Cathcart: 1 mile from Henderson on Cathcart

road. Story 2805. Clanwilliam: Langkloof, Schlechter 8050. Cradock: 10 miles E. of Cradock,

Jessop 656. De Aar: De Aar, Acocks 632. East London: East London, Thocle 6653 (STE). Fraser-

burg: Fraserburg, Net s.n. (STE). Gordonia: Gordonia district, Kotze 852. Graaff-Reinet : Graaff-

Reinet, Bolus 411 (BOL). Hankey: Hankey, Barker 7853 (NBG). Hay: Elmadale, Acocks 2231.

Heidelberg: 5 miles W. of Heidelberg, Jessop 591. Kentani: Columbia Mission Station, Pegler 1451

(BOL). Kimberley: 7 miles N.E. of Kimberley, Leistner <& Joynt 2650. King Williams Town: King

Williams Town, Sim 1060 (BOL). Komga: Komga, Flanagan 1793. Kuruman: 10 miles S. of

Kuruman, Louw 842. Ladismith: Prinspoort, Bond 306 (NBG). Laingsburg: Tweedside, Marloth

12076. Middelburg: Grootfontein, Theron 94. Montagu: Cogman’s Kloof, Barker 8625 (NBG).

Murraysburg: Roads Poort, Tyson 212 (GRA). Namakwaland: Doornpoort, Pillans 5466 (BOL).

Pearston: by the Blyde River, Burchell 2962 (K). Phillipstown : Potfontein Station, Schweickerdt

1196. Oudtshoorn: near dam, Britten 1624. Port Elizabeth: Cradock Place, Galpin 6439. Post-

masburg: west entrance to Langkloof, Leistner 1164. Prieska: near Prieska, Bryant J368. Prince

Albert: Farm Damascus, Marloth 1 1276. Queenstown: Long Hill, Galpin 2095. Riversdale: Lange-

berg, Muir 2529 (BOL). Robertson: Klaasvoogds, Van Niekerk 372 (BOL). Somerset East : Somerset

East, Van Niekerk 435 (BOL). Swellendam: National Bontebok Park, Liebenberg 6536. Taungs:

Kankatjies, Acocks 538. Uitenhage: near Uitenhage, Schlechter 2573. Uniondale: 8 miles from

Uniondale on Willowmore road, Fourcade 6256. Vryburg: Tiger Kloof, Brueckner 487. Worcester:

Veld Reserve, Van Breda 21.

Natal. — Camperdown: Nagle Dam, Wells 1361. Ubombo: Ubombo, Gerstner 3400 (NH).

Weenen: Mielietuin, West 1204.

47

Orange Free State. — Bloemfontein: south of Glen, Mostert 326. BoshofF: between Boshoff

and Windsorton Road, Burtt Davy s.n. Fauresmith: Fauresmith commonage, Leistner 1099. Kroon-

stad: Kroonstad, Pont 196. Parys: Parys, Rogers 5175. Trompsburg: Lucerne, Henrici s.n. Winburg:

Willem Pretorius Game Reserve, Bakker 1004.

Swaziland.- — Hlatikulu, Ingwavuma Poort, Ben Dlamini s.n.

Trasvaal. — Barberton: Barberton, Thorncroft 844 (NH). Brits: Jacksonstuin, Van Vuuren 292.

Groblersdal: 4 miles S. of Bethal Mission Station, Monnig 48. Fleidelberg: The Kloof, Thode 3435

(STE). Johannesburg: Milner Park, Moss 17583. Lichtenburg: Grasfontein, Sutton 461. Lydenburg:

Farm Parys, Barnard & Mogg 702. Marico: Zeerust, Thode 1508. Messina: Messina, Rogers 1-9297

(BOL). Middelburg: Tautesberg, Young A 163. Pietersburg: Reebok Vlei, Mauve 4295. Potchef-

stroom: Losberg, Theron 693 (NF1). Potgietersrus: Mosdene, Galpin 342. Pretoria: near Pretoria,

Schlechter 3584. Roodepoort: Spitskop, Archibald 2829 (GRA). Sibasa: 7 miles N.W. of Punda

Milia, Codd & De Winter 5556. Ventersdorp: Goedgedacht, Sutton 612. Vereeniging: Klipriviersberg,

Gilliland s.n. (BOL). Waterberg: Towoomba Pasture Research Station, Codd 6116. Zwartruggens:

Zwartruggens, Story 5638.

South West Africa. — Gobabis: Okasondana, Walter 4145 (M). Grootfontein : Otaviberge,

Rehm s.n. (M). Rehoboth: Floachanas, Dinter 1948 (SAM). Warmbad: Narudas-Kanus, Dinter

5168.

Bechuanaland. — Molepolole, Story 4869.

In the south-western Karroo, Little Karroo and as far east as the Albany district

there are two forms. One of these is erect and has spreading to ascending branches,

while the other, to which Baker gave the name A. burc/iellii, varies from erect to

climbing and usually has reflexed branches and spines. Evidence, based both on

herbarium material and on field observations, has shown that these forms are linked

by many intermediates and cannot be given taxonomic recognition. These intermediates

occur in the eastern regions of the distribution of the two forms but are absent in the

west. In the Transvaal there are also two forms which exist side by side. The larger

form is glabrous, and has black berries, while the smaller is pubescent, has a particularly

neat appearance, smaller cladodes, spreading branches and red berries. It was again

not found possible to separate these two forms when the variability of the plants of

this species from other areas of South Africa is taken into consideration. Two “ fox-

tail ” forms have been recorded. One of these, from the Laingsburg district, is only

about 25 cm high. It has been collected by Rogers (BOL 16322, Tvl. Museum 21 167).

The other, which is often over 1 m high, has been collected several times, for example

Mogg 22407 (PRE), in the Loskop Dam area in the Transvaal. Little-branched forms

occur in most areas where this species is found, and it is not felt that this character

should be regarded as taxonomically significant in this species.

4. A. glaucus Kies in Bothalia 6: 229, fig. 5 (1954); Solch, Beitr. FI. SLidwest-Afr.

36 (1961). Type: Fauresmith, Groenvlei, Kies 340 (PRE, holo.!).

Roots fibrous. Stems erect, up to 50 cm high, smooth, glabrous, woody, slightly

zigzagging. Branches spreading to ascending, solitary; stems and branches white at

first, becoming brown, often spine-tipped, but frequently damaged. Branchlets 1-3 cm

long, solitary or paired, straight, grooved, greenish, borne at the bases of spines or

branches, terminating in a spine. Lateral spines solitary, mostly less than 1 cm long,

occasionally up to 2 cm long, straight or recurved, slightly ascending or spreading,

pungent, bearing sterile nodes and often bearing lateral spines. Cladodes 2-10 mm

long, glaucous, fascicled, frequently 3-nate, terete, frequently up to 0-5 mm thick.

Leaves very reduced; lateral buds present at the bases of branches and spines.

Peduncles axillary, 2-nate, 2 mm long, borne on branchlets or spines, articulated near

the centre. Perianth segments similar, broadly oblong-obovate, spreading, 3 '5-4-0 mm

long, margins smooth, rather tough, green in bud, becoming greenish-white with a

green keel, with a bloom. Stamens about 3 mm long; filaments white, flattened

laterally; anthers oblong, about 1 mm long. Ovary shortly stalked; the ovules about

9 in each chamber; style branches shortly fused at the base. Fruit globose, 4 mm

diameter, containing a single seed, surrounded by a persistent perianth.

48

Found in the dry areas from the Oudtshoorn district in the south-west, northwards

to Keetmashoop and eastwards to Bloemfontein. It appears to have a strong preference

for brakish soils. Flowers have been recorded from October to December.

Cape. — Cradock: Jakkalsfontein, Acocks 17516. Hanover: Hanover, Sim s.n. Kimberley: Mollers

Pan, Acocks & Hafstrom H874; Leeuwpoort, Burtt Davy 9553. Oudtshoorn: Kruispad, Compton

21779 (NBG). Uniondale: bank of the Olifant’s River, Fourcade 6265 (BOL).

Orange Free State. — Bloemfontein: De Brug, Acocks 8464. Fauresmith: Groenvlei, Kies 340;

Heuningberg, Marais 124; Veld Reserve, Henrici 3981. Luckhoff: Rose Marie, Verdoorn 2148;

Henrici 4535. Trompsburg: Ventersvlei, Verdoorn 1153.

South West Africa. — Keetmanshoop : Great Karasberg, Noachabeb-Witmond, Ortendahl 425.

This species has similarities with A. suaveolens, but differs in having solitary spines,

fewer branches at each node, axillary peduncles, and larger, thicker perianth segments

which envelope the fruit.

5. A. africanus Lam., Encyc. 1: 295 (1783); Bresler, Diss. 9 (1826); R. & S.,

Syst. Veg. 7: 331 (1829); Bak. in Journ. Linn. Soc. Bot. 14: 619 (1875); FI. Cap. 6:

265 (1896); Marloth, FI. S. Afr. 4: pi. 20 (1915); Salter in FI. Cape Penins. 175

(1950); Kies in Bothalia 6: 177 (1951). Type: Cape without precise locality, Sonnerat

s.n. (P, holo.; PRE, photo.!).

Asparagus dependens Thunb., Prodr. 66 (1794); Bresler, Diss. 17 (1826); Kunth,

Enum. PI. 5: 74 (1850). Type: Cape of Good Hope, without precise locality, Thunberg

s.n. (UPS, holo.!; BOL!, PRE! photos). A. cooperi Bak. in Gard. Chron. 1: 818

(1874); Journ. Linn. Soc. Bot. 14: 613 (1875); FI. Cap. 6: 263 (1896); Flow. PI.

S. Afr. 13: t.503 (1933). Type: Cape without precise locality, Cooper s.n. (K, holo.!;

PRE, photo.!). A. multiflorus Bak. in Journ. Linn. Soc. Bot. 14: 610 (1875); FI.

Cap. 6: 261 (1896); Kies in Bothalia 6: 177 (1951). Type: “ Somerset division ”,

Bowker s.n. (K, holo.!; PRE, photo.!). A. asiaticus sensu Bak. in Journ. Linn. Soc.

Bot. 14: 618 (1875); FI. Cap. 6: 265 (1896). A. burkei Bak. in Journ. Linn. Soc.

Bot. 14: 607 (1875); FI. Cap. 6: 261 (1896). Type: South Africa, Crocodile River,

Burke s.n. (K, holo. ! ; PRE, photo. !). A. pilosus Bak. in Journ. Linn. Soc. Bot. 14: 610

(1875). Type: Bechuanaland, near Lake Ngami, McCabe 15 (K, holo.!; PRE, photo. !).

A. fleckii Schinz in Bull. Herb. Boiss. 1, 4, app. Ill: 44 (1896); Solch, Beitr. El.

Siidwest-Afr. 37 (1961). Type: S.W.A., Hoachanas, Fleck 901 (Z, holo.!; PRE,

photo.!). A.judtii Schinz in Bull. Herb. Boiss. 1, 4, app. Ill: 44 (1896); Solch, Beitr.

FI. Siidwest-Afr. 37 (1961). Type: S.W.A., Hoachanas, Fleck 901 (Z, holo.!). A.

conglomeratus Bak. in FI. Trop. Afr. 7: 428 (1898); in Vierteljahrsschr. Naturf. Ges.

Zurich 49: 176 (1904). Type: Ngamiland, Kwebe, Lugard 52 (K, holo.!; PRE,

photo.!). A. bechuanicus Bak. in FI. Trop. Afr. 7: 429 (1898). Type: Bechuanaland,

Nakalechwe, Lugard 25 (K, holo.!; PRE, photo.!). A. lugardii Bak. in FI. Trop.

Afr. 7: 431 (1898). Type: Bechuanaland, near Lake Ngami, Lugard 31 (K, holo.!;

PRE, photo.!). A. dinteri Engl. & Krause in Bot. Jahrb. 45: 155 (1910); Solch, Beitr.

FI. Siidwest-Afr. 36 (1961). Type: “ Bei Neitsas ”, Dinter 676 (B!, SAM!). A. patens

Krause in Bot. Jahrb. 51: 448 (1914). Type: S.W.A., Haribes, Engler 6570 (B!).

A. francisci Krause in Bot. Jahrb. 51: 449 (1914). Type: S.W.A., “ Zwischen

Omuramba und Omatako ”, Seiner 689a (B!). A. rivalis Burch, ex Kies in Bothalia

6: 179 (1951); Solch, Beitr. FI. Siidwest-Afr. 38 (1961). Type: Kosiefontein, Burchell

2587 (K, holo.!; PRE, photo.!).

Asparagopsis lamarckii Kunth, Enum. PI. 5: 87 (1850), nom. illegit. Type: as for

A. africanus. A. juniperina Kunth, Enum. PI. 5: 85 (1850). Type: Swartkops River,

Drege 8576. A. schlechtendalii Kunth, Enum. PI. 5: 90 (1850). Type: Olifants River,

Krebs s.n.

Roots fibrous. Stems woody, twining, either climbing to about 3 m or forming

low bushes up to about 1 m high, glabrous to pubescent, smooth or slightly grooved,

slightly zigzagging, grey-brown to greenish. Branches solitary, much-branched, similar

49

to the stems, but with a greater tendency to pubescence and grooving, usually spreading,

but in some forms — particularly the pubescent specimens — strongly reflexed. Cladodes

fascicled, up to 20-nate, but often about 8-nate, straight or slightly arcuate, subulate,

generally 5-10 mm long, ascending to spreading. Spines usually present on at least

the stems and larger branches, spreading to reflexed, usually straight, pale in many

specimens but frequently reddish brown, up to 1 cm or more long, usually with the

rest of the leaf forming well-developed spurs. Peduncles axillary and a few sometimes

terminal, up to 20-nate, but frequently 2- or 3-nate, up to 5-8 mm long, articulated

below the centre. Perianth segments similar, entire, oblong-obovate, 2 -5-3 -5 mm

long, white. Stamens nearly as long as the perianth segments. Style and ovary each

about 1 mm long. Fruit globose, fleshy, red, about 6 mm diameter, with a single seed.

This species usually occurs in bushy or wooded areas, where it climbs, but also

occurs in more open vegetation where it forms bushes or scrambles among rocks.

It grows mainly in the damper parts of South Africa, but also occurs in drier areas

and in South West Africa, Bechuanaland and further north. Flowers have been

recorded from August to May.

Cape. — Albany: Grahamstown, Galpin 284. Alexandria: Kowie Road from Southwell, Britten

2596. Aliwal North: Orange River, Gerstner 160. Barkly West: Vaal River at Sydney, Acocks

H830. Bathurst: Port Alfred, Rogers 16623 (BOL). Bedford: Bedford, Gane 118 (GRA). Bellville:

Tigerberg, Pi/Ians 8659 (BOL). Caledon: Baviaan’s Kloof, Genadendal, Gillett 855 (BOL). Cape:

Porcupine Buttress, Jessop 12 (BOL). Cathcart: 5 miles N. of Cathcart, Jessop 660. Cradock: 17

miles along Wapadberg road from Graaff-Reinet-Middelburg road, Jessop 653. East London: East

London, Bokelman 5-28 (NBG). Fort Beaufort : Beaufort, Cooper 523 (BOL). George: near George,

Barker 6856 (NBG). Gordonia: Kakoup, 50 miles S.W. of Olifantshoek, Leistner 2046. Graaff-

Reinet: near Graaff-Reinet, Bolus 132 (BOL). Heidelberg: 5 miles W. of Heidelberg, Jessop 589.

Herbert: St. Clair, without collector 237 (GRA). Humansdorp: Ratelsbosch, Fourcade 552. Kentani:

Kobongaba Hills, Pegler 1463. Kimberley: Christian’s Drift on Riet River, Leistner 1995. King

Williams Town: King Williams Town, Sim 1063 (BOL). Knysna: Leisure Island, Ta_ vlor 1019 (NBG).

Komga: Komga, Flanagan 299. Mafeking: Mafeking, Burtt Davy 11087. Middelburg: Groot

Vlei, Van Wyk’s Vlei, Archibald 3042 (GRA). Mossel Bay: Gouritz Valley, Horn s.n. Mount Currie:

Kokstad, Mogg 5177. Oudtshoorn: near Cango, Bolus 12369 (BOL). Paarl: Klein Drakenstein

at Salem, Bolus s.n. (BOL). Peddie: Fish River Valley below Breakfast Vlei, Acocks 1190. Port

Elizabeth: Cradock Place, Galpin 6408. Queenstown: Queenstown, Galpin 2152. Somerset East:

Zuurberg, Compton 20284 (NBG). Stellenbosch: Franschhoek, Schlechter 9218. Stockenstrom :

Hogs Back, Rattray s.n. (GRA). Stutterheim: Ripplemead, Kabousie, Hutton 482 (GRA). Tarkastad:

near Hofmeyer turning on Cradock-Tarkastad road, Jessop 655. Uitenhage: Enon, Thode A2769

(NH). Uniondale: between Krakeel and Joubertina, Jessop 595. Vryburg: Stella, 30 miles N. of

Vryburg, Hardy & Bayliss 1244. Wynberg: Orange Kloof, Wolley Dod 2470 (BOL).

Basutoland.— Leribe, Dieterlen 346.

Natal. — Bergville: Natal National Park, Edwards 962. Camperdown: Bothas Hill, Hutchinson,

Forbes & McClean 55 (NH). Durban: Berea, Forbes 383 (NH). Eshowe: Eshowe, Gerstner 2405.

Estcourt: Dalton Bridge, Acocks 10626. Himeville: 5 miles N. of Himeville, Jessop 664. Inanda:

Inanda, Wood 989 (NH). Klip River: near Colenso, Wood 4089 (NH). Lions River: 5 miles along

Merrivale-Boston road, Moll 937. Mtunzini: Umlalazi Nature Reserve, Ward 4275. New Hanover:

Albert Falls, Comins 84 (NH). Nongoma: Bangonoma, Gerstner s.n. (NH). Pietermaritzburg:

Pietermaritzburg, Schlechter 3297 (BOL). Pinetown: Gillitts, Lansdell s.n. (NH). Richmond:

Richmond, Schlechter 6717 (BOL). Umlazi: Isipingo, Wood 12480 (NH). Umvoti: Greytown, Wylie

s.n. Utrecht: Tweekloof, Altemooi, Thode A212.

Orange Free State. — Bethlehem: Kloof, Potgieter 81. Bloemfontein: Bloemfontein, Potts s.n.

(BOL). Boshoff: Smitskraal, Burtt Davy 10305. Fauresmith: Sleutelpoort, Henrici 2593. Harrismith:

Maccauvlei, Brandmuller 1 19. Kroonstad: Kroonstad, Powt 195. Winburg: Willem Pretorius Game

Reserve, Leistner 2975.

Swaziland. — Near Gollel, Compton 29105; Forbes Reef, Compton 30974; Stegi, Compton 26021.

Transvaal. — Barberton: Berea Hill, Galpin 622. Christiana: Bloemhof, Kameelpan, Theron

5633. Ermelo: Mavieriestad, Pott s.n. Heidelberg: Suikerbosrand, Mogg 18129. Johannesburg:

Zoo Kopjes, Moss 10498. Krugersdorp: Jack Scott Private Nature Reserve, Weds 2413. Letaba:

Tzaneen, Rogers 12506 (GRA). Lydenburg: Lydenburg, Thode A1648 (NH). Middelburg: Olifant’s

River Gorge, Mogg 22445. Nelspruit: 10 miles S. of Lower Sabie, Codd 6134. Pietersburg: Haenerts-

burg, Rogers 19041 (BOL). Pilgrim’s Rest: Sabie, Rogers 23090. Potchefstroom : Potchefstroom,

50

Burt/ Davy 48. Potgietersrus: Mosdene, Galpin 17346. Pretoria: Pretoria, Schlechter 3603. Sout-

pansberg: Spelonken, Jenkins s.n. Standerton: Standerton, Leendertz s.n. Ventersdorp: Goed-

gedacht, Sutton 522. Vereeniging: Lagerspoort, Mogg 19532. Wakkerstroom: Oshoek, Devenish

97. Warmbad: Warmbad, Leendertz s.n. Waterberg: banks of the Limpopo, Leipoldt 71 (BOL).

South West Africa. — Grootfontein: West Runtu “ in Okavongotal ”, Volk 1860a (M). Luderitz:

Aus, Peyer 15 (Z). Mariental: 60 miles from Mariental, Liebenberg 5154. Okahandja: Okahandja,

Hopfner c-65 (Z). Omaruru: Brandberg, Rodin 2713. Otjiwarongo: Okosongomingo, Volk 2568

(M). Rehoboth: Gravenstein, Volk 1 1 502 (M). Warmbad: Naruchas, Dinter 7214 ( M). Windhoek:

Voigtland, Walter 75 (M).

Bechuanaland. — Toteng, N.E. tip of Lake Ngami, Story 4690; 103 miles N.N.W. of Kan, De

Winter 7350.

Lamarck described A. africanus from a specimen which was sent from the Cape

by Sonnerat. The description mentions the angled stems and spine and cladode

characters which are typical of the form growing in the south-west Cape. This is

the area where Sonnerat is likely to have been able to have collected it. The holotype

is in the Lamarck Herbarium in Paris. Lamarck also cited Plukenet’s Phytographia,

t. 374 f. 4. This figure shows a slightly tortuose stem without spines, but Plukenet

mentioned the presence of spines in his description. The locality was given as Africa.

If this figure is intended to represent the Cape species, the shape of the cladodes and

absence of spines makes it a very poor likeness.

Kunth cited Drege 8576, which was collected at the Swartkops River, as the type

of Asparctgopsis juniperina. All the characters described are those of this species. In

particular, the spines, described as “ ferruginea ”, are very characteristic. Asparctgopsis

schlechtendalii Kunth is also likely to be a synonym for this species. The type was

collected along the Olifants River, but Kunth does not indicate which Olifants River

this is. The plant, is described as woody with flexuose branches. The cladodes and

axillary flowers are typical for A. africanus, but there is not enough information to

be absolutely certain.

Because various authors have used the name A. asiaticus L. for the species treated

here as A. africanus, it is desirable to discuss the typication of A. asiaticus here. Linnaeus

cited his Hortus Cliffortianus, page 122 (1737) in his description of A. asiaticus. In this

work, there is a reference to Plukenet’s Phytographia (1699) t. 15 f. 4. As there is

no specimen with this name in the Linnaean Herbarium, the Plukenet figure would

be regarded as the iconotype if there is no specimen in Herb. Sloane in the British

Museum upon which this illustration is based. Plukenet cited his species as “ e Maderas-

patans”; that is from the eastern coast of Peninsular India. Whether or not the

Indian species is synonymous with the South African plants called A. africanus in this

treatment would require further work on the material from India and Africa. There

are, however, several characters of Linnaeus’ description and of the figure which make

it very unlikely that these plants are conspecific. Linnaeus described the stems as

erect, and the figure also shows a rigidly erect stem. The figure also does not match

the South African species as it appears to have spines on the final branches. It therefore

seems better to regard this name as not applicable to the South African species until

work is done on the genus in a wider geographical context. This decision is in agree-

ment with that of Kies (1951).

Those South African Asparagi with fascicled, terete cladodes, stems tortuous

and not white and grooved, final branches and cladodes not horizontal as in A. setaceus

have been included in this species. The variation within these limits, as can be seen

from the description, is very great. It is possible that further field work would reveal

the limits of significant groupings within this definition, but, judging from the available

material and data, there seems to be no way of correlating any of the variable characters

in order to subdivide the group.

51

The form in the south-west Cape, A. africanus of Baker, has well-developed,

reddish spines, and tends to be less tortuous. This is replaced to the east by a more

pubescent form with straighter, reflexed branches and cladodes, referred to by Baker

as A. africanus var. dependens. More generally over most of the summer rainfall area

there is a less spiny, more tortuous form with narrower stems, branches and cladodes,

which was referred to A. asiaticus by Baker. This form becomes slightly herbaceous,

and may even produce annual stems.

In hot, dry areas specimens occur in which cladodes do not form, or are reduced,

towards the tips of the branches. This may be associated with the environment. Several

species have been based on specimens of this form, among them A. burkei Bak. and

A. judtii Schinz.

It is difficult to distinguish A. africanus from A. setaceus. The form of branching

and cladode arrangement which are so well known and characteristic of the horti-

cultural plants occurs in wild populations over a large part of South Africa. Inter-

mediate forms do occur, however, in which the branching shows some similarity with

A. africanus. When there is any doubt, these species may be separated by the terminal

flowers of A. setaceus, and axillary flowers of A. africanus.

6. A. setaceus ( Kunth ) Jessop, comb. nov. Asparagopsis setacea Kunth. Enum.

PI. 5: 82 (1850). Type: “ Drege, Herb. no. 8584. c.v.s. in Herb. reg. Berol. et Luc.”

(KIEL, lecto.!).

Asparagus declinatus sensu Bak. in Journ. Linn. Soc. Bot. 14: 609 (1875); LI. Cap.

6: 261 (1896). A. plumosus Bak. in Journ. Linn. Soc. Bot. 14: 613 (1875); Planchon

in Llor. des Serres ser. If, 23: t. 2413-2414 (1880); Bak. in FI. Cap. 6: 260 (1896).

Syntypes: Port Natal, Drege 4482 (K!); British Kaffraria, Cooper 202, partly, ( K ! ) ;

without locality, Gerrard & Me Ken 754.

Roots fibrous. Stems twining, smooth or grooved, glabrous, greenish to reddish-

brown. Branches similar to the stems, solitary. Cladodes linear, fine, arcuate, terete,

up to 10-nate, up to 10 mm long; final branches and cladodes all in the same plane,

horizontal (homalotropic). Spines not usually present, reflexed. Peduncles terminal,

usually solitary, rarely up to 3-nate, 2-5 mm long, articulated near the middle, pendent.

Perianth segments cream-white, oblong-obovate, entire, similar, about 3 mm long.

Stamens slightly shorter than the perianth segments; anthers ^-3 mm long. Styles

shortly divided, about 1 mm long; ovary about 1 mm long. Berry red, fleshy, up to

3-seeded, about 6 mm diameter.

This species occurs mainly in forests, but also in open country provided there is

sufficient support, from the Riversdale and Swellendam districts in the south-west,

through the eastern Cape and Natal into the Transvaal. To the north its range

continues into Rhodesia, and in the east into Mozambique. Llowers have been recorded

from September to April.

Cape. — Albany: Sandy Drift near Grahamstown, Daly 6 (GRA). Alexandria: Olifantshoek

between mouths of Bushman’s and Sunday’s Rivers, Ecklon & Zeyher s.n. Bathurst: Horseshoe on

Kowie River, Barker 9267 (NBG). George: Karatara, Drege 8584 (KIEL). Humansdorp: Ratelsbos,

Fourcade 377 (BOL). Kentani: woods near Ken tani, Pegler 345. King Williams Town : King Williams

Town, Sim 1043 (BOL). Knysna: Knysna, Marloth 7504. Komga: Komga woods, Flanagan 1711.

Queenstown: Umbombola Mountains, Galpin 2091. Riversdale: above Novo, Muir 2837 (GRA).

Somerset East: Boschberg, MacOwan 1917 (SAM). Stockenstrom: Katberg near Sanatorium,

Schonland 4307 (GRA). Swellendam: Grootvadersbos, Zeyher 4167 (BOL). Umtata: Convent of

the Holy Cross, without collector. Willowvale: Qora River mouth, Hilner 443.

Natal. — Durban: woods near Durban, Wood 437 (BOL). Eshowe: Eshowe, Lawn 730 (NH).

Estcourt: Giant’s Castle, Symons 40. Hlabisa: Hluhluwe Game Reserve, Ward 2159. Inanda:

Wood 1649 (BOL). Nkandla: Nkandla Forest, Nixon s.n. (NH). Ubombo: near turning to Mkuzi

Game Reserve, Jessop 668.

Swaziland.— Mbabane, Malagwane Hills, Ben Dlamini s.n. (NBG).

52

Transvaal. — Barberton, Thorncroft 2116. Belfast: Belfast, Leendertz 7953. Brits: Jacksonstuin,

Van Vuuren 65. Krugersdorp: 1 mile N.W. of Krugersdorp, Mogg 20305. Letaba: Grootbos

Government Forest Reserve, Sc/ieepers 896. Nelspruit: Numbi, Van der Schijff 1614. Pietersburg:

near Woodbush, Gillett 3192 (BOL). Pilgrim’s Rest: Mariepskop, Van der Schijff 5006. Pretoria:

Fountains Valley, Repton 717. Soutpansberg: Downs, Junod 4133. Ventersdorp: Goedgedacht,

Sutton 715.

In his description of Asparagopsis setacea, Kunth cited only one collecting, namely

Dregs 8584. However, he referred to two specimens, of which one was in Berlin,

and has now been destroyed, and one in “ Herb. Luc.”. This specimen is now in

Kiel. It is labelled Drege 8584b and was collected at Karatara in the George district.

Baker cited several specimens in his description of A. plumosus. Of these, Drege

4482 (K) from Port Natal, and Cooper 202, partly, (K) from British Kaffraria, have

been seen by the present author. A third specimen, Gerrard & McKen 754 without

precise locality, could not be found at Kew. Baker cited Asparagopsis setacea Kunth

under the synonymy of Asparagus declinatus L. He separated A. declinatus from

A. plumosus on the cladodes being filiform in A. plumosus but more robust in A.

declinatus. Asparagus declinatus was described by Linnaeus in his Species Plantarum

(1753) as being spineless, with declinate branches, and setaceous cladodes, and as

coming from Africa. Baker upheld this species in both the Journal of the Linnean

Society, and in the Flora Capensis, and cited MacOwan 1917 and Thunberg s.n. These

gatherings are included here in A. setaceus. It is possible that A. declinatus L. and

A. setaceus are conspecific, but the true identity of A. declinatus is obscure and it is

best discarded as a nomen dubium.

7. A. denudatus (Kunth) Bak. in Journ. Linn. Soc. Bot. 14: 606 (1875); FI. Cap.

6: 258 (1896). Type: Queenstown, Stormberg, Drege 3533 (BOL, lecto.!).

Asparagopsis denudata Kunth, Enum. PI. 5: 82 (1850).

Asparagus namaensis Schinz in Bull. Herb. Boiss. 1, 4, app. Ill: 44 (1896); Solch,

Beitr. FI. Siidwest-Afr. 37 (1961). Type: Great Namaqualand, Fleck 307a (Z, holo.!).

A. pearsonii Kies in Bothalia 6: 179 (1951); Solch, Beitr. FI. Siidwest-Afr. 38 (1961).

Type: Raman’s Drift, Pearson 4003 (K, holo.).

Roots fibrous. Stems erect to about 1 m or scrambling to nearly 2 m, zigzagging,

glabrous, smooth or wrinkled but never regularly grooved, green; the outer layers

easily separable from the aerial stems and branches, but not peeling. Branches usually

solitary but sometimes 2- or 3-nate, frequently — especially the more distal ones —

spreading or reflexed, not bearing branchlets, similar to the stems. Cladodes terete,

straight or slightly arcuate, up to 30 mm long, up to 5-nate laterally but usually 2- or

3-nate terminally (rarely up to 8-nate terminally), the outer layers often lost from the

distal parts of the cladodes after maturity. Spines developed only on the larger branches

and the stems, small, reflexed, straight, up to 3 mm long, pale coloured. Peduncles often

terminal and up to 4-nate or rarely more, axillarily only 2- or 4-nate, 2-6 mm long,

articulated near or below the centre. Perianth segments entire, similar, oblong to

oblong-obovate, greenish white to yellowish white, 2 -5-3-0 mm long. Stamens slightly

shorter than the perianth segments; anthers orange. Style about 1 mm long, shortly

divided; ovary about 1 mm long. Berry red, up to 3-seeded, about 4 mm diameter.

This species is found in two areas with rather different climates. In the north-west

it occurs over most of the drier areas of South West Africa, Bechuanaland and the

north-west Cape. In the south-east it is confined to Basutoland and the neighbouring

areas of the Cape and Orange Free State. Flowers have been recorded from October

to April in the south-east, but are apparently mainly restricted to the second half of

this period in the north-west.

53

Cape. — Aliwal North: Elandshoek, Bolus 151. Cathcart: Cathcart, Comins 1741. Gordonia:

Aughrabies Falls, Leipoldt 4437 (BOL). Herschel : Majuba Nek near Sterkspruit, Hepburn s.n. (GRA).

Queenstown: Long Hill, Galpin 2092 (BOL). Sterkstroom: Stormberg, Drege 3533 (BOL). Wode-

house: Buffelsfontein, Stretton 66.

Basutoland. — Leribe, mountain slopes, Dieterlen 828; Mafeteng, Tatai River, Dieterlen s.n.;

Mamathes, Jacot Guillarmod s.n.

Orange Free State. — Ficksburg: Westburg, Galpin 13881. Paul Roux: Rexford, Acocks 13174.

South West Africa. — Bethanie: Kuibis, Range 854 (SAM). Gibeon: Haribes, Volk 12168 (M).

Karibib: road to Roten Kuppe, Kinges 3184 (M). Keetmanshoop : 3-6 miles from Aroab on road

to Koes, De Winter 3457. Kaokoveld: Otjitundua, Barnard s.n. (SAM). Lude itz: Weissenborn,

Kinges 2404. Maltahohe: Duwisib, Volk 12482 (M). Okahandja: Osondu, Walter 1364 (M). Ovambo-

land: Odonga-Nukuambi, Rautanen 105 (Z). Swakopmund: Walvis Bay, Hardy & De Winter 1442.

Kunth based his description of this species on Drege 3533. The specimen in the

Berlin Herbarium has been destroyed and, as the specimen bearing this number in

the Bolus Herbarium matches Kunth’s description, it has been selected as a lectotype.

Asparagus denudatus was characterised mainly by the absence of cladodes, but

several specimens have been found in which a few cladodes are present. In all collec-

tions which include both flowering and sterile material, the sterile specimens invariably

have cladodes. This shows that the absence of cladodes is only a seasonal phenomenon.

The surface characters of the stems and the appearance of the spines are extremely

characteristic both of the typical material of A. denudatus , which was originally described

from the south-eastern Cape, and of typical A. namaensis, which was described from

South West Africa. These two species were separated from one another on the presence

or absence of cladodes. As it has been shown that this is not as significant a character

as previously thought, it is more satisfactory to combine these species.

8. A. virgatus Bak. in Saund. Ref. Bot. t. 214 (1870); Journ. Linn. Soc. Bot.

14: 606 (1875); FI. Cap. 6: 259 (1896). Type: Somerset East, Boschberg, Burchell

3197 (K, lecto.!).

Roots fibrous. Stems erect to 1 m, straight, glabrous, smooth, slightly herbaceous.

Branches similar to the stems, ascending, solitary, not bearing branchlets. Cladodes

up to 7-nate, but frequently 3-nate, up to 25 mm long, usually straight. Spines only

represented by a blunt spur, never pungent. Peduncles terminal and axillary, up to

6-nate, 5-9 mm long, usually articulated below the centre. Perianth segments oblong-

obovate, similar, entire, 3-4 mm long, pale yellowish green. Stamens slightly shorter

than the perianth segments. Style shortly divided, 1 mm long; ovary sometimes slightly

more than 1 mm long; cells 3- or 4-ovuled. Berry globose, 1-seeded, smooth or

wrinkled, about 5 mm diameter, red.

This species occurs in most parts of the wetter areas of the eastern half of South

Africa. It is particularly frequent on the margins of forests, but also grows in other

shady places. It also occurs in Rhodesia and Mozambique. Flowers have been

recorded mainly from September to January, but also in April.

Cape. — Aliwal North: Elands Hoek, Bolus 205 (BOL). Barkly East: Barkly East, Gerstner 180.

Cathcart: Inverthorn, Barker 3513 (NBG). East London: East London, Rattray s.n. (GRA). Kentani:

Kobongaba River, Pegler 1462. King Williams Town : King Williams Town, Sim 1050 (BOL). Komga:

Komga, Flanagan 1292. Mount Currie: Kokstad, Mogg 1805. Somerset East: Zuurberg, Holland

199. Stockenstrom: Katberg, Ecklon & Zeyher s.n. (BOL). Stutterheim: Donga Range, Acocks

9336. Uitenhage: Uitenhage, Schmutz s.n. (BOL).

Basutoland. — Maseru, Compton 22534 (NBG).

Natal.- — Bergville: Cathedral Peak, Esterhuysen 12878. Camperdown: Cato Ridge, Fairall 147

(NBG). Durban: Isipingo, Schlechter 2993 (BOL). Eshowe: near Reservoir, Lawn 554 (NH).

Estcourt: Estcourt, West 1334. Hlabisa: Hluhluwe Game Reserve, Ward 2283. Inanda: Inanda,

Wood 1373 (NH). Klip River: Van Reenen, Franks s.n. (NH). Lions River: Howick Falls, Barker

3567 (NBG). Ngwavuma: Lebombo Mountains, Gwalaweni Forest, Strey 4692. Nongoma: 11

miles N. of Nongoma, Jessop 672. Pietermaritzburg: Oribi, Lawson 641 (NH). Ubombo: near

turning to Mkuze Game Reserve, Jessop 667. Utrecht: Utrecht, Thode A1297.

54

Swaziland. — Mbabane, Fonteyn, Compton 25980 (NBG); Isateki Beacon, Compton 27313.

Transvaal. — Barberton: Barberton, Galpin 1155. Belfast: Belfast, Bruce 480. Brits: Jacksons-

tuin. Van Vuuren 211. Ermelo: Breyten, Burtt Davy s.n. Letaba: Duivelskloof, Scheepers 328.

Lydenburg: 4^ miles N. of Branddraai, Codd 6713. Marico: Zeerust, Leendertz 11324. Middelburg:

near Botsabelo, Marloth 1 1751. Pietersburg: Magoebaskloof, Papendorf 230. Pilgrim’s Rest: Nooit-

gedacht. Young A656. Potgietersrus: Naboomspruit, Galpin 343. Pretoria: Wonderboom Reserve,

Repton 2015. Rustenburg: Kloof, Lanham s.n. Soutpansberg: Soutpansberg, Burtt Davy 2654.

Wakkerstroom : Oshoek, Devenish 407. Warmbad: near Warmbad, Acocks & Hafstrom 196. Water-

berg: Zandspruit near Sandrivierspoort, Forssman s.n.

An illustration accompanies the type description. There is also a reference to

the manuscript name, A. sylxaticus Burch. There is a specimen in the Kew herbarium,

collected by Burchell on the Boschberg, bearing a label with the name A. sylvaticus

on it. As this specimen matches both Baker’s description and the figure, and it is

extremely likely that this is the source of the manuscript name to which Baker referred,

it is necessary to choose, in the absence of any other specimen, between this specimen

and the illustration for the lectotype of this species. The specimen is preferred.

De Winter 3909 from Okavongo (M, PRE) has been regarded as belonging to

this species, but has rather large thorns, and rather woody stems and branches. Its

true identity is obscure.

9. A. fasciculatus Thunb., FI. Cap. ed. 2: 329 (1818). Type: Cape without precise

locality, Masson s.n. (UPS, holo.!; BOL!, photos).

Asparagopsis consanguinea Kunth, Enum. PI. 5: 76 (1850). Type: Cape without

precise locality, Drege s.n. (7KIEL!, ?K!).

Asparagus consanguineus (Kunth) Bak. in Journ. Linn. Soc. Bot. 14: 615 (1875);

FI. Cap. 6: 260 (1896).

Roots slightly thicker than usual for this section, but not tuberous. Stems prostrate

or weakly climbing to about 1 m, glabrous, softly woody, usually light brown or less

often green, especially when young, to dark brown, zigzagging at the nodes. Branches

solitary, usually slightly reflexed; final branches not well enough defined to be regarded

as branchlets, very narrow, often consisting of a single internode with a fascicle of

cladodes at the base, or rarely with more than two or three internodes. Cladodes

often extremely fine, linear, arcuate, numerous in a fascicle, 15-30-nate or more, up

to 7 mm long, but often only about 4-5 mm long; the cladodes in each fascicle usually

not overlapping those of adjacent nodes. Spines may be present on the lower parts

of the main stems, but rarely exceeding 2-3 mm in length. Peduncles axillary or

terminal, 1-3-nate, variable in length, up to 5 mm long, articulated at the base of the

flower. Perianth segments oblong, reflexed, about 4 mm long, with a broad brownish

streak. Stamens as long as the perianth segments, with a spur at the base; anthers

yellow, about 0-5 mm long. Style about 2 mm long. Berry yellow, fleshy, globose,

wrinkled, normally 3-seeded.

This species occurs in the south-western Karroo, as well as parts of the western

Cape. It is frequent in the coastal vegetation of the Sand Veld, but has not been

recorded further south than Milnerton. Flowers have been recorded from March

to June.

Cape. — Calvinia: 5-) miles S. of Loeriesfontein, Acocks 14414. Cape: near Blaauwberg, Oliver s.n,

(BOL). Ceres: Muishond River, Esterhuysen 1805. Clanwilliam: Clanwilliam, Zeyher 1675 (STE).

Hopefield: Langebaan, Jessop 70 (BOL). Laingsburg: Whitehill, Compton 14645 (NBG). Malmes-

bury: S. side of Paardeberg, Van Niekerk 358 (BOL). Namakwaland: Witbank, Pi/tans 5113 (BOL).

Tulbagh: Saron, Schlechter 7890. Vanrhynsdorp: Bitterfontein, Ecklon & Zeyher 1675.

There appears io be no doubt from the protologue that this was the species which

Thunberg was describing. There are three specimens labelled A. fasciculatus in the

Thunberg Herbarium. Of these two do not match Thunberg’s description as they

55

have well developed spines. The third, however, matches in every character. This

specimen, on which is written: “ e. Cap. b. Spei. Mason”, belongs to A. fasiculatm

as contrued here. The collector was probably Francis Masson, who was in South

Africa at the same time as Thunberg.

Kunth based his species on a Drege specimen, but he did not give either the locality

or collecting number. In his description, he mentioned the apical articulation of the

peduncle, which is the best character of this species, the climbing habit, the lack of

spines, and the numerous, fascicled cladodes. Two specimens collected at the Cape

by Drege, and showing the above characters, have been examined by the author. The

specimen at Kiel lacks a number. As Kunth mentioned in his Enumeratio Plantarum

other specimens which are now in the Kiel Herbarium, it seems possible that this was

the specimen on which Kunth based his species. This specimen also lacks a locality.

A specimen in the Kew Herbarium was collected between the Kroin River and Petrus-

fontein on the Piketberg. It also lacks a number and may be an isotype.

In several respects A. fasciculatus does not seem to belong to the section Africani

but rather to the section Crispi. It has numerous cladodes, fibrous roots and few-

seeded fruits, characteristic of the section Africani, but the spurred anther filaments

and herbaceous habit would refer it to the section Crispi. On balance, it seems to be

best placed in the section Africani.

10. A. thunbergianus Schult. in R. & S., Syst. Veg. 7: 333 (1829); Bak. in Journ.

Linn. Soc. Bot. 14: 615 (1875); FI. Cap. 6: 262 (1896); Salter in FI. Cape Penins.

175 (1950). Type: Cape Peninsula, Thunberg s.n. (UPS, holo. ! ; BOL!, PRE!, photos).

Asparagus albus sensu Thunb., Prodr. 66 (1794). A. capensis sensu Marloth, FI.

S. Afr. 4: pi. 20 (1915).

Asparagopsis dregii Kunth, Enum. PI. 5: 84 (1850). Type: Cape of Good Hope.

Drege s.n. (?K, iso.!). A. thunbergii Kunth, Enum. PI. 5: 85 (1850), nom. illegit,

Type: as for A. thunbergianus. A. niveniana sensu Kunth, Enum. PI. 5: 88 (1850).

Roots fibrous. Stems erect to 1 m, or forming a dense bush up to 150 cm high

and 200 cm broad, in dense shade sometimes climbing to 150 cm, pale to dark brown,

glabrous, zigzagging. Branches glabrous, spreading or ascending, solitary, similar to

the stems. Cladodes fascicled, terete, slightly arcuate, 3-8 mm long, rarely up to

20 mm, ascending or spreading. Peduncles axillary or terminal, usually solitary, 3-6

mm long, articulated below the centre. Perianth segments spreading, white, obovate,

2-3 mm long. Stamens nearly as long as the perianth segments; anthers brown,

0-5 mm long. Styles about 1 mm long; ovary about 1 mm long. Berry globose,

up to 8 mm diameter, with a single seed, red.

This species occurs, usually in low bush, from the Cape Peninsula to Clanwilliam

in the north and Uitenhage in the east. Flowers have been recorded from January

to June.

Cape. — Bellville: Tigerberg, Pillans 8658 (BOL). Bredasdorp: Nachtwacht, Smith 3022A. Caledon:

Hermanus, Rogers 26572. Cape: above Camps Bay, Marloth 9492. Clanwilliam: Clanwilliam,

Leipoldt 408 (SAM). George: Montagu Pass near Herold, Jessop 592. Hopefield: between Hopefield

and Paternoster, Leipoldt 3895 (BOL). Humansdorp: along river, Britten 1 162. Knysna: Belvidere,

Duthie 763 (BOL). Malmesbury: Darling Flora Reserve, Rycroft \9[ 1 (NBG). Paarl: Dal Josaphat,

Tyson 858 (BOL). Piketberg: Het Kruis, Stephens & Glover s.n. (SAM). Riversdale: Albertinia

commonage, Muir 931. Simonstown: W. of Simonstown, Wolley Dod 933 (BOL). Somerset West:

Knorhoek, Jessop 77 (BOL). Stellenbosch: Stellenbosch, De Jongh s.n. Swellendam: Grootvadersbos,

Taylor 1042 (NBG). Tulbagh: Saron, Schiechter 7887. Uitenhage: Theescomb, Paterson 853 (BOL).

Wellington: Wellington, Thomson 153. Wynberg: Kirstenbosch, Middlemost 2008 (NBG).

Thunberg included Asparagus albus L. in his Prodromus plantarum Capensium.

In his herbarium, there is a sheet labelled “ Asparagus albus , p. Withering ", On this

sheet there is a specimen of A. thunbergianus Schult., but there is also a specimen of

56

another species — perhaps A. albus L. from Europe. The specimen of A. thunbergianus

is of a young shoot, but it shows the reflexed, solitary spines, smooth (somewhat

wrinkled during drying) stem and general appearance of a young shoot of A. thun-

bergianus. The description, however, applies equally well to either specimen. It

seems probable that Thunberg did not realise that there were specimens from different

countries on this sheet, but recognised the European specimen as A. albus L.

Schultes described A. thunbergianus as being from the Cape Peninsula. He

described it as a shrub up to two feet or more high with grooved stems and with solitary,

recurved spines and fascicled cladodes. He cited Thunberg’s A. albus L. as a synonym,

and as no other specimen on which he could have based his species is known to have

existed the Thunberg specimen may be regarded as the holotype.

Asparagopsis dregii was described by Kunth as having been collected at the Cape

of Good Hope by Drege, but this specimen was destroyed in Berlin. The branches

were described as smooth, glabrous and spinous. He regarded it as very similar to

Asparagopsis thunbergii (based on Asparagus thunbergianus Schult.) but the description

makes it very likely that these names were based on specimens of the same species.

A specimen in Kew was collected by Drege but, like the specimen which Kunth had,

had no number. It is labelled “ Table Mountain and Devils Mountain. Asparagopsis

dregei Kth.”. It seems very likely that this is an isotype.

Asparagus nivenianus Schult. was based on a specimen collected by Niven at the

Cape. This specimen has not been located and it is not possible to tell from the

description whether this was a specimen of A. thunbergianus or of A. africanus. This

name must therefore be regarded as a nomen dubium. Kunth transferred the epithet

to Asparagopsis and cited Ecklon 95b from the Cape as the only specimen he had seen

of Asparagopsis niveniana. A specimen of this collecting in the Munich Herbarium is

of A. thunbergianus.

11. A. mucronatus Jessop, sp. nov., A. thunbergiano Schult. afltnis, sed caulibus

ramisque maioribus, bracteis manifesto evolutis et ad basim ramulorum persistentibus

differt. Plate 1.

Radices non tuberosi. Caules recti ad 2 m, plerumque c. 1 m, basi bracteis

manifestissime evolutis ad 2 cm longis, glabri, costis magnis, basi recti, supra tortuosi,

cani vel fusci. Rami caulium partibus distalibus similes, rami minores nonnumquam

pubescentes vel scabridi, Ramuli fasciculati, ad 6-nati, pubescentes vel scabridi, recti

ad 6 cm longi; ramuli juvenes saepe bracteis manifesto evolutis et basi persistentibus

ad 5 mm longis. Cladodia recta vel arcuata, teretes, mucrone non-spinoso et non-

colorato, 3-8 mm longa, in ramulis orta, ad 10-nata. Spinae magnae, recurvae.

Pedunculi in ramulis axillares, 1-vel 2-nati, 4-7 mm longi, submedio articulati. Perianthii

segmenta oblongo-obovata, 3-4 mm longa, margine integro, similia. Stamina quam

perianthii segmenta paulum breviora; antherae £ mm longae, subglobosae. Ovarium

fere 1 mm longum; stylus breviter ramosus, c. 1 mm longus. Bacca globosa, succulenta,

deinde rugosa, rubra, fere 5 mm diam., seminibus 1 vel saepe 2.

Type: Cape: Middelburg district, Grootfontein, Theron 551 (PRE, holo.).

Roots fibrous. Stems erect to 2 m, usually about 1 m high, bearing very pronounced,

acute bracts up to 2 cm long at the base, glabrous, smooth, with large ridges, straight

at the base, becoming slightly zigzag, grey to brown. Branches similar to the distal

parts of the stems, the younger ones sometimes pubescent or scabrid. Branchlets

fascicled, up to 6-nate, pubescent or scabrid, straight, up to 6 cm long, often bearing

well-developed bracts up to 5 mm long when young and persisting towards the base.

As the stems are at least biennial, at times new and old branchlets occur simultaneously,

in which case the younger ones are paler, more pubescent, and bearing larger bracts,

and possibly peduncles. Cladodes straight or arcuate, terete, with a non-pungent,

57

colourless mucro, 3-8 mm long, borne on the branchlets, up to 10-nate. Peduncles

axillary on the branchlets, 1- or 2-nate, 4-7 mm long, articulated below the middle.

Perianth segments oblong-obovate, 3-4 mm long, entire, similar. Stamens slightly

shorter than the perianth segments; anthers £ mm long, rather globose. Ovary

about 1 mm long; style shortly divided, slightly less than 1 mm long. Berry globose,

fleshy, wrinkled when ripe, about 5 mm diameter, with a single or occasionally two

seeds, red.

This species is fairly common in at least some parts of the Karroo and Little

Karroo, but also occurs as far east as the Cathcart district. Flowers have been recorded

from October to December.

Cape. — Albany: Queens Road, Britten 5159 (GRA); Botha Hill, Dyer 1124 (GRA); Plutos Vale,

Jessop 637. Calvinia: Calvinia, Schmidt 358. Cathcart: about 5 miles N. of Cathcart, Jessop 661.

Graaff-Reinet : Graaff-Reinet, Henrici 4943. Hay: Blaauwbosput, Acocks 2011. Herbert: Valsch-

fontein, Acocks 2598; Heuningneskloof, Moran s.n. (BOL). Kimberley: Mauretzfontein, Acocks

2580; Acocks 4957 (BOL). Laingsburg: Matjiesfontein, Foley 62. Middelburg: Grootfontein,

Theron 551 (BOL). Montagu: between Dobbelaarskloof and Montagu, Esterhuysen 1872; 10 miles

N.E. of Montagu, Jessop 678. Port Elizabeth : Redhouse, Denman 282; Paterson 960 ( BOL) ; Paterson

977 (BOL). Vanrhynsdorp: Langeberge, Marloth 12953. Worcester; Veld Reserve, Olivier 276.

The greatest similarity is with A. thunbergianus. It is, however, a much larger

and more robust plant. It also differs in having scale leaves, and pubescence on the

final branches.

12. A. macowanii Bak. in Journ. Linn. Soc. Bot. 14: 609 (1875). Type: Albany,

Zeyher 879 (K, lecto.!; BOL!).

Roots fibrous. Stems erect, softly woody, glabrous, smooth or slightly grooved,

usually white, up to about 2 m high. Branches similar to the stems, solitary. Cladodes

fascicled, numerous, terete, arcuate, up to about 12 mm long. Spines not well-

developed, absent from final branches. Peduncles terminal or axillary, sometimes

solitary but usually up to 1 2-nate, articulated proximally. Perianth segments oblong

to oblong-obovate, 2-5 mm long, pale. Stamens slightly shorter than the perianth

segments; anthers less than 0-25 mm long; ovary 1-5-2 mm long, ovoid. Fruit not

known.

Southern Cape, eastwards through Natal, and in the eastern districts of the

Transvaal. Also in Mozambique.

For key to varieties see key to species.

(a) var. macowanii.

A. macowanii Bak. in Journ. Linn. Soc. Bot. 14: 609 (1875); FI. Cap. 6: 261

(1896).

The typical variety is characterised by its smaller size. The stems are rarely over

75 cm high. The stems are sometimes grooved at first, but soon become smooth.

This variety occurs in the southern Cape from Knysna to Komga, and inland

as far as Glen Grey. Flowers have been recorded from September to November and

in April and May.

Cape. — Albany: Howieson’s Poort, Zeyher 879 (BOL, K). Bathurst: Horseshoe on the Kariega

River, Jessop 616. Bedford: Bedford, Gone 88 (GRA). Glen Grey: White Kei Falls, Galpin 2504.

Knysna: Plettenbergbaai, Rogers 26141. Komga: Prospect Farm, Flanagan 296. Somerset East:

Somerset East, MacOwan s.n. (BOL, SAM); Rogers 161; 164 (GRA); Boschberg. Tyson 1878.

Baker, in his type description, cited Zeyher 879 from Albany and MacOwan 1917

from Somerset East. In the Flora Capensis, however, he removed the MacOwan

specimen to A. declinatus. It therefore seems best to regard the Zeyher specimen as

58

the lectotype. There are specimens of a MacOwan collecting of this species in the

Bolus and South African Museum herbaria. This collecting is not numbered but was

also collected in the Somerset East district. It seems possible that these are of the

same collecting as the specimen seen by Baker when he drew up the original description.

It would otherwise be hard to explain the epithet.

(. b ) var. zuluensis (N. E. Br.) Jessop, stat. nov.

A. zuluensis N. E. Br. in Kew Bull. 1912: 283 (1912). Type: Zululand, Wylie sub

Medley Wood 11969 (K, holo.!; NH!, PRE!).

This variety differs from the typical variety in being a much larger plant. The

stems are much-branched and up to 2 m high. The stems and branches are always

smooth and whitish. Spines are developed, although not large or pungent, on the

main stems and branches only.

Occurs from the Camperdown district in the south, through Zululand and into the

eastern Transvaal as far north as the Pilgrim’s Rest district. Flowers have been recorded

from September to November.

Natal. — Camperdown: Cato Ridge, McC/ean & Ogilvie s.n. (NH). Eshowe: Umlalazi Bridge,

Lawn 750 (NH). Hlabisa: Hluhluwe Game Reserve, Ward 4334. Mahlabatini: Umfolozi Game

Reserve, Ward 4379. Mtonjaneni: Mtonjaneni, Gerstner 3665. Mtunzini: Ngoya Forest Reserve,

Huntley 268.

Transvaal. — Barberton: Barberton, Thorncroft 16. Piet Retief: 13 miles from Pongola to Gollel,

Dyer & Verdoorn 5832. Pilgrim’s Rest: Eileen Orpen Dam near Tshokwane, Codd 6138.

N. E. Brown based A. zuluensis on a single collecting from Zululand by Wylie.

Medley Wood made the herbarium specimens which he distributed from a specimen

growing in the Durban Botanic Garden. A plant which is very likely the original

Wylie specimen is still growing in these gardens near the herbarium.

The differences between this and the type variety are not usually distinguishable

from herbarium material unless a note on the habit accompanies the specimen. The

distribution of the two varieties, however, makes it possible to identify with certainty

material of which the origin is known.

13. A. retrofractus L., Sp. PI. 313 (1753); Bresler, Diss. 10 (1826); R. & S., Syst.

Yeg. 7: 330 (1829); Bak. in Journ. Linn. Soc. Bot. 14: 621 (1875); FI. Cap. 6: 266

(1896); Salter in FI. Cape Penins. 176 (1950). Type: without locality or collector

(LINN No. 434.4, holo.; NBG!, PRE!, photos).

Asparagopsis retrofracta (L.) Kunth, Enum. PI. 5: 78 (1850).

Asparagus africanus Lam. var. microphyllus Bak. in FI. Cap. 6: 266 (1896). Type:

Rumman, Kosi Fontein, Burchell 2572 (K, holo.; PRE, photo.!). A. microphyllus

(Bak.) Kies in Bothalia 6: 178 (1951).

Roots fibrous. Stems scrambling or weakly climbing to 2 m or more, or forming

dense bushes up to 2 m or more high, white and grooved at first, but becoming brown

and smooth, zigzagging; swollen lateral buds are present at the nodes. Branches

similar to the stems, spreading or reflexed, not bearing branchlets, pubescent when

young. Cladodes fascicled, 6-30 mm long, arcuate, terete, spreading or ascending,

not all the same length. Spines sometimes present, strongly reflexed, occasionally

pungent, up to 6 mm long, never borne on the final branches. Peduncles axillary or

terminal, usually 2-6-nate, 5-40 mm long, articulated below the centre. Perianth

segments white with a green streak, narrow obovate, spreading, 3 mm long, margins

entire, similar. Stamens almost as long as the perianth segments; anthers orange.

Style divided for about half its length, 1 mm long; ovary slightly longer than the style.

Berry globose, 1-3-seeded, 5 mm diameter, orange; perianth generally persistent.

59

This species occurs mainly in the drier areas of South West Africa and the Karroo*

but extends into the winter-rainfall area as far as the Cape Peninsula. Flowers have

been recorded mainly from April to June, but also at other times.

Cape. — Barkly West: Klipvlei, Esterhuysen 2083 (BOL). Beaufort West: Sunnyside, Esterhuysen

4357. Bellville: Tigerberg, Pil/ans 4758 (BOL). Caledon: 12 miles west of Caledon, Salter 7215

(BOL). Calvinia: Calvinia, Schmidt 556. Cape Town: Porcupine Buttress, Pil/ans 4327 (BOL).

Carnarvon: 12 miles from Carnarvon on Van Wyks Vlei road, Acocks 1741. Ceres: Karroopoort,

Marloth 9008. Clanwilliam: Modderfontein, Gillett 3695 (BOL). Gordonia: Aughrabies Falls,

Jessop 351 (BOL). Graaff-Reinet : 14 miles north of Graaff-Reinet, Jessop 651. Herbert: Campbell

Kloof, Wilman 4069 (BOL). Laingsburg: Whitehall, Compton 8523 (NBG). Malmesbury: near

Darling, Bolus 12859 (BOL). Montagu: Cogman’s Kloof, Barker 959 (NBG). Namakwaland: Honde-

klip Bay, Pil/ans s.n. (BOL). Oudtshoorn: road to Armoed via Baakens Kraal, Britten 1706. Prieska:

Prieska district, Bryant J224. Stellenbosch: Papagaaisberg, Duthie 452 (STE). Vanrhynsdorp: Heeren-

logementsberg, Zeyher 1674 (BOL). Vryburg: Vryburg, Mogg 8976. Worcester: Welgevonden,

Esterhuysen 1903.

South West Africa. — Gibeon: Harris, Fleck 296a (Z). Keetmanshoop: Swartkop, Ortendahl 83.

Ludertiz: Aus, Peyer 15 (Z). Maltahohe: Duwisib, Volk 126363 (M). Warmbad: Kanus, Range

907 (SAM). Windhoek: Windhoek, Rogers 30480 (Z).

Linnaeus, in his type description, cited Plukenet, Phyt. t.375 f.3. This figure

shows the typical characters of A. retrofractus L. as construed here. There is a specimen

in the Linnaean Herbarium labelled A. retrofractus and probably received before

1753 (see Jackson, 1912). This specimen is therefore likely to be the holotype.

This species is replaced to the east by A. laricinus which, particularly on herbarium

material, is sometimes difficult to distinguish. Usually a less regular arrangement of

the cladodes distinguishes A. retrofractus, but the typically ascending arrangement

of the cladodes is not always apparent in A. laricinus, which can then be identified

by having small spines on the final branches, and by its more eastern distribution.

In the field, A. retrofractus can often be recognised as a climber or scrambler, while

A. laricinus is a large bush.

14. A. compactus Salter in Journ. S. Afr. Bot. 6: 165 (1940); FI. Cape Penins.

175 (1950); Kies in Bothalia 6: 179 (1951). Type: Cape Peninsula, Orange Kloof,

Wolley Dod 2521 (BOL, holo.!).

A. africanus Lam. var. wrightii Bak. in Journ. Linn. Soc. Bot. 14: 619 (1875); FI.

Cap. 6: 266 (1896). Type: Simons Bay, Wright 454.

Roots fibrous. Stems erect, normally about 50 cm long, zigzagging, woody,

white, minutely pubescent, grooved; outer layers sometimes lost leaving the stems

brown and smooth. Branches spreading to ascending, similar to the stems. Cladodes

fascicled, commonly up to 10-nate terminally, and fewer laterally, more or less straight,

ascending to spreading, 3-angled, rarely up to 1 mm thick, rigid, 10-30 mm long.

Spines pungent, spreading, recurved or reflexed, present on the aerial stems and all

branches. Peduncles axillary or terminal, 1- or 2-nate laterally, and up to 4- or 5-nate

terminally, 5-7 mm long, articulated below the centre. Perianth segments obovate,

similar, entire, white, about 3 mm long. Stamens slightly shorter than the perianth

segments, not spurred; anthers about f mm long. Style divided for nearly half

its length, about 1 mm long. Berry globose, fleshy, red, about 6 mm diameter, 1-3-

seeded.

This species is confined to the south-western Cape, occurring from near sea-level

to 3,000 ft or more on well-drained mountain slopes. Flowers have been recorded

in October, December, January and from March to May.

Cape. — Bellville: south side of Tigerberg, Pil/ans 8655 (BOL). Caledon: Swartberg near Caledon,

Schlechter 10364. Cape: Paarden F.iland, Salter 8268. Ceres: Milner Peak, Esterhuysen 7804 (BOL).

Malmesbury: near Kalabas Kraal, Salter 7198 (BOL). Paarl: above Franschhoek, Jessop 102 (BOL).

Piketberg: Porterville, Loubser 810 (BOL). Simonstown: Clovelly, Salter 8209 (BOL). Somerset

60

West: Knorhoek, Jessop 89 (BOL). Stellenbosch: Swartboskloof, Van Rensburg 268 (STE). Suther-

land: near Sutherland, Du Toit s.n. (BOL). Tulbagh: Sneeuwgat Valley, Great Winterhoek, Phillips

1880 (SAM). Wellington: Bains Kloof, Salter 7307 (BOL). Wynberg: Orange Kloof, Wolley Dod

2521 (BOL).

The holotype is cited by Salter as Wolley Dod 2521 (BOL). Salter regarded this

species as very probably identical with A. africanus var. wrightii Bak., which was based

on Wright 454. Baker’s description mentioned grooved branches. The cladodes

were given as very stout, angled, with a groove down each face, and the berries as

being larger than for the typical variety. This is not adequate to identify the species

with certainty, but with the locality, given as Simonstown, it is almost certainly this

species. According to Salter (1940), Professor Compton searched for the type in

Europe, but was unable to find it.

15. A. laricinus Burch., Trav. 1: 537 (1822); Bresler, Diss. 40 (1826); R. & S.,

Syst. Veg. 7: 337 (1829); Kunth, Enum. PI. 5: 75 (1850); Bak. in Journ. Linn. Soc.

Bot. 14:^620 (1875); FI. Cap. 6: 267 (1896); Solch, Beitr. FI. SUdwest-Afr. 37 (1961).

Type: Hay, between Griquatown and Wittewater, Burchell 1971 (K, holo.!; PRE,

photo. !).

Asparagus africanus Lam. var. concinnus Bak. in Journ. Linn. Soc. Bot. 14: 619

(1875); FI. Cap. 6: 266 (1896). Type: Pearston, Bruintjes Hoogte, Burchell 3020

(K, holo.!; PRE, photo.!). A. concinnus (Bak.) Kies in Bothalia 6: 178 (1951).

Root'> fibrous. Stems erect, forming much-branched shrubs up to 2 m high, but

usually only about 150 cm or less, often zigzagging or tortuous, woody, white, pubescent

and usually grooved when young, but soon losing the outer layers and becoming smooth

and darker. Branches similar, but retaining the grooved outer layers longer. Cladodes

fascicled, 15-60-nate laterally, more numerous terminally, more or less straight,

ascending, terete, fine but rigid, all approximately the same length at each node, 8-30

mm long. Spines pungent, ascending to recurved, usually present on all branches.

Peduncles axillary or terminal, commonly up to 7 or 8 terminally and 3 or 4 laterally,

up to 7 mm long, articulated below the centre. Perianth segments obovate, similar,

entire, white, 2-5-3 mm long. Stamens slightly shorter than the perianth segments,

not spurred; anthers about f mm long. Style divided for nearly half its length,

about 1 mm long. Berry globose, about 6 mm diameter, fleshy, red, 1-3-seeded.

This species has a wide distribution from the south-eastern Cape to South West

Africa, Bechuanaland and the Transvaal. Flowers have been recorded from September

to December.

Cape. — Albany: Mountain Drive, Grahamstown, Britten 2805 (GRA). Aliwal North: Elandshoek,

Bolus 204 (BOL). BarklyWest: Daniels Kuil, Esterhuysen 1153 (BOL). Cathcart: Fairford, Cotterell

126 (GRA). Fort Beaufort: Fort Beaufort, Zey/ier s.n. Flay: between Wittewater and Griquatown,

Burchell 1971 (K). Herbert: Heuningneskloof Station, Moran s.n. (BOL). Kimberley: Kimberley,

Moran s.n. (BOL). Mafeking: Mafeking, Burtt-Davy s.n. Mount Currie: Kokstad, Mogg 1806.

Port Elizabeth: Swartkops Estuary, Archibald 4914 (GRA). Port St. Johns: Eagles Nest, Schonland

4077. Somerset East: Somerset East, Bowker s.n. (BOL). Sterkstroom: Andriesberg, Galpin 2168.

Vryburg: Armoedsvlakte, Mogg 7922.

Batutoland. — Leribe, Dieterlen 406.

Natal. — Mooi River: Warley Common, Mogg 7152. Mtonjaneni: Melmoth, Lawn 1828 (NH).

Mtunzini: Ginginhlovu, Wylie s.n. (NH). Umzinto: Dumisa, Rudatis 1735.

Orange Free State. — Bloemfontein: Leeuwkop, Potts s.n. (BOL). Fauresmith: Veld Reserve,

Henrici 395 1 . Harrismith: Rensburgskop, Jacobsz 103. Kroonstad: Kroonstad, Pont 485. Senekal :

Doornkop, Goossens 649. Thaba Nchu: The Willows, Roberts 2992. Vredefort: near Vredefort,

Pole Evans 19474H. Wepener: sandstone hillside, Acocks 1172. Winburg: W. Pretorius Game

Reserve, Bakker 1006.

Transvaal. — Barberton: Malelane, Murphy s.n. Belfast: Dullstroom, Codd 5638. Bethal: Steen-

kool River, Weintraub s.n. Ermelo: Nooitgedacht, Henrici 1128. Germiston: Wattles, Moss 10503.

Heidelberg: Suikerbosrand, Mogg 18133. Johannesburg: near Johannesburg, Watt cfc Brandwyk

2207. Kroonstad: Kroonstad, Pont 485. Krugersdorp: 30 miles W.N.W. of Johannesburg, Mogg

61

24235. Lichtenburg: Witstinkhoutboom, Liebenberg 44. Lydenburg: Steenkampsberg, Young

A428. Marico: near Zeerust, Van der Merwe 128 (BOL). Pilgrim’s Rest: between Pilgrim’s Rest and

Sabie, Rogers 20564. Potchefstroom: Potchefstroom, Burtt-Davy 48. Pretoria: Irene. Rogers 11406

(BOL). Vereeniging: Vereeniging, Leendertz s.n. Wakkerstroom : Oshoek, Devenish 1157. Warmbad:

Warmbad, Leendertz s.n. Wolmaransstad: Boskuil, Sutton 56.

South West Africa. — Gobabis: Okasondana, Walter 4163 (M). Windhoek: Voigtland, Walter

116 (S.W.A. Herb.).

Bechuanaland. — Gaberones, Rogers 6254; Pharing, Kanye, Hillary & Robertson 466.

Burchell cited only his C.G. 1971 from between Griquatown and Wittewater.

This specimen, in the Kew Herbarium, is therefore taken as the holotype.

In some areas, particularly to the south and east of its distribution, plants occur

with rather small cladcdes, but otherwise the normal characters for this species. A

specimen of this form. Burchell 3020 from Bruintjes Hoogte, was described by Baker

as A. africanus var. concinmis. Kies raised this variety to specific status, but in the

present author’s opinion there are no grounds for giving this form taxonomic recog-

nition.

This species is similar in many respects to both A. retrofractus L. and A. macouanii

Bak. var. zuluensis (N. E. Br.) Jessop. The characters by which it can be distinguished

from A. retrofractus are given under that species. The regular arrangement of the

cladodes, the usually grooved stems, the development of spines on the final branches,

and the more western distribution serve to distinguish it from A. macowanii var.

zuluensis.

16. A. microrhaphis (Kunth) Bak. in Journ. Linn. Soc. Bot. 14: 612 (1875); FI.

Cap. 6: 262 (1896). Type: Queenstown, Drege 3534 (BOL, lecto.!).

Asparagopsis microrhaphis Kunth, Enum. PI. 5: 83 (1850).

Asparagus stellatus Bak. in Journ. Linn. Soc. Bot. 14: 612 (1875); FI. Cap. 6: 262

(1896). Syntypes: “Albert Div.”, Cooper 622 (K !) ; Aliwal North, Witteberge, Drege

8589 (K!).

Roots fibrous. Stems erect to 3 m, glabrous or minutely pubescent, ashen grey,

grooved but becoming smooth and brown, softly woody, straight, much-branched.

Branches similar to the stems, aiso branched, not bearing branchlets. Cladodes fascicled,

glabrous or minutely pubescent, usually up to about 8-nate but occasionally up to

25-nate, 1-4 mm long. Spines usually well developed on aerial stems and main

branches. Peduncles axillary, 1- or 2-nate, 5-6 mm long, articulated below the centre.

Perianth segments oblong-obovate, entire, similar, spreading, white, 3-4 mm long.

Stamens almost as long as the perianth segments. Styles shortly divided, about 1 mm

long. Fruit not known.

This species occurs in bush or on grassy slopes in the eastern Cape, Basutoland

and Natal. Flowers have been recorded in October and December.

Cape. — Herschel: Herschel, Gerstner 18. Mount Ayliff: Kokstad, Mogg 1912. Queenstown:

Table Mountain, Drege 3534 (BOL). Sterkstroom: Andriesberg, Galpin 2169. Wodehouse: Buffels-

fontein, St ret ton 65.

Basutoland. — Leribe, Dieterlen 301; Mafateng, Thababa-Morena, Dieterlen 1374.

Natal. — Bergville: Ndedema River Valley, Killick 1861. Estcourt: Giants Castle, Symons 38.

Lions River: Inhleyane, Moll 1286. Wakkerstroom: 7 miles N.E. of Wakkerstroom, Acocks 11525

(NH).

Kunth, in his description of Asparagopsis microrhaphis, cited Drege 3634. The

holotype is likely to have been destroyed in Berlin, but there is a specimen with this

number in the Bolus Herbarium. As it matches Kunth's description in every detail

this specimen has been selected as a lectotype.

62

A number of specimens, such as Cooper 622 and Drege 8589, are representatives

of a group possibly deserving varietal status. They have extremely small and very

numerous cladodes. This group was described by Baker under the binomial A. stellatus.

No specimens of this vegetative form have ever been collected in flower, however,

and it seems likely that this is only a seasonal form.

17. A. exuvialis Burch., Trav. 1: 432 (1822); Bresler, Diss. 40 (1826); R. & S.,

Syst. Veg. 7: 337 (1829); Kunth, Enum. PI. 5: 74 (1850); Bak. in Journ. Linn. Soc-

Bot. 14:" 608 (1875); FI. Cap. 6: 259 (1896); Schinz in Bull. Herb. Boiss. 1, 4, app-

III: 42 (1896); Salter in FI. Cape Penins. 175 (1950); Solch, Beitr. FI. Siidwest-A.fr-

36(1961). Type: “Along the Vaal River, Griqualand West ”, Burchell 1768 (K, holo.!;

PRE, photo. !).

A. ecklonii Bak. in Journ. Linn. Soc. Bot. 14: 615 (1875); FI. Cap. 6: 258 (1896).

Type: South Africa, without locality, Ecklon & Zeyher s.n. (TCD, holo.!; PRE,

photo.!; PRE, iso.!). A. hereroensis Schinz in Bull. Herb. Boiss. 1, 4, app. Ill: 43

(1896). Syntypes: Rehoboth, Fleck 295a (Z!); Harris, Fleck 549 (Z !).

Roots tuberous; the tubers almost sessile, about 5-10 cm long. Stems erect or

twining, usually about 50 cm high, but occasionally up to about 2 m, woody, glabrous,

smooth; the outer layers of the young stems pale, and peeling to leave the stems

darker, sometimes almost black, straight or zigzagging. Branches similar to the stems,

often spreading. Cladodes fine, terete, fascicled, from 5-30 mm long, deciduous.

Spines absent or poorly developed, and reflexed; the leaf firm and deltoid, of a rather

characteristic shape. Peduncles axillary, generally paired, but up to about 5-nate,

articulated near or below the middle, 3-5 mm long; the flowers opening in a racemose

sequence along the branches. Perianth segments oblong-obovate, entire, similar,

about 3-5-4 mm long, with a brown or purplish streak, spreading. Stamens not

spurred, nearly as long as the perianth segments; anthers yellow, about f mm long.

Style shortly divided, about 1 mm long; ovary about 1 mm long. Fruit globose,

turning black, dry, smooth; perianth not persistent.

This species is generally found in the Karroo and Kalahari, but does occur in

moist areas as well. It is distributed from the south-western Cape northwards to the

northern Transvaal, Bechuanaland and South West Africa. Flowers have been recor-

ded from October to April. In the Cape Peninsula, and probably other areas, the flo-

wers open in the morning, close at mid-day, and the following morning the next flo-

wer on the branch opens.

Cape.- Barkly West: Daniels Kuil, Esterhuysen 1149 (BOL). Bell vi lie : Kanonberg, Compton

115622 (NBG). Cape: Paarden Eiland, Salter 8294 (BOL). Clanwilliam: Lange Kloof, Schlechter

8051. Cradock: W. of Rayner's Koppie, Acocks 11926. Kimberley: Dorstfontein, Acocks 1404.

Kuruman: Kaap Plateau, Marloth 14070. Malmesbury: Darling Flora Reserve, Barker 8670 (NBG).

Namakwaland: Doornpoort, Pillans 5467 (BOL). Port Elizabeth: Redhouse, Paterson 963 (BOL).

Postmasburg: Postmasburg, Esterhuysen 1153. Prieska: Draghoender, Bryant 1174. Queenstown:

Queenstown, Rogers 2321 (GRA). Tulbagh: Wolseley, Schlechter 9949. Uitenhage: Uitenhage,

Ecklon & Zeyher 607. Vanrhynsdorp: Klawer, Andreae 411. Williston: Williston Station, Smith

2445. Worcester: near De Dooms, Bolus 13201. Wynberg: Princess Vlei, Jessop 88 (BOL).

Orange Free State. — Bloemfontein: Bestersput, collector unknown, 13. Fauresmith: Voelfontein,

Kies 338. Jacobsdal: near Heuningneskloof Station, Phillips 3480. Luckhoff: Rosemarie, Kalkbult,

Henrici 4260.

Transvaal. — Pilgrim’s Rest: 20 miles W. of Satara, Codd 6197. Potgietersrus: Naboomsrpuit,

Galpin 347. Pretoria: 6 miles W. of Hammanskraal, Kies 371. Sibasa: Punda Milia, Lang s.n.

Soutpansberg: Soutpan, Obermeyer, Schweickerdt & Verdoorn 51.

South West Africa. — Grootfontein : 157 miles E. of Grootfontein, Story 6199 (M). Karibib:

Marmorwerk, Volk 2757 (M). Luderitz: Aus, Dinter 6136 (B). Okahandja: Otjihua, Dinter 329

(SAM). Otjiwarongo: Okosongomingo, Volk 2523 (M). Outjo: Kamanyab and Kaross, Thorne

s.n. (SAM). Rehoboth: Rehoboth, Fleck 295a (Z). Warmbad: Sandfontein, Wilman 1602 (SAM).

Windhoek: Voigtland, Walter 2778 (M).

Bechuanaland. — Ngami See, Schinz 900 (Z); Mochudi, Marloth 950.

63

There is a form from the north-eastern Transvaal in which the steins are more

tortuous than usual, the flowers are more than 2-nate, and the perianth segments are

yellowish (e.g. Code I 6197, Obermeyer, Schweickerdt & Verdoorn 97, Long s.n. and

Van der Schijjf 911 ; all in PRE). This form does not seem to be sufficiently different

to warrant separate taxonomic status.

18. A. oxyacanthus Bak. in Journ. Linn. Soc. Bot. 14: 625 (1875); FI. Cap. 6:

270 (1896). Type: Somerset East, Bowker s.n. (K, holo.!; PRE, photo.!).

Tubers distant, about 6 cm long. Stems often unbranched, erect, smooth, straight,

up to about 50 cm long, pubescent, with numerous narrow, softly spinous leaf spurs,

grey to brown. Branches solitary, lacking branchlets, similar to the stems. Cladodes

solitary, oblong, up to 4 cm, rarely to 8 cm, long, with a median vein and thickened

margin, mucronate. Peduncles 2 mm long, solitary in the axils of large bracts, articulated

distally. Perianth segments obovate, obtuse, white, 2-3 mm long, spreading to ascending,

rather variable in width, but the inner whorl broader. Anthers large, up to 0-7 mm

long. Style branches free, short; ovary globose. Fruit not known.

This species is restricted to the area from the Bathurst district to the East London

district. It occurs mainly on rather dry slopes, particularly the slopes above the Fish

River. Flowers have been recorded from October to January.

Cape. — Albany: Overlooking Kaffir Drift, Leach & Bayliss 12646; Fraser’s Camp, Maguire 632

(NBG). Bathurst: Fish River, Dyer 3375; Schlechter 6105 (BOL); Kleinemonde, White 952 (SAM).

East London: East London, Rattray s.n. (GRA); Green Point, Smith 3765; East London Park,

Wood s.n. Peddie: Bergha River Valley opposite Wooldridge, Acocks 11885.

This is one of the “ fox-tail ” species of the section Racemosi, but can easily be

distinguished by the broad, mucronate cladodes, and the pubescent stem.

19. A. juniperoides Engl, in Bot. Jahrb. 10: 3 (1889); Schinz in Bull. Herb.

Boiss. 1, 4, app. Ill: 44 (1896); Solch, Beitr. FI. Slidwest-Afr. 37 (1961). Type:

S.W.A., Aus, Marloth 1538 (B, holo.!).

Tubers sessile or shortly stalked, fusiform. Stems erect, glabrous, grooved, straight,

annual, up to 1 m high. Branchlets narrow, numerous, up to 3 cm long, borne directly

on the stems; branchlets and cladodes ascending, forming a dense cylinder round the

stem. Cladodes solitary, flattened, usually very narrow, linear, often with ciliate

margins, up to 1 cm long. Leaves not forming spines. Peduncles axillary, solitary,

about 2 mm long, articulated distally. Perianth segments similar, oblong, with ciliate

margins, ascending, 6 mm long, white with a green streak. Stamens nearly as long

as the perianth segments; filaments not spurred; anthers minute, yellow. Ovary

globose, 1-0-1 -5 mm long. Berry globose, with one to three seeds, about 5 mm

diameter, red.

This species occurs from Tulbagh, up the west coast into South West Africa.

Flowers have been recorded from April to July.

Cape. — Clanwilliam: Lange Kloof, Schlechter 8028; Zeyher 4167 (STE); Graafwater, Zinn s.n.

(SAM). Namakwaland: Spektakel, Esterhuysen 5867 (BOL); Kamieskroon, Lewis 2146 (SAM);

Vioolsdrif, Middlemost s.n. (NBG); Doornpoort, Pillans 5379 (BOL); Witbank, Pillans 5139 (BOL).

Tulbagh: Tulbaghkloof, Zeyher s.n. (BOL). Vanrhynsdorp: 5 miles N. of Nuwerus, Compton 20571

(NBG); near Klawer Station, Marloth 7794 ; Heerenlogementsberg, Zeyher s.n. (BOL).

South West Africa. — Luderitz: Kuckaus, Dinter 3723; Spitskop, Merxmiiller & Giess 3418;

Tsirub, Range 207 4 (B).

20. A. minutiflorus ( Kunth ) Bak. in Journ. Linn. Soc. Bot. 14: 616 (1875); FI.

Cap. 6: 263 (1896). Type: Delagoa Bay, Forbes s.n. (B, holo.!; PRE, photo.!).

Asparagopsis minutifiora Kunth, Enum. PI. 5: 85 (1850).

7230691-3

64

Roots tuberous; tubers distant or lateral on the roots. Stems erect, almost

straight, softly woody, ridged, usually scabrid along the ridges, pale, up to about 50 cm

long. Branches usually absent. Branchlets straight usually fascicled, ascending,

branched, very narrow, spineless, ridged, scabrid, up to about 5 cm long. Cladodes

3-angled, up to about 10 mm long and about 0 2 mm thick, up to about 8-nate laterally

but more numerous terminally, slightly arcuate. Spines usually present on the stems,

slender, terete, pungent, more or less straight and spreading, the longest from 5-12

mm long, brown. Peduncles borne on stems and less often on branches, solitary or

paired, about 3 mm long. Pedicels usually 2- or 3-nate and terminal, but occasionally

lateral and solitary, borne in the axils of prominent bracts, about 1 mm long, articulated

near the middle. Perianth segments oblong-obovate, 2-3 mm long; the apex of the

inner whorl dentate. Stamens nearly as long as the perianth segments; anthers minute.

Style shortly divided, about 1 mm long; ovary about 1 mm long. Fruit with a smooth

surface, 1 -seeded, up to about 7 mm diameter.

This species has been recorded from a range of habitats, from stony hillsides to

dense thorn-scrub on the flats, widely spread over most of Natal, the eastern Transvaal

and Mozambique.

Natal. — Estcourt: Rondedraai, Acocks 10817. Ingwavuma: 10 miles S.E. of Ingwavuma, Codd

10296. Ngotshe: “ Ngotsche ”, Gerstner 2516 (BOL). Nongoma: Bangonomo, Gerstener s.n.

(NH). Ubombo: Mkuze Game Reserve, Gerstener 4597.

Swaziland. — Stegi, Tambuti Ranch, Compton 30940; St. Phillips Mission Station, Gerstner s.n.

(NH).

Transvaal. — Barberton: Avoca, Thorncroft 3006. Letaba: 3^ miles W. of Letaba Camp, Codd

& Dyer 4694. Pilgrim's Rest: 6 miles N.E. of Skukuza, Codd & Verdoorn 5484.

This and the following species have a unique type of inflorescence by which they

can be distinguished from any other species. In these the raceme has apparently been

so reduced as to leave only 2-4 flowers which are borne terminally. In a few specimen

there may be one or two lateral flowers as well showing what is probably an inter-

mediate stage between these species and the rest of the Section Racemosi.

21. A. rigidus Jessop , sp. nov., A. minutifloro Bak. affinis, sed cladodiis maioribus,

spinis minoribus, caulibus glabris differt. Plate 2.

Caules erecti, leviter tortuosi, lignei, costati, glabri, saepe virides, ad 60 cm longi.

Rami solitarii, leviter tortuosi, quam caules costatiores, brevissimi, inultum ramosi,

plus minusve ascendentes, ad 8 cm longi; rami ultimati non ramulos eximios lateraliter

facti. Cladodia valde 3-angulata, lateraliter ad 3-nata et terminaliter fere 6-nata,

plerumque at 10 cm longa et 0-5-0 -9 mm crassa, plus minusve recta, mucrone tereti

fuscoque; cladodiorum margines saepe costa subtiliter serrata cellarum dilatarum.

Spinae absentes vel ad 4 mm longae, reflexae. Pedunculi in ramis orti, solitarii vel

gemini, c. 3 mm longi. Pedicelli plerumque 2-4-nati terminales sed saepe in pedunculis

laterales, prope medium articulati. Perianthii segmenta oblongo-obovata, 2-3 mm

longa, integra. Stamina fere perianthii segmenta aequilonga; antherae c. 0-5 mm

longae. Stylus breviter ramosus, c. 1 mm longus. Ovarium c. 1 mm longum. Bacca

semine 1, c. 5 mm diam.

Type: Transvaal: Pilgrim’s Rest district, Mariepskop, Killick & Strey 2380

(PRE, holo.).

Stems erect, slightly zigzag, woody, ridged, glabrous, often green, up to 60 cm

long. Branches solitary, slightly zigzagging, more strongly ridged than the stems,

very short, much branched, more or less ascending, up to about 8 cm long; the final

branches not forming specialised branchlets. Cladodes strongly 3-angled, up to 3-nate

laterally and about 6-nate terminally, usually up to about 10 cm long and 0-5-0 -9

mm thick, more or less straight, with a terete, brown mucro; the margins of the cladodes

65

frequently bearing a minutely serrated ridge of enlarged cells. Spines absent or up

to 4 mm long, reflexed. Peduncles borne on the branches, solitary or paired, about

3 mm long. Pedicels usually 2- to 4-nate and terminal but often lateral on the peduncles,

articulated near the middle. Perianth segments oblong-obovate, 2-3 mm long, entire.

Stamens nearly as long as the perianth segments; anthers about 0-5 mm long. Style

shortly divided, about 1 mm long; ovary about 1 mm long. Fruit 1 -seeded, about

5 mm diameter.

This species grows among scrub in rocky areas. Although it is mainly restricted

to the south-eastern parts of the Transvaal, there is a single record from the Petersburg

district. Flowers have been recorded only in November.

Transvaal. — Barberton: Barberton, Pott 5546; Saddleback Mountain, Thorncroft 790 (NH).

Pietersburg: The Downs, Rogers 21982. Pilgrim’s Rest: Mariepskop, Kitlick & Strey 2380; Van der

Schijff 5589; 8 miles N. of Sabie on Graskop road, Codd & de Winter 3313.

This species resembles the preceeding in the nature of the inflorescence, but can

be separated by the fewer, larger cladodes, smaller spines, and glabrous stems.

22. A. densiflorus ( Kunth ) Jessop, comb. nov.

Asparagopsis densiflora Kunth, Enum. PI. 5: 96 (1850). Type; Cape, Lalande s.n.

(P, lecto. ; PRE, phioto.!).

Asparagus sarmentosus sensu Bak. in Journ. Linn. Soc. Bot. 14: 625 (1875); FI.

Cap. 6: 270 (1896). A. myriocladus Bak. in Journ. Bot. Lond. 18: 43 (1889); FI.

Cap. 6: 271 (1896). Type: Inanda, Medley Wood 355 (K, holo. !; PRE, photo.!;

NH!). A. sprengeri Regel in Acta Horti Petrop. 11: 302 (1890); Bak. in FI. Cap.

6: 271 (1896). Type: described from a cultivated specimen from Durban. A. myersii

Hort., nomen nudum.

Roots tuberous; the tubers either borne directly on the rhizome or laterally on

the roots. Stems erect or weakly decumbent, straight or slightly tortuous, finely ridged,

glabrous, 30-60 cm long, green to brown. Branches similar to the stems, but not

usually formed. Branchlets numerous, grooved, occasionally over 10 cm long, spreading

or ascending. Cladodes flattened or rarely 3-angled, slightly arcuate, 5-15 mm long,

weakly to strongly mucronate, commonly solitary, but sometimes ternate or numerous,

with a distinct midrib when flattened. Spines usually well-developed, but not often

pungent, usually absent from the final branches, reflexed, the leaf attenuate. Peduncles

rather shorter than the branchlets, occasionally not differentiated. Pedicels solitary,

generally articulated near the centre. Perianth segments more or less similar, oblong-

obovate, obtuse, spreading, white or pale pink, about 2-5 mm long. Stamens three

quarters of the length of the perianth segments; anthers small. Styles divided for

about half their length. Berry globose, about 5 mm diameter, 1 -seeded, red.

This species grows in a range of habitats, mainly in the coastal areas of the south-

eastern Cape and Natal, from coastal dunes to open rocky places or woods. It has

also been recorded from Inhaca Island. Flowers have been recorded from October

to March and in July.

Cape. — Albany: Howieson’s Poort Hills, Galpin 3084. Alexandria: Alexandria Forest, Jessop

612. Bathurst: Port Alfred, Galpin 2948. Bedford: Bedford, Potts 1730 (BOL). East London:

East London beach, Bokelman 3-28 (NBG). Kentani: Kentani, Pegler 1482 (BOL). Keiskammahoek:

Keiskammahoek, Stayner 53 (GRA). Kirkwood: between Rautenbach’s Drift and Addo Drift,

Bitrchell 4208 (K). Komga: Komga, Flanagan 221 1 . Mount Ayliff: Kokstad, Tyson 31 58. Mqanduli:

Koffiebaai, Van der Schijff 5447. Peddie: Kaffir Drift, Compton 17829 (NBG). Port Elizabeth:

Redhouse, Paterson 531 (BOL). Queenstown: Umbombola Mountain, Galpin 2094 (GRA). Somerset

East: Kommadagga, Burchell 3302 (K). Uitenhage: Swartkops River, Ecklon & Zeyher 1061.

Natal. — Durban: near Durban, Medley Wood 7559 (NH). Eshowe: Eshowe, Lawn 781 (NH).

Hlabisa: Hluhluwe Game Reserve, Ward 2941. Inanda: Inanda, Medley Wood 1626 (BOL). Lower

Tugela: Tugela Beach, Johnson 408 (NBG). Mahlabatini: Mahlabatini, Gerstner 3525 (NH). Port

66

Shepstone: Beach Terminus, Thode 3429 (STE). Ubombo: Mkuze Game Reserve, Ward 3984.

Umzinto: Scottburgh, Thode 3424 (STE). Utrecht: Utrecht, Thode A1298. Weenen: Griffins Hill,

Acocks 11413 (NH).

Orange Free State. — Without precise locality, Cooper 3603 (K).

Transvaal. — Barberton: Sheba, Thorncroft 14.

Linnaeus described A. sarmentosus as having solitary, linear-lanceolate cladodes;

as being flexuose, and as having recurved spines. He cited Hermann’s Hortus Academici

Lugduno-Batavus t. 650, which shows the tuber characters. Another plate, t. 63,

shows the aerial parts. There is a specimen, LINN No. 434. 15, labelled A. sarmentosus

in the Linnaean Herbarium, but this does not fit the description as it has fascicled

cladodes. There is no specimen of this species in the Hermann Herbarium in Leyden,

and the figures must be regarded as iconotypes. Both Linnaeus and Hermann gave

the source of the material as Ceylon. According to Trimen (1898) there are two species

to which this name might apply in Ceylon, A.falcatus L. and A. gonocladus Bak. These

are both much-branched twiners, and it is unlikely that either could by synonymous

with A. sarmentosus sensu Bak. from South Africa. The name A. sarmentosus L.

can therefore not be applied to the South African species.

As Regel described A. sprengeri from Durban, and as he mentioned that it had

racemes and a striate stem, it is very likely that this name is synonymous with A.

densiflorus as construed here. Regel also mentioned that this species had been introduced

into Europe through horticulture. The plant generally known in horticulture as A.

sprengeri fits Regel’s description and plants of this form grow near Durban, making

it very probable that this is the correct application of the binomial. Another cultivated

form of A. denstflorus is known as A. myersii (often misspelt meyeri ), but this name has

never been validly published.

This is an extremely variable species, but no practical way of subdividing it has

been found. It can be identified by the fact that it never climbs to more than a little

over a metre, and then only weakly twining, not twisting to any extent round its support.

The racemes may be obscured by the development of cladodes at the nodes, but are

never reduced to the form found in A. minutiflorus and A. rigidus. The stems are

grooved or ridged, but always glabrous, and branches are usually absent, although

branchlets are well developed.

23. A. saundersiae Bak. in Journ. Bot. Lond. 18: 42 (1889); FI. Cap. 6: 269

(1896). Type: Natal, Saunders s.n. (K, holo.!; BOL!).

Stems woody, erect to climbing, up to 2 m high, hardly zigzagging, slender, glabrous,

smooth, yellowish or golden-brown. Branches similar to the stems, wide spreading

to ascending, usually solitary, lacking spines. Cladodes fascicled, about 6-8-nate,

less than 4- mm brod, angled, subulate, ascending or spreading, up to 1-5 cm long,

slightly arcuate. Spines usually well-developed, absent from the final branches, up

to 1 cm or more long, pungent, spreading, slightly recurved. Peduncles 1-4-nate,

usually about 4 cm long. Pedicels solitary, 5-6 mm long, articulation variable. Perianth

segments serrate, obovate, about 3 mm long, white. Stamens nearly as long as the

perianth segments; anthers minute, globose, yellow. Ovary stalked; style shortly

divided. Fruit not known.

This species occurs mainly at the margins of forests in the south-eastern and

eastern parts of South Africa, possibly as far north as Kenya. It occasionally grows

in the open, even on dry, exposed slopes. Flowers have been recorded from November

to May.

Cape. — Lusikisiki : between Port St. Johns and Lusikisiki, Hutchinson 1780. Mqanduli: Mqanduli,

Pegler 599. Port St. Johns: about 5 miles N.E. of Port St. Johns, Jessop 663.

67

Natal. — Camperdown: Camperdown, Carnegie s.n. (NH); Franks s.n. (NH); Botha’s Hill, Medley

Wood s.n. (NH). Eshowe: Eshowe, Gerstner 2841 (BOL). Hlabisa: near Hluhluwe, Wells 2039;

Hlabisa, Ward 2911. Mtonjaneni: Umhlatuzi Road, Lawn 331 (NH). Nongoma: Nongoma, Gerstner

4423 (NH); 11 miles N. of Nongoma, Jessop 670. Pinetown: Marionhill, Kolbe s.n. (BOL). Rich-

mond: Richmond Road, Schlechter 6740.

Swaziland. — Isateki Beacon, Compton 27312.

Transvaal. — Letaba: Duiwelskloof, Scheepers 1059.

This species is rather similar to A. buchananii Bak. The main difference is that

A. buchananii is a climber with very large spines which are characteristically flattened

at the base. The yellowish colour of the stems and branches of A. saundersiae often

distinguishes it from A. buchananii which usually has rather browner stems and

branches.

24. A. buchananii Bak. in Kew Bull. 1893: 211 (1893); FI. Trop. Afr. 7: 434

(1898). Type: Nyasaland, Buchanan 1503 (K, lecto. !; PRE, photo.!); Buchanan 757.

Stems woody, climbing to at least 9 metres, tortuous and often strongly zigzagging,

glabrous, very variable in colour, smooth. Branches similar to the main stems.

Branchlets often well-defined, solitary, lacking spines, spreading to ascending. Cladodes

fascicled, about 6- to 8-nate, less than mm broad, angled, terete or grooved, subulate,

ascending or spreading, up to 1 • 5 cm long, slightly arcuate. Spines well-developed,

absent from final branches, up to 4 cm or more long on the main stems, dorsally

flattened towards the base, usually reflexed. Peduncles 1- to 4-nate, 2-4 cm long,

usually solitary. Pedicels solitary, 3-5 mm long, articulation variable. Perianth

segments serrate, obovate to oblong-obovate. 1 • 5-3 mm long, white. Stamens nearly

as long as the perianth segments; anthers minute, globose, yellow to brown. Ovary

stalked, style slightly divided. Berries fleshy, about 5 mm diameter, 1- or 2-seeded, red.

This species is a forest climber from Natal in the south to at least as far north

as Nyasaland. Flowers have been recorded from April to June, and in September

and November.

Natal. — Mapumulo: near Thringspost, Mauve 4207. EJbombo: Makatini Flats, Vahrmeyer 432.

Transvaal. — Barberton: Sheba Valley, Thorncroft 3006. Groblersdal: Loskopdam, Mauve 4278.

Lydenburg: Bothas Hoelc, Repton 5947; Sekhukhuneland, Barnard 448. Nelspruit: Numbi, Van der

Schijff 29. Petersburg: Thabina, Schweickerdt 1019; Molopo Reserve, Gerstner 5322. Potgietersrus :

Ihlenfeldt 2067. Sibasa: Punda Milia, Codd 5292. Soutpansberg : Elim, Obermeyer s.n. Wakker-

stroom: Slangapies Mountains, Burtt Davy 1937. Warmbad: 5 miles N. of Warmbad, Vahrmeyer

& Tolken s.n. Waterberg: near Elmeston, Meeuse 10537.

South West Africa. — Grootfontein : 8 miles E. of Runtu, De Winter 3176a. Ovamboland: Oshi-

kango, De Winter & Giess 7054.

Baker cited Buchanan 1503 and Buchanan 757, both from the Shire highlands in

Nyasaland, in his description of A. buchananii. It is proposed that Buchanan 1503

(K) be the lectotype.

This species, particularly if the characters of the main stems are not known, may

be difficult to distinguish from A. saundersiae. The characters of use in distinguishing

these species are given under A. saundersiae. It may also be difficult to distinguish

from the northern forms of A. aethiopicus, which have flattened cladodes and generally

lack the large spines. These spines are the most characteristic feature of the plant,

and may reach more than 4 cm and be extremely pungent and strongly reflexed. The

flattened base is conspicuous even in fairly small spines, but may occur to a small

degree in A. aethiopicus.

25. A. aethiopicus L., Mant. 32 (1767). Type: Cape without precise locality

(LINN No. 434.6, neo.; NBG!, PRE!, photos).

68

Tubers stalked, borne laterally on the main roots, about 30 cm long, fusiform

to ovoid. Stems climbing to about 7 m or erect to 1 m, woody, terete, glabrous.

Branches grooved, spreading or recurved, similar to the stems. Branchlets sometimes

well-defined, grooved, straight. CJadodes flattened, linear, mucronate in many speci-

mens, 3-6-nate or more numerous, 10-40 mm long, 1-2 mm broad, with a single vein.

Spines usually present at all nodes, but often absent on final branches, spreading or

reflexed, often pungent, up to 1 cm long or occasionally more. Leaves and bracts

sometimes ciliate. Peduncles usually 3-15 cm long, paired, unbranched. Peduncles

usually single or paired, 2-4 mm long, usually articulated near the centre, but variable.

Perianth segments obovate, about 3 mm long, often with a ciliate margin, spreading,

white. Stamens almost as long as the perianth segments; anthers orange, 0-5 mm

long. Style short, 0-6 mm long; ovary about 1 mm long. Berry globose, 5-7 mm

diameter, with 1-3 seeds, red; perianth not persistent.

Mainly a forest climber, this species also occurs in coastal vegetation and in parts

of the Little Karroo and Namakwaland. Distributed over a large area from the

south-western Cape to central Africa.

For key to the varieties see key to species.

(a) var. aethiopicus.

A. aethiopicus L., Mant. 32 (1767); R. & S., Syst. Veg. 7: 335 (1829); Bak. in

Journ. Linn. Soc. Bot. 14: 626 (1875); FI. Cap. 6: 271 (1896); Salter in FI. Cape

Penins. 175 (1950). A. lanceus Thunb., Prodr. 66 (1794). Type: without precise

locality, Thunberg s.n. (UPS, holo. !; BOL!, PRE!, photo.).

Asparagopsis lancea (Thunb.) Kunth, Enum. PI. 5: 106 (1850). A. aethiopica (L.)

Kunth, Enum. PI. 5: 95 (1850).

In this variety the stems are grooved. The cladodes appear to be more opaque

and tend to be a little broader and shorter than average.

This variety is confined to the Cape Province, occuring mainly in the coastal

forests from the Cape Peninsula to Humansdorp. Flowers have been recorded from

January to June and in October.

Cape. — Bredasdorp: Uilenkraal Forest, Taylor 286 (NBG). Caledon: Danger Point, Pil/ans 9541

(BOL). Cape: Camps Bay, Pappe s.n. (SAM). Clanwilliam: near Lambert’s Bay, Leipoldt 4244

(BOL). George: Wilderness, Van Niekerk 211 (BOL). Humansdorp: Ratelsbosch, Fourcade 245

(BOL). Knysna: The Point, Taylor 1331 (SAM). Ladismith: Anysberg Dam, Stokoe 8520 (BOL).

Namakwaland: Witbank, Pidans 5121 (BOL). Riversdale: Bottebersfontein, Muir 1421. Robertson:

Robertson, Schmidt 24. Simonstown: Clovelly, Walgate s.n. (BOL). Swellendam: Bonnievale,

Mar loth 12006. Wynberg: Houtbaai, Compton 15598 (NBG).

Linnaeus described A. aethiopicus as having solitary, recurved spines, angled

branches and linear-lanceolate cladodes. He also mentioned that it came from the

Cape, and was similar to A. falcatus, but differed in having fewer cladodes, reflexed

spines, and flexuose, angled branches. The specimen in the Linnaean Herbarium

bears the letters “ H.U.”, indicating a cultivated specimen from the Uppsala Gardens,

as does the description in the Mantissa, and shows the typical cladodes and stem

characters of this variety. The Linnaean specimen is possibly from the same plant as

that described, but is unlikely to have been the actual specimen described, as Jackson

(1912) does not indicate that this specimen was in the Enumeration of 1767 of the

specimens in the Linnaean Herbarium. This specimen is therefore selected as the

neotype.

In the south-western Karroo and in the Little Karroo there are plants which

combine some of the characters of this species and A. racemosus. Both have grooved

branches and the form of branching characteristic of the south-western Cape specimens

of A. aethiopicus. Where the cladodes are flattened, specimens are placed in this

69

species. In flat-cladode forms from the Swellendam area (e.g. Stokoe 8520 from the

Anysberg Dam) there are exceptionally well-developed spines present on the final

branches. In the Laingsburg area (e.g. Jessop 355 from near Matjesfontein) the same

development of spines occurs, but the cladodes are three-angled, and these plants

must therefore be placed in A. racemosus.

(, b ) var. angusticladus Jessop, var. nov.

A. ternifolius sensu Hooker fil. in Bot. Mag. t. 7728 (1900). A. sprengeri sensu

Wright in Bot. Mag. t. 8052 (1906).

Haec varietas caulibus levibus, cladodiis longioribus angustioribus et statu sicco

leviter perlucidis distinguitur. Plate 3.

Type: Transvaal: Middelburg district, Olifants River Gorge, Mogg 22470 (PRE,

holo.).

This variety is distinguished by the smooth stems, and the longer and narrower

cladodes which appear to be slightly translucent when dry.

It is found over most of the Transvaal, but occurs also in Natal, Swaziland,

the extreme eastern parts of the Cape Province, and in a large part of east Africa.

Flowers have been recorded in September and October.

Cape. — Kentani: Kentani, Pegler 840. King William’s Town: King William’s Town, Sim 1061

(BOL).

Natal. — Lower Tugela: Darnall, Schmidt 11 (NH). Mtunzini: Tugela Beach, Johnson 408 (NBG).

Utrecht: Kaffir Drift, Thode A250.

Swaziland. — Mbabane, Dalriach, Compton 30739 (NBG); Hlatikulu, Compton 28903 (NBG);

Mpisi, Compton 30112 (NBG); Stegi, Compton 28020 NBG).

Transvaal. — Barberton: Barberton, Thorncroft 843 (NH). Brits: Weltevreden 61, Mogg 20391.

Carolina: Arnhemburg, Roberts s.n. Heidelberg: Kuilfontein, Mogg 20121. Johannesburg: 9

miles N.W. of Johannesburg, Mogg 19188. Krugersdorp: Jack Scott Private Nature Reserve, Wells

2412. Middelburg: Olifants River Gorge, Mogg 22470. Nelspruit: Amajuba Mountain, Liebenberg

3118. Piet Retief: 27 miles S.E. of Piet Retief, Codd 10268. Pretoria: Pienaar’s River area, Repton

3727. Soutpansberg: Elim, Obermeyer 592.

Baker described A. aethiopicus var. ternifolius in 1871. This variety has been

transferred to A. falcatus. J. D. Hooker, in Bot. Mag. t. 7728, raised this variety to

specific rank, but it is clear that the plant on which the illustration was based is referable

to the variety described above. Wright, in 1906, described a plant which was also

illustrated in the Botanical Magazine, and considered it to be A. sprengeri, but referred

to A. ternifolius as being a later synonym. Again, the figure is of a climber and must

be referred to this variety.

26. A. falcatus L., Sp. PI. 313 (1753). Type: Burmann, Flora Zeylanica t. 13, f. 2.

(1737)

Stems climbing or scrambling, tortuous, smooth, glabrous, pale, much-branched.

Branches usually solitary, spreading, similar to the stems. Cladodes flattened, straight

or falcate, with a prominent vein, up to 9 cm long and 7 mm broad, dark green. Spines

well-developed, recurved, pungent, strong, often borne on vegetative branchlets; the

rest of the leaf not usually developed. Peduncles up to 4-nate laterally, more numerous

terminally, usually about 3-5 cm long, unbranched or branched. Pedicels 4-8 mm

long, articulated proximally, 1- or 2-nate. Perianth segments oblong to obovate, the

inner whorl slightly narrower than the outer, 2-5^1 mm long, cream to white. Stamens

almost as long as the perianth segments; anthers small and globose. Style short;

ovary about 1 mm long. Berry fleshy, about 5 mm diameter, 1- or 2-seeded, red.

This species occurs in the eastern Cape and northwards through east Africa to

Asia. Flowers have been recorded in South Africa from September to February.

70

For key to the varieties see key to species.

(a) var. falcatus.

A. falcatus L., Sp. PI. 313 (1753); Bresler, Diss. 2 (1826); Kunth, Enum. PI. 5:

71 (1850); Bak. in Journ. Linn. Soc. Bot. 14: 626 (1875); FI. Cap. 6: 271 (1896).

The typical variety has larger cladodes, which are usually 3-5-9 cm long and

more or less falcate. The inflorescence is simple.

This variety occurs over the entire range of the species, mainly in forest. Flowers

have been recorded from December to February in South Africa.

Cape. — East London: East London, MacOwan 2413 (BOL). Kentani: Kentani, Pegler 1335.

Port St. Johns: Port St. Johns, Galpin 3491.

Natal. — Durban: Berea , Medley Wood 150. Eshowe: Eshowe, Lawn 73 (NH). Ingwavuma:

Kosi Bay, Vahrmeyer 494. Lower Tugela: Tugela Beach, Johnson 607 (NBG). Mtunzini: Ngoya

Forest, Wells & Edwards (A (Nil). Nkandla: Qudeni Forest, Davis 119 (NH). Pinetown: Umgababa,

Bayer 862. Ubombo: Makatini Flats, Vahrmeyer 429.

Linnaeus described A. falcatus as having solitary spines, and ensiform, falcate

leaves. He reported its origin as being from Ceylon. He also cited Burmann’s Flora

Zeylanica t. 13, f. 2 (1737). This figure must be regarded as an iconotype in the

absence of a specimen from which it was drawn. This specimen should be in Paris

according to Lanjouw and Stafleu (1954), but is apparently missing from the collection.

According to Jackson (1912), a specimen did not appear in the Enumerations of the

Linnaean Herbarium. Moreover, the specimen in the Linnaean Herbarium, LINN

No. 434.3, has short cladodes, and is in fact A. aethiopicus var. aethiopicus. The

specimen in the Linnaean Herbarium cannot, therefore, be regarded as the type. Four

specimens from Ceylon, the type locality for A. falcatus , were loaned to the author

by the Director of the Kew Herbarium. These specimens match several specimens,

such as Thode A1563 (PRE) from Zululand and Goldsmith 123/62 (PRE) from Melsetter,

exactly. In a few respects most Ceylon specimens are not a perfect match of the South

African plants, however. The racemes are shorter, being only from 1-2 cm long,

and the cladodes are longer, from 4-12 cm, and narrower, often only about 2 mm broad.

These characters cannot, however, be regarded as being of significance taxonomically.

( b ) var. ternifolius {Bak.) Jessop, comb. nov.

A. aethiopicus L. var. ternifolius Bak. in Saund. Ref. Bot. t. 261 (1871). Type: Natal,

Cooper sub Saunders 1448 (K, holo. !; PRE, photo.!). A. aethiopicus L. var . natalensis

Bak. in FI. Cap. 6: 272 (1896). Type: Natal, Inanda, Medley Wood 1351 (K, holo.!;

PRE, photo.!; NH!).

This variety has shorter cladodes than the typical variety, being up to about

35 mm long. The inflorescence is branched.

This variety does not occur in the Cape Province and probably does not extend

as far north as the typical variety. It habit is similar, but it does not appear to climb

to such a height, perhaps only two or three metres. It tends to occur in drier areas

and the height may be associated with this. Flowers have been recorded from September

to December.

Natal. — Camperdown: Drummond, Gerstner 2421. Hlabisa: Hluhluwe Game Reserve, Ward

1535. Inanda: Umzinyati Falls, Medley Wood 1357 (SAM). Lower Tugela: Tugela Valley below

Sans Souci, Edwards 1685. Mtonjaneni: Mkubaan Valley, Acocks 12936. Mtunzini: Mandini,

Gerstner 2817 (NH). Nongoma: 1 1 miles N. of Nongoma, Jessop 671. Umfolozi: about 3 miles N.W.

of Empangeni, Jessop 665. Ubombo: Mkuze, Galpin 13649. Umvoti: between Greytown and Tugela

Ferry, Edwards 922.

Swaziland. — Ingwavuma Poort, Compton 28903; Hlatikulu, Sipopaneni, Murdoch 125 (NBG);

Blue Jay Ranch, Compton 29570 (NBG).

71

Transvaal. — Barberton: Komatipoort, Van der Schijff 1130. Waterberg: near Elmeston, Mecuse

£c Strey 10430.

The climbing species of Asparagus with racemes and flattened cladodes can be

readily divided into four groups, but the status which should be alotted to these is

not easily decided. It has been decided that the most natural grouping is to place

the two groups in which the cladodes dry very dark, often almost black, together under

A. falcatus. These two can be separated with certainty only because of the difference

in inflorescence. Although plants with branched racemes tend to have shorter cladodes

than those with simple racemes, there is a complete intergradation in the cladode length

of these varieties. The other two groups, in which the cladodes dry a lighter colour,

are placed as varieties of A. aethiopicus and are differentiated from each other on the

basis of the stems being grooved or smooth.

27. A. aspergillus Jsssop, sp. nov., A. racemoso Willd. affinis, sed caulibus scabridis,

rectioribus cinereisque, antheris fuscis differt. Plate 4.

A. racemosus sensu Solch, Beitr. FI. Siidwest-Afr. 38 (1961).

Caules ascendentes vel erecti ad 2 m, lignosi, sulcati vel costati, scabridi, cani.

Rami caulibus similes sed saepe sulcatiores et scabridiores, solitarii, divaricati vel

ascendentes. Ramuli 1-3-nati, plus minusve divaricati, recti, sulcatissimi. Cladodia

fasciculata, multa, angulata, recta vel arcutata, ad 4 cm longa et mm crassa, sed

plerumque c. 1-5-2 cm longa, plerumque flore absentes. Spinae in caulibus praecipue

ramis et plerumque ramulis, sed non in pedunculis, pungentes, divaricatae vel leviter

reflexae, leviter recurvae, ad 1 cm longae, pallide fuscae. Pedunculi gemini, nonnunquam

cladodiis instructi. Peclicelli solitarii vel interdum gemini, 1-5-2 -5 mm longi, prope

florem articulati. Perianthii segmenta oblongo-obovata, integra, 2-4 mm longa, alba.

Stamina quam perianthii segmenta paulo breviora; antherae fuscissimae. Stylus

1-1-2- mm longus; rami c. mm longi. Baccus globosus, seminibus 1 vel 2. c. 6 mm

diam., ruber, pericarpio rugoso.

Type: Transvaal: Letaba district, near Birthday Road at Mamaranga, Breyer

in TRV 19063 (PRE, holo.).

Stems twining or erect to 2 m, woody, grooved or ridged, scabrid, pale grey.

Branches similar to the stems, but often more prominently grooved and scabrid.

solitary, spreading to ascending. Branchlets 1-3-nate, more or less spreading, straight,

strongly grooved. Cladodes fascicled, numerous, angled, straight or arcuate, up to

i mm thick and 4 cm long, but usually about 1-5-2 cm long, usually absent during

the flowering period. Spines present on the main stems, branches and usually on

the branchlets, but not on the peduncles, pungent, spreading or slightly reflexed, slightly

recurved, up to 1 cm long, pale brown. Peduncles paired, sometimes bearing cladodes,

usually 2-5 cm long, but up to 15 cm, not usually bearing spines. Pedicels solitary

or less often paired, 1-5-2 -5 mm long, articulated near the flower. Perianth segments

oblong-obovate, entire, 2-4 mm long, white. Stamens a little shorter than the perianth

segments; anthers very dark coloured. Style 1-1-} mm long; branchlets about -} mm

long. Berry globose, 1- or 2-seeded, about 6 mm diameter, red, with a wrinkled

pericarp.

This species occurs in the northern Transvaal, Bechuanaland, northern South West

Africa and Rhodesia. Flowers have been recorded in October and June.

Transvaal. — Letaba: near Birthday Road at Mamaranga, Breyer'm TRV 19063. Messina: Zeerust,

Tokwe Farm, Breyer s.n.; Messina, Rogers 19418. Pilgrim’s Rest: 20 miles N. of Satara, Van der

Schijff 518.

South West Africa. — Caprivi: east of Cuando River, Curson 900. Grootfontein: Oshikango,

De Winter & Giess 7053. Kaokoveld: Orumane, Abner A23 (M); 15 miles E. of Ohopuho, Merxmuller

1517.

72

This species was confused with A. racemosus Willd. by Solch (1961), but is clearly dis-

tinct. The stems differ in being grey, scabrid, and straighter. The spines tend to be rather

more slender. The anthers are very dark, almost black, giving the inflorescence the

appearance of being covered by the fruiting bodies of an Aspergillus.

28. A. racemosus Willd., Sp. PI. 2: 152 (1799). Type: India, in herb. Willdenow

(B, holo.; PRE, photo.!).

A. tetragonus Bresler, Diss. 27 (1826); Wright in Bot. Mag. t. 8288 (1909). Type:

South Africa, Mundt & Maire s.n. A. racemosus Willd. var. tetragonus (Bresler) Bak.

in Journ. Linn. Soc. Bot. 14: 624 (1875); FI. Cap. 6: 269 (1896).

Asparagopsis floribunda Kunth, Enum. PI. 5: 98 (1850), nom. illegit. Type: as for

A. racemosus. A. subquadrangularis Kunth, Enum. PI. 5: 94 (1850). Type: “ Eckl.

et Zeyh. Herb. Cap. n. 171. (v.s. in Herb. Luc.)”.

Stems twining, climbing, woody, usually grooved, glabrous. Branches similar to

the stems, solitary, spreading to ascending. Branchlets usually 1- to 3-nate, spreading

or ascending, straight, grooved, spineless. Cladodes fascicled, numerous, angled,

straight, up to 1 mm broad, generally 1-2 cm long, green at first but turning greyish.

Spines pungent, usually about 3-4 mm long, brownish to greyish. Peduncles paired,

up to 10 cm long. Pedicels 1- or 2-nate laterally, more numerous terminally, 2-3 mm

long, articulated near or below the centre. Perianth segments similar, obovate-oblong,

2 -5-3-0 mm long. Stamens not appreciably shorter than the perianth segments;

anthers about ^ mm long. Style short, shortly divided. Berry globose, 1 -seeded,

about 6 mm diameter, with a wrinkled pericarp, red.

This species is distributed from the southern Cape through east Africa and into

Asia. It climbs or scrambles to four metres, but usually only about two metres, in

bush or forest. Flowers have most frequently been recorded from March to July in

South Africa.

Cape. — Albany: Queensroad, Schouland 4357 (GRA). Alexandria: De Bega, Archibald 5908.

Bathurst: Port Alfred, Schouland s.n. (GRA). Beaufort West: Sunnyside, Esterhuysen 4359. Cathcart:

5 miles N. of Cathcart, Jessop 658. Cradock: 10 miles east of Cradock, Jessop 654. East London:

near Nahoon Mouth, Galpin 5737. Heidelberg: 5 miles W. of Heidelberg, Jessop 590. Humansdorp:

Gamtoos Gorge, Compton 23448 (NBG). King William’s Town: King William’s Town, Sim 1052.

Knysna: Blaauwkrantz, Dyer 2029 (GRA). Komga: Prospect Farm, Flanagan 299 (SAM). Ladismith:

Sandberg W. of Calitzdorp, Wurts 1371 (NBG). Middelburg: Doornberghoek, Acocks 8665. Mossel-

baai: Gouritz Valley, Horn s.n. Port Elizabeth: near Port Elizabeth, Rodin 1228 (BOL). Riversdale:

near Kweekkraal, Muir 3140. Somerset East: Cookhouse, Schlechter 2750. Uitenhage: Redhouse,

Rogers 2793 (SAM). Umtata: near Umtata, Acocks 10910. Uniondale: Kouga River, Compton

5214 (BOL). Willowmore: Willowmore, Marloth 14129.

Natal. — Estcourt: Pasture Research Station, West 364. Lions River: 5 miles along Merrivale-

Boston road. Mol! 934. Ubombo: near the turning to Mkuze Game Reserve, Jessop 669. Utrecht:

4 km from Utrecht on Knight’s Pass road, Story 1766. Weenen: Mielietuin, West 1209.

Swaziland. — Ingwavuma Poort, Compton 28579 (NBG).

Transvaal. — Lydenburg: 16 miles beyond Malipsdrif, Obermeyer & Verdoorn 22; Erasmus Pass,

Strey 3719.

Willdenow described A. racemosus from India. The South African material

differs from most specimens examined from India by the author in having well-defined

branchlets, grooved stems and straighter cladodes, but all these characters are so

variable as not to warrant even varietal status when the whole range of variation over

the range of the species is considered. For example, one specimen from Madras,

Jacob 17549 (K), differs from South African specimens only in having spines on the

final branches. Walker 56 (K) from Ceylon matches the South African plants even

more closely, as it has grooved stems, and lacks the spines on the final branches.

Bresler considered that the South African plants were specifically different, and named

them A. tetragonus. Baker reduced this species to varietal status.

73

Asparagopsis floribunda Kunth is a superfluous name for A. racemosus Willd.

Asparagopsis subquadrangularis was described as having setaceous cladodes, grey stems

and pedicels 2 mm long. The inflorescence is a raceme. Kunth cited the specimen

as “ Eckl. et Zeyh. Herb. Cap. no. 171. (v.s. in Herb. Luc.)”. According to De Candolle

(1880), the Luca specimens were at Kiel, but this specimen appears to have been

destroyed. It is very likely that this name is referable to A. racemosus, although no

authentic material has been seen.

29. A. obermeyerae Jessop, sp. nov., A. racemoso Willd. affinis, sed caulibus erectis

et canis, ramis canis, floribus axillaribus differt. Plate 5 and 6.

Caules erecti, lignosi, glabri, plus minusve recti; caules juvenes cani vel luteo-

cani sulcati, maturi fusci leves; ad fere 1 m alti. Rami caulibus similes, fasciculati,

constanter cani et sulcati, simplices vel ramis paucis brevibus, ad 15 cm longi. C/adodia

fasciculata, multa, angulata, plus minusve recta, 12-15 mm longa, c. 0-2 mm crassa,

statu sicco fuscescentes. Spinae pungentes, fere rectae, patentes, caulibus ramisque

ortae, ad 15 mm longae, cauli concolores. Pedicelli solitarii vel gemini, ramis orti,

fere vel supra medio articulati, 2-3 mm longi. Perianthii segmenta obovata vel oblongo-

obovata, 2-5-3 mm longa, margine integro, similia, margine pallido. Stamina quam

perianthii segmenta breviora; antherae c. £ mm longae. Stylus c. {- mm longus, semi-

divisus. Baccus globosus, seniine 1, c. 8 mm diam., levis, perianthio persistenti.

Type: Transvaal: Pietersburg district, near Boyne on road to Wolkberg, Mauve

4442 (PRE, holo.).

Stems erect, woody, glabrous, more or less straight, pale grey or yellowish-grey

and grooved when young, but becoming brown and smooth, up to almost 1 m high.

Branches similar to the stems, fascicled, persistently greyish and grooved, simple or

with a few short branches, up to 15 cm long. Cladodes fascicled, numerous, angled,

more or less straight, 12-15 mm long, about 0-2 mm thick, darkening with drying.

Spines pungent, almost straight, spreading, borne on branches, articulated near or

above the middle, 2-3 mm long. Perianth segments obovate or oblong-obovate, 2 • 5-3

mm long, entire, similar, with a pale margin. Stamens shorter than the perianth

segments; anthers about y mm long. Style about § mm long, divided for about half

its length. Berry globose, 1 -seeded, about 8 mm diameter, smooth, with a persistent

perianth.

This species has been recorded only from dry, bushveld slopes in the Pietersburg

district. Flowers have been collected in October.

Transvaal. — Pietersburg: Subiaco Mission, Molepo Reserve, Gerstner 5359; Naauwpoort, Breme-

kamp & Schweickerdt s.n.; Chunes Poort, Hafstrom & Acocks 197; 10 miles E. of Boyne, Mauve 4294;

Mauve 4442.

While this species most nearly resembles A. racemosus, it differs in having axillary

flowers, in its erect habit, and in the pale grey, grooved stems and branches. This

last character is shared with several species, such as A. compactus, in the section Africani.

It also resembles the Africani in having axillary flowers, but the fascicled branches

place it rather in the Racemosi.

It is named after Mrs. Mauve who has taken a particular interest in the genus,

and collected the type material of this species.

30. A. krebsianus {Kunth) Jessop, comb. nov.

Asparagopsis krebsiana Kunth, Enum. PI. 5: 93 (1850). Type: Cape, without

precise locality, Krebs s.n.

74

Tubers stalked, borne laterally on the main roots, about 30 cm Jong, fusiform.

Stems erect or scrambling, slightly zigzagging, woody, smooth or slightly ridged, grey-

green, glabrous. Branches solitary, almost always grooved, much-branched, similar

to the stems. Cladodes fascicled, three-angled, rigid, mucronate, 15-20 mm long or

sometimes only 1 cm or less, linear. Spines present on all stems and branches, pungent,

slightly recurved, browner than the stems and branches; the rest of the leaf not con-

spicuous. Peduncles 1-4-nate, with or without spines, up to 10 cm long, articulated

near the centre; the proximal half greatly thickened. Perianth segments oblong to

obovate, the outer whorl narrower, 2-5-4 mm long, white, reflexed near the centre

of each segment. Stamens nearly as long as the perianth segments. Style slightly

less than 1 mm long, divided for about half its length; ovary about 1 mm long. Berry

smooth, 1-seeded, red; perianth not persistent.

A. krebsianus may scramble among rocks or on bushes in wetter areas, but is

generally more or less erect and usually occurs on fairly dry slopes. It has been recorded

from the eastern Cape and parts of the Transvaal. Specimens, which may be referrable

to this species, have been collected in South West Africa. Flowers have been collected

in most of the summer months from October to April.

Cape. — Albany: Grahamstown, Daly & Sole s.n. ; Penther 1462 (BOL). Alexandria: Alexandria,

Archibald 5488. Cathcart: Inverthorn, Barker 3469 (NBG). Komga: Komga, Flanagan Till. Peddie:

Peddie, Barker 7821 (NBG). Victoria East: Tukulu, Acocks 11145.

Transvaal. — Brits: Jacksonstuin, Codd 6805. Krugersdorp: Baviaanspoort, Smith 351. Lyden-

burg: Driekop, Mogg & Barnard s.n. Pretoria: Pretoria Koppies, Leendertz 531; Derdepoort,

Robertson 123.

Asparagopsis krebsiana is based on a specimen collected by Krebs at the Cape.

The Krebs specimens were mainly at Kiel according to De Candolle (1820). A large

part of the Kiel Herbarium has been destroyed, and it does not now have the specimen.

Most of the plants on which Kunth worked were in Berlin, but there is no specimen

there either. The plant was described as having three-angled cladodes, spinous final

branches, racemes, and pedicels 3 mm long. As Krebs did collect in the eastern Cape

where the species described above occurs, and as A. krebsianus as construed here does

match Kunth’s description in every detail, there seems little doubt as to the correct

application of the epithet. It is proposed, should an isotype of Krebs’ specimen not

be found, that Flanagan 2377 (PRE) from Komgha be taken as the neotype. Duplicates

of this collecting have been seen by the author in the Bolus and Albany Museum

Herbaria.

The type of A. confertus Krause was Engler 6764 from Schakalskuppe in the

Luderitz district. A specimen from the Berlin-Dahlem Herbarium of this collecting

has been examined by the author, and appears to be identical with A. krebsianus

although the cladodes are a little shorter than the average, being about 10-15 mm

long. It seems likely that despite the considerable gap between this locality and the

nearest known for A. krebsianus it should be referred to this species. Other specimens

of this form are Range 1715 (B, SAM) from the type locality and Dinter 6155 (B)

from Aus.

31. A. acocksii Jessop, sp. nov., A. racemoso Willd. affinis, sed caulibus erectioribus,

levibus et pubescentibus differt. Plate 7.

Caules lignei, erecti vel scandentes, tortuosi, teretes, pubescentes, leves, cani.

Rami divaricati vel ascendentes, recti vel leviter tortuosi, plerumque solitarii sed aliquando

ad 3-nati. Cladodia subulata, plerumque ad 6 mm longa, arcuata vel recta, acuminata,

multi-nata, glabra. Spinae manifeste evolutae, leviter recurvae, omnino in caulibus

et ramulis vegetativis. Pedunculi in paribus, plerumque fere 3 cm longi, aliquando

spinosi. Pedicelli 1— 3-nati, 3-4 mm longi, fere 1 mm e tamo articulati. Perianthii

75

segmenta similia, oblongo-obovata, fere 3 mm longa, margine integra. Stamina quam

segmenta perianthii leviter breviora; antherae fere \ mm longae. Stylus brevis, semi-

divisus; ovarium fere sessile, ad fere 2 mm longum. Bacca fere 6 mm longa, semine

solitario, laevigata.

Type: Cape: Cradock district, 12 miles N.N.E. of Fish River Station, Acocks

16296 (PRE, holo.).

Stems erect to scrambling, zigzagging, woody, terete, smooth, grey, densely

pubescent. Branches spreading to ascending, straight or slightly zigzagging, usually

solitary but sometimes up to 3-nate. Cladodes subulate, usually up to 6 mm long,

arcuate or straight, acuminate, many in a bundle, glabrous. Spines well-developed,

slightly recurved, present on all stems and vegetative branches. Racemes paired,

usually about 3 cm long, sometimes bearing spines. Pedicels 1 -3-nate, 3-4 mm long,

articulated about 1 mm from the branch. Perianth segments similar, oblong-obovate,

about 3 mm long, entire. Stamens slightly shorter than the perianth segments; anthers

about mm long. Style short, divided for about half its length; ovary almost sessile,

up to about 2 mm long. Fruit about 6 mm diameter, 1-seeded, smooth.

This species does not appear to be common anywhere, but is spread over a large

part of the drier southern areas oi the Cape. In the west it is known from Namakwaland

and in the east there are single records from both Zululand and the Transvaal. Despite

this disjunct distribution, the characters are sufficiently distinct to make it very likely

that the specimens are all correctly included in this species. Flowers have been collected

from October to January and in March.

Cape. — Beaufort West: Sunnyside, Esterhuysen 5036 (BOL). Bedford: 13 miles N.N.E. of Cook-

house, Acocks 11908. Cradock: 12 miles N.N.E. of Fish River Station, Acocks 16296. Graaff-Reinet:

Wimbledon, Henrici 4964. Namakwaland: Holgat, Pillans 5178 (BOL). Port Elizabeth: Redhouse,

Paterson 978 (BOL). Uitenhage: Aloes, Drege s.n.

Natal. — Ingwavuma: Ndumu Game Reserve, Ward 2034.

Transvaal. — Lydenburg: 2 miles N. of Ohrigstad, Young A616.

This is one of the most densely pubescent species in South Africa. The normally

strongly arcuate cladodes and recurved spines are also characteristic. The nearest

species is A. racemosus from which it can be distinguished by the pubescence, the smooth

stems, and more erect habit. Another species with many similarities is A. nelsii, which

has a whitish stem, well-defined branchlets and yellowish flowers.

32. A. crassicladus Jessop, sp. nov., A. racemoso Willd. affinis, sed ratnulis minus

manifestis, cladodiis crassioribus, racemis pedicellisque brevioribus differt. Plate 8.

Caules lignei, tortuosi, teretes, plerumque pubescentes, cani vel cano-fusci, ali-

quando cortice cano exuviales. Rami divaricati vel ascendentes, leviter tortuosi, 1-

vel 3-nati. Cladodia transversa sectioni ovalia, acuminata, 12-25 mm longa, interdum

scabrida, lateraliter 1- ad 3-nata, terminaliter ad 12-nata. Spinae manifeste evolutae,

reflexae, omnino in caulibus et ramulis vegetativis. Racemi in paribus dispositi, ad

6 cm longi. Pedicel/i 2-6-nati, 3-4 mm longi, c. 1 mm e ramo articulati. Segmenta

perianthii similia, oblonga, aliquando leviter obovata, 3 mm longa, lacticolorata.

Stamina f perianthii segmentorum aequantia; antherae minutae. Ovarium 1 mm

longum, breviter stipitatum: stylus fere 0-5 mm longus, semi-divisus.

Type: Cape: Uniondale district, Kouga, Esterhuysen 4670 (BOL, holo.).

Stems woody, tortuous, terete, usually slightly pubescent, grey or brownish grey,

sometimes with the outer layers peeling. Branches spreading or ascending, slightly

tortuous, 1- or 3-nate. Cladodes oval in cross section, acuminate, 1-3-nate laterally,

up to 12-nate terminally, sometimes scabrid, 12-25 mm long. Spines well-developed,

reflexed, sometimes persistent on all vegetative stems and branches. Racemes paired,

76

up to 6 cm long. Pedicels 2-6-nate, 3-4 mm long, articulated about 1 mm from the

peduncle. Perianth segments similar, oblong, sometimes slightly obovate, 3 mm long,

cream coloured. Stamens three quarters of the length of the perianth segments; anthers

minute. Ovary shortly stalked, 1 mm long; style about half as long, divided for about

half its length. Fruit not known.

This species is a climber to about 2 m on bushes in the southern Cape from tne

Uniondale district in the west to at least as far east as the Somerset East district.

Flowers have been recorded from December to February.

Cape. — Alban> : Sushington Valley, Britten 2724 (GRA); Penrock Farm, Dyer 1193 (GRA).

Alexandria: 9 miles from Salem towards Alexandria, Jessop 609. Bathurst: Horseshoe on the Kariega

River, Jessop 617. Kirkwood: Addo Reserve, Repton 5674. Somerset East: 1 mile W. of Bruintjes-

hoogte, Jessop 645. Uniondale: Kouga, Esterhuysen 4670 (BOL); Ecklon & Zeyher s.n. (BOL).

This species is very similar in general habit to A. racemosus , but sometimes has

spines on the final branches and diffeis in the shape of the cladodes and in the length

of the peduncles and pedicels. There is little differentiation of branchlets.

33. A. nelsii Schinz in Bull. Herb. Boiss. 1, 4, app. Ill: 44 (1896); Solch, Beitr.

FI. Siidwest-Afr. 38 (1961). Type: S.W.A., ‘ Hereroland ”, Nets 20 (Z, holo.!).

Stems erect, somewhat zigzagging, smooth, sometimes slightly pubescent, grey-

white becoming brown as the outer layers disintegrate, generally up to about 50 cm

high. Branches spreading to ascending, solitary except on the older parts of the plant

where they may be paired, similar to the stems. Branchlets usually a pale yellow

to pale brown, solitary except on the older parts where they may be paired. In young

plants solitary herbaceous branchlets occur on solitary woody branches, but in the

older parts, branches and branchlets become more numerous and may become indis-

tinguishable from one another. Cladodes narrow, terete, fascicled, 10-20 mm long,

appearing after the flowers. Spines well-developed on all stems, branches, branchlets

and peduncles, spreading to recurved; the larges ones grey with brown apices; the

smaller brown. Peduncles 3-5 cm long, unbranched, or up to about 10 cm long, and

branched, pale grey to yellowish, solitary or up to about 10-nate. Pedicels articulated

once just below the centre, or sometimes twice, 2-5 mm long, up to 4-nate. Perianth

segments similar, free or fused for up to about one fifth of their length, entire, 3-4 mm

long, oblong-obovate, yellowish. Stamens slightly shorter than the perianth segments.

Style and ovary each about 1 mm long; ovules 8 or 9 per cell. Berry globose, about

5 mm diameter, 1- or rarely 2-seeded, red.

This species is widespread and fairly frequent in the dry Karriod and Kalahari

areas of the northern Cape, Bechuanaland and South West Africa. Flowers have been

recorded from August to October.

Cape. — Gordonia: 11 miles N.E. of Twee Rivieren, Leistner 2899. Hay: 8 miles E. of Postmasburg,

Acocks s.n. Kenhardt: near Pofadder, Jessop 352 (BOL). Kuruman: Tellery Pan near Kuruman

River, Leistner 2167.

Transvaal. — Soutpansberg: Hackthorne Farm, Code I 4336.

South West Africa. — Gibeon: Haribes, Volk 12188 (M). Grootfontein: Onguma, Walter 416

(M). Keetmanshoop : Gariganus, Acocks 15656. Maltahohe: Duwisib, Volk 12685 (M). Okahandja:

Okahandja, Dinter 282 (SAM). Otjiwarongo: Quickborn, Bradfield 89. Outjo: Outjo, Thorne s.n.

(SAM). Ovamboland: Oshikango, Loeb 447. Rehoboth: Rehoboth, Fleck 212 (Z). Warmbad:

Warmbad, Wandres s.n. (Z). Windhoek: Windhoek, Rogers 29773 (Z).

Bechuanaland. — Toteng, Story 4661; Ngamiland, Curson 556.

34. A. striatus (Linn, f) Thumb., Prodr. 65 (1794); Bak. in Journ. Linn. Soc. Bot.

14: 621 (1975): FI. Cap. 6: 267 (1896); Solch, Beitr. FI. Siidwest-Afr. 39 (1961).

Type: Cape of Good Hope without precise locality, Thunberg s.n. (UPS, holo.!).

77

Dracaena striata Linn, f., Suppl. 204 (1781). D. erecta Linn, f., Suppl. 204 (1781);

R. & S., Syst. Veg. 7: 346 (1829). Syntypes: Cape of Good Hope without precise

locality, Thunberg s.n. (UPS, a!, b!). D. stricta Schult. in R. & S., Syst. Veg. 7: 346

(1829), in errore D. striata.

Asparagus erectus (Linn, f.) Thunb., Prodr. 65 (1794); Bak. in Journ. Linn. Soc.

Bot. 14: 621 (1875); FI. Cap. 6: 267 (1896). A. engleri Krause in Bot. Jahrb. 51:

450 (1914). Type: S.W.A., Great Karasberg, Englsr 6651 (B, holo.!).

Myrsiphyllum striatum (Linn, f.) Kunth, Enum. PI. 5: 110 (1850). M. erectum

(Linn, f.) Kunth, Enum. PI. 5: 109 (1850).

Tubers borne laterally on the roots. Stems erect, glabrous or minutely pubescent,

woody, grooved, almost straight, greenish. Branches ascending, similar to the stems,

but often more zigzagging, not bearing branchlets. Cladodes terminal and 2- or 3-nate

and lateral and solitary in most specimens, very variable in shape from linear to almost

lanceolate, 10-40 mm long, 1-5 mm broad, rarely almost terete, very firm and rigid.

Spines often not developed, small, reflexed, hardly ever pungent: the remaining part

of the leaf with an entire margin. Peduncles in terminal fascicles varying from 1-2

or less often up to 15 or more, about 5 mm long, articulated below the centre. Perianth

segments 3-4 mm long, oblong-obovate with a serrate apex or not, similar. Stamens

not spurred, almost as long as the perianth segments; anthers about £ mm long. Style

branches very short; style about 1 mm long. Berry fleshy, about 5 mm diameter,

generally 1- or 2-seeded, red.

This species occurs in dry areas, particularly on rocky hills, over much of the Cape

Province east of Laingsburg. There are also a few records from the Orange Free

State and South West Africa. Flowers have been recorded from August to January

and in March.

Cape. — Adelaide: 3 miles from Adelaide on Bedford road, Jessop 644. Albany: Grahamstown,

Schlechter 2633 (GRA). Alexandria: Bushman’s River Poort, Galpin 2976. Aliwal North: Orange

River, 3 miles down from Aliwal North, Gerstner 62. Bathurst: Hopewell, Acocks 11128. Beaufort

West: Sunnyside, Esterhuysen 4356 (BOL). Calvinia: Calvinia, Schmidt 232. Clanwilliam: Brak-

fontein, Ecklon & Zeyher s.n. (BOL). Cradock: Cradock, Killick 796. De Aar: near De Aar,

Schweickerdt 1206. Graaff-Reinet : near Graaff-Reinet, Bolus 516 (BOL). Hay: Floradale, Esterhuysen

s.n. Humansdorp: Kabeljaauw, Burtt-Davy s.n. Kirkwood: Addo Reserve, Repton 5703. Kuruman:

Lohatlha, Esterhuysen 2414 (BOL). Laingsburg: Ngaap Kop, Compton 9242 (NBG). Middelburg:

Middelburg, Theron 454. Murraysburg: The Cave, Tyson 242 (GRA). Oudtshoorn: near Cango,

Britten 1771 (GRA). Port Elizabeth: Swartkops River, Zeyher 480. Prieska: Prieska, Bryant J330.

Queenstown: Oxton Manor, Galpin 7373. Riversdale: Riversdale, Bolus s.n. (BOL). Somerset

East: Boschberg, MacOwan 1113 (SAM). Uniondale: Kouga near Misgund, Compton 7483 (NBG).

Uitenhage: The Aloes, Drege 3164 (GRA). Victoria West: Slypfontein, Rehm s.n. Willowmore:

Willowmore, Watt & Brandwyk 2396.

Orange Free State. — Fauresmith: Botanical Reserve, Pole Evans 1794. Thaba Nchu: Thaba

’Nchu, Roberts 2945.

South West Africa. — Warmbad: Kanus, Dinter 3072 (SAM). “ Grosse Namaland ”, Fleck

306a (Z).

Thunberg did not cite a Linnaean synonym for A. striatus but there is no reasonable

doubt that the omission of Dracaena striata was accidental, as all other species collected

at the Cape by Thunberg and regarded as Dracaena species by Linnaeus are cited.

D. striata and D. erecta were described in the same work, and the epithet striatus is

preferred by the present author as it is in more common use.

Linnaeus separated these two species on D. erecta being herbaceous and D. striata

being woody, with striate cladodes. Neither of these characters has any significance

as both are woody, although often green, and have striate cladodes. Baker (1896),

who realised the worthlessness of these characters, separated his concept of these two

species on whether the flowers were solitarW (in A. erectus) or not. This character

does not seem to have any taxonomic significance.

78

Schultes cited both A. striatus and D. striata as synonyms for his D. stricta, the

spelling of which is, therefore, very likely to have been an orthographic error.

35. A. subulatus Thunb., Prodr. 66 (1794); R. & S., Syst. Veg. 7: 326 (1829);

Kunth, Enurn. PI. 5: 74 (1850); Bak. in Journ. Linn. Soc. Bot. 14: 612 (1875); FI.

Cap. 6: 262 (1896). Type: Cape of Good Hope without precise locality, Thunberg

s.n. (UPS, holo.!; BQLb PRE!, photos).

Tubers borne laterally on the roots. Stems erect, slightly zigzagging, glabrous

or minutely pubescent, woody, grooved, greenish. Branches reflexed or at least

spreading, similar to the stems, but more zigzagging. Cladodes often paired terminally

and single laterally, but sometimes up to 10 or more terminally and paired laterally,

terete, grooved, up to about 2 ■ 5 cm long and 1 mm broad, rather firm. Spines rudi-

mentary or absent, the remainder of the leaf not scarious. Peduncles in terminal

fascicles varying from 1-15, 3-5 mm long, articulated below the centre. Perianth

segments 3-4 mm long, oblong-obovate, sometimes much narrowed towards the apex,

entire, similar. Stamens not spurred, almost as long as the perianth segments; anthers

about \ mm long. Style branches rather short; style about 1 mm long. Fruit about

5 mm diameter, fleshy, 1-3-seeded; perianth persistent and conspicuous.

This species occurs mainly in dry areas, especially on rocky slopes, mainly in

the south-eastern parts of the Cape, but also in parts of the Transvaal, Natal and

Mozambique; flowers have been recorded from September to December.

Cape. — Albany: Sandy Drift near Grahamstown, Daly 70 (SAM). Alexandria: valley south of

Signal Hill, Gatpin 2923. Bathurst: West bank of Kowie, Britten 1875. Butterworth: overlooking

the Butterworth River, Pegler 2078 (BOL). Cathcart : near the Swart Kei River, 15 miles N. of Cathcart,

Roberts 1827A. East London: Chalumna causeway, Barker 6966 (NBG). Fort Beaufort : Fort Beau-

fort, Taylor s.n. (GRA). Humansdorp: Zeekoe River mouth, Fourcade 5802 (NBG). Kentani:

Kentanij Pegler 2078. King William’s Town: King William’s Town, Sim 1047 (BOL). Kirkwood:

Addo National Park, Archibald 3781. Komga: Kabousie River near Komga, Flanagan 2584 (BOL).

Peddie: 15 miles from Peddie towards East London, Barker 3969 (NBG). Port Elizabeth: Swartkops

River, Ecklon & Zeyher 250. Queenstown: Newstead below Otterburn, Acocks 12158. Uitenhage:

Aloes, Drege 3162. Umtata: near Umtata, Acocks 10971. Victoria East: Breakfast Vlei, Barker

2834 (NBG).

Natal. — Camperdown: Umgeni Valley, Forbes 1257 (NH). Lower Umfolosi: Umfolosi Game

Reserve, Ward 1665 (NH). Mahlabatini: Mahlabatini, Gerstner 2825. Msinga: 3 miles S. of Tugela

Ferry, Codd 6336. Ngotche: about -V mile from Pongola Bridge on Magudu road, Edwards 3193.

Nongoma: 11 miles N. of Nongoma, Jessop 673. Ubombo: Mkuze Game Reserve, Ward 3991.

Umzinto: Umpambinyoni River, T/iode 3430 (STE).

Transvaal. — Barberton: Budd’s Farm near Kaap Muiden, Thorncroft 3065 (NH). Lydenburg:

Lulu Mountains, Mogg 16890. Pilgrim’s Rest: 1 mile N.E. of Skukuza, Codd 5718. Soutpansberg:

Ingwe Motel, Jacobsen 1876.

The only significant variation in this species is in the number of cladodes in a

bundle. Although there are few species of Asparagus in South Africa in which both

solitary and numerous cladodes occur on different specimens, this character has been

found so variable in this species as not to be of any value taxonomically.

36. A. scandens Thunb., Prodr. 66 (1784); Bresler, Diss. 5 (1826); R. & S., Syst.

Veg. 7: 325 (1829); Bak. in Saund., Ref. Bot. t. 21 (1869); Journ. Linn. Soc. Bot.

hT 622 (1875); FI. Cap. 6: 268 (1896); Salter in FI. Cape Penins. 174 (1950).

Syntypes: Cape of Good Hope without precise locality, Thunberg s.n. (UPS, a!, b ! ;

BOL!, PRE!, photos).

Asparagus pectinatus Delile in Red., Lil. t. 407 (1813). Type: “ l’orangerie du

jardin la Malmaison ”.

Asparagopsis scandens (Thunb.) Kunth, Enum. PI. 5: 78 (1850).

79

Roots tuberous; the tubers 2-3 cm long, about 0-5 cm broad, borne on long

stalks. Stems weakly scrambling or climbing to 2 m, perennial, glabrous, green,

tortuous, not zigzagging at the nodes. Branches wide-spreading or ascending, angled.

Cladodes ternate, one conspicuously longer than the other two, spreading in the same

plane on all ultimate branches on the same branch, linear-lanceolate, falcate, with a

single prominent vein, bright green, 0-5-1 -5 cm long, 0-8-1 -6 mm broad. Leaves

much-divided, scarious, not spine-forming. Peduncles axillary, usually solitary, occas-

ionally 2-3-nate, 8-12 mm long, articulated in the distal half. Perianth segments

obovate, the outer whorl slightly smaller, the margins entire or minutely toothed,

spreading, 3-4 mm long, white or rarely pinkish. Stamens nearly as long as the perianth

segments; filaments not spurred; anthers yellow, \ of the length of the filaments.

Style 1 mm long; ovary 1 mm long, with 3 ovules per chamber. Berry globose,

1 -seeded, red.

This species grows in shady places where there is perennial moisture, from the

Cape Peninsula in the west to Zitzikama in the east. Generally it is found on coastal

mountains below about 2,000 feet. Flowers are produced irregularly, but have been

recorded in January, September and October.

Cape. — Caledon: Betty’s Bay, Topper 88 (NBG). George: Forest between Wilderness and George,

Story 3561. Humansdorp: Holman's Bos, Britten 1325 (GRA). Knysna: Blaaukrantz Pass, Galpin

4738. Paarl: Franschhoek, Phillips 1345 (SAM). Riversdale: Grootvadersbos, Zeyher 8570b (STE).

Stellenbosch: Jonkershoek, Rodin 3233. Swellendam: Strawberry Hill, Van der Merwe s.n. (STE).

Tulbagh: Tulbaghkloof, Ecklon & Zeyher s.n. Wynberg: Devils Peak, Wolley Dod 1739 (BOL).

37. A. ramosissimus Bak. in Gard. Chron. n. s., 2: 6 (1874); Journ. Linn. Soc.

Bot. 14: 622 (1875); FI. Cap. 6: 268 (1896). Type: “Cooper in Hort. Saunders

5102”.

A. scandens Thunb. var. deflexus Bak. in Journ. Linn. Soc. Bot. 14: 622 (1875);

FI. Cap. 6: 268 (1896). Type: Somerset East, Boschberg, MacOwan 1771 (BOL!,

SAM!).

Stems weak and scrambling or climbing, up to 1 m long, probably usually perennial,

herbaceous. Stems and branches grooved, glabrous, slightly zigzagging, greenish.

Ultimate branches reflexed at the base, similar to the other branches. Cladodes angled

or flattened, linear, 3-nate, 5-15 mm long. Leaves not spine-forming. Peduncles

solitary, 2-10 mm long, articulated distally. Perianth segments similar, entire, obovate,

about 3 mm long, spreading, white. Stamens nearly as long as the perianth segments;

filaments not spurred; anthers up to 1 mm long. Style sometimes at least, unbranched

about 1 mm long; ovary about 1-5 mm long. Berry globose, fleshy, up to nearly,

1 cm diameter, 1- or 2-seeded, red.

This species occurs in moist shady places from the Worcester district through the

southern Cape and Natal up to the eastern Transvaal. Flowers have been recorded

from September to January and in June.

Cape. — Heidelberg: between Port Beaufort and Riversdale, Lewis 5941 (NBG). George: Cradock-

berg, Esterhuysen 30168. Keiskammahoek : Cala Forest Reserve, Story 3269 (GRA). King William’s

Town: Pirie, Flanagan 2235. Mount Ayliff: Insiswa Mountains, Schlechter 6439. Mount Currie:

Mount Currie, Stephany 33 (GRA). Riversdale: Kampscheberg, Muir 2977 (BOL). Somerset East:

Boschberg, MacOwan 1771 (SAM). Stockenstrom: Hogsback, Barker 1901. Stutterheim: Fort

Cunynghame, Galpin 2439. Swellendam: Tradouw Pass, Walgate 914 (NBG). Victoria East: Victoria

East, Rattray 338. Worcester: Welgevonden, Esterhuysen 1902 (BOL).

Basutoland. — Leribe, Pitzeng, Dieterlen 707.

Natal. — Bergville: Ndedena, Esterhuysen 28502 (BOL). Estcourt: Cathkin Park, Howlett 47.

Orange Free State. — Bethlehem: Golden Gate National Park, Roberts 3229.

Swaziland. — N.W. of Mbabane, Compton 30552.

80

Transvaal. — Barberton: Ivy Range, Moodies, Thorncroft 338. Ermelo: Spitskop, Pott 15118.

Pilgrim’s Rest: Graskop, Ga/pin 14531. Pretoria: Wonderboomspruit, Mogg 14754. Wakkerstroom:

Oshoek, Devenish 206.

Baker described A. ramosissimus as “ Wide climbing, leaves obscurely spurred at

the base, cladodes 3- to 8-nate, flattened, linear, falcate, acute, -j- to f inches long,

spreading. Flowers solitary at the tips of the branchlets There are two characters

in this description which are rare in the specimens examined by the present author.

These are the 3- to 8 -nate cladodes and the terminal flowers. Both of these characters

do occur, however, and there is no other species which fits this description. In 1875,

Baker cited “ Cooper in Flort. Saunders 5102 ”, but added that the description was

made from a figure by Fitch. This drawing does not appear to have been published,

and Moss (Ms. in National Herbarium, Pretoria) could not find a specimen of the

species among the specimens in Cooper’s complete set at Kew.

In 1896, Baker wrote that his description of A. ramosissimus was from a living

plant in Wilson’s garden at Reigate. It is likely that no specimen has been preserved,

and no other specimen was cited in any of Baker’s works.

38. A. crispus Lam., Encyc. 1: 295 (1783); Bresler Diss. 22 (1826); R. & S.

Syst. Veg. 7: 326 (1829); Kunth, Enum. PI. 5: 73 (1850); Bak. in Journ. Linn. Soc.,

Bot. 1 4 : 607 (1875); FI. Cap. 6: 259 (1896); Marloth, FI. S. Afr. pi. 20(1915); Salter

in FI. Cape Penins. 174 (1950). Type: “ L’lsle de France”, collector unknown, (P,

holo.; PRE, photo.!).

A. flexuosus Thunb., Prodr. 66 (1794); Kunth, Enum. PI. 5: 74 (1850). Type:

Cape of Good Hope without precise locality, Thunberg s.n. (UPS, holo.!; BOL!,

PRE!, photos). A. decumbens Jacq., Hort. Schoenbr. 1: 51, t. 97 (1797); Bresler,

Diss. 21 (1826). Type: Cape of Good Hope without precise locality, collector unknown.

Asparagopsis decumbens (Jacq.) Kunth, Enum. PI. 5: 77 (1850).

Roots tuberous; the tubers borne on short stalks, 3-6 cm long, 0-5-1 -5 cm

broad. Stems annual, weak, straggling or climbing, usually up to 1 m long, much-

branched, angled, zigzagging at the nodes, glabrous, green. Branches reflexed at the

base, curving upwards, similar to the stems. Cladodes ascending, ternate, except

where a normal branch is formed when they are paired, borne at nodes towards the

ends of branches, soft or slightly firm, slightly arcuate, flattened or angled, 3-9 mm long,

less than 1 mm broad and usually less than \ mm broad. Leaves frequently forming

soft spine-shaped processes, grey. Peduncles solitary, axillary, 6-1 1 mm long, articulated

distally. Buds cylindrical. Perianth segments reflexed from their centres, but forming

a cylinder proximally; the base of the flower truncate; segments similar, entire, linear-

oblong, whitish with a pale brown or green streak, 4-5 mm long. Stamens about

3 mm long, orange; filaments broadened towards their bases, and with minute spurs.

Style 1-5 mm long; ovary 1-5 mm long, with 2-4 ovules in each chamber. Berry

ovoid, 3-9-seeded, 8-15 mm long, pale coloured; style persistent and conspicuous;

perianth persistent.

This species grows in a wide range of habitats from coastal sands to rich mountain

soils, but has a rather restricted distribution. It occurs in the south-western Cape from

St. Helena in the north, round the coast as far as Swellendam. It also grows as far

inland as Worcester. Flowers have been recorded from June to October.

Cape. — Caledon: Van Niekerk 739 (BOL). Cape: Robben Island, Walgate 494 (NBG); Signal

Hill, Dimmer 1600 (SAM). Clanwilliam: between the Olifant’s River and Brakfontein, Ecklon &

Zeyher s.n. Humansdorp: mouth of Klein River, Zeyher 41 55 (BOL). Malmesbury: Darling Flora

Reserve, Rycroft 1931 (NBG). Paarl: Tigerberg, Pillans 8657 (BOL). Robertson: Robertson, Van

Niekerk 383. Simonstown: Lakeside Plateau, Jessop 62 (BOL). Somerset West: Knorhoek, Jessop

61 (BOL). Stellenbosch: Jonkershoek, Duthie 527 (STE). Tulbagh: Saron, Schlechter 7892 (GRA).

Worcester: Karroo Garden, Leighton 1037 (NBG). Wynberg: Kirstenbosch, Salter 7356 (BOL).

81

Lamarck described this species from a specimen in the Lamarck Herbarium. This

specimen is described as having been collected on Mauritius. The only Mauritian

species is A. umbellu/atus Bresler, which differs in having fascicled cladodes, flowers

not solitary, peduncles articulated near the centre and a globose fruit. The locality

must, therefore, either be wrongly given or the plant must have been cultivated.

The illustration by Jacquin includes an accurate drawing of the flower, which

shows the truncate base and cylindrical proximal part characterising A. crispus Lam.

39. A. asparagoides (L.) Wight in Century Dictionary II: 845 (1909); Salter in

FI. Cape Penins. 173 (1950). Type: Tilli, Cat. Plant Horti Pisani t. 12, f. 1,2 (1723).

Medeola asparagoides L., Sp. PI. 339 (1753); Delile in Red., Lil. t. 442 (1816). M.

angustifolia Mill., Gard. Diet. ed. 8 (1768). Type: Tilli, Cat. Plant. Horti Pisani t. 12

“ myrti folio angustiore ” (1723).

Dracaena medeoloides Linn, f., Suppl. 203 (1781). Type: Cape of Good Hope

without precise locality, Thunberg s.n. (UPS, holo.!; BOL!, PRE!, photos). D. vo/u-

bilis Linn, f., Suppl. 204 (1781). Type: Cape of Good Hope without precise locality,

Thunberg s.n. (UPS, holo.!; BOL!, PRE!, photos).

Asparagus volubilis (Linn, f.) Thunb., Prodr. 66 (1794); Bak. in Journ. Linn. Soc.

Bot. 14: 628 (1875); FI. Cap. 6: 272 (1896). A. medeoloides (Linn, f.) Thunb., Prodr.

66 (1794); Marloth, FI. S. Afr. 4: pi. 20 (1915). A. krausii Bak. in Journ. Linn. Soc.

Bot. 14: 628 (1875); FI. Cap. 6: 272 (1896), nom illegit. Type: as for Myrsiphyllum

krausianum Kunth. A. krausianus (Kunth) MacBride in Contrib. Gray Herb. 56:

17 (1794); Salter in FI. Cape Penins. 174 (1950), sub A. kraussianus. A. kuisibensis

Dinter in Fedde, Rep. 29: 270 (1931). Type: S.W.A., Kuisib, Tjuezu in Herb. Dinter

4698 (B, holo.!). A. ovatus Salter in Journ. S. Afr. Bot. 6: 167 (1940); FI. Cape

Penins. 174 (1951); Dyer in Flow. PI. Afr. 29: t. 1146 (1943). Type: Cape, Rugby,

Salter 8214 (BOL, holo!.; NBG!, PRE!). A. multitub erosus Dyer in Bothalia 6: 442

(1954). Type: Ceres, Karoopoort, Marloth 9006 (PRE, holo.!; STE!).

Ruscus volubilis Thunb., Prodr. 13 (1794); Kunth, Enum. PL 5: 276 (1950). Type:

Cape of Good Hope without precise locality, Thunberg s.n. (UPS, holo.!; BOL!,

PRE!, photos).

Myrsiphyllum asparagoides (L.) Willd. in Ges. Naturf. Fr. Berl. Mag. 2: 25 (1808);

Kunth, Enum. PI. 5: 105 (1850); Hook. f. in Bot. Mag. t. 5584 (1866). M. angusti-

folium (Mill.) Willd. in Ges. Naturf. Fr. Berl. Mag. 2: 25 (1808); Kunth, Enum. PI.

5: 106 (1850). M. falciforme Kunth, l.c., 107 (1850). Type: Cape without precise

locality, Drege 2704a in “Herb. Luc.” (K, iso.!). M. gramineum Kunth, l.c., 108

(1850). Type: “ M. striatum Schlechtend. in Herb. reg. Berol., excl. synon. (v..s\) etl

Hort. Berol. 1847 ”. M. krausianum Kunth, l.c., 107 (1850). Type: Wynberg, Con-

stants, Krauss 1333.

Hecatris asparagoides (L.) Salisb., Gen. Plant. 66 (1866).

Tubers varying from almost sessile to 7 cm distant, usually 5-15 cm long. Stems

climbing to 3 m, scrambling or under exceptional circumstances erect, tortuous, glabrous,

terete or angled, annual. Branches similar to the stems, spreading or ascending.

Cladodes broadly ovate to lanceolate, with an obtuse base, sessile, 1-7 cm long, up

to about 7 prominent veins and numerous smaller ones. Leaves scarious, not forming

spines. Peduncles 1- or 2-nate, 3-8 mm long, articulated near the flower. Perianth

segments similar, entire, linear-oblong, 5-6 mm long, spreading at first, later reflexed

from the centre of each segment, white. Stamens nearly as long as the perianth

segments; anthers 0-5 mm long; filaments with two basal spurs. O vary 2 mm long;

style 2-3 mm long, the branches sometimes only loosely fused; ovary chambers with

4-9 ovules. Berry globose, 6-10 mm diameter, up to 8-seeded, red; perianth persistent.

82

It is distributed from the south-western Cape through the eastern parts of South

Africa and north to Kenya and the Congo. The flowering period is mainly June to

September, but varies according to the time of the rainfall. In the Transvaal, flowering

usually occurs in January and February.

Cape. — Albany: Frasers Camp, Maguire 636 (NBG). Alexandria: near Sandflats, Burtt Davy

14249 (BOL). Barkly East: Barkly Pass, Rattray 7324. Bathurst: Kowie River, MacOwan 1920

(SAM). Bellville: De Grendel, Esterhuysen 23057 (BOL). Bredasdorp: Strandkloof, Compton 14778

(NBG). Caledon: Uilenkraal, Taylor 1 584 (SAM). Calvinia: near Middelpost, Salter 3484 (BOL).

Cape: near Rugby, Salter 8214. Ceres: Karroopoort, Mar/oth 9006. Clanwilliam: Brakfontein,

Schlechter 7974. Colesberg: Arundel, Acocks 17992. East London: near Cove Rock, Galpin 3102,

Fort Beaufort : near Fort Beaufort, Story 2226. George: Wilderness, Van Niekerk 251 (BOL). Graaff-

Reinet: Graaff-Reinet, Bolus 463 (BOL). Humansdorp: Ratelsbos, Fourcade 53 (BOL). Kentani.

Kentani, Pegler 1240 (BOL). King William’s Town: Amabele, De Vries 23. Kirkwood: Addo

Bush, Mar loth 6439. Knysna: Knysna Village, Fourcade 5885 (BOL). Komga: Prospect Farm,

Flanagan 315. Laingsburg: White Hill, Compton 11200 (NBG). Malmesbury: Darling Flora Reserve,

Rv croft 1969 (NBG). Montagu: Montagu, Levyns s.n. (BOL). Namakwaland: Silverfontein near

O’okiep, Drege 2704b (K). Peddie: near Wooldridge, Acocks 1 1886. Piketberg: Avontuur Mountain,

Pillans 7571 (BOL). Port Elizabeth: Perseverence, Paterson 67 (GRA). Queenstown: Invana,

Cooper 328 (BOL). Riversdale: Vet Rivier, Muir 353. Simonstown: Smitswinkel, Wolley Dod 3009

(BOL). Somerset East : Boschberg, MacOwan s.n. (SAM). Somerset West: Fir Grove, Parker 4485

(BOL). Stellenbosch: near Mill Stream, Duthie 1479 (STE). Stockenstrom : Andriesberg, Galpin

2536 (GRA). Tulbagh: Saron, Schlechter 4876 (GRA). Uitenhage: Enon, Thocle A2111. Uniondale:

near Laudina Store, Acocks 14643. Vanrhynsdorp: Giftberg, Phillips 7619 (SAM). Wellington:

Wellington, Thomson s.n. Worcester: De Dooms, Geriche s.n. Wynberg: Kirstenbosch, Salter

7596 (BOL).

Basutoland. — Leribe, Dieterlen 102.

Natal. — Bergville: Cathedral Peak Forest Reserve, Killick 1723. Camperdown: Mid Illovo:

Thode 3432 (STE). Eshowe: Eshowe, Gerstner 3754 (NH). Inanda: Inanda, Medley Wood 623

(NH). Kranskop: Kranskop, Thode 3419 (STE). Lions River: Dargle Forest, Mol! 1233. Ngotche,

Ngome Forest, Gerstner 4396. Pietermaritzburg: Allerton, Mogg 6620.

Orange Free State. — Bethlehem: Clarens, Van Hoepen s.n. Bloemfontein: Glen, Mostert 363.

Lindley: near Lindley, Phillips s.n. (BOL).

Swaziland. — Near Hlatikulu, Compton 26393 (NBG).

Transvaal. — Barberton: Umvoti Creek, Galpin 857. Belfast: near Machadadorp, Bruce 481.

Ermelo: Ermelo, Rogers 14782 (BOL). Johannesburg: Jeppe, Gilmore 520. Krugersdorp: near

Krugersdorp, Mogg 23103. Letaba: Duiwelskloof, Scheepers s.n. Lichtenburg: Grasfontein, Sutton

581. Middelburg: Boschhoek, western Steenkampsberg, Young A384 (NH). Petersburg: Blaauw-

berg, Dyer 9152. Pilgrim’s Rest: Mariepskop, Van der Schijjf 4784. Potchefstroom : Nooitgedacht,

Cohen s.n. Wakkerstroom : Wakkerstroom, Beeton 124 (SAM).

South West Africa. — Grootfontein: Otavibirge, Dinter 743 (SAM); Otavi, D inter 5547 (B).

Linnaeus, in his Species Plantarum, cited M. A. Tilli’s “ Catologus Plantarum

Horti Pisani ” (1723) t.12, f.l & 2. There is no specimen labelled Medeola asparagoides

in the Linnaean Herbarium so that Tilli’s figure, which shows specimens with solitary,

many-veined cladodes and tortuous stems should be regarded as the iconotype, as

this was the only illustration cited.

Miller regarded the specimen in Tilli’s plate with larger cladodes as the type of

Linnaeus’ species, and coined the name Medeola angustifolia for the form with smaller

cladodes shown in the other figure on the same plate.

Dracaena volubilis Linn. f. was based on a Thunberg specimen from the Cape.

The specimen in the Linnaean Herbarium, a photograph of which has been examined

by the author, has ternate cladodes as opposed to the alternate cladodes mentioned

in the type description and according to Jackson (1912), it does not occur in the

enumerations of the Linnaean Herbarium. It can, therefore, not be considered as

a type. The sheet in the Thunberg Herbarium labelled Asparagus volubilis has three

specimens which agree with the younger Linnaeus’ description of Dracaena volubilis

and with the present author’s concept of Asparagus asparagoides. Thunberg's

A. volubilis was described in the same words as Dracaena volubilis Linn, f., and

83

reference was made to “ Linnaeus Syst. 334 The only reference to this species earlier

than Thunberg’s Prodromus appears to be the Supplementum of the younger Linneaus.

Stearn (1961) mentions that both Linnaeus the younger and Thunberg played a part

in adding to the elder Linnaeus’ manuscript of this work. It is therefore very likely

that “ Linnaeus Syst. 334 ” refers to this manuscript.

15 20 50 40 50 70 Km

CLADO&E LENGTH

Fig. 3. — Scatter diagram and histograms showing cladode length and breadth, and the relation of

these characters to one another, in Asparagus asparagoides

84

The holotype of Myrsiphyllum falciforme was cited by Kunth as being in “ Herb.

Luc.”. This specimen was, therefore, likely to have been in Kiel, but is not there now,

and has probably been destroyed. The specimens cited by Kunth under his M.

gramineum are all likely to have been destroyed in Berlin. He described this species

as a climber with flattened, solitary cladodes, and there can, therefore, be no reasonable

doubt as to the correct application of the name. Krauss 1333, which is the type of

M. kraussianum, has not been traced. Kunth’s description, however, clearly places

it in the synonymy of this species. He spelt the collector’s name Krause, but the correct

spelling is Krauss, and the correct spelling of the binomial should have been M.

kraussianum.

There is considerable variation in the size and shape of the cladodes of A. aspara-

goides as construed by the present author. Both A. krausianus and A. volubilis were

described from specimens with lanceolate cladodes, while A. ovatus and A. multi-

tuberosus were described from specimens with ovate cladodes. A. asparagoides appears

to have been described from an illustration showing cladodes of an ovate-lanceolate

foim in the left hand figure and lanceolate cladodes in the right hand figure. As A.

kraussianus sensu Salter and A. asparagoides sensu Salter (1950) are frequently found

in similar environments and sometimes near one another, it seems likely that their

difference is genotypic, and not merely the effect of different habitats. In order to

determine the taxonomic value of cladode shape in this group in South Africa, the

scatter diagram and histograms (Fig. 3) were drawn. These show that there is no

justification for subdivision of A. asparagoides , as construed here, on the basis of cladode

shape or size. In the eastern Cape there is a narrow-cladode form with a strong garlic-

like smell. Without any other subsidiary character, however, it was felt that this form

could not be given taxonomic recognition.

The other important character which has been used for the separation of species

in this complex is the tuber arrangement. The tubers of A. ovatus were described as

distant as opposed to those of A. asparagoides sensu Salter (1950), which were sessile.

The tuber character of A. ovatus may be associated with the loose sand in which the

plants occur. The typical form of A. asparagoides with sessile tubers generally occurs

in soils of a heavier texture. Parker collected specimens which he claimed had both

sessile and distant tubers and, in the experience of the present author, it is, often

impossible to define the difference between sessile and distant tubers as there is a complete

gradation between these two forms. There is also a gradation between the rooting

system in A. multitub erosus, where the tubers are very numerous and only about 5 cm

long, and that in A. asparagoides sensu Salter in which there are about twelve tubers,

each about 10-15 cm long. The specimens which have been placed in A. multitub erosus

come from dry areas with very hard soils such as occur on the Giftberg near Van-

rhynsdorp.

40. A. undulatus {Linn. /.) Thunb., Prodr. 66 (1794); Bak. in Journ. Linn. Soc.

Bot. 14: 628 (1875); FI. Cap. 6: 273 (1896); Marloth, FI. S. Afr. 4: pi. 20 (1915);

Salter in FI. Cape Penins. 173 (1950); Solch, Beitr. FI. Siidwest-Afr. 39 (1961). Type:

Cape of Good Hope without precise locality, Thunberg s.n. (UPS, holo.!; BOL!,

PRE!, photos).

Dracaena undulata Linn, f., Suppl. 203 (1781); R. & S., Syst. Veg. 7: 346 (1829).

Myrsiphyllum undulatum (Linn, f.) Kunth, Enum. PI. 5: 109 (1850).

Asparagus klinghardtianus Dinter in Fedde, Rep. 29: 270 (1931). Type: S.W.A.,

Klinghardtgebirge, Dinter 3998 (B, holo.!; PRE!).

Tubers distant, 0-8-15 mm broad, 25-50 mm long. Stems erect, frequently with

minute obtuse processes giving them a rough surface, annual, up to 60 cm high, but

usually less, angled, the edges often produced into wings in the upper parts, green.

85

Branches similar to the stems, ascending. Cladodes solitary, sessile, ovate to lanceolate,

undulate and often folded towards the tips, acutely attenuate, with prominent veins

which are sometimes produced into longitudinal wings or ridges, scabrid, 8-40 mm

long. Leaves deltoid, not forming spines. Peduncles 1-3-nate, 5-10 mm long, articu-

lated in the distal third. Perianth segments similar, entire, linear-oblong, the distal

half spreading or reflexed, 6 mm long, purplish or greenish. Stamens almost as long

as the perianth segments; filaments spurred; anthers 0-5 mm long. Ovary 2 mm long;

styles 2-5 mm long. Berry 5-6 mm diameter, 1-3-seeded, red; perianth persistent.

This species is particularly abundant on the flat sandy area stretching from the

Cape Peninsula north to near Citrusdal. It also occurs in the drier areas of Nama-

kwaland and S.W.A Flowers have been recorded from July to October.

Cape. — Bellville: Tigerberg, Pillans 8660 (BOL). Calvinia: Glen Lyon, Lewis 5726 (NBG). Cape:

Signal Hill, Thode 6079 (STE). Clanwilliam: Clanwilliam, Schlechter 8416. Hopefield: near Hopefield,

Bolus 12858 (BOL). Malmesbury: near Groenkloof, Bolus 3690 (Z). Namakwaland: Witbank,

Pillans 5222 (BOL). Piketberg: south of Piketberg, Barker 5775 (NBG). Stellenbosch: near Libertas,

Duthie 542 (STE). Tulbagh: Tulbagh New Kloof, Compton 11708 (NBG).

South West Africa. — Grootfontein: Otavi, Dinter 5547 (Z). Luderitz: Klinghardtgebirge,

Dinter 3998. “Ex reg. : Hereroland ”, Dinter 623 (Z).

There is a certain amount of variation in cladode breath of specimens from different

areas. For example, the specimens from north of Citrusdal have particularly narrow

cladodes. There is, however, a complete gradation between the different cladode

shapes.

Doubtful Species

A. confertus Krause in Bot. Jahrb. 51: 449 (1914); Solch, Beitr. FI. Siidwest-Afr.

40 (1961). Type: Luderitz, Schakalskuppe, Engler 6764 (B, holo.!).

Material matching the type shows many similarities with A. krebsianus, but differs

mainly in having shorter cladodes, which are about 10-15 mm long. These specimens

have been collected only in the Luderitz district of South West Africa. A. krebsianus

has not been recorded closer to this than the central districts of the Transvaal. In

view of the paucity of material, the smaller cladodes and the considerable gap between

this population and A. krebsianus, it is considered best not to refer these specimens

to that species. Similarly it does not seem justified to uphold A. confertus without

further material.

South West Africa. — Luderitz: Aus, Dinter 6155 (B); Schakalskuppe, Engler 6764 (B); Range

1715 (B, SAM).

A. declinatus L., Sp. PI. 313 (1753). Type: No specimen or earlier description

cited.

As there is no specimen in the Linnaean Herbarium, as Linnaeus did not cite any

earlier work on which this binomial could have been based, and as the description,

which does not even give the geographical area from which the plant came, is com-

pletely inadequate for identifying the species, the name must be regarded as a nomen

dubium. Baker (1875) used the name for a form which is treated here as A. setaceus,

and A. declinatus sensu Baker is therefore discussed under that species.

A. nivenianus Schult. in R. & S., Syst. Veg. 7: 331 (1829). Type: Cape without

precise locality, Niven s.n.

The type specimen of this binomial has not been located. It may have been

destroyed in Berlin. From the description it is not possible to tell whether the Niven

specimen should be referred to A. africanus or A. thunbergianus, and the name must

therefore be regarded as a nomen dubium. Kunth, when he made the combination

in Asparagopsis, cited Ecklon 95b from the Cape. This specimen is referable to

A. thunbergianus.

86

References

ARBER, A., 1924. Myrsiphyllum and Asparagus: a morphological study. Ann. Bot. 38: 635-659

ARBER, A., 1925. Monocotyledons: a morphological study. London: Cambridge University

Press.

BAKER, J. G., 1875. A Revision of the Genera and Species of Asparagaceae. Journ. Linn. Soc.

Bot. 14: 508-632.

BAKER, J. G., 1896. Asparagus in: FI. Cap. 6: 256-274. London: L. Reeve & Co.

BRESLER, M., 1826. Generis Asparagi historia naturalis atque medici. Berlin.

DE CANDOLLE, A., 1880. La Phytographie ou l’art de decrire les Vegetaux. Paris: G. Masson,

Editeur Libraire de FAcademie de Medecine.

JACKSON, B. D., 1912. Index to the Linnaean Flerbarium. Suppl. to Proc. Linn. Soc. Lond.,

124th session.

KAUSSMANN, B., 1955. Histogentische Untersuchungen zum Flachsprossproblem. Bot. Studien 3,

KIES, P., 1951. Notes on Asparagus africanus and related Species. Bothalia 6: 177-181.

KUNTH, K. S., 1850. Enumeratio Plantarum 5. Stuttgart and Tubingen: J. G. Cotta.

LANJOUW, J. & STAFLEU, F. A., 1954. Index Herbariorum, pt. II, Collectors, 1st instalment.

LINNAEUS, C., 1753. Species Plantarum, ed. 1. Stockholm: Laurentius Salvius.

PHILLIPS, E. P., 1951. Genera of South African Flowering Plants. Pretoria: Government Printer.

SALTER, T. M., 1950. Asparagus in: FI. Cape Penins. CaDe Town and Johannesburg: Juta &

Co., Ltd.

SCHLITTLER, J., 1953. Bliitenartikulation und Phyllokladien der Liliaceae organophylogenetische

betrachtet. Fedde, Rep. 55: 154-258.

SCHLITTLER, J., 1959. Die Asparageenphylokladien erweisen sich auch ontogenetisch als Blatter.

Bot. Jahrb. 79: 428-446.

SOLCH, A., 1961. Beitrage zu einer Flora Stidwest-Afrikas. Munich: A. Solch.

STEARN, W. T., 1961. In: Cat. Books R. M. M. Hunt 2, 1: civ.

ROEMER, J. J. & SCHULTES, J. A., 1829. C. Linne, Systema Vegetabilium 7. Stuttgart: J. G.

Cotta.

TRIMEN, H., 1898. Handbook of the Flora of Ceylon. London: Dulau & Co.

Index

Asparagopsis aethiopica (L.) Kunth, 68

albus (L.) Kunth, 39

consanguinea Kunth, 54

decumbens (Jacq.) Kunth, 80

densiflora Kunth, 65

denudata Kunth, 52

dregii Kunth, 55

floribimda Kunth, 72

juniperina Kunth, 48

krebsiana Kunth, 73

lamarckii Kunth, 48

lancea (Thunb.) Kunth, 68

microrhaphis Kunth, 61

minutiflora Kunth, 63

niveniana sensu Kunth, 55, 85

passerinoides Kunth, 43

retrofracta (L.) Kunth, 58

scandens (Thunb.) Kunth, 78

schlechtendalii Kunth, 48

setacea Kunth, 51

spinescens (Schult.) Kunth, 46

stipulacea (Lam.) Kunth, 45

subquadrangularis Kunth, 72

thunbergii Kunth, 55

triancantha (Willd.) Kunth, 46

zeyheri Kunth, 46

Asparagus acocksii Jessop, 74, pi. 7

aethiopicus L., 67, 71

TlllUpiLLlb U/, / I

var. aethiopicus, fig. 1, 68, 70

var. angusticladus Jessop, 69, pi. 3

var. natalensis Bak., 70

var. ternifolius Bak., 69, 70

africanus Lam., 33, 48, 85

var. concinnus Bak., 60

var. deoendens (thunb.) Bak., 51

var. microphylhis Bak., 58

var. wright ii Bak., 59

albus sensu Thunb., 55

albus L., 55

asiaticus sensu Bak., 48

asiaticus L., 50

asparagoides ( L .) Wight, 33, 35, 39, 81

aspergillus Jessop, 71 pi. 4

bechuanicus Bak. 48

buchananii Bak. 67

burchellii Bak., 45

burkei Bak., 48

capensis L., 31, 33, 39, 43, 45

capensis sensu Marloth, 55

var. capensis, 43

var. litoralis Suess. & J. Karl., 44

compactus Salter, 33, fig. 1, 59, 73

87

concinnus Kies, 60

confertus Krause, 74, 85

conglomerate Bak., 48

consanguineus (Kunth) Bak., 54

cooper i Bak., 48

crassicladus Jessop, 75, pi. 8

crispus Lam., 33, 80

declinatus sensu Bak., 51, 57

declinatus L., 85

decumbens Jacq., 80

densiflorus (Kunth) Jessop, 39, 65

densus Bak., 45

denudatus (Kunth) Bak., 33, fig. 1, 52

dependens Thunb., 48

dinteri Engler & Krause, 48

ecklonii Bak., 62

engleri Krause, 77

erectus (Linn, f.) Thunb., 77

exuvialis Burch., 33, 35, 37, 62

falcatus L., 66, 69

var. falcatus, 70

var. ternifolius (Bak.) Jessop, 70

fasciculatus Thunb., 37, 39, 54

fleckii Schinz, 48

flexuosus Thunb., 80

francisci Krause, 48

glaucus Kies, 35, 47

gonocladus Bak., 66

hereroensis Schinz, 62

intangibilis Dinter, 46

judtii Schinz, 48

juniperoides Engler, 35, 38, 63

klinghardtianus Dinter, 84

krausii Bak., 81

krausianus (Kunth) MacBride, 81

krebsianus (Kunth) Jessop, 39, 73, 85

kuisibensis Dinter, 81

lanceus Thinb., 68

laricinus Burch., 59, 60

lugardii, Bak., 48

macowanii Bak., 57

var. macowanii, 57

var. zuluensis (N. E. Br.) Jessop, 58, 61

medeoloides (Linn, f.) Thunb., 81

meyeri nom. nud., 66

microphyllus (Bak.) Kies, 58

microrhaphis (Kunth) Bak., 61

minutiflorus (Kunth) Bak., 33, 63, 64, 66

mucronatus Jessop, 56, pi. 1

multiflorus Bak., 48

multituber osus Dyer, 81

myersii nom. nud., 65

myriocladus Bak., 65

namaensis Schinz, 52

nelsii Schinz, 33, 75, 76

nelsonii Bak., 43

nivenianus Schult., 85

obermeyerae Jessop, 73, pi. 5, 6

officinalis L., 37, 39

omahekensis Krause, 46

ovatus Salter, 81

oxyacanthus Bak., 63

patens Krause, 48

pearsonii Kies, 52

pectinatus Redoute, 78

pi /osus Bak., 48

plumose Bak., 51

racemose sensu Solch, 71

racemosus Willd., 68, 71, 72, 73, 74, 75

var. tetragonus (Bresler) Bak., 72

ramosissimus Bak., 79

retrofractus, L., 33, 39, 58, 61

rigidus Jessop, 33, 64, 66, pi. 2

rivalis Kies, 48

sarmentosus sensu Bak., 65

sarmentosus L., 66

saundersiae Bak., 66, 67

scandens Thunb., 33, 78

var. de fie xus Bak., 79

setaceus (Kunth), Jessop, 33, 51, 85

spinescens Schult., 45

spinosissimus O. Kuntze, 46

sprengeri Regel, 65

sprengeri sensu Wright, 69

stellate Bak., 61

stipulaceus Lam., 33, 39, 44

stipulaceus sensu Bak., 46

striatus (Linn, f.) Thunb., 35,^76

suaveolens Burch., 45, 48

subulatus Thunb., 78

sylvaticus Burch. Ms., 54

ternifolius sensu Hooker f., 69

tetragonus Bresler, 72

thunbergianus Schult., 39, 55, 57, 85

triacanthus Burm. f., 43

triacanthus Willd., 45

umbellulatus Bresler, 81

undulatus (Linn.f.) Thunb. , 35, 84

virgatus Bale., 53

volubilis (Linn, f.) Thunb., 81

zuluensis N. E. Br., 58

Dracaena erecta Linn, f., 77

medeoloides Linn, f., 81

striata Linn, f., 77

stricta Schult., 77

undulata Linn, f., 84

volubilis Linn, f., 81

Hecatris asparagoides (L.) Salisb., 39, 81

Medeola angustifolia Miller, 81

asparagoides L., 39, 81

Myrsiphyllum angustifolium (Miller) Willd., 81

asparagoides (L.) Willd., 39, 81

erectum (Linn, f.) Kunth, 77

falciforme Kunth, 81

gramineum Kunth, 81

krausianum Kunth, 81

striatum (Linn, f.) Kunth, 77

undulatum (Linn, f.) Kunth, 84

Ruscus volubilis Thunb., 81

x

89

Plate I. — Asparagus mucronatus Jessop (Theron 551, holotype).

7230691

90

Plate 2. — Asparagus rigidus Jessop (Kil/ick & Strey 2380, holotype).

91

/5 /.

'f/'C ‘

r*fort 4

Plate 3. — Asparagus aethiopieus L. var. angusticladus Jessop (Mogg 22470, holotvpe).

92

Plate 4. — Asparagus aspergillus Jessop ( Breyer in TRV 19063, holotype).

93

Plate 5. — Asparagus obermeyerae Jessop ( Mauve 4442, holotype).

94

Plate 6.

Asparagus- obermeyerae Jessop ( Hafstrom & Acocks\91).

95

N A noNAI. (IEKBARILrM,

PRETORIA.

DISTRICT <\

CM,- ' Hf;€fee ■'/■ Vo.

Ycm. Name

Locality

Habitat

Dcscriyjion etc.

SSi)V

NAS ION M

N \TION M

III- iUlMUl M, I*Rl TORI

> j-' '

•Ki -C rt H j

. i, v IbAlo

Plate 7. — Asparagus acocksii Jessop ( Acocks 16296, holotype).

96

UNIVERS'TY OF C A°F TOWN

. , (. U \ •< u • M Arm '«M:M P

l).u-niMi. ,.i il* ii ssor i" '■

Plate 8. — Asparagus crassicladus Jessop ( Esterhuysen 4670, holotype).

97

Bothalia 9, 1: 97—112

Notes on South African Cruciferae

by

W. Marais

Abstract

A new genus in the section Arabideae, sixteen new species and four new varieties are described,

and eleven new combinations made: — Heliophila arenaria Sond. var. acocksii — var. glabrescens (Schulz)

(H. sabulosa Schltr. var. glabrescens Schulz),' — var. agtertuinensis (Schulz) (H. maximilianii Schltr.

var. agtertuinensis Schulz), H. diffusa (Thunb.) DC. var. flacca (Sond.) (H. flaeca Sond.), H. elata

Sond. var. pillansii, H. latisiliqua E. Mey. ex Sond. var. macrostylis (E. Mey. ex Sond.) (H. macrostylis

E. Mey. ex Sond.), H. linearis (Thunb.) DC. var. linearifolia (Burch, ex DC.) (H. linearifolia Burch,

ex DC.), — var. reticulata (Eckl. & Zeyh.) ( H . reticulata Eckl. & Zeyh.), H. meyeri Sond. var. minor,

H. pusilla L.f. var. maerosperma, — var. setacea (Schltr.) (H. setacea Schltr.), — var. lanceolata (Adamson)

(H. lanceolata Adamson), H. alpina, H. cedarbergensis, H. cuneata, H. esterhuyseniae, H. eximia, H.

filieaulis, H. katbergensis, H. obibensis, H. promontorii, H. rimicola, H. schulzii, Lepidium divarication

Ait. subsp. trifurcum (Sond.) ( L . trifurcum Sond.), L. basuticum, L. suluense, L. transvaalense, Thlas-

peocarpa nama uensis, APLANODES, type A. doidgeana, A. sisymbrioides (Schltr.) ( Heliophila sisym-

brioides Schltr.) Brachycarpaea juncea (Berg.) ( Cleome juncea Berg.).

In continuation of my studies on the South African Cruciferae for the Flora of

Southern Africa (see Bothalia 8: 166-169, 1964), further combinations and new

descriptions are required.

In the lists of specimens cited, no herbarium reference is given for collections

represented in the National Herbarium, Pretoria.

HELIOPHILA

Heliophila arenaria Sond. in Abh. Ges. Naturw. Hamb. 1: 262 (1846). Type:

Drege 7568 (K!).

(а) var. arenaria.

H. arenaria Sond., l.c. (1846); FI. Cap. 1: 53 (1860). H. maximilianii Schltr. in

Bot. Jahrb. 27: 136 (1899). Type: Schleehter 8229 ( B ! ). H. sabulosa Schltr., l.c. 138

(1899). Type: Schleehter 8339 ( B ! ). H. lightfootii Phillips in Ann. S. Afr. Mus. 9:

111 (1913). Type: Phillips 7683 (SAM!).

(б) var. acocksii Marais , var. nov., a typo siliquis crassioribus tnarginibus fere

rectis et stylis in fructu longioribus differt.

Type: Calvinia, Lokenburg, Acocks 19750 (PRE, holo. ; K).

Cape. — Vanrhynsdorp: Tigerberg, Lewis 4959 (SAM); Urionskraal, Barker 8554 (NBG). Calvinia:

Lokenburg, Acocks 19750; Leistner 317; between Nieuwoudtville and Oorlogskloof, Leipoldt 3960;

between Nieuwoudtville and Graskop, Lewis in BOL 19823 (BOL); Botterkloof Pass, Maguire 196,

p.p. (NBG); Lewis 3124 (SAM); 3125(SAM); Compton 20898 (BOL, NBG). Clanwilliam: Brandewyn

River, Maguire 245 (NBG); Pakhuis, Compton 19972 (NBG); between Pakhuis and Wupperthal,

Bolus s.n. (BOL); Leipoldt 3968; between Pakhuis and Bidouw, Bolus 8922; Wupperthal, Thode

1960; between Nardouws Pass and Brakvlei, Marais 1436.

(c) var. glabrescens ( Schulz ) Marais, comb. nov.

H. sabulosa var. glabrescens Schulz in Bot. Arch. 31: 541 (1931). Type: Stephens

& Glover 8705 (K!).

( d ) var. agtertuinensis (Schulz) Marais, comb. nov.

H. maximilianii var. agtertuinensis Schulz, l.c., 31: 540 (1931). Type: Schlechter

10858 (B!).

Heliophila diffusa (Thunb.) DC., Syst. 2: 685 (1821). Type: Thunberg Herb.

No. 14956.

(a) var. diffusa.

H. diffusa (Thunb.) DC., l.c. (1821); Sond. in Abh. Ges. Naturw. Hamb. 1: 220

(1846); FI. Cap. 1: 42 (1860). H. peltaria DC., l.c. (1821); Sond. in Abh. Ges.

Naturw. Hamb. 1: 222 (1846); FI. Cap. 1: 43 (1860). Type: Thunberg Herb. No.

14958. H. capensis (L.f.) C. A. Smith in Kew Bull. 1931 : 155 (1931), non (L.f.) Kuntze

(1898).

Lunaria diffusa Thunb., Prodr. 107 (1800). L. pinnata Thunb., l.c. (1800). Type:

Thunberg Herb. No. 14958.

Peltaria capensis L.f., Suppl. 296 (1781). Type: LINN 829.3 or 829.4.

(b) var. flacca (Sond.) Marais, stat. nov.

H. flacca Sond. in Abh. Ges. Naturw. Hamb. 1: 223, t.22, f.l (1846); FI. Cap.

1: 43 (1860). Type: Ecklon & Zeyher 69 (K!).

Heliophila elata Sond. in Abh. Ges. Naturw. Hamb. 1: 247 (1846). Type: Drege

7566a (K!).

(o) var. elata.

H. elata Sond., l.c. (1846); FI. Cap. 1: 49 (1860).

(b) var. pillansii Marais, var. nov., a typo petalis et filamentis lateralibus exappen-

diculatis, sepalis interioribus basi non saccatis, foliis saepe lobatis, indumento plerumque

densiore et longiore differt.

Type: Cape: Piketberg District, between Avontuur and Zebra Kop, Pillans 7603

(BOL, holo. ; GRA, K).

Cape. — Piketberg: between Avontuur and Zebra Kop, Pillans 7603 (BOL, GRA, K); PHlans 7506

(BOL); Kapteinskloof, Pillans 8017. Clanwilliam: Edwards in BOL 14450; 14451 (BOL); Elands-

kloof, Lewis in BOL 27109 (BOL); Pakhuis Pass, Compton 6950 (BOL, NBG); Lewis in BOL 27125

(BOL); Middelburg, Esterhuysen 1119 (BOL, K); Compton 12693 (NBG); Esterhuysen 2468; Sneeu-

berg, Esterhuysen 13099 (BOL).

Heliophila latisiliqua E. Mey. ex Sond. in Abh. Ges. Naturw. Hamb. 1 : 224

(1846). Type: Drege 585 (B!).

(a) var. latisiliqua.

El. latisiliqua E. Mey. ex Sond., l.c. (1846); FI. Cap. 1: 43 (1860).

Several-stemmed herbs with erect or procumbent stems up to 35 cm high. Leaves

1-5 cm long, the lobes 0-2-2 cm long. The flowers are bigger than in var. macrostylis

but, though the filaments are shorter, the anthers are longer. Distributed from the

Hex River Mts. to Calvinia.

(b) var. macrostylis (E. Mey. ex Sond.) Marais stat. nov.

Heliophila macrostylis E. Mey. ex Sond. in Abh. Ges. Naturw. Hamb. 1 : 225 (1846);

FI. Cap. 1: 43 (1860). Type: Drege s.n. (K!).

Stout erect herbs up to 90 cm high, usually with a single leafy stem. Leaves up

to 10 cm long, the lobes 2-5-4 cm long. The flowers are small, but the filaments are

proportionately much longer than those of var. latisiliqua. Confined to Namaqualand.

99

Heliophila linearis ( Thunb .) DC., Syst. 2: 697 (1821). Type: Thunberg Herb.

No. 15167 (UPS!).

(a) var. linearis.

H. linearis (Thunb.) DC., !.c. (1821); Sond. in Abh. Ges. Naturw. 1 : 238 (1846);

FI. Cap. 1: 47 (1860). H. linearifolia var. hirsuta Burch, ex DC., l.c. 692 (1821).

Type: Burchell 5518 (K.!). H. falcata Eckl. & Zeyh., Enura. No. 77 (1834-5). Type:

Ecklon & Zeyher 77 (K!). H. linearifolia var. pilosiuscula Sond. in Abh. Ges. Naturw.

Hamb. 1: 253 (1846); FI. Cap. 1: 51 (1860), partly, as to Burchell 5518. H. tenuis

N. E. Br. in Kew Bull. 1894: 99 (1894). Type: Pappe s.n. (K!).

Cheiranthus linearis Thunb., Prodr. 108 (1800). Type: Thunberg Herb. No. 15167.

(b) var. linearifolia {Burch, ex DC.) Marais, comb. nov.

H. linearifolia Burch, ex DC., Syst. 2: 692 (1821), excluding var. hirsuta-, Sond.

in Abh. Ges. Naturw. Hamb. 1: 252 (1846); FI. Cap. 1: 51 (1860), partly, excluding

var. pilosiuscula, and var. filifolia as to its basionym. H. linearifolia var. dolichocarpa

Schulz in Bot. Arch. 31 : 540 (1931). Type: Rogers s.n. (K!). H. elongcita Adamson

in FI. Cape Penins. 417 (1950), partly, excluding basionym of var. filifolia, non

(Thunb.) DC.

(c) var. reticulata {Eckl. & Zeyh.) Marais, comb. nov.

H. reticulata Eckl. & Zeyh., Enum. No. 90 (1834-5); Sond. in Abh. Ges. Naturw.

Hamb. 1 : 243 (1846); FI. Cap. 1 : 48 (1860). Type: Ecklon & Zeyher No. 90 (SAM!).

El. christieana Schulz in Bot. Arch. 31: 530 (1931). Type: Christie 49 (B!). H.

cornigerci Fourcade in Trans. Roy. Soc. S. Afr. 21 : 90 (1932). Type: Eourcade 1815a

(BOL!).

Heliophila meyeri Sond. in Abh. Ges. Naturw. Hamb. 1: 226 (1846). Type:

Drege s.n. (K!).

{a) var. meyeri.

H. meyeri Sond., l.c. (1846); FI. Cap. 1: 44 (1860). H. dentifera Sond. in Abh.

Ges. Naturw. Hamb. 1: 219 (1846); FI. Cap. 1: 42 (1860); Adamson in FI. Cape

Penins. 415 (1950). Type: Ecklon & Zeyher s.n. (SAM!).

{b) var. minor Marais, var. nov., a typo floribus et fructibus in partibus omnibus

minoribus, filamentis omnibus basi dilatatis papillatis sine appendice papillata differt.

Type: Cape: Swellendam District, Bonnievale, Leighton in BOL 21146 (BOL,

holo.).

Cape. — Swellendam: Bonnievale, Leighton in BOL 21146 (BOL). Worcester: Audensberg, Compton

9762 (NBG); Esterhuysen 3428 (K); Louwshoek Mts., Stokoe in SAM 59125 (SAM).

These plants differ from var. meyeri mainly in size. Since the appearance of the

plants is reminscent of some of the other species with smaller white flowers like H.

pendula and H. diffusa, rather than of var. meyeri, it is desirable to give it a name.

Heliophila pusilla L.f., Suppl. 297 (1781). Type: Thunberg Herb. Linn. 840.4.

{a) var. pusilla.

El. pusilla L.f., l.c. (1871); Sond. in Abh. Ges. Naturw. Hamb. 1: 210 (1846); FI.

Cap. 1: 40 (1860); Adamson in FI. Cape Penins. 414 (1950). H. divaricata Banks

ex DC., Syst. 2: 687 (1821); Sond. in Abh. Ges. Naturw. Hamb. 1 : 236 (1846); FI.

Cap. 1: 46 (1860). Type: Masson s.n. (BM!).

100

( b ) var. macrosperma Marais , var. nov., a typo floribus majoribus, filamentis

lateralibus papillatis, ovarii stylo longiore, ovulis paucioribus, siliquis latioribus et

seminibus majoribus differt.

Type: Cape: Stellenbosch District, Prior s.n. (K., holo.).

Cape. — Stellenbosch: Prior s.n. (K); Golf Course, Strey 665; Stellenbosch flats, Garside 158;

410 (K); Vanderstel, Smith 3207; Faure, Parker 3973 (BOL, K); Firgrove, Compton 13453 (BOL,

NBG); Elsenburg, Marioth 9281, partly ; 9281b; Simondium, Morrison in STE 1 1 109 (STE). Tulbagh:

De Hoek, Saron, Barker 5838 (BOL, NBG). Wellington: Compton 11624 (NBG). Without exact

locality: between Paarl and Pont, Drege 7555.

(c) var. setacea ( Schltr .) Marais, comb. nov.

H. setacea Schltr. in Bot. Jahrb. 49: 417 (1913). Type: Schlechter 4992 (B!).

(d) var. lanceolata ( Adamson ) Marais, comb. nov.

H. lanceolata Adamson in J. S. Afr. Bot. 7: 197 (1941). Type: Adamson 2973

(BOL!).

Heliophila alpina Marais, sp. nov. distincta.

Herba perennis parva, 10-15 cm alta, caulibus pluribus procumbentibus, partibus

junioribus puberulis. Folia 5-9 mm longa, pinnatim 5-7 lobata, stipulis minutis.

Inflorescentia pseudoterminalis, pauciflora; pedicelli breves, ± interpetiolares, bracteis

2 minutis subtendentes, saepe rami floriferi laterales pedicellos superantes. Sepala

4-5-5 mm longa, oblonga, puberula; exteriora 2 apice cucullata, breviter corniculata,

interiora 2 basi saccata. Pet ala 6-7 mm longa, 2-5-3 mm lata, anguste obovata.

FUamenta 2-4-3 mm et 3-4 mm longa. Antherae 1-1 -2 mm longae. Ovarium anguste

oblongo-ellipticum, replo minute puberulo; stylus longus; stigma breviter bilobatum;

ovula 8-12. SUiquae 1-1-5 cm longae, 3-4 mm latae, oblongae vel lineari-oblongae,

brevissime stipitatae; valvae 1-nervae; stylus 2-3 mm longus, robustus, apicem versus

attenuatus. Semina 2 -3-2 -8 mm longa, 1-5-1 -7 mm lata, ambitu oblongo-ovata,

papillosa.

Type: Basutoland: Thaba Ntlenyana, Guillarmod 2358 (PRE, holo.).

Procumbent, several-stemmed perennial herbs. Stems 10-15 cm long, the young

parts puberulous, glabrescent. Leaves 5-9 mm long, pinnately 5-7-lobed with linear,

entire lobes, probably = L fleshy, minutely puberulous, glabrescent, minutely stipulate.

Inflorescence pseudo-terminal, few-flowered, the pedicels ± interpetiolar, subtended by

2 minute bracts, often overtopped by lateral flowering branches. Sepals 4-5-5 mm

long, ovate-oblong to oblong-elliptic, puberulous; the outer 2 cucullate, shortly horn-

tipped; the inner 2 saccate and with broad membranous margins. Petals 6-7 mm

long, 2-5-3 mm broad, narrowly obovate, cuneate, very minutely puberulous at the

base outside. Filaments 2-4-3 mm and 3-4 mm long, the 2 short ones slightly thickened

at the base and with a glandular area. Anthers 1-1-2 mm long Ovary narrowly oblong-

elliptic, subsessile, finely puberulous on the replunr; style long; stigma shallowly

2-lobed; ovules 8-12. Fruits 1—1-5 cm long, 3-4 mm broad, oblong or linear-oblong,

very shortly stipitate; valves 1 -nerved, net-veined, bulged by the seeds; stipe and

replum puberulous; style 2-3 mm long, stout, tapering. Seeds 2 -3-2 -8 X 1-5-1 -7

mm, oblong-oval, subcompressed; seed-coat minutely papillose.

A very distinctive plant found on screes and steep grassy slopes at altitudes between

9.800 and 11,000 ft. Collected in fruit with a few late flowers in January; in fruit

only in March.

Cape. — Barkly East: Ben McDhui, Galpin 6573.

Basutoland. — Thabana Ntlenyana, Guillarmod 2358.

101

Heliophila cedarbergensis Marais, sp. nov., a H. scoparia Burch, habitu patenti

vel patulo, inflorescentiis intercalaribus, fructibus brevioribus seminibus paucioribus

differt.

Fruticulus patens, dense virgatus, glaber. Folia 0-5-3 cm longa, 1-2-5 mm lata,

oblanceolata vel anguste elliptico-lanceolata, levia vel cartilagineo-papillata, stipulis

minutis. Injiorescentia intercalaris. Sepala 4-5-5 3 mm longa, elliptica vel elliptico-

obovata; exteriora 2 apice cucullata, breviter corniculata, interiora 2 basi saccata.

Petala 10-11-5 mm longa, 4-5-5 mm lata, obovato-spathulata, longe ungulata. Fila-

menta exappendiculata. Antherae 1-5-1 -8 mm longae. Ovarium lanceolatum, stipi-

tatum, ovulis 10-12. Siliquae 2-4 cm longae, 2- 5-3 -5 mm latae, lanceolatae, breviter

stipitatae; valvae 1-nervatae; stylus 2-5-5 mm longus, subulatus. Semina 2-5 mm

longa, 1-6-1 -8 mm lata, ovalia, compressa, papillata.

Type: Cape: Clanwilliam District, Cedarberg, Wolfberg, Esterhuysen 20566

(BOL, holo.; K, PRE).

Trailing or densely twiggy, woody shrublets; branches glabrous with red-brown

bark, smooth or finely cartilaginous-papillate. Leaves 0-5-3 cm long, 1-2-5 mm

broad, oblanceolate to narrowly elliptic-lanceolate, acute or acuminate, mucronate,

smooth or finely cartilaginous-papillate, minutely stipulate. Inflorescence few- to

several-flowered, terminal or apparently so but soon over-topped by new leafy growth

of the central axis, i.e., intercalary. Pedicels 0-9-1 -6 cm long, erect or ascending,

subtended by 2 minute filiform bracts. Sepals 4 -5-5 -3 mm long, 1 -8-2-3 mm broad,

elliptic to elliptic-obovate; outer 2 cucullate and shortly horn-tipped, with mem-

branous margins, inner 2 membranous, saccate. Petals 10-11-5 mm long, 4-5-5 mm

broad, obovate-spathulate, rounded, long-clawed. Filaments 3-5 mm and 4 -5-5 -5

mm long. Anthers 1-5-1 -8 mm long. Ovary lanceolate, stipitate; style long, stigma

capitate; ovules 10-12. Fruits 2-4 cm long, 2 -5-3 -5 mm broad, lanceolate, shortly

stipitate; valves flat, 1 -nerved, slightly bulged by the seeds; style 2-5-5 mm long,

stoutly subulate. Seeds 2-5 mm long, 1 -6-1 -8 mm broad, oval, compressed, papillate.

Growing among rocks and boulders or on shady ledges at an altitude of 4,000-

5,500 ft. Collected in flower in October; flowers described as white or pink.

Cape. — Clanwilliam: Cedarberg, Wolfberg, Esterhuysen 20566; Esterhuysen 22448 (BOL, K);

slopes below Wolfberg Cracks, Esterhuysen 29974 (BOL); Tafelberg, Esterhuysen 30000 (BOL).

Heliophila cuneata Marais, sp. nov., a H. integrifolia L. planta suffruticosa perenni,

floribus multo majoribus, fructibus latioribus et seminibus majoribus; a H. lineari

(Thunb.) DC. forma foliorum, fructibus multo latioribus stylis longioribus, et ovulis

paucioribus differt.

Frutex diffusus usqae 1-3 m altus, ramis junioribus sparse pilosis. Folia 3-6-5

mm longa, 1 • 5-3 cm lata, cuneata vel obovato-cuneata, in parte superiore 5-9 dentata,

superne sparse vel dense pilosa vel hirsuta, subtus minus dense pilosa vel glabra.

Racemus terminalis, multiflorus, initio contractus, denique elongatus et laxus. Sepala

7-9-5 mm longa, lineari-oblonga, dense pilosa; exteriora 2 apice cucullata; interiora

2 basi saccata, marginibus membranaceis. Petala 10-16-5 mm longa, 6-5-11-5 mm

lata, obovata, rotundata, breviter unguiculata, appendicibus papillosis. Filamenta

4-5-5 mm, 5-5-7 mm longa, breviora 2 appendicibus magnis instructa. Antherae

3-3-5 mm longae. Ovarium oblongum, stylo longo gracili, ovulis 6-8. Siliquae 6-8

cm longae, 7-8 mm latae, lineari-oblongae; valvae 3-nervae; stylus 1 • 1-1 -4 cm longus,

robuste subulatus. Semina 6 mm longa, 5 mm lata, subrotundata, compressa, alata.

Type: Cape: Stellenbosch District, Swartboskloof, Taylor 5839 (PRE, holo.).

Straggling shrub up to 1-3 m high with long weak branches; young branches

sparsely pilose. Leaves 3-6-5 cm long, 1-5-3 cm broad, cuneate or obovate-cuneate,

gradually narrowed below, 5-9 toothed or lobulate-dentate in the upper half; sparsely

102

to densely pilose or hirsute on the upper surface, less densely hairy to glabrous

underneath. Racemes terminal, many-flowered, contracted in flower; elongate and

very lax in fruit; rhachis glabrous. Sepals 7-9-5 mm long, linear-oblong, densely

pilose; outer 2 cucullate; inner 2 saccate with broad membranous margins. Petals

10-16-5 mm long, 6-5-11-5 mm broad, obovate, rounded, shortly clawed, each with

a papillate appendage. Filaments 4-5 • 5 mm and 5 • 5-7 mm long, the short ones each

with a large appendage. Anthers 3-3-5 mm long. Ovary oblong with a long, slender

style, as long as or longer than the ovary; ovules 6-8. Fruits 6-8 cm long (including

the style), 7-8 mm broad, linear-oblong with straight margins; valves flat, strongly

3-nerved (sometimes with 2 or 4 additional weaker veins); style 1-1-1 -4 cm long,

stoutly subulate. Seeds 6 mm long, 5 mm broad, subcircular, winged.

The flowers are blue with a white centre, produced in spring. Known only from

the one locality where it was first recorded in 1960, growing among boulders in scrub

vegetation.

Cape. — Stellenbosch: Jonkershoek, Swartboskloof, Taylor 5839; Marais 1451 (K); Van der Merwe

22-58; Van der Merwe s.n.

Heliophila esterhuyseniae Marais, sp. nov., a H. dregeana Sond. floribus majoribus,

fructibus non moniliformibus differt.

Fruticulus lignosus confertus, erectus vel patens, glaber, 0-6-1 m altus. Caules

glabri, angulati vel anguste alati. Folia 1-4 cm longa, 1 • 5-4 mm lata, lineari-lanceolata

vel elliptico-lanceolata, sessilia, levia vel marginibus costaque subtiliter cartiligineo-

papillatis, stipulis minutis. Inflorescentia intercalaris, floribus pluribus, pedicellis bracteis

minutis 2 subtendentibus. Sepala 5 • 3-6 mm longa, ovata vel obovata, membranacea,

3-nervata; interiora 2 basi saccata. Petata 11-14 mm longa, 6- 5-7 -5 mm lata,

rotundato-obovata, unguiculata. Filamenta exappendiculata. Antherae 1-5-2 mm

longae. Ovarium lanceolatum, ovulis 4-8. Siliquae 0-8-2 -8 cm longae, 2-5-4 mm

latae, lanceolatae, breviter stipitatae, erectae; valvae 1-nervatae, denique asperae et

naviculares; stylus 1 - 5—5 mm longus, robuste subulatus. Semina 3-5-4 mm longa,

3-3-2 mm lata, citriformia, papillosa.

Type: Cape: Paarl District, Wemmershoek Pk., Esterhuysen 11290 (BOL, holo.).

Dense, erect or trailing, woody shrublets 0-6-1 m high. Stems angled or narrowly

winged by the decurrent leaf-margins. Leaves 1-4 cm long, 1-5-4 mm broad, linear-

lanceolate to elliptic-lanceolate, sessile, smooth or the margins and the midrib, which

is prominent on the lower surface, finely cartilaginous-papillate; subtended by 2 minute

stipules. Inflorescence intercalary, few- or several-flowered, each pedicel subtended by

2 minute bracts; the old pedicels from previous seasons’ flowers persistent. Sepals

5-3-6 mm long, 2-3-3 mm broad, membranous, 3-nerved; outer 2 obovate, cucullate,

shortly horn-tipped ; inner 2 ovate, strongly saccate. Petals 1 1-14 mm long, 6 • 5-7 • 5 mm

broad, subcircular-obovate, clawed. Filaments 4-4-3 mm and 4 -5-4 -8 mm long.

Anthers 1-5-2 mm long. Ovary lanceolate, shortly stipitate; style long, slender;

ovules 4-8. Fruits 0-8-2 -8 cm long, 2-5-4 mm broad, lanceolate, shortly stipitate,

erect; valves 1 -nerved, at first smooth, later rough, i ridged and warted, bulged

by the seeds, eventually boat-shaped; style 1-5-5 mm long, stoutly subulate. Seeds

3-5-4 mm long, 3-3-2 mm broad, citriform, papillose.

Growing on rock ledges in partial shade, old screes, at foot of cliff's or on stream-

banks. The flowers are pale blue, mauve or white, produced in spring.

Cape. — Ceres: Witelskloof, Esterhuysen 26396 (BOL); Skurfdeberg, Esterhuysen 14724 (BOL).

Worcester: Slanghoek Mts., Esterhuysen 22306 (BOL, K); Stettynskloof, Esterhuysen 17629 (BOL);

Hex River Mts., Morainekloof, Esterhuysen 28073 (BOL). Paarl: Wemmershoek Pk., Esterhuysen

11290 (BOL). Caledon: Fransch Hoek Mts., Roesbos Pk., Esterhuysen 29422 (BOL). Paarl-

Worcester: Du Toitskloof, Esterhuysen 30359 (BOL).

103

Heliophila eximia Marais, sp. nov. distincta.

Herba perennis, scandens?, glabra. Folia 4-5-6- 5 cm longa, 1-2 cm lata, elongato-

cuneata prope apicem 3-7 dentata, in petiolum attenuata. Racemus terminalis,

contractus, pauci- ad multi-florus. Sepala 3-3 • 8 mm longa, oblonga; interiora 2 leviter

saccata. Petala 6 mm longa, 4 • 5-4 • 8 mm lata, rotundato-obovata, breviter unguiculata.

Filamenta subulata. Antherae 1 -8-2 mm longae. Ovarium anguste oblongum, breviter

stipitatum; stylus brevis; stigma capitatum, magnum; ovula 8-12. Fructus ignotus.

Type: Cape: Namakwaland district, Hellskloof, Hall 789 (NBG, holo.).

Perennial scandent herbs?, glabrous. Leaves 4 -5-6 -5 cm long, 1-2 cm broad,

elongate-cuneate, 3-7-toothed near the apex, narrowed into a petiole. Racemes terminal,

few- to many-flowered, contracted. Sepals 3-3-8 mm long, 1-5 nun broad, oblong;

outer 2 cucullate, shortly horn-tipped; inner 2 slightly saccate with broad membranous

margins. Petals 6 mm long, 4 -5-4 -8 mm broad, obovate-circular, shortly clawed.

Filaments 2-2-4 mm and 2- 5-2- 8 mm long, subulate. Anthers 1 • 8-2 mm long. Ovary

narrowly oblong, shortly stipitate; style short; stigma large, capitate. Fruits not

known.

Known from only one gathering mentioned above which was collected on 19th.

September, 1953, in flower; the flowers are white. The affunity of this plant is obscure,

all the more so in the absence of ripe fruits.

Heliophila filicaulis Marais, sp. nov., a H. tulbaghensi Schinz floribus majoribus,

fructibus compressis differt.

Fruticulus multo ramosus tenuis; rami glabri, striis minutis cartilagineis, plerumque

valde moniliformibus. Folia 1-3 cm longa, lineari-filiformia, acuta, mucronata, stipu-

lata. Racemus terminalis, pauci ad pluri-florus. Sepala 5-5 • 5 mm longa, anguste

oblonga, membranacea; interiora 2 basi saccata. Petala 10-11-5 mm longa, 3 -8-4 -3

mm lata, oblongo-obovata, unguiculata. Filamenta exappendiculata. Antherae 1-7-

1-9 mm longae. Ovarium lanceolatum, stipitatum; stylus longus; ovula 4-8. Siliquae

1-5-3 cm longae, 1 -6-1-8 mm latae, lineares, compressae, moniliform.es, stipite 0-5-2

mm longo; stylus 2-5-3 mm longus, linearis, robustus. Semina 2 -2-2 -3 mm longa,

1-3-1 -4 mm lata ovata, compressa, papillosa.

Type: Cape: Worcester District, Hex River Mts., Esterhuvsen 15936 (BOL,

holo.; K. NBG. PRE).

Much-branched slender shrublets with wiry branches; branches glabrous with

minute cartilaginous lines which are mostly strongly beaded, these disappearing with

age. Leaves 1-3 cm long, linear-filiform, acute, mucronate, subtended by 2 minute,

filiform stipules. Racemes terminal, few- to several-flowered. Pedicels 0-6-1 cm

long, erect or ascending, usually subtended by 2 minute bracts. Sepals 5-5 • 5 mm long,

1-4-2 mm broad, narrowly oblong, membranous; outer 2 cucullate; inner 2 saccate.

Petals 10-11-5 mm long, 3 -8-4 -3 mm broad, oblong-obovate, narrowly clawed.

Filaments 4-5 mm and 5 • 5-6- 5 mm long. Anthers 1 • 7-1 • 9 mm long. Ovary stipitate,

lanceolate; style long; ovules 4-8. Fruits 1-5-3 cm long, 1-6-1 -8 mm broad, linear,

compressed, moniliform with narrow waists between the seeds; stipe 0-5-2 mm long;

style 2-5-3 mm long, linear, stout. Seeds 2 -2-2 -3 mm long, 1-3-1 -4 mm broad,

ovate, flat, papillose.

Growing on southern, south-eastern and eastern aspects in shelter of rocks.

Collected in flower in October; flowers blue to pinky-blue.

Cape. — Worcester: Hex River Mts., Esterhuvsen 15936; Outer Sanddrift Peak, Esterhuysen 29777

(BOL).

7230691-4

104

Heliophila katbergensis Marais, sp. nov., a H. rigidiuscula Sond. habitu, petalis

exappendiculatis et seminibus paucioribus differt.

Herba perennis. Caules plures, procumbentes, striis minutis cuticularibus. Folia

1-1 • 8 cm longa, 1 • 8-2 • 3 mm lata, oblanceolata vel oblanceolato-elliptica, integra,

acuta, stipulis 2 minutis. Racemus terminalis, laxus. Sepala 3 • 5-4 mm longa, oblonga;

interiora 2 basi saccata. Petala 6 • 5-7 • 5 mm longa, 3 • 5-4 • 5 mm lata, late obovata. Fila-

menta subulata, breviora 2 appendiculata. Antherae 1-8-2 mm longae. Ovarium

ellipticum, stipitatum, ovulis 4-6. Fructus immaturus 1-6-1 -7 cm longus, 4-5-5 mm

latus, oblongus, pedicellis recurvis 1 -5-2 cm longis; valvae planae, 1-nervae, reticulatae;

stylus 3 -5-6 -5 mm longus, robustus, c. 1 mm crassus.

Type: Cape: Stockenstroom District, Katberg, Hutchinson 1614 (PRE, holo.;

BOL, K).

Procumbent, many-stemmed, glabrous herbs. Stems with minute cuticular ridges.

Leaves 1-1-8 cm long, 1-8-2 -3 mm broad, oblanceolate or oblanceolate-elliptic, entire,

acute, mucronate, subtended by 2 minute stipules. Racemes terminal, lax, many-

flowered. Sepals 3-5-4 mm long, oblong, membranous-margined; outer 2 thickened

at the apex; inner 2 saccate. Petals 6 -5-7 -5 mm long, 3 -5-4 -5 mm broad, broadly

obovate. Filaments subulate, the short ones each with an appendage. Anthers 1 • 8-2

mm long. Ovary elliptic, stipitate; ovules 4-6. Immature fruit 1-6-1 -7 cm long,

4-5-5 mm broad, oblong, on recurved pedicels 1-5-2 cm long; valves flat, 1 -nerved,

reticulately veined; style 3 -5-6 -5 mm long, stout, c. 1 mm thick.

Known from only the one gathering from the top of Katberg; collected in flower

on 27th November, 1928. The flowers are described as deep mauve. Like so many

other of Hutchinson’s specimens, this one was misidentified and in “A Botanist in

Southern Africa ” he refers to it as Heliophila glauca Burch.

Heliophila obibensis Marais, sp. nov., a H. amplexicauli L.f. foliis angustis linearibus,

floribus multo minoribus, petalis et filamentis exappendiculatis; a H. pectinata Burch,

ex DC. foliis simplicibus, siliquis latioribus lineari-oblongis differt.

Herba annualis, erecta, parva, glabra. Folia linearia, integra. Racemus terminalis,

pluriflorus. Sepala 1-7-2 mm longa, anguste oblonga. Petala 2 -5-3 -5 mm longa,

1 • 2 mm lata, anguste oblongo-obovata. Filamenta 1 • 8-2 mm et 2-2 ■ 3 mm longa,

filiformia. Antherae 0 ■ 5-0 • 6 mm longa. Ovarium stylo brevi robusto, stigmate capitato

magno, ovulis 8-12. Siliquae 1-2-1 -7 cm longae, 2-2-5 mm latae, lineari-oblongae,

submoniliformes; valvae planae, 1-nervae, reticulatae; stylus c. 0-5 mm longus.

Semina ignota.

Type: South West Africa: Luderitz District, Obib, Merxmuller & Giess 3340

(M, holo.).

Erect, branched, glabrous, annual herbs up to 15 cm high. Leaves up to 2 • 5 cm

long, 0-7-0 -8 mm wide, linear, simple, each subtended by 2 small filiform stipules.

Pedicels each subtended by 2 small filiform bracts; in flower slender, erect, 2-4 mm

long; in fruit spreading or ± recurved, 5-6 mm long. Sepals 1 • 7-2 mm long, 0- 6-0- 8

mm broad, narrowly oblong; the outer 2 ± cucullate; the inner 2 slightly saccate.

Petals 2 -5-3 -5 mm long, 1-2 mm broad, narrowly oblong-obovate, clawed. Filaments

1-8-2 mm and 2-2-3 mm long, filiform. Anthers 0-5-0 -6 mm long. Ovary linear-

oblong; style short, stout; stigma large, capitate; ovules 8-12. Siliquae 1-2-1 -7 cm

long, 2-2 -5 mm broad, linear-oblong, submoniliform; valves flat, 1 -nerved, reticulately

veined; style c. 0-5 mm long. Seeds not seen.

This distinctive species is known from only the one gathering made on the 1st

of September, 1963, in quartzite screes on the slopes north of Obib. The small white

flowers turn pinkish with age.

105

Heliophila promontorii Marais, sp. nov., a H. macowaniana Schltr. planta fere

glabra, floribus majoribus caeruleis vel lilacinis siccitate roseis, siliquis sursum versus

curvatis stylis inconspicuis parvis differt.

Herba annualis, tenuis, sterea; caules basin versus fusci; nodi inferiores minute

puberuli, reliqui glabri. Folia integra vel bipinnatifida, linearia vel lineari-filiformia.

Sepala 2-8-4 mm longa, oblonga, interiora 2 basi saccata. Petala 5 -5-8 -5 mm longa,

2-5-6 mm lata, obovata, exappendiculata vel appendicibus minutis, caerulea vel lilacina,

siccitate rosea. Filamenta lateralia plerumque appendicibus minutis. Ovarium lineare,

ovulis 16-25. Siliquae 2-3 cm longae, 0-8-1 -2 mm latae, lineares, tenues moniliformes

sursum versus curvatae; stylus 1-1 -6 mm longus, parvus, subfusiformis vel subclavatus.

Semina 1 mm longa, 0-7 mm lata, ovalia anguste marginata.

Type: Cape Peninsula, Kenilworth, Bolus 7922 (PRE, holo.; BOL, K).

Slender wiry annual herbs 6-25 cm high. Stems dark in the lower part only or

all over; lower part of stem, sometimes only a few nodes, minutely puberulous, rest

of the plant glabrous. Leaves 1-5-5 -5 cm long, simple or 3- to 7-lobed, rarely

subbipinnately lobed, subtended by 2 minute stipules; lobes linear or linear-filiform.

Racemes terminal, lax, few-flowered. Pedicels 0-7-1 -2 cm long in flower, slender;

0- 9-1 -3 cm long in fruit, spreading; subtended by 2 minute bracts. Sepals 2-8-4 mm

long, 0-9-1 -5 mm broad, oblong; outer 2 cucullate, thickened at the tip; inner 2

saccate with broad membranous margins. Petals 5 -5-8 -5 mm long, 2-5-6 mm broad,

obovate to circular-obovate, clawed, without or with a very small appendage. Fila-

ments 1 -8-2-8 mm and 2-3-1 mm long, the short ones usually with a small appendage.

Anthers 1-1 ■ 5 mm long. Ovary linear; stigma capitate; ovules 16-25. Fruits 2-3 cm

long, 0-8-1 -2 mm broad, linear, shallowly moniliform, curving upwards; valves

1- nerved; style 1-1-6 mm long, subfusiform or subclavate. Seeds broadly oval,

narrowly margined.

This is Ft. trifida sensu Adamson in FI. Cape Peninsula and sensu Schulz in

Botanisches Archiv 31: 542 (1931), a species which has not yet been given a name.

True Ft. trifida Thunb., an illegitimate name, is a synonym of H. pinnata L.f. (see

Bothalia 8, 2: 167, 1965). Some of the specimens have been identified as H. tenuifolia

Sond. which is, however, a synonym of H. digitata L.f.

The flowers are described as blue, blue with a white eye, or mauve with a white

centre. They all dry pink. The flowering period is from September to November.

The species is confined to the Peninsula, usually found on sand dunes and flats.

Cape. — Peninsula: Kenilworth, Bolus 7922; Salter 5654 (BM, BOL, K); Bolus in PRE 28953;

Levyns 5200; Rondebosch, Bolus 3965 (BOL); Vlakkenburg, Compton 20177 (BOL, NBG); Smith’s

Farm, Salter 7044 (BOL); Smitswinkel, Salter 234/3 (BM); Rooihoogte, Leighton in BOL 27350

(BOL); Compton 13781 (NBG). Without exact locality; Zeyher 41 (BOL, K, SAM); Wright s.n.

(K); Grey s.n. (K).

Heliophila rimicola Marais, sp. nov., a Ft. esterhuyseniae Marais fructibus com-

pressis et seminibus compressis levibus differt.

Fruticulus perennis, ramis rigidis. Folia 0-8-4 cm longa, 1-2-8 mm lata, lineari-

oblonga vel oblanceolata, acuta, crassa, stipulis 2 minutis. Inflorescentia intercalaris

vel ramulis lateralibus abbreviatis terminalis, pauciflora, rare floribus axillaribus.

Pedicelli 1-2-3 cm longi, erecti, apice in fructu valde incrassato, bracteis 2 filiformibus

minutis subtendentibus. Sepala 5-7 mm longa, ovata vel obovato-oblonga, exteriora

2 cucullata. Petala 1-1-1 -5 cm longa, 5-7 mm lata, obovato-spathulata. Filamenta

exappendiculata. Antherae 1-8-2 -7 mm longae. Ovarium lanceolatum, stipitatum;

stylus tenuis; ovula 4-8. Siliquae 1-5-3 cm longae, 2-7-4 mm latae, lanceolatae,

longe acuminatae, compressae, stipite 0-5-2 mm longo; valvae 1-nervae; stylus 2-3-5

mm longus, tenuis, apicem versus attenuatus. Semina 2-1-3 mm longa, 1-8-2 -5 mm

lata, subrotunda vel oblongo-rotunda, compressa, levia.

106

Type: Cape: Ladismith District, Swartberg, Towerkop, Esterhuysen 26800 (BOL,

holo.).

Glabrous, perennial shrublets with stiff branches; young branches striate with

ridges from the thickened leaf-base; older branches smooth, round, with a light brown

bark. Leaves 0-8-4 cm long, 1-2-8 mm broad, linear-oblong, linear-oblanceolate or

oblanceolate, acute or shortly acuminate, fleshy, subtended by 2 minute stipules.

Inflorescence intercalary or terminal on abbreviated side-branches, rarely with some

axillary flowers. Pedicels 1-2-3 cm long, erect, the apex thickened and becoming

strongly swollen in fruit, subtended by 2 minute filiform stipules. Sepals 5-7 mm

long, 1 • 5-3 • 5 mm broad ; outer 2 ovate or obovate-oblong, cucullate, thickened at

the apex; inner 2 ovate to elliptic, membranous. Petals 1-1-1 -5 cm long, 5-7 mm.

broad, obovate-spathulate, clawed. Filaments 4-4-8 mm and 5 -3-6 -5 mm long.

Anthers 1 -8-2-7 mm long. Ovary lanceolate, stipitate; style slender; ovules 4-8.

Fruits 1-5-3 cm long, 2-7-4 mm broad, lanceolate, long acuminate, flattened; stipe,

0-5-2 mm long; valves 1 -nerved; style 2-3-5 mm long, slender, tapering. Seeds

2-1-3 mm long, 1-8-2 -5 mm broad, subcircular or oblong-subcircular, flattened,

smooth.

Apparently always found in moist crevices on rock faces at an altitude of 6-7,000

ft. The flowers are described as white, light purplish or tinged with purple. Flowering

specimens have been collected during December; specimens with ripe fruits during

March and April.

Cape. — Ladismith: Swartberg, Towerkop, Esterhuysen 18516 (BOL); Esterhuysen 26800 (BOL);

Esterhuysen 30672 (K). Prince Albert: Swartberg, Ridge Peak, Andreae 1223; Primos 55; Stokoe

1752; 1796.

Heliophila schulzii Marais, sp. nov., a H. lactea Schltr. foliis minute stipulatis,

pedicellis bracteis 2 similibus subtendentibus, sepalis puberulis differt.

Herba annualis, 15-45 cm alta. Folia 5-12 cm longa, integra vel 3-lobata, lobis

linearibus vel lineari-filiformibus, supeme minute puberula, stipulis minutis caducis.

Sepala 4-5-6 mm longa, oblonga, breviter puberula; interiora 2 basi saccata. Petala

0-8-1 -2 cm longa, 3 -5-7 -8 mm lata, obovata, appendiculata. Filamenta 3 -5-4 -5 mm

et 4-5 mm longa. brevioribus 2 appendiculatis. Antherae 1-5-2 mm longae. Ovula

(36-) 44-70 (-80). Siliquae 3 • 5-6 cm longae, 1-1 ■ 8 mm latae, lineares, margine recto,

denique pendulae; valvae 1-nervae, glabrae vel puberulae; stylus 1 -8-3-5 mm longus.

Type: Cape: Naniakwaland district, Bowesdorp, Stokoe in SAM 59126 (SAM,

holo.).

Slender or stout annual herbs 15-45 cm high, simple or with many ascending

branches; basal part of stem and sometimes the axils of the branches shortly puberulous.

Leaves 5-12 cm long, entire or 3-lobed, the lobes linear or linear-filiform, the upper

leaves entire, shorter; upper surface minutely puberulous; stipules minute, caducous.

Racemes lax. Pedicels shortly puberulous at the base, with 2 minute bracts; in flower

slender, erect, in fruit 1-1-5 cm long, widely spreading but finally sharply reflexed.

Sepals 4-5-6 mm long, 1-2-1 -9 mm broad, narrowly oblong or oblong-elliptic, with

membranous margins, shortly puberulous; outer 2 cucullate, thickened or very shortly

horn-tipped; inner 2 saccate. Petals 0-8-1 -2 cm long, 3 -5-7 -8 mm broad, obovate,

rounded, cuneate or clawed, each with an appendage. Filaments 3 -5-4 -5 mm and

4-5 mm long, the short ones appendaged. Anthers 1-5-2 mm long. Ovary linear;

stigma large, capitate; ovules (36-) 44-70 (-80). Fruits 3-5-6 cm long, 1-1-8 mm

broad, linear with straight margins; valves 1-nerved, strongly bulged by the seeds,

glabrous or puberulous all over or just near the base; style 1 -8-3-5 mm long, straight

or very slightly swollen near its base, or subclavate. Seeds 0-9-1 -4 mm long, 0-7-1

mm broad, subcircular to oblong, emarginate.

107

The flowering time is from August to October and the flowers are blue or purple.

Apparently confined to the Kamiesberg.

Cape. — Namakwaland: Brakdam, Schlechter 11147; Maguire 958 (NBG); Kamiesberg, De Kom,

Leipoldt 3914; Leipolclt 3915 (BOL); between Kamieskroon and Leliefontein, Salter 1506 IK); Kamies-

kroon, Herre in STE 11786 (STE); Macdonald 62 (BM); Bowesdorp, Stokoe in SAM 59126 (SAM);

Garies, Compton 11565 (NBG); Henrici 2145; Spektakel, Morris in BOL 5580 (BOL); between

Swartdoring River and Groen River, Drege (BM); Grootvlei, Compton 17285 (BOL, NBG).

LEPIDIUM

Lepidium africanum (Burm.f) DC., Syst. 2: 552 (1821), quoad synon. tandem,

excl. specim. et descr. ; non auct.

Thlaspi africanus Burm.f., Prodr. FI. Cap. 17 (1768). Type: Burmann in Herb.

Delessert (Gl).

Lepidium divaricatum sensu DC., l.c., 543 (1821). quoad specim. Burm.; sensu auct.

pro parte.

In Burmann’s herbarium there are three specimens with the name Thlaspi africanus.

Burmann’s diagnosis: “ Siliculis subovatis, foliis lanceolatis serrulatis glaberrimus ”,

agrees very well with one of them. This specimen, however, is the one which De

Candolle excluded from Lepidium africanum when he made the combination. On

p. 543 under L. divaricatum he writes: “ Haec species est in herb. Burmanni confusa

cum Lepidio suo africano, ad quod revera accedit ”.

Thus, the name L. africanum must now be applied to plants which, since De

Candolle’s treatment, have been included in L. divaricatum Ait. The plants which

I include in L. africanum have mostly been put into L. divaricatum subsp. linoides.

In my opinion the type of L. linoides Thunb. does not belong here, but should go into

L. divaricatum sens, strict. The plants that we used to know as L. africanum must

now be called L. capense Thunb.

Lepidium capense Thunb., Prodr. 107 (1800). Type: Thunberg Herb. No. 14761

(UPS!).

L. decumbens Desv. in Journ. Bot. Appl. 3: 165, 176 (1814). Type: Herb. Desvaux

(ex cult.) (P, holo.!; BM!). L. africanum (Burm. f.) DC., Syst. 2: 552 (1821), pro

parte, quoad specim. et descr., excl. synon. Burm.f. — var. typicum Thell. in Viertelj.

Naturf. Ges. Ziir. 51: 187 (1906), excl. synon. Burm.f. — var. capense (Thunb.) Thell.,

l.c. — var. serratum Thell., l.c., 188 (1906). Type: Thunberg Herb. No. 14792

(UPS!). — var. serratum f. glabratum Thell., l.c., 189 (1906). Type: Ecklon & Zeyher

No. 43, p.p., not seen. — var. burchellii Thell., l.c. (1906). Type: Burchell 137 (K!).

L. africanum auct.

My concept of this species does not differ from that of Thellung except that I

do not uphold all his varieties. Only the name, and thus the synonymy, has changed.

Lepidium divaricatum Ait., Hort. Kew. ed. 1, 2: 375 (1789).

(a) subsp. divaricatum.

L. divaricatum Ait., l.c. Type: “Lepidium divaricatum Banks’ ex Hort. Kew”

in Herb. Delessert (G ! ). — subsp. eu-divaricatum Thell. in Viertelj. Naturf. Ges. Ziir.

51: 167(1906). Type as above. L. linoides Thunb., Prodr. 107 (1800). Type: Thunberg

Herb. No. 14776 (UPS!). L. divaricatum subsp. linoides (Thunb.) Thell., l.c. (1906).

— subsp. linoides var. typicum Thell., l.c., 168 (1906). Type as above. L. subdentatum

108

Burch, ex DC., Syst. 2: 545 (1821). Type: Burchell 1299 (K!). L. linoides var. subden-

tatum (DC.) Sond. in FI. Cap. 1: 28 (1860). L. divaricatum subsp. linoides var.

subdentatum (DC.) Thell., l.c. (1906). L. pinnatum sensu Sond. l.c., 30 (1860), pro

parte, quoad syn. L. divaricatum.

The type specimen is a cultivated plant, preserved in Herb. Delessert in Geneva,

with large, broad, pinnately lobed leaves like those of Bolus 25 from Graafif-Reinet.

They do not represent the usual form of the species; the majority are fairly slender

plants with small, narrower leaves.

(. b ) subsp. trifurcum (Sond.) Marais , comb. nov.

L. trifurcum Sond. in Linnaea 23: 4 (1850); FI. Cap. 1: 30 (1860). Type: Zeyher

213 (K!, PRE!).

Thellung, in Vierteljahreschrift der Naturforschenden Gesellschaft, Zurich 51 : 171

(1906), interpreted L. trifurcum incorrectly, probably because Sonder had, in Flora

Capensis, misidentified Drege 7542a with his species. Rehmann 3033, 3034 and 3219

and Drege 7542a from Vienna, cited by Thellung, all belong to L. schinzii Thell. 1

have not seen the sheet of Drege 7542a from Herb. Delessert which is also cited by

Thellung.

Lepidium ecklonii Schrad., Ind. Sem. Hort. Goetting. 3 (1830). Type: ex Hort.

Goettingen (LE!).

L. divaricatum subsp. ecklonii (Schrad.) Thell. in Viertelj. Naturf. Ges. Ziir. 51:

169 (1906). L. sylvaticum Eckl. & Zeyh., Enum. 6, No. 37 (1834-5). Type: Ecklon

& Zeyher 37 (LE!). L. pinnatum Eckl. & Zeyh., l.c. 7, No. 45 (1834-5), non Thunb.

L. hirtellum Sond. in FI. Cap. 1: 30 (1860). Type: Ecklon & Zeyher 45 (LE!). L.

capense sensu Sond., l.c. 29 (1860), pro parte, quoad syn. L. ecklonii Schrad.

Lepidium basuticum Marais , sp. nov., a L. desertorum Eckl. & Zeyh. siliculis magnis

ellipticis stylis in emarginaturis inclusis differt.

Herba perennis. Caules pilis ± falcatis dense induti. Folia basilaria rosulata,

petiolata, 8-10-jugata, pinnata, pinnis lobatis vel dentatis. Folia caulina lobis pauci-

oribus angustioribus et minus lobulatis; folia superiora minora, oblanceolata, 3-fida

vel integra. Racemi fructiferi 1-5 cm longi, densi. Sepala 0-8-0 -9 mm longa, oblonga.

Petala 0. Stamina 2. Nectaria 0-4-0 -5 mm longa, filiformia. Siliculae 2-8-3 mm

longae, 1 -7-1 -9 mm latae, ellipticae, leviter emarginatae, stylo in emarginatura incluso.

Semina 1 — 1-4 mm longa, 0-5-0-6 mm lata.

Type: Basutoland: Butha Buthe, Pela Tseou Streams, Guillarmod 2073 (Hb.

Guillarmod, holo.).

Perennial herb; stems prostrate or erect up to 30 cm high; with a long taproot.

Stems densely ± retrorsely set with flattened ± sickle-shaped hairs. Basal leaves in

a rosette, petiolate, up to 12 cm long, pinnate, 8-10-jugate, the pinnae variously lobed

or dentate; both surfaces of petiole and lower surface of the rhachis and midribs set

with short, flattened hairs. Cauline leaves pinnately lobed with fewer, narrower and

less lobulate lobes, upwards smaller, the uppermost oblanceolate, 3-fid to entire.

Racemes terminal, in fruit 1-5 cm long, dense; rhachis puberulous. Pedicels 1-5-3 -5

cm long, wide-spreading, puberulous all round. Sepals 0-8-0 -9 mm long, oblong

with membranous margins, subpersistent. Petals 0. Stamens 2; filaments subulate,

thickened at the base. Nectaries 0-4-0 -5 mm long, filiform. Siliculae 2-8-3 mm long,

1-7-1 -9 mm broad, elliptical, shallowly emarginate with the style shorter than or as

long as the sinus. Seeds 1—1-4 mm long, 0-5-0-6 mm broad, brown.

109

A distinctive plant. The leaves are reminiscent of those of L. desertorum, and

so are the short fruiting racemes. The long filiform nectaries are found in both species.

Differs from L. desertorum in the large elliptical siliculae.

A high mountain species found between 9,200 and 10,300 feet.

Basutoland. — Butha Buthe, Pela T eou St earns, Guillarmod 2073; Lehaha-la Sekhonyana, Guil-

larmod 348; Maluti Mts., Dieterlen 1260.

Cape. — Barkly East: Doodmanskrans, Galpin 6574, p.p.

Lepidium suluense Marais, sp. nov. distincta.

Herba annua vel perennis, 30-60 cm alta. Caules pilis d= clavatis brevibus puberuli.

Folia caulina inferiora sub-bipinnatipartita, lobis patulis vel plusminusve recurvis

angustis, acutis, lineari-lanceolatis. Folia superiora pinnatipartita, lobis brevibus, folia

omnia glabra vel aliquando petiolis sparse ciliolatis. Racemus fructiferus elongatus,

laxus. Sepala 0- 7-0- 8 mm longa, oblonga. Petala 0 -2 m longa. Stamina 2. Nectaria

parva, triangularia. Siliculae 2-2-3 mm longae, 1-7-1 -8 mm latae, ovato-oblongae

vel ovatae, emarginatura satis profunda, lobis saepe plusminusve convergentibus;

stylus quam emarginatura brevior. Semina 1-3-1 -4 mm longa, 0-6-0 -7 mm lata.

Type: Natal: Hlabisa District, Hluhluwe, Ward 4577 (PRE, holo.).

Annual? or biennial herbs 30-60 cm high with spreading branches in the upper

part. Stems puberulous with short dr clavate hairs. Lower stem leaves sub-bipinnati-

partite with spreading or dr recurved, narrow, acute, linear-lanceolate lobes, or some

few leaves oblanceolate, pinnatifid. Upper leaves pinnatipartite with short lobes; all

leaves completely glabrous or occasionally the petioles sparsely ciliolate. Racemes

terminal, in fruit elongate, lax; rhachis puberulous. Pedicels puberulous on the upper

surface; in fruit 1-5-3 -5 mm long, slender, ascending or arcuate. Sepals 0-7-0 -8

mm long, oblong with membranous margins. Petals 0-2 mm long. Stamens 2; fila-

ments subulate. Nectaries small, triangular. Siliculae 2-2-3 mm long, 1-7-1 -8 mm

broad, ovate-oblong or ovate, fairly deeply emarginate with the lobes frequently d:

converging; style shorter than the sinus. Seeds 1 -3-1 -4 mm long, 0-6-0- 7 mm broad,

red-brown.

A distinctive species, apparently not a weed, growing in sandy soil in light shade

in open savanna.

Natal. — Hlabisa: Hluhluwe, Ward 4577. Ntambanana, Salberg sub Curson 15.

Mozambique. — Lourenco Marques: Gomes e Sousa 3654 (K); Khocene, Junod 314 (Z).

Synonymy: L. divaricatum subsp. eu-divaricatum var. dissectum Thell. in Viertelj.

Naturf. Ges. Ziir. 51: 166, 167 (1906). Type: Junod 314 (Z, holo.!).

Lepidium transvaalense Marais, sp. nov. distincta.

Herba perennis parva; caules plures, 15-25 cm alti, sparse vel dense retrorso-

pubescentes. Folia basilaria et caulina inferiora 2-5-5 cm longa, angusta, 7-9 pinnati-

lobata, lobis brevibus oblanceolatis, vel folia oblanceolata breviter lobata. Folia

caulina media 0-5-3 cm longa, 3-7 pinnatilobata, lobis brevibus acutis puberulis cilio-

latis. Folia superiora acute dentata, lineari-oblanceolata vel cuneato-oblanceolata.

Racemus fructiferus densus. Sepala 0 • 6-0 • 8 longa, oblonga. Petala 0 • 3-0 • 4 mm longa.

Stamina 2. Nectaria parva, triangularia. Siliculae 1-8-2 -5 mm longae, 1-4-1 -7 mm

latae, ovato-oblongae vel subrotundo-oblongae, leviter emarginatae, stylo in emargina-

tura incluso. Semina 0-9-1 mm longa, 0-4-0 -6 mm lata.

Type: Transvaal: Bethal District, Leendertz 3574 (PRE, holo.).

110

Small perennial herbs, several-stemmed from a woody crown, or sometimes with

branched woody stems, 15-25 cm high, sparsely to densely retrorse pubescent. Basal

and lower stem-leaves 2 • 5-5 cm long, narrow, pinnately 7-9-lobed with short (0 • 5 cm),

lanceolate lobes, or leaves broadly oblanceolate, shortly lobed. Middle stem leaves

0- 5-3 cm long, pinnately 3-7-lobed with short sharp lobes, puberulous (especially

on the lower surface), ciliolate. Upper leaves sharply toothed, linear-oblanceolate to

cuneate-oblanceolate. Racemes terminal, in fruit dense, 4-8 (-12) cm long. Pedicels

puberulous on the upper surface, ascending or arcuate. Sepals 0-6-0 -8 mm long,

oblong, membranous-margined. Petals 0-3-0 -4 mm long, white. Stamens 2; fila-

ments subulate. Nectaries small, triangular. Siliculae 1 -8-2-5 mm long, 1 -4-1 -7 mm

broad, ovate-oblong to subcircular-oblong, shallowly emarginate, the style included

in the sinus. Seeds 0-9-1 mm long, 0-4-0 -6 mm broad, brown.

Transvaal. — Bethal: Leendertz 3574 (PRE); Machadodorp, Young in TRV 27231. Ermelo: Burtt

Davy 7721; Henrici 1278. Heidelberg: Leendertz 2639; Burtt Davy 9145. Standerton: Jenkins

in TRV 10043. Volksrust: Mogg in T.D.A. Herb. 9687 (K). Potchefstroom : Liebenberg 1019.

Natal. — Estcourt: Edwards 432 (NU). Pietermaritzburg: Wilms 1855 (K).

Orange Free State. — Harrismith: Witzieshoek, Junod in TRV 17369.

Basutoland. — Leribe: Dieterlen 137b.

Cape. — ? Prieska: Bryant B 7.

The habit of this species is distinctive, but its relationship is not clear. Included

by Thellung in his wide concept of L. divarication , which is here treated as several

distinct species.

The majority of the specimens are remarkably uniform. Junod in TRV 17369,

Mogg in T.D.A. Herb. 9687, Wilms 1855 and Dieterlen 137b differ in the broader,

cuneate-oblanceolate leaves. Edwards 432, collected in a very advanced state, obviously

belongs here but is much more robust than the other specimens.

THLASPEOCARPA

Thlaspeocarpa namaquensis Marais, sp. nov., a T. capensi C.A. Sin. floribus fruc-

tibus et seminibus minoribus, racemis fructiferis laxis differt.

Herba annua; caules et rami minute puberuli. Folia 3-10 cm longa, pectinate

3-9-lobata, lobis linearibus. Racemi fructiferi laxi. Sepala 1-5-2 mm longa, 0-8-1

mm lata, puberula vel glabra, interiora 2 basi leviter saccata. Petala 2-3 ■ 5 mm longa,

1- 3-1 -8 mm lata, obovata. Filamenta breviora minute appendiculata. Antherae

0- 3-0 -5 mm longae. Pedicelli fructiferi 1-1-5 cm longi, patuli, gracillimi. Siliculae

1 — 1-3 cm longae, 0-8-1 cm latae, subrotundae vel subrotundo-obovatae, compressae,

pendulae; valvae 1-nervae, membranaceae, maturitate reticulatae. Semina 4-7-5 mm

diametro, rotundata, compressa, late alata.

Type: Cape: Namakwaland District, Steinkopf, Acocks 19536 (PRE, holo.).

Annual herbs 15-35 cm high, branched in the upper part; stems and branches

minutely puberulous. Feaves 3-10 cm long, pectinately 3-9-lobed, the lobes broadly

linear, acute, puberulous. Racemes soon lengthening, lax in fruit. Sepals 1 • 5-2 mm

long, 0-8-1 mm broad, puberulous or glabrous, membranous-margined; the inner

2 slightly saccate. Petals 2-3-5 mm long, 1-3-1 -8 mm broad, obovate, rounded,

cuneate. Filaments of the short stamens with minute appendages. Anthers 0-3-0 -5

mm long. Fruiting pedicels 1-1-5 cm long, wide-spreading, very slender. Siliculae

1- 1-3 cm long, 0-8-1 cm broad, subcircular or subcircular-obovate, compressed,

pendulous; valves 1 -nerved, thin, finely but conspicuously reticulately veined at

maturity. Seeds 4-7-5 mm in diameter, circular, broadly winged with a narrow sinus.

The flowering time is from August to September. Apparently confined to northern

Namaqualand. The plants are more strongly pigmented than in T. capensis , the whole

plant usually having a purplish tinge. Differs from T. capensis in the small fruits and

seeds, the lax fruiting racemes, and in the flowers being smaller in all their parts.

Cape. — Namakwaland: Kasteelpoort, near Klipfontein, Bolus 6502; Bolus in Herb. Norm. Austro-

Afric. No. 1102; Steinkopf, Acocks 19536. Without exact locality: Stokoe in PRE 29579.

APLANODES, gen. nov.

Aplanodes Marais, gen. nov. (Cruciferae-Arabideae), Cardamini L. affinis, sed

foliis simplicibus vel pinnatifidis, indumento pilorum ramosorum, fructuum valvis

costatis manifestis differt.

Herbae perennes, pilis irregulariter ramificatis. Folia ovata, petiolata, inferiora

basi cordata vel pinnatifida lobis brevibus patentibus. Racemi terminales. Sepala

patentia, interiora 2 leviter saccata. Petala obovata, unguicularia, alba usque ad

purpurata, leviter emarginata. Filamenta plana vel basi crassa. Nectaria lateralia

circum bases filamentorum brevium annulatiformia, in annulo nectariis 2 medianis

loratis extra filamenta longa conjunctiva. Ovarium sessile; ovula in quoque loculo

6-18; stigma capitatum, brevissime 2-lobatum. Siliquae lineares, valvis planis 1-nervatis

reticulatis acutis. Semina uniseriata, late ovata, alata, cotyledonibus accumbentibus.

Type species: A. doidgeana Marais.

Perennial herbs with irregularly branched hairs. Leaves ovate, petiolate, the lower

ones cordate at the base or pinnatifid with short spreading lobes. Racemes terminal.

Sepals spreading, the inner 2 slightly saccate. Petals obovate, clawed, slightly emar-

ginate, white to purple. Filaments flattened or thickened at the base. Lateral nectaries

ring-shaped, closed round the bases of the short filaments; united into a ring with

the ribbon-shaped median nectaries which are outside the long filaments. Ovary

sessile; ovules 6-18 per locule; stigma capitate, very shallowly 2-lobed. Siliquae

linear, the valves flat, 1 -nerved, net-veined, acute. Seeds in 1 row, broadly oval,

winged; cotyledons accumbent.

Species 2, natives of South Africa.

The name is derived from the negative of the Greek word P/anodes, “ to wander

about ”, hence “ not to wander about ”, i.e. not an introduced plant as I first thought

it might be. Also, the type species has never been described before (cf. Planodes Greene

ex Schulz in Pflanzenfam. 17b: 541, 1936).

Aplanodes doidgeana Marais, sp. nov., species typica.

Herba perennis, ad 75 cm alta. Caules sparse vel dense puberuli, pilis irregulariter

ramificatis. Folia simplicia, pilis ramificatis dense puberula, subtus pilis plerumque

longioribus minus ramosis, petiolis basi leviter dilatatis; basalia ovato-rotundata vel

reniformi-rotundata, basi cordata, obtusa, crenata, conferta, rosulata, mox caduca;

folia caulina inferiora basi leviter cordata; superiora basi cuneata, apicibus obtusis

vel acutis, dentata vel acute serrata, ovato-oblonga vel ovata. Pedice/li fructiferi 1-1*7

cm longi, graciles, puberuli, late recurvato-patentes. Sepala 3-5-5 mm longa, puberula;

exteriora 2 cucullata, corniculata; interiora 2 basi leviter saccata. Petala 6-9-5 mm

longa, 2 -5-4 -5 mm lata, obovato-cuneata, unguiculata, leviter emarginata. Filamenta

2 -5-3 -5 mm et 3-5-5 mm longa, longioribus 4 basi applanatis, brevioribus 2 basi

incrassatis. Antherae 1-5-1 -7 mm longae. Ovarium puberulum vel glabrum; ovula

in quoque loculo 6-10. Siliquae 3-5-5 cm longae, 2-5 mm latae, lineares; valvae

1-nervae; stylus 4-6 mm longus, robustus. Semina 3-5 mm longa, 2-3 mm lata, late

ovalia, alata.

112

Type: Natal: Bergville District, Cathkin Peak Hostel, West 15 (PRE, holo.).

Perennial branched herbs up to 75 cm high. Stems thinly to densely puberulous

with irregularly branched hairs; some hairs longer, less branched to simple. Leaves

simple, densely puberulous with branched hairs, lower surface usually with longer,

less branched hairs, petiolate, the petiole slightly dilated at the base; basal leaves

petiolate, the petiole slightly dilated at the base; basal leaves ovate-circular or reniform-

circular, cordate at the base, obtuse, crenate, crowded, rosulate but dying off early;

lower stem leaves shallowly cordate, the upper cuneate at the base, obtuse to acute,

dentate to sharply and irregularly or doubly serrate, ovate-oblong to ovate. Racemes

dense in flower, lax in fruit; rhachis puberulous. Fruiting pedicels 1-1-7 cm long,

slender, widely recurved-spreading, puberulous. Sepals 3-5-5 mm long, puberulous,

membranous-margined; outer 2 narrowly oblong, ± cucullate, horn-tipped; inner

2 ovate, ± saccate. Petals 6-9-5 mm long, 2 -5-4 -5 mm broad, obovate-cuneate,

clawed, shallowly emarginate. Filaments 2 -5-3 -5 mm and 3-5-5 mm long; long

ones flattened at the base; short ones thickened at the base. Anthers 1-5-1 -7 mm

long. Ovary linear-oblong, glabrous or with coarse hairs on the valves; ovules 6-10

per locule. Siliquae 3-5-5 cm long, 2-5 mm broad, linear; valves 1-nerved, net-veined;

style 4-6 mm long, stout; stigma capitate, shallowly 2-lobed. Seeds 3-5 mm long,

2-3 mm broad, broadly oval winged.

Named after Dr. Doidge who first collected the species in 1920. The flowers

are described as white, but some of the dried specimens show a definite pinkish tinge.

Collected in flower from October to May.

Natal. — Bergville: National Park, Galpin 10317 (K); Cathedral Peak Forestry Stn., Killick 1862;

Cathkin Peak Hostel, West 15; Tugela Valley, Doidge s.n.; Acocks & Hafstrom 524.

Since the description was drawn up, I have received from the Government

Herbarium, Stellenbosch, a specimen of Thode 7778, collected in the Injasuti Valley,

altitude 6-7,000 ft., in November 1890. This specimen differs from all the others in

having only simple hairs, and somewhat narrower, ovate-lanceolate upper stem leaves.

Its flowers are described as dull lilac. This specimen consists of only the upper part

of the plant, and appears rather more robust than the other specimens. All other

characters are the same.

Aplanodes sisymbrioides ( Schltr .) Marais, comb. nov.

Heliophila sisymbrioides Schltr. in Journ. Bot. 35: 340 (1897). Type: Queenstown,

Galpin 2260 (B !).

An insufficiently known species, differing from A. doidgeana in the pinnatifid

leaves.

BRACHYCARPAEA

Brachycarpaea juncea {Berg.) Marais, comb. nov.

Cleome juncea Berg., Descr. PI. Cap. 164 (1767); Thunb., Prodr. 109 (1800). Type:

Grubb s.n. (SBT!). C. laxa Thunb., l.c. (1800). Type: Thunberg, Herb. no. 15423!

Fleliophila flava L.f., Suppl. 297 (1781). Type: Thunberg in Herb. Linn. no. 8407!

Brachycarpaea varians DC., Syst. 2: 699 (1821); Sond. in FI. Cap. 1: 33 (1860).

B. varians var. flava (L.f.) DC., l.c. (1821); Sond., l.c. (1860). B. varians var. pur-

purascens DC., l.c. (1821); Sond., l.c. (1860). Type: Herb. Burmann sub. Polygala

bracteolata. B. polygaloides Eckl. & Zeyh., Enum. no. 51 (1834-5). B. linifolia Eckl.

& Zeyh., l.c. no. 53 (1834-5). B. laxa (Thunb.) Sond., l.c. (1860). B. laxa var.

stricta Sond., l.c. (1860). B. flava (L.f.) Druce in Rep. Bot. Exch. Cl. Br. 1. 1916:

610(1917). B. capensis Fourc. in Bot. Surv. Mem. 20: 36 (1941), excl. synon. Linn.

Coronipus anomalus Spreng., Syst. Veg. 2: 853 (1825), nom. illeg.

113

Bothalia 9, 1: 113— 121.

Notes on Boerhavia in Southern Africa

by

L. E. Codd

Abstract

A key is provided to the six species recognized in South Africa, of which B. pterocarpaS. Wats,

and B. erecta L. are introduced weeds from America. B. deserticola (South West Africa) is described

as new.

During the preparation of a handbook on the more common weeds of South

Africa, it became necessary to review the names in use for the species of Boerhavia

present in this country. Many of the species are cosmopolitan weeds and, as is common

experience in such cases, their nomenclature presents some difficulty. Thus, although

the available herbarium material may be sorted into groups of specimens, it is not

always easy to assign a name to each group. For assistance in this respect I am

grateful to our Liaison Officer at Kew, Dr. O. A. Leistner, who, in turn, received

valuable advice from Mr. R. D. Meikle, the author of the treatment of Boerhavia in

the second edition of the Flora of West Tropical Africa, Vol. 1, p. 177 (1954).

In the Flora Malesiana 6, 3: 452 (1964), Stemmerik takes a broad view of the

genus and reduces Commicarpus Standi, to synonymy. This view seems unnecessarily

drastic as the two genera can be distinguished chiefly by the character of the fruits,

although florally they are very close. In South Africa, six species of Boerhavia and

four species of Commicarpus are recognized. The present study is concerned with

Boerhavia sens, strict.

Boerhavia fruits (anthocarps) are very small but, under magnification, often provide

useful diagnostic features. When a clear-cut fruit character is associated with a

relatively constant facies, it is considered that such a combination provides grounds

for separate taxonomic status, especially when the type of pubescence is also relatively

constant within the group of specimens.

BOERHAVIA

Boerhavia I., Sp. PI. 3 (1753); Gen. PI. ed. 5: 9 (1754); Willd., Sp. PI. 1: 19 (1797) (as

“ Boerhaavia ”) ; Choisy in DC., Prodr. 13, 2: 449(1852); Heimerl in Pffanzenfam. 3,

lb: 26 (1889); 16c: 117 (1934); Bak. & Wright in FI. Trop. Afr. 6, 1 : 2 (1909);

T. Cooke in FI. Cap. 5, 1: 393 (1910); Standley in N. Amer. FI. 21: 204 (1918);

Burtt Davy, FI. Transv. 1 : 209 (1926); Cufodontis in Bull. Jard. Bot. Brux. 23 (Suppl.):

76: (1953) (as “Boerhavia”); Meikle in FI. W. Trop. Afr. ed. 2, 1: 177 (1954);

Stemmerik in FI. Males. 6, 3: 452 (1964).

A small genus containing several cosmopolitan weeds and other species of more

restricted distribution. Six species are recognized in South Africa, one of which is

subdivided into three varieties. Probably three or four species are indigenous and

there is a possibility that, in certain cases, there may be indigenous and introduced

populations.

Synonymy was investigated only to the extent necessary to ascertain, as far as

possible, the correct name for each taxonomic unit recognised in South Africa.

114

B. diffusa var. diffusa is reported to be a good fodder plant, but the sticky fruits

x>f this, and certain other species, are generally regarded as being objectionable,

especially in wool-producing areas.

Fruit eglandular:

Fruit 3-4-winged 1. B. pterocarpa

Fruit 5-ribbed:

Fruit truncate at the apex, ribs obscurely undulate; perianth purple, up to 2-5 mm long;

leaves broadly ovate; stems puberulous 2. B. erecta

Fruit rounded at the apex, ribs smooth: perianth white to pale pink, 3-5 mm long; leaves

ovate or ovate-elliptic to ovate-lanceolate; stems puberulous to glandular-hispid

5. B. deserticoia

Fruit glandular:

Flowers in ultimately lax, much-branched cymes; leaves usually more than 2-5 cm long:

Leaves broadly ovate to subrotund; stems glabrous to densely glandular-hirsute:

Stems usually glabrous; young leaves fringed with long septate hairs, otherwise glabrous,

usually not wrinkled below; flowers deep purple 3a. B. diffusa var. diffusa

Stems puberulous to glandular hirsute; leaves thick-textured, often wrinkled and whitish

below, usually puberulous to hispid; flowers white to deep pink:

Stems puberulous to hispid often with scattered multicellular hairs; leaves usually thinly

hispid below, glabrescent 3b. B. diffusa var. hirsuta

Stems and leaves densely hirsute, viscid, with stipitate glands often present 3c. B. diffusa

var. viscosa

Leaves ovate-elliptic to lanceolate; stems densely glandular-hirsute 4. B. hereroensis

Flowers in axillary clusters or short cymes up to 2-5 cm long; leaves small, usually less than

2-5 cm long 6. B. repens

1. B. pterocarpa S. Wats, in Proc. Am Acad. 17: 376 (1882); Standley in N.

Amer. FI. 21: 209 (1918). Type: from Arizona, U.S.A.

B. cordobensis Kuntze, Rev. Gen. 3, 2: 264 (1898). Type: from Argentinia, S.

America.

Annual herb. Stems prostrate, puberulous to hispid with scattered long septate

hairs. Leaves broadly ovate to subrotund; blade 2-5-4 cm long and 1-5-2 -5 cm

broad, sparsely hispid on the margins and nerves below, glabrous above, apex acute

to obtuse, base truncate; petiole 0-5-2 -5 cm long. Inflorescence cymose, usually

freely branched: peduncles glandular-puberulous. Flowers small, in many-flowered

glomerate capitula, colour not recorded on specimens collected in South Africa.

Anthocarp eglandular, obconical-clavate, 3-4-winged, broadly truncate at the apex

(Fig. 1:2).

A weed, widespread in the warmer parts of America. First recorded in South

Africa from the Kimberley District in 1919 and now known to occur in several districts

in central and northern Cape Province and western Orange Free State. Common

and evidently spreading along roads and in gardens.

Cape. — Beaufort West: Adamson D162; Theron 1615; Nelspoort, Lewis 4015 (SAM); [Fraserburg

Road, C. A. Smith 2497. Graafif Reinet: Swart River, Maguire 725 (NBG). Kimberley: Leistner

938; Schmidt’s Drift, Jan. 1919, Wilman in BOL 15854 (BOL), in SAM 13307 (SAM); Kalk Drift,

Brueckner 780. Prieska: Bryant J198; Wilman in BOL 3047 (BOL). Prince Albert: Prince Albert

Road, Compton 23903 (NBG). Steytlerville: Gill 55. Taung: Parker s.n.

Orange Free State. — Fauresmith: Leistner 1084. District unknown: Pocock in SAM 52266

(SAM); Mostert 1217.

Superficially similar to B. diffusa var. diffusa but is a smaller plant and may readily

be separated by the eglandular fruits, truncate at the apex, with 3 or 4 wing-like ridges

(see Fig. 1). The margins of the ridges are smooth, not undulate as in the next species.

115

The South African specimens cited above are an excellent match of the type of B.

cordobensis Kuntze, which comes from the Argentine. In addition, a specimen of

Leistner 1084 was sent to the Gray Herbarium of Harvard University where it was

reported to be an almost perfect match of the type of B. pterocarpa S. Wats.

2. B. erecta L., Sp. PI. 3 (1753); Standley in N. Amer. FI. 21 : 210 (1918); Meikle

in FI. W. Trop. Afr. ed. 2, 1: 178 (1954); Woodson et al. in Ann. Miss. Bot. Gard.

48: 51 (1961). Type: America, LINN 9-1 (PRE, photo.).

Annual herb. Stems decumbent to prostrate, puberulous, with scattered long

septate or gland-tipped hairs, sometimes viscid. Leaves ovate to broadly ovate, sparsely

hispid to subglabrous on the margins and on the nerves below, glabrous above, apex

acute to obtuse, base rounded to truncate; blade 2-4-5 cm long and 1 -5-3 cm broad;

petiole 1-2 cm long. Inflorescence cymose, usually freely branched; peduncles glandular-

puberulous. Flowers minute, magenta, in glomerate, 4-10-flowered capitula. Anthocarp

eglandular, clavate, broadly obtuse at the apex, tapering to the base, 5-ribbed, the ribs

slightly undulate along the margins (Fig. 1:1, la, b, c, d).

A weed, widespread in the warmer parts of America. First recorded in South

Africa from the Bloemfontein District in 1934. Now known, in addition, from Pretoria

and several localities in the eastern Cape Province. It also occurs in tropical Africa.

Cape. — Bedford: 5 miles S.E. of Daggaboers Nek, Acocks 15802; Baviaansrivier, Theron 407.

Somerset East: Marais s.n. Uitenhage: Weed Inspector s.n.

Orange Free State. — Bloemfontein: Tempe Farm, 1934, Motsamoi 4958.

Transvaal. — Pretoria: Repton 4856; Walters in PRE 29073; Codd 10514.

May be recognised by the eglandular, 5-ribbed fruits, truncate at the apex, and

with obscurely undulate margins.

3. B. diffusa L., Sp. PI. 3 (1753). Type: India, LINN 9-3 (PRE, photo.).

Stems decumbent to prostrate, diffusely branched, glabrous or variously pubescent

from minutely puberulous, with or without long brown septate hairs, to hispid or

glandular-hispid, sometimes viscid. Leaves broadly ovate to subrotund, glabrous,

except for a fringe of septate hairs on the margins, to sparingly or densely hispid, apex

obtuse to rounded, base truncate to subcordate; blade 3-5-5 cm long and 2 -5-4 -5 cm

broad; petiole 1-3 cm long. Inflorescence cymose, ultimately diffusely branched;

peduncles glabrous to puberulous. Flowers small, whitish to red or purple, in 2-5-

flowered capitula. Anthocarp glandular, clavate, 5-ribbed, rounded at the apex, tapering

to the base; ribs entire, not undulate.

Three varieties recognized in South Africa, differing mainly in pubescence of stems

and leaves. For key to varieties, see key to species.

(a) var. diffusa.

B. diffusa L., Sp. PI. 3 (1753); Choisy in DC., Prodr. 13, 2: 452 (1852); Cufodontis

in Bull. Jard. Bot. Brux. 23 (Suppl.): 76 (1953), pro parte, excl. syn. B. repens I..;

Meikle in FI. W. Trop. Afr. ed. 2, 1: 178 (1954). B. repens var. diffusa (L.) Hook.f.

in FI. Brit. Ind. 4: 709 (1885); Bak. & Wright in FI. Trop. Afr. 6, 1 : 5 (1909); T.

Cooke in FI. Cap. 5, 1: 394 (1910). B. adscendens Willd., Sp. PI. 1: 19 (1797); Bak.

& Wright in FI. Trop. Afr. 6, 1 : 4 (1909); T. Cooke in FI. Cap. 5, 1: 395 (1910).

Type: from Guinea (Herb. Willd.).

Stems glabrous or minutely puberulous with a few long brown septate hairs at

the nodes; leaves glabrous except for scattered long septate hairs on the margins;

underside not markedly wrinkled; flowers mauve, reddish or purple (Fig. 24).

116

117

A weed, widespread in the tropics of both hemispheres, possibly introduced into

South Africa; recorded from the coastal areas of Natal and from eastern and northern

Transvaal.

Natal. — Durban: Medley Wood 7199; 9345; 12220; Isipingo, Ward 586. Pietermaritzburg:

Umgeni Dam, Barker 5221 (NBG). Umlazi: Amanzimtoti, Medley Wood 13206.

Transvaal. — Barberton: Pott 5475; Kaapmuiden, Rogers 30050. Nelspruit: Liebenberg 2499.

Letaba: Modjadji's Reserve, Krige 126 ; Duiwelskloof, Scheepers s.n. Pretoria: Wonderboom,

C. A. Smith 2292; 2298; Wasserfal & Van Niekerk s.n. Sibasa: Punda Milia, Lang in TRV 32168;

13 miles N.E. of Sibasa, Codd 6883.

( b ) var. hirsuta Heim, in Bot. Jahrb. 10: 9 (1889). Type: “ Kuruman ”, Mar/oth

1102 (on the PRE specimen the locality of this number is givep as “ Namaqualand,

Ga Mhana Peak ”).

B. schinzii Heim, in Ver. Bot. Verein. Brandenburg 31 : 222 (1889). Type: Ovambo-

land, Schinz 741 (Z, holo.; BOL!; PRE, photo.). — — forma cmgustcita Heim., l.c.

223 (1889). Type: Hereroland, Liideritz 174. — — forma vi/losa Heim., l.c. 223 (1889).

Syntypes: Hereroland, Omaruru, Schinz 742 (Z; PRE, photo.); Liideritz s.n. B.

marlothii Heim, in Bot. Jahrb. 10: 9 (1889). Type: Otjimbingwe, Marloth “ 1372 ”

(this is the number cited with the type description on p. 9 though, later in the same

publication, Heimerl changed the number to 1342; on the PRE specimen, the original

number 1342 has been crossed out and replaced by 1372). B. diffusa forma subhirsuta

Heim, in Bull. Herb. Boiss. 5, Appendix 3: 67 (1897). Type: Amboland, Hopfner

66. B. bracteata T. Cooke in Kew Bull. 1909: 421 (1910); FI. Cap. 5, 1: 394 (1910);

Burtt Davy, FI. Transv. 1: 209 (1926). Syntypes: Transvaal, Barberton, Galpin 1240

(PRE); Bechuanaland, Chue Valley, Burchell 2381 (PRE); Natal, Tugela, Gerrard

1787. B. diffusa sensu Burtt Davy, l.c. 209 (1926). B. coccinea Mill. var. a Meikle

in FI. W Trop. Afr. ed. 2, 1 : 177 (1954).

Stems hispid, often with numerous long brown septate hairs, sometimes glabrescent

with age; leaves with scattered hispid hairs on the nerves below, with septate or hispid

hairs on the margins, usually glabrous above, thick-textured; lower surface usually

whitish and wrinkled; peduncles glabrous; flowers white, pink or mauve, rarely

pinkish-purple.

Widespread in the drier parts of tropical Africa and America. This is the com-

monest member of the genus in South Africa, occurring in northern Natal, Swaziland,

Transvaal, northern Cape Province and South West Africa. It is probably indigenous

but behaves as a weed, spreading on bare areas, the sticky fruits being dispersed by

grazing animals.

Natal. — Ngotshe: Pongola Bridge, Gerstner 2460.

Swaziland. — Hlatikulu: Big Bend, Compton 30913.

Transvaal. — Barberton: Avoca, Galpin 1240; Kaapmuiden, Rogers 24286: Komatipoort ,Schlechter

11874. Lydenburg: Sekukuniland, Barnard & Mogg 981. Witrivier: Kruger National Park, near

Skukuza, Letty 31 ; Codd & De Winter 5010; Lower Sabie Rest Camp, Van der Schijffl99. Pretoria:

Pienaars River, Schlechter 4221; Leendertz 774. Sibasa: 32 miles N.E. of Punda Milia, Codd 5981.

Soutpansberg: Wylliespoort, Breyer in TRV 22025; Rodin 4234; Waterpoort, Bremekamp &

Schweickerdt 298; Soutpan, Schweickerdt & Verdoorn 551; Messina: Messina, Rogers 19273; 19433;

19434; 21191; 21226; Dongola area, Codd 4385.

Cape. — Kimberley: Windsorton, Marloth 4823; Leeuwpoort, Acocks 8578. Namakwaland: Ga

Mhana Peak, Marloth 1102. Postmasburg: Klapin, Leistner 2065. Kuruman: Silk 1341; Chue

Valley, Burchell 2381.

South West Africa. — Bethanien: Kinges 2074; Buchholzbrunn, Pearson 3664 (BOL). Groot-

fontein: Schoenfelder 551. Kaokoveld: Namutoni, Barnard 550 (SAM); Kunene Banks, Barnard

551 (SAM). Karibib: Seydel9\l\ 1335; Otjimbingwe, Marloth 1404 (SAM); 1372. Keetmanshoop:

Great Karasberg, Pearson 8473 (BOL). Rehoboth: Bullspoort, Tdlken & Hardy 662. Okahandja:

Dinter 357 p.p.; Merxmiiller & Giess 1020; near Okahandja, Tdlken & Hardy 723; 22 miles from

Okahandja on road to Kalkfield, Tdlken & Hardy 730. Okavango Native Territory: Cigarette,

118

Maguire 2317 ; 6 miles E. of Tondoro Mission, De Winter 3968. Outjo: Thorne in SAM 31701 (SAM);

near Kamanjab, De Winter & Leistner 5118; near turn-off to Welwitschia on Fransfontein road to

Outjo, Tolken & Hardy 846; 20 miles from Fransfontein on road to Outjo, Tdlken & Hardy 859.

Ovamboland: Schinz 741 (BOL). Windhoek: Gil/man 152 (SAM); Wilman in BOL 15227 (BOL).

Bechuanaland. — Ngamiland, Van Son in TRV 28953.

Intermediates between this variety and the previous one occur in the northern

and eastern Transvaal, but the great majority of specimens can be allocated without

difficulty.

Pearson 9176 (BOL) from Ababes, Rehoboth District, probably is a form of

B. diffusa var. hirsuta, although it is described as being a “ 3 ft. bush The leaves

(immature?) are only 1-1 -4 cm long and 0-6-1 cm broad; the inflorescence is diffusely

branched, but there are only 1 or 2 flowers to each ultimate peduncle. However, the

anthocarps and the pubescence on stems and leaves agree well with B. diffusa var.

hirsuta.

(i c ) var. viscosa (Lag. & Rodr.) Heim, in Beitr. Syst. Nyct. 27 (1897). Type: from

South America.

B. viscosa Lag. & Rodr. in Anal. Cienc. Nat. 4: 256 (1801). B. coccinea Mill. var.

viscosa (Lag. & Rodr.) R.M. Mosc. in Cat. FI. Doming. 1: 180 (1943); Meikle in

FI. W. Trop. Afr. ed. 2, 1: 177 (1954).

Stems densely glandular-hispid, viscid; leaves densely hispid on both surfaces;

peduncles glandular-puberulous; flowers pink to reddish-purple.

Indigenous in dry, rocky places in northern Cape Province and South West Africa.

Described originally from South America and also present in tropical West Africa.

Cape. — Gordonia: near Nakop Station, Acocks 18813. Hay: Magoras, Acocks 1985. Prieska:

between Prieska and Marydale, Acocks 1757; 16 miles E. of Draghoender, Codd 1237.

South West Africa. — Keetmanshoop : Great Karasberg, Pearson 8174 (BOL); near Narubis,

Wilman 328 (BOL). Maltahohe: Dinter 2002 (2 sheets in SAM dated 16/3/1911 and 31/3/1911).

4. B. hereroensis Heim, in Bot. Jahrb. 10: 9 (1889); Bak. & Wright in FI. Trop.

Afr. 6, 1 : 3 (1909). Type: Otjimbingwe, Marloth 1403 (PRE, SAM).

Stems suberect to erect, 30-40 cm long,g landular-hirsute, viscid. Leaves ovate-

elliptic or ovate-lanceolate to lanceolate; blade 2-3 cm long and 0-5-1 -5 cm broad,

glandular-hispid on both surfaces, apex acute, base rounded; petiole 1-2 cm long,

glandular-hispid. Inflorescence cymose, freely branched; peduncles glandular-puber-

ulous. Flowers solitary or in 2-4-flowered capitula; perianth 2-2-5 mm long, purple.

Anthocarp glandular, elliptic-clavate, 5-ribbed, rounded at the apex, margin of ribs

entire (Fig. 2: 5).

Indigenous in the northern Cape Province and South West Africa, usually in

stony places.

Cape. — Gordonia: near Riemvasmaak, Acocks 16391.

South West Africa. — Bethanien: Kuibis, Range 922 (SAM). Kaokoveld: Kapupa Valley, Story

5889. Karibib: Kinges 3136; Seydel 1436; Otjimbingwe, Marloth 1403; Small Spitzkop, Tolken

& Hardy 810. Keetmanshoop: Great Karasberg, Pearson 8474 (BOL). Okahandja: Dinter 357

(SAM), p.p. Omaruru: Brandberg, Merxmiiller & Giess 1663.

The pubescence is similar to that found in B. diffusa var. viscosa, but the leaves

are constantly narrower in relation to their length, and the stems tend to be more

ascending.

119

5. B. deserticola Codd, sp. nov., B. hereroensi Heim, affinis, sed fructibus eglan-

dulosis et perianthio majore differt.

Caules decumbentes 40-80 cm longi; rami puberuli vel glanduloso-hirsuti. Folia

opposita, ovata vel ovato-lanceolata, subglabra vel glanduloso-hispidula, 3-5-5 cm

longa, 1-3-2 -8 cm lata, apice acuta, basi cuneata vel obtusa. Inflorescentia 15-30 cm

longa; bracteae lineari-lanceolatae, acuminatae. Flores in cymis multi-floris dispositi;

perianthium album 3-5 mm longum. Fructus 5-costatus, eglandulosus, elliptico-clavatus,

3 • 5-4 mm longus.

Type: South West Africa: Kaokoveld Reserve, Orupembe Waterhole near Sani-

tatas, 5th May, 1957, De Winter & Leistner 5722 (PRE, holotype; K, M).

Perennial herb. Stems woody at the base, decumbent, 40-80 cm long, divaricately

branched, puberulous to glandular-hirsute and viscid. Leaves opposite, ovate to

ovate-lanceolate, subglabrous to glandular-hispidulous and viscid; petiole 0-5-2 -5 cm

long; blade 3-5-5 cm long and 1-3-2 -8 cm broad, apex acute, minutely apiculate,

base cuneate to rounded; lower surface usually paler and somewhat wrinkled. Inflore-

scence elongate, divaricately branched, 15-30 cm long; rhachis glandular-hispidulous

or puberulous: bracts linear-lanceolate, gradually acuminate, 3-5 mm long. Flowers

glomerate in dense, many-flowered cymes; bracteoles ovate-lanceolate, acuminate,

2 mm long; perianth white to pale pink, 3-5 mm long, 5-lobed; stamens 4, anthers

just exserted at anthesis; style filiform, 6 mm long; stigma peltate. Anthocarp 5-ribbed,

eglandular, 3 • 5^1 mm long, elliptic-clavate, rounded at the apex, ribs entire, conspicuous

(Fig. 2: 1, la, b, 2, 2a).

Known as yet from two restricted areas in South West Africa, one in the Kaokoveld

near Sanitatas, and the other in the Omaruru District, inhabiting dry watercourses

and rocky places.

South West Africa. — Omaruru: Brandberg, Boss in TRV 36251; Merxmiiller & Giess 1645; 8

miles from Uis on road to Welwitschia, Tdlken & Hardy 831 ; 28 miles from Uis on road to Welwitschia,

Tdlken & Hardy 836. Kaokoveld Reserve: Orupembe Waterhole, near Sanitatas, De Winter & Leistner

5722; Anabib (Orupembe), Story 5733.

In general facies this species comes nearest to B. hereroensis Heim, but differs in

the eglandular anthocarps and longer perianth, which is white (rarely pale pink) instead

of deep pink to purple. In the field, the two may also be distinguished by the difference

in habit, B. hereroensis having semi-erect to erect stems, forming a bushy, viscid, herb

about 30 cm tall, while the stems of B. deserticola are decumbent, up to 80 cm long,

ascending at the ends, and are less conspicuously viscid.

The specimens now included in B. deserticola vary considerably in pubescence.

Merxmiiller & Giess 1645 comes nearest to B. hereroensis in this respect, having glandular-

hirsute stems with leaves glandular-hispidulous (glandular-hispid in B. hereroensis).

At the other extreme is the specimen selected as the type, namely De Winter & Leistner

5722, with minutely puberulous stems, glandular-puberulous inflorescence-branches

and peduncles, and subglabrous to minutely puberulous leaves. This gathering was

made on “ outcrops of volcanic origin (amygdaloidal basalt and a lava-like rock)

with overlying limestone ”, and the stems and leaves of the dried specimen have a

greyish-green appearance.

At first, consideration was given to maintaining these two gatherings as distinct

varieties, but the two recent collections Tdlken & Hardy 831 and 836 are somewhat

intermediate in the character of the pubescence. These two gatherings and Merxmiiller

& Giess 1645 are found on granite formation, in dry watercourses and among rocks.

120

Fig. 2.— 1, la, Boerhavia deserticoIa,xl (De Winter & Leistner 5722), lb, fruit, x 6. 2, B. deserticola',

inflorescence, X 6 ( Merxmiiller & Giess 1645); 2a, fruit, x 6. 3, B. repens, fruit, x 6

( Bryant 198a). 4, B. diffusa var. diffusa, fruit, x 6 (Pott 5475); 5, B. hereroensis, fruit, x 6

( Tolken & Hardy 810).

121

6. B. repens L., Sp. PI. 3 (1753); Meikle in FI. W. Trop. Afr. ed. 2, 1 : 177 (1954).

Type: Egypt, LINN 9-8 (PRE, photo.).

B. repens var. minor Del., FI. Egypte Icon, t.3, 1 (1813). Type: It is not known

if a specimen exists, but the illustration will suffice. B. diffusa var. minor (Del.) Cuf.

in Bull. Jard. Bot. Brux. 23 (Suppl.): 77 (1953).

Stems prostrate, pallid, glabrous. Leaves small, broadly ovate or subrotund to

ovate-lanceolate; blade 1-2-5 cm long and 0-6-1 -5 cm broad; petiole 0-5-1 -4 cm

long. Flowers in axillary clusters, in short cymes up to 2-5 cm long, rarely solitary,

often in 2-4-flowered capitula on axillary peduncles, white, pink or mauve. Anthocarp

glandular, elliptic-clavate, rounded at the apex, 5-ribbed, margin of ribs entire (Fig. 2:3).

A species of tropical Africa and Asia, probably indigenous in southern Africa,

where it occurs in dry water-courses and at the sides of pans in northern Cape Province

and South West Africa. Although locally common, it does not appear to be a potential

weed.

Cape. — Gordonia: Obobogorap, Leistner 1783; Gemsbok National Park, Leistner 1048; 1889.

Hay: Acocks 1978. Kenhardt: near Kakamas, Acocks 16367. Namakwaland: near Springbok,

Krapohl in PRE 23295. Prieska: Bryant 198a.

Sooth West Africa. — Gibeon: Mariental, Dinter 1978 (SAM). Karibib: Farm Otjimbojo, Kinges

3380; Farm Ameib, Tolken & Hardy 767. Keetmanshoop : De Winter 3348; Liebenberg 5193;

Seeheim, Dinter 2957 (SAM); Pearson 3724 (BOL); near Asab Tolken & Hardy 637. Windhoek:

Avisdam, Liebenberg 4494.

The type of B. repens L. appears, from the photograph in PRE, to be an immature

specimen, with the result that the interpretation of it has varied considerably. It has

been united with B. diffusa L., both as B. diffusa var. repens and as B. repens var.

diffusa.

The specimens cited above are an excellent match of the plant described by Delile

as B. repens var. minor Del., and it is clear that Meikle interpreted B. repens in the

Flora of West Tropical Africa ed. 2, 1 : 177 (1954) on the basis of var. minor Del. It

is convenient to follow this interpretation here. The South African specimens, whether

they should be correctly known as B. repens L. or B. repens var. minor Del., are certainly

distinct, at the specific level, from those cited under B. diffusa and its varieties, and

from the other South African species.

.

123

Boihalia 9, 1: 123 — 151.

New and Interesting Records of African

Flowering Plants

by

Various Authors

CELASTRACEAE

The Cassine Complex

Dr. N. K. B. Robson, in Bol. Soc. Brot. 39: 5-55 (1965), published an analysis

of the genera allied to Cassine L. occurring in the Flora Zambesiaca area and included

those found in the neighbouring territory of Southern Africa. He deserves credit for

his meticulous study, which has brought together a great deal of data. However,

many of his conclusions do not commend themselves to the present author, who prefers

the broader concept presented by Ding Hou in Flora Malesiana 6, 2: 284 (1962).

An important contribution by Robson concerns the typification of Cassine L.

On the evidence produced, it appears that the genus should be typified by the Cape

species, C. peragua L., Sp. PI. 268 (1753) (= C. capensis L.), an opinion that will

probably be generally accepted. Differences of opinion may arise, however, regarding

his conclusion to restrict the genus to the two Cape species, C. peragua L. and C.

parvifolia Sond. This conclusion, he argues, follows from an attempt to classify the

species into “ natural ” groups, leading him to uphold the following five genera in

addition to Cassine sens. str. : Elaeodendron Jacq.f. ex Jacq. (in which he includes

Lauridia Eckl. & Zeyh.), Crocoxylon Eckl. & Zeyh. (in which he includes Pseudocassine

Bredell), Hartogia L.f., Mystroxylon Eckl. & Zeyh. and Allocassine N. Robson.

Certain of these genera, such as Mystroxylon and Allocassine sens, str., may well

be “ natural ” ones in the sense that they appear to be relatively discrete units in an

intricate and reticulate pattern. The question arises whether any useful purpose is

served by attempting to split up a heterogeneous but easily recognized group into a

number of units based on characters which appear to be matters of degree. The result,

as applied to the South African species, is unsatisfactory and, until more information

of a fundamental nature is forthcoming, which supports this treatment throughout

the entire range of the group, it is preferred to adopt a broad concept of Cassine and

to include within it the genera Elaeodendron , Lauridia , Crocoxylon , Pseudocassine and,

probably, Mystroxylon.

The genus Pseudocassine, with its 3-merous flowers, has previously been regarded

as an aberrant member of the Celastraceae, in which family flowers with three stamens

are exceptional. When seen in perspective, it may be interpreted as the product of

an evolutionary trend within the Cassine complex towards reduction in numbers of

flower parts.

This broad concept of Cassine calls for certain combinations, which are now

effected.

Cassine reticulata {Eckl. & Zeyh.) Codd, comb. nov.

Lauridia reticulata Eckl. & Zeyh., Enum. PI. Afr. Austr. 124 (1834-35).

Elaeodendron reticulatum (Eckl. & Zeyh.) Ettingshausen in Denkschr. K. Akad.

Wiss. Wien 13: 58 (1857); Robson in Bol. Soc. Brot. 39: 39 (1965).

124

Cassine transvaalensis (Bunt Davy) Codd , comb. nov.

Salacia( ?) transvaalensis Burtt Davy in Kew Bull. 1921: 51 (1921).

Pseudocassine transvaalensis (Burtt Davy) Bredell in S. Afr. J. Sci. 33: 330 (1937).

Crocoxylon transvaalsense (Burtt Davy) Robson in Bol. Soc. Brot. 16: 41 (1965).

Hartogia L.f. (1781) is an illegitimate name, being antedated by Hartogia L.

(1759). Consideration may be given to including it in Cassine , but it shows certain

anomalous features which first require thorough investigation. It differs from Cassine

mainly in the endosperm being scanty or absent(?), while the fruit is relatively thin-

walled like that of typical C. aethiopica. Although the fruit usually contains one seed,

a second seed may partially or fully develop. It may be noted that a similar situation

arises in C. tetragona , included by Robson as a second species in his genus Allocassine.

C. tetragona appears to be out of place in Allocassine. The similarities which it shows

to the typical species, A. laurifolia (Harv.) Robson, are suggestive of convergent evolution

rather than a natural relationship. Allocassine sens. str. may well be a good genus,

but it is preferred to return A. tetragona to Cassine as it is clearly related to C. aethiopica.

The Status of the Genus Lydenburgia

Another point on which I must differ from Robson is in connection with his

decision to erect the genus Lydenburgia Robson, based on L. cassinoides Robson, a

species found in Sekukuniland, Lydenburg District. It so happens that I recently

studied a range of material of this species and was in the process of describing it in

the genus Catha. As stated by Robson, it differs from typical Catha in its longer seeds

which are trigonous and possess a small aril, lacking the wing-like aril of typical Catha.

In floral characters and general facies it closely resembles Catha edulis and I have no

hesitation in reducing Lydenburgia to synonymy.

Catha Forsk. ex Scop., Introd. Hist. 228 (1777).

Lydenburgia Robson in Bol. Soc. Brot. 39: 34 (1965).

Catha cassinoides (Robson) Codd, comb. nov.

Lydenburgia cassinoides Robson in Bol. Soc. Brot. 39: 35 (1965).

L. E. Codd

CUPRESSACEAE

Notes on Widdringtonia

In the course of a revision of the genus Widdringtonia Endl. for the Flora of

Southern Africa, attention was given to the typification of W. juniperoides (L.) Endl.,

the name generally applied to the South African “ Cedar ” occurring on the Cedarberg

in the Clanwilliam District of the Cape Province. As a result, the conclusion was

reached that the Linnaean basionym, Cupressus juniperoides L., should be rejected

as a nomen dubium, a view already expressed by Stapf in Flora Capensis 5, 2: 23 (1933).

Unfortunately, Stapf did not carry his opinion to the conclusion required by the Inter-

national Code of Botanical Nomenclature, and he upheld the name W. juniperoides

Endl., Syn. Conif. 34 (1847), non (L.) Endl. Endlicher validly effected the combination

W. juniperoides (L.) Endl., taking up the Linnaean epithet, and enlarging the concept

of the species to include certain specimens from the Cedarberg.

Linnaeus described Cupressus juniperoides in his Species Plantarum ed. 2, 1422

(1763), basing his description on two-year-old plants, originating, according to him,

from “ Caput Bonae Spei ”. There is no evidence that any subsequent author saw

a specimen of the plant described by Linnaeus, and no specimen exists in the present

Linnaean Herbarium. Judging from historical records, it is very unlikely that any

collector visited the Cedarberg at such an early date. If the seed did come from the

125

Cape it is more probable that the species concerned would be the one readily available

at the Cape, which was adequately described a few years later as Thuia cupressoides

L., Mantissa Plantarum 125 (1767). The name W. cupressoides (L.) Endl. has generally

been accepted as the correct one for the S.W. Cape plant. Up to the time of Endlicher,

botanists had occasionally applied the epithet “juniperoides ” to the S.W. Cape plant

e.g. specimens collected by Oldenland (BM) and Burniann (G). The epithet “juni-

peroides ” thus, as the oldest name, should probably be applied to the S.W. Cape plant

known as W. cupressoides. However, this might cause confusion, so it has been decided

to reject the epithet altogether, on the grounds that the epithet is inadequately described

and typified, and might even belong to a genus other than Widdringtonia. If this

course is adopted, W. cupressoides (L.) Endl. remains the correct name for the S.W.

Cape plant, while a name other than W. juniperoides becomes necessary for the Clan-

william “ Cedar ”. Although numerous synonyms with copious ecological and economic

notes exist, none of them appears to be supplied with a botanical description. Thus

the species is now redescribed and typified.

Widdringtonia cedarbergensis Marsh, sp. nov., W. schwarzii (Marloth) Mast,

affinis, sed seminibus minoribus angustissimis alatis et cicatriculis minoribus differt.

Arbor vel frutex 5-20 m altus. Folia adulta squamiformia, appressa, decussata,

ovata. Coni masculi parvi, 4 mm longi; squamae c.12, peltatae, late ovatae, decussatae.

Coni feminei globosi; valvae 4, lignae, rugosae, verrucis ad margines regulariter dispo-

sitis. Semina ovoidea, obscure alata, trigona; seminis cicatricula c. 4-5 mm lata,

6 mm longa. Cotyledones post germinationem 35 mm longae, 5 mm latae.

Callitris arborea Schrad. ex Drege, Zwei Pflzgeogr. Doc. 73 (1843), nom. nud.;

Dallimore in Kew Bull. 35: 222 (1913), nom. nud.; Stapf in FI. Cap. 5, 2: 24 (1933),

in synon. C. ecklonii Schrad. ex Pappe, FI. Cap. Med. Prodr. ed 1 : 25 (1850), nom.

nud. C. juniperoides sensu Durand and Schinz, Consp. FI. Afr. 5: 951 (1894); Engl.,

Pflzw. Afr. 2: 89 (1908); Marloth, Kapland 167 (1908).

Widdringtonia wallichii Endl., Syn. Conif. 34 (1847), nom. nud.; Stapf in FI.

Cap. 5, 2: 23 (1933), in synon. W. juniperoides sensu Endl., Syn. Conif. 32 (1847);

sensu Stapf in FI. Cap. 5, 2: 23 (1933); sensu Hubbard in S. Afr. Journ. Sc. 33: 572

(1937); sensu Smith in Journ. S. Afr. For. Add. 25 (1955); sensu Chapman in Kirkia

1: 138 (1960-61). W. wallichiana Gordon, Pinet. suppl. 107 (1875), nom. nud.

Type: Clanwilliam, Cedarberg Mts., near Middelberg West Peak, 17.6.1965,

Liickhojf in PRE 29824 (PRE, holo.).

Tree normally 5-7 m high, occasionally attaining a height of about 20 m with stem

diameter up to 2 m; crown pyramidal when young, spreading with age; bark on

young trees reddish grey, thin, fibrous, flaking off annually. Leaves of two types;

juvenile leaves on seedlings and young trees needle-like, spreading, 1-2 cm long and

up to 2 mm broad; adult leaves scale-like, appressed, ovate, 2-4 mm long, usually

strictly decussate, semi-circular in transverse section, adnate at the base, free portion

often much shorter than adnate portion. Male cones 2 mm long, mostly on short,

lateral branchlets; scales usually 6 pairs, decussate, coriaceous, peltate, broadly ovate,

acuminate, with 4 pollen sacs at the base of each scale. Mature female cones subglobose,

diameter about 2-5 cm, usually consisting of 4 (rarely 5 or 6) woody valves; valves

rough, with regular verrucae along the margin. Seeds ovoid, obscurely winged,

trigonous; seed scar about 4-5 by 6 mm. Cotyledonary leaves about 35 mm long

by about 5 mm broad. Plate 1.

Found on the Cedarberg Mts. near Clanwilliam in the Cape Province; occurs

singly or in groups over some 30 miles on rocky outcrops at an altitude of between

3,000 and 5,000 feet.

126

Cape. — Clanwilliam: Cedarberg Mts., Coetzee s.n. (PRE); Compton 12776 (NBG); Drege s.n,

(BM, G, L, P, S); Forester Bath 5535 (FD Herb., S); Forester s.n. (L, PRE); Galpin s.n. (PRE);

Kappler 9626 (FD Herb.); Leipoldt 1649 (BM, G, K, P, Z); Liickhojf s.n. (PRE): MacOwan in BOL

27689 (BOL); Marloth 3086 (PRE); Mogg 2010 (PRE); Pocock 522 (STE); Wallich s.n. (BM, K);

Ecklon & Zeyher 74-3 (GRA, L, MO, W, Z; this gathering is attributed to Drege in E and G).

A detailed study of the genus revealed that the species can be divided into two

main groups on the basis of the valve structure of the female cone and the shape of the

adult leaves. One of these groups consists of the two species W. schwarzii and W.

cedavbergensis , which differ only in their seed characteristics. The other group consists

of the species W. cupvessoides in the broad sense, which is now interpreted as including

the following species which were kept separate by Stapf, l.c. : W. stipitata Stapf, W.

dracomontana Stapf and W. whytei Rendle. Although local geographical forms in

this group can sometimes be recognised, the degree of variation is such that no constant

differences could be found on which separate specific or even subspecific status could

be justified.

J. A. Marsh

ERICACEAE

Two New Erica Species from the Drakensberg

Erica dominans Killick, sp. nov., E. glaphyrae Killick affinis, sed foliis adpressis,

inflorescentiis plerumque terminalibus, sepalis basi manifeste imbricatis, ovario ovoideo-

globoso distinguitur.

Fruticulus erectus vel suberectus, 15-45 cm altus; ramuli dense virgati, juvenes

albo-tomentosi, maturi glabri. Folia constanter 3-nata, conferta, adpressa vel interdum

nonnihil suberecta, recta, elliptica, 1-9-3 -3 mm longa, 0-7-0 -9 mm lata, supra plana

vel concava, infra convexo-triangularia, sulcata, praeter margines petiolosque glabra,

olivaceo-viridia, nitida. Flores in acervis cernuis terminalibus vel interdum in axillis

foliorum superiorum dispositi; pedicelli recurvati, 0-7-1 -8 mm longi, albo-tomentosi;

bracteae 2, 1 mm longae, marginibus ciliatis. Sepala 4, cinnamomea erubescentia

vel rubra, basi manifeste imbricata, ovata, 1-1-1 -2 mm longa, 0-5-0 -7 mm lata

(interiora saepe exterioribus angustiora et crassiora), concava, dorse carinata et in

parte superiore sulcata, cartilaginea, marginibus hyalinis ciliatis. Corolla erubescens

vel purpurata accedens (ex Lubkeo brunnea, sed flores speciminis ejus probabiliter

veteres et sicci), cyathiformis, basi nonnihil angusta, 2 -0-2 -5 mm longa; tubus inter

sepala ventricosus; lobi obtusi, 0-5-0 -9 mm longi. Stamina 8, filamentis capillaribus

1 mm longis; antherae 1 mm longae, basi setis (haud caudis) 0-6 minutis instructae.

Ovarium ovoideo-globosum, nonnunquam apice subumbonatum, glabrum, stylo tenui

1-1-1 -2 mm longo, stigmate cyathiformi-peltato incluso vel raro exserto (in Schwei-

ckerdtio s.n. et Lubkeo 275 longe exserto).

Type: Basutoland, Quachas, summit of Drakensberg in Organ Pipes Pass area,

c. 9,800 feet, 2 October 1964, Killick 3519 (PRE, holo.). Fig. 1: 2.

Erica dominans is probably the most common species of Erica on the summit of

the Drakensberg between 9,800-10,500 feet and is often dominant over large areas,

hence the epithet dominans. It is either purely dominant or co-dominant with Heli-

chrysum trilineatum var. tomentosum in what has been described as alpine heath

(Killick, 1963). E. dominans commences flowering in July, reaching a peak in October.

The range of this species is along the Drakensberg from Doodmans Krans Mountain

(not traceable on maps, but probably in the Mount Fletcher District) in the south to

Mont aux Sources in the north with several records from the northern part of Basutoland.

Basutoland. — Leribe : Butha Buthe area, Nchoatle/Khotjoane Pass, Jacot Guillarmod 4004; Khatibe

Camp, above Tlolohatze River Valley, Lubke 275; Maluti Mountains, Staples 19; Ntibokho Valley,

Jacot Guillarmod 296. Quachas: Tsanatalana Valley, Killick 1728; Organ Pipes Pass area, Killick

3519; 3538; 3539; 3540.

127

Fig. 1. — 1, Erica alticola, flowering branch (Thode A1620). 2, E. dominans, flowering branch ( Killick

3519). 3, E. glaphyra, flowering branch ( Killick 3541). a, portion of branchlet with leaf,

x 4; b, leaf, x 6; c, flower, X 10; d, pistil, x 12; e, stamen, anterior, lateral and dorsal

views, x 10.

128

Natal. — Bergville: Mont aux Sources, Edwards 360; Ga/pin 9497; Hutchinson, Verdoorn & Forbes

117; Schweickerdt s.n.; Sidey 2016; Castle Buttress, Esterhuysen 10185; Cleft Peak, Esterhuysen

12892; Organ Pipes Pass, Schelpe 847. Estcourt: Giants Castle, Ndnni s.n. Mount Fletcher?: Dood-

mans Krans Mountain, Ga/pin 6764.

Below the summit of the Drakensberg, between 6,500-8,500 feet, there occurs

what may prove to be a form of E. dominans. It is larger in all parts than typical

E. dominans. This taxon is represented in the National Herbarium by Killick 3525,

Esterhuysen 7939, 8828, 10187, 10197, 15471, 15556, 17360, 17364 and 17365. However,

careful field observations with particular reference to altitudinal variation are necessary

before its status can be definitely established.

Erica glaphyra Killick, sp. nov., E. dominanti Killick affinis, sed foliis patentibus’

inflorescentiis pseudo-spicatis vel interdum terminalibus, sepalis basi plerumque liberis,

ovario depresso-globoso distinguitur.

Fruticulus erectus vel suberectus, 15-50 cm altus; ramuli subvirgati, juvenes

puberuli, maturi glabri. Folia constanter 3-nata, patentia vel interdum suberecta

(sed petioli semper erecti), recta vel leviter incurvata, elliptica, 2- 1-3-7 mm longa,

0-5-0 -9 mm lata, supra plusminusve plana, infra convexa, sulcata, praeter margines

petiolosque glabra, olivaceo-viridia vel atro-viridia, nitida. Flores in pseudo-spicis ad

1 cm longis vel interdum in acervis cernuis terminalibus dispositi; pedicelli recurvati,

0-8-2 -5 mm longi; bracteae 2, 1 mm longae, puberulae. Sepala 4, cinnamomea vel

erubescentia vel rubra, basi libera vel interdum leviter imbricata, ovata, 11-1-2 mm

longa, 0-4-0 -6 mm lata (interiora saepe exterioribus angustiora et crassiora), concava,

dorsaliter carinata et in parte superiore sulcata, nonnihil cartilaginea, marginibus

hyalinis nonnunquam ciliatis. Corolla plerumque vinosa, 1- 5-2-0 mm longa, cyathi-

formis; tubus inter sepala ventricosus; lobi obtusi, 0-7-0 -8 mm longi. Stamina 8,

filamentis 0-8 mm longis, antheris 0-6-0 -8 mm longis basi setis (haud caudis) 0-6

minutis instructis. Ovarium depresso-globosum, glabrum; stylus crassus 0-7-1 -2 mm

longus, stigmate cyathiformi-peltato incluso vel leviter exserto. Fig. 1 : 3.

Type: Basutoland, Quachas, summit of Drakensberg in vicinity of Castle Buttress,

10,000 feet, 11 July 1952, Killick 1729 (PRE, holo.).

Erica glaphyra is less frequent than E. dominans and is rarely dominant in alpine

heath. In the Organ Pipes Pass area it occurs on south-facing slopes often in the shelter

of boulders at the edge of the escarpment. Miss E. Esterhuysen of the Bolus Herbarium,

who has made several collectings of this specie s in different parts of the Drakensberg

records E. glaphyra chiefly from south- and south-west-facing slopes. It would appear

to prefer the more mesic slopes — unlike E. dominans, which occurs on the drier north-

facing slopes and on horizontal portions of the summit. The flowering periods of

the two species are practically the same with E. glaphyra starting to flower a little later

than E. dominans. The epithet glaphyra refers to the polished or glossy appearance

of the plant. E. glaphyra has been recorded along the Drakensberg from Cathkin

Peak in the south to Mont aux Sources in the north between 8,000-10,000 feet.

Basutoland. — Quachas: Summit of Drakensberg, Castle Buttress, Killick 1729; 3520; 3541;

3542; 3543.

Natal. — Bergville: Cathedral Peak, Esterhuysen 15466; Cleft Peak, Esterhuysen 12891; Mitre,

Esterhuysen 14538; Mont aux Sources, Esterhuysen 15554; The Bell, Esterhuysen 14467. Estcourt:

Cathkin Peak, Esterhuysen 17359.

The two Erica species described above were first collected by the author in 1952,

while engaged on a botanical survey of the Cathedral Peak area. In the National

Herbarium the specimens collected were found to match a number of specimens

determined as forms of E. alticola Guth. & Bol., a Transvaal species described from

“ near Devil’s Kantoor ”. It was considered, however, that the Drakensberg material

could be divided into two distinct species differing from E. alticola. In order to reach

Table 1. — Main differences between Erica dominans , E. glaphyra and E. alticola

129

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a definite conclusion, the author in October 1964 collected 100 random samples of

the two species in the Organ Pipes Pass area with the object of applying some statistical

test of resemblance. The specimens, together with those available of E. alticola, were

examined and the resulting data discussed with a statistician. After careful consideration

it was decided that there was no point in applying statistical tests, because the data showed

so clearly the presence of three distinct entities. Some of the Drakensberg specimens

examined showed evidence of hybridization. A summary of the main differences

between E. dominans, E. glaphyra and E. alticola as revealed by the investigation is

given in Table 1. See also Fig. 1.

The possibility of two such closely related species occurring in the same area may,

at first sight, seem questionable: but, as already pointed out, they do occupy different

ecological habitats. This is probably another example of ecological isolation, where

“ closely related species are kept apart by the spatial pattern which results from their

differing ecological requirements ” (Cain, 1944). Hybridization will occur at the

ecological boundaries.

The precise position of E. dominans and E. glaphyra in the subgenus Euerica is

difficult to establish. Although obviously closely related, they possess inflorescences

which could refer them to two different sections. E. dominans with its usually terminal

inflorescence could be placed in the section Arsace, while E. glaphyra which usually

has a short pseudo-spike could be placed in Chlorocodon. The fact that both species

sometimes have the opposite inflorescence suggests that they occupy a position inter-

mediate between the two sections.

References

CAIN, S. A., 1944. Foundations of Plant Geography. Harper and Brothers: New York.

KILLICK, D. J. B., 1963. An Account of the Plant Ecology of the Cathedral Peak Area of the Natal

Drakensberg. Mem. Bot. Surv. S. Afr. 34.

D. J. B. Killick

GRAMINEAE

A New Combination in Hyparrhenia

Hyparrhenia pilosissima {Hack.) J. G. Anderson , comb. nov.

Andropogon pilosissimus Hack, in DC., Monogr. Phan. 6: 690 (1889). Type: Natal,

Drakensberg, Biggarsberg, Rehmann 7109 (W, holo.!). A. audits Stapf in FI. Cap.

7: 357 (1898). Syntypes: U itenhage, Ecklon & Zeyher 626 ; Queenstown, near Shiloh,

Baur 897; Thaba Nchu, Burke 427!; Zeyher 1799!; Griqualand East, grassy slopes

near Clydesdale, Tyson 3105; Natal, Rietvlei, Buchanan 222.

Hyparrhenia aucta (Stapf) Stent in Both. 1: 249 (1924); Chippindall in Gr. &

Pastures S. Afr. 1 : 514 (1955).

J. G. Anderson

POLEVANSIA De WINTER, GEN. NOV.

Polevansia De Winter, gen. nov. (Gramineae-Chlorideae) Willkommiae Hack, affinis,

sed lemmate fertili exaristato, lemmate paleaque subcoriaceis (haud membranaceis),

anthoecii callo brevi obtuso differt.

Spiculae a latere visae anguste oblongae, exaristatae, dorsaliter compressae, adaxi-

ales, solitariae. distichae, alternae, contiguae, oblique vel demum horizontaliter patentes;

rhachilla inter glumam superiorem et anthoecium disarticulans, supra anthoecium haud

producta. Anthoecium 1, hermaphroditum; callus brevissimus, obtusus, minute

pilosus. Glumae persistentes, inaequales; inferior circiter \ superiorem aequans,

131

explanata, lanceolata vel anguste ovata, obtusa, enervis, membranacea; superior

lemma aequans, explanata oblongo-lanceolata, obtusa, tenuiter coriacea, dorsaliter

rotundata, 1-nervis. Lemma spiculam aequans, dorsaliter plus minusve rotundatum,

marginibus incurvis, explanatum, lanceolatum vel anguste ovatum, obtusum, tenuiter

coriaceum, 3-nerve, nervibus lateralibus in triente superiore evanescentibus, nervo

medio in mucrone brevissimo excurrente, prope margines infra medium breviter

pubescens, ceterum glabrum. Palea lemma fere aequans, a dorso visa lanceolata,

membranacea, 2-nervis, nervibus ciliatis. Lodiculae 2, oblique cuneatae truncatae,

glabrae. Stamina 3. Ovarium glabrum; styli terminates, liberi; stigmata plumosa,

albida, e anthoeciis lateraliter exserta.

Gramen perenne humile, mattam rhizomatibus et stolonibus ramosis brevibus

formans; culmi graciles, erect i vel geniculati, simplices, paucinodosi; ligulae brevis-

simae, truncatae, inembranaceae, glabrae; laminae lineares, in apicem obtusum gradatim

attenuatae, breves, planae, rigidiusculae, marginibus cartilagineis; racemi plures, in

axi primario inflorescentae contractae orti.

Type species: P. rigida De Winter.

A monotypic genus known only from Basutoland.

This genus is named in honour of Dr. I. B. Pole Evans well known for his valuable

contributions to South African botany. The spelling Polevansia is preferred to the more

unwieldy alternative Pole-Evansia.

The position of Polevansia as a member of the tribe Chlorideae can be well

substantiated. Certain characters, however, such as the single floret which usually

is devoid of a prolongation of the rhachilla, and the adaxially positioned lower glume,

can be regarded as somewhat unusual for this tribe. These characters it has in common

with several other genera of the Chlorideae. Willkommia Hack., Microcldoa R. Br.

and Catalepis Stapf et Stent exhibit both one-flowered spikelets and adaxially placed

lower glumes, while Craspedorhachis is one-flowered but has the glumes laterally

appressed to the rhachis. The other characters of Polevansia are typical of the Chlori-

deae. A study of the leaf anatomy has supplied additional evidence confirming its

position in the tribe. The silicified cells of the epidermis are suborbicular to squarish

in shape and the alternate cells are deeply ripple-walled. The epidermal hairs are

two-celled and club-shaped and typical especially of the Chlorideae and Zoiseae (Fig. 3).

The cross-section of the leaf shows a radially arranged chlorenchyma, one layer of

cells thick. The motor cells, which are characteristic in shape, occupy at least half

the thickness of the leaf (Fig. 3). For comparison of its anatomy with that of other

genera in the Chlorideae, the figures given by Prat, in Ann. Sc. Nat. bot. Ser. 10, 18:

165-258 (1936) can be used.

Polevansia rigida De Winter, sp. nov. Gramen perenne, humile, infra 10 cm alta,

mattam rhizomatibus et stolonibus ramosis brevibus formans; culmi graciles, erecti

vel geniculati, simplices, paucinodosi; innovationes intra-vaginales, breves, dense

foliosi; vaginae leves. glabrae, marginibus tenuiter membranaceae; ligulae brevissimae,

truncatae, membranaceae; laminae lineares in apicem obtusam gradatim attenuatae,

breves, planae. lnflorescentia contracta, racemis spiciformis secundis breviter pedicel-

latis. Spiculae a latere visae angustae oblongae, exaristatae dorsaliter compressae

adaxiales, solitariae, alternatim distichae, contiguae. Anthoecium 1, hermaphroditum

callus obtusus, minute pilosus. Glumae persistentes, inaequales; inferior enervis;

superior l-nervis. Lemma spiculam aequans, 3-nerve, prope margines infra medium,

breviter pubescens, ceterum glabrum. Palea lemma fere aequans, a dorso visa lanceolata,

membranacea, inter carinas dorsaliter ciliata. Lodiculae 2. Stamina 3. Ovarium

glabrum. (Fig. 2).

132

Type: Basutoland, near top of Lekhalabatesi Valley, Pole Evans 12 (PRE,

holo.).

Perennial, about 10-0 cm high, matforming with a branched system of relatively

slender rhizomes; culms ascending or prostrate, densely sheathed, rooting at the

nodes; innovations arising from the underground rhizomes and the prostrate culms,

intravaginal, short, densely leafy, ascending to erect. Leaf-sheaths chartaceous, pallid,

rather loose, 1 -0-1 -5 cm long, striate, glabrous; margins thinly membranous, glabrous.

Auricles rounded, brownish, glabrous or scabrid. Collar glabrous. Ligule a short

membranous minutely fimbriate rim. Leaf-blade linear, 1 • 0-3 • 0 cm long, 1 • 5-2 • 0 mm

broad, flat, tapering to an obtuse apex, with a few scattered long hairs above, glabrous

below; nerves numerous, arranged closely together, raised below, slightly raised above,

midnerve more or less prominent below; margin narrow, cartilaginous, scabrid.

Inflorescence rigid, oblong, spikelike, consisting of a number of short, dense, appressed,

secund racemes, arranged alternately on the axis, 2 -0-3-0 cm long and 0-8-1 -5 mm

wide ; rhachis and branches triangular in cross-section, scabrid on the angles, otherwise

glabrous. Spikelets 3 • 5^4 - 5 mm long, solitary, dorsally compressed, pedunculate,

alternate, lower glume always adaxial to the rhachis; peduncles 0-5-1 -5 mm long,

triangular in cross-section, scabrid on the angles; rhachilla disarticulating above the

glumes, not produced beyond the base of the floret. Floret solitary, hermaphrodite;

callus short, obtuse, shortly hairy. Glumes persistent, unequal; lower 1-5-2 -5 mm

long, lanceolate to narrowly ovate, subhyaline, subcoriaceous, nerveless, apex obtuse

truncate or emarginate ; upper 3 • 5-4 • 5 mm long, lanceolate (when flattened), greenish,

subcoriaceous, one-nerved, slightly keeled towards the apex, scaberulous on the keels

Fig. 2. — Polevansia rigida. A, Side view of spikelet showing adaxial lower glume, X 10; B, lower

glume, X 10; C, upper glume, x 10; D, lemma, X 10; E, palea, X 10; F, lodicules,

X 10; G, stamens and gynoecium, X 10 ( Pole Evans 12).

133

Fig. 3. — Polevansia rigida: A, cross section of the leaf; B, abaxial epidermis of the leaf: C, diagram

of the cross section of a leaf. AB, abaxial epidermis; AD, adaxial epidermis; BH, two-

celled hair; C, chlorenchyma : LC, long ripple- walled cell; MC, motor cell; SC, silicified

cell; S, mechanical tissue or stereome; ST, stoma: 1 VB, first order vascular bundle; 3VB,

third order vascular bundle ( Pole Evans 12).

134

and upper part of midnerve, apex obtuse. Lemma 3 -5-4 -5 mm long, narrowly

lanceolate-oblong (when flattened), 3-nerved, midnerve excurrent into a very small

mucro, lateral nerves evanescent (occasionally a few short inconspicuous nerves present

at the base between the midnerve and lateral nerves) ; lateral nerves hairy in the lower

half. Palea 3 -5-4-0 mm long, narrowly oblong-elliptic, two-keeled, subhyaline, sub-

coriaceous; apex truncate; margins hairy in the lower half. Lodicules 2, free, small,

truncate, 0-25 mm long. Stamens 3, anthers cultrate. Ovary glabrous, oblong, styles

distinct, plumose, Caryopsis unknown. Fig. 2.

This species has been collected only in two different valleys in the Quachas District

of Basutoland. It is reported to be a pioneer on old lands by Pole Evans. After the

discovery of the new genus by Pole Evans, a special search was undertaken by Killick,

who found it only once, growing on a rocky outcrop. Killick found Catalepis gracilis

in large quantities as a pioneer of abandoned lands. Since Polevansia resembles Catalepis

in some respects the latter may have been mistaken for Polevansia. The ecology of

the species should therefore be more thoroughly investigated.

Basutoland. — Near top of Lekhalabatesi Valley, colonizing worn out native lands, 17.2.1950,

alt. 8,500 ft, Pole Evans 12; Mokhotlong River Valley, abundant on rock outcrop, 1.5.1953, alt.

8,000 ft, Killick 1983.

B. de Winter

Styppeiochloa De Winter, gen. nov.

Styppeiochloa De Winter, gen. nov. Danthoniae DC. affinis, sed lemmatibus

breviter trifidus nervis 3 prominentibus etiam plerumque basi nervis 1-4 brevibus,

hilo lineari f- caryopsin subaequanti differt.

Spiculae breviter aristatae anguste elliptico-oblongae vel elliptico-oblongae, latera-

liter compressae, solitariae, ramis paniculae sparse ramosis, appressis. Rhachilla

supra glumas et inter anthoecia disarticulans. Anthoecia 2-5, hermaphrodita vel

supremum sterile vel ad lemma redactum, callus brevis truncatus, dense pilosus.

Glumae persistentes, subaequales vel inaequales, lanceolatae, 1-3-nervatae (raro 5-ner-

vatae nervis lateralibus tenuissimis), arista breve vel nulla, margine interdum mucrone

singulari vel duobus. Lemmata apice trifida, basin versus marginibus et dorso pilosa,

lanceolata 3-5-nervata (vel 7-nervata, nervis lateralibus tenuissimis) nervo medio in

aristam scabram rigidam brevem excurrenti, utrinque nervo singulari in mucronem

vel in aristam brevem excurrenti. Paleae anguste lanceolatae, lemmatibus longiores

vel breviores, inter Carinas duas basin versus paucipilosae, spica bifida, carinis scabris.

Lodiculae 2, connatae vel liberae, parvae truncatae interdum mucronatae, 2-5-nervatae.

Stamina 3, flavida vel purpurea. Ovarium ellipitico-oblongum glabrum, appendice

nodosa inter bases stylorum, stylis distinctis, stigmatibus plumosis. Caryopsis elliptica,

embryone quarta parte grani longitudinis, hilo lineari tribus quartibus grani longi-

tidinis.

Gramen perenne dense caespitosum, basi dura fibrosa pilosa, innovationibus

intravaginalibus, culmis erectis gracilibus glabris paucinodosis, foliis convolutolinearibus,

paniculis sparse ramosis, ramis singulis vel binis laxis vel contractis.

Type species: Styppeiochloa gynoglossa (Goossens) De Winter.

A monotypic genus from Southern Africa.

135

C

Fig. 4. — Styppeiochloa gynoglossa: A, cross section of the leaf; B, abaxial epidermis; C, diagram

of the cross section of a leaf. AB, abaxial epidermis; AD, adaxial epidermis; BH, two-

celled hair; CC, suberized cell; C, chlorenchyma; LC, long ripple-walled cell; MC, motor

cells: OU, one-celled retrorse hair; SC, mechanical tissue or stereome; S, silicified cell;

1 VB, first order vascular bundle: 3 VB, third order vascular bundle (Killick 1013).

7230691-5

136

Spikelets shortly awned, narrowly elliptic-oblong to elliptic-oblong, laterally

compressed, appressed to the contracted branches of the panicle. Rhachilla disarti-

culating above the glumes and between the florets. Florets 2-5, hermaphrodite,

uppermost floret sterile or reduced to the lemma. Callus short, truncate, densely

pilose. Glumes persistent, subequal to unequal, lanceolate, 1-3-nerved (rarely 5-nerved

with the lateral nerves short) apex trilobed or lobes absent and apex acute. Lemma

with a tri-lobed apex, lanceolate, 3-5-nerved (rarely 7-nerved with two lateral nerves

very short), median nerve and one lateral nerve on each side of the median excurrent

into a mucro or short awn. Palea narrowly lanceolate, subequal to the lemma.

Lodicules 2, small, truncate, 2-5-nerved. Stamens 3, yellow or purplish. Ovary elliptic-

oblong, glabrous, with the style bases knob-like. Caryopsis elliptic; embryo \ the

length of the grain; hilum linear, J the length of the grain.

Dense sclerophyllous tufts with hard fibrous basal sheaths forming a thick dense

mat.

The name of the genus is a combination of the Greek words “ stuppeion ” (fibre)

and “ chloa ” (grass), an allusion to the tough fibrous base of the type species.

Styppeiocliloa gynoglossa, the only species of the new genus, was first described

as Crinipes gynoglossa by Goossens in Kew Bull. 1934: 200 (1934), although at the time

Goossens thought that it might represent an undescribed genus. Specimens were sent

to Kew for an opinion on the generic affinities and, since it was similar in several

respects to the genus Crinipes Hochst., which was then only known from a few collec-

tions, and because the material was inadequate for a complete study. Professor Goossens

was advised to include it in Crinipes. Since then much more material of true Crinipes

and of species which are related to it, but could hardly be included in Crinipes, has

come to light. For this reason a re-assesment of generic delimitations in this group

was undertaken by Hubbard with the result that the South African species was excluded

from Crinipes and is here described as a distinct genus.

An investigation of the leaf anatomy of Styppeiocliloa gynoglossa has shown that

the genus does not belong in the Eragrosteae (in which, as a member of the genus

Crinipes , it had been placed) but is more closely allied to Danthonia and its allies in the

Danthoniae. The silicified cells are dumb-bell-shaped (Fig. 4) and quite different

from the typical silicified cells of Eragrostis and its allies, which are more or less round

or square in shape and usually have accompanying cells associated with them. Moreover

the chlorenchyma is not radially arranged round the vascular bundles in a single row

of cells, as in the Eragrosteae. The leaf is of the “ Danthonoid ” types where the

chlorenchyma is more or less continuously distributed between the bundles, the motor

cells being comparatively small and not forming divisions between the vascular bundles

as in the “ Eragrostoid ” type (Fig. 4). Bicellular hairs similar to those of Danthonia

are also present (Fig. 4). It has even proved somewhat difficult to find good characters

to distinguish this species from members of Danthonia. The very reduced awns and

the rather long linear hilum together with the different facies distinguishes this genus

from typical Danthonia. The hilum in all the members of Danthonia investigated

proved to be rather short, usually narrowly elliptic in outline, or not very well defined.

None of the species investigated had a distinctly linear hilum such as possessed by

Styppeiocliloa.

Styppeiochloa gynoglossa ( Goossens ) De Winter, comb. nov.

Crinipes gynoglossa Goossens in Kew Bull. 1934: 200(1934); Chippindall in Grasses

and Pastures of S. Afr. 1 : 123 (1955). Type: Transvaal, Barberton District, Kaapsche

Hoop, Pole Evans 1022 (PRE, holo.!; K).

S. gynoglossa occurs all along the Drakensberg escarpment from the Impendhle

district in Natal northwards through Swaziland to the Eastern Transvaal and the

eastern districts of Rhodesia. At Namaacha on the border of Swaziland it is found

within the territory of Mozambique. It prefers seepage areas where the rock is overlain

137

by a thin layer ot soil. The extremely tough fibrous bases of these plants form a thick

mat so resistant that a pickhandle is easily broken in trying to lift it. Some of the stands

are several acres in extent. It is found mainly at high altitudes usually between

5,000-6,000 ft. above sea level. At Millers Falls in Swaziland and the nearby Namaacha

in Mozambique the altitudes are 4,500 and 2,000 ft. above sea level respectively. The

rainfall in the greater part of its range exceeds 880 mm p.a.

SOUTH AFRICA. Transvaal.— Ermelo: Pont s.n. (K). Barbe ton: Kaapsche Hoop, Pole

Evans 1022. Lydenburg: Steenkampsberg, 29 miles W. of Kydenburg, shallow moist seepage area

over quartzite slabs, Codd 8219. Pietersburg: Blaauberg on way to Trig. Beacon, Codd & Dyer 9071.

Pilgrim's Rest: 7 miles from Graskop on road to Vaalbank, De Winter & Codd 220 \ Mariepskop,

plateau near beacon. Van der Schijff 6344; Driekop Gorge near Graskop, De Winter & Codd 211.

Natal.— Bergville: Cathedral Peak Forest Research Station, Killick 1013; 1237; 1248; 1548;

1565; 1789. Impendhle: along streambank, Huntlev 481 ; 798. Utrecht: farm Naauwhoek, Devenish

935; 172.

SWAZILAND. Mbabane: Millers Falls, Compton 26188.

MOZAMBIQUE. Sul do Save. — Lourenco Marques: Namaacha, Myre & Carvalho 300; 1312.

RHODESIA. Melsetter: Chimanimani, rocks west of airfield, Goodier & Phipps 20.

S. gynoglossa is extremely closely allied to the Madagascan Redfieldia hitchcockii

A. Camus and the latter may on further investigation prove to belong to the genus

Styppeiochloa. It certainly does not belong to the monotypic American genus Redfieldia

Vasey, from which it differs in several important points.

B. de Winter

A New Species of Eragrostis

Eragrostis moggii De Winter , sp. nov., E. glandulosipedatae De Winter affinis, sed

spiculis minoribus et angustioribus lemmatium nervis conspicuis differt.

Gramen perenne, decumbens vel suberectum (rare erectum). Culmi geniculati,

3-multinodosi, multiramosi, saepe fasciculati. ad 90 cm longi, glabri, saepe purpurascenti.

Vaginae laxae, glabrae, marginibus pilosis, carinis costisque glanduloso-punctatis.

Ligula dense ciliata. Laminae planae vel involutae, infra glabrae supra minute scabridae.

Panicula angusta, plus minusve contracta vel patens, ramis solitariis vel binis. Spiculae

plumbeae, distincte pedicellatae, 4-11-florae, 3-7 mm longae, 1-1 -5 mm latae; pedicelli

glande annulari instructi. Glumae subequales vel nonnihil inaequales purpurascentes,

ovales, apice obtusae vel subtruncatae, 1-nervatae, carinis scabridiusculis. Lemmata

subcoriacea, nitida. late ovata, obtusa, ca 1 - 5 mm longa, 1 0 mm lata, nervis 3 con-

spicuis. Paleae anguste ovato-ellipticae, 1-3-1 -4 mm longae, bicarinatae, truncatae,

membranaceae. Lodiculae 2, carnosae, cuneiformes. Stamina 3, antheris anguste

oblongis ca 0-7 mm longis. Ovarium oblongum, stylis 2. Caryopsis nonnihil lateraliter

compressa, apice et basi subtruncata, opaca, nonnihil rugosa, ca 6 cm longa.

Type: Mocambique, Inhaca Island, Mogg 26816 (PRE, holo.; K, SRGH).

Perennial up to 90 cm high (usually 30-40 cm), decumbent, geniculate-ascending,

or erect. Culms somewhat branched to much-branched and occasionally fascicled

from the nodes, rarely prostrate and rooting from the nodes which bear dense fascicles

of culms, few- to many-noded, internodes glabrous and smooth, flushed with purple

or pallid, easily compressed. Leaf-sheaths usually shorter than the internodes, lax

and slipping from the culms leaving them bare, glabrous except for the ciliate margins,

somewhat striate, with glands on the ribs upwards, often purplish. Collar conspicuous,

yellowish, glabrous or hairy on the back, the auricles conspicuously bearded. Ligule

long ciliate. Lecf-hlades expanded or rolled, up to 3 mm wide, up to 20 cm long,

138

tapering to a fine point, glabrous and smooth below, finely ribbed above with the ribs

minutely scaberulous. Panicle narrow and contracted to effuse and open, branches

single or paired, stiff, glabrous, finely ribbed. Pedicels of the spikelets slender, stiff,

furnished with a distinct annular gland. Spikelets lead-coloured (rarely pallid), narrowly

oblong, 4-7 mm long and 1 -0-1 • 5 mm wide, 4-13-flowered, rhachilla not disarticulating

the lemmas deciduous, the paleas remaining attached to the rhachilla. Florets herma-

phrodite. Glumes 1 -nerved, subequal or somewhat unequal, broadly ovate-oblong,

obtuse, finely scaberulous on the keels, subcoriaceous, usually distinctly purplish.

Lemmas about 1 -5 mm long and about 1 -0 mm wide, broadly elliptic-ovate in outline,

very finely scabridulous, 3-nerved, nerves prominent, apex obtuse to subtruncate.

Palea equalling the lemma or slightly shorter, membranous, subhyaline, bicarinate,

margins inflexed, keels scaberulous. Lodicules truncate, cuneate, somewhat fleshy,

small. Stamens 3; anthers about 0- 7 mm long, cultrate, pale yellow. Ovary glabrous;

styles free; stigmas 2, plumose. Caryopsis somewhat laterally compressed, apex and

base subtruncate giving it a nearly square outline seen in side view, smooth, whitish-

opaque; embryo half to slightly less than half the length of the grain, brownish; hilum

punctiform, basal.

Two varieties can be distinguished.

Inflorescence effuse and open, the branches and branchlets spreading var. moggii

Inflorescence narrow and contracted the branches appressed to the rhachis var contracta

var. moggii.

Inflorescence effuse and open, the branches and branchlets spreading.

Occurs in Mozambique particularly in the vicinity of Lourenco Marques and

islands off the coast. It has also been collected in northern Zululand. It prefers sandy

soils and often occurs in open forest.

SOUTH AFRICA. Natal, — Ubombo: Mtenzi Game Reserve, Tinley 596.

MOZAMBIQUE. Sul do Save. — Lourenco Marques: Costa da Sol, Balsinhas 261; Myre 864; Inhaca

Island, Mogg 27205; 26816; 29354; 26870 (pro parte); 26746 (J); Gillard J. 34664 (J); Weintraub

J. 20469 (J); Mauve & Verdoorn 103; Bazaruto Island, Mogg 28512 (J); Masinga, Schlechter 12127.

Maputo. — Catuane, Myre & Balsinhas 585.

Inhambane. — Inharrime, De Lomos & Balsinhas 1 74 ; Velho, Gomes e Sousa 2136.

var. contracta De Winter, var. nov., inflorescentia angusta contracta, rami ramulique

rhachi adpressi.

Type: Mozambique, Inhaca Island, Mogg 26947 (PRE, holo.; K).

This variety has been recorded from the vicinity of Lourenco Marques and has

the same ecological preferences as the typical variety.

MOZAMBIQUE. Sul do Save. — Lourenco Marques: between Lourenco Marques and Costa da Sol,

Schweickerdt 1910; Inhaca Island, Mogg 26870 (pro parte); 26947; 26870; 29362; 27571; 30069

(J); 30119 (J).

The two varieties often occur together and one collection, Mogg 26870, consisted

of specimens bearing either narrow and contracted or divaricate open inflorescences.

So far the species has been collected mainly in the surroundings of Lourenco Marques.

The Zululand specimen is the only record from South Africa and shows that the species

is probably much more widely spread than the present records indicate.

This species seems to be allied to E. glandulosipedata de Winter, with which it

agrees in the annular glands on the pedicels, the glands on the leaf-sheaths and the

general configuration of the spikelets. It is, however, smaller in all the parts of the

139

spikelet and very easily distinguished. E. moggii could be confused with E. lehmanniana

which it resembles generally in habit but the latter has no glands on the pedicels and

the nerves of the lemmas are not as prominent nor are the glumes purplish.

LILIACEAE

B. de Winter

New Species of Kniphofia

Names are required for three undescribed species of Kniphofia which are included

in check-lists being compiled for the territories of Swaziland by Prof. R. H. Compton

and of Basutoland by Mrs. A. Jacot Guillarmod, respectively. The three species are

described here, together with a fourth species which was recently collected in the

Mountain Zebra National Park, Cradock District, Cape Province.

K. ritualis Code! , sp. nov., K. sarmentosae (Ait.) Kunth affinis, sed foliis marginibus

scabridulis, floribus longioribus differt.

K. sarmentosa sensu Phillips in Ann. S. Afr. Mus. 16: 294 (1917).

Planta 40-80 cm alta. Folia carinata, glauca, 40-90 cm longa, 1 • 2-2 • 4 cm lata,

margine conspicue serrulata. Inflorescentia densa, rhomboideo-cylindrica vel ovoidea,

9-14 cm longa, 4-5 cm lata, gemmis pendulis corallinis ve! rubris, floribus maturis

flavo-virentibus. Bracteae lineari-lanceolatae, acutae vel acuminatae, 10-12 mm

longae, 2 mm latae, minute denticulatae. Pedicelli 1-2 mm longi. Perianthium

subcylindricum, 2-5-3 cm longurn. Stamina inclusa vel vix exserta. Stylus 3 mm

exsertus. Capsu/a subglobosa, 7-8 mm longa. Plate 2.

Type: Basutoland, Berea District, Bitsolebe area, alt. 6,500 ft, 22/1/1957, Marais

1327 (PRE, holo.).

Plants solitary or in small groups. Leaves in 3 or 4 ranks, 8-12 per peduncle,

at first erect, later bending at the middle, soft in texture, glaucous, 40-90 cm long and

1-2-2 -4 cm broad near the base, deeply keeled; margin conspicuously and regularly

serrulate. Peduncle subequal or shorter than the leaves, 40-80 cm long. Inflorescence

rhomboid-cylindrical to ovoid, dense, 9-14 cm long and 4-5 cm in diameter; buds

and flowers pendulous; buds coral or salmon to orange-red, becoming greenish-

yellow as the flowers open. Bracts lanceolate to linear-lanceolate, tapering to the

acute or acuminate apex, 10-12 mm long and 2 mm broad; margin usually minutely

denticulate. Pedicels 1-2 mm long at flowering stage, elongating to 5 mm in the

fruiting stage. Perianth subcylindrical, 2-5-3 cm long, very slightly constricted above

the ovary, 3 mm wide at the base, increasing gradually to 5 mm wide at the mouth;

lobes ovate, rounded, 3-3-5 mm long, not spreading. Stamens 6, of two lengths;

anthers not or scarcely exserted at anthesis, later withdrawn into the perianth tube.

Style equal to the stamens at anthesis, later included or exserted by up to 3 mm.

Capsule subglobose, not markedly triquetrous, 7-8 mm long.

Common in Basutoland and the adjoining areas of the eastern Orange Free State,

mainly on sandstone formation, in shallow soil in rock crevices, in dry water courses

and on grassy slopes, at altitudes of 5,500 to 9,000 ft. The flowering time is from

January to March.

Basutoland. — Mafeteng: Malealea, Munro s.n. (cultivated in Pretoria). Maseru: Thaba Bosigo,

Van der Merwe 1156; Jordan Valley, Guillarmod 1671; Senqunyane Valley, Guillarmod 2256. Berea:

below Molimo-Nthuse, Marais 1326; Bitsolebe area, grassy slope among shrubs, Marais 1327 (PRE,

holo.); at foot of Pulane Mt., Bruce 360; Pulane area, Bruce 361 ; 362; near Mateka, Bruce 364;

Thaba Chitja, Bruce 365; Mamathes, Bruce 390. Leribe : Dieterlen 174; 174a.

Orange Free State. — Ficksburg: farm Wintershoek, near Kirklington, Codd 10540; 10541; farm

Franschhoek, 20 miles N.E. of Ficksburg, Codd 10549. Fouriesburg: 23 miles N.E. of Ficksburg,

Bruce 343; 12 miles S. of Clarens, Bruce 393; near Golden Gate, 11 miles E. of Clarens, Codd 10537.

Senekal: Goossens 838. Thabanchu: Thabanchu Mt., Roberts s.n.

140

Phillips, l.c., misidentified this species as K. sarmentosa (Ait.) Kunth, a species

largely restricted to the Roggeveld area of the Cape Province. He records the Sotho

common names as “ Leloele or Lelole ” and states that: “A decoction prepared from

this species is drunk to cure pains in the shoulders. Women, during the times girls

are undergoing initiation rites, make use of this plant for some purpose, but this being

one of the sacred rites of the tribe, no further information could be obtained

K. sarmentosa and K. ritua/is are similar in having conspicuously long, narrow

bracts. The latter species can, however, readily be distinguished by the markedly

serrulate leaves and the longer perianth, 2-5-3 cm long, as against 2-2-5 cm long

in K. sarmentosa.

K. ritualis also shows an affinity to K. caulescens, but there are several characters

distinguishing them. K. caulescens is one of the few caulescent species in the genus

and is gregarious, forming large groups. With its markedly glaucous leaves and

attractive inflorescences grading from coral or red buds to lemon-coloured floweis, a

group in flower is an impressive sight. The perianth (2 -2-2 -4 cm) is distinctly shorter

than in K. ritualis , while the stamens are exserted by 8-13 mm at anthesis. It is

interesting to note that the natives regard K. caulescens as a charm against lightning

and frequently plant the species near their huts. Phillips, l.c., who misidentified this

species as K. tysonii Bak., also records that the natives bathe themselves with a decoction

from the plant in order to protect themselves against lightning.

K. hirsuta Codd , sp. nov., K. ritualis Codd affinis, sed foliis hirsutis differt.

Planta 45-60 cm alta. Folia carinata, erecta, 40-60 cm longa, 1-5-2 -5 cm lata,

hirsuta, margine conspicue serrulata. lnflorescentia densa, rhomboideo-cylindrica vel

ovoidea, 8-12 cm longa, 4-4-5 cm lata, gemmis pendulis corallinis vel rubris, floribus

maturis flavo-virentibus. Bracteae lineari-lanceolatae, acuminatae, 10-14 mm longae,

2 mm latae, minute denticulatae vel integrae. Pedicelli 2-3 mm longi. Perianthium

subcylindricum, 2 -2-2 -7 cm longum. Stamina inclusa vel vix exserta. Stylus 3 mm

exsertus. Capsula subglobosa, 7-9 mm longa. Plate 3.

Type: Basutoland. Berea District, Mamalapi, alt. 8,500 ft, 20/1/1957, Guillarmod

& Marais 1307 (PRE, holo.).

Plants usually solitary. Leaves in 3 or, rarely, 4 ranks, 8-12 per peduncle, eiect,

soft in texture, dull green, 40-60 cm long and 1 -5-2-5 cm broad near the base, deeply

keeled., hirsute along the nerves on both surfaces; margin conspicuously and regularly

serrulate. Peduncle equal to or longer than the leaves, 45-60 cm long. Inflorescence

shortly rhomboid-cylindrical to ovoid, dense, 8-12 cm long and 4-4-5 cm in diameter;

buds and flowers pendulous; buds orange or dull coral to salmon-pink tipped green,

becoming dull greenish-yellow as the flowers open. Bracts linear-lanceolate, tapering

to the acuminate apex, 10-14 mm long and 2 mm broad; margin minutely denticulate

to entiie. Pedicels 2-3 mm long at flowering stage, elongating to 4 mm in fruit. Perianth

subcylindrical, 2 -2-2 -7 cm long, very slightly constricted above the ovary, 3-5 mm wide

at the base, increasing gradually to 5 mm wide at the mouth; lobes ovate, rounded,

2-5 mm long, not spreading. Stamens 6, of two lengths, not or scarcely exserted at

anthesis, later withdrawn into the perianth tube. Style equal to the stamens at anthesis,

later exserted by up to 3 mm. Capsule subglobose, not markedly triquetrous, 7-9

mm long.

Found in the mountains of Basutoland on grassy slopes and stream banks, at

altitudes of 8,000 to 9,000 ft. The flowering takes place in December and January.

Basutoland. — Berea District, Mamalapi, in grassveld near rocks, alt. 8,500 ft, 20/1/1957, Guillarmod

& Marais 1307 (PRE, holo.); Guillarmod & Marais 1308; Mamalapi, December 1948, Compton 21266.

141

K. hirsuta is closely allied to K. ritua/is, having similar inflorescence, perianth

and bract characters, but the former species is unique in the genus in having pubescent

leaves. The only other species which approaches it in this respect is K. stricta Codd,

which sometimes has a few scabrid hairs on the nerves. K. hirsuta was apparently

discovered in December 1948 by Prof. R. H. Compton at Mamalapi at an altitude

of about 8,500 ft. Since then, good flowering and fruiting material was collected at

the same locality by Mrs. A. Jacot Guillarmod and Mr. W. Marais, who report that

it tends to flower earlier than K. ritua/is and to grow at higher altitudes (up to 9,000 ft),

where it grows in mountain grassland, associated with Danthonia-Erica-Cliffortia

scrub of rocky slopes or streams.

K. umbrina Codd, sp. nov., K. typhoidi Codd affinis, sed foliis conspicue carinatis,

floribus et bracteis longioribus diflert.

Planta 70-90 cm alta. Folia carinata, erecta, apice reflexa, flaccida, 45-70 cm

longa, 1-2 cm lata, margine carinaque levi. Inflorescentia densa, subcylindrica, 7-15

cm longa, 1-5-2 cm lata, gemmis patulis, floribus maturis primum patulis demum

pendulis, umbrinis vix aromaticis. Bracteae lineari-lanceolatae, acuminatae, 9-1 1 mm

longae, 1-5 mm latae, integrae. PediceUi 1-5 mm longi. Perianthium cylindricum,

8-9 mm longum. Stamina 2 mm exserta. Stylus 3 mm exsertus. Capsula ovoidea-

triquetra, 5-6 mm longa. Plate 4.

Type: Swaziland, 4 miles S. of Forbes Reef, 20/2/1951, Bruce 272 (PRE, holo.).

Plants usually in small groups. Leaves not distichous, 6-8 per peduncle, at first

erect, later reflexed, mid-green, somewhat flaccid, 45-70 cm long and 1-2 cm broad,

tapering towards a rather blunt apex, V-shaped in cross section, breaking up into a

few persistent fibres at the base; margin and keel smooth. Peduncle overtopping

the leaves, 70-90 cm tall. Inflorescence subcylindrical, tapering slightly towaids the

apex, dense, 7-15 cm long and 1 -5-2 cm in diameter; buds spreading, flowers eventually

pendulous; buds purplish-brown, flowers reddish-brown to dull red, slightly scented,

drying blackish. Bracts linear-lanceolate, acuminate, 9-11 mm long and 1-5 mm

broad; margin entire. Pedicels 0-5 mm long. Perianth cylindrical, not constricted

above the ovary, 8-9 mm long, 1 -5 mm wide at the base, increasing to 2-5 mm at the

mouth; lobes ovate, 1 mm long, not spreading. Stamens 6, of two lengths, exserted

by up to 2 mm at anthesis. Style exserted by 1-2 mm at anthesis and eventually by

3 mm. Capsule ovoid-triquetrous, 5-6 mm long.

Known from only a small area a few miles south of Forbes Reef in Swaziland,

at an altitude of about 4,500 ft, where it grows in or near grassy vleis. The flowering

time is Februaiy to early March.

Swaziland. — Four miles S. of Forbes Reef. 20/2/1951, Bruce 272 (PRE, holo.); 6/3/56, Codd 9521 ;

12 miles N. of Mbabane on Forbes Reef road, 19/2/51, Reynolds 5866; near Forbes Reef, 21/2/58,

Compton 27552; 8/2/62, Compton 31290.

With its reddish-brown, slightly scented flowers, this species shows an obvious

relationship to K. tvphoides Codd (Notosceptrum natalense Bak.), but there are several

clear distinguishing characters. The leaves are not distichous or glaucous, as in K.

typhoides, and they are distinctly keeled. In addition, the perianth is longer (4 -5-6 -5

mm long in K. typhoides ) and more pendulous, while the floral bracts are longer (4-6

mm long in K. typhoides) and are acuminate at the apex, not rounded as in K. typhoides.

In view of the restricted distribution of this species and the growing pressure of

agricultural activity, it is desirable that adequate steps be taken to ensure that it is

protected from eradication.

142

K. acraea Codd, sp. nov., K. breviflorae Bak. affinis, sed inflorescentiis densioribus:

bracteis latis ovatis differt.

Planta 50-70 cm alta. Folia nervosa, carinata, 50-70 cm longa, 1-1-5 cm lata,

margine minute denticulata. Inflorescentia densa, ovoidea vel subcylindrica, 6-12 cm

longa, 2 -2-2 -5 cm lata, gemmis patulis viridi-luteis, floribus maturis primum patulis

demum pendulis luteis. Bracteae latae ovatae, obtusae, 4-5 mm longae, 2-2-5 mm

latae, minute eroso-denticulatae. Pedicelli 1-5-2 mm longi. Perianthium breviter

cylindricum, 8-9 mm longum, 2 mm latum. Stamina 2-3 mm exserta. Stylus 3-4 mm

exsertus. Fructus non visi. Plate 5.

Type: Cape Province, Cradock District, National Mountain Zebra Park, on top

of Bankberg, alt. about 6,000 ft, 6/3/1964, Liebenberg 7120 (PRE, holo.).

Plants in small groups. Leaves about 12 per peduncle, 50-70 cm long and 1-1 -5

cm broad, folded along the midrib, nerves conspicuous; margin minutely and somewhat

distantly denticulate, keel smooth. Peduncle subequal to the leaves, 50-60 cm long.

Inflorescence dense, ovoid to subcylindrical, tapering to the apex in the young stage,

6-12 cm long and 2-2-2- 5 cm in diameter; buds greenish-yellow, spreading; mature

flowers yellow, at first spreading, later deflexed. Bracts broadly ovate, obtuse, 4-5 mm

long and 2-2-5 mm broad, chartaceous, erect and overlapping the buds in the young

stage, later reflexed; margin minutely erose-denticulate. Pedicels 1-5-2 mm long.

Perianth shortly cylindrical, not constricted above the ovary, 8-9 mm long, about 2 mm

wide at the base, widening slightly to 2-5-3 mm wide at the mouth; lobes bioadly

ovate to oblong, rounded to acute, 1 • 5-2 mm long. Stamens exserted by 2-3 mm at

anthesis. Style equal to the stamens at anthesis, eventually exserted by 3-4 mm.

Fruit not seen.

Known only from the above gathering, consisting of 3 specimens, collected on

top of the Bankberg, in a seepage area among rocks. It is a very distinctive species,

resembling K. breviflora Harv. ex Bak. in the length and colour of the flowers, but

its real relationship is not clear.

L. E. Codd

PODOCARPACEAE

The Typification of Podocarpus elongatus

The taxon was first described by Solander in Aiton’s Hortus Kewensis ed. 1, 3:

415 (1789) as Taxus elongata. It is mentioned that the plant is a native of the Cape

of Good Hope and that it was introduced in 1774 by Thomas Lucas.

Two sheets at the British Museum (Natural History) were annotated by Solander

as Taxus elongata. The first was collected by Masson at the Cape but no exact locality

or date aie indicated. As Masson undertook his last collecting trip at the Cape during

1774 the specimen could not have been gathered later than that year. Mounted on

the sheet are two branches with male cones and both represent the Cape plant.

On the second sheet three specimens are mounted, all bearing male cones. One

of them is annotated by Solander: “ Hort. Dni. Lucas Lee in Kent 1777 ”. (“ Dni.”

stands for “ Domini ” and Lee is the village in Kent in which the Lucas garden was

situated). The two other specimens are labelled in a different hand.: “ Hort. Dni.

Salisbury 1785 ” and “Hort. Dni. Vere in Kensington Gore 1788 ”. These three

specimens are referable to Podocarpus macrophyllus (Thunb.) Don subsp. maki Sieb.,

an Asiatic species which closely resembles P. elongatus.

Stapf in Flora Capensis 5, 2 (Suppl.): 8 (1933) cites (among others) the Masson

specimen and the specimens from the gardens of Lucas, Salisbury and Yere as P.

elongatus. In a typewritten report on the South African species of Widdringtonia

and Podocarpus (27/5/1921), he expresses the opinion that the specimen from the garden

of Lucas in the BM is the one that Solander described in the Hortus Kewensis.

143

It appears, however, that Solander included a mixture in his concept and there

is no conclusive evidence that his description of Taxus elongata is based solely or even

mainly on the Lucas specimen. The plant mentioned in the Hortus Kewensis was

introduced in 1774 whilst the “ Hort. Lucas ” specimen was collected in his garden

in 1777. As most plants were brought to England as seed or cuttings it is unlikely

that a specimen of Podocarpus which was introduced in 1774 could have borne male

cones in 1777.

The Hortus Kewensis refers to a plant in the Lucas garden which is a native of

the Cape of Good Hope, while the “ Hort. Lucas ” specimen is undoubtedly of Asiatic

origin. It thus appears that Lucas had both species in his garden. The source of the

Asiatic species is not known but the date 1774 given for the introduction of the Cape

species points to Masson who sent large botanical collections from the Cape to England

during that year. Salisbury mentions in a note in Burchell's Memoranda Botanica

(MS. in Kew Library) that he had a specimen of “ Taxus elongata ” growing in his

garden which Masson had collected at the Cape.

Gray (Journ. Arn. Arb. 34: 166, 1953) pointed out that the specimens from the

gardens of Lucas, Salisbury and Vere in the BM do not represent the Cape species

and argued that the Lucas specimen can thus not be taken as the type of Podocarpus

elongatus. She proposed two “ lectotypes ” : as male type she chose the Thunberg

specimen in UPS and as female type the Sonnerat specimen in P. Obviously this

treatment cannot be accepted. Apart from the fact that a name cannot be typified

by more than one specimen, neither of the specimens chosen by Gray beat s Solander’s

handwriting and it is unlikely that he saw either of them. Instead, the Masson specimen

in the BM is proposed as lectotype of the name Podocarpus elongatus. The specimen

was identified by Solander and although no specimens are explicitly mentioned in

the Hortus Kewensis there is every reason to assume that the author saw it before

publishing the description. The only alternative lectotype is one of the specimens

from the gardens of Lucas, Salisbury or Vere and this would involve transference of

the name P. elongatus to the Asiatic species presently known as P. macrophyllus subsp.

maki whilst the Cape plant would require a new name.

I am grateful to Dr. L. Codd of the Botanical Research Institute, Pretoria, and to

Mr. A. A. Bullock of Kew who provided valuable advice.

O.A. LE STNER

STERCULIACEAE

A New Species of Dombeya and a New Variety of Dombeya rotundifolia

Dombeya autumnalis Verdoorn, sp. nov. D. rotundifolia (Hochst.) Planch, affinis

sed plantis plerumque fruticosis caulibus gracilioribus, cortice levioie, floribus et foliis

plerumque in auctumno coaetaneis, ovario breve stellato-pubescente nunquam setosi

diffeit.

Frutex vel arbuscula, 1-6-5 m altus, caulibus pluribus gracilibus ad 5 cm diam.,

cortice leviore. Ramuli graciles, foliati, breve stellato-pubescentes. Folia suborbiculata,

1-5-5 cm longa, 1-5 cm lata, utrinque dense vel sparse stellato-pubescentia, subtiliter

cienato-dentata, subtus obscure reticulato-nervosa petiolis gracilibus 2-12 cm longis

stellato-pubescentibus. Cymae in axillis foliorum super iorum, quam folia subtendentia

longiores vel breviores, pedunculis gracilibus c 2-5 cm longis stellato-pubescentibus,

pedicellis giacilissimis 1-1-5 cm longis stellato-pubescentibus. Bracteolcie lineari-

naviculares, c. 2-5 mm longae, utrinque stellato-pubescentes. Calyx basi rotundatus

obtusus, 0-5 mm connatus, lobis reflexis c. 4 mm longis 2 mm latis, extus stellato-

pubescentibus. Petala alba, c 7 mm longa, 5 mm lata, persistentia. Stamina basi

144

connata in tubum c 1 mm longum, filamentis ad 2-5 mm longis, antheris c 1 mm longis

stami aodiis c 5 mm longis. Ovarium globosum, c 3 mm diam., breve stellato-tomentosum

(non setosum). Stylus glabrescens vel minute stellato-pubescens, c 2 mm longus in

ramos 3 c 2 mm longos divisus. Capsula c 5 mm diam., stellato-tomentosa.

Type: Lydenburg, Penge Mine, Verdoorn 2470 (PRE, holo.).

Shrub or small tree 1-5-5 m tall; stems several, slender, up to about 5 cm diam.;

bark compaiatively smooth, stringy. Branchlets slender, leafy, new growth shortly

stellate pubescent, the pubescence formed of short spreading hairs from a scaly base.

Stipules caducous, deltoid or linear-subulate from a deltoid base, densely pubescent.

Leaves more or less orbicular, on flowering branches 1-5-5 cm long, 1-5 cm bioad,

densely to sparesely stellate-pubescent on both surfaces, finely crenate-dentate on the

margins, reticulate veins rather obscure beneath; petiole slender, 2-12 cm long, stellate-

pubescent. Cymes in the axils of the upper leaves, often overtopping them, sometimes

shorter; peduncles slender about 2-5 cm long, pubescent; pedicels very slender 1-1-5

cm long, pubescent. Bracts 7 or 2 at the calyx-base, 1 further down or all 3 scattered

on the pedicel, linear-navicular, about 2-5 mm long pubescent on both surfaces. Calyx

rounded at the base, united for 0- 5 mm; lobes reflexed about 4 mm long, 2 mm broad,

dorsally stellate-pubescent. Petals persistent, white turning cinnamon rufous with

age, about 7 mm long, 5 mm broad. Stamens united at the base for more oi less 1 mm;

filaments with the longest about 2-5 mm long; anthers up to 1 mm long; staminodes

about 5 mm long. Ovary globose about 3 mm diam. shortly stellate-tomentose (not

setose); style glabrous or minutely stellate pubescent, about 2 mm long, branches

about 2 mm long; ovules 2 in a cell. Capsule about 5 mm diam., stellate-tomentose.

Recorded from the eastern Transvaal in mountainous country on mesophytic,

well-wooded slopes among rocks and in riverine bush.

Transvaal. — Lydenburg: Abel Erasmus Pass, Schlieben & Strey 8387; Codd 10027; near Penge

Mine, Codd & Dyer 7737; Codd & Verdoorn 10488; Repton 5936 (partly); Verdoorn 2470; near

Weltevreden Asbestos Mine, Verdoorn 2471. Letaba: Dublin Mine, Miller 4271.

The only other species in southern Africa, with suborbicular leaves is D. rotundifolia

which has a wide distribution stretching from Natal and the Tiansvaal northwards

into east tropical Africa and westwards through Bechuanaland to South West Africa.

D. autumnalis differs from D. rotundifolia (as it occurs in the eastern and central areas

of southern Africa) in that it flowers in late summer and autumn (with its leaves) while

D. rotundifolia flowers in spring on more or less leafles branches. From all forms

including those in South West Africa, our species differs mainly in that its slender

stems do not develope a lough bark. If collector’s notes about bark are lacking on

a specimen this feature can be determined by the examination of a cut bianchlet,

because the cortex in D. autumnalis appears as a thin, solid outer layer while in D.

rotundifolia it is a thicker, porous layer which evidently gives rise to the rough bark

on the old stems. A further diagnostic feature is that the pubescence on the ovary

of D. rotundifolia, throughout the length and breadth of its distribution, is setose as

well as stellate-tomentose, the bristle-like hairs standing more or less erect, and reaching

a length of up to 1 mm while in D. autumnalis the ovary is stellate-tomentose and not

setose as well. Other small differences can be found such as the distinctly reticulate

veining on the under surface of the leaf in D. rotundifolia while on our species the

reticular veining, if any, is obscure.

Dombeya rotundifolia ( Hochst .) Planch, var. velutina Verdoorn, var. nov., a typica

ovario breve stellato-tomentoso non setoso foliis utrinque velutino-tomentosis differt

Frutex elatus ad 5 m altus caulibus pluribus virgatis, cortice rugosa, ramulis

molliter et breve tomentosis Folia suborbiculata vel oblongo-orbiculata, 3-9 cm

longa, 2 -5-9 -5 cm lata, utrinque velutino-tomentosa, crenato-dentata, cordata, 5-7

145

palmato-nervosa, petiolis 1-2 cm longis molliter tomentosis. Cymae ramulis lateialibus

et terminalibus confertae, pedunculis 10-15 mm longis, pedicellis 7-10 mm longis.

Calyx c. 6 mm longus, extus dense et breve tomentosus. Petala c. 7 mm longa. Stamina

basi in tubum brevissimum connata, 0-5 mm longum; filamenta c. 2-5 mm longa;

staminodia c.5 mm longa. Ovarium dense et breve stellato-tomentosum (non setosum).

Stylus stellato-pubescens.

Type: South West Afiica. Rehoboth, Buellsport, Aub Schlucht, Strey 2010 (PRE,

holo., BOL, NBG).

Tall shrubs with several virgate stems, up to 5 m tall; bark rough, new growth

softly and shortly tomentose. Leaves suborbicular oi broadly oblong-orbicular,

broadest in the upper half 3-9 cm long, 2 -5-9 -5 cm broad, velvetty tomentose on

both surfaces (the tomentum made up of minute stellate-pubescent scales, the hairs

short and silky), crenate dentate, cordate at the base, palmately 5-7-nerved ; petiole 1-2 cm

long, softly tomentose. Cymes crowded on lateral and terminal branchlets; peduncles

10-15 mm long; pedicels 7—10 mm long. Calyx about 6 mm long, densely and shortly

tomentose without. Petals about 7 mm long. Stamens united at the base for about

0-5 mm. filaments unequal lengths, about 2-5 mm long, staminod.es about 5 mm long.

Ovary densely and shortly stellate-tomentose (not setose); style 2-3 mm long, stellate-

pubescent.

Occurs along the banks of the permanent stream at Aub Schlucht in the Naukluft

Mountains, South West Africa.

S.W.A. — Rehoboth: Naukluft Mountains, farm Buellsport, Aub Schlucht, Strey 2010; 2328;

Tolken & Hardy 666.

In the common and widespread species, D. rotundifolia, three features, in the

gross morphology, have been found to be constant and of diagnostic value. They

are the suborbicular leaves, the rough bark and the pubscence on the ovary which

is stellate-tomentose and setose. Microscopically this pubescence consists of stellate

and tufted hairs, the tufted being in the upper part of the ovary giving the bristly or

setose appearance. D. rotundifolia thus characterised occurs widespread in South

West Africa although not in the optimum form found so commonly in the eastern

and central regions of tropical and subtropical Africa. In South West Africa it generally

occurs as a group of low bushy shrubs and only where conditions are favourable does

it grow up into the characteristic, rough-barked tree.

In a valley in the Naukluft Mountains, Rehoboth district, along a permanent

stream the variety here described, grows. It has the suborbicular leaves and rough

bark of typical D. rotundifolia but the ovary is constantly shortly stellate tomentose,

not setose. Investigation has shown that the habit of these plants also differs from

that of typical D. rotundifolia as found in the rest of South West Africa. The plants

along the stream are all tall, several-stemmed shrubs. These two differences together

with a third, the velvetty texture of the pubescence seem to justify at least the varietal

rank here given this form of D. rotundifolia.

I. C. Verdoorn

147

Plate \.—Widdringtonia cedarbergensis Marsh (Luckhoff in PRE 29824, holotvpe)

7230691

148

Plate 2. — Kniphofia ritualis Codd ( Marais 1327, holotype, in PRE)

149

?P-gJ3363 — JM7-&— 144-Hon *. ^

NATIONAL HERBARIUM,

PRETORIA.

DISTRICT /,

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Plate 3 .—Kniphofia hirsute Codd ( Guillarmod

& Marais 1307, holotype, in PRE)

150

Plate 4. — Kniphofia umbrina Codd ( Bruce 272, holotype, in PRE)

151

Plate 5 .—Kniphofia acraea Codd ( Liebenberg 7120, holotype, in PRE)

7230691

153

Bothalia. 9, 1: 153-156.

The Genus Ceramium in South Africa

by

R. H. Simons

Abstract

A revision of the genus Ceramium in South Africa has been undertaken. The morphology of

the genus is discussed, and a key to the ten species which are recognised is provided. The following

new taxa and new combinations are published: Ceramium arenarium, C. papenfussianum, C. diaphanum

(Lightf.) Roth var. capense, — var. pulchellum (Kuetz.) (Hormoceras pulchellum Kuetz.), C. centro-

ceratiforme.

Ceramium capense Kuetz., which was transferred to Microcladia by Papenfuss, is regarded as

belonging to Ceramium.

Introduction

The Delimitation of Ceramium and Allied Genera

The genus Ceramium (family: Ceramiaceae), although used in a restricted sense

for over one hundred years, is still not fully understood. Dixon (1960) studied the

British forms recently and his observations together with those of Feldmann-Mazoyer

(1940) are detailed and informative. But certain problems, some highlighted by South

African forms, remain and, although the present author has attempted to resolve some

of them, it cannot yet be claimed that the genus Ceramium is satisfactorily delimited.

One of these problems concerns the attachment of tetrasporangia. According

to Dixon (1960), and some other authors, they arise from unspecified cells of the cortex.

The inference is that any cortical cell may act as the supporting cell of a tetrasporangium.

The present author was able to trace the origin of many tetrasporangia in five out of

the ten species listed in this paper. In every case they were attached to pericentral cells.

No other supporting cell was indicated. Where the actual derivation was not directly

observable the tetrasporangia always occupied positions such that attachment to the

pericentrals was not only feasible but highly likely. If this derivation is as constant as

the present author suggests it might be then it must be regarded as of fundamental

importance and any interpretation of Ceramium should take it into account. Obviously

more information is required on this subject before its value can be properly assessed.

The occurrence in South Africa of Ceramium centroceratiforme Simons, sp. nov.,

raises the second problem dealt with here, namely, that of the status of the genus

Centroceras Kuetz. From time to time species such as this one have been described

which link Ceramium and Centroceras, and opinions have been divided as to their

autonomy. Most authors have accepted them as different although some have done

so without much conviction. The tendency of Ceramium centroceratiforme Simons

to have an orderly arrangement of cortical cells but at the same time to have distinct

genicules somewhere along its length puzzled the author because the species appeared

to belong to both genera equally well. A solution seemed to be to regard Centroceras

as synonymous with Ceramium as some other authors have done. It was decided

instead to make a closer study of the South African form of Centroceras clavulatum

(Ag.) Mont., the type species of Centroceras, to discover whether there was any justifi-

cation for its separation from Ceramium. Certain facts emerged from this investigation

which the writer regards as notable.

154

First, in C. clavulatum the regular arrangement of basipetal secondary cortical

cells in a single layer is the result of their exclusively transverse segmentation. The

resultant cortical files of any single articulation develop equally by what appears to be

simultaneous segmentation of their apical cells. In species of Ceramium, on the other

hand, there is no notable regularity of segmentation of cortical cells either in time or

space: apical cells of the laterals of limited growth divide transversely, longitudinally

and obliquely. Secondly, in C. clavulatum acropetal development of cortex is virtually

absent; instead there is almost radial development to form a girdle of tissue raised

slightly from the surface of the thallus at every node. As a result of this, all genicules

retain their individuality and there is no confluence of cortical tissue from adjacent

axial cells. In contrast to this, completely corticated species of Ceramium have a tissue

which is continuous between successive nodes. Thirdly, in male plants of C. clavulatum,

spermatangia were found confined to the terminal part of each fertile articulation,

i.e. on the radially developed girdle of tissue at each node.

South African species of Ceramium bearing spermatangia have not been found

by the writer, but Dixon (1960) and others have reported on various other species

in which they found spermatangia scattered haphazardly over the surface of the cortex.

Of the characteristics of Centroceras listed above, the most significant seems to be

that relating to the development of the basipetal cortex of each articulation. The

contrast of the latter with what occurs in species of Ceramium is sufficient, in the present

author’s opinion, to justify the separate generic status of Centroceras Kuetz., as typified

by C. clavulatum (Ag.) Mont. The other differences seem to corroborate this view.

As to the affinities of C. centroceratiforme Simons: its cortical development is like

that of species of Ceramium, to which genus it, therefore, undoubtedly belongs.

A third problem has resulted from the recognition by Papenfuss (1940) of South

African material as a species of Microcladia. The main reason for doing this seems

to have been a well developed multi-layered cortex. Feldmann-Mazoyer (1940),

writing of Microcladia , claimed that the genus was little understood and contained

unrelated species. She suggested that the only justification for removing M. glandulosa

(Soland.) Grev. to a separate genus was its unique arrangement of cells of different

sizes in the innermost layer of the cortex. The latter characteristic leads to a flattened

bilateral thallus. The present author’s study of Microcladia capensis (Kuetz.) Papenf.,

the South African species referred to above, revealed that the thallus is not bilateral

nor does it possess any other characteristic to exclude it from Ceramium. This species

is dealt with in this paper as C. capense Kuetz.*

Some Morphological Features of South African Species of Ceramium

Commenting on the glandular cells so often found in species of Ceramium, Dixon

(1960) stated that these cells are rather short-lived. Tt is interesting to note, therefore,

that similar cells in one or two South African species are sometimes very persistent.

In C. glanduliferum Kylin, for example, they are often obtrusive and so long-lived

that the specific epithet was suggested by their presence. Glandular cells in C. papen-

fussianum Simons, sp. nov., were found to be almost as persistent but not nearly so

obvious as those of the previous species.

* Since writing this article the author has seen the paper by Max Hommersand on the ’'Morphology

and Classification of some Ceramiaceae and Rhodomelaceae" (University of California Publications in

Botany Vol. 35, No. 2, pp. 165-366, published in 1963). In this paper Hommersand discusses the

genera Ceramium , Microcladia and Centroceras ; he upholds the separate status of these three.

Microcladia produces upwardly and outwardly directed cortical filaments only which entirely

enclose the axial cells. This interpretation makes no mention of bilateral symmetry but does not

affect my placing of C. capense in the genus Ceramium.

Hommersand’s observations on the derivation of cortical filaments in Centroceras are essentially

similar to mine and are apparently sufficient reason for a generic separation.

Ceramium poeppigiana Grunow, removed to the genus Rheinboldiella and not considered in my

paper, is returned by Hommersand to the genus Ceramium.

155

Several South African species have extremely thick-walled axial cells. The mature

axial cells of C. papenfussianum Simons (Fig. 2: lc), for instance, have such thick

lamellate walls that the central lacunae are very narrow. The thickening appears to

develop centripetally in clearly demarcated layers on the inner faces of the cell walls,

and the result is a likeness to the annual rings of higher plants. This characterictic

no doubt leads to the comparative rigidity of these diaphanous plants.

Intercellular hyphae ramifying within the cortex of C. planum Kuetz. have been

seen in no other South African species. Their presence adds greatly to the girth of

an already thick cortex in the mature parts and renders the thallus very cartilaginous

and difficult to dissect when fresh. The origin of these hyphal filaments was not traced,

but they appeared sometimes to arise from a cingula-like thread of similar material

encircling a median constriction of an axial cell. Attachment of this cingula to the

axial cell was at times evident. Perhaps this species belongs to Campylaephora.

Intracellular hyphae ( Thylles of Feldmann-Mazoyer, 1940), such as those described

by Dixon (1960), have possibly been seen in only one South African species. However,

they were not entirely convincing.

Terminology of Morphological Features

The joints between adjacent axial cells are called nodes. The cortex consists of,

primarily, the pericentral cells differentiated at the distal end of every axial cell, and,

secondarily, the laterals of limited growth remaining closely appressed to the axial

cells. The laterals of limited growth arise by segmentation of apical cells cut off acro-

and basi-petally from the pericentrals. A cortical zone of common origin is referred to

as a genicule and uncorticated zones are inter genicules. Unless otherwise mentioned,

length is the dimension in the plane of the long axis of the plant, and width is measuied

at right angles to this.

CERAMIUM

Ceramium Roth in Cat. Bot. 1: 148 (1797).

Dictiderma Bonnemaison (1822).

Hormoceras Kuetz. (1841).

Gongroceras Kuetz. (1841).

Echinoceras Kuetz. (1841).

Acanthoceras Kuetz. (1841).

Chaetoceras Kuetz. ( 1 847), non Chaetoceros Ehr. ( 1 844), nee Chaetoceras Kuetz* (1849).

Trichoceras Kuetz. (1849).

Celeceras Kuetz. (1849).

Pteroceras Kuetz. (1849).

Ceramothamnion Richards (1901).

Plants of pseudodichotomously or irregularly branched uniaxial filaments, partly

or completely corticated by closely appressed cells in one or more layers, radially

symmetrical in transverse section, cells of inner layers larger than those of outer, arising

as laterals of limited growth from pericentrals, and cut off aero- and basi-petally by

random segmentation. Pericentrals attached to, and forming a ring around the distal

end of every axial cell. Tetrasporangia immersed in or exserted from cortex in vicinity

of nodes. Cystocarps naked, subtended by one or more involucral branches. Sper-

matangia scattered over surface of cortex (not seen in South African species).

Without any implication as to primitiveness or derivation, the species listed here

are arranged in a series decided, primarly, by the extent of tetrasporangial immersion

in the cortex and, secondarily, by the degree of cortication.

156

Genicules distinct in lower part of plant:

Intergenicules below equal to or longer than genicules:

Tetrasporangia unilateral or whorled, exserted 1. C. glanduliferum

Tetrasporangia whorled, partly or entirely immersed:

Tetrasporangia immersed partly:

Pericentrals at bases of genicules 3. C. tenerrimum

Pericentrals median in genicules; genicules contiguous in upper part of filaments

4. C. arenarium

Tetrasporangia immersed entirely:

Tetrasporangia patent through outer cortex; pericentrals frequently exposed and cortical

cells often transversely elongated 5. C. papenfussianum

Tetrasporangia not visible through outer cortex; pericentrals also occluded:

Plants flaccid, branches ultimately incurved strongly 7 (a) C. diaphanum var. capense

Plants setaceous, ultimately branches almost straight; apical cells patent 7 (, b ) C. diaphanum

var. pulchellum

Intergenicules all shorter than genicules:

Cortical cells randomly disposed 6. C. atrorubescens

Cortical cells somewhat ordered into longitudinal and transverse rows in lower parts of

genicules 9. C. centroceratiforme

Genicules not distinct in lower parts of plant:

Genicules distinct above; branch apices digitate 2. C. planum

Genicules nowhere distinct:

Branching more or less alternate:

Branching pinnate, racemo-corymbose 2. C. planum

Branching pseudodichotomous, alternate; adventitious branches frequently secund

10. C. obsoletum

Branching on all sides, profuse 8. C. capense

1. C. glanduliferum Kylin in Lunds Univ. Arsskr. N.F. Avd. 2, 34, 8: 14, fig.

7A-B (1938).

Holdfast : prostrate filaments with distinct but weakly developed genicules of one

half or one third the length of intergenicules; genicules often with single whorl of

large terminal, exserted glandular cells; rhizoids numerous at nodes. Erect system :

filaments up to 1-5 cm high, pseudodichotomously branched, adventitious branches

rare, 180 p wide at base tapering gradually to about 60 p terminally with slightly

incurved apices; axial cells up to one-and-a-half times longer than broad; genicules

shortly [separated above but widely so below, often up to twice as wide as long, some-

times even shorter and then pericentrals often at bases, cells angular minute (about

10 p X 5 p) most often with much larger and very conspicuous ovate (up to 30 p X 18/*),

exserted glandular cells in an apical whorl; number of pericentrals variable, up to nine,

often superficially visible through the cortex, more or less spherical; tetrasporangia

totally exserted. mostly unilateral on inner faces of upper dichotomies; cystocarps

not seen. Plate 1: 1.

Cape. — Mossel Bay: Tergniet, Simons 634. Knysna: Beacon Isle, Isaac B.942. East London:

Ecol. Sttrv* L66. Willowvale: Dwessa, Isaac B. 1013. Elliotdale: Bomvanaland, Flanagan s.n.

(Tyson s.n. sub BOL 27476).

Natal. — Durban: Ecol. Surv. D. 100.

Typically, this species is readily identifiable by its conspicuous glandular cells

but, when the latter are absent, it is not easily recognisable. The variable position

of the pericentrals in the genicules. however, is a fairly reliable characteristic.

* The abbreviation “ Ecol. Surv .” used in citations throughout this paper refers to material in

the University of Cape Town collection from the ecological survey of marine, littoral fauna and flora

of South Africa. This survey was directed during the period 1931-1940 by Prof. T. A. Stephenson and

is now being continued under the guidance of Prof. J. H. Day, both Professors acting in their capacities

as Head of the Department of Zoology, Cape Town University.

157

2. C. planum Kuetz. in Sp. Alg. 687 (1849); Tab. Phyc. 13: 5, pi. 1 1, fig. c-d (1863);

Papenf. in J. S. Afr. Bot. 17: 177 (1952).

C. cancellation Ag. in Syst. Alg. 136 (1824); Sp. Alg. 2: 145 (1828), non C. cancel-

latum (L.) DC. in Bull. Sci. Soc. Philom. Paris 3, 94: 264 (1804-1805). C. flexuosum

(Kuetz.) Grunow in Alg. Nov. 64 (1867), non C. flexuosum Ag. in Syst. Alg. 141 (1824).

Pteroceras cancellation (Ag.) Kuetz. in Sp. Alg. 690 (1849); Tab. Phyc. 13: 8,

pi. 22, fig. a-b (1863). P. flexuosum Kuetz. in Sp. Alg. 690 (1849); Tab. Phyc. 13:

8, pi. 22, fig. c-e (1863).

Holdfast: short haustorium-like rhizoids produced at base of erect filaments

attach plants to others. Erect system: up to 15 cm high, terete, about \ mm wide;

branching almost alternate and more or less distichous, racemo-corymbose; cortication

either complete or interrupted in upper parts, multi-layered below, large-celled inner

layer often with intercellular ramifying hyphae; pericentrals six, almost spherical;

tetrasporangia exserted, frequently bilateral, or hemicyclic, attached to pericentrals;

cystocarps surrounded by five or six involucral branches. Plate 1 : 2.

Cape. — Namaqualand: Port Nolloth, Simons 273; 331; Ecol. Surv. N. 19; Strong s.n. (Tyson

s.n. sub BOL 27435); Tyson s.n. sub BOL 27431; Buffels River, Ecol. Surv. BR2H. Clanwilliam:

Lamberts Bay, Ecol. Surv. B50. Malmesbury: Steenberg’s Cove, Ecol. Surv. C4E; Saldanha Bay,

Simons 594; Dassen Island, Tyson s.n. sub BOL 27429. Peninsula: Table Bay, Tyson s.n. sub BOL

21 All to 27428 incl., 27433, 27434, 27436, to 27448 incl.; Oudekraal, Ecol. Surv. A13; Kommetjie,

Simons 25; False Bay, Ecol. Surve. F. 115 in part sub Simons 655.

The numerous pinnate branches, arranged more or less distichously on the main

filaments, make this plant unique amongst the South African Ceramium species,

separating it clearly from C. capense Kuetz. especially. These two species have cortices

which at times are almost indistinguishable superficially. In longitudinal section their

thalli can be distinguished because the inner cells of C. capense are all more or less

elongated longitudinally whereas those of C. planum are more polygonal, those in the

middle of an articulation being more or less ovate and orientated so that their long

axes are radial to the adjacent axial cell. The ramifying hyphae present in C. planum

have not been seen in C. capense.

3. C. tenerrimum {Mart.) Okam. in Icon. Jap. Alg. 4: 112, pi. 179 fig. 1-7 (1921);

Feldmann-Mazoyer in Ceramiacees de la Mediterranee 289 (1940).

Hormoceras tenerrimum Mart, in Preuss. Exped. ost-Asien, Bot. Teil: Tange: 146,

pi. 13, fig. 2 (1866).

Holdfast: prostrate lower parts of erect filaments with distinct very short genicules;

numerous rhizoids attach plant to rock or other algae. Erect system: up to 2 cm high

and 200 p thick, branching pseudodichotomous, ultimate branches inrolled ; adventitious

branches occasional; genicules distinct throughout, short, up to three times wider

than long; pericentrals up to seven at bases of genicules; glandular cells present,

inconspicuous; tetrasporangia in single whorls on genicules of penultimate branches,

apically exserted, attached to pericentrals; cystocarps of three or four gonimolobes

subtended by about five branches up to five times longer than height of gonimolobes.

Fig. 1: 1; Plate 1: 3.

Cape. — Namaqualand: Port Nolloth, Simons 303; Hondeklip Bay, Ecol. Surv. F1B5B; Groen

River, Ecol. Surv. GR4F. Clanwilliam: Lamberts Bay, Ecol. Surv. B70. Malmesbury: Ysterfontein,

Ecol. Surv. YZ1F. Caledon: Flermanus, Simons 539; 540; F.ansk aal, Simons 647. Knysna:

Robberg, Isaac B. 788.

Although present among the older collections, this species was overlooked by

other workers. The specimens cited here compare well with one from the Mediter-

ranean supplied to the present author by Mme Feldmann. The most striking thing

about this species, compared with other South African species, is the basal position

of the pericentrals in almost every genicule. Cell rows initiated basipetally seldom

project below the bases of their pericentrals and are generally only one cell long.

158

Fig. 1. — 1, Ceramium tenerrimum, surface view of a genicule; la, longitudinal section through a

genicule; lb, transverse section through the pericentrals of a tetrasporiferous genicule 2, C.

arenarium, longitudinal section through thallus near the apex of a branch; 2a, longitu-

dinal section through a genicule of the mature portion of a thallus; 2b, transverse sec-

tion through the pericentrals of a genicule.

a.c. — axial cell; c. — cortical cell; g.c. — glandular cell; p. — pit connection; p.c. — pericentral

cell; t. — tetrasporangium.

159

Besides Japan, the Mediterranean and the Cape, there appears to be only one

other recorded locality for this species, viz. N. Pacific (Dawson, 1962, p. 49). Such

disjunctions seem strange and it is probable that C. tenerrimum will be found elsewhere.

4. C. arenarium Simons, sp. nov.

Thallus usque ad 3 cm altus, pileum formans, inferne circa 150 p latus pro parte

repens cum rhizoidibus multis ex geniculis excurrentibus, sursum quoque ramorum

gradatim gracilior; rami ultimi circinati; genicula superne contigua, inferne multo

distincta intergeniculis quam genicula triplo longioribus; cellulae pericentrales c. 7

cellulis corticalibus occlusae, plus minusve in medio geniculorum dispositae; tetras-

porangia in ramulis superioribus monoverticillata dimidio superiore exserto; cysto-

carpia subapicalia, geminata ramulis nunc paucis nunc saepius quinque vel sex incurvatis

involucrata. Fig. 1: 2; Plate 1: 4.

Type: Namaqualand, Port Nolloth, Isaac B.676 sub Simons 632 (PRE, hole., spirit

material only).

Holdfast : prostrate basal filaments with many rhizoids at each genicule entwining

neighbouring filaments and attaching plant to substratum; genicules distinct, half as

long as wide; intergenicules up to four times longer but of same width as genicules.

Erect system : up to 3 cm high, filaments dichotomously branched, frequently producing

adventitious branches in the axils in a plane at right angles to dichotomies, 150 y thick

or more, narrowing slightly at each dichotomy to about 100 y subapically, ultimately

circinately inrolled; pericentrals about seven completely obscured in the mature

parts, more or less median in genicule; genicules contiguous in upper half of filaments,

separated below and then about one-and-a-half times wider than long, intergenicules

becoming progressively longer abapically until about three times longer than genicules;

tetrasporangia in a whorl at each of the upper genicules, upper half exserted when

mature; cystocarps subapical, two gonimolobes, involucral branches up to six, some-

times absent.

South West Africa. — Luderitz, Simons 432.

Cape. — Namaqualand: Port Nolloth, Isaac B. 662; B. 676; Simons 632. Clanwilliam: Lamberts

Bay, Ecol. Surv. B. 13. Penin ula: Table Bay, Simons 117.

The characteristic circinate apices, the contiguous genicules above and the more

or less median pericentrals separate this species from C. tenerrimum (Mart.) Okam.

with which it shares the habit of partially exserted tetrasporangia. The epithet chosen

for this species is suggested by its sandy habitat amongst other low-growing turf-

forming algae.

Transverse sections of axial cells frequently showed thick-walled cellular inclusions.

Their true shape could not be deduced nor could their origin because longitudinal

sections provided no evidence of their presence. There appeared to be seven such

bodies circular in outline and closely appressed to the inner wall of the axial cells in

which they were seen. Their diameter was approximately equal to half the greatest

dimension of the pericentrals. It is possible that these structures may be similar to

the intracellular filaments referred to by Dixon (1960, p. 338) and Feldmann-Mazoyer

(1940, p. 161).

5. C. papenfussianum Simons , sp. nov.

C. pulchellum sensu Kylin in Lunds. Univ. Arsskr. N.F.Avd.2, 34, 8: 14 fig. 7C-E

(1938); non Hormoceras pulchellum Kuetz. (=C. diaphanum war. pulchellum (Kuetz.)

Simons, comb, et stat. nov., see p. 162).

160

Plantae parvae usque ad 2-5 cm altae, in aliis algis habitae, ramosissimae paene

fastigiatae, ramis supra hamatis; genicula usque ad 150 p. lata, omnibus praeter

terminalia distinctis; cellulae pericentrales plerumque 6 patentes; cellulae corticales

angulatae, basi-petalibus praecipue in geniculis superioribus saepe transverse elongatis;

tetrasporangia monoverticillata, immersa, patenti; cystocarpia ramulis 5 vel 6 involu-

crata. Fig. 2: Plate 2: 1.

Fig. 2. — 1, Cevamium papenfussianum, surface view of two genicules; la, transverse section through

the pericentrals of a mature genicule; lb, longitudinal section through a tetrasporiferous

genicule showing attachment of tetrasporangium to pericentral cell; lc, transverse section

through intergenicule showing curious wall thickening.

a.c. — axial cell; c. — cortical cell; p.c. — pericentral cell; t. — tetrasporangium; t.p.c. — tetras-

poriferous pericentral cell.

Type: Peninsula, Sea Point, Simons 694 (PRE, holo.; iso. in spirit collection).

Holdfast : prostrate lower filaments with widely spaced genicules from which

numerous rhizoids arise to attach plant to others. Erect system', filaments up to 3 cm

high, branching pseudodichotomous, almost fastigiate, terminal branches more or less

straight but hooked inwards at the tips; adventitious branches absent; genicules distinct

except terminally, widely separated below, about as wide as long; intergenicules up

to four times longer than wide; pericentrals usually six, median in genicules, often

clearly visible between outer smaller cortical cells; cortical cells angled, basipetal

cells frequently transversely elongated especially in upper genicules; glandular cells

sometimes conspicuous as an apical whorl on genicules, distinctly angled, and of same

size as other cortical cells; tetrasporangia whorled, subapical, immersed but patent,

as high as parent genicules, attached to pericentrals; cystocarps surrounded by five

or six involucral branches up to twice as high as gonimolobes.

Cape. — Namaqualand: Port Nolloth, Simons 275; 302. Malmesbury: Paternoster, Simons 482;

Ecol. Surv. P10A; Saldanha Bay, Simons 571; 572; 576; 581; 586; 587; 597; 609; 611; 619.

Peninsula: Sea Point, Simons 694; Levyns 0170 ( Tyson s.n. sub BOL 27470); St. James (False Bay),

Ecol. Surv. F115. Caledon: Hermanus, Simons 538; 541; 542; 545. Mossel Bay: Tergniet, Isaac

B. 949; B. 950. Knysna: Robberg, Isaac B. 806. Port Elizabeth, Ecol. Surv. E75. East London,

Ecol. Surv. L66. Komga: Cape Morgan, Flanagan 111 ( Tyson s.n. sub BOL 27471). Port St. Johns,

Isaac B. 834. Locality not stated: Flanagan s.n. ( Tyson s.n. sub BOL 27460-27463 inch). Collector

unknown ( Tyson s.n. sub BOL 27467; 27468).

161

C. papenfussianum is common on both the west and south coasts of the Cape.

The occurrence of the transversely elongated cortical cells caused earlier workers to

assign this species to C. gracillimum Grift', et Harv., but these two species differ in the

positions of their tetrasporangia, those of C. papenfussianum being immersed while

those of C. gracillimum are exserted. A more recent confusion has been that of Kylin

(1938) who misidentifiea this species as Hormoceras pulchellum Kuetz. The following

characteristics of H. pulchellum Kuetz. can be deduced from Kutzing’s (1849, 1862)

description and/or figures: first, the patent and somewhat diverging apical cells;

secondly, the completely obscured pericentral cells; thirdly, the presence of many

and often opposite adventitious branches. None of these characteristics is possessed

by C. papenfussianum.

J. G. Agardh (1851) assigned H. pulchellum Kuetz. to Ceramium diaphanum (Lightf.)

Roth because he could find no essential difference between the two. The present author

agrees with Agardh to the point of including H. pulchellum Kuetz. in the species C.

diaphanum (Lightf.) Roth, but considers it sufficiently different from the typical form

to be treated as a separate variety. In this paper H. pulchellum Kuetz. is placed as

C. diaphanum var. pulchellum (Kuetz.) Simons, comb. et. stat. nov. (p. 162).

The presence of the ring of glandular cells sometimes seen girdling the upper

margin of the genicules of C. papenfussianum might confuse the observer but, unlike

the relatively large ovate glandular cells of C. glanduliferum Kylin, those of this species

are angled and no bigger than normal cortical cells.

The specific epithet for this species was chosen in honour of Prof. G. F. Papenfuss,

who has contributed so much to the knowledge of South African marine algae.

6. C. atrorubescens Kylin in Lunds Univ. Arsskr. N.F. Avd. 2, 34, 8: 15, fig. 7F

(1938).

Holdfast : prostrate lower parts of filaments with numerous rhizoids arising from

narrowly separated genicules. Erect system : up to about 3 cm high and 500 p thick

below, narrowing upwards to about 100 p: branching pseudodichotomous, adventitious

branches at right angles in dichotomies, often opposite; branches ultimately somewhat

incurved; genicules variable in length, somewhat wider than long and scarcely separated

above, up to one-and-a-half times longer than wide with intergenicules varying greatly

in length below, acropetal development of cortex greater than basipetal, upper margin

of genicule undefined; pericentrals more than eight, superficially scarcely visible;

tetrasporangia immersed, patent, in a single whorl on every genicule throughout almost

entire length of erect filaments; cystocarps subapical, subtended by four or five

involucral branches somewhat longer than height of gonimolobes. Plate 1 : 5.

South West Africa. — Swakopmund, Simons 641; “ Walfisch Bay”, Cleverly s.n. ( Tyson s.n. sub

BOL 27473; 27474); Luderitz, Simons 417; Isaac B. 754.

Cape. — Namaqualand: Port Nolloth, Ecol. Surv. N42.

This species has a superficial resemblance to C. circinatum (Kuetz.) J. Ag., but

its genicules are, as it were, inverted. It is also very like C. arenarium Simons, sp.

nov. The chief difference between C. arenarium and C. atrorubescens is that the

first has genicules in which the pericentrals are always median, whereas those of C.

atrorubescens seldom are. Other differences between them are the circinate tips, the

continuous cortex above, the often half-exserted tetrasporangia and the production

of rhizoids for the greater length of the filaments of C. arenarium , all of which charac-

teristics contrast with their counterparts in C. atrorubescens.

7. C. diaphanum (Lightf.) Roth in Cat. Bot. 3: 154 (1806); Feldman-Mazoyer

in Ceramiacees de la Mediterranee 306 (1940).

1 62

Holdfast4, prostrate filaments with distinct genicules from which many rhizoids

arise; litho- or epi-phyte. Erect system: genicules distinct throughout, often wider

than intergenicules; seven pericentrals completely obscured by outer cortical tissue of

large and small cells; tetrasporangia immersed, often inconspicuous; cystocarps

subapical, subtended by involucral branches.

(a) var. capense Simons, var. nov. Plate 1 : 6.

A var. typico Feldmann-Mazoyer (1940), tetrasporangiis plus minusve geniculorum

maturorum dimidio superiori circumscriptis, ramis ultimis valde incurvatis differt.

Type: Hopefield, Schaapen Island (Saldanha Bay), Simons 614 (PRE, holo.,

spirit collection only).

Differs from var. typicum Feldmann-Mazoyer (1940) by its tetrasporangia being

more or less confined to upper half of mature genicules and its strongly incurved branch

apices.

Holdfast: prostrate filaments with distinct genicules; numerous rhizoids from

genicules attach plant to substratum. Erect system: up to 5 cm high, intergenicules

about 250 p thick in lower parts; genicules tumescent in lower parts, as much as 450 p

wide medianly and equally high, becoming relatively less swollen distally and ultimately

scarcely wider than the intergenicules at about 50 p; branching mainly pseudodichoto-

mous but adventitious branches common, branches ultimately strongly incurved;

tetrasporangia immersed and obscured in upper halves of mature genicules; cystocarps

not seen.

Known only from the single record cited above. This plant, like var. pulchellum

(Kuetz.) Simons, stat. nov., shows an arrangement of cortical cells essentially similar

to that described and figured for C. diaphanum (Lightf.) Roth by Feldmann-Mazoyer

(1940). The other shared characteristics of seven pericentrals and immersed tetras-

porangia have convinced the present author that it should be assigned to C. diaphanum.

Many varieties of this species have been recognised and these two (var. capense and

var. pulchellum) are sufficiently different from other varieties and from each other to

deserve separate varietal rank. These two differ from each other mainly in texture

and in the shapes of their ultimate branches.

( b ) var. pulchellum {Kuetz) Simons, comb, et stat. nov.

Hormoceras pulchellum Kuetz. in Sp. Alg. 676 (1849); Tab. Phyc. 12: 23, pi. 75,

fig. d-i (1862).

Ceramium pulchellum (Kuetz.) Kylin in Lunds Univ. Arsskr. N.F. Avd. 2, 34,8: 14,

fig. 7C-E (1938).

Holdfast: plant epiphytic with basal haustorial attachment and prostrate filaments

provided with rhizoids at genicules which are distinct and somewhat swollen. Erect

system: filaments up to 7 cm high, about 400 p below tapering ultimately to patent

relatively large apical cells about 10 p wide, much branched both pseudodichotomously

and adventitiously; adventitious branches arising from genicules singly, oppositely

or in occasional whorls, strongly divergent at first but soon becoming parallel with the

parent axis, branched, all tapering upwards and downwards; ultimate dichotomies

slightly incurved but apical cells remaining patent; genicules distinct throughout

with intergenicules about as long as genicules; tetrasporangia immersed in whorls in

penultimate and older branches; cystocarps surrounded by between three and six

involucral branches somewhat mnger than the height of the gonimolobes. Fig. 3: 1;

Plate 2: 2.

Cape. — Hopefield: Schaapen Island (Saldanha Bay), Simons 633. Peninsula: Table Bay, Tyson

s.n. sub BOL 27475, 27477 to 27482 inch

163

164

This variety differs from var. diaphanum by its relatively large, patent, divergent

apical cells.

All the Tyson specimens cited under this variety were designated “ C. kylinii sp.

nov. ined. G. F. Papenfuss”; a further two specimens similarly designated are of

C. atrorubescens Kylin.

8. C. capense Kuetz. in Linnaea 15: 740 (1841); Sp. Alg. 686 (1849); Tab. Phyc.

13: 3, pi. 5, fig. c-e (1863).

Microcladia capensis (Kuetz.) Papenf. in Bot. Not. 223 (1940).

Holdfast: epiphytic plant attached by basal haustorial rhizoids. Erect system-

plant up to 20 cm high and 1 mm thick near base, tapering downwards slightly, and

upwards gradually to about 50 p subapically; apically strongly incurved; main

branching pseudodichotomous, adventitious branches frequent, often in whorls, at

almost every articulation; cortication complete, multilayered when mature, outer

layer of very small cells, inner layer of large elongated or isodiametric cells; seven

pericentrals; articulations half as long as wide, cylindrical, translucent in upper parts,

lower down one-and-a-half times longer, medianly constricted; tetrasporangia whorled,

completely immersed, attached to pericentrals, often in two transverse ranks; cystocarps

clasped by five or six involucral branches. Fig. 3: 2.

South West Africa. — Luderitz: Elizabeth Bay, Simons 257.

Cape. — Namaqualand: Port Nolloth, Simons 278; 313; Isaac B. 658; Ecol. Surv. N16; N20A ;

N40; Groen River, Ecol. Surv. GR4E. Clanwilliam: Lamberts Bay, Ecol. Surv. B12; B13; B39;

B51; B52. Malmesbury: Paternoster, Ecol. Surv. P6A; Saldanha Bay, Simons 570; Isaac B. 31.

Peninsula: Table Bay, Tyson s.n. sub BOL 27485-27491 inch; Kommetjie, Levyns 0269 ( Tyson s.n.

sub BOL 27492).

This species differs from C. rubrum (Huds.) Ag. in having seven pericentrals

instead of eight (Feldmann-Mazoyer, 1940). Very similar to C. vimineum J. Ag.,

C. capense differs by its more extensive branching and rather shorter mature articu-

lations. True nodes, indicated by the point of origin of lateral branches, alternate

with false ones marked by transverse constrictions of the frond especially in the older

parts.

The description given above applies to the plant designated by Papenfuss as

Microcladia capensis, but it does not seem to apply well to Ceramium capense Kuetz.

which Papenfuss has designated as the basionym of this species. For instance, the

articulations of C. capense are all half as long as wide and the cortical cells are larger

(than those of C. rubrum). The present author is of the opinion that C. capense Kuetz.

is a form of C. obsoletum Ag., but on the available evidence hesitates to make the change

because it would necessitate the erection of a new name for M. capensis Papenf. The

missing type of C. capense must be found and re-examined before this matter can

be satisfactorily resolved.

9. C. centroceratiforme Simons, sp. nov.

Fila prostrata, intricata, pulvinum parvum formantia; genicula distincta, pellucida,

rhizoidibus numerosis: cellulae pericentrales in dimidio superiore geniculorum; corticales

sub pericentralibus plus minusve longitudinaliter vel transverse seriatae, omnibus sub-

pellucidis; fila erecta, pseudo-dichotomoso-ramosa, usque ad 2 cm alta, 300 p lata,

interdum ramis adventitiis instructa; rami ultimi suberecti, breves, divaricati et fere

triangulares; genicula superiora contigua vel fere contigua, inferiora plus minusve

distincta ; pericentrales 6 vel 7, in partibus superioribus medianae, in partibus inferioribus

in dimidio superiore geniculorum dispositae; cellulae corticales plus minusve in stratis

transverse et longitudinaliter Centrocerati similes dispositae, praecipue in partibus

165

geniculorum infra pericentrales; tetrasporangia verticillata subterminalia omnino

immersa vix manifesta, pericentralibus orta; cystocarpia bilobata uno ramulo subtensa.

Plantae subflavae vel galbanae. Fig. 3: 3; Plate 2: 3.

Type: Caledon, Hermanus, Simons 555 (PRE, holo., spirit collection only).

Holdfast : prostrate filaments matted together forming a small cushion, rhizoids

abundant at distinct genicules; pericentrals in upper half of genicules; cortical cells

below pericentrals arranged more or less in longitudinal and transverse rows, all more

or less colourless. Erect system : up to 2 cm high and 300 n thick, branching dichoto-

mous, adventitious branches occasional, ultimate branches short, more or less straight

and somewhat triangular in contour; upper genicules contiguous or almost so, lower

genicules the same or distinct and scarcely different from those of prostrate filaments,

longer than intergenicules; cortex, in vicinity of articulations and sometimes elsewhere,

two-layered; pericentrals six or seven, median in upper parts, in lower parts occur

in upper half of genicules; cortical cells arranged more or less in transverse and longi-

tudinal rows similar to those of the genus Centroceras, especially in parts of genicule

below pericentrals; tetrasporangia immersed, whorled, subterminal, scarcely discer-

nible, attached to pericentrals; cystocarps two-lobed, subtended by only one ramule.

Plant yellowish or greenish-yellow.

Cape. — Caledon: Hermanus, Simons 489; 555; 557. Knysna: Noetzie, Isaac B. 755; Robberg,

Isaac B. 939; B. 941. Willowvale: Dwessa, Isaac B. 1014.

The superficial appearance of this species is so like that of a Centroceras that

sometimes only the distinct genicules of the prostrate filaments disclose its identity.

Mostly, however, the genicules in other parts of the plant are also distinct. The author’s

reasons for assigning this species to Ceramium and not to Centroceras have been given

in the introduction to this paper.

10. C. obsoletum Ag. in Sp. Alg. 2: 145 (1828); Kuetz. in Sp. Alg. 687 (1849);

Tab. Phyc. 13: 4, pi. 10, fig. a-d (1863); Papenf. in J. S. Afr. Bot. 17: 178 (1952).

C. fnrcellatum Kuetz. in Sp. Alg. 687 (1849); Tab. Phyc. 13: 4, pi. 11, fig. a-b

(1863).

Holdfast : epiphytic on other algae, haustorial rhizoids arising from the base.

Erect system : somewhat rigid and setaceous, up to 20 cm high, 700 /u thick below,

tapering slightly to base and upwards gradually; branching pseudodichotomous,

often diverging strongly; adventitious branches sparse, but numerous and secund

when fertile; cortex continuous throughout, one or more layered, inner layer of large

cells of almost hexagonal outline with small angular cells seeming to cover only the

interstices between the larger cells; pericentrals nine or ten, more or less spherical

and largest of the cortical cells; plant translucent throughout, articulations as long

as cr shorter than width; tetrasporangia immersed in adventitious stichidia-like branches

and in penultimate branches; cystocarps on short adventitious branches subtended

by about five involucral branches. Plate 2: 4.

Cape. — Peninsula: Kommetjie, Simons 692. Knysna: Robberg, Ecol. Surv. RR1C. Bathurst:

Kowie (Shark’s Bay), Tyson s.n. sub BOL 27483. East London: Ecol. Surv. LI 14. Komga: Cape

Morgan, Flanagan 56 ( Tyson s.n. sub BOL 27484).

There is a confusing amount of variation in the habit of this species: in well-

developed plants the pseudodichotomies diverge at right angles to one another, whilst

in other plants they converge somewhat. Ultimately, however, they are never inrolled.

The most consistent features of this plant are the large cortical cells outlined by very

much smaller cells and the translucent thallus. When dry the plant is more or less

opaque.

166

References

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AGARDH, C. A., 1828. Species Algarum. 2. Greifswald.

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AGARDH, J. G., 1894. Analecta Algologica. Continuatio 2. Lunds Univ. Arsskr. 30.

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167

5 6

Plate 1.— 1, Ceramium glanduliferum, terminal fertile dichotomies, X 60 ( Isaac B 942)' la part of

mature filament (refractive cells are the glandular cells), x 60; 2, 2a, C. planum , completely

and partially corticated terminal portions of racemes, X 10 (2a, Simons 655); 3, C tener-

rimum, terminal and part of prostrate filament with rhizoids, X 30 ( Simons 647)- 4 C

arenarium, terminal portion of erect system (tetrasporiferous), X 30 ( Simons 632); 5’ C

atrorubescens, portion of a mature filament with two fertile laterals, each indicating a node

x 30 (Ecol. Sin v. N, 42); 6. C. diaphanum var. capense, terminal portions of laterals one

attached to a mature genicule, X 30 (Simons 614).

7230691

168

Plate 2. — 1, Ceramium papenfussianum, terminal tetrasporiferous portion of erect system, x 30

( Simons 609); la, transverse section through distal end of an axial cell showing several

pit-connections to the ring of pericentrals, X 500 (Ecol. Surv. F. 115); 2, C. diaphanum

var. pulchellum, general habit, X 2; 2a, detail of tetrasporiferous plant (swollen genicules

indicate presence of tetrasporangia), X 30 (Simons 633); 3, C. centroceratiforme, terminal

portions of filaments and portion of more mature thallus, X 30 ( Isaac B. 939); 4, C. obso-

letum, terminal portions of filaments (slight swelling of some genicules indicate presence of

tetrasporangia), X 15 (Ecol. Surv. L. 114),

169

Botha Ha 9, 1:169-171

A New Species of the Dictyotales from

South Africa

by

R. H. Simons

Whilst examining some seaweeds from Natal, I came across a few specimens

collected by Mr. M. Meyer, Lecturer at the Botany Department of the University

of Natal, which had the appearance of a dichotomising Zonaria. The specimens were

mounted on a single sheet and came from Kosi Bay in the District of Ingwavuma.

The characteristics of these plants were unlike any Zonaria previously described from

South Africa and it was therefore decided to make a detailed study of them.

Externally, the plant looks like a small Dilophus suhrii (Kutz.) Papenf. with its

dichotomising, strap-shaped branches with distinctly darker margins. Near the base

of the plant, multicellular brown rhizoids are produced, forming a stupose mat. The

resemblance to Dilophus ends here because the Kosi Bay plant has a terminal meristem

instead of a single apical cell. The fronds develop zonately, a dark transverse band

marking the termination of a period of growth. Scattered over both surfaces,

particularly towards the distal parts of the fronds, were raised sori of two kinds: one

containing nothing but multicellular uniseriate filaments (Fig. 1, C) and the other

containing large oval bodies, which were empty or had undivided contents, together

with occasional marginally disposed uniseriate filaments as in the other type of sorus.

The large oval bodies were sessile. Neither type of mature sorus was found with a

complete indusium but the remnants of the ruptured cuticle were always visible at

the margins of the sectioned sori (Fig. 1, C). The multicellular uniseriate filaments

are interpreted as hairs (paranemata) because they show no signs of being fertile.

The oval bodies appear to be fertile since they were often observed to be without

contents. Because their contents, when present, were undivided they are almost

certainly oogonia. No sporangia or male sori were found.

Closely appressed tubular skeletons of a Bryozoan forming, as it were, an additional

tissue were often seen. The Bryozoan, so far as could be seen, was confined

to the ventral surface of the fronds. Although the presence of the Bryozoan

did not seem to prevent sori from developing, the sori on the affected surface were

not as frequent as on the other surface.

The tissue is differentiated into four : two epidermal layers, a cortex and a

medulla (Fig. 1, A, B, and C). The dorsal epidermis consists of a single layer of

rather small sub-isodiametric cells, whereas the ventral epidermis has somewhat

longer cells. A central medulla of a single layer of elongated and relatively high cells

is contained within a dorsal and ventral multi-layered cortex. The medullary cells

differ from the cortical cells only by their greater height. The cells of the internal

tissue are almost always twice as wide as the epidermal cells (Fig. 1, B) except that the

subepidermal layer, on the dorsal side only, sometimes has cells of the same width

as the epidermis. On the other hand cells of the ventral epidermis are occasionally

also twice as wide as those of the dorsal epidermis. Thus, the differentiation into

dorsal and ventral surfaces is quite clear. In longitudinal section (Fig. 1, E) the origin

of new tissue in the zonate bands is seen to be from the medulla and inner cortex, the

superficial remnant layers forming flaps of truncated tissue heavily pigmented on the

upper and lower surfaces.

170

A

Fig. 1. — Pocockiella dichotoma Simons: A, L.S. of mature frond; B, T.S. of mature frond; C, T.S.

of frond showing hairs; D, L.S. through an oogonium; E, L.S. of frond through zone of

regeneration.

c: cortex; de: dorsal epidermis; e: epidermis; h: hairs; m: medulla; nf: new

frond; o: oogonium; of: old frond; ve: ventral epidermis.

As to the relationships of this plant, its zonate development and terminal meristem

indicate that it is allied to species of Zonaria. Besides, new tissue originates from the

middle layers of the frond (Fig. 1, E) as in Zonaria (Simons, 1964). it does not agree

in all features, however with the typical Zonaria construction. The absence of a stalk

cell at the base of an oogonium is found only in Zonaria tournefortii and Pocockiella

variegata ( =Gymnosorus variegatns) amongst the Dictyotales (fide Papenfuss, 1943).

171

Secondly, the presence of the medulla as distinct from the cortex is not found in species

of Zonaria. Thirdly, there is a tendency towards dorsiventrality of the thallus as

evidenced by the longer ventral epidermal cells. Finally, the arrangement of hairs in

discrete sorus-like groups instead of transverse bands is not found in species of Zonaria.

A plant which differs from typical Zonaria in like manner to the above is Pocockiella

variegata on which the genus Pocockiella Papenfuss is based. The present plant could

therefore be placed in Pocockiella. The characteristic separating Pocockiella from

Zonaria is the absence of paraphyses in its sporangial sori. These structures, unfor-

tunately, are not present on the Kosi Bay specimens. Papenfuss in his discussion

(l.c., p. 465) expressed the opinion that the peculiar internal differentiation into

cortex and medulla found in P. variegata was diagnostic and on that basis transferred

Gynmosorus nigreseens (Sond.) J. Ag. to Pocockiella. On the same basis, therefore,

the present plant should be regarded as a Pocockiella and it is described in this

genus below. The specific epithet refers to its distinctive dichotomous habit.

Pocockiella dichotoma Simons sp. nov.

Thallus dorsiventralis, ramosus, ramis linearibus, pseudodichotomis, margine

integerrimis, zonatis, inferne rhizoides gerens, ex quattuor stratis, stroma dorsali

cellularum subisodiametricarum, strato pleiostromatico corticali, stroma cen-

trali medullari et stroma ventrali cellularum sub-elongatarum, constans; cellulis

corticalibus medullaribusque aeque elongatis sed medullaribus in altitudine

majoribus, utriusque quam cellulis superficialibus duplo latioribus et ad usque octem

longioribus; paranemata et oogonia in soris discretis nudis; oogonia is sessilia; sporangia

et antheridia ignota.

Type: Natal: Ingwavuma, Kosi Bay, Meyer sub Simons 625 (PRE, holo.).

References

PAPENFUSS, G. F., 1943. Notes on Algal nomenclature. IF Gynmosorus J. Agardh. Amer. J.

Bot., 30 (7): 463-468.

SIMONS, R. H., 1964. Notes on the species of Zonaria in South Africa. Bothalia 8: 195-200.

173

bothalia 9, 1 : 173-1 82

A Virus Disease of Datura Spp.