ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 31,2

Oct. 2001

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

Obtainable from the National Botanical Institute, Private Bag XlOl, Pretoria 0001, Republic of

South Africa. A catalogue of all available publications will be issued on request.

BOTHALIA

Bothalia is named in honour of General Louis Botha, first Premier and Minister of Agriculture of

the Union of South Africa. This house journal of the National Botanical Institute, Pretoria, is

devoted to the furtherance of botanical science. The main fields covered are taxonomy, ecology,

anatomy and cytology. Two parts of the journal and an index to contents, authors and subjects are

published annually.

Three booklets of the contents (a) to Vols 1-20, (b) to Vols 21-25 and (c) to Vols 26-30, are available.

STRELITZIA

A series of occasional publications on southern African flora and vegetation, replacing Memoirs of

the Botanical Survey of South Africa and Annals of Kirstenbosch Botanic Gardens.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

The memoirs are individual treatises usually of an ecological nature, but sometimes dealing with

taxonomy or economic botany. Published: Nos 1-63 (many out of print). Discontinued after No. 63.

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

A series devoted to the publication of monographs and major works on southern African flora.

Published: Vols 14-19 (earlier volumes published as Supplementary volumes to the Journal of

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with accompanying text. The plates are prepared

mainly by the artists at the National Botanical Institute. Many botanical artists have contributed to

the series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian Condy, Betty Connell, Stella

Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700 plates), Claire Linder-

Smith and Ellaphie Ward-Hilhorst. The Editor is pleased to receive living plants of general interest

or of economic value for illustration.

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

An index to Vols 1^9 is available.

FLORA OF SOUTHERN AFRICA (FSA)

A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia

and Botswana. The FSA contains descriptions of families, genera, species, infraspecific taxa, keys

to genera and species, synonymy, literature and limited specimen citations, as well as taxonomic

and ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the informa-

tion is presented in the form of tables and photographic plates depicting fossil populations. Now

available:

Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidiiun, 1983, by J.M. & H.M.

Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidiiun), 1989, by J.M.

& H.M. Anderson.

Prodromus of South African Megalloras. Devonian to Lower Cretaceous, 1985, by J.M. &

H.M. Anderson. Obtainable from: A. A. Balkema Mai'keting, Box 317, Claremont 7735,

RSA.

Towards Gondwana Alive. Promoting biodiversity and stemming the Sixth Extinction, 1999,

by J.M. Anderson (ed.)

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 31,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

i5§OTANICAL

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XlOl, Pretoria 0001

ISSN 0006 8241

October 2001

Editorial Board

D.F, Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBl, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

Acknowledgements to referees

Archer, Mrs C. National Botanical Institute, Pretoria, RSA.

Balkwill, Prof. K. University of the Witwatersrand, Johannesburg, RSA.

Beyers, Dr J.B.P. National Botanical Institute, Cape Town, RSA.

Braggins, Dr J. Auckland, New Zealand.

Bredenkamp, Mrs C.L. National Botanical Institute, Pretoria, RSA.

Bremer, Prof. K. University of Uppsala, Uppsala, Sweden.

Brown, Dr E. Royal Botanic Gardens, Sydney, Australia.

Brownsey, Dr P.J. Herbarium, National Museum, Wellington, New Zealand.

Cutler, Dr D.F. Royal Botanic Gardens, Kew, UK.

De Paiva, Dr J.A.R. University of Coimbra, Coimbra, Portugal.

Farjon, Dr A. Royal Botanic Gardens, Kew, UK.

Furuki, Dr T. Natural History Museum & Institute, Chiba, Japan.

Geerinck, Dr D. National Botanical Garden, Meise, Belgium.

Henderson, Ms F. Agricultural Research Council, Pretoria, RSA.

Hill, Dr K.D. National Herbarium of New South Wales, Sydney, Australia.

Jordaan, Ms M. National Botanical Institute, Pretoria, RSA.

Jovet-Ast, Dr S. Biarritz, France.

Kativu, S. National Herbarium, Harare, Zimbabwe.

Leistner, Dr O.A. National Botanical Institute, Pretoria, RSA.

Manning, Dr J.C. National Botanical Institute, Cape Town, RSA.

Moll, Prof. E.J. SAWC, Hoedspruit, RSA.

Nelson, Dr E.C. Outwell, Wisbech, UK.

Nevling, Dr L.I. Illinois Natural History Survey, USA.

Newton, Prof. F.E. Kenyatta University, Nairobi, Kenya.

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

Parris, Dr B. Fern Research Foundation, Bay of Islands, New Zealand.

Patterson-Jones, Dr D. National Botanical Institute, Cape Town, RSA.

Poes, Prof. T. Eszterhazy College, Eger, Hungary.

Retief, Ms E. National Botanical Institute, Pretoria, RSA.

Rourke, Dr J.P. National Botanical Institute, Cape Town, RSA.

Roux, Dr J.P. National Botanical Institute, Cape Town.

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

Schutte-Vlok, Ms A. Oudtshoorn, RSA.

Weberling, Prof. Dr F. University of Ulm, Germany.

Welman, Ms W.G. National Botanical Institute, Pretoria, RSA.

CONTENTS

Volume 31,2

1. Studies in the genus Riccia (Marchantiales) from southern Africa. 25. A new species in subgenus Ric-

ciella, section Ricciella. S.M. PEROLD 151

2. Five new species of Erica (Ericaceae) from the Swartberg Range, Western Cape, South Africa

and a note on E. esterhuyseniae. E.G.H. OLIVER and I.M. OLIVER 155

3. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 1. The genus Aneura

and its local representative. S.M. PEROLD 167

4. Three new species of Tritoniopsis (Iridaceae: Crocoideae) from the Cape Region of South Africa J.C.

MANNING and P. GOLDBLATT 175

5. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 2. The genus Riccardia

and its type species. R. multifida. with confirmation of its presence in the region. S.M. PEROLD 183

6. Notes on African plants;

Acanthaceae, Thymelaeaceae, Lamiaceae. New records and distributional disjunctions from South

Africa, Zimbabwe and Mozambique. T.J. EDWARDS, A.J. BEAUMONT and D. STYLES 199

Colchicaceae. A remarkable new species of Androcymhium from Northern Cape. South Africa.

J.C. MANNING and P. GOLDBLATT 203

Dennstaedtiaceae-Pteropsida. Hypolepis villoso-viscida new to the Flora of southern Africa. J.P.

ROUX 195

Eabaceae. The identity of Melolohium lampolobiim (Papilionoideae) A. MOTEETEE and B-E.

VANWYK 209

Gentianaceae. A new' species of Sebaea from the Swartberg Range, Western Cape. South Africa.

I.M. OLIVER and J.B.P. BEYERS 207

Hyacinthaceae. The generic delimitation within Hyacinthaceae, a comment on works by E. Speta.

B. STEDJE 192

Iridaceae. Two new renosterveld species of Crocoideae from South Africa. J.C. MANNING and

P. GOLDBLATT 189

Portulacaceae. Talinum panicidatum. a naturalized weed in South Africa. E.M.A. STEYN and

G.F. SMITH 195

Pteridophyta. New distribution records of southern African Pteridophyta. J.E. BURROWS and

S.M. BURROWS 205

Zamiaceae. Encephalartos relictiis: a new species from southern Africa. P.J.H. HURTER and H.F.

GLEN 197

7. Taxonomic significance of inflorescences, floral morphology and anatomy in Passerina (Thymelaea-

ceae). C.L. BREDENKAMP and A.E. VAN WYK 213

8. Miscellaneous notes;

Aloaceae. Are ovules and seeds in Lomatophyllum Willd. (Aloe sect. Lomatophyllum sensu auct.)

anatropous and exarillate? E.M.A. STEYN and G.F. SMITH 237

9. Obituary; Heinrich Johann Wilhelm Giess ( 1910-2000). HERTA KOLBERG 241

10. Book review 245

11. National Botanical Institute South Africa; administration and research staff, 31 March 2001, publications

1 April 2000-31 March 2001. Compiler; B.A. Momberg 247

12. Guide for authors to Bothalia 265

New species, subspecies and combinations in Bothalia 31,2 (2001)

Androcymbium asteroides J.C. Manning & Goldhlatt, sp. nov., 203

Encephalartos rdictus P.J.H.Hnrter, sp. nov., 197

Erica blaerioides E.G.H.Oliv., sp. nov., 163

Erica blaerioides subsp. hirsuta E.G.H.Oliv., subsp. nov., 165

Erica chionodes E.G.H.Oliv., sp. nov., 161

Erica dolflana E.G.H.Oliv., sp. nov., 155

Erica oreotragus E.G.H.Oliv., sp. nov., 160

Erica taylorii E.G.H.Oliv., sp. nov., 157

Freesia fucata J.C. Manning & Goldhlatt, sp. nov., 189

Ixia atrandra Goldhlatt & J.C. Manning, sp. nov., 191

Melolobium lampolobum (E.Mey.) A.Moteetee & B-E.van Wyk, comb, nov., 210

Riccia sibayenii Perold, sp. nov., 151

Sebaea amicorum I.M.Oliv. <&. Beyers, sp. nov., 207

Tritoniopsis hicoXor J.C. Manning & Goldhlatt, sp. nov., 175

Tritoniopsis flava J.C. Manning & Goldhlatt, sp. nov., 178

Tritoniopsis toximontana J.C. Manning & Goldhlatt, sp. nov., 178

Bothalia31,2: 151-154(2001)

Studies in the genus Riccia (Marchantiales) from southern Africa. 25.

A new species in subgenus Ricciella, section Ricciella

S.M. PEROLD*

Keywords: Marchantiales, Riccia sibayenii Perold, section Ricciella (A.Br.) Bisch., subgenus Ricciella, southern Africa

ABSTRACT

Riccia sibayenii Perold is a new species, recently collected in Mpumalanga, near Sibayeni and has been named for this

town, which is close to the Mozambique border and is in the summer rainfall area. R. sibayenii is distinguished by lime-

green, medium-sized to quite large thalli with a chlorophyllose dorsal epidermis, air pores that soon enlarge and lead into

mostly 6-sided air chambers below; ventrally with a thin median layer of storage tissue; ventral scales absent and the

rhizoids all smooth. The spores are winged and polar, the distal face completely or incompletely reticulate and the proxi-

mal face ornamented with fine to coarse, sometimes spinose granules.

Riccia sibayenii Perold, sp. nov.

Thalli (iioici, annui, subdense turbati. rosulas non for-

mantes, prasini sine ullo colore purpureo, ramis usque ad

16 vel 17 mm longis, 1.05-3.75 mm lads, oblongis vel

apicem versus angustatis. Cavemulae supra ports apert-

entes, apicem versus parvis, proximaliter maiortbus, aetate

disintegrantes; tela penarta solum in parte media cartnata,

4—6 cellulas profunda. Squamae ventrales absentes, rhi-

zoideae omnes glabrae. Capsidae exsertae, ventraliter ape-

rientes. Sporae 77.5-100.0 pm diametro. trtangulart-glo-

bosae, atrobrunneae, alls cum duabus sertebus granularum

secus marginem, anisopolares, superficie distali cum alve-

olis completis vel incompletis, cristis cum duabus sertebus

granularum grossarum asperatis; superficie proximali cum

nota triradiata prominenti, in quaque superficula cum

granulis tenuibus vel grossis dispersis omata. Numerus

chromosomatum ignotus.

TYPE. — Mpumalanga, 2531 (Komatipoort): ± 10 km

south of Komatipoort, between the Kaftan fence and

Komati River, near Sibayeni town, altitude 270 m, on

black loamy soil, (-DB ), 2000-07- 1 2, Jean J. Meyer 2875

(PRE, holo.). Figure 1.

Thalliis medium-sized to fairly large, annual, quite

densely massed in patches, not forming rosettes, some-

times with branches overlying, occasionally covering old

thalli from previous season, usually closely adherent to

substrate, lime-green, without any purple coloration;

branches never simple, mostly twice to several times fur-

cate (Figure 2A), total length up to 16-17 mm long, seg-

ments narrowly to moderately divergent, 3-7 x

1.05-3.75 mm, 700-1250 pm thick medianly, thinner

toward winged margins, in section 1.5-3 times wider

than thick; oblong to tapering toward apex, shortly once

or twice emarginate, flat to slightly concave above, not

medianly grooved, margins winged, blunt, rather irregu-

larly scalloped, flanks sloping obliquely, ventral face

rounded and keeled, giving rise to perennating stolons.

* National Botanical Institute, Private Bag XIOI, 0001 Pretoria.

MS. received: 2001-03-02.

scales absent; when dry, margins sometimes slightly

raised, not inflexed.

Dorsal epidermis (Figures 2B; 3B) covering air cham-

bers flat, delicate and translucent, cells unistratose, 4—6-

sided, 50-80 x 42.5-67.5 pm; air pores distally small

(Figure 3A), soon enlarging to 80-125 x 50-75 pm or

more in older parts, irregular in shape, bordering cells not

differentiated (Figure 3C), breaking down with age and

air chambers below becoming exposed. Assimilation tis-

sue up to 875 pm thick, occupying most of thickness of

thallus, air chambers 12-14 across width of thallus, from

above mostly penta- or hexagonal, but in cross section

polygonal and variable in size, the enclosing chlorophyl-

lose, unistratose walls with cells 80.0-87.5 x 75-100 pm,

vertically to obliquely arranged, here and there intersect-

ing, air chambers then in 2 or 3 layers, except at margins,

where only 1 layer deep; storage tissue confined to medi-

an keeled part, 4—6 cell layers or 200-250 pm thick, cells

dorsiventrally flattened, 50.5-62.5 x 77.5-102.5 pm,

sometimes with smaller cells wedged in between. Ventral

scales absent. Rhizoids all smooth, 15-35 pm wide.

Dioicous. Antheridia serially arranged in 2 or 3 rows

(Figure 3D) and submerged along middle of thallus

FIGURE 1. — Locality of Riccia sibayenii in southern Africa.

152

Bothalia 31.2 (2001 )

FIGURE 2. — Riccia sihayenii. A, plant with several branches; B, epi-

dermal cells and air pore (hatched) overlying one air chamber,

the other partly exposed, as seen from above; C, c/s male

branch through air chambers and antheridial cavity; D, c/s near

apex of female branch with air chambers and capsule; E, c/s

nearer to base of female branch with air chambers and capsule.

A-E, ./../. Meyer 2H75. Scale bars: A, 2 mm; B, 50 pm; C-E, I

mm.

(Figure 2C), cavities 330-350 x 310-410 jam, necks aris-

ing above from small hollows and projecting up to 230

pm above thallus. Archegonia arranged in a row along

middle of segments, sometimes obliquely orientated,

necks ± 160 pm long (Figure 3F), dark red. Capsules

protruding and opening ventrally (Figure 2D, E), up to 6

in an acropetal row along branch, 450-575 pm diam.,

hardly visible from above (Figure 3E), containing up to

± 350 spores each, those in the youngest capsule near

apex sometimes still immature. Spores 77.5-100.0 pm

diam., triangular-globular, anisopolar, dark brown, semi-

transparent to opaque; wing ± 7.5 pm wide, at marginal

angles width ± 10 pm and perforated by single large pore

(Figure 4D), margin finely crenulate with 2 rows of gran-

ules, one along edge and the other immediately inward;

ornamentation on the 2 faces quite different; distal face

(Figure 4A-D) completely to incompletely reticulate:

with up to 5 ± complete alveoli, 12.5-17.5 x 12.5-22.5

pm, across distal face and totalling ±15, otherwise with

3 or 4 larger, central alveoli, ± 25 x 17.5 pm, surrounded

by smaller ones that may be confluent or separated,

ridges ± 5 pm high, thick and roughened by 2 rows of

granules, coarser than marginal ones, not raised at nodes

and not extending across wing; proximal face (Figure

4E, F) with prominent triradiate mark ± 2.5 pm high, all

or some arms extending across wing, each facet orna-

mented with scattered, fine to coarse granules, some single

and spinose, others joined together. Chromosome num-

ber. unknown, as material no longer living.

Thallus sections from the dry plants that were soaked

up, proved quite difficult to prepare. Hopefully, living

plants will still come to hand in the future. So far, the

species has only been collected once (Figure 1). It grows

on black, loamy soil (pH7) derived from weathered

basalt of the Letaba Formation (Lebombo group) of the

Karroo Sequence (Visser et al. 1984), in full sun on a

gentle slope in a valley that becomes waterlogged during

the summer months, when most of the rain falls. The

vegetation type of vascular plants in the area is Lebombo

Arid Mountain Bushveld (Low & Rebelo 1996).

The locality is described as disturbed. As it is close to

the Mozambique border, there has been an influx of peo-

ple fleeing from the war and its aftermath. It is possible

that the recent floods in the wake of cyclones Hudah and

Eline, may have spread the spores of this species further

southward, as it must be more widespread than this sin-

gle, though quite copious collection suggests. The area

is, however, very undercollected as far as bryophytes are

concerned, in fact, none have previously been recorded

for the quarter degree grid 253 IDB.

Although related to R. rubricollis, which was collect-

ed by Duthie in 1929 at Knysna, this new species differs

from it by being more delicate and by totally lacking any

purple colouration due to anthocyanin pigmentation on

the dorsal surface of the thallus, as well as in the

antheridial necks. Furthermore, the spores of/?, sibayenii

have double rows of granules along the wing margins

and on the alveolar walls on the distal face. The proximal

face is ornamented with spinose granules and not with

smooth-walled alveoli, which are present in R. rubricol-

lis on both faces (Perold 1991). In addition, R. sibayenii

grows in the subtropics, which has summer rain, not in

Bothalia 31.2 (2001)

153

FIGURE 3. — Riccia sibayenii. A. distal branches of thallus with air pores gradually increasing in size more proximally; B. pores and dorsal epi-

dermal cells; C. single pore and surrounding dorsal cells; D. 2 or 3 rows of antheridia near apex of male branch; E. toward apex of female

branch with capsule only faintly visible from above; F, protruding archegonial neck (no longer erect). A-F, J.J. Meyer 2875. A, x 8.3; B,

X 58; C. X 202; D. X 20; E, X 18.4; F, X 133.

FIGURE 4. — Riccia sibayenii. Spores. A-C. distal face; C, side view; D, part of distal face, wing margin and pore; E, proximal face with triradi-

ate mark; F, side view of proximal face. A-F. J.J. Meyer 2875. A. x 418; B, x 500; C, x 514; D. x 661; E, x 403.7; F, x 399.

the temperate southern Cape, known for its fynbos and

rain almost all year round.

Because its sporangia protrude and open ventrally, R.

sibayenii is referred to subgenus Ricciella (A.Br.) Bisch.,

section Ricciella, together with R. stricta and R. piirpuras-

cens. R. nibricollis was referred to subgenus Ricciella,

section Spongodes, because the sporangia are deeply em-

bedded in the thallus and open dorsally (Perold 1995).

ACKNOWLEDGEMENTS

1 wish to sincerely thank Dr S. Jovet-Ast for kindly ref-

ereeing this paper, as well as my colleague, Mr J.J. Meyer,

154

Bothalia 31,2 (2001 )

for collecting this new species. Dr H.F. Glen is thanked for

translating the diagnosis into Latin. I also extend my grati-

tude to Ms G. Condy for the drawings, to Mrs A.

Romanowski for developing and printing the photographs,

and to Ms D. Maree for typing the manuscript.

REFERENCES

LOW. A.B. & REBELO, A.G. 1996. Vegetation of South Africa,

Lesotho and Swaziland. Department of Environmental Affairs

& Tourism, Pretoria.

PEROLD, S.M. 1991. Studies in the genus Riccia (Marchantiales) from

southern Africa. 22. R. nihricollis, now validated, typified and

described. Bothalia 21: 51-54.

PEROLD, S.M. 1995. The taxonomic history of the Ricciaceae

(1937-1995) and a classification of sub-Saharan Ricciae.

Bothalia 25: 211-231.

VISSER, D.J.L. et al. 1984. Geological map of the Republics of South

Africa, Transkei, Bophuthatswana, Venda and Ciskei and the

Kingdoms of Lesotho and Swaziland. The Government Printer,

Pretoria.

Bothalia 31.2: 155-165 (2001)

Five new species of Erica (Ericaceae) from the Swartberg Range,

Western Cape, South Africa and a note on E, esterhuyseniae

E.G.H. OLIVER* and I.M. OLIVER*

Keywords: Erica L., taxonomy, new species. South Africa, Swartberg, Western Cape

ABSTRACT

Descriptions are provided of five new species of Erica L, from the Klein and Groot Swartberg Mountains in Western Cape,

South Africa. The two known populations off. dolfiana E.G.H.Oliv. are confined to the stony, southern slopes of two well-

separated areas. E. taylorii E.G.H.Oliv. is also known from two well-separated populations on the Swartberg but with the

others in the Cederberg. The other three new species are allied to E. esterhuyseniae Compton: E. chionodes E.G.H.Oliv.,

with brilliant white flowers, occurs only in seepage zones in two well-separated areas, E. oreotragus E.G.H.Oliv. (syn. E.

esterhuyseniae Compton var. trimera Compton) with pale pink to white flowers and E. blaerioides E.G.H.Oliv. with white

flowers, are widespread in the central and western Swartberg Range.

I. Erica dolfiana E.G.H.Oliv., sp. nov., in genere

affmitate dubia, sed foliis quadrinatis petiolo longo

bractea bracteolisque comparate longis plus minusve

sepala aequantibus sed remotis, corolla urceolata glabra

alba parva, antheris glabris calcaribus longis tenuibus fil-

amentis tenuibus, stigmate subpeltata ad peltata semi-

nibus vulgo ericoideis reticulatis dignoscenda. Eigura 1 .

TYPE. — Western Cape, 3322 (Oudtshoom); Oudts-

hoom, Groot Swartberg, peak just north of Tierberg,

2 010 m, (-AD), 17 October 2000, E.G.H. & I.M. Oliver

11688 (NBG, holo.; BM, BOL, K. MO, NY, P, PRE, S).

Erect, compact shrub up to 300(-500) mm tall, single-

stemmed reseeder. Branches', main branches up to 50 mm

long, erect, with numerous secondary branches 2-5 mm

long on every node towards apex and a few sometimes

distally, all ending in an inflorescence; all branches with

short, spreading to reflexed hairs with or without a few

stouter, longer gland-tipped hairs admixed. Leaves 4-nate,

subspreading, imbricate, ovate to elliptic, 2. 0-3. 3 x

1.0-1. 3 mm, adaxially flattened and abaxially rounded

with acute margins, sparsely puberulous with a few long,

stouter gland-tipped hairs interspersed abaxially, ciliate

with a few long, gland-tipped hairs, sulcus narrow and

open at base; petiole ± 1.3 mm long, puberulous.

Inflorescence: flowers 4-nate in 1 whorl at apex of main

and secondary branches, these packed in terminal, capi-

tate or longer spike-like synflorescences; pedicel ± 2.2

mm long, loosely puberulous, white to pale green, some-

times tinged red; bract partially recaulescent from basal to

middle position, oblong, ± 1 .4 x 0.4 mm, with very short

subapical sulcus, white and slightly hyaline, sometimes

with pale green tip, sparsely puberulous in upper 'U-fi,

margins with gland-tipped hairs and one apically, glands

white or pink; bracteoles 2, median to just below calyx in

position, otherwise like bract. Calyx 4-lobed, fused ± Vg

at base; lobes adpressed to corolla, oblong, ± 2.0 x 0.5

mm. subacute, sulcus in upper half, short and narrow.

* Compton Herbarium. National Botanical Institute, Private Bag X7,

7735 Claremont. Cape Town.

MS. received: 2(301-04-02.

white to pale green or reddish, more so at apex, puberu-

lous in upper 'A-'A, margins with short gland-tipped

hairs, glands red. Corolla 4-lobed, campanulate to slight-

ly urceolate, ± 3.4 x 2.2 mm, glabrous, white or tinged

pink; lobes ± 0.7 x 1.3 mm, rounded to emarginate, mar-

gins entire to slightly erose. Stamens 8, included, free; fil-

aments narrowly linear, ± 1.8 mm long, with slight apical

S-bend, glabrous, white; anthers bipartite, oblong in

adaxial view, dorsally attached near base, appendiculate;

thecae erect adpressed, ovate to elliptic in lateral view, ±

0.9 X 0.5 mm, smooth dark brown, spurs narrowly lance-

olate, ± 0.6 mm long, with 2 or 3 teeth, white, pore ± 'A

length of theca; pollen in tetrads. Ovary 4-locular, broad-

ly obovoid-ellipsoid, ±1x1 mm, emarginate, puberu-

lous, green, with small dark-green nectaries around base;

ovules 9-15 per locule, laterally spreading; style ± 2.7

mm long, exserted, pale greenish red, glabrous; stigma

subpeltate to peltate, dark red. Emit a dehiscent capsule,

± 1.1 X 1.5 mm, valves splitting for V4 their length and to

± 45° angle, septa equally divided on columella and

valves. Seeds ellipsoid, ± 0.4 x 0.25 mm, slightly alveo-

late, orange, testa cells 60-75 x 25-30 pm, anticlinal

walls jigsawed, inner periclinal walls pitted. Elowering

time: October to early December. Figure 1 .

Erica dolfiana is characterized by 4-nate leaves with

relatively long petioles, relatively large bract and bracte-

oles about the size of the sepals, but remote from them,

sepals white with long-stalked red marginal glands, urce-

olate glabrous white corolla, which is relatively small for

the genus, glabrous anthers with long narrow appendages

and thin filaments, enlarged subpeltate to peltate stigma,

and the standard reticulate/alveolate ericoid-type seed.

E. dolfiana shows no close affinity to any other species.

It may, however, be related to E. lignosa H.A. Baker, a

widespread Swartberg species, and to E. jugicola

E.G.H.Oliv. ined., a more restricted new species. Both have

similar long-stalked red glands on the bract, bracteoles and

sepals, but both have larger, pink flowers (very much larg-

er in the latter) which are mostly finely hairy. They also

have larger anther appendages and a small simple stigma.

All three species are sympatric on the Blesberg.

156

Bothalia 31,2 (2001 )

FIGURE I. — Erica dolfiana. A, flowering branch, natural size; B, stem with leaves removed; C, leaf; D, flower; E, bract; F, bracteole; G, sepal;

H, flower opened laterally to show androecium and gynoecium with ovary opened laterally; 1, stamen, back, side and front views; J, gynoe-

cium; K, capsule; L, seed; M, testa cells. A-J, drawn from the type collection, Oliver & Oliver 11688-, K-M, from Oliver 11576. Scale

bars: B-K, 2 mm; L, 1 mm; M, 50 pm.

This new species is named after Dolf Schumann

(191 8-) senior author of that very fine illustrated book,

Ericas of South Africa (Schumann & Kirsten 1992). He

made collections of this and the next species on a joint

trip with Jan Vlok in 1985. This is the second species of

Erica that is named after him, the other being E. schu-

mannii E.G.H.Oliv. (Oliver & Oliver 1998a).

Erica dolfiana is known thus far from only two locali-

ties— the small peak just north of Tierberg and the ridges

west and east of Blesberg, some 40 kms to the east

(Figure 2). In the latter locality, Pienaar, on the label of

his specimen 67, noted that west of the peak the species

was fairly common on the upper northern slopes. Having

visited Blesberg peak in July and January, out of the

flowering season, we too saw numerous young plants but

only in fruit.

The species is confined to southern and northern

slopes at high altitudes where it grows in rocky/stony

places with very short alpine fynbos. It is very common

in the type locality where it is the dominant species of

Erica. On the open slopes the shrublets, together with the

other plants, are small and compact, whereas the few

FIGURE 2. — Known distribution of

Erica dolfiana, ★; E. tay-

lorii, O.

Bothalia 31,2 (2001)

157

shrubs, among larger rocks in rocky areas, are looser and

up to 500 mm tall.

Vlok noted that the flowers gave off puffs of pollen

when disturbed. This was confirmed by us. The dull

white colour of the flowers renders them relatively

insignificant and, with the slightly enlarged stigma and

somewhat reduced nectaries, suggest both insect- and

wind-pollination strategies for the species (Rebelo et al.

1985). However, the stickiness of the flower heads,

caused by the glands on the bracts, bracteoles and calyx

segments, may discourage foraging insects from clam-

bering around the inflorescences.

Erica dolfiana would seem to be another of those

species in transition from insect- to wind-pollination,

with its small white or dull-coloured flowers as found in

so many other wind-pollinated Erica species. In our

experience, size differences between the flowers of

species point to different pollination syndromes, hut not

necessarily to different affinities. The other two unde-

scribed species in this possible alliance have been

observed by us to be pollinated by long-tongued flies.

Paratype material

WESTERN CAPE. — 3322 (Oudtshoom): Great Swartberg, peak

due south of highest point of jeep track, 2 000 m, (-AD), 5-11-1985.

Schumann 390 (NBG); peak behind Tierberg, 5800 ft. (-AD), 5-11-

1985, Vlok 1266 (K. NBG, NY. PRE); Blesberg. neck just E. 2 000 m,

(-BC), 13-07-2000, fruiting, Oliver 11576 (NBG); ibid., 6-01-2001,

fruiting, Oliver 11764 (NBG); ibid., neck just W, 5900 ft. (-BC). 12-11-

1976, Pienaar 67 (NBG).

2. Erica taylorii E.G.H.OIiv., sp. nov., pilis longis

in marginibus foliorum, bracteae, bracteolarum et sepa-

lorum, bractea bracteolisque magnis subpetaloideis,

antheris calcaribus brevibus crassis, ovario ramisque

puberulis dignoscenda. Figura 3.

TYPE. — Western Cape, 3322 (Oudtshoom): Oudts-

hoom, Groot Swartberg, peak just north of Tierberg,

2 010 m, (-AD), 17 October 2000, E.G.H. & l.M. Oliver

11691 (NBG, holo.; BM, K, MO, NY, P, PRE).

Erica oresigena Bolus var. mollipila Bolus in Guthrie & Bolus: 129

(1905). Types: Worcester Div., on the Matrons Berg, Bolus s.n.

(BOL?); Clanwilliam Div.; Sneeuwkop. Cederberg Range near

Wuppertal, Bodkin sub Bolus 6492 (BOL!).

Illustration.; Schumann & Kirsten: 112, fig. 49. 50 (1992).

Low, prostrate, spreading shmblet up to 200 mm tall

and 0.5(-l,2) m across, single-stemmed reseeder.

Branches: main branches mostly prostrate, numerous old

and woody; numerous secondary branches ± 100-300

mm long, erect, with occasional ±5-10 mm long tertiary

branches, all ending in a florescence, intemodes very

short, all covered with fine dense short retrorse hairs,

occasionally interspersed with few plumose hairs on

older branches. Leaves 3- or 4-nate, imbricate, oblong,

3-5 X 1 mm, erect, adaxially flattened, abaxially round-

ed, with acute margins, finely hairy on both sides and

with short, stout, plumose hairs on margins, sulcus nar-

row, closed at base; petiole 0.6-0. 8 mm long, pubemlous

and edged with a few short, stout, plumose hairs.

Inflorescence: flowers 3 or 4 in l(-3) whorls, terminal on

secondary and tertiary branches; pedicel 2. 0-3. 5 mm

long, finely hairy with retrorse hairs and occasionally

short, stout, plumose hairs interspersed, reddish; bract

partially recaulescent from 'A-'A way up pedicel, oblong

to lanceolate, 3. 5-4. 5 x 1.0-1. 2 mm, cream to reddish

pink, sometimes with green mainly in upper half,

glabrous to finely hairy, with long, plumose marginal

hairs, sulcus narrow, Vz length of bract; bracteoles 2,

placed Y4 way up pedicel, 3.0 x 0. 7-1.0 mm, otherwise

like bract. Calyx 4-partite, segments slightly imbricate,

broadly lanceolate to ovate, 3. 5^. 5 x 1.0-1. 5 mm, red-

dish pink, glabrous to finely hairy, margins with long,

plumose hairs, sulcus '/z length of segment. Corolla 4-

lobed, 6-9 x 3-5 mm, ovoid-urceolate, finely puberulous

to glabrous, pink to dark pink, hairs white; lobes spread-

ing-recurved, rounded, 1. 5-2.0 x 1.8-2. 5 mm, erose.

Stamens 8, included, free; filaments linear, ± 4 mm,

slightly widening to base, with slight bend below anther,

glabrous, white; anthers bilobed, narrowly ovate to

oblong in adaxial view, dorsally attached near base,

appendiculate; thecae ovate to elliptic in lateral view,

1 .0-1.2 X 0.5-0.6 mm, erect, brown, very finely aculeate,

pore small V4-V5 length of theca, spurs subspreading,

abaxially ± 0.6 x 2.5 mm, occasionally toothed, pink;

pollen in tetrads. Ovary 4-locular, broadly obovoid to

globose, 2-3 x 2-3 mm, emarginate, covered with dense,

short hairs, red, with dark nectaries around base; ovules

40-50 per locule, spreading laterally from full-length

placenta; style included, + 4.5-5 .0 mm long, glabrous,

dull red; stigma capitate, dark red. Emit a dehiscent cap-

sule. ± 2.5 X 3.5 mm, valves splitting Y4 their length and

to 45° angle, septa only on valves, placenta large, con-

voluted and warty. Seeds ellipsoid, ± 0.6 x 0.35 mm,

shallowly reticulate-alveolate, brown; testa cells ± 100 x

40 pm, anticlinal walls jigsawed, inner periclinal walls

densely pitted. Elowering time: October to December.

Figure 3.

In Elora capensis Bolus (1905) described the variety

mollipila of his E. oresigena, based on collections from

Matroosberg and the Cederberg. Hugh Taylor, who stud-

ied the vegetation of the Cederberg, expressed surprise,

since he knew E. oresigena and did not think that Bolus’

variety mollipila belonged to the same species. It is a

mystery to us why Bolus did not describe the variety

mollipila as a separate species, since our studies have

shown that the two taxa differ in many critical characters

which indicates that this taxon is more closely related to

species other than E. oresigena. The only similar charac-

ters are those of the anthers. E. oresigena is an upright,

woody, rounded bush 0.5-1. 5 m tall. It is distinguished

by its long pedicels, entire corolla lobes, short, woolly

hairs on the pedicel and sepals, large dark red glands on

the margins of the bract, bracteoles, sepals and young

leaves, and by the ovary which is glabrous.

Erica taylorii, here described de novo as a distinct

species, is related to the group of species which is char-

acterized by plumose hairs and large, coloured, petaloid

bract and bracteoles. The group includes the E.

goatcheriana L. Bolus complex, E. modesta Salisb. and

the recently described E. schumannii E.G.H.OIiv. (Oliver

& Oliver 1998a) and E. kirstenii E.G.H.01iv.(01iver &

Oliver 2000). E. taylorii differs in having very few, rela-

tively short (not numerous and dense), plumose hairs on

158

Bothalia 31,2 (2001)

FIGURE 3. — Erica taylorii. A, flowering branch, natural size; B, stem with leaves removed; C, leaf; D, flower; E, flower with corolla and ovary

cut open laterally; F, bract; G, bracteole; H, sepal; I, stamen, front, back and side views; J, gynoecium; K, seed; L, testa cells; M, typical

multiseriate plumose hair, A-J, drawn from the type collection, Oliver & Oliver II69I\ K & L, drawn from Oliver 10007a. Scale bars:

B-J, 2 mm; K, 1 mm; L, M, 100 pm.

the pedicel, and flowers in one, sometimes up to three

whorls. This latter character relates the species to two

species with plumose hairs and umbel-like inflores-

cences, namely E. cernua C.V.Montin and E. sphaero-

cephala J.C.Wendl. ex Benth. The former has a globose

corolla and larger anther appendages, whereas the latter

has much reduced bracteoles and sepals (not large and

petaloid), and long-aristate leaves, bract, bracteoles and

sepals.

This new species is named after the vegetation ecolo-

gist, the late Hugh Taylor (1925-1999), a colleague and

friend of many years who had a profound knowledge of

the Cape Flora (McDonald et al. 2()()0). He collected

many fine herbarium specimens, including one of this

species, made during his major vegetation survey of the

Cederberg Mountains.

Erica taylorii occurs in two widely separated areas,

the Swartberg Range above Ladismith and Oudtshoorn,

and the Cederberg Mountains east of Clanwilliam

(Figure 2). It is variable in the number of leaves per

whorl, number of flowers per inflorescence and in the

size of the corolla, which can be hairy or glabrous.

The Cederberg populations have 3- or 4-nate leaves,

whereas the Swartberg populations have only 4-nate

leaves. Most plants have a hairy corolla with the varia-

tion in indumentum not linked to any specific area. The

totally glabrous form occurs in Schumann 380 with the

Bothalia 31,2 (2001)

159

type collection having a few flowers with some hairs on

the corolla lobes and slightly more in Vlok 1263. The

Seweweekspoort populations are all finely puberulous.

All the collections from the Cederberg have a hairy

corolla, except for a few branches on Bond 1403, which

have very short hairs only on the corolla lobes.

The eastern populations of the species are confined to

the Swartberg Range in two widely separated areas, one

above Ladismith along the Klein Swartberg in the region

of Seweweekspoort Mtn, and the other some 90 kms to

the east in the Tierberg area above Oudtshoorn. It is sur-

prising that no collections have been made on any of the

numerous intervening peaks where suitable habitats

surely occur, despite visits to these peaks by Stokoe,

Esterhuysen, Taylor and ourselves. We have selected our

recently collected material near Tierberg as the type of

the species, since it provided adequate isotypes and

excellent fresh material for us to study.

The type population was very localized on a small,

very low rocky ridge on the southern slopes, just below

the summit of the minor peak north of Tierberg. A few

plants were also found on nearby rocky outcrops. The

plants grew in crevices on the rocks and between rocks

and spread out over the surface for up to half a metre.

They were woody and in some cases had gnarled stems.

The open slopes surrounding the low ridge were covered

with numerous plants of E. dolfiana (see above) in very

short, alpine-like vegetation.

On the peaks just west of Seweweekspoort, E. taylorii

is apparently more common, judging by the collection

records of ‘frequent’. It was collected there by Andreae,

Barnard, Primos and Stokoe on the first expedition to the

Klein Swartberg in 1928 (Linder et al. 1993). On Hoeko

Peak we noted numbers of low. spreading plants growing

among short restiads and in rock crevices on northern

slopes, but only in fruit. Andreae recorded the plants as

'low spreading shrublets on ground or adpressed to rocks

on flat open places’. Vlok noted that plants were ‘fre-

quent on southern slope between large rock blocks’,

which could explain why he recorded the height of the

plants as ± 1.2 m, if they grew up among the rocks.

The populations in the Cederberg Mtns occur on the

highest peaks from Krakadouw to Sneeuberg, a distance

of 30 kms. Pocock, who made numerous collections of

plants in the Cederberg in the mid- 1920s, noted on her

collection 650 ‘found on all the high peaks but nowhere

abundant except on Krakadouw Great Peak’.

While looking for E. cedromontana E.G.H.Oliv.

(Oliver & Oliver 1998b) on the Langberg, we searched

for E. taylorii too, but did not locate any plants. Thus

details of the habit and habitat are taken from other col-

lectors’ notes. Pocock recorded plants as ‘low shrubs

3-15 inches tall, growing on small vlaktes and crannies

among rocks on Krakadouw’ and ‘stunted shrub in

cracks in rocks’ on Sneeukop summit. Taylor recorded it

as growing ‘in small Ischyrolepis cur\’iramis sand flat on

bedrock’ and Stehle gave ‘on rocks’.

Esterhuysen gave a detailed record of the species on

the Sneeuberg when she collected some old fruiting

material in April. She wrote ‘plants were common and

conspicuous on the south side in rocky places — mostly

growing from rock crevices in flat or sloping rock and

spreading out over rocks forming low dense masses, pro-

tected from fires’. She also made the interesting obser-

vation that the plants were rooting along the stems.

The record by Harry Bolus of a Bolus s.n. specimen

from Matroosberg, a syntype of his E. oresigena var.

mollipila, was regarded by us as doubtful. This collection

was not filed under E. oresigena in BOL, but was noted

on the sheet as having been removed by Elsie

Esterhuysen to E. maderi where, unfortunately, it cannot

now be found. The collection must have consisted of a

flowering branch 100 mm long, deduced from the clear

imprint on the sheet where it had formerly been mount-

ed and annotated by Bolus, and certainly gives the

impression of being E. maderi, which is common on the

lower slopes of Matroosberg. A collection of that species

from Matroosberg by Alfred Bolus is housed in SAM.

The other syntype of the variety, which was collected by

Bodkin from the Cederberg, consists of three small

branchlets in poor condition with just six buds and six

mature flowers.

Paratype material

WESTERN CAPE. — 3219 (Wuppertal); Krakadouw Peak, summit.

5900-6700 ft. (-AA), Pocock 650 (NBG); Groot Krakadouw, summit

massif. 1 700 m, (-AA), 22 November 1987, Taylor 11905 (NBG);

Sneeukop, (-AC), Bodkin sub Bolus 6492 (BOL); ibid., 5500-6300 ft,

6-04-1969. fruiting, Esterhuysen s.n. (BOL); ibid.. 6300 ft. 13-10-

1923. Pocock 214 (NBG); Langberg, 6000 ft. 15-12-1941, Bond 1403

(NBG); ibid.. 15-12-1941, Esterhuysen 7306 (BOL, SAM); Sneeuberg

Peak, 6200 ft, (-AC), 2-\l-\969^ Stehle 310 (NBG); Wolfterg Arch

area. (-AD). 28-11-1959. Jessop sub Oliver 324 (NBG). 3321

(Ladismith); Toverkop. 6000-7000 ft, (-AC). 15-12-1956, Esterhuysen

26759 (BOL); Seven Weeks Poort Mtn, 2 300 m, (-AD), 26-12-1928,

Andreae 1181 (BOL, NBG); ibid.. 5000-7000 ft, 12-1928. Barnard in

SAM46306 (SAM); ibid., 1 800-2 100 m. 26-12-1928, Primos 16

(BOL. NBG); ibid., 7200 ft, 26-12-1928, Stokoe 1878 (BOL); ibid.,

7500 ft, 5-01-1981, Vlok 140 (NBG); Hoeko Peak, 1 900 m, (-AD), 4-

02-1992, fruiting flowers only, Oliver 10007a (NBG). 3322

(Oudtshoorn); Great Swartberg, peak north of Tierberg, 6250 ft, (-AD),

5-11-1985. Vlok 1263 (NBG); ibid.. 1 900 m, 5-11-1985, Schumann

389 (NBG. PRE).

THE ERICA ESTERHUYSENIAE COMPLEX

The following three new species form a complex with

E. esterhuyseniae Compton. They are all confined to

high altitudes along the Klein and Groot Swartberg

ranges and in some places grow sympatrically.

Erica esterhuyseniae was described by Compton in

1941 (Compton 1941). In the protologue he divided the

species into two varieties, var. a tetramera and var. 6

trimera based on the distinction of 4-nate versus 3-nate

leaves. According to the current nomenclatural rules he

should have retained the epithet ‘esterhuyseniae' under

the autonym rule for the var. a (Oliver & Oliver 1994).

We support Compton’s recognition of two separate

taxa. but with all the additional material available to us,

feel that there are sufficient grounds for separating them

at specific level. The number of leaves per whorl is a

clear distinction, but one which Compton felt was of

‘less systematic importance’ in E. esterhuyseniae. There

160

Bothalia 31.2 (2001)

FIGURE 4. — Known distribution of

Erica oreotragus, •, and E.

esterhuyseniae, O.

are, however, several other eharacters which he did not

notice and which can be used to distinguish the two taxa

(see below).

3. Erica esterhuyseniae Compton: 193 (1941) pro

E. esterhuyseniae var. a tetrarnera', Dulfer; 75 (1965); E.

esterhuyseniae var. esterhuyseniae, E.G.H.Oliv. &

I.M.Oliv.: 27 (1994). Type: Cape Province. Uniondale

Division: Kamanassiberg [Kammanassie] (Mannetjies-

berg), 1 700 m, 1-02-1941, Compton 70557 (NBG!).

Illustration: Compton: 194. fig. 3 (left), 1-12 (1941 ).

Erica esterhuyseniae is characterized by the 4-nate

leaves which are relatively broad, open-backed and

curve inwards, and the long, narrow, almost acuminate,

corolla lobes '13-/2 length of corolla. It is allied to E.

oreotragus and E. hlaerioides (see below).

The species occurs on the far-eastern portion of the

Swartberg Range around Blesberg, on the Karoo insel-

berg — the Kammanassie Mtns just to the south, and on

the western half of the Kouga Mtns (Figure 4). In all

localities it grows on the south-facing upper slopes in

sandy loamy soil, or in areas where there is more mois-

ture in winter, which is the case with our recent collec-

tion, 11799, from the drier foothill ridge just north of the

main range of Blesberg.

Specimens examined

WESTERN CAPE.— 3322 (Oudt.shoorn): Blesberg, (-BC), 17-10-

1955, Esterhuvsen 24917 (BOL); ibid,, 2 040 m, 13-07-2000 (fruiting),

Oliver 11549 (NBG); ibid,, 1 960 m, 7-01-2001, Oliver 11779 (NBG);

ibid., northern foothill ridge, 1 840 m, 7-01-2001, Oliver 11799 (NBG);

Great Swartberg at Kolberg, 1 500 m, (-BD), 20-03-1983, Schumann

93 (NBG); Kammanassie Mtns, Buffelsberg, 5000 ft, (-DB), 29-01-

1929. Viviers & Vlok 37 (NBG); Kammanassie Mtns, Mannetjiesberg,

(-DC), 1-02-1941, Compton /0557 (NBG); ibid., 5000 ft, 1-02-1941,

Esterhuysen 4762 (BOL, K, PRE); ibid., 1 800 m, 9-01-2001, Oliver

1 183 1 (NBG). 3323 (Uniondale): Hoopsberg, 5000 ft, (-CB), 12-03-

1966, Kourke 399 (NBG); Kouga Mtns, Saptoukop. 1 530 m, (-DA),

16-12-1991, Oliver 9938a (NBG).

4. Erica oreotragu.s E.G.H.Oliv., sp. nov., Ericae

esterhuy.seniae Compton alTinis sed ab ea foliis ternatis

angustioribus sulco minus aperto, lobis corollae angus-

tioribus, calcaribus antherarum latioribus differt. Figura 5.

E. esterhuyseniae Compton var. trimera Compton: 193 (1941);

Dulfer: 75 (1965); E.G.H.Oliv. & I.M.Oliv.: 27 (1994). Type: Cape

Province. Oudtshoorn Division: Swartberg Pass, 2 000 m, 28-12-1941.

Bond 866 (NBG, holo.!; BOL!, PRE).

Illustration: Compton: 194, fig. 3 (right), 1-10 (1941).

Frect, sparse to suberect and compact, shrublet, 300-

400 mm high, single-stemmed reseeder. Branches:

numerous main branches 50-100 mm long, terminating

in an inflorescence, internodes 2-10 mm long, occa-

sional short, secondary branches 2-5 mm long with or

without terminal inflorescence; all branches covered

with dense, short, spreading hairs or with short and long

hairs intermixed. Leaves 3-nate, erect incurved, broadly

ovate to elliptic, 2. 0-3. 2 x 0.8-1. 3 mm, subacute, adax-

ially flat, abaxially rounded, margins acute, densely

hairy on both surfaces, sulcus partially open, closed at

base; petiole ± 1 mm long, hairs all over or only on mar-

gin. Inflorescence: flowers 3-nate in 2, rarely 3, whorls

in umbel-like group at apices of main and secondary

branches; pedicel ±1.2 mm long, glabrous to finely and

shortly hairy; bract partially recaulescent, in mid posi-

tion on pedicel, narrowly lanceolate to linear, 0.7-1. 6 x

0.1 mm, variable in size in an inflorescence, glabrous

but ciliate along margin, esulcate, greenish red to

whitish; bracteoles 2, in mid position, minute or reduced

to just a tuft of hairs within an inflorescence, esulcate.

Ccdyx 4-partite, very slightly fused at base; segments

lanceolate to ovate, 1.5-1. 8 x 0.5 mm, subacute, puberu-

lous to glabrescent, ciliate with long, spreading hairs

and occasionally 1 or 2 minute subsessile to sessile

glands, sulcus fairly broad, 73-72 length of lobe. Corolla

4-lobed, urceolate, ± 3.3 x 2.0 mm, with 4 bulges at

base, densely hairy with short and long spreading hairs

admixed, pale pinkish to white; lobes spreading, ± 1 mm

long, broadly rounded, subacute to obtuse, margins

minutely crenulate to entire. Stamens 8, included, free;

filaments ± 1.5 mm long, widening towards base, with

marked apical S-bend; anthers bilobed, elliptic in out-

line in adaxial view, dorsally attached near base, appen-

diculate; thecae erect adpressed, narrowly ovate-elliptic

in lateral view, ± 0.8 x 0.3 mm, glabrous, smooth.

Bothalia 31.2 (2001) 15]

FIGURE 5. — Erica oreotragus. A. flowering branch, natural size; B, stem with leaves removed; C, leaf; D, flower; E, corolla, F, bract; G. bracte-

ole; H. sepal; 1. stamen, side, front & back views; J, gynoecium; K. ovary opened laterally. L, capsule; M. seed; N. testa cells. All drawn

from the type collection. Scale bars: B-H. 2 mm; 1-M. 1 mm.

brown, pore ± 'A length of theca; spurs ± 0.6 x 0.2-0. 1

mm, attenuate from broad flat base, unevenly serrate to

entire and shortly ciliate, brown; pollen in tetrads.

Ovary 4-locular, globose. ±1x1 mm, obtuse, densely

and shortly hairy, with distinct nectaries around base;

ovules ± 9 per locule. pendulous from placenta in upper

-/s; style included, terete, glabrous, white; stigma simple,

rounded, reddish. Fruit a dehiscent capsule, ± 1.7 x 2.2

mm. valves fused up to 7s their length spreading to 45°,

equally on valves and columella. Seeds ± 0.6 x 0.3 mm,

4-ridged, abaxially rounded, laterally and adaxially flat-

tened, reticulate, orange-yellow, testa cells 100-125 x

27 pm, with undulate thick anticlinal walls, periclinal

walls with a few very small pits. Flowering time:

December to March. Figure 5.

This new species and the other two new species

described here, namely E. blaerioides and E. chionodes,

are closely allied to E. esterhuyseniae. All three new

species, however, have 3-nate leaves, whereas in E.

esterhuyseniae they are 4-nate.

Erica oreotragus occurs on the Klein Swartberg and

on the western and central portions of the Groot

Swartberg (Figure 4) with no areas of overlap with E.

esterhuyseniae, which occurs much further east. It grows

sympatrically with E. blaerioides in the Botes Nek area

(see below).

The epithet for this, the commoner of the two former

varieties of E. esterhuyseniae, is derived from Oreo-

tragus oreotragus. the scientific name of the small klip-

springer buck which is often seen in the Cape folded

mountains. We who have the privilege of knowing the

remarkable Elsie Esterhuysen (1912-) and of having

been out in the mountains collecting with her, will

always have visions of a metaphorical klipspringer hop-

ping from rock to rock and darting around the slopes,

selecting choice plants for the collecting bag and press.

Elsie is in her element roaming these Cape mountains.

This epithet is chosen in fond remembrance.

Paratype material

WESTERN CAPE. — 3321 (Ladismith): Klein Swartberg. N slopes

of Hoeko Peak. 2 080 m, (-AD), 3-02-1992, Oliver 10004 (NBG. PRE);

Seven Weeks Poort Mtn. 7000 ft. (-AD), 12-1928. Primos 24 (BOL.

NBG, PRE); Calitzdorp area, Groot Swartberg in Botes Nek area.

6000 ft [1 830 m], (-BD), 31-12-1969. Oliver 3068 (NBG). 3322

(Oudtshoom): Swartberg Pass, 1 590 m. 6-02-1986, Brusse 4866 (NBG.

PRE); ibid., 5000 ft. 25-01-1941, Esterhuysen 4513 (BOL); ibid.,

5000 ft, 25-01-1941. Esterhuysen 4772 (BOL); ibid., 28-01-1961,

Esterhuysen 28806 (BOL, K. PRE); ibid.. S side, 1 500 m, (-AC), 01-

1979. Oliver 7407 (NBG); ibid,, 03-1942. Stokoe in SAM55440 (SAM).

Swartberg, without precise locality, 01-1935. Stokoe 6668 (BOL).

5. Erica chionodes E.G.H.Oliv., sp. nov., synflor-

escentibus longis densis spicatis, floribus niveis puberu-

162

Bothalia 31.2 (2001)

FIGURE 6. — Erica chionodes. A, tlowering branch, natural size; B, lateral flowering branchlet; C, stem with leaves removed; D. leaf; E, flower;

E. flower, half section; G, bract; H, bracteole; I, calyx lobe; J, stamen, back, front and side views; K, ovary and stigma; L, capsule with

one valve removed; M, seed; N, testa cells. All drawn from the type collection. Scale bars: B-E, L, 2 mm; G-K, M, 1 mm; N, 100 pm.

lis, ramis puberulis, foliis abaxialiter semipatentibus,

calcaribus antherarum longis angustis, ovario hinsuto

dignoscenda. Figura 6.

TYPE. — Western Cape, 3322 (Oudtshoorn): Oudts-

hoorn, Groot Swartberg, peak just north of Tierberg,

small kloof WNW of peak, 1 720 m, (-AD), 17 October

2000, E.G.H. & l.M. Oliver 11699 (NBG, holo.; BM,

BOL, K, MO, NY, P, PRE, S).

Illustration: pro E. esterhuvseniae, Schumann & Kirsten: 1 16. fig,

70.71(1992).

Erect, sparse to compact shrub up to 600 mm tall, sin-

gle-stemmed reseeder. Branches', main branches erect

30-50(-80) mm long, with determinate, occasionally

continuing growth; secondary branchlets very short, ±

1-2 mm long on every node; tertiary branchlets occa-

sional, ± 1 mm long on previous year’s secondary

branchlets; all branches pilose to hirsute with spreading

hairs. Leaves 3-nate, subspreading, narrowly lanceolate-

oblong, 2. 5-3. 5 X 0.4- 1.2 mm, adaxially Oat, abaxially

rounded with rounded margins, partially open-backed,

sulcus broad, open at base, pilose to hirsute all over

except in sulcus; petiole 0.5 mm long, pilose.

Inflorescence: Powers l(-3)-nate on ends of secondary,

and occasionally tertiary, branchlets, crowded along and

towards ends of main branches in den.se spike-like syn-

llorescences l00(-200) mm long; pedicel 1.7 mm long,

pilose, pale red; bract partially recaulescent about Ve way

up pedicel, narrowly lanceolate, 0.4 x 0.15 mm, not sul-

cate, pilose, white; bracteoles 2, like bract but placed

slightly higher up pedicel and slightly smaller. Calyx 4-

lobed, 0.9 x 0.5 mm, tube about 'A length of calyx; lobes

ovate, adpressed to corolla, with very short, subapical,

wide sulcus, puberulous, very pale green to white.

Corolla 4-lobed, urceolate, 2. 8-3. 8 x 2.0 mm, puberu-

lous, white; lobes 0.8 x 1.0 mm, rounded, erose. Stamens

8, included; filaments free, linear, ± 2 mm long, with

slight apical S-bend, glabrous, white; anther bilobed,

dorsally attached near base, ovate in adaxial view, appen-

diculate; thecae erect adpressed, ovate-elliptic in lateral

outline, ± 0.5 x 0.3 mm, aculeate, brown, spurs irregu-

larly linear-lanceolate, ± 0.7 mm long, slightly longer

than thecae, pendulous, with few lateral teeth, yellowish,

pore '/i-'/z length of theca; pollen in tetrads. Ovary 4-

locular, broadly obovoid, ± 1 .0 x 0.9 mm, rounded, fine-

ly hairy all over, green, with reddish green apical portion

and dark red nectaries around base; ovules ± 6 per locule,

pendulous from placenta in upper half of locule; style

manifest, 2 mm long, dull reddish; stigma truncate to

sub-capitulate, dull red. Fruit a dehiscent capsule, ± 1.5

X 1.8 mm, valves fused ± 'U at base, opening to ± 45°

angle; septa about equally .split on valve and columella.

Seeds ellipsoid, ± 0.5 x 0.4 mm, slightly flattened later-

ally, testa medium hard in texture, orange, reticulate-

alveolate, cells 60-75 x 35^5 pm, anticlinal walls

undulate to slightly jigsawed, inner periclinal wall pitted

Bothalia 31.2 (2001)

163

FIGURE 7. — Known distribution of

Erica chionodes. ■; E. blae-

rioides subsp. blaerioides,

•; E. blaerioides subsp. hir-

suta. O.

with fairly large pits occasionally coalescing. Flowering

time: September to November. Figure 6.

Erica chionodes is distinguisheti by the long, dense,

spike-like synflorescences with pure white, finely hairy

flowers, partially open-backed leaves, long, narrow, pen-

dulous anther appendages and hairy ovary. It is closest to

E. oreotragus E.G.H.Oliv. (see above) which is common

along the full length of the Swartberg Range and has

flowers in terminal 6-8-flowered heads on main branch-

es (not in spike-like synflorescences), glabrous, mostly

shorter pedicels, longer sepals which are about half the

length of the corolla and have fewer longer hairs, rela-

tively long corolla lobes, and anther appendages shorter

than the thecae and broader and hairy. It is also allied to

the new species, E. blaerioides (see below).

The species was studied by us in a large kloof on the

northwest side of Tierberg above the Cango Caves, Oudts-

hoom (Figure 7), where we saw just two populations.

Both are growing in seepage areas with a predominance

of low, matted species of Restionaceae. The profusion of

brilliant white flowers contrasted dramatically against

the dark restiads and was reminiscent of patches of snow,

hence the epithet, chionodes - like snow (Greek).

The type population was undoubtedly the one seen by

the first and only other collectors of the species, Vlok &

Schumann in 1985. In the kloof the species was common

and grew together with another very striking, but pink-

flowered species, E. nervata Guthrie & Bolus. The pop-

ulation was studied on a hot sunny afternoon when sev-

eral flies and beetles were seen to visit the flowers. The

flowers gave off no detectable scent. The population

lower down and across the valley (11705) was growing

in a pure stand of restiads in a seepage area close to the

main stream and had older flowers.

There is a record from Toverberg above Ladismith in

the Klein Swartberg by Esterhuysen (Esterhuysen

26768) — ‘S slopes near base of shale band, slightly

marshy’. No colour is recorded for the flowers. This

specimen fits the species, but is more compact in growth

with shorter synflorescences and has 4-nate leaves. The

large gap between the two areas is similar to that in the

previous species, E. taylorii, described here, and is also

surprising, although in this case the wet seepage habitat

is very much scarcer than the rocky one of the latter.

Paratype material

WESTERN CAPE. — 3322 (Oudtshoom): Swartberg Mtns. main

kloof running north just NW of Tierberg. sides of main stream.

1 400 m. (-AC), 17-10-2000. Oliver 11705 (E. NBG. W); next to track

near Tierberg. 4600 ft. (-AD). 5-11-1985. Vlok & Schumann 1264

(BM. BOL, k. NBG. NY. PRE).

6. Erica blaerioides E.G.H.Oliv., sp. nov., Ericae

oreotragus E.G.H.Oliv. affmis, sed ab ea synflorescen-

tibus breviter spicatis, bractea bracteola subaequanti dif-

fer!. Eigura 8.

TYPE. — Western Cape, 3321 (Ladismith): Calitzdorp

area, Groot Swartberg in Botes Nek area, 6000 ft

[1 830 mj, (-BD), 31 December 1969, Oliver 3049

(NBG, holo.; BM, BOL, K, NY, PRE).

Erect, compact, much branched shrublet to 0.3 m,

rarely 1.2 m tall, with thin twiggy branches, single-

stemmed reseeder. Branches: numerous, thin; main

branches 20-50(-100) mm long, with continuous apical

growth, secondary branches 1-5 mm long ending in an

inflorescence, all branches covered with simple short

spreading hairs, intemodes 1.0-1. 5 mm long. Eeaves 3-

nate, erect, slightly incurved, occasionally spreading

when older, lanceolate to elliptic, 1.7-2. 2 x 0. 6-0.8 mm,

adaxially flat, abaxially rounded, with subacute margins,

partially open-backed to narrowly sulcate, sulcus open at

base, puberulous to sparsely so when young with longer

apical tuft, glabrescent with age; petiole ± 0.5-0. 9 mm

long, glabrous, ciliate. Inflorescence: flowers 3 in 1

whorl at apex of secondary branches; pedicel ± 2. 0-3. 5

mm long, glabrous to puberulous; bract partially

recaulescent, way up pedicel, narrowly oblong to

lanceolate, 1.4 x 0.2-0.3 mm. white, glabrous or sparse-

ly puberulous, ciliate, sulcus very small or absent;

bracteoles 2, slightly above bract, otherwise same as

bract. Calyx 4-partite, very slightly fused at base; seg-

ments not imbricate, mostly adpressed to corolla, lance-

olate to broadly so, ± 1.4— 2.2 mm long, acute, white,

glabrous or puberulous, ciliate with hairs and a few non-

sticky short-stalked glands, sulcus ± V3 length of sepal.

164

Bothalia 31,2 (2001)

FIGURE 8. — Erica bUierioides subsp. htaerioides. A, tlowering branch, natural size; B, llowering branchlet with single inllorescence; C, stem

with leaves removed; D, two leaves; E, flower; F, flower, cut open longitudinally; G, bract; H, bracteole; 1, sepal; J, stamen, back, side and

front views; K, gynoecitim; capsule, with one valve removed; M, seed; N, testa cells. Subsp. hirsuta. O, leaf; P, flower; Q, stamen; R,

gynoecium. A-N. drawn from the type of the species, Oliver 3049\ O-R, from the type, Oliver 5529. Scale bars: B-1, O, P, 2 mm; J-M.

Q. R, I mm; N, 50 pm.

narrow. Corolla 4-lobed, narrowly urceolate, 3. 2-3.4 x

1 .7-2.0 mm, sparsely puberiiloiis to puberulous, white;

lobes SLiberect to recurved, narrowly deltoid to broadly

so, 0.6-0. 9 X 0.5 mm, subacute to rounded, margins

erose. Stamens 4-8, free, included; filaments oblong,

1 .2-1.5 X 0.2 mm, slightly widened in middle or towards

base, with marked apical S-hend, glabrous, white;

anthers bipartite, erect, lanceolate to oblong in adaxial

view, dorsal ly attached near base, appendiculate; thecae

erect adpressed, curved oblong in lateral view, 0. 6-0.9 x

0.2-0. 3 mm, aculeate, brown, pore ± '/i length of cell,

appendages irregularly shaped, 0. 3-0.6 x 0. 1-0.2 mm,

irregularly lacerate, puberulous; pollen in tetrads. Ovary

4-locular, broadly obovoid to globose, 0.8-1 .0 x 0.7 mm,

very slightly emarginate, puberulous to puberulous in

upper half only, with well developed nectaries around

ba.se; ovules 8-12 per locule, pendulous from placenta

almost the length of columella; style included, straight.

Bothalia 31,2 (2001)

165

1. 5-2.0 mm long, glabrous; stigma simple truncate. Fruit

a dehiscent capsule, valves splitting nearly to base and to

± 45°, delicate, not hard and woody, septa only on the

valves. Seeds elongate ovoid, ± 0.6 x 0.3 mm, orange,

testa cells 75-100 x 25 pm, anticlinal walls slightly

thickened, straight to very slightly and irregularly undu-

late, no pits in inner periclinal walls. Figure 8.

The species occurs at high altitudes only in the west-

ern parts of the Groot Swartberg Range and eastern Klein

Swartberg (Figure 7).

Erica blaerioides is closely related to the previous

two species. It has spike-like synflorescences similar to

those of E. chionodes, but differs in these being looser

and much shorter, in having longer sepals, a differently

shaped corolla which is more sparsely hairy with longer

hairs, differently shaped ovary, stamens in a lower posi-

tion in the corolla and very different seeds. It is also

allied in many respects to E. oreotragus, but differs in the

form of its inflorescence which is spike-like and not

umbel-like, and by its subequal bract and bracteoles.

This new species is divided into two subspecies on a

number of characters, which show slight disjunctions —

stamen number, size of the leaf sulcus, hairiness of the

bract, bracteoles and sepals, and size of the sepals and

anther appendages. The two taxa are allopatric. The type

of the species and typical subspecies has been chosen

from among the field collections made by the first author

in the eastern populations.

subsp. blaerioides

Stamens always 4; bract, bracteoles and sepals

glabrous but ciliate; leaves partially open-backed; longer,

narrower sepals (1. 6-2.0 mm long); broader, larger

anther appendages (Figure 9A-N). Flowering time:

December and January.

The typical subspecies has been recorded from the

summit ridge of the Groot Swartberg from Botes Nek to

above Kliphuisvlei (Figure 7) where it occurs on cool

south-facing slopes. In the type locality the species

occurred sympatrically with E. oreotragus and was easi-

ly recognizable as a distinct species mainly on the differ-

ences in the inflorescence types.

subsp. hirsuta E.G.H.Oliv., subsp. nov., a sub-

specie typica foliis minus sulcatis, omnibus partibus

floris (staminibus exclusis) puberulis, sepalis brevioribus

et latioribus (± 1.4 mm longis), staminibus 4-8 cal-

caribus angustioribus differt. Figura 90-R.

TYPE. — Western Cape, 3321 (Ladismith): Klein

Swartberg, Seweweekspoort peak, 7500 ft [2 280 m],

(-AD), 5-01-1981, Vlok 139 (NBG).

This subspecies is distinguished from the typical sub-

species in having leaves not quite as open-backed, all

parts of the flower (excluding stamens) puberulous,

shorter and broader sepals (±1.4 mm long) and 4-8 sta-

mens with narrower appendages (Figure 90-R).

Flowering time: December and January.

The subspecies has been recorded from the high alti-

tudes of the eastern end of the Klein Swartberg

Mountains Just west of Seweweekspoort (Figure 7)

where it has been recorded growing with E. oreotragus.

It was first collected by two of the first group of botani-

cal explorers of that rugged high area in December 1928,

Andreae and Stokoe (Linder et al. 1993) (see also E. tay-

lorii above). This area is about 40 kms west of the popu-

lations of subsp. blaerioides. Other than the peak just

east of Seweweekspoort which we have investigated, the

intervening mountain range has not to our knowledge

ever been botanised. However, this range decreases in

altitude down to the large Gamkaskloof and would there-

fore most probably not provide the type of high altitude,

cool slopes required by the species.

Paratype material

subsp. blaerioides

WESTERN CAPE. — 3321 (Ladismith): Oudtshoom, Groot Swart-

berg, summit ridge above Kliphuisvlei. 1 900 m, (-BD). 1-01-1975,

Oliver 5529 (NBG, PRE); ibid., 6500 ft [1 980 m], 17-01-1954, Taylor

1087 {BOD.

subsp. hirsuta

WESTERN CAPE. — 3321 (Ladismith): Klein Swartberg, Klein

Swartberg Peak [?Seweweekspoortberg], 6700 ft [2 040 m], (-AD), 28-

12-1928, Andreae 1280 (BOL, PRE); ibid., 6800 ft [2 070 m], 28-12-

1928, Stokoe 1768 (NBG. PRE), as 1768a (BOL, PRE); Sewe-

weekspoort Peak, (-AD), 12-1928, Stokoe 6669 (BOL, K, PRE).

REEERENCES

BOLUS, H. 1894. Contributions to the Flora of South Africa. 1. Erica-

ceae. Journal of Botany 1894: 234—290.

GUTHRIE. F. & BOLUS, H. 1905. Erica. In W.T. Thistleton-Dyer, Flora

capensis 4: 4—315.

COMPTON, R.H. 1941. Plantae novae africanae. Journal of South

African Botany 1\ 193-195.

DULFER. H. 1965. Revision der sUdafrikanischen Arten der Gattung

Erica L. 2. Annalen des Naturhistorischen Museums in Wien

68: 25-177.

LINDER. H R, VLOK, I.H., MCDONALD, D.J., OLIVER, E.G.H.,

BOUCHER, C., VAN WYK, B.-E, & SCHUTTE, A. 1993. The

high altitude flora and vegetation of the Cape Floristic Region,

South Africa. Opera Botanica 121: 247-261.

MCDONALD, D.I.. BOUCHER, C., & OLIVER, E.G.H. 2000. Obi-

tuary: Hugh Colin Taylor (1925-1999). Bothalia 30: 1 15-119.

OLIVER, E.G.H. & OLIVER, I.M. 1994. Studies in the Ericoideae

(Ericaceae). XIV. Notes on the genus Erica. Bothalia 24:

25-30.

OLIVER, E.G.H. & OLIVER, I.M. 1998a. Erica schumannii, a new

mat-forming species from South Africa. Yearbook of the Heather

Society 1998: 32-38.

OLIVER, E.G.H. & OLIVER, I.M. 1998b. Three new species of Erica

(Ericaceae) from South Africa, Novon 8: 267-274.

OLIVER, E.G.H. & OLIVER, I.M. 2000. Erica kirstenii, a new rock-

loving species from South Africa. Yearbook of the Heather

Society 2000: 57-62.

REBELO, A.G., SIEGFRIED, W.R. & OLIVER, E.G.H. 1985.

Pollination syndromes of Erica species in the south-western

Cape. South African Journal of Botany 51: 270-280.

SCHUMANN. D, & KIRSTEN, G. 1992. Ericas of South Africa. Fern-

wood Press, Cape Town.

Bothalia 31,2: 167-173 (2001)

Studies in the liverwort family Aneuraceae (Metzgeriales) from

southern Africa. 1. The genus Aneura and its local representative

S.M. PEROLD*

Keywords: Aneuraceae, Aneura pingiiis (L.) Dumort., Aneuroideae, Riccardia Gray, southern Africa, taxonomic problems

ABSTRACT

This is the first of several projected articles dealing with the southern African Aneuraceae H.Klinggr. The family is

cosmopolitan and comprises two subfamilies, only one of which, the Aneuroideae, with two genera, Aneura Dumort.

and Riccardia Gray, occurs locally. Because of the taxonomic problems associated with this family and the scarcity of

fruiting material, only one taxon in the genus Aneura, A. pinguis (L.) Dumort.. has so far been accepted in this treat-

ment. This is not meant to imply, however, that A. pseiidopinguis Herzog does not occur in southern Africa; only, that

the specimens I have examined, all belong to A. pinguis. A description and illustrations of A. pinguis. together with a

distribution map are provided.

INTRODUCTION

One of the relatively smaller families within the order

Metzgeriales, is the cosmopolitan Aneuraceae H.Klinggr.

It is divided into two subfamilies, which together com-

prise only four genera, but more than 200 species world-

wide. Only one subfamily, the Aneuroideae (Schuster

1992). with two genera, occurs in southern Africa:

Aneura Dumort. with one locally, infrequently collected

species, and the more common Riccardia Gray, with sev-

eral species. These two hygrophilous genera are related.

Initially, and up to the middle of the 20th century, they

were treated as synonyms, ‘sometimes using the former

name and sometimes the latter, for the same taxon’

(Grolle 1993).

The genera Aneu/'fl and Riccardia may be separated as

follows:

Aneura

Thalli prostrate, closely adherent to substrate, fleshy,

bright green and with somewhat greasy lustre, robust,

2^(-7) mm wide, ligulate; apices rounded or scarcely

emarginate; branching subsimple, developing few lateral

pinnae; oil bodies in cells numerous, clear and small,

2. 5-7. 5 pm diam.; in cross section thallus (9)10-15 (or

more) cells thick medianly, thallus gradually becoming

thinner toward undulate or plicate margins. Rhizoids

restricted to median ventral surface. Gemmae absent in

local species.

Dioicous; male branches with 2-A- irregular rows of

antheridia; female branches rudimentary with inflores-

cences sessile and ventrally hidden in deep lateral notch,

archegonia in 2 or more rows; shoot-calyptra large,

clavate, rounded at summit, corona inconspicuous; seta

thick, 9-16 cells diam.; capsule valves with thickenings

along longitudinal walls of epidermal cells on both adax-

ial and abaxial surfaces; spores 18.0-27.5 pm diam.

* National Botanical Institute, Private Bag XlOl, 0001 Pretoria.

MS. received: 2001-05-23.

Riccardia

Thalli prostrate, rarely erect, pale to deep green, rela-

tively delicate and narrow, 0. 5-2.0 mm wide, linear to

Ungulate; apices emarginate, rounded or truncate;

branching freely, mostly uni- to rarely multipinnate; oil

bodies sometimes absent in epidermal cells, otherwise

1-3(4) per cell, brown or black, large, 7.5-20.0 x 5-9

pm, ovoid, spherical or ellipsoid; in cross section axis up

to 9 cells thick, margins plane. Rhizoids scattered on

ventral surface. Gemmae endogenously produced on

dorsal surface of thallus and 2-celled.

Dioicous, autoicous or heteroicous; male branches lat-

eral, linear, with 2 regular rows of antheridia in 2-8 pairs;

female branches lateral, well developed, with 2 rows of

archegonia; shoot-calyptra medium-sized, clavate or

cylindrical, corona conspicuous; seta slender, 4 cells

diam.; capsule valves with thickenings confined to adax-

ial faces of longitudinal radial walls of epidermal cells.

In both genera there is little anatomical differentia-

tion, usually without any distinction between the wings

and midrib or nerve (the epithet Aneura, signifies the

absence of a nerve). Because of their simplicity in organ-

ization, there are few distinctive characters. Moreover,

frequent variability, apparently environmentally induced,

in those few features that are available, make them diffi-

cult to study. Taxonomically it is widely accepted that the

Aneuraceae is one of the most troublesome families

among the Hepaticae.

The long, narrow thalli of Riccardia species in particular,

often grow in intricately intertwined, densely overlying

mats, with more than one species sometimes being pre-

sent. The branches are rather brittle, and unravelling and

cleaning them for study needs to be done with care and

is time-consuming. Sterile plants are of little value, as

gametangial and sporophytic characters are necessary for

the correct placing of specimens; unfortunately, they are

frequently without sporophytes, although they may pro-

duce gametangia. Furthermore, as they soon disappear,

the oil bodies in samples from fresh plants, that are

preferably kept in a refrigerator at 5°-6°C, need to be

168

Bothalia 31,2 (2001)

photographed and studied as soon as possible after col-

lection. Data from disintegrating oil bodies can be mis-

leading.

It is also recommended that plants should be studied

in cultivation.

Meenks ( 1987) is of the opinion that herbarium collec-

tions are often useless, particularly if they are more than

20 years old. Type specimens are, however, far older gen-

erally, yet one needs to study them to compare them with

fresh, living collections, however difficult that may be.

HISTORICAL NOTES

The Aneuraceae have been relatively poorly studied

in southern Africa. Lehmann (1829) described our first

new local taxon, Aneura (sub Jungermannia) fastigiata,

collected by Ecklon on Devil’s Peak and Table Mountain

and compared it to A. (sub J.) multifida, which he also

reported from Table Mountain, as well as two varieties of

it. Gottsche et al. (1844-1847) accepted the presence of

A. fastigiata, but compared it to A. palmata. They also

briefly mentioned Ecklon’s collections of A. pinnatifida

and A. multifida from Promontorium Bonae Spei

(Gottsche et al. 1844-1847: 788).

No further species were recorded from tropical or

southern Africa (Jones 1956), until Stephani (1886)

reported R. pinguis L. from S. Tome. During 1890-1893

and 1917 Stephani described more than 20 new species

from the African continent and the Mascareignes. Gola

(1914) and Pearson (1922) each described one new

species from Africa. Sim (1926) described no new

species in his treatment, and only briefly mentioned A.

fastigiata and A. compacta. Then, in 1952, S.W. Amell

described five new species from southern Africa. In

1963, he placed one of them, R. submarginata, in syn-

onymy under the Brazilian species R. pseudopinguis

Herzog (1942), although he had previously agreed with

Jones (1956) that it was identical with R. pinguis. Jones

(1956) was very conservative in his treatment of

Riccardia species in tropical Africa. He placed some of

Stephani’s species in synonymy and made several new

combinations, transferring four of Stephani’s Aneura

species to Riccardia, as the distinctions between the two

genera had become generally accepted by then. Vanden

Berghen (1972) reported the presence of A. pseudopin-

guis from Shaba, Kafubu and Lufira in Zaire (Demo-

cratic Republic of Congo).

In 1981, Jones drew attention to an inversion of spe-

cific epithets that had occurred on the herbarium labels

of the type specimens of A. congoana Steph. and A. con-

goensis Steph. Later, Jones & Harrington (1983) report-

ed R. angusticosta and R. limhata from Sierra Leone and

Ghana, as well as an unidentified Riccardia species from

Ghana. Gradstein et al. (1983) announced the presence

of A. pseudopinguis in West and East Africa, and in

the same article Poes transferred it from Riccardia to

Aneura.

The last treatment of East African Aneuraceae was by

Meenks & Poes ( 1 985), as a large number of unidentified

specimens and new information had accumulated in the

meantime. Although they present a key to the African

species of the family, and give extensive lists of speci-

mens examined, literature records and distribution maps,

only one species, R. compacta, is illustrated. Illustrations

by Amell (1952, 1963) are generally poor and Stephani’s

leones (1985) are not freely available. Schuster (1992)

remarked, with some justification, that African taxa of

the Aneuraceae remain poorly known, as Arnell’s treat-

ment is far from satisfactory.

It is accordingly considered necessary to revise at

least the southern African members of the Aneuraceae,

which often represent southern extensions of range, and

to give good illustrations wherever possible. This publi-

cation is the first in a series of projected papers on the

local Aneuraceae.

METHODS AND MATERIAL STUDIED

With some exceptions, the same methodology as was

employed in my treatment of southern African Fossom-

hronia species (Perold 1997), is followed here: 1, sam-

ples of field-collected specimens were fixed in EAA on

the day of collection; 2, more samples were selected for

oil body study and cultivation and initially kept at

5°-6°C in transparent screw boxes, lined with damp fil-

ter paper; and 3, the rest of the specimens, destined to be

held in the herbarium, were allowed to air-dry rapidly.

Only traditional morphological-anatomical and histolog-

ical methods were employed. Ear less extensive use of

the SEM was made in this study, however, as the spores

of our local Riccardia species do not display marked dif-

ferences in their ornamentation. The drawings were

based on photographs taken with a camera mounted on a

Nikon light microscope. The Degree Reference System

(Edwards & Leistner 1971) was again used for recording

distribution data.

Specimens e.xamined

(held at PRE, unless otherwise indicated)

S.VV. Amell 1552, 1766 (types of R. submarginata BOL); 1782

(BOL); 1803, 1844 (S).

Burgoyne 2462.

Cholnoky s.n. (S) KwaZulu-Natal; grid unknown and not indicated

on Figure 2.

Koekemoer 2063.

Perold 3774, 3823. Perold & Koekemoer 4304, 4483, 4504, 4514.

Aneuraceae H.Klinggr., Die hoheren Cryptogamen

Preussens: 11 (1858); Mull.Erib.: 492 (1951-1958);

S.W.Amell; 84 (1963); Euruki: 306 (1991); R.M.Schust.:

545 (1992); Paton: 537 (1999); Bednarek-Ochyra et al.;

178 (2000). Type; Aneura Dumort.

Plants thalloid, prostrate or sometimes ascending,

bright green with greasy lustre or light to dark green,

robust and fleshy. Ungulate to sublinear, 2^(-6) mm

wide or narrow and thin, relatively delicate, mostly lin-

ear, 0. 5-2.0 mm wide, apices rounded or emarginate,

margins undulate, sometimes flat or plane and entire.

Branching virtually simple to sparse, branches laterally

and irregularly lobed or freely pinnate to quadripinnate.

Bothalia 31,2 (2001)

169

occasionally crowded. Dorsal epidermal cells generally

somewhat smaller than internal cells, not otherwise dif-

ferentiated, usually thin-walled, occasionally with

minute trigones, 5-7-angled. Oil bodies in each cell,

either numerous and then small and hyaline, or only

1-3(4), and then large, finely granular and brown or

black, rarely absent. Cross section of main axis plano-

convex to concavo-convex with weakly recurved mar-

gins, or flattened to biconvex; pluristratose medianly,

usually 9-15 cells thick, gradually thinning out laterally

to margins, acute, winged, or obtuse, without discrete

midrib, internally homogeneous, lower cells sometimes

with basidiomycete fungal hyphae. Mucilage papillae

often in 2 alternate ventral rows, one on each side of mid-

line and/or clustered at apex of thallus. Rhizoids restrict-

ed to ventral median area or scattered over ventral sur-

face of thallus. Asexual reproduction absent or by 2-

celled, endogenously produced gemmae.

Dioicous, monoicous or synoicous. Sexual branches

lateral from main thallus, short and determinate in

length; male branches single or in sympodial pairs or in

threes, antheridia in up to 4 irregular rows or in 2 regular

rows of antheridial chambers, each containing a single

antheridium, dorsally exposed; female branches within

deep lateral notch of thallus or almost sessile laterally,

archegonia in 2 or more rows, protected by papilliform

or finger-like paraphyses. Shoot-calyptra large, clavate

or cylindrical, with wall several cell layers thick, surface

almost smooth or bearing papillose outgrowths. Seta

thick. 10-12 cells diam., or slender and only 4 cells

diam. Capsule ellipsoidal, 4-valved, wall bistratose,

outer cell layer with bands of thickening on adaxial and

abaxial radial walls; inner cell layer with bands of thick-

ening on adaxial and abaxial radial and inner tangential

walls, or else, outer cell layer with bands of thickening

on adaxial radial and inner tangential walls, without

bands of thickening on inner cell layer. Spores relatively

small, up to 27.5 pm diam., finely papillose, or only up

to ± 20 pm diam., minutely scabrate. Elaters up to ± 350

pm long, uni-spiral, band reddish brown, 7.5-10.0 pm

wide, tapering at both ends. Elaterophores in fascicles at

apices of valves.

Aneura Dumort., Commentationes Botanicae: 1 15

(1822); Hewson: 184 (1970); Furuki: 308 (1991);

R.M.Schust.: 551 (1992); Paton: 537 (1999). Type spe-

cies: Aneura pinguis (L.) Dumort.

Jungermannia pinguis L.: 1136 (1753).

Trichostyliiim Corda: 116 (1835); Nees: 475 (1838);

R.M.Schust.: 53 (1958).

Riccardia subgenus Trichostyliiim Mizut. & S.Hatt.:

35 (1957).

Thalli prostrate, clear green and greasy, robust, fleshy,

dorsally flat or slightly concave, gradually thinning out

from thick median region to margins, smooth, up to 35

mm long, 2^(-6) mm wide. Ungulate to sublinear,

apices rounded or slightly emarginate, margins undulate,

sometimes flat. Branching sparse and irregular. Oil bod-

ies (2-)5-24(-67) in all dorsal epidermal and inner cells.

subglobose or ovoid, 2. 5-5.0 pm diam., hyaline. Cross

section of thallus (9)10-15 cells thick medianly, acute at

margins. Mucilage papillae only on ventral surface of

thallus, in 2 rows, one on each side of midline and clus-

tered at branch apices. Rhizoids restricted to median area

of ventral surface of thallus. Asexual reproduction ab-

sent, gemmae unknown in local species.

Dioicous. Heterothallic. Male thalli somewhat small-

er, antheridial branches single or sympodially paired and

antheridia in up to 4 irregular rows, separating walls usu-

ally bistratose, dorso-lateral wing up to 6 cells wide.

Eemale branches shielded and hidden in deep, lateral

notches of thallus, paraphyses around and between

archegonia. Shoot-calyptrae large, ± 10 mm long, fleshy,

wall ± 500 pm thick, corona inconspicuous. Setae mas-

sive, 10-12 cells diam., spirally twisted. Capsules ellip-

soidal, 4-valved, bistratose, epidermal cells with nodular

thickenings of both adaxial and abaxial radial longitudi-

nal walls; inner cell walls with numerous annular or

semiannular (tangential) bands. Spores finely papillose,

20.0- 27.5 pm diam. Elaters up to 350 pm long, 10 pm

wide, unispiral.

Aneura pinguis (L.) Dumort., Commentationes

Botanicae; 115 (1822); Dumort.: 86 (1831); Gottsche et

al.: 493 (1844-1847); Steph.: 762/272 (1899); Mull.Frib.:

331 (1908); Brown & Braggins (with question mark): 117

(1989); Furuki: 311 (1991); R.M.Schust.: 555 (1992);

Paton: 539 (1999). Jungermannia pinguis L.: 1136

(1753). Riccardia pinguis (L.) Gray: 683 (1821);

Mull.Frib.: 494 (1951-1958); S.W.Amell: 30 (1956);

E.W.Jones: 84 ( 1956). Trichostyliiim pinguis R.M.Schust.:

53 (1958). Type: ‘Habitat in Europae paludibus’, sin.

coll.. Dill, in ‘Historia Muscorum: 509. Lichenastrum

no. 42. t. 74, fig. 42 (1741)’ fOXF, syn.; H-SOL,

isosyn.].

R. viridissima Schiffn.: 176 ( 1898). A. viridissima (Schiffn.) Steph.:

763/273 (1899). Type: Java. Prov. Batavia, in monte Megamendon,

Schiffner 235, syntype (JE, L. PR. PRC, W), fide Furuki 1991.

A. laurentiana Steph.: 32 (1917). R. laurentiana (Steph.) Demaret:

306 (1942). Type: Congo, env. d'Eala, Laurent 64 (G).

R. submarginata S.W.Amell: 139 (1952). Types: Deepwall Forest

Reserve. V2 mile south of Forest Station, S.W. Arnell 1552, 1766

(BOLI; S).

A complete list of synonyms is provided in R.M. Schuster (1992:

555, 556).

Thalli prostrate, in patches, bright green, with greasy

lustre, smooth, yellow when dried, opaque, becoming

translucent toward margins, otherwise fleshy, robust,

axis plane to slightly concave, subsimple, Ungulate to

sublinear, 15-35 mm long, 2-6(-7) mm wide, apices

rounded, margins entire, undulate to somewhat crisped

or lobed. Branching sparse, not pinnate, often short,

sometimes long, inegularly furcate. Dorsal epidermal

cells (Eigure IH) from above, 4—6(7)-sided, thin-walled,

50.0- 87.5 X 37.5-62.5 pm; subdorsal cells (Eigure II)

1 12.5-175.0 X 87.5-132.5 pm; subventral cells 100-160

X 50-75 pm; ventral epidermal cells 62.5-100.0 x 37.5-

50.0 pm. Oil bodies in all cells, several to numerous,

(2-)5-24(-67), small, 2. 5-5.0 pm diam., spherical to

subspherical or ovoid, faintly granular, inconspicuous.

170

Bothalia 31,2 (2001)

FIGURE 1. — Aneura pinguis. A, male thallus with antheridial branches; B, female thallus with calyptra, shortened seta and capsule, old, collapsed

calyptra on upper right, very young calyptra opposite it. C-I, thallus: C, c/s; D, E, c/s marginal part; F, c/s median part; G, margin from

above; H, dorsal epidermal cells with oil bodies; I, subdorsal cells with oil bodies. J, rhizoid with ramified tip and internal mycorrhizal

hypha. K-M, male branches: K, partly from side, only one wing shown; L, c/s; M, wing. N, O, young female branches: N, ventral view,

located at notch in thallus margin; O, lateral view. P, paraphysis. Q-T, capsule: Q, valve with elaterophore at tip. R, c/s part of bistratose

wall, with thickenings: abr, abaxial radial; adr, adaxial radial; mw, median; it, inner tangential; ot, outer tangential. S, thickenings of outer

cell layer; T, thickenings of inner cell layer. U, calyptra; V, c/s calyptra; W, c/s seta; X, spore; Y, elater. A, B, N, S, T, W-Y, Perold &

Koekemoer 4504\ C, D, F, H, I, K-M, 0-R, U, V, Perold <6 Koekernoer 45I4\ E, G, Perold & Koekemoer 4483; J, Perold 3823. Scale bars:

A, B, U, 2 mm; C, K, L, 500 pm; D, E, G, .1, S, T, 100 pm; H, 1, R, Y, 50 pm; Q, I mm; F, M-P, V, W, 250 pm; X, 25 pm.

Bothalia 31,2 (2001 )

171

colourless, scattered or occasionally aggregated in clus-

ters. Marginal cells (Figure IG): outermost row unis-

tratose, rectangular, 47.5-75.0 x 25^5 jam, inner cells

angular, bistratose, 42.5-55.0 x 37.5-62.5 |jm. Cross

section of thallus (Figure 1C) plano-convex to concavo-

convex, with wings weakly recurved, medianly pluris-

tratose (Figure IF), 9-15 cells (or 460-650 pm) thick,

gradually becoming thinner laterally, epidermal cells

chlorophyllose, 37.5-50.0 pm high, shallower than inter-

nal, hyaline cells, but not forming a distinctive layer;

margins acute, bordered by bistratose wing, 3 cells wide

plus a single conical cell (Figure ID, E), ± 50 pm high.

Mucilage papillae clustered at ventral apices of branch-

es and spaced along 2 rows, one on each side of midline,

clavate, 105-114 x 42.5-51.0 pm, not persistent.

Rhizoids restricted to ventral median area, 17.5-25.0 pm

wide, sometimes ramified at tips and occasionally with

mycorrhizal hyphae (Figure IJ). Asexual reproduction

by gemmae absent.

Dioicous. Male plants (Figure lA) somewhat smaller

than female plants, axis up to 17 x 3. 5^. 2 mm and medi-

anly 9-11 cell layers thick, in close proximity to female

plants, virtually touching; antheridial branches (Figure

IK) lateral, solitary, sublinear, 2.8 mm long or more, ±

1.15 mm wide and ± 850 pm thick (Figure IL), scal-

loped, bistratose wings (Figure IM), 300-500 pm or 6-8

cells wide, a continuation of thallus margin; otherwise

divided into 2 or 3 branches, from near base or further

along, up to 2.4 mm wide, antheridial chambers very

irregularly arranged, not in rows, 2-4 across width of

branch; antheridia subglobular, 200-250 pm wide, cavi-

ty walls between them bistratose; after antheridial pro-

duction, branch may continue vegetative growth. Female

plants (Figure IB) with archegonial branches very short,

ventral to lateral notch in thallus margin, at base of deep

sinus and obscured by reflexed thallus folds on each side

(Figure IN), archegonia in rows, hidden by densely

crowded paraphyses, up to ± 500 pm long, composed of

single strands of cells joined end to end (Figure lO) or

partly of 3 or more rows of laterally joined cells (Figure

IP). Calyptra clavate, large (Figure lU), 10-12 mm

long, ± 2 mm diam., wall 8 or 9 cell layers (+ 500 pm)

thick, cells in cross section (Figure IV) 5-7-sided,

65-100 X 62.5-75.0 pm. outermost cells of wall in sur-

face view long-rectangular, 150-170 x 60-75 pm, in

lower part of calyptra often developing rhizoids, but

becoming nearly smooth with age, corona inconspi-

cuous, base partly sheathed by adjacent, suberect thallus

margins. Seta up to 21 mm long, somewhat spirally

twisted, 750 pm or 15 cells diam. in cross section (Figure

IW), marginal cells small, 32.5-47.5 x 25-40 pm, inner

cells round, 55-75 pm diam. Capsules oblong-ovoid,

reddish brown, 2500-3250 pm long, with 4 valves, each

one ± 1050 pm (or ± 42 cells) wide, bistratose, to either

side of midline of every valve, thickenings laid down as

a mirror image; epidermal (outer) cells in external longi-

tudinal view (Figure IS) with nodular thickenings, cells

usually narrowly rectangular, 162.5-232.5 x 15.0-37.5

pm, in cross section (Figure IR), thickenings only rarely

extending slightly across outer tangential walls, mostly

on adaxial radial and abaxial radial walls; inner cells in

internal longitudinal view (Figure IT) less regular, taper-

ing or with oblique end walls, 142.5-220.0 x 32.5-67.5

pm, in cross section (Figure IR) adaxial radial longitudi-

nal walls all develop bands which extend across inner

tangential walls and connect with abaxial radial bands on

opposite side of cell, thickenings therefore ± U-shaped

(Figure IR). Spores 20.0-27.5 pm diam., dark brown,

papillose (Figure IX). Elaters (Figure lY) 212.5-350.0

X 10 pm, with single spiral band, ± 10 pm broad, pink-

ish red, tapering to ends, these without spiral.

Elateropliores 1000-1445 x 220 pm, one at tip of each

valve (Figure IQ). Chromosome no.: n = 10 (Fritsch

1982).

Aneura pinguis is subcosmopolitan in its distribution

and has been reported from many localities worldwide;

South America and Australia are, however, excluded. It

is also present in southern Africa, but is infrequently col-

lected (Figure 2). It grows on soil or rotting wood, in per-

manently damp areas, often fed by seepages.

DISCUSSION

Aneura specimens have recently been collected at a

few localities in Mpumalanga, Eastern Free State,

KwaZulu-Natal, Western and Eastern Cape, but Perold

& Koekemoer 4405 & 4415 from Garcia’s Pass, near

Ladismith in Western Cape, are the only specimens that

are fertile, with both mature male plants bearing ripe

antheridia on lateral branches, as well as female plants

with calyptrae and ripe capsules. Samples of these have

been preserved in EAA for future reference. These spec-

imens have been referred to Aneura pinguis because of

the following characters: 1, the large, bright green, rather

greasy-looking, fleshy and opaque thalli; 2, thalli in cross

section medianly up to 15 cell rows thick; 3, each cell

with several to many minute, colourless oil bodies; 4,

2— f rows of antheridial chambers; 5, spores that are

20.0-27.5 pm diam. On one side of the same male thal-

lus of Perold & Koekemoer 4514, there are a few short,

ligulate male branches, bearing the antheridia in 2 or 3

rows, whereas on its opposite side, there are several fur-

cate male branches, some of which are wider and

rounder at the apices, with up to 4 irregular rows of

numerous antheridia.

Meenks & Poes (1985) have drawn up a short table

differentiating between A. pinguis and A. pseudopinguis.

FIGURE 2. — Distribution of Aneura pinguis in southern Africa.

172

Bothalia 31,2 (2001 )

but there appears to be some overlap in the characters

assigned to these two species in our local specimens.

Meenks & Poes regard A. pinguis as having opaque thal-

li, medianly (9)10-20 cells thick, with relatively short

male lobes, which seldom branch and contain 2-5(6)

pairs of antheridia; the spores are 20-25 pm diam. On

the other hand, A. pseudopinguis thalli are, according to

them, translucent, medianly only 5-8(9) cells thick, the

male branches are longer, often with 3 (seldom 4) rows

or 7-10(-20) pairs of antheridia, and the spores are

13-16 pm diam.

In the specimens they studied, Gradstein et al. (1983)

regarded the identity as confirmed in ‘only the records of

plants possessing either male branches or spores’. They

also report that in African samples of A. pseudopinguis

‘usually only one sex is found in a particular locality’.

This is contrary to the findings in our two fertile collec-

tions refeiTed to above, and may provide further evi-

dence that we are dealing with A. pinguis.

In 1952 Arnell reported his new Riccardia siibmar-

ginata from Deepwall Forest Reserve. There were no

mature capsules with spores, but the median cross sec-

tions of the thalli were given as 15 cell rows thick. In my

examination of Arnell’s specimens the median cross sec-

tions of the thalli ranged from 8-13 cell rows thick.

As already remarked in the paragraphs on ‘Historical

notes’, Arnell had admitted to Jones (1956) that, the

Deepwall specimens were identical to R. pinguis,

although later, Arnell (1963) placed R. submarginata in

synonymy under R. pseudopinguis. Arnell was familiar

with European representatives of R. pinguis, as he had

described it in 1956 in his Flora Fennoscandia. Arnell

(1963) also gave vegetative characters, based on the size

of the thallus cells in R. pinguis, but these characters

appear to be in contradiction with his own account of

Scandinavian A. pinguis, as noted by Gradstein et al.

(1983).

Vanden Berghen (1972) reported the Congoan pres-

ence of A. pseudopinguis from Shaba, Kafubu in Lufira.

Hodgetts et al. (1999) also reported A. pseudopinguis

from Lesotho, northeast and west of New Oxbow Lodge,

but gave no details of their plants. Herzog (1942) de-

scribed his new species, R. pseudopinguis, from Brazil as

follows: thallus is medianly 6-8 cells thick in cross sec-

tion, male branches are elongated, ligulate, sometimes

furcate at the base, and the antheridia are in 8-10 serial

pairs, whereas the spores are 13-16 pm in diameter. He

suggested that previous collections from neighbouring

areas, which had been assigned to R. pinguis, should be

checked against his new species. In 1956, Jones placed

R. submarginata in synonymy under R. pinguis. In his

plants from West Africa, however, the central portion of

the thallus is 5-7 or occasionally 9 cells thick, which

would place them nearer to A. pseudopinguis. Wigginton

& Grolle (1996) observed that, ‘A. pseudopinguis is

sometimes considered to be conspecific with A. pin-

guis'. Schuster (1989) noted that, ‘The status of A. pin-

guis s. lat. badly needs world-wide study. Ever since

Showalter (1926, 1928) showed that, so-called ‘races’

of the holarctic populations were, in some cases inter-

sterile, the matter of how to define this species has been

highly uncertain’. Lurthermore, it seemed most unlike-

ly to Schuster that a species could occur in the Arctic

polar desert of Ellesmere Island and also in tropical and

subtropical areas. He concluded this paragraph with the

following observation, ‘Recent study has also shown

that the tropical American-African plants formerly

referred to A. pinguis represent A. pseudopinguis Herz’.

According to Meenks & Poes (1985), however, both

taxa occur in East Africa, as well as in West Africa

(Gradstein et al. 1983), but A. pinguis appears to be rarer

in tropical Africa than A. pseudopinguis. In southern

Africa, A. pinguis is very infrequently collected and fer-

tile plants are the exception by far, rather than the rule;

this scarcity of fruiting material further complicates the

study of these plants. To quote Proskauer (1971),

"Riccardia pinguis, a species which we have long known

to present a nightmarish problem ..., probably requiring

more than a single lifetime of research for a minimum

understanding’.

Stephani (1899) placed R. latissima (Spruce) Schiffn.

in synonymy under R. pinguis. However, Meenks & De

Jong (1985) note that, the oil bodies in A. latissima are

bright brown, not clear and colourless, as they are in A.

pinguis. Eurthermore, a chromosome no. n = 8, has been

reported by Nakata & Inoue (1989) for A. latissima,

whereas the chromosome no. n = 10 is given for A. pin-

guis. Stephani’s synonymy of A. latissima under A. pin-

guis, is, accordingly not accepted here.

ACKNOWLEDGEMENTS

I wish to thank the curators of BOL and S for the loan

of specimens; the curator of PRE, Ms M. Koekemoer, for

all her help with fieldwork; Dr T. Furuki and Dr O.A.

Leistner for their helpful comments; the artist, Ms G.

Condy; the photographer, Mrs A. Romanowski and the

typist, Ms D. Maree.

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Bothalia 31,2: 175-181 (2001)

Three new species of Tritoniopsis (Iridaceae: Crocoideae) from the

Cape Region of South Africa

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

Keywords: Iridaceae, new species, pollination, South Africa, taxonomy, Tritoniopsis L. Bolus, T. bicolor J.C. Manning & Goldblatt, T. flava

J.C. Manning & Goldblatt, T. toximontana J.C. Manning & Goldblatt, Western Cape

ABSTRACT

Three new species of the largely Western Cape genus Tritoniopsis L, Bolus are described, bringing the number of species

in the genus to 24. Tritoniopsis bicolor and T. flava are newly discovered, narrow endemics of the Bredasdorp Mountains

and the Kogelberg Biosphere Reserve, respectively, in the southwestern Cape. Both of these are areas of high local

endemism. T. toximontana. known since at least 1965 but misunderstood, is restricted to the Gifberg-Matsikamma

Mountain complex of northern Western Cape. Notes on the pollination biology of the species are provided.

INTRODUCTION

The genus Tritoniopsis L. Bolus (Iridaceae: Croco-

ideae) is a near-endemic of the Cape Floristic Region

(Goldblatt & Manning 2000a), with a single species, T

cajfra (Ker Gawl. ex Baker) Goldblatt, extending east-

wards to East London. The genus is a particularly char-

acteristic member of the Cape Flora as almost all of the

species are restricted to acidic, nutrient-poor soils

derived from sandstones of the Cape system. Originally

established to accommodate the single species Tritoniop-

sis lesliei L. Bolus (Bolus 1929), the genus was subse-

quently combined with Exohehea Foster (Phillips 1951;

Lewis 1959), and finally expanded to include the species

previously placed in Anapalina N.E.Br. (Goldblatt

1990). The 21 species currently recognised exhibit a

wide diversity in floral morphology but are united by

several highly unusual characters. The leaves of Trito-

niopsis, which are hysteranthus in all species, are unlike

most other genera of Iridaceae subfamily Crocoideae

(Goldblatt et al. 2000) in lacking a well-developed

pseudomidrib or central vein, having instead one to

seven equally prominent main veins. Flowering in most

species is delayed until summer or autumn, and although

green leaves are often evident at flowering time, these

sometimes represent vegetative growth derived from the

daughter corm. The contemporary appearance of leaves

and flowering shoots from successive generations is

unknown among hysteranthus Iridaceae and is correlated

with the persistence of corms from previous years’

growth and the late flowering habit. In other hysteran-

thus taxa of Iridaceae, leafing of the daughter corm typi-

cally occurs only after the flowering shoot of the parent

corm has died down. Alternatively, as in some species of

Gladiolus L., leaves are produced on a separate sterile

shoot later in the season. Tritoniopsis is also unusual in

having the inner floral bracts longer than the outer and

not evidently notched at the apices. The flowers,

although highly diverse in form, are always strongly

* Compton Herbarium, National Botanical Institute, Private Bag X7,

Claremont 7735, Cape Town.

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

P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2001-03-07.

herkogamous. This spatial separation of pollen and style

is achieved through the recurving of the filaments a few

days after anthesis, whereby they carry the anthers well

away from the style, the branches of which only become

receptive after the filaments have diverged. This charac-

teristic, noted by Lewis (1960) in her revision of the

species placed in Anapalina, was apparently overlooked

in her earlier account of Tritoniopsis s.s. (Lewis 1959)

but is characteristic of all species of the genus. Another

distinctive characteristic of most of the species is the way

in which the tepals gape at the tips in bud, several days

before anthesis actually occurs. This feature has not been

commented on before and is otherwise unknown in the

family. Finally, the seeds of almost all species of

Tritoniopsis are tetrahedral or pyramidal in shape and

typically more or less winged on the angles. Species in

which the seeds are larger and more prominently winged

invariably produce much-inflated capsules. This range of

autapomorphies accord with indications that the genus is

taxonomically isolated within subfamily Crocoideae.

Several of the species have restricted ranges and the

three described here are no exception. One species, T

toximontana, appears to be restricted to the Gifberg and

Matsikamma massif, while T. bicolor and T flava are

each known from single populations in the Bredasdorp

Mountains and Kogelberg Biosphere Reserve in the

southwestern Cape respectively. Both of these are areas

of high local endemism (Goldblatt & Manning 2000a).

Tritoniopsis bicolor J.C. Manning & Goldblatt, sp.

nov., plantae 200-400(-600) mm altae usitate eramosae,

cormo globoso 20-30 mm diam. tunicis fibris brunneis

dense implexis conferto, foliis 5-9 inferioribus (4)5-7

basalibus viridis vel extemis siccis ad tempore floredi in

pseudopetiolum subteretem 30-35 mm longum infime

abrupte contractis, lamina anguste lanceolata ( 85-) 1 00- 1 50

X (5-)6-10 mm prominente 1-2(3 )-nervata, foliis caulinis 1

vel 2 subsquamosis, spica 20-35-flora, bracteis infime

viridibus supeme siccis brunneisque, externa 7-10 mm

longa acuta, interna 8-1 1 mm longa obtusa, floribus zygo-

morphis mo.schatis vel pungentibis odoris flavis tepalibus

tribus infemis marroninus suffusis in medio atrostriatis ore

tubi et basibus tepalarum minute papillosis, perianthii tubo

2-5 mm longo, tepalis inaequalibus marginibus undulato-

176

Bothalia 31,2 (2001 )

crispis dorsale 14—16 x 3^ nim inferioribus tribus 9-15

mm infemis connatis laminis 9-10 mm longis, filamentis

6-7 mm longis, antheris ± 3 mm longis apiculis ± 0.5 mm

longis ferentibus, styli ramis ± 1 mm longis.

TYPE. — Western Cape, 3419 (Caledon): western end

of Bredasdorp Mtn, plateau west of Grootkop and

Normanskop, (-BD), 6 Dec. 2000, Manning 2286 (NBG,

holo.; K, MO, NBG, PRE, iso.).

FIGURE 1 . — Tritouiopsis hicolor.

A, corm; B. flowering spike

and leaves. C-E, flower: C,

front view; D, side view; E,

half flower. F, capsules; G,

seed. Scale burs: A-F, 10

inm; G, 10 mm. Artist: John

Manning.

Bothalia 31.2 (2001)

177

Plants 200^00(-600) mm high. Conns globose,

20-30 mm diam., tunics of densely matted fine brown

fibres with well-developed neck, corms from previous

seasons persistent. Leaves 5-9, lower (4)5-7 basal, green

at flowering or the outermost drying, contracted abrupt-

ly into semiterete pseudopetiole 30-35 x 1.0-2. 5 mm,

blade narrowly lanceolate, acuminate, with 1 or 2(3)

main veins, (85-)100-150 x (5)6-10 mm, cauline leaves

evidently 1 or 2, represented by minute scales, lower

cauline leaf concealed by fibrous corm neck. Stem mod-

erately slender to robust, decurved just above ground

level neck and then suberect or weakly inclined,

unbranched, 2. 5^.0 mm diam. near base. Spike densely

20-35-flowered, 80-100(-150) mm long; bracts green at

base but dry and brown above, leathery, oblong, outer

bracts 7-10 mm long, acute, inner bracts 8-1 1 mm long,

obtuse. Flowers zygomorphic, yellow, three lower tepals

each flushed maroon in lower half and with darker medi-

an streak, producing strong, acrid or musty scent; peri-

anth tube 2-5 mm long; tepals unequal, dorsal slightly

larger, margins crisped or undulate, base of lower tepals

and mouth of tube minutely papillate, dorsal tepal

ascending and erect to slightly recurved above, narrowly

oblanceolate, obtuse, 14-16 x 3-4 mm, upper laterals

spreading horizontally, strongly clawed and recurved

above, claw involute, 2-4 mm long, blade oblanceolate

to obovate, obtuse, 7-9 x 2-3 mm, lower 3 tepals fused

to one another for 1.0- 1.5 mm, with or without short,

chanelled claw to 1.5 mm long, blade oblong, truncate,

9-10 X 2.0—4. 5 mm. Filaments arcuate but later erect,

6-7 mm long, base of median filament minutely papil-

late; anthers yellow, 3 mm long with short apiculus 0.5

mm long. Ovary ellipsoid, ± 4 mm long; style ± 9 mm

long, dividing opposite anther bases, branches ultimately

1 mm long. Capsule ovoid to broadly ellipsoid, inflated,

23-28 mm long. Seeds tetrahedral, 10-12 x 10 mm, red-

dish brown, winged on angles and conspicuously crested

on faces. Flowering time: December. Figure 1.

Distribution and biology. Tritoniopsis bicolor has

been collected only once from the Bredasdorp Moun-

tains on the summit plateau west of Normanskop (Figure

2), flowering in the late spring after a fire the previous

summer. The plants are restricted to seasonally water-

logged flats where they are fairly common growing in

deep, humus-enriched sands. The smaller, cream-flow-

ered species, T. unguicularis (Lam.) G.J. Lewis, occupies

the adjacent drier, rocky slopes and the occasional hybrid

plant was recorded at the interface between the popula-

tions. A few plants of T. parviflora (Jacq.) G.J. Lewis

were found in fruit within a few hundred metres of the

locality, also on drier slopes. Several individuals of the

solitary bee Amegilla spilostoma (Apidae: Anthophori-

nae) were seen and captured while they visited flowers of

T. bicolor.

History, this species was brought to our attention by a

local botanist. Nick Helme, who collected it during a

survey of portion of the Farm Pofadderskloof.

Diagnosis and relationships: T. bicolor is most likely

to be confused with T. parviflora and the two are clearly

closely related. The flowers in both species are moderate-

ly-sized with a musky to acrid scent and are coloured

bright yellow with the lower tepals prominently marked

FIGURE 2. — Distribution of Tritoniopsis bicolor. O; T. flava. •; and

T. toximontana. ▲. in Western Cape.

and flushed with maroon; the floral tube in both species is

very short; the base of the tepals and mouth of the tube are

minutely papillate; and the capsules of both species are

conspicuously inflated. Tritoniopsis pan’iflora differs in

its more prominently bilabiate flowers in which the lower

three tepals are joined together in a distinct palate, 3—4

mm long, and in its much longer filaments, 12-14 mm

long, that arch well over the lower lip. In T. bicolor the

lower tepals are joined together for l.(3-L5 mm without

forming a well-defined lower lip and the anthers are held

much closer to the mouth of the tube on short filaments

6-7 mm long. Tritoniopsis bicolor is distinctive in its

leaves, which have a conspicuous semiterete pseudopeti-

ole. 30-35 mm long, expanding abruptly into a narrowly

lanceolate blade, 6-10 mm wide. The leaves of T. parx’i-

flora are linear or linear-lanceolate, 2-5 mm wide and

although narrowed below are not contracted into a dis-

tinct pseudopetiole. The seeds of T. bicolor, measuring

10-12 mm long, also seem to be slightly larger than those

of T. par\'iflora, which never exceed 9 mm in length.

Both species are restricted to acidic soils derived from

sandstone rocks of the Table Mountain Series and

although they both appear to flower only after a burn,

they differ markedly in other aspects of their ecology and

pollination biology. Tritoniopsis pan’iflora is widespread

on dry, stony slopes in shallow sandy soils, from

Citrusdal in the north to Bredasdorp in the south. It is

unique among African Iridaceae in secreting floral oils

from epithelial elaiophores on the perianth. Oil secretion

is an adaptation to pollination by the large oil-collecting

bee Rediviva gigas (Mellittidae) (Manning & Goldblatt

in press) which has been captured while visiting and pol-

linating flowers of T. parviflora. Tritoniopsis bicolor,

however, is restricted to a seasonally waterlogged site on

the plateau of the Bredasdorp Mountains where it grows

in deep, humus-enriched sands (pH 4.5). It is pollinated

by the nectar-collecting bee Amegilla spilostoma. T.

bicolor is evidently a local segregate of T. pan’iflora that

has speciated on the specialised, seepage habitat on the

Bredasdorp Mts at the extreme southern limits of the

range of T. pan’iflora. The papillae on the palate and

mouth of the tube that characterise both T. bicolor and T.

parviflora are unique in the genus and are a significant

synapomorphy for them. While the papillae are function-

178

Bothalia 31,2 (2001 )

al in oil secretion in T. paniflora, tests for floral oils in

T. bicolor proved negative or at most equivocal. This

suggests that the secretory function of the papillae has

been suppressed in T. bicolor and that they remain as

vestiges of an evolutionary origin from 71 parx’iflora. It is

possibly significant in this context that Rediviva gigas is

not known from south of Kleinmond (Whitehead &

Steiner 1993) and is therefore apparently absent from the

Bredasdorp Mts. A few plants of 71 parviflora with well-

developed fruits were, however, collected within a few

hundred metres of the population of 71 bicolor indicating

that a pollinating agent was active at the locality. Given

the similarity in floral form between the two species it is

likely that Amegilla spilostoma visits both species, but

the difference in flowering time between the two at this

locality prevents any possible hybridization.

Tritoniopsis flava J.C. Manning & Goldblatt, sp.

nov., plantae 500-600 mm altae eramosae, cormo glo-

boso 20-35 mm diam. tunicis fibris rubro-brunneis dense

implexis, foliis 10, inferioribus 7 basalibus viridis ad

tempore floredi in pseudopetiolum complanatum infime

contractis, lamina anguste lanceolata 130-190 x (3-)5-8

mm prominente 2-3-nervata, foliis caulinis 3 supernis

duobus squamosis inferno ad 60 mm longo vaginante,

spica ± 25-flora, bracteis ovatis obtusis viridibus infime

siccis brunneisque superne, externa 6-9 mm longa, inter-

na 7-10 mm longa, floribus zygomorphis probabiliter

odoris flavis tepalibus tribus infemis pallide flavis in

medio marroninus striatis, perianthii tubo 4-5 mm longo,

tepalis inaequalibus anguste unguiculatis dorsale 15-16

X 6 mm oblanceolato infime perangusto inferioribus

tribus ± 2.5 mm connatis infime laminis ±10x6 mm, fil-

amentis ± 10 mm longis, antheris ± 5 mm longis apiculis

± 0.5 mm longis ferentibus, styli ramis ± 1 mm longis ad

apicem dilatatis.

TYPE. — Western Cape, 3418 (Simonstown): Kogel-

berg State Forest, along footpath and next to stream,

(-BD), 10 Dec. 1991, Kruger 147 (NBG, holo.).

Plants 500-600 mm high. Conns globose, 20-35 mm

diam., tunics of densely matted reddish brown fibres

with short neck. Leaves 10, lower 7 basal, green at flow-

ering, narrowed into flattened pseudopetiole below,

blade narrowly lanceolate, 130-190 x (3-)5-8 mm,

acuminate, with 2 or 3 main veins, cauline leaves 3 but

upper 2 scarcely evident and represented by minute

scales, lowermost reddish brown, lanceolate, up to 60

mm long, clasping throughout. Stem robust, decurved

slightly just above ground level, unbranched, 4-5 mm

diam. near base. Spike densely ± 25-flowered, 150-180

mm long; bracts green at base but dry and brown above,

rigid and leathery but papery along margins, ovate, outer

bracts 6-9 mm long, obtuse, inner bracts 7-10 mm long,

obtuse. Flowers zygomorphic, yellow, three lower tepals

each paler towards base with small maroon median

streak in lower third; perianth tube slightly curved and

widening to throat, 4-5 mm long; tepals unequal, dorsal

slightly larger, oblanceolate and strongly narrowed in

lower third, obtuse, 15-16 x 6 mm, upper laterals

clawed, claw channelled, 5 mm long, blade obovate,

obtuse, 8-9 x 5 mm, lower 3 tepals fused to one another

for ± 2.5 mm, strongly clawed, claws channelled, 2 mm

long, blades obovate-pandurate, truncate, ± 10x6 mm.

Filaments arcuate but later recurving, ± 10 mm long;

anthers yellow, ± 5 mm long with short apiculus 0.5 mm

long. Ovary ellipsoid, ± 4 mm long; style ± 9 mm long,

dividing opposite anther bases, branches 1 mm long,

expanded apically. Capsule and seeds unknown.

Flowering time: December.

Distribution and biology: Tritoniopsis flava is known

from a single population in the Palmiet River Valley

(Figure 2). The plants grow in seasonally damp soils

along a stream and flower only in the season following a

burn. The vegetation at the site regenerates rapidly and

within a few seasons the area is covered with a dense

growth of woody vegetation.

History: this enigmatic species is known from a single

collection made in 1999 during a survey of the flora of

the Kogelberg Biosphere Reserve. The plants appeared

along a small tributary of the Palmiet River following a

fire and have not been seen since. The vegetation across

the Palmiet River from this site burned in the summer of

2000 but several visits to apparently similar streams

opposite the original site failed to rediscover the species.

Diagnosis and relationships: T flava is one of a small

group of closely allied species that includes 71 caledo-

nensis (R.C. Foster) G.J.Fewis, 71 unguicularis, 71 pand-

flora and 71 bicolor. The alliance is characterised by rel-

atively small, fragrant, cream-coloured to yellow flowers

with abruptly clawed tepals in which the claws are chan-

nelled or involute, and by the short style branches. The

species are distinguished on details of flower colour, size

and proportions, and by leaf shape and venation. The

alliance is restricted to the mountains of the southwest-

ern Cape between the Cederberg and Bredasdorp. 71

flava is most likely to be confused with 71 parviflora or

71 bicolor on account of the size of its flowers but is dis-

tinguished from these species by its robust habit, nar-

rowly lanceolate leaves mostly 5-8 mm wide with 2 or 3

main veins, and by the lack of extensive maroon col-

oration on the lower tepals. It also lacks the papillate

mouth of the tube and palate that characterises the flow-

ers of these two species. All three of these species appear

to flower only in the season following a summer burn.

Tritoniopsis toximontana J.C. Manning & Gold-

blatt, sp. nov., plantae 300-650 mm altae usitate

eramosae vel 1-2-ramosae, cormo globoso 30-50 mm

diam. tunicis fibris dense implexis rubrobrunneis confer-

to, foliis 4 vel 5 inferioribus 1-2 basalibus siccis ad tem-

pore florendi in pseudopetiolum teretem 100-150 mm

longum abrupte infime contractis, lamina linearo-lanceo-

lata 200-300 x 5-10 mm prominente trinervata, foliis

caulinis 3 vaginantibus omnino, spica 5-9-flora, bracteis

viridibus infime siccis roseis superne, externa 10-12 mm

longa acuta, interna 14-16 mm longa acuminata, floribus

zygomorphis inodoris pallide roseis tepalibus inferne

albis atrorubrisque notatis basibus filamentarum et ore

tubi atrorubris coloratis, perianthii tubo 20-21 mm

longo, tepalis inaequalibus dorsale 23-25 x 7-9 mm

inferioribus tribus 22-25 mm longis infime 5-6 mm con-

natis, filamentis 13-15 mm longis, antheris 7-8 mm

longis apiculis ± 2 mm longis usitate bifurcatis feren-

tibus, styli ramis 3-4 mm longis ad apicem parum

dilatatis.

Bothalia 31.2 (2001)

179

TYPE. — Western Cape, 3118 (Vanrhynsdorp):

Gifberg, near top of pass, sheltered sandstone slope in

arid fynbos, (-DD), 22 Mar. 2000, Manning 2234 (NBG,

holo.; K, PRE, iso.).

Plants 300-650 mm high. Corm globose, 30-50 mm

diam., tunics of densely matted reddish brown fibres

with thick neck to 150 mm long. Leaves 4 or 5, the

lower 1 or 2 basal, dry at flowering, contracted abruptly

into slender, terete pseudopetiole 100-150 mm long,

blade linear-lanceolate, 200-300 x 5-10 mm, attenuate,

oblique at base, with 3 main veins, abaxial vein diverg-

ing first, cauline leaves 3, reddish brown, lanceolate,

clasping throughout, lowermost to 60 mm long, upper

reduced and bract-like. Stem slender, straight or slightly

flexuose, usually unbranched, rarely with one or two

single-flowered branches; branches filiform and

suberect. Spike laxly 5-9{-12)-flowered, 300-600 mm

long, lateral spikes 1 -flowered; bracts green at base but

dry and flushed pink in upper half at flowering, soft and

more or less membranous, oblong, outer bracts 10-12

mm long, acute, inner bracts 14-16 mm long, acumi-

nate. Flowers zygomorphic, unscented, pale pink, three

lower tepals each with dark red median streak outlined

in white and flanked at base by dark pink streak, upper

laterals usually with darker median streak near apices,

mouth of tube and bases of filaments flushed dark red;

perianth tube straight or slightly curved, very slightly

widened to throat. 20-21 mm long; tepals unequal, dor-

sal slightly ascending and recurved above, others direct-

ed forward below and recurved above, dorsal largest,

oblanceolate, 23-25 x 7-9 mm, narrowed below, upper

laterals 22-25 x 3. 5-5.0 mm. lower tepals as long as

upper laterals but usually slightly narrower, fused to one

another for 5-6 mm. Filaments arcuate but later recurv-

ing, 13-15 mm long; anthers purple. 7-8 mm long

including recurved, usually bifid apiculus ± 2 mm long.

Ovary ellipsoid, ± 4 mm long; style 35-37 mm long,

dividing slightly beyond anther apices, branches ulti-

mately 3-4 mm long, slightly expanded apically.

Capsule ovoid to ellipsoid, somewhat inflated, 15-23

mm long. Seeds tetrahedral, ±6x4 mm, reddish brown,

winged on angles and rugose on faces. Flowering time:

March to early May. Figure 3.

Distribution and biology: Tritoniopsis toximontana is

known only from the Gifberg and Matsikamma

Mountains (Figure 2), where it is quite common in arid

fynbos on the summit plateau, although apparently

absent from the highest reaches of the Matsikamma. It is

possible that the species extends northwards along the

Bokkeveld Mountains but a search there in March 2000,

failed to discover the species. The plants grow tightly

wedged among sandstone boulders in sheltered situa-

tions, often on cooler slopes. The leaves are dry at flow-

ering time or sometimes those of the next season’s

growth are emergent.

The pink, long-tubed perianth with dark red throat and

tepal markings of T. toximontana are typical of plants

adapted to pollination by long-proboscid nemestrinid

and tabanid flies in South Africa (Goldblatt & Manning

2000b). The floral tube is shorter than is usual among

such plants, however, and T. toximontana is visited and

pollinated by bees in the genus Amegilla (Apidae:

Anthophorinae) as well as a moderate-sized nemestrinid

fly in the genus Prosoeca (unpublished observations).

The nemestrinid was also observed visiting the pink

flowers of Brunsvigia striata in the same area. Both of

these plant species appear to belong to an autumn-flow-

ering guild that is pollinated by this as yet unnamed

species of long-proboscid fly.

Historyr the species was first collected by Nan

Horrocks, an assistant in the Compton Herbarium, in

1965, several years after Lewis’ revision of the genus

(Lewis 1959). Although Lewis subsequently saw this

collection, she identified it as T. ramosa var. unguicula-

ta (Baker) G.J. Lewis, apparently on the basis of the long

perianth tube. Tritoniopsis toximontana was collected

only twice in the thirty years following Lewis’ death in

1967. The most recent collection, by Cape Town envi-

ronmentalist and botanist Nick Helme, alerted us to the

fact that it represented an undescribed species. Trito-

niopsis toximontana is quite common on the Gifberg

plateau and its late flowering habit must account for the

fact that it has not been collected more frequently from a

locality that is relatively well-visited by botanists.

Diagnosis and relationships: the long-tubed pink

flowers of T toximontana are superficially similar to

those of T revoluta (Burm.f.) Goldblatt and T flexuosa

(L.f.) G.J. Lewis but these species are both characterised

by large, usually firm floral bracts, 15-35 mm long. The

bracts of T toximontana are softer and membranous and

only 12 — 15 mm long. Although the leaves in most

species of Tritoniopsis are narrowed towards the base of

the blade into a flattened pseudopetiole, there are only

four in which the pseudopetiole is terete and completely

lacking in marginal flanges of blade tissue. This shared

character seems to indicate that T burchellii (N.E.Br.)

Goldblatt, T. triticea (Burm.f.) Goldblatt, T flexuosa and

T toximontana comprise a monophyletic group. How-

ever. the differences between the species in bract shape

and texture, and particularly in the form of the cauline

leaves suggests that this relationship, if correct is not a

close one. The closely related red-flowered T. burchellii

and T triticea are unusual in the genus in their aristate

cauline leaves and small, woody capsules and cannot be

confused with any other species. Among the pink-flow-

ered species with terete leaf bases, T toximontana differs

from T flexuosa not only in the bracts but also in the leaf

The leaf blade in T flexuosa is unique in the genus in

being oblong and at most 60 mm in length, whereas the

blade in T toximontana is narrowly lanceolate-attentuate

and 200-300 mm long.

Additional material examined

WESTERN CAPE. — 3118 (Vanrhynsdorp): Matsikamma

Mountains. Op de Berg 314, just west of Kdelkop Trig Beacon, (-DB),

1 Apr. 1997, Helme 1254 (NBG); Gifberg Mtn top. (-DD), 7 May

1965. Horrocks 205 (NBG); 12.7 km from Zandkraal turnoff at top of

Gifberg Pass, (-DD), 30 Mar. 1982, Snijman 595 (NBG).

ACKNOWLEDGEMENTS

We are grateful to Nick Helme for bringing two of the

new species to our attention, Mark and Ameida Johns for

180

Bothalia 31,2 (2001 )

FIGURE 3. — Tritoniopsis toximon-

tana. A, conn; B, flowering

spikes and leaves. C. D,

flower: C, front view; D, half

flower. E. capsules; E, seed.

Scale bars: A-E, 10 mm; F, 5

mm. Artist: John Manning.

help in the field, and to both the Western Cape Nature

Conservation Board and the Northern Cape Nature

Conservation Service for collecting permits.

REFERENCES

BOLUS, L. 1929. Plants — new and noteworthy. South African

Gardenin!> & Country Life 19: 123.

GOLDBLA'IT. P. 1990. Status of the southern African Anapalina and

Antholyza (Iridaceae) genera, based solely on characters for

bird pollination, and a new species of Tritoniopsis. South

African Journal of Botany 56: 577-582.

GOLDBLATT, P. & MANNING, J.C. 2000a. Cape plants. A conspec-

tus of the Cape Flora of South Africa. Strelitzia 9.

GOLDBLATT, P. & MANNING, J.C. 2000b. The long-proboscid fly

pollination system in southern Africa. Annals of the Missouri

Botanical Garden 87: 146-170.

GOLDBLATT, R, BERNHARDT, P, & MANNING, J.C. 2000.

Adaptive radiation of pollination mechanisms in the African

genus Ixia (Iridaceae: Crocoideae). Annals of the Missouri

Botanical Garden 89: 564—577.

LEWIS, G.J. 1959. South African Iridaceae. The genus Tritoniopsis.

Journal of South African Botany 25: 319-355.

Bothalia 31.2 (2001)

181

LEWIS, G.J. 1960. South African Iridaceae. The genus Anapalina.

Journal of South African Botany 26: 51-72.

MANNING, J, & GOLDBLATT, P. In press. The pollination of

Tritoniopsis parviflora (Iridaceae) by the oil-collecting bee

Rediviva gigas (Hymenoptera: Melittidae): the first record of oil-

secretion in African Iridaceae. South African Journal of Botany.

PHILLIPS, E.P. 1951. The genera of South African flowering plants,

edn 2. Memoirs of the Botanical Survey of South Africa No.

25.

WHITEHEAD, V.B & STEINER, K.E. 1993. A new Rediviva bee

(Hymenoptera: Apoidea: Melittidae) that collects oil from

orchids,. African Entomology I: 159-166.

Bothalia 31,2: 183-187 (2001)

Studies in the liverwort family Aneuraceae (Metzgeriales) from

southern Africa. 2. The genus Riccardia and its type species, R. multi-

fida, with confirmation of its presence in the region

S.M. PEROLD*

Keywords: Aneuraceae, Riccardia Gray, Riccardiaceae, R. midtifida (L.) Gray, R. saccatiflora (Steph.) S.W.Amell, southern Africa, type species

ABSTRACT

A generic description of Riccardia Gray is presented, as well as confirmation of the local presence of its type species,

R. midtifida (L.) Gray, which is described and illustrated. Amell (1952; 1963) incoiTectly identified his collections of this

species as R. saccatiflora (Steph.) S.W.Amell; some notes on it are added.

INTRODUCTION

In my previous account of the Aneuraceae (Perold

2001), I distinguished between the two locally occurring

genera, Aneitra Dumort. and Riccardia Gray, and briefly

traced the taxonomic history of the family in our region,

as well as that of the areas to the north. Short descriptions

of the family and the genus Aneitra were presented, as

well as a detailed description and an illustration of A.

pingiiis (L.) Dumort.

In the present publication a short account of the genus

Riccardia is given, as well as a detailed description and

illustration of R. midtifida (L.) Gray. Lehmann (1829)

had previously reported the presence of this species and

two of its varieties on Table Mountain, and Gottsche et

al. (1844-1847: 788) mentioned its occurrence at the

Cape. Arnell ( 1952, 1963), however, did not allude to it

at all, having misidentified his collections of it as R. sac-

catiflora. Reports of its local occurrence by Muller

(1951-1958) and by Smith (1990), were probably based

on Lehmann’s work and the brief reference to it by

Gottsche et al.

Riccardia Gray, A natural arrangement of British

plants: 679, 683 (1821) as ‘Riccardiiis’, correction by

Trevis.: 785 (1874) to 'Riccardia'-, Hewson: 75 (1970);

Brown & Braggins: 25 (1989); Luruki: 323 (1991);

R.M.Schust.: 579 (1992); Baton: 540 (1999); Bednarek-

Ochyra et al.: 178 (2000). Lectotype: Riccardia midtifi-

da (L.) Gray {Jimgermannia multiflida L.).

Synonyms are given in Hewson ( 1970); Brown & Braggins (1989)

and R.M. Schust. ( 1992).

Thalli prostrate to ascending, pale to dark green, cell

walls sometimes turning brown, rather delicate, somewhat

fleshy or not, short or long, length up to 40 mm, sometimes

more, narrow, 0.5-2.0 mm wide, lingulate to linear, apices

truncate, rounded or emarginate, margins obtuse to acute.

Branching lateral, pinnate to quadripinnate, rarely multipin-

nate, stolons sometimes present. Dorsal epidermal cells

* National Botanical Institute, Private Bag XlOl, Pretoria 0001.

MS. received; 2001-07-02.

thin-walled, smaller than internal cells. Oil bodies 1-3 in

each epidemial cell, sometimes more, rarely absent, ovoid,

spherical or ellipsoid, 12.5-22.5 x 10.0-12.5 pm, also pre-

sent, although larger, in internal cells, composed of numer-

ous fine globules, opaque, black or brown. Cross section of

branches plano-convex, biconvex or concavo-convex,

rarely circular, 3-9 cells thick medianly, at mai'gins acute to

winged or not. Mucilage papillae ventral, in 2 acropetal

rows, one on each side of midline and at branch apices.

Rliizoids usually present, numerous on ventral surface, uni-

cellulai'. Ase.xiial reproduction by 2-celled gemmae, endoge-

nously produced by epidermal cells, frequently present.

Monoicous, dioicous or synoicous. Male branches

lateral on main axis or on pinnae/primary branches, linear

to ligulate, sometimes in sympodial clusters, antheridia

in 2 regular rows, in chambers usually separated by uni-

stratose cell walls, dorso-lateral wing up to 8 cells wide.

Female branches short, lateral on main axis or on pinnae,

archegonia in 2 rows, protected by fringe of finger-like

or lamellate paraphyses. Shoot-calyptra large and fleshy,

up to ± 5 mm long, 0.7-1 . 1 mm wide, wall 5 or more cell

layers thick, surface smooth to rough, apical cells, i.e.

corona, poorly to well developed. Seta in cross section

with 4 inner and 12 outer cell rows (4 cells diam.).

Capsule ellipsoidal, 4-valved, wall bistratose, cells of

external layer with thickenings on adaxial radial and

inner tangential walls, cells of inner layer mostly without

thickenings, although the adaxial radial wall is some-

times slightly thickened. Spores spherical, finely

scabrate, small, 9-15 pm diam. Elaters 120-350 pm

long, 8-10 pm wide, unispiral.

The genus Riccardia, was named in honour of Vincento

Riccardi, a resident of Llorence during the early years of

the 18th century, and a donor to Micheli’s (1729) Nova

plantaruni genera.

Riccardia multiflda (L.) Gray, A natural arrange-

ment of British plants 1: 684 (1821); A. Evans: 22, 23

(1937); Mull.Lrib.: 499 (1954); R.M.Schust.: 316 (1987);

R.M.Schust.: 618 (1992); Baton: 541 (1999). Type: Europe.

Aneitra midtifida Dumort.; 115 (1822); Mull.Frib.; 336 (1908);

Macvicar; 54 (1926).

3 mm

184

Bothalia 31,2 (2001)

RGURE 1. — Riccardia multifida. A, thallus with axis and lateral branches, 2- or 3-pinnate; B, median dorsal epidermal cells (solid lines) of ultimate

.segment of axis without oil bodies, but present in large subdorsal cells (stippled lines); C, ultimate pinnule, some marginal cells with oil bodies,

medianly opaque; D, margin of ultimate pinnule, oil bodies mostly absent in outer, but present in inner cell rows; E, F, c/s axis at ± middle of

ultimate segment; G-J, c/s ultimate pinnules; K, ventral surface of part of pinnule, with mucilage papillae at apex. L, M, antheridial branch: L,

from above; M, c/s. N, synoicous branch; O, gynoecial branch with paraphyses; P, Q, calyptra: Q, c/s wall. R, c/s seta; S, capsule with 4 dehisced

valves. T-V, valve: T, cells of epidermal layer in external longitudinal view; U, cells of inner layer in internal longitudinal view; V, c/s part of

bistratose wall (thickenings partly after Evans 1937). Wall: mw, median; ot, outer tangential; it, inner tangential; adr, adaxial radial; abr, abaxial

radial. W1 , W2, spores; X, elater. PeroU & Koekemoer 4449-4452-. A, C, F-M, P, W1 , W2, X, 4449; B, 4452; D, R, 4450; E, N, O, Q, S-V, 4451.

Bothalia 31,2 (2001)

185

Jimgermannia midtifida L.; 1136 (1753).

For a complete list of synonyms, Schuster ( 1992) may be consulted.

Thalli prostrate to suberect, in loosely or densely over-

lying and intricately intertwining mats, rather light green,

sometimes at maturity cell walls brown, opaque, with nar-

row, translucent margins (Figure lA); when dry, dark

green; medium-sized to large, flat and flabellate above.

Main axis dorsally plane to convex or slightly concave,

often retaining its dominance, up to 27 mm long, 0.82-1 .05

mm wide, apically rounded or slightly emarginate. evenm-

ally narrowing somewhat toward base, margins acute and

unistratose for a width of 1 or 2 cells. Primary branches

lateral, opposite or subopposite, 0.8-1. 4 mm between suc-

cessive ones, obliquely spreading, often overlapping, regu-

larly pinnate with pinnae well developed, linear, distally

with 2 or 3 paired pinnules, proximally with 4 or some-

times even 5 paired pinnules, older, primary branches/pin-

nae up to 4.7 mm long, 590-650 pm wide, secondary

branches/pinnules 0.75-1.6 mm long, 475-530 pm wide;

from base of main axis or from apices of secondary branch-

es, flagelliform branches sometimes produced. Dorsal epi-

dermal cells in median part of apical segment of main axis

(Figure IB), from above 5-7-sided, thin-walled,

62.5- 105.0 X 25.0-37. 5(-55.0) pm, subdorsal cells larger,

4—7-sided, 155.0-262.5 x 62.5-87.5 pm, subventral cells

170.0-237.5 X 60-85 pm, ventral epidermal cells 75-100

X 30.0-47.5 pm. Oil bodies rare or absent in dorsal and

ventral epidermal cells, absent in up to 70% of marginal

cells, present in internal cells, 1 or occasionally 2 per cell,

15-25 X 7.5-12.5 pm, subspherical, ellipsoidal or ovoid,

with rounded ends, light brown, opaque, composed of

numerous small globules. Margins of ultimate pinnules

crenulate (Figures 1C, D; 2A), outer cells smaller than

intramarginal ones, from above 35-55 x 35.0-47.5 pm, ±

subquadrate, with free walls bulging, intramarginal cells

often obliquely orientated, polygonal, 72.5-100.0 x

47.5- 52.5 pm, generally with 1 oil body each. Cross sec-

tion of main axis at middle of ultimate segment (Figure IE,

F) plano-convex, or weakly concavo-convex, sometimes

biconvex, 180 pm or 6(7) cell rows thick medianly, gradu-

ally tapering to unistratose margins, 1 or 2 cells wide, dor-

sal cells ± 27.5 pm thick, internal cells 37.5-50.0 pm thick;

cross section of ultimate pinnule of primary branch (Figure

IG-J ) 100 pm or 3 or 4 cells thick medianly, pellucid, unis-

tratose margins 2 or 3 cells wide. Mucilage papillae ventral

(Figure IK), clustered at slightly emarginate apex of

branch and in 2 spaced, acropetal rows, one on either side

of midline, 300-360 pm between successive ones, reddish,

club-shaped, 2-celled. above 45.0-62.5 x 25.0-32.5 pm,

narrowing to inconspicuous basal cell, non-persistent.

Rhizoids ventral on axes, sometimes also on pinnae, hya-

line, 7.5-15.0 pm wide. Asexual reproduction by gemmae

seemingly absent.

Heteroicous, with male, female and rarely synoicous

branches. Antheridial branches lateral, near base of pri-

mary pinnae, on one or both sides, occasionally at tips

of secondary pinnules, oblong-linear (Figure IL), ± 700

pm long, 350 pm wide, in cross section 240 pm high

(Figure IM), margins crenulate, with single, erect layer

of swollen cells, 62.5-77.5 x 45-50 pm, bearing 5 or

more pairs of antheridia, separated by unistratose cell

layer; very rarely synoicous (Figure IN), basally with

paired antheridia and apically with a few archegonia and

FIGURE 2. — Riccardia multifida: A, pinnule (living), with outwardly

bulging marginal cells along both margins; B, fully rehydrated

pinnule, (collected 1951), with marginal cells not bulging out-

wardly. R. saccatiflora: C, fully rehydrated pinnule (collected

late 1800s), with much smaller marginal cells not bulging out-

wardly, only one margin shown. A, Perold & Koekemoer 4449\

B, S.W.Amell 1952 (S); C, Rodriguez G10699 (G). A-C, x 1 14.

short, laciniate cilia at tip. Gynoecial branches short,

arising opposite base of primary branch on main axis,

or toward base of main axis, sometimes toward middle

of primary branch, between pinnae of 1st and 2nd

186

Bothalia 31,2 (2001 )

orders, or between pinnae of 2nd and 3rd orders; para-

physes between and surrounding archegonia (Figure

10), up to 437.5 pm long, composed of single strands of

9 or more cells, joined end to end; sometimes up to 3

female branches in a sympodial cluster. Calyptra

(Figure IP) clavate, 3. 2-5. 3 x 0.7-1. 1 mm, cross section

of wall up to 225 pm or 4-6 cells thick (Figure IQ), out-

ermost row with scattered, thick-walled papillae, 65-90

X 25-65 pm, extending upwards to apex, where they are

grouped together, forming a corona. Seta ± 12 mm long,

± 280 pm wide, with 4 inner and 12 outer cell rows, i.e.

4 cells diam. (Figure IR). Capsule ± cylindrical,

1200-1450 pm long, with 4 dehisced valves (Figure

IS) , 320-375 pm or 12-15 cells wide, bistratose; cells

of epidermal layer in external longitudinal view (Figure

IT) , 100.0-142.5 X 25-35 pm, with straight or oblique

end walls, and strong nodular thickenings; in cross sec-

tion (Figure IV) cells ± rectangular in shape, thicken-

ings on adaxial radial (adr) and inner tangential (it)

walls sometimes extending slightly to outer tangential

(ot) walls, mostly bands on one side of median wall

(mw) alternating with those on the other side, i.e. form-

ing a mirror image; inner cells in internal longitudinal

view (Figure lU), 1 12.5-132.5 x 25-35 pm, never with

nodular thickenings; in cross section (Figure IV) cells

faintly thickened on adaxial radial (adr) and sometimes

extending slightly on inner tangential (it) walls. Spores

(Figure IWl, W2) 15-20 pm diam., translucent, finely

scabrate, pinkish, with inner, iiregularly shaped, green

areas. Elaters (Figure IX) 245-390 pm long, 10-14 pm

wide, with single spiral band, tapering to unspiralled

tips. Elaterophores at valve apices (Figure IS), in per-

sistent tufts. Chromosome no.: n = 20.

Riccardia midtifida is a widespread species or species

complex which Schuster (1992) says, extends from the

northern Spruce-Fir zone southwards. According to

Schuster, two subspecies occur in North America, i.e. R.

midtifida subsp. midtifida, with a more northerly range

along the western states, and R. midtifida subsp. synoica

with a more southerly range along the eastern states.

Apart from occupying a different geographical range, the

latter subspecies is also distinguished from the former

autoicous one, by most of its inflorescences being bisex-

ual and by its preference for growing on rotting wood. R.

midtifida subsp. midtifida is further known from

Newfoundland, Greenland, Iceland, British Isles,

Macaronesia, Europe, North Africa, Madagascar,

Sikkim-Himalaya, West Java (Meijer 1959) and China

(Yunnan). Another subspecies, R. midtifida subsp. de-

crescens, is reported from Japan by Furuki (1991).

Reports of R. midtifida from the Falklands have been

excluded according to Engel (1990). As far as is known,

it also does not occur in Australia, New Zealand or in

South America, although there is an unconfirmed report

from the Andes (Gottsche 1864). Muller (1951-1958)

remarked that, ‘Die Angaben von der sudlichen Hemi-

sphare gehdren aber vielleicht nicht zu R. midtifida.'

Wigginton & Grolle (1996) also do not include it in their

Catalogue of the Hepaticae and Anthocerotae of suh-

Saharan Africa. Jones (1956), however, observed that a

specimen collected from the ‘Foresta di Tusu, Mt Kenya’

and described by Gola (1914), appeared to him to be

identical with R. midtifida. Schuster (1992), on the other

hand, thinks that R. midtifida may be the only Riccardia

species with a reasonably certain bipolar range, as a col-

lection from Campbell Island, south of New Zealand,

differs only slightly from holarctic plants, although he

concedes that it needs more study.

Schuster (1992), with his extensive knowledge and

customary astuteness, expressed the opinion that, of the

species that Arnell (1952, 1963) reported from South

Africa, R. saccatiflora ‘seems virtually inseparable from

R. midtifida'. After thorough examination of Arnell’s

collections, I share this view, although it must be admit-

ted that Arnell’s drawings do not show the regular bi- and

tripinnate branching generally exhibited by R. midtifida.

However, the narrow translucent margins of the ultimate

pinnules of his specimens are conspicuous (Figure 2B),

and may have misled him into thinking he was dealing

with R. saccatiflora. Otherwise, it is surprising that Arnell

made this mistake, as he should have been familiar with

R. midtifida, having treated it for the Illustrated moss

flora of Eennoscandia (Arnell 1956).

Study of the type and five other Rodriguez collections

of R. saccatiflora from Reunion (Bourbon), as well as

Stephani’s leones (1985) nos. 000229 and 000230,

showed that the unistratose marginal cells of the ultimate

pinnules form a distinct wing, but are small (Figure 2C);

they measure 27.5-32.5(-45.0) x 10-20(-35) pm, where-

as the intramarginal cells are only slightly larger,

37.5^2.5 X 25-30 pm. In cross section the ultimate pin-

nules are 4 or 5 cell layers or 100 pm thick medianly and

the main axes 8 cell rows or 300 pm thick. Stephani

(1890) referred to the marginal cells as ‘multo-minores’.

Arnell (1963) reported the oil bodies of his ‘/?. saccat-

iflora' collections to be ‘dark, large, composed of very

small droplets, one per cell’. In their examination of R.

saccatiflora specimens, Meenks & Poes (1985) found

that their oil body data agreed well with Amell’s observa-

tions, but added that the oil bodies seem ‘rather persistent,

since they still occur in some older herbarium collec-

tions’. This is not the case in R. midtifida, where the oil

bodies are not persistent. Meenks & Poes (1985) only cite

the literature records from South Africa given by Arnell

(1963: 92). Their reports of R. saccatiflora from East

Africa are, however, accepted for the time being.

Paton (1999) states that, in British R. multifida, the

occasional synoicous inflorescence occurs on the same

thallus as unisexual and paroicous inflorescences, but

that such populations are not referable to the American

subspecies synoica R.M.Schust., in which nearly all the

inflorescences are bisexual. The same criteria apply to

the South African plants, where synoicous inflorescences

are very rare.

The new Perold & Koekemoer collections of R. multi-

fida are from Diepwalle Eorest Station, near Knysna, in

the southeastern part of Western Cape (Figure 3). This is

the same vicinity as Gouna Forest, Bracken Hill, Buffels

Nek and Garden of Eden, from where Arnell (1952)

recorded several of his so-called R. saccatiflora gather-

ings, which have now been transferred to R. multifida.

R. multifida specimens are distinguished by the fol-

lowing characters: 1, the thalli are markedly feathery,

with rather crowded and generally very regular, pinnate

Bothalia 31,2 (2001)

187

branching; 2, the ultimate and lateral pinnules are thin,

linear and paralled-sided, with the outer 3 cell rows unis-

tratose and translucent on either side of the thicker,

opaque, median region; 3, the ultimate pinnules have

crenulate distal margins, with the marginal cells bulging

externally and smaller than the intramarginal cells; 4, the

oil bodies are mostly absent from the marginal and epi-

dermal cells of the median region, but present in the inter-

nal cells; 5, the epidermal cells of the median part are

generally narrower than in allied species, i.e. 25.0-37. 5(-

55.0) pm wide; 6, the plants are rarely synoicous.

ECOLOGY

Riccardia midtifida plants grow on soil, twigs, dead

leaves or decaying wood in forested areas at streambanks

or seepages, i.e. in constantly wet or damp sites in ± dif-

fused light.

ACKNOWLEDGEMENTS

The curators of BOL. G and S are thanked for the loan

of specimens. I also express my sincere gratitude to Ms

M. Koekemoer, curator of PRE, for all her help with

fieldwork, and to the referees for their helpful sugges-

tions and advice. The artist. Ms G. Condy, the photogra-

pher, Mrs A. Romanowski and the typist, Ms D. Maree

are thanked for their valued contributions.

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EVANS, A.W. 1937. The structure of the capsule wall in certain species

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MEENKS, J.L.D. & POCS, T. 1985. East African Bryophytes IX.

Aneuraceae. Abstracta Botanica 9: 79-98.

MEIJER.W. 1959. Notes on species of Riccardia from their type local-

ities in western Java. Journal of the Hattori Botanical Labo-

ratory 21: 73, 74.

MICHELL PA. 1729. Nova plantarum genera. Paperini, Florence.

MULLER, K. (Mull.Frib.) 1905-1916. Die Lebermoose Deutschlands,

Oesterreichs und der Schweiz. Dr L. Rabenhorst's Krypto-

gamen-Flora 2, edn 6, part 1: 1-870. Geest & Portig, Leipzig.

MULLER. K. (Mull.Frib.) 1951-1958. Die Lebermoose Europas. Dr

L. Rabenhorst's Kryptogamen-Flora 6, edn 3: 492-505. Geest

& Portig, Leipzig.

PATON, J.A. 1999. The liverwort flora of the British Isles. Harley

Books, Colchester.

PEROLD, S.M. 2001. Studies in the liverwort family Aneuraceae

(Metzgeriales) from southern Africa. I. The genus Aneura and

its local representative. Bothalia 31: 167-173.

SCHUSTER. R.M. 1987. Studies on Metzgeriales: 1. North American

Aneuraceae. Journal of the Hattori Botanical Laboratory 62:

299-329.

SCHUSTER, R.M. 1992. The Hepaticae and Anthocerotae of North

America 5: 1-854. Field Museum of Natural History, Chicago.

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University Press, Cambridge.

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tation Company, Leiden.

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italiane. Rendiconti Reale Istituto Dnnhardo di Scienze e Lettere,

Ser. 2,7: 776-786.

WIGGINTON, M.J. & GROLLE, R., Supplemented by GYARMATl,

A. 1996. Catalogue of the Hepaticae and Anthocerotae of sub-

Saharan Africa. Bryophytorum Bibliotheca 50. Cramer, Berlin,

Stuttgart.

SPECIMENS EXAMINED

Riccardia midtifida

Arnell ISIS Buffels Nek, IS92», 1731 Gouna For. (BOL), 1839

Diepwalle For. Res.(BOL), 1843** (BOL. PRE), 1844* *, 184S (BOL),

1918 (PRE), 1929 (BOL), I9S2 (S), 1974 Bracken Hill For. (BOL),

2079, 2133 Garden of Eden (BOL).

Perold & Koekemoer 4449, 44S0, 44S1, 44S2, Diepwalle Forest

Reserve, Knysna District, Western Cape.

Riccardia saccatifiora

Rodriguez G010697 (G) (type specimen), G010698 (G), G010699

(G), G010700 (G), G010702 (G), G010703 (G) Bourbon (Reunion).

• on decaying wood.

* ‘A mile south of Diepwalle Forest Reserve.

Bothalia31,2: 189-212 (2001)

Notes on African plants

VARIOUS AUTHORS

IRIDACEAE

TWO NEW RENOSTERVELD SPECIES OF CROCOIDEAE FROM SOUTH AFRICA

INTRODUCTION

Southern Africa and the Cape Floral Region in partic-

ular is a major centre of diversity for Iridaceae.

Approximately 70% of the species of Iridaceae recorded

from southern Africa occur in the Cape Region and most

are endemic to it (Goldblatt 1991; Goldblatt & Manning

2000). The family is especially well represented in the

west and southwest of the region, which has a true sum-

mer-dry climate (Manning et al. in press). Local ende-

mism is a common feature among the species in the

region. Iridaceae display a high degree of substrate

fidelity (Goldblatt & Manning 1996) and edaphic and

topographic diversity appear to have played a significant

role in speciation in the subcontinent (Goldblatt &

Manning 1998). Although the great majority of species

recorded from the Cape Floral Region are typically

restricted to fynbos vegetation, the vegetation type

known as renosterveld historically dominated much of

the Cape lowlands. This fine-leaved shrubland occurs on

nutrient-intermediate, fine-grained clays derived from

shales of the Malmesbury and Bokkeveld Formations

and is particularly rich in geophytic species. Now large-

ly transformed by agricultiue (Low & Rebelo 1996),

renosterveld was commonly found along the coastal

forelands as well as in inland drainage basins that are

often isolated from one another. The drainage basin of

the Breede River, which forms the region known as the

Worcester Karoo, is rich in local endemics. Among these

are Ixia collina, known only from near Aan de Dooms,

southeast of Worcester, 7. vanzijliae, distributed between

Worcester and Roberston, and Sparaxis maculosa,

restricted to the valley of the Hoeks River, a minor tribu-

tary of the Breede River, near Villiersdorp in the south-

western comer of the Worcester Karoo. Two additional

local endemics restricted to the renosterveld in this area

have since been discovered and are described here. It is

becoming evident that the Worcester Karoo, known

locally as the Bosjesveld at least since 1815 when the

explorer William Burchell recorded the name (Burchell

1822), is a distinct enclave of endemism for Iridaceae

and especially the genus Ixia. Fortunately extensive

stretches of indigenous renosterbos shrubland still

remain undisturbed in this area although it is slowly

being removed to establish orchards and vineyards.

Freesia fucata J.C. Manning & Goldblatt, sp. nov.

Plantae 150-300 mm altae, cormo conico 15-20 mm

diam. tunicis pallide reticulatis, folds 5-6 inclinatis vel

suberectis linearibus 80-250 x 4-5 mm, caule usitate

ramoso minute papillate, spica inclinata 4-9-florum,

floribus (30-)35-47 mm longis fragrantibus albis

exteme atropurpureis suffusis tepalis inferioribus auran-

tiacis notatis, tubo perianthii 20-30 mm longo infime

filiforme 6-7 mm longo, tepalis inaequalibus, dorsale

grandiore suberecto ovate 11-18 x 8-11 mm, inferi-

oribus recurvatis, filamentis 13-15 mm longis, antheris

7-8 mm longis, stylo diviso prope apicem antherarum.

TYPE. — Western Cape, 3319, Villiersdorp Dist., 5.5

km along Doornrivier Road off Villiersdorp- Worcester

Road, Farm Die Hoek, (-CD), 10-07-2000, Manning

2271 (NBG, holo.; MO, PRE, iso.).

Plants 150-300 mm high. Conn conic, 15-20 mm

diam., tunics pale, medium-textured, reticulate, accumu-

lating with age and forming neck around base of stem.

Leaves 5 or 6, inclined to suberect, linear, tapering

above, acute, 80-250 mm long, usually about as long as

stem, 4-5 mm wide, cataphylls flushed deep purple.

Stem minutely papillate, flexed outward above upper-

most sheath and then suberect, usually with one or two

branches. Spike lightly decumbent, 4-9-flowered; bracts

soft, herbaceous, green with hyaline margins, tricuspi-

date, 6-10(-13) mm long; inner bracts slightly shorter,

forked. Flowers (30-)35-47 mm long, strongly violet-

scented, white flushed dull purple on reverse of tepals,

base of lower tepals yellow-orange with dark midline;

perianth tube 20-30 mm long with basal narrow part 6-7

mm long, widening rather abruptly into wider, flared

upper portion; tepals unequal, the dorsal largest,

suberect, ovate, 1 1-18 x 8-11 mm, upper laterals slight-

ly narrower, oblong, lower tepals recurved, lower laterals

narrowly ovate, 10-14 x 6-8 mm, lower median oblong,

10-16 X 5-6 mm. Filaments arcuate, inserted at base of

widened upper portion, included, 13-15 mm long;

anthers 7-8 mm long. Style dividing at or slightly beyond

anther apices, branches deeply forked. Capsule oblong,

three-lobed, (6-)9-12 x (6-)7-10 mm. Seeds globose

with inflated chalaza and raphe, rugose, glossy reddish

brown, 2. 5-3.0 mm diam. Flowering time: July. Figure 1.

Distribution and biology: the species is known from

several populations in the valley of the Hoeks River, in

the southwestern comer of the Worcester Karoo, south of

Villiersdorp (Figure 2). It occurs on clay soils derived

from shales of the Bokkeveld series in open shmbland

dominated by the renosterbos, Elytropappus rhinocerotis.

History: first collected at the type locality in 1975 by

the Worcester dentist, I.B. Walters, who had a keen inter-

est in the local flora and collected extensively in the

Worcester Valley, this highly fragrant species is known

only from this small area.

190

Bothalia 31,2 (2001)

FIGURE 1. — Freesia fucata. A,

whole plant; B, flowering

spike; C, D, flower: C, front

view; D, half flower. E, outer

bract (left) and inner bract

(right); F, capsules; G, seed.

Scale bar: A-F, 10 mm; G, 1.5

mm. Artist: John Manning.

Diagnosis and relationships: F fucata has the minute-

ly papillate-puberulous stem, green bracts and narrowly

ovate lower tepals characteristic of section Viridibractea

(Goldblatt 1982). Within the section it is somewhat inter-

mediate between F. alba (G.L.Mey.) Gumbleton and F

caryophyllacea (Burm.f.) N.E.Br. It has the narrow, erect

leaves of the former and the bilabiate, white flowers

flushed with purple of the latter. F. fucata is, however,

unique in the genus in its distinctly tricuspidate outer

bracts. F. alba, which is characterized by its almost

actinomorphic perianth, is essentially a species of the

southern Cape coastal forelands, occurring along forest

margins or in open scrub in stony, sandstone-derived

soils. F. fucata and F caryophyllacea, in contrast, both

occur in renosterbos shrubland on stony clay soils. The

rather variable F. caryophyllacea typically has more or

less prostrate, usually rather broad, typically 10-16 mm

wide, lanceolate and obtuse leaves. Populations in the

west, centred around Caledon and Bredasdorp, however,

have acute and often quite long leaves, 5-9 mm wide, but

these are never as narrow or as long as in F fucata. The

Bredasdorp populations, which are found on limestone

outcrops, were regarded by Goldblatt (1982) as a distinct

species, F. elimensis L. Bolus, of possible hybrid origin

between F. alba and F. caryophyllacea. Subesquent col-

lections, however, suggest that it is no more than a local

edaphic form of F. caryophyllacea (Goldblatt &

Manning 2000). The possibility that F. fucata represents

a hybrid between F. alba and F. caryophyllacea can be

discounted since neither of the putative parents has the

curious tricuspidate outer bracts of F. fucata, nor has

either been collected in the immediate vicinity.

Bothalia 31,2 (2001)

191

FIGURE 2. — Distribution of both Freesia fucata and I.xia atrandra in

Western Cape.

The more common and widespread F. refracta (Jacq.)

Klatt, which is well known from the Breede River Valley,

also resembles F. fucata in its upright habit and narrow

leaves and the first collection of the new species was

assigned to it in error. F refracta is easily identified,

however, by its dry and membranous floral bracts and

smaller flowers, 25-35(^0) mm long, with the dorsal

tepal more prominent than the relatively small, cordate

lower tepals that have their margins conspicuously

curved upwards.

Additional material examined

WESTERN CAPE. — 3319 (Worcester): near Die Hoek. Doom-

rivier, (-CD), 23-07-1975. Walters 157 (NBG).

Ixia atrandra Goldblatt & J.C. Manning, sp. nov.

Plantae 250-500 mm altae, cormo globoso tunicis

reticulatis, folds (3)4 lanceolatis basalibus 6-10 mm

latis, caule simplice vel 1-ramoso, spica congestis

(3)4-10-florum, bracteis scariosis translucentibus 5-6

mm longis, floribus camels vel cremeis centro atrocolore

hypocrateriformibus, tubo perianthii filiforme 6-9 mm

longo, tepalis obovatis 11-14 x 6-8 mm, filamentis

2. 5-3.0 mm longis ascendentibus, antheris oblongo-

sagittatis 5-6 x 1. 5-2.0 mm thecis latrorsibus nigris,

pollinibus flavis, stylo diviso ad apicem tubi. Figure 3.

TYPE. — Western Cape, Villiersdorp Dist., about 5

km along Doomrivier Road off Villiersdorp- Worcester

Road, Farm Die Hoek, (-CD), 3-10-2000, Goldblatt 11604

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

Plants 250-500 mm high. Corms globose, 15-20 mm

diam., tunics of medium-textured, wiry reticulate fibres.

Cataphylls membranous, upper one reaching shortly

above ground level and then flushed reddish purple.

Leaves usually 4, sometimes 3, all basal, lanceolate,

6-10 mm wide, usually reaching to about middle of

stem, uppermost leaf usually entirely sheathing, margins

and midrib hyaline, lightly thickened, plane or lightly

twisted in upper half. Stem erect, unbranched or with one

FIGURE 3. — Ixia atrandra. A, corm and leaves; B, flowering spike; C,

anthers and style branches. Scale bar: A-C, 10 mm. Artist: John

Manning.

ascending branch, 0. 5-0.8 mm diam. below base of

spike. Spike crowded, (3)4- 10- flowered; bracts scarious,

translucent, veins more or less pink above, outer 5-6 mm

long, obscurely three-dentate, inner bract about as long

as outer or slightly shorter, bicuspidate. Flowers

192

Bothalia 31,2 (2001)

hypocrateriform, pink or rarely cream-coloured, lower

third of tepals yellowish or pinkish ochre-coloured with

an outer band of deep lilac, unscented; perianth tube fili-

form and clasping style for its entire length, 6-9 mm

long, widened only in upper 1 mm; tepals obovate, some-

what narrowed into short claw below, spreading with dis-

tal margins curving upward, 11-14 x 6-8 mm. Filaments

inserted at apex of tube and occluding throat, blue to

brownish, 2. 5-3.0 mm long, weakly diverging, free or

connate at base; anthers oblong-sagittate, connective

broad and thecae restricted to margins, dehiscing lateral-

ly, 5-6 X 1. 5-2.0 mm (at anthesis), initially erect, later

diverging and becoming slightly twisted, black; pollen

yellow. Ovary ovoid. 2 mm long; style straight and erect,

dividing at or just above mouth of tube, branches blue,

arching outward, 3^ mm long, ultimately reaching base

of anthers. Flowering time: late September and October.

Distribution and biology: currently known from two

populations in the valley of the Hoeks River in the south-

western corner of the Worcester Karoo (Figure 2), Ixia

atrandra grows on clay soils derived from shales of the

Bokkeveld series in open shrubland dominated by the

renosterbos, Elytropappus rliinocerotis. The flowers of I.

atrandra display the typical adaptations for pollination

by monkey beetles (Scarabaeidae: Rutelinae). These

include the congested, subcapitate inflorescence, bright-

ly coloured bowl-shaped flowers with central dark mark-

ing, lack of nectar or fragrance and dark anthers. The

broadened connectives are part of this syndrome and are

matched by remarkably similar sagittate anthers in some

beetle-pollinated species of Babiana, particularly B. vil-

losa, and B. melanops.

History: this species was first collected in October

1996 as a voucher for studies on beetle-pollination, but

the material was inadequate for formal description. A

later collection from nearby serves as source of the type

material.

Diagnosis and relationships: Ixia atrandra appears to

be most closely allied to /. rouxii G.J. Lewis, with which

it shares pink flowers with a large central stain, tepals

which are narrowed and almost clawed below and black

anthers with widened connectives. I. rouxii is restricted

to the upper reaches of the Breede River and the con-

tiguous Berg River, between Porterville and Wolseley.

An unusual feature of both species, but particularly

prominently developed in 1. atrandra, is the presence of

dark blue amber-like deposits on the anther connective.

These appear to be hardened secretions from the epider-

mis. I. atrandra differs from I. rouxii in its four (occa-

sionally three), lanceolate leaves 6-10 mm wide, typical-

ly unbranched stem (or with a single, ascending branch)

and distinctive oblong-sagittate anthers with a broad

connective about 1 .5 mm wide and the thecae dehiscing

laterally. I. rouxii typically has five to six (rarely four)

almost linear leaves 2-3(-6) mm wide and is distinctive

in its sharply spreading branches, and while the anther

connectives are widened, they are less than 1 mm wide,

and the anthers dehisce outwards. A third species with

pink or white flowers with a prominent dark centre and

black anthers, 7. versicolor G.J. Lewis, is probably also

part of this alliance. A local endemic restricted to damp,

gravelly renosterveld flats at Gordon’s Bay, it is distin-

guished from the other two species by its conspicuously

thickened, crenulate leaf margins and elliptic tepals that

are not narrowed into claws at the base. These three

species appear to be geographic segregates restricted to

lowland renosterbos shrubland.

Additional material examined

WESTERN CAPE. — 3319 (Worcester): Brandvlei hills south of

Worcester, clay ground in renosterveld, (-DD), 26-10-1996, Goldblatt

10568 (MO).

REFERENCES

BURCHELL, W.J. 1822. Travels in the interior of southern Africa.

Reprinted in 1953 by The Batchworth Press, London.

GOLDBLATT, P. 1982. Systematics of Freesia Klatt (Iridaceae).

Journal of South African Botany 48: 39-91.

GOLDBLATT, P. 1991. An overview of the systematics, phylogeny and

biology of the African Iridaceae. Contributions from the Bolus

Herbarium 13: 1-74.

GOLDBLATT, P. & MANNING J.C. 1996. Two new edaphic endemic

species and taxonomic changes in Gladiolus (Iridaceae) of

southern Africa, and notes on Iridaceae restricted to unusual

substrates. Novon 6: 172-180.

GOLDBLATT. P. & MANNING, J.C. 1998. Gladiolus in southern

Africa. Femwood Press, Cape Town.

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

of the Cape Flora of South Africa. Strelitzia 9.

LOW, A.B. & REBELO, A.G. 1996. Vegetation of South Africa,

Lesotho and Swaziland. Department of Environmental Affairs

& Tourism, Pretoria.

MANNING, J., GOLDBLATT, P. & SNIJMAN. D. in press. Cape

bulbs and their allies. Timber Press, Oregon.

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

* National Botanical Institute, Private Bag X7, 7735 Claremont, Cape

Town.

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

PO. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2000-10-23.

HYACINTHACEAE

THE GENERIC DELIMITATION WITHIN HYACINTHACEAE, A COMMENT ON WORKS BY F. SPETA

INTRODUCTION

The definition of genera and the assignment of

species to genera within the family Hyacinthaceae, or

subfamily Scilleae of the family Liliaceae, have troubled

taxonomists since Linnaeus. The group is poor in quali-

tative characters, which has made it difficult to maintain

stable genera based on good diagnostic characters.

Species have often been moved from genus to genus

either due to different opinions on generic delimitation or

to misinterpretation of characters. Recently two works

by Speta (1998a, b) have appeared that have addressed

Bothalia 31.2 (2001)

193

generic delimitation within Hyacinthaceae. The first is a

paper called ‘Systematische Analyse der Gattung Scilla

L. s.l. (Hyacinthaceae)’ (Speta 1998a), the second is the

treatment of Hyacinthaceae for ‘The families and genera

of flowering plants’ (Speta 1998b). In these works the

author has published a new generic delimitation within

the family. As I have worked on sub-Saharan genera of

Hyacinthaceae for several years (Stedje 1987, 1988,

1996. 1997, 1998, 2000; Stedje & Thulin 1995, and ref-

erences therein), both as a biosystematist, and a molecu-

lar and floristic taxonomist contributing to new floras, I

feel the need to comment on Speta’s (1998a, b) new

generic delimitations. A review of different taxonomists’

opinions on the generic delimitation of Hyacinthaceae is

given in Stedje (2001).

GENERAL COMMENTS ON THE WORKS OF SPETA ( 1998a. b)

In Speta (1998a) the family Hyacinthaceae is split

into five subfamilies of which four are new: Chloro-

galoideae Speta, Oziroeideae Speta, Urgineoideae Speta

and Omithogaloideae Speta. Of these, Oziroeideae,

Urgineoideae and Hyacinthoideae are treated further

with descriptions and species lists of selected genera.

Later DNA analysis excludes Chlorogaloideae from

Hyacinthaceae (Pfosser & Speta 1999). Oziroeideae is

confined to South America, and I will not deal with it fur-

ther here. Several of the genera in Speta (1998a) are

described as new to science. There is extensive splitting

of the ‘traditional’ genera Urginea Steinh. (or Drimia

Jacq. if a wider generic circumscription is used), Orni-

thogalum L. and Scilla L. into newly described genera or

reinstated ones. These new genera and generic reinstate-

ments are presented practically without any discussion or

justification. The groupings are said to have support in

DNA sequence data, but no cladograms or reference to

publication of these data are included. Later, in 1999, a

paper including a phylogenetic analysis of Hyacin-

thaceae based on chloroplast DNA sequences appeared

(Pfosser & Speta 1999). Unfortunately only a few sub-

Saharan species are included here, and for most genera

occurring in Africa south of the Sahara, only one species,

if any, is analysed. When two or more species are ana-

lysed. the genera are, with one exception (Drimiopsis),

para- or polyphyletic. Furthermore no diagnostic key is

provided. A key is given in Speta (1998b), but in that

work several of the genera of Speta (1998a) are not

included (see below for further comments on this issue).

The definition of the genera of Speta’s (1998a) new treat-

ment of the family Hyacinthaceae is in other words very

unclear. Unfortunately, throughout this work virtually no

discussion is given prior to his conclusions, making a

meaningful discussion between taxonomists difficult.

Furthermore, insufficient details are given of the materi-

al studied: no indication of the type of material seen,

whether herbarium specimens or living plants, or how

many specimens were studied for which species.

COMMENTS ON THE SUBFAMILY URGINEOIDEAE

The treatment of this subfamily is rather confusing.

Altogether 12 genera are listed as belonging to the sub-

family Urgineoideae in the introductory part of Speta

(1998a: 53). Two of these genera (Bowiea Hook f. and

Schizobasis Baker) are not treated further in the part con-

taining generic descriptions, while three genera are

described here under subfamily Urgineoideae, but not

listed on p. 53: the reinstated genus Fusifiliim Raf., and

the new genera Igidia Speta and Urginavia Speta. In

Speta (1998b) four of the genera of Speta (1998a) are not

included. These are Fusifiliim, Urginavia, Cliarybdis

Speta and Ebertia Speta. They may be included under

what in the diagnostic key of Speta (1998b) is referred to

as Urginea sd. If the manuscript of Speta (1998b) was

completed before the manuscript of Speta (1998a), one

would expect to find a key to these critical genera includ-

ed in the latter. This would help to clarify the distinctions

between these genera.

Drimia sensu lato as a ‘Monstergattiing ’

Botanists have in recent years advocated either a wide

(Jessop 1977; Steam 1978; Stedje 1987, 2000) or a nar-

row (Stilton 1976; Obermeyer 1980, 1981; Hilliard &

Burtt 1982) circumscription of the genus Drimia. I have

discussed this issue in detail elsewhere (Stedje 1987,

2000), and will not repeat that discussion here. Speta

(1998a) calls Drimia sensu lato a ‘Monstergattung’. He

does not define this expression, but his intention is clear-

ly uncomplimentary. Drimia sensu lato consists of up to

100 species, a number quite modest when compared with

some other Angiosperm genera. There is of course vari-

ability within the genus, but not to the extent that it forms

a ‘rag-bag’ taxon with highly heterogeneous elements.

The genus is well defined by its winged seeds, an autapo-

morphy for Drimia s.l., and the spurred bracts, a synapo-

morphy of Drimia, Bowiea and Schizobasis.

Where have all the Drimia species gone?

Under each of the genera provided with descriptions

in Speta (1998a), a list of species is given. The species

list of the genus Ledebouria Roth is said to be incom-

plete, but this is not said of any of the other genera. One

may therefore interpret those other species lists as

intended to be complete. For Africa south of the Sahara

this leaves us with 15 species of Drimia (sensu lato)

recognized by either Jessop (1977) or Stedje & Thulin

(1995), which have not been taken into account by Speta

(1998a). Does this mean that the 15 species are not

regarded as belonging to Hyacinthaceae any more, or are

they simply forgotten? If so, how can 15 species be

neglected and why are names that Jessop (1977) regarded

as synonyms of some of these species included? How

can one possibly fit these forgotten species into genera

when, as mentioned above, the circumscription of the

genera is unclear and no key is given?

One species or three genera?

In the case of the genus Avonsera, discussed later, two

apparently discordant species were combined in a single

genus. There are also, in Speta’s work (Speta 1998a),

cases where apparently closely related species are segre-

gated into different genera. Four synonyms of Drimia

modesta (Baker) Jessop (sensu Jessop 1977) have here

been placed in three different genera, Urgineopsis

Compton, Thuranthos C.H.Wright and Fusifilum. The

194

Bothalia 31,2 (2001)

species are U. salted Compton, T. revoluta (Duthie) Speta,

F. dregei (Baker) Speta and F. gracilis (Duthie) Speta. As

already mentioned, different botanists must be allowed to

have different opinions as long as appropriate documenta-

tion is presented. This is, however, not done here.

COMMENTS ON THE SUBFAMILY ORNITHOGALOIDEAE

Several of the genera recognized in this subfamily

have previously been included in the genus Orni-

thogaliim L. The sub-Saharan Ornithogalum is, for

example, split into the genera Stellarioides Medicus,

Coilonox Raf., Eliokannos Raf. and Zahariadia Speta, a

new genus which is possibly monotypic. The first three

genera appear to coincide in part with Obermeyer’s

(1978) subgenera even if it seems that only a part of

Obermeyer’s subgenus Urophyllon is accommodated by

the genus Stellarioides. Where the rest of the subgenus

Urophyllon is to be placed, is unclear. The subfamily

Omithogaloideae is split into the tribes Dipcadieae and

Omithogaleae. These tribes are based on seed characters,

which are mainly quantitative, and they key out at lead

13 in Speta (1998b). The alternatives of this lead are not

very clear and it should be possible to simplify them. The

same applies to lead 12 which also refers to seed charac-

ters and is even less clear. The two alternatives of lead 1 3

are, for example: 1 ) Seeds flattened, orbiculate, D-

shaped, or elongate; tepals green, brown or yellow, or

whitish, with a green streak, versus 2) Seeds minute,

comma-shaped, glabrous or shortly pilose, rarely edged,

or large, elongate, with irregular edges, or globose to

ellipsoid; tepals white, with or without a green stripe, or

yellow or orange. Within this subfamily both tribes and

genera are defined largely on the basis of quantitative

characters of the seeds. For me it is difficult to under-

stand the necessity of this excessive splitting. Take, for

example, three species, the sub-Saharan O. temiifolium

F.Delaroche, the Moroccan O. sessiliflorum Desf. and

the Mediterranean O. narbonense L. These three species

are very similar in all characters, except possibly for

some differences in quantitative morphological charac-

ters between the two Mediterranean species, and in the

Mediterranean species having more turgid seeds than O.

temdfoliiim. I am not absolutely sure in which genera to

put these species as Speta does not treat them specifical-

ly. O. sessiliflorum in particular, does not quite fit in any-

where. As I interpret Speta’s new system, however, these

three species are put into two different tribes, Dipcadieae

and Omithogaloideae, and three different genera

Stellarioides, Cathissa Salisbury and Lonocomelos Raf.

respectively.

COMMENTS ON THE SUBFAMILY HYACINTHOIDEAE

For this subfamily the treatment is far less confusing

than for subfamily Urgineoideae, but there are some points

to be commented on. Altogether 38 genera are listed as

belonging to the subfamily Hyacinthoideae. Of these, just

21 are treated with full de.scriptions in the following text,

and there is no obvious reason for omitting the 1 7 others.

One of these 17 genera, Namopliila U. & D.Muller-

Doblies, is not mentioned at all in Speta (1998b).

The genus Avonsera Speta

The new genus Avonsera Speta is based on two

species, A. convallarioides (Perrier) Speta and A.

lachenalioides (Baker) Speta (Speta 1998a). A. conval-

larioides is a species endemic to Madagascar, originally

described in the genus Ornithogalum. Obermeyer (1978),

in endorsing the placement of the species in Ornitho-

galum, writes: ‘the characters of the flowers, capsules

and seeds agree with Ornithogalum ’. A. lachenalioides is

confined to KwaZulu-Natal and Eastern Cape and was

placed in Drimiopsis by Jessop (1972). It appears strange

to base a genus on two species with a rather disjunct dis-

tribution that have recently been placed in two distantly

related genera by two competent South African botanists.

There may of course be good reasons for doing so, but

one would expect a short discussion of the issue. I have

not seen such in either Speta (1998a) or Speta (1998b).

CONCLUSION

Speta’s (1998a, b) treatment of Hyacinthaceae is an

unfortunate case of excessive splitting of the family

which will create more confusion than clarity. Many of

the genera, especially the new ones are monotypic. His

conclusions have no clear proven basis, neither in cladis-

tic analyses, nor in more classical criteria for generic

delimitation such as giving emphasis to qualitative mor-

phological characters and using minor characters only to

preserve genera already recognized. A full review on this

issue is given by Stuessy (1990, and references herein).

Speta’s generic delimitation might fit the Mediterranean

and European species of the family, but I see substantial

difficulties when applied to sub-Saharan species.

REFERENCES

HILLIARD, O.M. & BURTT, B.L. 1982. Notes on some plants of

southern Africa chiefly from Natal: IX. Notes from the Royal

Botanic Garden Edinburgh 40: 247-298.

JESSOP, J.P, 1972. Studies in the bulbous Liliaceae in South Africa: 2.

Drimiopsis and Resnova. Journal of South African Botany 38:

151-162.

JESSOP, J.P, 1977, Studies in the bulbous Liliaceae in South Africa: 7.

The taxonomy of Drimia and certain allied genera. Journal of

South African Botany 43: 265-319.

OBERMEYER, A. A. 1978. Ornithogalum: a revision of the southern

African species. Bothalia 12: 323-376.

OBERMEYER, A. A. 1980. The status of Urginea epigea. Bothalia 13:

139.

OBERMEYER, A. A. 1981. A reappraisal of Urginea altissima.

Bothalia 13: 452, 453.

PFOSSER, M. & SPETA, F, 1999. Phylogenetics of Hyacinthaceae

based on plastid DNA sequences. Annals of the Missouri

Botanical Garden 86: 852-875.

SPETA, F. 1998a. Systematische Analy.se der Gattung Scilla L. s.l.

(Hyacinthaceae), Phyton {Horn} 38: 1-141.

SPETA. F. 1998b. Hyacinthaceae. In K. Kubitzki, The families and

genera of vascular plants 3: 261-285. Springer- Verlag, Berlin.

STEARN, W.T. 1978, Mediterranean and Indian species of Drimia

(Liliaceae): a nomenclatural survey with special reference to

the medicinal squill, D. maritima (syn. Urginea maritima).

Annales Musei Goulandris 4: 199-210.

STEDJE, B. 1987. A revision of the genus Drimia Jacq.

(Hyacinthaceae) in East Africa. Nordic Journal of Botany. 1:

655-666.

STEDJE, B. 1988. Cytotaxonomic studies within Hyacinthaceae with

special emphasis on karyotype evolution. Ph.D. thesis,

University of Oslo, Norway.

Bothalia31,2 (2001)

195

STEDJE, B. 1996. Hyacinthaceae. In R.M. Polhill, Flora of tropical

East Africa. Balkema, Rotterdam.

STEDJE, B. 1997. Hyacinthaceae. In S. Edwards, Sebsebe Demissew

& I. Hedberg, Flora of Ethiopia and Eritrea 6: 138-147. The

National Herbarium, Addis Ababa.

STEDJE, B. 1998. Phylogenetic relationships and generic delimitation

of sub-Saharan Scilla L. (Hyacinthaceae) and allied African

genera as inferred from morphological and DNA sequence data.

Plant systematics and evolution 211: 1-11.

STEDJE, B. 2000. Evolutionary relationships of the genera Drimia.

Thuranthos, Bowiea and Schiiobasis, elucidated by morpholog-

ical and chloroplast DNA sequence data. In K.L. Wilson & D.A.

Morrison, Proceedings of the Second International Conference

on Comparative Biology of the Monocotyledons, Sydney,

Australia, 1998, Vol. 1. Systematics and evolution of Monocots:

414-417. CSIRO Publishing, Collingwood, Australia.

STEDJE, B. in press. Generic delimitation of Hyacinthaceae, with spe-

cial emphasis on sub-Saharan genera. Proceedings of the 16"'

AETEAT Congress, 28 August to 2 September 2000. Systematics

and geography of plants 1 1 .

STEDJE. B, & THULIN, M. 1995. Synopsis of Hyacinthaceae in East

and North-East Africa. Nordic Journal of Botany 15: 591-601.

STIRTON, C.H. 1976. Thuranthos: notes on generic status, morpholo-

gy, phenology and pollination biology. Bothalia 12: 161-165.

STUESSY, TF. 1990. Plant taxonomy: the systematic evaluation of

comparative data. Columbia University Press, New York.

B. STEDJE*

* University of Oslo, Botanical Garden and Museum, P.O. Box 1172

Blindem, N-0318 Oslo, Norway. E-mail: brita.stedje@nhm.uio.no

MS. received: 2000-11-02.

DENNSTAEDTIACEAE-PTEROPSIDA

HYPOLEPIS VILLOSO-VISCIDA NEW TO THE FLORA OF SOUTHERN AFRICA

Recently, a Hypolepis collection from Genadendal

in the Western Cape, differing from H. sparsisora

(Schrad.) Kuhn came to my attention. This collection

differs in the distribution of hairs on the lamina, the

presence of glandular and receptacular hairs, and larger

stomata and spores. To determine whether the plants

may be recent introductions and to determine the size of

the population, I have subsequently visited the location.

A review of herbarium collections revealed that this

species, although scarce, also occurs elsewhere in

South Africa, but has always been erroneously deter-

mined as H. sparsisora. A morphological study showed

that these plants are synonymous with H. villoso-visci-

da (Thouars) Tardieu, a species also occurring on the

South Atlantic island groups of Gough and Tristan da

Cunha.

Key to the South African species of Hypolepis

Lamina with acicular and oblong hairs confined to axes and

veins; receptacle nude; stomata 28-(37. 11)^6 pm long;

spores 24— (29.04 )-38 x 15-(19.63)-26 pm ....//. sparsisora

Lamina with acicular and glandular hairs (rarely also oblong

hairs) on axes, veins and lamina surfaces; receptacle

usually with uniseriate hairs; stomata 38-(50.33)-64

pm long; spores 32-(38.01 )^6 x 20-(23.95)-30 pm . .

H. villoso-viscida

Hypolepis villoso-viscida (Thouars) Tardieu, Elore

de Madagascar et de Comores 5,1: 6, fig. 1, t. 3-5 (1958).

Polypodium villoso-viscidum Thouars: 33: (1808). Type:

Tristan d’ Acunha, Aubert du Petit-Thouars s.n. (P, holo.).

Cheilanthes viscosa Carmich.; 511 (1818). Type; Tristan da Cunha,

Carmichael s.n. (K, holo.; BM, iso.).

Specimens e.xamined

EASTERN CAPE. — 3225 (Somerset East); Somerset East,

Boschberg, 760 m, Nov. 1875, (-DA), MacOwan 1575 (SAM).

WESTERN CAPE. — 3418 (Simonstown); Orange Kloof, W-facing

slopes, ± 220 m, 24 Nov. 2000, (-BD), Roilx 3023 (NBG). 3419

(Caledon); Genadendal, Baviaans River, north bank alongside road

upstream of weir, 280 m, 2 June 2000, (-BA), Boucher 6515 (NBG);

Genadendal, Baviaans River, above 2nd weir, ± 300 m, 7 July 2000,

(-BA), Roux 3007, 3009, 3010, 3011 (NBG).

REFERENCES

CARMICHAEL, D. 1818. Some account of the island of Tristan da

Cunha and its natural productions. Transactions of the Einnean

Society of London 12; 483-513.

TARDIEU-BLOT, M.L. 1958. Polypodiacees (sensu lato). Dennstaedti-

acees-Aspidiacees. In H. Humbert, Flore de Madagascr et des

Comores. Earn. 5,1; 1-391. Paris.

THOUARS, L.M. AUBERT DU PETIT. 1808. Esquisse de laflore de

ITsle de Tristan dAcunga. Paris.

J.P. ROUX*

* Compton Herbarium, National Botanical Institute, Private Bag X7,

7735 Claremont, Cape Town.

MS. received; 2000-09-01.

PORTULACACEAE

TALINUM PANICULATUM, A NATURALIZED WEED IN SOUTH AFRICA

Talinum Adans. is a medium-sized genus of semi-suc-

culent herbs and shrubs with annual branches sprouting

from a perennial base with tuberous roots. These fleshy

underground parts tend to draw the attention of succulent

plant collectors, who would include the plants in the

broadly conceived group of caudiciform succulents

(Smith et al. 1997). About 50 species are recognized in

Talinum (Von Poellnitz 1934), occurring mostly in

Africa, Australia, parts of Asia and North and South

America. In the African species the pedicels are always

swollen below the fruit and are more or less recurved

when fruiting (Tolken 1969). Eive species are indigenous

in South Africa, restricted to summer rainfall areas

(Tolken 1969). Talinum paniculatum (Jacq.) Gaertn. is

196

Bothalia 31,2 (2001)

FIGURE 4. — Talinum paniculatum: A, habit of plant in fruit, x 0.6; B,

open flower as seen from above, x 5; C, flower in side view

with one petal removed, x 5; D, capsular fruit x 3.7; E, dissect-

ed capsule showing multiple seeds with free-central placenta-

tion, X 3.7. Artist: Marietjie Steyn.

one of two alien species of Portulacaceae that has

become naturalized in the flora of southern Africa

(Jordaan 1997). This report forms part of ongoing efforts

to document, describe and illustrate naturalized succu-

lents in southern Africa.

Known as ‘American star-flowers’ or ‘pink star-flow-

ers’, T. paniculatum is native to the plains from Texas to

Arizona, south and central Florida, Mexico, the West

Indies and South America (Gray 1895; Von Poellnitz

1934). Its occurrence in Sri Lanka and in Malaysia

(Singapore), Indonesia (Java), Thailand and China, prob-

ably resulted from early cultivation and a subsequent

escape into the wild (Von Poellnitz 1934). In South

Africa, the species was recorded about thirty years ago as

a garden escape in a few places, but did not then seem to

spread as a weed (Tolken 1969). It has since become

established as a troublesome weed in domestic and pub-

lic gardens in and around Pretoria and along the Western

Cape coast with its Mediterranean-type climate. The

plants are not easy to eradicate; although the aerial parts

tend to die off in winter, the species is perennial through

a thickened rhizome and fleshy roots (Figure 4). It also

flowers and fruits prolifically and produces a multitude

of wind-dispersed microscopic seeds that spread and

germinate easily.

2406000-600 Talinum paniculatum (Jacq.) Gaertth,

De fructibus et seminibus plantarum 2: 219, t. 128 (1791);

Poelln.: 1 (1934); Adams: 267 (1972). Type: not desig-

nated.

Portulaca paniculata Jacq.: 22 (1760) non L.: 640 (1762); Jacq.:

148 (1763); Jacq.: 71, t. 151 (1772-1773).

Portulaca patens L.: 242 (1771); Ruclingia patens (L.) Ehrh.: 135

(1788); Helianthemoides patens (L.) Med.: 95 (1789); Talinum patens

(L.) Willd.: 863 (1799); A. Gray: 265 (1895); Claytonia patens (L.)

Kuntze: 56 (1891). Type: not known.

T. reflexum Cav.: 1, t. 1 (1791); Sims: t.1543 (1813); Portulaca

reflexa (Cav.) Haw.: 141 (1803); Claytonia reflexa (Cav.) Kuntze: 57

(1891). Type: not known.

T. sarmentosum Engelm.: 153 (1850); Claytonia sarmentosa

(Engelm.) Kuntze: 57 (1891); T. reflexum var. sarmentosum (Engelm.)

Small: 415 (1903); T. paniculata var. sarmentosum (Engelm.) Poelln.:

123 (1933). Type: not known.

For a more complete list of synonyms, see Von

Poellnitz (1934: 11).

Glabrous herb with annual branches developing from

perennial base with tuberous roots. Stems erect, terete, up

to 0.4 m long. Leaves alternate to subopposite, petiolate,

glabrous, somewhat succulent; blade obovate or oblance-

olate, varying to spatulate, 40-110 x 15-45 mm, apex

acute to rounded, margins entire; petiole swollen at base;

exstipulate. Inflorescence terminal, panicle-like, com-

pound, dichasial, up to 450 mm long; bracteoles minute,

± 1 mm long, membranous, lanceolate, early deciduous.

Flowers bisexual, star-shaped, ± 2.5 mm diam., opening

mid-afternoon, closing at dusk; pedicels filiform, ±3.5

mm long. Sepals 2, opposite, green, glabrous, 1 mm

long, acuminate, recurved in open flower. Petals 5,

oblong with rounded apices, spreading, pink to rose-

coloured, tardily deciduous from capsule. Stamens

numerous, ± 1.5 mm long; filaments dorsifixed; anthers

opening with slits. Ovary superior, spherical, 1 mm

diam., unilocular, multi-ovular, placentation free-central;

style terete, articulating at base; stigma with 3 spatulate

lobes. Capsule spherical, 2.5 mm diam., thin-walled,

bright orange when young, turning to brown during

ripening, opening with 3 longitudinal slits. Seeds numer-

ous, lens-shaped, 0.8-0. 9 mm long, short-beaked with

small, white aril at hilum, seed coat shiny black, tuber-

culate. Figure 4. Chromosome number unknown.

Flowering time'. September to April.

Vouchers: Barker 4439 (PRE); Barrett 333 (PRE); Pegler 1587

(PRE); G.K Smith & E.M.A. Steyn 4 (PRE).

Illustrations: Jacq.: t. 151 (1772-1773) as P. paniculata; Cav.: t.I

(1791), as T. reflexum; Sims: t. 1543 (1813), as 71 reflexum.

Common names: American star-flower (Von Poellnitz

1934: 1) or pink star-flower (Gray 1895).

Bothalia 31,2 (2001 )

197

REFERENCES

ADAMS. C.D. 1972. Flowering plants of Jamaica: 267. University

Press. Glasgow.

CAVANILLES. A.J. 1791. Taliniim reflexion. leones el descriptiones

plantariim 1: 1, t. 1. (Reprint 1965). Cramer, New York.

EHRHART. J.F. 1788. Beitrdge zur Naturkunde 2: 135.

ENGELMANN. G. 1850. Portulacaceae. Boston Journal of Natural

History 6: 153.

GAERTNER, J. 1791. Talinwn Adans. De fructibus et seminus plan-

tarum 2: 219. t. 128. fig. 13. Guilielmi Henrici Schrammii,

Tubinge.

GRAY, A. 1895. Talinwn patens Willd. Synoptical flora of North

America 1. Part 1, Fascicle 1: 265. American Book Co., New

York.

HAWORTH. A. H. 1803. Portulacca (sic.). Miscellanea naturalia: 141.

Taylor, London.

JACQUIN. N.J. 1760. Enumeratio systematica plantarum: 22.

Lugduni, Batavorum.

JACQUIN, N.J. 1763. Selectarum stirpium americanarum historia:

148. (1971 Facsimile). Hafner, New York.

JACQUIN, N.J. 1772-1773. Hortus botanicus vindobonensis 2: 71, t.

151. Kaliwoda, Vindobonae.

JORDAAN. M. 1997. Portulacaceae. In G.F. Smith, E.J. van Jaarsveld,

T.H. Arnold, F.E. Steffens, R.D. Dixon & J.A. Retief, List of

southern African succulent plants: 145. Umdaus Press. Pretoria.

KUNTZE. O. 1891. Portulacaceae. Revisio generum plantarum 1: 56,

57. Kommisionen. Leipzig.

LINNAEUS, C. 1762. Species plantarum, Vol. 1, edn 2: 640.

Laurentius Salvius, Stockholm.

LINNAEUS, C. 1771. Mantissa plantarum: 242. (1961 Facsimile).

Cramer. Weinheim.

MEDIKUS, F.K. 1789. Philosophische Botanik 1: 95. Neue Hof- und

Akademische Buchhandlungen, Mannheim.

SIMS, J.K. 1813. Talinum reflexion. Reflex-flowered Talinum. Curtis’s

Botanical Magazine 37: t. 1543.

SMALL, J.K. 1903. Talinum Adans. Flora of the southeastern United

States: 415. Small. New York.

SMITH, G.F., VAN JAARSVELD, E.J., ARNOLD, T.H., STEFFENS,

F.E., DIXON, R.D. & RETIEF, JA. (eds). 1997. List of south-

ern African succulent plants: ii. Umdaus Press, Pretoria.

TOLKEN. H.R. 1969. The genus Talinum (Portulacaceae) in southern

Africa. Bothalia 10: 19-28.

VON POELLNITZ, K. 1933. Zur Kenntnis der Gattung Talinum

Adans. (Portulacaceae). Berichte der Deutschen Botanischen

Gesellschaft 51: 123.

VON POELLNITZ, K. 1934. Monographic der Gattung Talinum

Adans. Repertorium specierum novarum regni vegetabilis 35:

1-34.

WILLDENOW, C.L. 1799. Talinum patens. Species plantarum 2, Part

2: 863. Nauk, Berlin.

E.M.A. STEYN* and G.FSMITH*

* National Botanical Institute, Private Bag XlOl, 0001 Pretoria.

MS. received: 2001-04-06.

ZAMIACEAE

ENCEPHALARTOS RELICTUS: A NEW SPECIES FROM SOUTHERN AFRICA

In a continuing attempt to document biodiversity in the

African Zamiaceae, an evaluation of material from

Swaziland (Hurter 1993) has led to the conclusion that

there exists a distinct undescribed species, that may already

be extinct in the wild. Due to the fact that no new material

has been forthcoming in more than 20 years it was decided

that this relict species should be described for posterity.

Encephalartos relictus P.J.H. Hurter, sp. nov., E.

heenanii R. A. Dyer foliis rigidis caesiis similis, sed pinnis

linearo-lanceolatis adscendentis inflexis, microsporo-

phyllis ovatis tomentosis, et habitatione nemorali differt.

Strobili glabrous, scale facets smooth, light greenish

yellow. Megastrobili unknown. Microstrobili 1-3 per

trunk, subconical, 200-240 x 120-150 mm, stalked on

peduncle 30-50 mm long. Median microsporophylls

spreading, more or less at right angles to axis, lamina

oblong, tapering to base, ± 35^0 mm long, 30-35 mm

wide and 10-15 mm high, margins contracted to pedi-

cel, bulla with terminal facet projecting slightly as

drooping lip-like structure, edges verrucose, microspo-

rangia separated from lateral margins. Figure 5.

Diagnostic features and affinities

TYPE. — Swaziland: Siteki, Farm Muti-Muti, (leaf

and part of male cone), 15-03-1971, J.J.P. du Preez s.n.

(PRE33123, holo.).

Plant arborescent, suckering from base. Trunk up to

2.5 m long, 400^50 mm diam., leaf bases persistent,

crown and cataphylls tomentose, golden brown, becom-

ing subglabrous with age. Leaves numerous in dense,

spreading crown, rigid, subsessile, waxy blue-grey in

colour, 1.0-1.2(-1.4) m long, pinnae ascending. Petiole

apparent, woolly, becoming subglabrous with age,

except pulvinus. Rachis straight, woolly, becoming

subglabrous with age, apex slightly incurved. Pinnae

woolly, becoming glabrous with age, entire, veins

raised abaxially, margins slightly thickened, inflexed,

directed towards apex of leaf at an angle of ± 60° to

rachis, opposing leaflets inflexed, set at an angle of ±

40° to each other and orientated succubously, proximal

pinnae gradually reduced to a few prickles. Median

leaflets oblong-lanceolate, pungent, 200-250 x 14-17

mm, margins entire, 20-25 prominent veins abaxially.

E. relictus superficially resembles E. heenanii

R. A. Dyer (Dyer 1972), on account of its stiff waxy blue-

grey leaves and pinnae with the veins prominently

raised abaxially. However, it differs markedly from E.

heenanii in morphology, habit and habitat. E. relictus

used to occur in mixed deciduous woodland (Figure 6),

whereas E. heenanii occurs in high rainfall, high alti-

tude, sour grassland. The important morphological dif-

ferences between the two species are summarized in the

following table.

TABLE 1 . — Differences between E. heenanii and E. relictus

198

Bothalia 31,2 (2001)

FIGURE 5. — Encephalartos relictus

RJ.H. Hurter. A-C, micro-

sporophyll: A, abaxial view;

B, frontal view; C, side view.

D, median pinna, adaxial

view; E, F, median section of

leaf showing inflexed nature

of pinnae; G, median pinna,

abaxial view and orientation

towards apex of leaf; H, peti-

ole and proximal pinnae.

Scale bars: A-H, 30 mm.

Artist: S.J. Burrows.

FIGURE 6. — Encephalartos relictus

in habitat (photo: J.J.P. du

Preez).

Bothalia 31,2 (2001 )

199

FIGURE 7. — Encephalartos relictus microstrobili (photo: J.J.R du

Preez).

Geographical distribution

As far as is known, this species used to grow only at

a single locality in Swaziland, at an altitude of 1 000 m

(Figure 8). Its present conservation status code (lUCN

1994) is ExW.

Other specimen examined

SWAZILAND. — Siteki, Farm Muti-Muti, P.J.H. Hurter 95s/hl

(GLOW).

FIGURE 8. — Known former distribution of Encephalartos relictus.

The species no longer occurs in the wild (Red List category

ExW).

ACKNOWLEDGEMENTS

The research of which this publication is a result was

financed by the National Botanical Institute. The artist is

Sandie Burrows. The director of the National Botanical

Institute (PRE) is thanked for permission to use the

accompanying photographs.

REFERENCES

DYER, R.A. 1972. A new species of Encephalartos from Swaziland.

Bothalia 10: 539-546.

HURTER, P.J.H. 1993. Focus on: Encephalartos heenanii.

Encephalartos No. 40: 4—7.

lUCN 1994. lUCN Red List Categories. lUCN, Gland, Switzerland.

P.J.H. HURTER* and H.F. GLEN**

* Lowveld National Botanical Garden, P.O. Box 1024, 1200

Nelspruit, South Africa.

** National Botanical Institute, Private Bag XlOl, 0001 Pretoria.

MS. received: 2000-03-08.

NEW RECORDS AND DISTRIBUTIONAL DISJUNCTIONS FROM SOUTH AFRICA,

ZIMBABWE AND MOZAMBIQUE

INTRODUCTION

Floristic evidence for the continuity of vegetation

types often lingers in the form of relicts which inhabit

isolated refugia. This evidence can be ambiguous in fam-

ilies with adaptations to long distance dispersal.

However, in a number of families, vicariance is the only

logical explanation for distributional anomalies. This

paper records disjunctions in Acanthaceae, Thymelaea-

ceae and Lamiaceae, all families renowned for the

parochial dispersal of seeds and fruits.

ACANTHACEAE

Recent research (Edwards & Harrison 1998) revealed

a distributional extension to the range of Pseuderan-

themum hildebrandtii Lindau (Acanthaceae) which reit-

erates the floristic links between tropical East Africa and

northern KwaZulu-Natal. This species is associated with

woodlands and provides compelling evidence of the con-

tinuity of dense savanna and forest habitat between the

eastern seaboard of South Africa and populations in

Tanzania. Acanthaceae have explosive fruits with elastic

200

Bothalia 31,2 (2001)

FIGURE 9. — Recorded distribution of Metaningia pubinervia.

funicles (retinaculae) which are effective in local disper-

sal but are not suitable for long distance dispersal. It was

therefore exciting to find another species of forest

Acanthaceae, Metarungia pubinen’ia, which corrobo-

rates the above pattern.

According to Baden (198 1 ) the genus Metarungia com-

prises three species, two of which, M. galpinii (Baden)

Baden and M. longistrohus (C.B. Clarke) Baden, are en-

demic to the subcontinent and the type species, M.

pubinervia (T.Anderson) Baden, is common in tropical

East Africa, but has been recorded in southern Mozam-

bique. The phytogeography of Metarungia is intriguing

in its inconsistency. M. longistrobus occurs in asso-

ciation with forest patches in Mpumalanga, southwest

Mozambique and Swaziland and, although the species

occasionally ventures into stony grasslands with steep

slopes, it is unable to survive fire. Similarly, M. galpinii

is limited to forest habitats around East London, Eastern

Cape, but in cultivation both species are more vigorous

in exposed situations. There is no doubt that M. galpinii

and M. longistrobus are sister species, the plants have

very similar habit, creamy mustard corollas of similar

size, similar androecia, gynoecia, inflorescence struc-

ture, fruits and seeds. The major differences which sep-

arate them relate to indumental differences, bract size,

the loss of bracteoles in M. galpinii, and the much small-

er calyces of M. longistrobus. If one accepts that these

are vicarious sister species, then their current parochial

distributions (Baden 1981) are remnants of the range of

their hypothetical parent. This distributional disjunction

provides compelling evidence of an ancient forest link

between the afromontane and riverine forests of

Mpumalanga, in the north, and the coastal forests

around East London. Neither species has yet been

recorded from the intervening forests of KwaZulu-

Natal. Extending this line of reasoning to M. pubinervia

(Figure 9), allows one to postulate past forest linkage

between the southern (Krantzkloof) and northern

(Tanzanian) populations.

Metarungia pubinervia (T.Anderson) Baden in

Kew Bulletin 39: 638 (1984). Macrorungia pubinervia

(T.Anderson) C.B.Clarke: 255 (1900); Brenan & Green-

way: 13 (1949); Dale & Greenway: 17 (1961); Agnew: 607

(1974); Baden: 148 (1981). Rungia pubinervia T.Ainderson:

46 (1863). Himantochilus pubinerx’ia (T.Anderson) Lindau:

373 (1895a). Anisotes pubinervis (T.Anderson) Heine: 189

(1966). Type: Malawi, Mt Chiradzulu, Kirk s.n. (K!, lecto.).

Himantochilus marginatus Lindau: 60 (1894); Lindau: 346 ( 1895b).

Type: Tanzania, Usambara. Holst 9063 (HBG, lecto.; G, P, M, Z, W).

Shrubs 1-5 m tall; stems initially hairy, often with

aerial roots. Leaves elliptic to narrowly obovate,

140-250 X 40-100 mm, sparsely strigose to glabrescent,

apex acute to acuminate, veins sericeous when young,

becoming glabrescent, acarodomatia present in vein

axes; petiole 15-60 mm long. Inflorescences axillary,

usually on leafless stems, 15-50 mm long; peduncles sel-

dom exceeding 2 mm long, glabrescent; bracts tightly

imbricate; lower bracts sterile, ovate to elliptic, 5-12 x

3-5 mm, apex slightly acuminate, margin slightly scari-

ous; fertile bracts broadly ovate, 10-15 x 6-9 mm, apex

acuminate, margin markedly scarious, pink to red;

bracteoles absent. Calyx 9-12 mm long, membranous;

lobes pink, 7-9 x 2-3 mm, midveins green, extending

into an apiculus. Corolla cylindrical, red; tube 9-1 1 mm;

upper lip 28-31 x 8 mm, cucullate; lower lip coiled at

anthesis, 20 x 3 mm. Stamens 2; filaments 20 mm long,

cream-coloured, epipetalous, attached in throat; anther

thecae oblique, 2-3 mm long. Ovary 2. 5-3.0 x 1.5 mm,

glabrescent; style 27-30 mm long, glabrous; nectarifer-

ous cupule 1-2 mm tall.

In the Krantzkloof population bracteoles were not

observed. However, Baden (1981) records the occasion-

al occurrence of linear bracteoles in tropical populations

of M. pubinervia. In a generic context, the fugitive

occurrence of bracteoles in this species is interesting

because this character is used to discriminate between M.

longistrobus and M. galpinii.

Throughout its range M. pubinerx’ia occurs between

500 and 2 000 m in riverine and evergreen forest, in light

shade. The Krantzkloof population is limited to scree

along the base of sandstone cliffs. This unstable substrate

reduces the establishment of the tree canopy and maintains

the higher light intensities in which the species thrives.

This population is fairly large (several hundred adult indi-

viduals and an abundance of seedlings) and the plants,

although conspicuous, are in a fairly remote section of the

gorge and have consequently escaped detection.

Specimen examined

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Krantzkloof

Nature Reserve, (-DD), Styles, Johnson & Edwards s.n. (K, NH, NU,

PRE).

Bothalia31,2 (2001)

201

THYMELAEACEAE

The poor capacity for long distance dispersal is com-

mon to a number of families, including Thymelaeaceae.

Most South African species of Gnidia are grassland

inhabitants and, due to the continuity of this habitat, rela-

tively few species display disjunct distributions. Gnidia

denudata is anomalous in this regard. The species has a

tree-like habit and occurs in forest margins, lacking the

resprouting caudex which enables grassland taxa to sur-

vive fire. The remaining Gnidia species are shorter,

multi-stemmed shrubs or herbs inhabiting fynbos, grass-

land or subalpine zones. Lindley (1823) described G.

denudata, the specific epithet referring to the lack of

lamina hairs, which usually distinguishes this species

from G. imbricata L.

Gnidia denudata LindL, Botanical Register 9: t. 757

(1823); Spreng.: 152 (1827); Meisn.: 441 (1840); Drege:

123 (1843); Meisn.: 585 (1857); C.H.Wright: 63 (1915). G.

tomentosa sensu Hook.: t. 2761 (1827) non Thunb. Type:

South Africa, in nemorosis prope George, IV. C. b.,

Drege (NY, iso!).

Shrub to small tree up to 3 m tall (Wright 1915).

Branches slender, erect to laxly arching, and pilose when

young, glabrescent with age. Leaves simple, exstipulate,

3-5-veined from base, clustered towards ends of branches,

pilose when young. Inflorescences terminal or lateral clus-

ters comprising 4—6 mbular, pale flowers; involucral bracts

green, foliaceous, smaller than leaves, as long as hypanthia.

Flowers 4-merous, sweetly scented at night; petaloid scales

paired, narrowly oblong, alternating with calyx lobes.

Stamens 8, in 2 rows in mouth and throat of tube.

The pilose young stems, leaves and flowers, together

with the comparatively rare condition of broadly foli-

aceous leaves that are three to five-veined from the base,

distinguish this species.

Some confusion has arisen regarding the nomenclature

of G. denudata and G. tomentosa. Linnaeus (1753)

described G. tomentosa from the mountains of the Cape

Peninsula and Worcester areas. This shrub attains about 1

m in height and bears slightly scabrid, semi-coriaceous

leaves which have verrucose surfaces: a result of raised

epidermal cells around the bases of hairs. Flowers, ± 6, are

produced only in terminal inflorescences. Hooker (1827)

misapplied G. tomentosa to plants of G. denudata. Leaves

of G. denudata lack the coarse texture and verrucose sur-

faces typical to G. tomentosa. In addition, the narrowly

oblong petaloid scales of G. denudata distinguish it from

G. tomentosa which has broad fleshy scales.

The distribution of this species shows a clear disjunc-

tion (Figure 10). It occurs south of Oudtshoom and in the

Knysna District of Western Cape and in the Willowmore

District of Eastern Cape, where it inhabits margins of

afromontane forest (Lubke & McKenzie 1996).

Additional, less precise, distribution records (omitted

from Figure 10) come from Western and Eastern Cape,

the Swellendam Mountains; near Touws River, and the

forest at Elands River, Tsitsikamma, Humansdorp

(Wright 1915). The population from Inhaca Island,

FIGURE 10. — Recorded distribution of Gnidia denudata, •, (O dis-

puted record); Plectranthus dolomiticus, ▲.

Mozambique, is highly disjunct. A specimen housed in

the Geneva Herbarium and apparently collected by

Drege from Port Natal (Durban) is probably erroneously

labelled, as this species does not appear in any of his list-

ing of collections from the province (Drege 1843). The

specimen Coetzee s.n., collected in KwaZulu-Natal (and

without any further locality details) in 1943, appears to

be the only other collection from this province, which is

curious in the light of the common occurrence of Cape

relicts on the Natal Group Sandstone of the eastern

seaboard (Van Wyk 1990). The species also appears to

be absent from the mainland of Mozambique. Wright

(1915) regarded the specimen, Rogers 20682 (STE), col-

lected near Grahamstown, as a doubtful record, but in the

light of the Inhaca population, this may warrant further

investigation.

Specimens examined

WESTERN CAPE. — 3322 (Oudtshoom): Saasveld Forestry

College, (-DC), Beaumont & Smith s.n. (NU); Karatura River, (-DD),

Schlecliter 5887 (C). 3323 (Willowmore): Diepwalle Forest Reserve,

(-CC), Bos 735 (STE). 3423 (Knysna): between Kruisfontein and

Bracken Hill, (-AA), Dahlgren & Peterson 1484 (GB),

EASTERN CAPE.— 3323 (Willowmore): Camel Pile, (-DD),

Esterhuysen 27327 (GB). 3326 (Grahamstown): Grahamstown, (-BC),

Rogers 28682 (GRA).

KWAZULU-NATAL. — Precise locality unknown: Coetzee s.n. (J),

Drege s.n. (G).

MOZAMBIQUE. — 2632 (Bela Vista): Inhaca Island, (-BB),

Maguire 35 (J),

LAMIACEAE

Plectranthus dolomiticus was described in 1984 by

Codd. At that time, and in the subsequent FSA account

(Codd 1985), the species was known only from the type

locality near Penge Mine in Mpumalanga. Recently, a

second population of the species was recorded from

granite domes in Zimbabwe (Figure 10). While the geol-

ogy of the respective sites is different, the physical

aspects are very similar. Both populations occur in arid

woodland on rock outcrops where plants occur in

humus-filled crevices. During the dry season the species

withers aerially and the populations rely on subter-

202

Bothalia 31,2 (2001)

ranean tubers for sustenance. The disjunction, approach-

ing 700 km, is considerable for Plectranthus species

which have no mechanism to aid long-distance disper-

sal. It is therefore likely that additional populations of

the species occur(ed) in the intervening rocky savanna.

Codd’s (1985) interpretation of alliances, between P.

dolomiticus and the remainder of the genus, was that its

closest relative is P. petiolaris. Considerable similarity

exists between the flowers of these two species, both of

which are sigmoid, with similar proportions and deep

purple colouring. Recently a third species, which dis-

plays a very similar corolla, was discovered in

Zimbabwe (Edwards, Paton & Crouch 2000). This last

species, P. porphyranthus T.J. Edwards & N.Crouch, is

without doubt convergent in corolla form for it belongs

to section Coleoides, whereas the former species are

members of section Plectranthus. The convergence is

probably being driven by widespread anthophorid bees,

of the genus Amegilla, which are the primary pollina-

tors. Amegilla caelestina (Cockrell), A. fallax (Smith)

and A. bothai (Meadt-Woldo) were recorded on P por-

phyranthus, and A. caelestina, A. bothai and A. mimad-

vena (Cockrell) are the primary pollinators of P. petio-

laris. The proposed alliance of P. dolomiticus with P.

petiolaris (Codd 1985), based on corolla characters,

may need reviewing in light of the convergent corolla

design of P. porphyranthus.

Specimens examined

Plectranthus dolomiticus

ZIMBABWE.— 1832 (Juliasdale), Pine Lake Inn, (-BA), Vos 339

(NU).

Plectranthus porphyranthus

ZIMBABWE. — 2029 (Harare), Masvingo, (-BB) Richmond Farm,

Crouch 800 (E, K, NH, NU, PRE); Harare, 3 miles [4.8 km] east of

Zimbabwe (ruins), Leach 8043 (PRE).

ACKNOWLEDGEMENTS

The Natal University Research Fund and the National

Research Fund are thanked for their financial assistance.

The following herbaria C, G, GB, GRA, J, K, NU, PRE

and STE are thanked for the loan of herbarium material

and the National Botanical Institute, Pretoria, is thanked

for access to the Mary Gunn Library. Neil Crouch is

acknowledged for his endless enthusiasm in tracking

down obscure references. Christina Potgieter is thanked

for contributing insect data on the pollination of

Plectranthus petiolaris. Rod Edwards is acknowledged

for his contribution in the discovery of the Metarungia

pubinervia population.

REFERENCES

AGNEW, A.D.Q. 1974. Acanthaceae. Upland Kenya wild flowers'.

573-61 1. Oxford University Press.

ANDERSON, T. 1863. African Acanthaceae. Proceedings of the Linnaean

Society, Botany 7: 13-54.

BADEN, C. 1981. The genus Macrorungia (Acanthaceae), a taxonomic

revision. Nordic Journal of Botany 1: 638.

BADEN, C. 1984. Metarungia. a valid name for Macrorungia auctt.

(Acanthaceae). Kew Bulletin 39: 638.

BRENAN, J.P.M. & GREENWAY, P.J. 1949. Checklist of the forest trees

and shrubs of the British Empire, Tanganyika Territory 5,2:

1-319. Imperial Forestry Institute, Oxford.

CLARKE, C.B. 1900. Acanthaceae. Flora of tropical Africa 5: 1-255.

Reeve, London.

CODD, L.E. 1984. A new species of Plectranthus (Lamiaceae). Bothalia

15: 142, 143.

CODD, L.E. 1985. Plectranthus (Lamiaceae). Flora of southern Africa

28,4: 137-172.

DALE, I.R. & GREENWAY, P.J. 1961 . Kenya trees and shrubs. Hatchards,

London.

DREGE, J.F. 1843. Zwei pflanzengeographische Documente. Flora 2:

44-229. Leipzig/Regensburg.

EDWARDS, T.J. & HARRISON, E.R. 1998. New records from KwaZulu-

Natal, South Africa. Bothalia 28: 187-190.

EDWARDS, T.J., PATON, A. & CROUCH, N. 2000. A new species of

Plectranthus (Lamiaceae) from Zimbabwe. Kew Bulletin 55:

459-464.

HEINE, H. 1966. Acanthaceae. Flora du Gabon 13: 3-241. Paris.

HOOKER, J.D. 1827. Gnidia tomentosa. Botanical Magazine 54: t. 2761 .

London.

LINDAU, G. 1894. Acanthaceae africanae II. Botanische Jahrbucher

20: 1-78.

LINDAU, G. 1 895a. Acanthaceae. Die Pflanzenwelt Ost-Afrikas und der

Nachbargebiete Deutsch Ost-Afrika 5,C: 365-374.

LINDAU, G. 1895b. Acanthaceae. Die natiirlichen Pflanzenfamilien

4,3b: 274-354. Leipzig.

LINDLEY, J. 1823. G?tidia denudata. Botanical Register 9: t. 757. London.

LINNAEUS, C. 1753. Gnidia tomentosa. Species plantarum, edn 1.

Salvius, Stockholm.

LUBKE, R. & MCKENZIE, B. 1996. In A.B. LOW & A.G. REBELO,

Vegetation of South Africa, Lesotho and Swaziland. Department

of Environmental Affairs and Tourism, Pretoria.

MEISNER, C.F. 1840. Synopsis Thymelaearum, Polygonearum et

Begoniarum Africae Australis, imprimus a cl. J.J. Drege lec-

tarum. Linnaea 14: 385-516.

MEISNER, C.F. 1857. Thymelaeaceae. In A. P.D. de Candolle, Prodro-

mus systematis naturalis regni vegetabilis 14: 493-605. Treuttel

& Wiirtz, Paris, Strasburg, London.

SPRENGEL, K. 1827. In C. Linnaei, Systema vegetabUium, Curare

posteriores 152. Gottingen.

VAN WYK, A.E. 1990. The sandstone regions of Natal and Pondoland:

remarkable centres of endemism. Palaeoecology of Africa and

the surrounding islands 21 : 243-257. Balkema, Rotterdam.

WRIGHT, C.H. 1915. Thymelaeaceae. In W.T. Thiselton-Dyer, Flora

capensis 5,2: 63, 64. Reeve, Ashford, Kent.

T.J. EDWARDS*!, A.J. BEAUMONT* and D. STYLES*

* School of Botany and Zoology, University of Natal, Private Bag XOl ,

3209, Scottsville.

t Author for correspondence.

MS. received: 2000-10-26.

Bothalia31.2 (2001)

203

COLCHICACEAE

A REMARKABLE NEW SPECIES OF ANDROCYMBIUM FROM NORTHERN CAPE. SOUTH AFRICA

INTRODUCTION

A member of the Old World lilialean family Colchi-

caceae (Nordenstam 1998), Androcymbium Willd. is a

largely African genus comprising some 40 species, main-

ly of arid and semi-arid areas. It has its greatest diversi-

ty in southern Africa where the majority of species, at

least three quarters of the total, occur (Muller-Doblies

1984; Meyer 2000). Just three of these extend into tropi-

cal Africa, one as far north as Ethiopia (Baker 1897;

Sebsebe Demissew 1997). Two more species are endem-

ic to the Canary Islands and a further six are distributed

along the arid fringe of the south Mediterranean littoral

(Pedrola-Monfort & Caujape-Castells 1998). The genus

is distinguished from the other truly cormous members

of the family by its condensed or rarely elongated stem,

and stellate-campanulate flowers with free tepals borne

in the axils of large, leaf-like bracts.

The genus Androcymbium was divided by Krause

(1921) into three sections based on the shape of the

tepals. The largest of these sections, sect. Androcymbium

(= Cymbanthes) is characterized by auriculate, deeply

cucullate tepal limbs and is almost entirely southern

African with a few species extending into tropical Africa,

Ethiopia and Eritrea. The remaining sub-Saharan

species, along with the more northern members of the

genus, have flat or shallowly concave tepal limbs with-

out distinct basal auricles. Relationships between these

species are difficult to infer. Krause (1921) recognized

the somewhat heterogeneous nature of the group in erect-

ing the monotypic sect. Dregeocymbium to accommo-

date the rather aberrant southern African A. dregei Presl.

This species is distinguished from the other taxa with

plane tepals by its well-developed aerial stem and

extremely short stylules. The other species with flat

tepals known to him, which included the Mediterranean

taxa plus two of the sub-Saharan species, A. bellum

Schltr. & Krause and A. roseurn Engl., were accommo-

dated in sect. Eryllirostictus.

The distinction between sections Dregeocymbium and

Erythrostictus was blurred by the subsequent discovery

of two species from southern Namibia and northern

Namaqualand, A. exiguum Roessler and A. cruciatum U.

& D.Miill-Doblies, which are apparently intermediate

between them (Roessler 1974; MUller-Doblies & Muller-

Doblies 1984). The two sections were consequently

combined by Miiller-Doblies & Muller-Doblies (1984).

Section Erythrostictus thus currently comprises four

species from southern Africa and about six species from

the Canary Islands and the Mediterranean. Among the

southern African species of sect. Erythrostictus, three are

restricted to the winter rainfall region of Western and

Northern Cape and southern Namibia, and only one, A.

roseurn, was previously known to occur in the summer

rainfall zone of southern Africa. This unusual species

occurs in northern Namibia, Angola and Botswana. It is

readily distinguished from all other southern African

species by its rosulate, multifoliate habit and prominent-

ly exposed pinkish flowers that are not concealed by the

leaf-like bracts. It was a great surprise, therefore, to

encounter a similar species in the Great Karoo near De

Aar. Closer examination revealed that although allied to

A. roseurn, it is quite distinct from that species.

Androcymbium asteroides 7. & Gold-

blatt, sp. nov., haec species habitu caulescente, foliis

rosulatis multis atque tepalomm limbis oblongis exauricu-

latis Androcymbio roseo Engl, similis, sed ab eo foliis

lanceolatis (3-) 10- 18 mm lads atque floribus omnino

albis supra apicem plantae vix vel baud protuberantibus

tepalorum breviorum latiorumque ungue ±7x3 mm ac

limbo ovato 8-9 x 4-6 mm differt.

TYPE. — Northern Cape, 2922 (Prieska): Prieska com-

monage, in vlei ground, 06-05-1928, (-DA), Bryant sub

Marloth 13570 (PRE, holo.).

Stemless perennial with indistinguishable intemodes

and leaves in basal rosette. Cotw ovoid, 10-15 mm

diam.; tunics dark brown, coriaceous. Cataphyll single,

70-110 mm long, translucent, papery in texture. Leaves

± 8, spreading, lanceolate, (60-)9()-l 10 x (3-) 10-1 8 mm,

but expanding up to 25 mm wide at base, attenuate,

amplexicaul, upper leaves widening more conspicuously

towards base than lower, spirally inserted, glaucous

green, lightly canaliculate with depressed median vein,

margins ciliolate, grading into floral leaves; fertile floral

leaves shorter and broader at base than foliage leaves,

partially encircling flowers but not obscuring them,

suberect or incurved. Elowers 2-6, sessile, not or slight-

ly protruding above crown of plant; perianth ± 10 mm

diam., white, unscented; tepals unequal, inner slightly

smaller than outer, (1 3-) 15- 17 mm long, limb flat or

lightly concave, ovate, 8-9 x 4—6 mm, claw oblong-

cuneate, 6-8 x 2-3 mm. Stamens included; filaments

arcuate, 4-5 mm long, greenish yellow; anthers 2 mm

long, yellow; nectary wider than filament base, ovate

with central keel, 1.2 mm diam., greenish yellow. Ovary

obovoid, 3-lobed, 5-6 mm long, white flushed green

above, stylules 6 mm long, white, stigmas elongate, 1

mm long. Capsule subglobose, coriaceous, 17 mm long.

Seeds subglobose, 2.0-2. 5 mm diam., dark brown, testa

rugulose. Elowering time: May to June. Figure 11.

Distribution and biology: the species is largely restrict-

ed to the southern edge of the interior plateau of southern

Africa at altitudes of ± 1 400 m (Figure 12), and is not

uncommon southeast of De Aar around Hanover and

Richmond in Northern Cape. It occurs in seasonally

waterlogged depressions in clay soils, usually derived

from dolerite. These depressions, due to the particular

properties of the doleritic clays, are locally moist for much

longer than the surrounding flats and constitute a special-

ized, highly localized habitat that is occupied by several

dwarf geophytes apart from Androcymbium asteroides.

Among these are Massonia comata, Polyxena ensifolia

(Hyacinthaceae) and Moraea falcifolia (Iridaceae). These

204

Bothalia 31,2 (2001)

FIGURE 1 1 . — Androcymbium aster-

oides. A, whole plant with

corm; B, individual flower

and subtending leaf; C, outer

tepal and stamen; D, gynoe-

cium. Scale bars: 10 mm.

Artist: John Manning.

plants share a suite of morphological characters, including

the stemless habit with spreading or prostrate leaves, sub-

terranean ovaries and pale-coloured flowers apparently

adapted to a generalist pollination strategy. Their growth

cycle is closely linked to the local climate and they sprout

rapidly in response to the late summer and autumn rains

that characterize the climate of the area. They flower soon

after sprouting, in late autumn or early winter, before tem-

peratures drop too low for active growth.

History, this singular species seems to have been first

collected by E.G. Bryant in 1925. Bryant, a mining engi-

neer with a great interest in fodder plants and conserva-

tion, moved to Prieska in 1920, where he collected exten-

sively. This first collection was sent to the Bolus

Herbarium, where it remained among the incertae.

FIGURE 12. — Distribution of Androcymbium asteroides in Northern

Cape.

Bryant was clearly intrigued by the species as he made

several collections of it over the years. The second of

these, which serves as the type, he sent to Rudolf

Marloth, who identified it as A. roseum. This determina-

tion was then applied to the few subsequent collections

that have been made.

Diagnosis and relationships'. Androcymbium aster-

oides is clearly allied to A. roseum, with which it shares a

multifoliate, rosette habit and more or less flat or shallow-

ly concave tepal limbs lacking prominent basal auricles. It

is distinguished from A. roseum by the broader, lanceolate

leaves with the blades usually at least 10 mm wide towards

the base and smaller, pure white flowers that do not pro-

trude much above the crown of the plant. The tepal claws

are oblong, 6-8 mm long and the limbs ovate, 8-9 x 4-6

mm. A roseum is characterized by linear-lanceolate leaves

up to 5 mm wide and pinkish or pink-streaked flowers that

protrude conspicuously above the crown of the plant. The

tepal claws are narrowly oblong, 10-12 mm long and the

narrowly oblong-elliptic limbs are 13-16 x 3^ mm. A

fine illustration of A. roseum accompanies the account of

the species by Dyer (1956).

These two species show a marked similaiity to the

Mediterranean species of Androcymbium in their multifoli-

ate, rosette habit, monomorphic leaves and flat tepals. It is

likely, however, that most if not all of these are plesiomor-

phic states and the exact relationship between the

Mediterranean and Macronesian species and the other

African members of sect. Erythrostictus has yet to be

demonstrated. A resolution of the broad patterns of rela-

tionships between the species of Androcymbium, especial-

ly those of section Erythrostictus, is an essential prerequi-

site to understanding the historical biogeography of the

genus.

Bothalia31,2 (2001)

205

Additional material examined

NORTHERN CAPE. — 2922 (Prieska): Prieska, 05-1925, (-DA),

Bryant s.n. (BOL); Prieska, 10-1928, Bryant s.n. (PRE); Prieska, 07-35,

Bryant 1128 (PRE). 3023 (Britstown): De Aar, Quaggafontein, sandy

karoo in open patches, 06-05-1946, (-DB), Acocks 12604 (PRE). 3122

(Loxton): Meltonwold. northwest of Graskop, dry vlei, 13-05-1976,

(-DB), Thompson 3065 (PRE). 3123 (Victoria West): Richmond, 3 km

SE of town along road to Graaff-Reinet, seasonally waterlogged

drainage line in dolerite, 28-04-2001, (-BD), Manning 2322 (NBG); 27-

05-2001, Snijrnan 1815 (NBG).

ACKNOWLEDGEMENTS

Our thanks to C. Archer (PRE) and T. Trinder-Smith

(BOL) for expediting the loan of material from their

respective herbaria, B. Momberg, and the referees for

their efficiency, Roy Gereau for his incomparable Latin

diagnosis, and Dee Snijrnan for kindly collecting further

material. Material for study was collected on a permit

provided by Northern Cape Department of Nature and

Environmental Conservation.

REEERENCES

BAKER, J.G. 1897. Liliaceae. Flora of tropical Africa 7: 559, 560.

DYER, R.A. 1956. Androcymbium roseutn. The Flowering Plants of

Africa 31; t. 1225.

KRAUSE, K. 1921. Revision der Gattung Androcymbium Willd.

Notizblatt des Botanischen Gartens and Museums zu Berlin-

Dahlem 70: 512-526.

MEYER, N. L. 2000. Colchicaceae. In O. A. Leistner, Seed plants of south-

ern Africa: families and genera. Strelitzia 10.

MULLER-DOBLIES, U. & MULLER-DOBLIES, D. 1984. Zur Kennt-

nis der Gattung Androcymbium (Colchicaceae) im siidlichen

Afrika: Zwei Synonyma und fiinf neue Arten. Willdenowia 14:

179-197.

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

and genera of vascular plants III Lilianae (except Orchi-

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PEDROLA-MONFORT, J. & CAUJAPE-CASTELLS, J. 1998. The

Androcymbium species of the Canary Islands. Curtis’s Botan-

ical Magazine 15: 61-11 .

ROESSLER, H. 1974. Die Gattung Androcymbium Willd. (Liliaceae)

in Sudwestafrika. Mitteilungen der Botanischen Staatssamm-

lung, MUnchen 11: 545-566.

SEBSEBE DEMISSEW, S. 1997. Colchicaceae. Flora of Ethiopia and

Eritrea 6: 184-189.

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

* Compton Herbarium, National Botanical Institute, Private Bag XI,

Claremont 7735, Cape Town, South Africa.

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

Garden, PO. Box 299, St. Louis, Missouri 63166, USA.

PTERIDOPHYTA

NEW DISTRIBUTION RECORDS OF SOUTHERN AFRICAN PTERIDOPHYTA

Recent travels by the authors have resulted in the col-

lection of a number of new national and provincial

records of pteridophytes from the southern African sub-

region which are not reflected in the most recent revi-

sions and regional checklists. The Malawian records

update Burrows & Burrows (1993); the South African

records update Burrows (1990) and Roux (1995), while

the Zambian records update Komas (1979). All speci-

mens are housed in the Buffelskloof Herbarium,

Lydenburg, South Africa; duplicates, where distributed,

are indicated.

MALAWI

Anogramma leptophylla (L. ) Link, Filicum species

in horto regio botanico berolinensis cultae; 137 (1841);

Schelpe: 99, t. 32 (1970); J.E.Burrows: 114, t. 26 (1990).

Type: South Europe, Toumefort 5337 (P, lecto.).

MALAWI. — 1033: (North), Nyika National Park, S slopes of

Nganda Hill, 2 480 m, (-BD), 31 March 2000, J.E. & S.M. Burrows

6850 (Buffelskloof Herb., MAL, NBG, SRGH, UZL).

This very small, ±10 mm tall annual fern, frequently

mistaken for sporelings of other ferns, was found grow-

ing in the lee of large boulders in montane grassland. It

is fairly widespread in Africa although, due to its

extremely small size, it is frequently overlooked. It has

been collected in South Africa and Zimbabwe to the

south, and northwards in Tanzania and Kenya.

Asplenium erectum Bory ex Willd. in Species

plantarum edn 5: 328 (1810); N.C.Anthony & Schelpe:

190 (1986); J.E.Burrows: 223, t. 51 (1990). Type:

Reunion, Bory s.n. (B-W19906, lecto.).

var. erectum.

MALAWI. — 1033: (North), Nyika National Park, Zovochipolo

Forest, 2 200 m, (-DA), 2 April 2000, J.E. & S.M. Burrows 6893

(Buffelskloof Herb.).

This specimen was found growing near a small stream

in the deep shade of evergreen montane forest. In addi-

tion to being a new record for Malawi, it is significant

that, in Africa, this variety was only previously known

from South Africa. It is not known from Zimbabwe or

southern Malawi and therefore represents an unusually

disjunct record into tropical Africa.

Cyathea mossambicensis Baker in Annals of

Botany 5: 185 (1891); Schelpe: 72 (1970); J.E.Burrows:

86, t. 18 (1990). Type: Mozambique, Namuli, Last s.n.

(K, holo.).

MALAWI. — 1134: (North), Mzuzu, ± 15 km from Mzuzu on the

Nkhata Bay road, (-CA), 14 May 1999, J.E. Burrows 6484

(Buffelskloof Herb.).

Plants were found growing in dense riverine forest

along a small stream, near the summit of the western

206

Bothalia 31,2 (2001 )

escarpment of the Lake Malawi rift above Nkhata Bay,

an area of unusually high rainfall in Malawi (Chapman &

White 1970). This is the first record of this taxon from

Malawi; elsewhere C mossamhicensis occurs in eastern

Zimbabwe, Mozambique and Tanzania.

Ophioglossum convexum J.E. Burrows in Bothalia

19: 167, t. 1 (1989). Type: South Africa, Mpumalanga,

Lydenburg Dist., Coromandel Farm, J.E. Burrows 3683

(PRE, holo.!; BOL!, K!).

MALAWI. — 1033: (North), Nyika National Park, Wowve River,

2 030 m. (-BD). 4 Apr. 2000, J.E. & S.M. Burrows 6909 (Buffelskloof

Herb., MAL. PRE, UZL).

This very small fern (trophophore 10-13 mm long) is

distinguished by its small suborbicular trophophores

which are held ± oppressed to the ground. It was found

growing in montane grassland on alluvial soils near a

river, growing in association with O. lusoafricanum

Welw. ex Prantl. It is currently known from South Africa

and Zimbabwe and this record therefore extends its

known distribution well into tropical Africa.

SOUTH AFRICA

Thelypteris dentata (Eorssk.) E.P.St.Johii van

buchananii Schelpe in Journal of South African Botany

31: 265 (1965), excl. Buchanan 8 [vide Holttum: 142

(1974)]; Schelpe: 198 (1970); J.E.Burrows: 264 (1990).

Type: Mozambique, Manica e Sofala, Garuso, ‘Jaegers-

berg’ (Bandula Mt), Schelpe 5599 (BOL, holo.!).

KWAZULU-NATAL.— 2831 (Nkandla): Ongoye Forest, (-DC), 18

Feb. 1996, J.E. & S.M. Burrows 5944 (Buffelskloof Herb.).

Plants were found growing on a deeply shaded forest

floor in evergreen forest. Previously this variety was

only known from Zimbabwe, Mozambique and Malawi.

Van huchananii is distinguished from the typical variety

by having two or more pairs of veins anastomosing

below the sinus between the ultimate pinna lobes (com-

pared to I'A pairs, or 3 veins, in van dentata).

Thelypteris hispidula (Decne.) C.EReed in Phyto-

logia 17: 283 (1968); J.E.Burrows: 264, t. 64 (1990);

J.E.Burrows & S.M. Burrows: 95 (1993). Christella his-

pidula (Decne.) Holttum: 312 (1976). Type: Indonesia,

Timor, Guichenot s.n. (P, holo.).

Thelypteris quadrangularis (Fee) Schelpe: 196 (1964); Schelpe:

266 (1965); Schelpe: 195 (1970). Type: Guyana, Leprieur 182 (P,

?holo.; NY).

Christella hilsenhergii (C.Presl) Holttum: 142 (1974). Thelypteris

hilsenhergii (C.Presl) Panigrahi: 369 (1975). Type: Mauritius, Sieber

s.n. (PRC, holo.'; K, P).

KWAZULU-NATAL. — 2831 (Nkandla): Ongoye Forest, (-DC), 18

Feb, 1996, J.E. & S.M. Burrows 5932 (Buffelskloof Herb.).

Growing in evergreen forest, in deeply shaded sites

along a stream. This is the first record of T. hispidula

from South Africa; previously this species had been

recorded as far south as Chirinda Forest in southeastern

Zimbabwe.

Polypodium vulgare L., Species plantarum 2: 1085

(1753); Schelpe & N.C. Anthony: 157 (1986); J.E.Bur-

rows: 193, t. 195 (1990). Type: Europe (not found).

MPUMALANGA. — 2530 (Lydenburg): Lydenburg Dist., Moko-

bulaan Plantations, (-BA), 2 100 m, 26 Feb. 2000, J.E. Burrows, A. &

J. Nel 6716 (Buffelskloof Herb., J, NBG, PRE).

Polypodium vulgare is widespread throughout the

temperate zone of the northern hemisphere. Somewhat

anomalously it also occurs on the South Atlantic island

of Kerguelen, and in temperate South Africa, extending

from the mountains of the Western Cape to the high

Drakensberg Mountains of KwaZulu-Natal (Schelpe &

Anthony 1986). This new record extends the distribution

of the species northwards well beyond its current range.

Polystichum wilsonii H. Christ in Botanical Ga-

zette 51: 353 (1911); J.P.Roux: 149, t. 51 (1998). Type:

China, Szechuan Province, Mupin, Wilson 2614 (BM,

holo.).

Polystichum alticola Schelpe & N.C, Anthony in Anthony & Schelpe:

144 (1982); Schelpe & N.C. Anthony: 257 (1986): J.E.Buitows: 317

( 1990). Type: South Africa, Western Cape, Ladismith, Swartberg, Tower-

kop, Esterhuy.sen 26699 (BOL, holo.; C. G, GH, K, M, MO, NBG, P. PRE).

MPUMALANGA. — 2530 (Lydenburg); Lydenburg Dist., Mokobu-

laan Plantations, (-BA), 2 100 m, 26 Feb. 2000, J.E. Burrows, A. & J.

Nel 6717 (Buffelskloof Herb., NBG).

This species occurs from the southern Cape moun-

tains through the mountains of the Drakensberg to the

Wakkerstroom area. Eurther north, P. wilsonii is repre-

sented in Zimbabwe by three collections. It then ‘jumps’

to northern Tanzania, Kenya, Ethiopia and westwards to

Cameroon and Bioko (Roux 1998). It is found also

through much of eastern Asia as far as China. The above

collection serves to close the preceived gap in its distrib-

ution between the Drakensberg Mountains of South

Africa and the Eastern Highlands of Zimbabwe.

ZAMBIA

Mohria marginalis (Savigny) J.PRoivc in South Afri-

can Journal of Botany 56: 401 (1990). Type: Reunion

(‘Bourbon’), Commerson s.n. (P, lecto.).

Mohria hirsuta J.P.Roux; 435 (1984). Type; South Africa. Free

State, Qwa Qwa, Tsheseng, near cave on road to The Sentinel, Roilx

907 (NBG. holo.; K, PRE).

ZAMBIA. — 1 124: North-Western Province, Luakela River, on road

from Mwinilunga to Ikalenge, (-CB), 4 March 1999, J.E. & S.M.

Burrows 6433 (Buffelskloof Herb.. NBG).

Plants were found growing in miombo (Brachystegia-

Julbernardia-Isoherlinia) woodland near a river, on a

rocky outcrop in light shade. This is 1210he first record

of this species from Zambia and the first record of any

species of Mohria from the western half of Zambia

(Kornas 1979). The woodland habitat of this collection is

anomalous for this taxon which is otherwise a montane

species (Roux 1995).

Ophioglossum ammophilum C.D. Adams in

Annals and Magazine of Natural History, ser. 12, 7: 874

(1954). Type: Nigeria, Accra Dist., near Labadi, near

Kapeshi Lagoon, Adams GC4642 (BM, holo.!; K!).

ZAMBIA. — 0932: Northern Province, 52 km from Nakonde on the

Mbala road, (-AB), 14 Feb. 1995, J.E. & S.M. Burrows 5796 (Buffels-

kloof Herb., K).

Bothalia31,2 (2001)

207

Plants were found growing on the outer margins of a

shallow seasonally inundated pan over laterite, in miom-

bo woodland, in association with Ophioglossum costa-

tum R.Br., O. lusoafricanum Welw. ex Prantl, O. thoma-

sii R.T.Clausen and O. vulgatum L. subsp. africanum

Pocock ex J.E. Burrows var. africanum. This is the first

record of this species for the southern African subregion;

previously it was known only from Nigeria, Sudan and

Ethiopia. Ophioglossum ammophilum is very similar to

O. gomezianum Welw. ex A. Braun but is distinguished

from that species in the field by its somewhat orange

tinge to the sporophore and the few pale basal-medial

cells on the tropophore, creating the impression of a

pseudo-midrib.

ACKNOWLEDGEMENTS

The Fern Society of South Africa, particularly Allan

and Jolanda Nel, is acknowledged for their part in the

discovery of the two new Mpumalanga records; the

Parker and Engelbrecht families for access to

Mokobulaan; and J.P. (Koos) Roux for confirming the

identification of Mohria marginalis. The Southern

African Botanical Diversity Network (SABONET) is

thanked for providing the opportunity to visit northern

Malawi.

REFERENCES

ADAMS. C.D. 1954. New species of ferns from the Gold Coast.

Annals and Magazine of Natural History. ser.l2, 7: 874.

ANTHONY, N.C. & SCHELPE, E.A.C.L.E. 1982. New species and

new combinations in African Pteridophyta and Orchidaceae.

Contributions from the Bolus Herbarium No. 10: 143-164.

BAKER, J.G. 1891. A summary of the new ferns which have been dis-

covered or described since 1874. Annals of Botany 5: 185.

BURROWS. J.E. 1989. New taxa, combinations and records of Pteri-

dophyta from southern and central Africa. Bothalia 19;

167-174.

BURROWS. J.E. 1990. Southern African ferns and fern allies. Frand-

sen Publishers, Sandton.

BURROWS, J.E. & BURROWS, S.M. 1993. An annotated checklist of

the pteridophytes of Malawi. Kirkia 14: 78-99.

CHAPMAN, J.D. & WHITE, F. 1970. The evergreen forests of Malawi.

Commonwealth Forestry Institute. University of Oxford.

CHRIST, K.H.H, 1911. Fihces wilsonianae. Botanical Gazette 5 1 : 345-359.

HOLTTUM, R.E. 1974. Thelypteridaceae of Africa and adjacent

islands. Journal of South African Botany 40: 123-168.

HOLTTUM. R.E. 1976. The genus Christella Leveille, sect. Christella.

Studies in the family Thelypteridaceae, XL Kew Bulletin 31:

293-339.

KORNAS, J. 1979. Distribution and ecology of the pteridophytes in

Zambia. Panstwowe Wydawnictwo Naukowe, Krakow.

LINK, J.H.F 1841 . Filicum species in horto regio botanico beroliinen-

sis cultae. Berolini.

LINNAEUS, C. 1753. Species plantarum 2, edn 1. Holmiae.

PANIGRAHI. G. 1975. Notes on certain taxa of Thelypteris (Thelyp-

teridaceae) of Asia, II. Phytologia 31: 369-372.

REED, C.F 1968. Index thelypteridis. Phytologia 17: 249^66.

ROUX, J.P. 1984. Mohria hirsuta, a new fern species from the

Drakensberg. Journal of South African Botany 50: 435^141.

ROUX, J.P. 1990. A new species and combination in Mohria

(Schizaeaceae: Pteridophyta). South African Journal of Botany

56: 399^02.

ROUX. J.P. 1995. Systematic studies in the genus Mohria (Pterido-

phyta: Anemiaceae). VI. Taxonomic review. Bothalia 25: 1-12.

ROUX, J.P. 1998, A taxonomic revision of Polystichum (Pteropsida:

Dryopteridaceae) in Africa and neighbouring islands. Ph.D.

thesis. University of Pretoria, Pretoria.

SCHELPE, E.A.C.L.E. 1964. Pteridophyta collected on an expedition

to northern Mozambique. Journal of South African Botany 30:

177-200.

SCHELPE. E.A.C.L.E. 1965. A review of the southern African species

of Thelypteris. Journal of South African Botany 31: 259-269.

SCHELPE, E.A.C.L.E. 1970. Pteridophyta. In A.W. Exell & E. Launert,

Flora zambesiaca. Crown Agents, London.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. Flora

of southern Africa. Botanical Research Institute, Pretoria.

WILLDENOW, C.L. 1810. In Carol! a Linne, Species plantarum edn 5.

Berolini.

J.E. BURROWS* and S.M. BURROWS*

* Buffelskloof Herbarium, RO. Box 710, 1 120 Lydenburg, South Africa.

MS. received: 2000-09-26.

GENTIANACEAE

A NEW SPECIES OF SEBAEA FROM THE SWARTBERG RANGE, WESTERN CAPE, SOUTH AFRICA

Sebaea amicorum I.M.Oliv. & Beyers, sp. nov., in

genere distincta foliis lanceolatis, floribus 4-meris,

calyce minute carinata, antheris glande apicali minutissi-

ma, stylo sine tumore stigmatico dignoscenda. Figura 13.

TYPE. — Western Cape, 3322 (Oudtshoom): Oudts-

hoom, Groot Swartberg, neck just SE of Witberg on

Angeliers Berg 187, 1 710 m, (-AC), 18 October 2000,

E.G.H. & I.M.Oliver 11716 (NBG, holo.; K).

Delicate herbaceous biennial, or perennial (?), vary-

ing from small, ± 50 mm tall and 2-4-branched to

100-200 mm tall and many branched, main stem 10-150

X ± 2 mm. Branches: few to many, thin, delicate, erect,

long main and secondary branches all glabrous with 2-A

longitudinal ridges, intemodes 4—10 mm long. Leaves

well developed, scattered along stem, opposite, spread-

ing, basal ones often reflexed, sessile, narrowly lanceo-

late, 5-12 X 1-2 mm, acute, flat, glabrous, midrib slight-

ly prominent on abaxial surface only, margins micro-

scopically dentate or entire. Inflorescence 1 -flowered or

a lax dichasium of 1st and 2nd order, sometimes 3rd

order, at ends of all branches; 1st order peduncle 10-30

mm long, 2nd order peduncles 5-25 mm long, 3rd order

peduncles 5-10 mm long; flowers 4-merous, ± 13 mm

long, yellow; bracts subtending flower subspreading ±

3. 5-5.0 X 1.0 mm, leaf-like; pedicel very short, ± 0.5 mm

long. Calyx 4-partite, free, adpressed to corolla, seg-

ments elliptic to ovate, ± 7.0 x 2. 6-3. 5 mm, acute, veins

not visible, slightly keeled abaxially, keel 0.2-0.3 mm

broad, green centrally, becoming hyaline towards mar-

gins, glabrous, margins entire. Corolla 4-lobed; tube ± 4

mm long, shorter than lobes; lobes elliptic to ovate,

8.0-9.0 X 4.5 mm, subacute. Stamens 4, inserted in corol-

208

Bothalia 31.2 (2001)

FIGURE 13. — Sebaea amicorum. A, flowering branch; B, stem with leaves; C, closed flower; D. bracts; E. sepal; F. corolla, opened and viewed

adaxially; G. stamen, back, side and front views; H. gynoecium; I, seed. All drawn from the type collection. Scale bars; B. D. E. G. FI. 2

mm; C, F, 4 mm; 1. 0.5 mm. Artist; I.M. Oliver.

la sinuses; filaments ± 1 .5 mm long; anthers ± 2.4 mm

long, apex slightly recurved with very small sessile

round apical gland, basal glands absent, thecae fully

adnate. Ovary 2-locular, ellipsoid, 2. 5^.0 x 1.2-2. 5

mm, glabrous, with numerous ovules; style 6. 2-8. 5 mm

long, filiform, with no stigmatic swelling; stigma capi-

tate, ± 0.4 X 0.5 mm, 2-lobed. Seeds ± ovoid, ± 0.4 x 0.2

mm, with many frilly small wings. Flowering time:

October to December. Figure 13.

This new species is at first glance very distinct from

all others in the genus mainly due to its leaves which are

narrowly lanceolate and evenly distributed on the

branches and to the large flowers. It is related to S. capi-

tcita Cham. & SchltdI. and S. laxa N.E.Br. which also

have 4-merous flowers.

Sehaea capitatci is a more robust plant with thicker

stems and with a different leaf shape — elliptic-ovate or

ovate, cordate, 3-14 mm broad, with a distinct petiole,

whereas S. amicorum has naiTOwly lanceolate leaves 1-2

mm broad, which are sessile. The sepals of S. capitata

have a keel which is ± 1.2 mm broad compared to the

very naiTow keel, ± 0. 2-0.3 mm broad, in the new

species. S. capitata has inflorescences which are dense

and compact with short peduncles, whereas E. amicorum

has a loose open inflorescence. The anthers are similar in

both species in that they have a minute apical gland, but

the new species has much longer thecae — 2.4 mm com-

pared to 1 .5 mm long. S. capitata has a style without or

rarely with a stigmatic swelling and in S. amicorum there

is no swelling.

Well-developed plants of S. laxa appear very similar

to those of E. amicorum in that they are delicate and

somewhat bushy, with similar lax secondary branching

and with the inflorescences having similar secondary

branching. However, the leaves of S. laxa are ovate, 2-6

mm broad, l-3-nerved abaxially, even though they are

similarly scattered over the whole plant. The flowers of

S. laxa are much shorter and more delicate. Both species

have an equally narrow keel down the sepals. Major dif-

Bothalia 31.2 (2001)

209

900 - 1500 m h

r~l 300 • 900 m .

I 1 Under 300 m |

FIGURE 14. — Known distribution

of Sebaea amicoriim.

ferences between the two species occur in the anthers

with 5. la.xa having short thecae 1.0-1. 2 mm long, the

apex much recurved with a large apical gland hanging on

a short thread-like stalk. There is also a stigmatic

swelling on the style in 5. laxa which is absent in the new

species.

Apart from one unnamed collection, Esterhuysen

36120a, from Towerkop [Toverkop] in the Klein

Swartberg, which has 5-merous flowers and a large api-

cal and basal gland on the thecae and ovate leaves, no

other Sebaea species has been recorded from the very

long Swartberg Mountain Range from near Ladismith to

almost Willowmore. 5. amiconim is represented by sev-

eral collections from this Range but in two distinct

areas — the Klein Swartberg above Ladismith and the

region just east of the Swartberg Pass (Figure 14). Two

of the peaks cited by Esterhuysen and Stokoe,

Koudeveldberg (Koudeberg) and Krevasberg, cannot be

located on any map.

In the Flora of southern Africa, Marais & Verdoom

(1963) cite under S. capitata var. sclerosepala (Schinz)

Marais a collection, Esterhuysen 28035, from the Cocks-

comb in the Great Winterhoek Mtns. This was found to

be clearly the same as the new species, but is far removed

from the other two areas. This feature is not surprising in

the genus Sebaea in which many species have very wide

and disjunct distributions.

The species has been recorded by Esterhuysen from

ledges, the base of cliffs and shady gullies mostly on the

southern slopes of the Klein Swartberg at altitudes

between 5500 and 6500 ft [1 670 and 1 980 m]. Linder

recorded ‘shady, mossy S-facing ledges’. The type popu-

lation was found on a moist stony S-facing slope just

below the summit ridge in short grassy/restioid vegeta-

tion that had been burnt the previous year. The

Cockscomb collection came from ‘steep rocky SE

slopes, marshy spot in gully’.

This new species is named amicorum = of the friends,

because we have been closely involved as colleagues in

the herbarium for the last 15 years.

Paratype material

WESTERN CAPE.— 3321 (Ladismith): Klein Swartberg,

Towerkop, 5500-6000 ft [1 670-1 830 m], (-AC), 16-12-1956,

Esterhuysen 26748 (BOL); ridge E of Towerkop, 2 000 m, (-AC), 7-

02-1992, Linder 5518 (PRE, photocopy); E. of Towerkop facing

Elandsberg, 6000-6500 ft [1 830-1 980 m], (-AC/AD), 29-03-1964,

Esterhuysen 30665 (BOL); ridge from Kouveldberg to Towerkop, ±

6500 ft [1 980 m], (-AC/AD), 31-04-1964, Esterhuysen 30669 (BOL);

between Towerkop and Elandsberg/Koudeveldberg. 6000 ft [1 830 m],

(-AC/AD), 28-03-1959, Esterhuysen 28244 (BOL). 3322

(Oudtshoorn): Swartberg Pass area, Krevasberg, summit, (-?AC), 12-

1942, Stokoe 8996 (BOL).

EASTERN CAPE. — 3324 (Steytlerville); Great Winterhoek Mtns,

Cockscomb. 5500 ft [1 670 m], (-DB), 30-1 1-1958, Esterhuysen 28035

(BOL).

REFERENCE

MARAIS, W. & VERDOORN, I.C. 1963. Sebaea. Elora of southern

Africa 26: 171-211.

I.M. OLIVER* and J.B.P. BEYERS*

* Compton Herbarium, National Botanical Institute, Private Bag XI,

7735 Claremont, Cape Town. MS. received: 2001-05-16.

FABACEAE

THE IDENTITY OE MELOLOBIUM LAMPOLOBUM (PAPILIONOIDEAE)

The genus Melolobium is currently being revised and

a comprehensive survey of the literature and herbarium

specimens has revealed a poorly known species that was

recently re-collected. The species was listed by Bentham

(1844) as a synonym of M. collinum Eckl. & Zeyh. and

by Harvey (1862) as a variety of M. microphyllum Eckl.

& Zeyh., but our studies have shown it to be distinct. The

identity of the plant became quite apparent when the

original material was discovered in the Paris Herbarium.

One of Drege’s types clearly shows the diagnostically

different pods of the species. When E. Meyer (Eeb.1836)

described the genus Sphingium, it had already been

described a month earlier as Melolobium by Ecklon &

Zeyher (Jan. 1836). In subsequent publications some of

Meyer’s names appear to have been overlooked.

There are two Drege’s specimens in the Paris Her-

barium. Drege 6463 (Figure 15 A) is a flowering speci-

men with no locality label, Drege s.n. (Figure 15B) is a

fruiting specimen with the locality label ‘Kendo’ in

210

Bothalia 31,2 (2001)

Mulolohiiim lampolnhiim (E Mey ) A Moteelce & B-E.

Dct. A Moteetee

05/02/2001

Wyk

A

Melolobhim lampolohimi (E Mey.) A, Molcclec & B-E. van

Del. A. Moteetee

0.5/02/2001

Wyk

B

FIGURE 15. — Type specimens of Melolohiwn lampolobum in the Paris Herbarium. A. flowering specimen, Drege 6463 (syntype); B, fruiting

specimen Drege s.n. (lectotype).

Drege’s handwriting. Because of the diagnostic impor-

tance of the fruit and the unambiguous type locality

details, the latter is here chosen as lectotype.

Melolobium lampolobum (E.Mey.) A.Moteetee &

B-E.van Wyk, comb. nov. Type: Western Cape, 3322

(Oudtshoom): ‘Kendo’, (-BD), Drege s.n. (P!, specimen

with pods, lectotype, here designated); Drege 6463 (P!,

specimen with flowers, syntype).

Sphingium lampolobum E.Mey.: 67 (Feb. 1836). M. microphyUwn

L. var. lampolobum (E.Mey.) Harv,; 79 (1862).

Woody, strongly spinescent shrublet up to 0.6 m high.

Leaves stipulate; leaflets oblong to broadly obovate,

(3-)5-9 X 2-A mm, glabrous, apex somewhat mucronate

to emarginate; petiole 2-5 mm long; stipules semi-cor-

date to auriculate, 1-4 x 1-2 mm. Inflorescence slender,

terminal, 40-60 mm long, many-flowered raceme

(10-16 flowers); llowers 7-10 mm long; bracts ovate-

lanceolate, 2-3 X 0.5-1 .0 mm; bracteoles narrowly ovate

to lanceolate, 2-3 x 0.5-0. 7 mm. Calyx shortly bilabiate,

glandular with sessile glands, very sparsely hairy; upper

lobes acute, 5-6 mm long, apices obtuse, upper sinus 2-3

mm deep; lower lobes acute, 6-7 mm long, apices

obtu.se, lower sinuses 0.2-1 .0 mm deep. Corolla yellow;

standard suborbicular, 6-8 x 4-5 mm, with well-devel-

oped, channelled claw, 2-3 mm long; wing petals

oblong, sculptured, 7-8 x 2-3 mm, with linear claw 3—4

mm long; keel petals shortly half-oblong, apically round-

ed, 3^ X 2-3 mm, with linear claw 3— f mm long.

Amlroeciwn monadelphous, split on its upper side, con-

sisting of four long, basifixed anthers and six short, dor-

sifixed anthers (alternating with the carinal, intermediate

one). Gynoecium narrowly oblong, 3. 0-3. 5 mm long,

hairy, with 4 or 5 ovules; style curved, 3^ mm long.

Endl broadly falcate, strongly compressed, 12-18 x 2-4

mm, without glands, almost glabrous, surface distinctly

shiny, 2-4-seeded; seeds discoid, light brown, 2. 3-2. 5

mm diam. (side view). Figure 16.

Diagnostic characters: the shiny pod of the aptly

named M. lainpolohinn is a very useful diagnostic feature.

Unlike many other related species, the pods are devoid of

both sessile and stalked glands. The glabrous, obcordate

leaflets and general morphology indicate an affinity to M.

exudans Harv., the only other species in the entire genus

with glabrous leaves. M. lampolobum differs from the lat-

ter in the strongly spiny, curved branches (slightly spiny

in M. exudans), pubescent stems (glabrous in M. exudans)

and glabrescent pods (glandular and hairy in M. exudans).

Because of its branching pattern and the rigid spines, this

species can be confused with M. candicans (E. Mey.)

Eckl. & Zeyh., from which it differs in the dark brown

and pubescent branches (distinctly white-tomentose in

most forms of the latter), larger leaves, falcate, shiny pods

(straight and densely hairy in M. candicans) and longer

inflorescences with more flowers.

Distribution and habitat: the known distribution of M.

lampolobum is shown in Figure 17. It occurs at lower alti-

tudes near Robertson and further northeast in the moun-

tains of the Little Karoo in the southern part of Western

Cape, extending from the western end of Anysberg along

the Klein and Groot Swartberg Mountains as far east as

Oudtshoom. The plants grow in mountain karoo scrub

and on rocky slopes, at altitudes of 900 to over 1 500 m.

Additional specimens e.xamined

WESTERN CAPE. — 3319 (Worcester): Vrolijkheid. Robertson.

(-DD). Van der Merwe 3001 (PRE). 3320 (Montagu): Karoo Garden.

Whitehill. (-BA). Compton 11213 (NBG): western end of Anysberg at

Booplaas Earm. (-BC). C.M. van Wyk 1080 (PRE); 16.6 km SSE of

Laingsburg. (-BD). B-E. van Wyk2143, 2145 (JRAU). 3321 (Ladismith):

Ladismith. (-AD), Baylis.s 2817 (NBG); Groot Swardberg. (-BD),

Marshall 234 (JRAU. PRE); Swartberg Mountains next to Gamkaskloof,

(-BD), Vlok 1489 (PRE); range N of Sandberg. (-DA), Wurt.s 1393

(NBG).

ACKNOWLEDGEMENTS

We thank the curators of the cited herbaria for loans

of specimens. The Government of Lesotho and the

National Research Foundation are thanked for financial

support.

Bothalia 31,2 (2001)

211

FIGURE 16. — Melolobium lampolobum. A. abaxial view of leaf with stipules; B. adaxial view of young leaf with stipules; C. lateral view of

flower; Dl, abaxial view of bract; D2 abaxial view of bracteoles; E. calyx opened out with upper lobes to left; F, standard petal; G, wing

petal; H. keel petal; II, long, basifixed anther; 12, carinal (intermediate) anther; 13, short, dorsifixed anther; J, pistil; K, lateral view of pod.

Scale bars: A-K. 1 mm.

HEIGHT ABOVE SEA LEVEL

EM Over 1500 m

900 - 1500 m

300 - 900 m

I I Under 300 m

0 20 40 60 00 100 km

FIGURE 17. — The known distribu-

tion of Melolobium lampolo-

bwn.

212

Bothalia 31,2 (2001)

REFERENCES

BENTHAM, G. 1844. Enumeration of Leguminosae, indigenous to

southern Asia and central and southern Africa. In J.D. Hooker.

London Journal of Botany 3: 350-353.

ECKLON, C.F. & ZEYHER, C.L.R 1836. Emimeratio plantaruin

Africae australis extratropicae: 188-190. Perthes & Besser,

Hamburg.

HARVEY, W.H. 1862. Leguminosae. In W.H. Harvey & O.W. Sender,

Flora capensis 2: 77-81. Hodges Smith, Dublin.

MEYER, E.H.F. 1836. Commentariorum de plantis Africae aus-

tralioris: 65-67. Leopoldum Voss, Leipzig.

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

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

2006 Auckland Park, Johannesburg.

MS. received: 2001-03-19.

Bothalia 31,2: 213-236 (2001)

Taxonomic significance of inflorescences, floral morphology and

anatomy in Passerina (Thymelaeaceae)

C.L. BREDENKAMP* and A.E. VAN WYK**

Keywords: anatomy, androecium, exotegmen, flower, gynoecium, morphology, Passerina, phylogeny, taxonomy, Thymelaeaceae, vasculation

ABSTRACT

Comparative studies were undertaken on the inflorescence, bracts and floral morphology of all taxa of the genus

Passerina L. in southern Africa. Information is given in tabular form and a key based on bract morphology is pre.sented.

Floral morphology supported the status of the infrageneric taxa and also proved to be of taxonomic significance in the

genus. Controversy surrounding the interpretation of a number of floral morphological structures in Passerina has been

resolved. Morphological and anatomical evidence allowed a re-interpretation of the structure of the receptacle, hypanthium

and sepals, ovary type and position, structure of the seed coat, ovule type and position, obturator, fruit and seed. On this

basis an authentic generic description of the floral morphology was compiled. Passerina is distinguished by the following

set of characters, a very short floral receptacle, tubular hypanthium, petaloid calyx, absence of petals and petaloid scales,

diplostemonous dimorphic androecium, extrorse anthers, superior ovary, anatropous, ventrally epitropous ovule, an obturator

of elongated cells, a 1 -seeded berry or an achene and tegmic seed with nuclear endosperm becoming cellular throughout.

On this basis the flower in Passerina is considered a phylogenetically advanced structure, supporting the view that the genus

is advanced within the Thymelaeoideae. The proposed taxonomic relationship between Thymelaeaceae and Malvales is

confirmed by floral morphological evidence.

CONTENTS

Introduction 213

Materials and methods 216

Terminology 216

Phylogeny 216

Results 216

Inflorescences 216

Bracts 218

Key to taxa based on bract characters 218

Generic description of floral morphology 219

Generic description of floral anatomy (as seen in

transverse section) 219

Floral morphology at species level 223

Discussion 223

Inflorescence 223

Bracts 223

Floral morphology and anatomy at generic level . 223

Floral and fruit morphology at species level .... 232

Less important taxonomic characters 234

Taxonomic relationships 234

Speculations on phylogeny 235

Systematic value 235

Conclusions 235

Acknowledgements 236

References 236

INTRODUCTION

The infrageneric taxonomy of Passerina L. is a prob-

lem, due to the apparent lack of marked morphological

differences between species. The latest revision by

Thoday (1924), is now mostly outdated.

* National Botanical Institute. Private Bag XlOl. 0001 Pretoria.

** H.G.W.J. Schweickerdt Herbarium. Department of Botany, Univer-

sity of Pretoria, 0002 Pretoria.

MS. received: 2000-08-22.

Most species of Passerina are endemic to the Cape

Floristic Region and adapted to a Mediterranean or semi-

Mediterranean climate. The distribution of P. sp. nov. 4

and P montana extends eastwards and northwards along

the eastern mountains and Great Escarpment of southern

Africa, predominantly summer rainfall areas. The

species are apparently all wind-pollinated.

The controversy surrounding the interpretation of

certain floral structures in the genus became obvious

during the present study. Heinig (1951) did not include

Passerina in her study of the floral morphology of the

Thymelaeaceae. From the sexual system of Linnaeus (1781,

1784) to Domke (1934), floral morphology played an

integral part in the intrafamilial classification of the

Thymelaeaceae and in the circumscription of the family

(Table 1). Between ± 1960 and 1996, vast leaps were

taken in the classification of the flowering plants by

making use of anatomical, floral, palynological, embryo-

logical and chemical evidence. During this period, dis-

agreement on the circumscription of the Thymelaeaceae

was common (Table 2). Since 1990, many higher-level

phylogenetic relationships in angiosperms were based on

evidence from rbcL and 18S nuclear ribosomal DNA

sequence data. In many cases these studies confirmed

previously proposed phylogenetic patterns and trends

within the family, based on morphological evidence; in

other cases, however, profound changes in the circum-

scription and relationships of the Thymelaeaceae were

suggested (Table 3).

The present paper emanated from a monographic

study of the genus currently in progress. Available evi-

dence suggests the presence of at least four new species

and four new subspecies, to be added to 16 previously

recognized species, mostly endemic to southern Africa

(Thoday 1924; Bond & Goldblatt 1984) (Table 4). Here

we report on a comparative morphological study of the

inflorescence, bract, flower, fruit and seed. Specific

Classitication of Passerimi within the Thymelaeaceae according to different authors, based mainly on floral, fruit and .seed morphology

214

Bothalia 31.2 (2001)

black, crustaceous testa,

micropyle beak-like

TABLE 2, — Classification of Passerina within tlie Thymelaeaceae according to different authors, based on morphological, anatomical, palynological, embryological and chemical characters

Bothalia31,2 (2001)

215

216

Bothalia 31,2 (2001)

TABLE 3. — Classification of the Thymelaeaceae according to different authors, based on molecular data

* Angiosperm Phylogeny Group.

results in leaf anatomy, indicating the arrangement of

taxa in Passerimi (Bredenkamp & Van Wyk 2001), are

associated with floral morphological structures as well as

fruit and seed types in the present study,

MATERIALS AND METHODS

As far as possible, material was collected from at least

five different localities for every taxon. Live and pre-

served (dried and in liquid preservatives) material of all

the species, subspecies and varieties in Passerimi was

studied (Table 4). Illustrations were made from herbari-

um material by means of a drawing tube. Measurements

were taken by using a dissection microscope and a cali-

brated eyepiece.

Flowers were fixed and stored in a 0. 1 M phosphate-

buffered solution at pH 7.4, containing 2.5% formalde-

hyde, 0.1% glutaraldehyde and 0.5% caffeine [modified

Karnovsky fixative; Karnovsky (1965)]. Light micro-

scopy (LM) was used to study the floral anatomy of P.

ericoides (Bredenkamp 956, 962), which has fleshy fruit

and P. vulgaris (Bredenkamp 944, 951) with dry fruit. As

the flowers are quite small, whole flower buds, flowers

directly after anthesis and young fruits enveloped in the

floral tube were washed in water, dehydrated and embed-

ded in glycol methacrylate (GMA) following the meth-

ods of Feder & O’Brien ( 1968). Embedded floral mater-

ial was serially sectioned from the base of the receptacle

to the anthers. Sections were stained in toluidine blue

‘O’, subjected to the periodic acid-Schiff’s (PAS) reaction

and mounted in Entellan (Art. 7961, E. Merck, Damstadt).

Terminology

Terminology is used according to the following

authorities: inflorescences (Weberling 1983, 1989;

Weberling & Herkommer 1989); hypanthium (Bunniger

1972; Dahlgren 1975a, b, 1980; Dahlgren & Thorne 1984);

stamen morphology (Heinig 1951; Fahn 1967; Noel 1983);

gynoecium (Heinig 1951; Davis 1966; Comer 1976); fruit

(Spjut 1994).

Floral envelope

The authors regard the floral envelope (‘outer floral

whorl’) as a hypanthium (fused calyx and androecium),

differentiating into four petaloid sepals and a diploste-

monous androecium, arising from the hypanthium rim at

the separation of the sepals. For the description of colour,

texture and measurement of total floral length, only the

hypanthium and sepals are considered — the stamens are

excluded.

Phylogeny

Speculations on phylogeny are based on prevailing

family characters representing the ancestral state and

derived characters, indicating a reduction in tissue at

genus and species levels, considered as possibly

advanced. A cladistic analysis based on anatomical and

morphological characters in Passerina is scheduled for

the final stages of the study.

RESULTS

Inflorescences

Polytelic synflorescences present in all species; main

florescences and co-florescences spicate (Figures 1 A; 2A),

often extended, forming multiflowered politelic flores-

cences in most species, sometimes strongly reduced

(P. hurchellii) (Table 5). Proliferating spikes with inflores-

cence apex growing out and returning to vegetative growth,

common; main florescences and co-florescences subtenni-

nal. Spikes sometimes artificially resembling terminal sub-

capitulate inflore.scences, but each characterized by two

terminal leaves with axillary blind-ending mdimentary

flowers, enveloping minute growing point (P. montana, P.

paleacea, P. glomerata, P. sp. nov. 3); proliferating growth

less common in subcapitulate inflorescences.

Bothalia 31.2 (2001)

217

TABLE 4. — Passerina specimens examined and housed at PRE

Species

burchellii Thoday

comosa C.H. Wright

drakensbergensis Hilliard

& B.L.Burtt

ericoides L.

falcifolia C.H. Wright

filifonnis L. snbs.p. filifbnnis

filiformis L. subsp. nov.

galpinii C.H. Wright

Collector

Bredenkamp 1545

Bolus 687*. Stokoe 2542

Thoday 212

Andreae 1288*,

MacDonald 2125

Edwards 974

Bredenkamp 1018, 1019.

1020. 1021*

Bredenkamp 956'

Bredenkamp 962'*

Taylor 4042

Bredenkamp 917*

Bredenkamp 915

Tyson 1449

Boucher 2833

Bredenkamp 1039*

Schlechter 5125*

TaMor 1542

Bredenkamp 946*

Bredenkamp 932

Bredenkamp 933

Bredenkamp 923

Bredenkamp 988

Bredenkamp 994

Bredenkamp 1002

Bredenkamp 984, 985*

Bredenkamp 977

Bredenkamp 973*

glomerata Thunb. subsp. nov. Stokoe 8040

Esterhuvsen 28587*

glomerata Thunb. subsp.

glomerata

montana Thoday

obtusifolia Thoday

paleacea Wikstr.

paludosa Thoday

pendula Eckl. & Zeyh.

rigida Wikstr.

rubra C.H.Wright

vulgaris Thoday

sp. nov. 1

sp. nov. 2

sp. nov. 3

sp. nov. 4

Giess 13136

Bredenkamp 1024

Bredenkamp 1025

Bredenkamp 889. 890

Bredenkamp 893*

Bredenkamp 971

Bredenkamp 967

Bredenkamp 1033, 1034

Bredenkamp 929

Bredenkamp 919*

Bredenkamp 960*

Bredenkamp 961

Bredenk inp 952

Bredenkamp 950

Bredenkamp 949

Bredenkamp 940

Bredenkamp 1035

Jangle 156*

Thoday 100

Fourcade 3043

Bredenkamp 908. 909*

Ward 7211

Bredenkamp 1013*

Bredenkamp 899

Bredenkamp 898

Bredenkamp 897

Bredenkamp 911

Bredenkamp 914*

Bredenkamp 905

Bredenkamp 900

Bredenkamp 926

Bredenkamp 907

Bredenkamp 901

Bredenkamp 95h

Bredenkamp 944'*

Bredenkamp 924

Locality

WESTERN CAPE. — 3319 (Worcester): Jonaskop. (-DC).

WESTERN CAPE. — 3419 (Caledon): Baviaanskloof, Genadendal. (-BA).

WESTERN CAPE. — 3320 (Montagu): Montagu District, near Concordia. (-CD).

WESTERN CAPE. — 3321 (Ladismith): Seweweekspoort. (-AD).

KWAZULU-NATAL. — 2828 (Bethlehem): Royal Natal National Park. (-DB).

KWAZULU-NATAL. — 2829 (Harrismith): Ndedema Gorge. Cathedral Peak Forest

Reserve. (-CD).

WESTERN CAPE. — 3318 (Cape Town): Milnerton. (-CD).

WESTERN CAPE. — 3418 (Simonstown): Cape Maclear, (-AD).

WESTERN CAPE. — 3419 (Caledon): Pearly Beach. (-CB).

WESTERN CAPE. — 3323 (Willowmore): Gouna State Forest. (-CC).

WESTERN CAPE. — 3324 (Steytlerville): opposite Tsitsikama Lodge. (-CD).

WESTERN CAPE. — 3423 (Knysna): Knysna. (-AA).

WESTERN CAPE. — 3118 (Van Rhynsdorp): Clanwilliam Dist.. Diepkloof S of Verlore-

vlei. (-AD).

WESTERN CAPE. — 3318 (Cape Town): Signal Hill. (-CD).

WESTERN CAPE. — 3218 (Clanwilliam): Alexander's Hoek. (-BC).

WESTERN CAPE. — 3218 (Clanwilliam): Malmesbury Div.. Vredenburg. Steenberg's

Cave. (-CC).

WESTERN CAPE. — 3420 (Bredasdorp): De Hoop Nature Reserve. (-AD).

WESTERN CAPE. — 3421 (Riversdale): Riethuiskraal, (-AD).

WESTERN CAPE. — 3421 (Riversdale): Still Bay. (-AD).

WESTERN CAPE. — 3422 (Mossel Bay): Mossel Bay, (-AA).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains. Eikeboom, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains. Kromrivier, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains. Algeria. (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Citrusdal. Piekenierskloof Pass. (-CD).

WESTERN CAPE. — 3219 (Wuppertal): Groenfontein. (-DC).

WESTERN CAPE. — 3319 (Worcester): Tulbagh. (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains.

WESTERN CAPE. — 3319 (Worcester): Hex River Mountains.

NAMIBIA. — 2217 (Windhoek): Auas Mountains. Molteblick, (-CA).

MPUMALANGA.— 2430 (Pilgrim’s Rest): World’s View, (-DD).

MPUMALANGA. — 2430 (Pilgrim’s Rest): God’s Window. (-DD).

FREE STATE. — 2828 (Bethlehem): Golden Gate National Park. (-DA).

FREE STATE. — 2927 (Maseru): Ladybrand. (-AB).

WESTERN CAPE. — 3319 (Worcester): Karoo National Botanical Garden. (-CB).

WESTERN CAPE.— 3319 (Worcester): Jonaskop. (-CD).

WESTERN CAPE. — 3321 (Laingsburg): Seweweekspoort. (-AD).

WESTERN CAPE.— 3321 (Ladismith): Rooiberg, (-CB).

WESTERN CAPE. — 3322 (Oudtshoom): Perdepoort. (-CD).

WESTERN CAPE. — 3418 (Simonstown): Kommetjie. (-AB).

WESTERN CAPE. — 3418 (Simonstown): Cape Maclear, (-AD).

WESTERN CAPE. — 3418 (Simonstown): Harold Porter National Botanical Garden, (-BD).

WESTERN CAPE. — 3420 (Bredasdorp): De Hoop Nature Reserve, (-AD).

WESTERN CAPE. — 3420 (Bredasdorp): Waenhuiskrans, (-CA).

WESTERN CAPE.— 3421 (Riversdale): Puntjie, (-AC).

WESTERN CAPE. — 3418 (Simonstown): Rondevlei Nature Reserve, (-BA).

WESTERN CAPE. — 3418 (Simonstown): Rondevlei Nature Reserve, (-BA).

WESTERN CAPE. — 3418 (Simonstown): Riet Valley, Cape Flats, (-BA).

EASTERN CAPE. — 3324 (Steytlerville): Zuur Anys, (-CB).

EASTERN CAPE. — 3325 (Port Elizabeth): Groendal Nature Reserve, (-CB).

KWAZULU-NATAL.— 2832 (Mtubatuba): St Lucia Park, (-AD).

KWAZULU-NATAL. — 3130 (Port Edward): Umtamvuna River Mouth, (-AA).

EASTERN CAPE. — 3326 (Grahamstown): Kenton-on-Sea, (-DA).

EASTERN CAPE. — 3326 (Grahamstown): Port Alfred, (-DB).

EASTERN CAPE.— 3327 (Peddie): Kleinmond West, (-CA).

EASTERN CAPE, — 3424 (Humansdorp): Jeffreys Bay, (-BB).

EASTERN CAPE. — 3324 (Steytlerville): en route to Kareedouw, (-CD).

EASTERN CAPE. — 3325 (Port Elizabeth): Colchester. (-DB).

EASTERN CAPE. — 3326 (Grahamstown): Grahamstown. (-AD).

WESTERN CAPE. — 3321 (Ladismith): en route to Korenthe River Dam. (-CC).

EASTERN CAPE. — 3325 (Port Elizabeth): Groendal Nature Reserve, (-CB),

EASTERN CAPE. — 3326 (Grahamstown): Grahamstown, (-AD).

WESTERN CAPE. — 3418 (Simonstown): Betty’s Bay, Harold Porter NBG, (-BD).

WESTERN CAPE.— 3420 (Bredasdorp): Cape Infanta. (-BD).

WESTERN CAPE.— 3422 (Mosselbaai): Kleinbrak. (-AA).

Goldblatt & Manning 8627 WESTERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment. (-AB).

Bredenkamp 1044, 1046,

1047

Oliver 9281*

Esterhuvsen 12189, 26859*

Stokoe 9302

Schlechter 5846

Esterhuvsen 10734*

Esterhuvsen 28006*

Killick 238

Bredenkamp 1016, 1017

Van Wvk & Bredenkamp

1, 1012

Bredenkamp 1327*

Gillett 4537

Keet s.n.

WESTERN CAPE. — 3319 (Worcester): Waboomberg. Ceres, (-AD).

WESTERN CAPE. — 3218 (Clanwilliam): Northern Cederberg Mountains, (-BB).

WESTERN CAPE. — 3322 (Oudtshoom): Swartberg Pass, Prince Albert area, (-AC).

WESTERN CAPE. — 3322 (Oudtshoom): Montagu Pass, (-CD).

EASTERN CAPE. — 3323 (Willowmore): Kouga Mountains. (-DA).

EASTERN CAPE. — 3324 (Steytlerville): Cockscomb, Uitenhage area, (-BD).

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Table Mountain, (-CB).

KWAZULU-NATAL.— 3030 (Port Shepstone): Oribi Gorge, (-CB).

KWAZULU-NATAL. — 3130 (Port Edward): Umtamvuna River Bridge. (-AA).

KWAZULU-NATAL. — 3130 (Port Edward): Mkambati Nature Reserve, (-AC).

WESTERN CAPE. — 3422 (Mossel Bay): en route to Knysna from George, (-BB),

WESTERN CAPE.— 3423 (Knysna): Knysna. Redlands. (-AA).

Illustrated specimens; • specimens used for light microscopy.

218

Bothalia 31,2 (2001)

FIGURE 1 . — Passerina ericoides.

A-G, Bredenkamp 962. A,

young inflorescence; B, leaf

exposing adaxial surface; C,

bract exposing adaxial sur-

face; D, fruiting branchlet; E,

flower enveloped by bract; E,

fleshy one-seeded berry; G,

seed with black tegmen and

white fragment of endocarp.

A, D, X 10; B, C, E-G, x 20.

Scale bars: 10 mm.

Bracts

Arrangement subterminal in proliferating spikes,

pseudoterminal in subcapitulate spikes; youngest bracts

closest to growing point, enlarging along florescence

axis; bracts enveloping flowers and fruits (Figures lA,

D, E; 2 A, D), largest after anthesis of flowers, becom-

ing more coriaceous and rounded at fruit set, gradually

acquiring leaf shape at proximal end of florescence

axis, decussate, sessile, imbricate, often conspicuously

enlarged, resulting in longer spikes (P.falcifolia, P.fili-

forrnis, P. sp. nov. 4, P. rubra). Lamina inversely ericoid;

adaxial surface (inside) concave, facing inflorescence

axis, abaxial surface (outside) convex; cymbiform (boat-

shaped) or helmet-shaped; in outline (plane shape), ob-

long, lanceolate, ovate and obovate to widely ovate and

obovate, rhombic and narrowly obtrullate to obtrullate

(Figures 3; 4; Table 5); bracts without leaf-like point,

length X 'L width (2.5-)3.5^.5(-5.5) x (0.9-)1.0-1.5(-2.4)

mm; bracts with leaf-like point, length x 'A width (4.0-)

5.1-6.3(-7.3) X (1.4-)1.5-2.0(-2.6) mm; outside glabrous,

seldom tomentose {P. comosa, P. sp. nov. 3), inside, base

or midrib tomentose with glabrous wings or completely

comose, tomentose, villous or setose; trichomes nong-

landular, uniseriate, mostly strongly spiralled, white;

coriaceous or chartaceous, rugose or smooth, ± succu-

lent or thin, outline of epidermal cells often macroscop-

ically visible; often ± 3-5-ribbed or reticulately veined

on each side of main vein. Wings absent, or bordering

lamina or distinct from lamina and conspicuously

rounded or bullate; mostly glabrous, coriaceous, charta-

ceous or membranous, often straw-coloured and rosy-

tinged. Base sessile or cuneate. Midrib strongly devel-

oped, forming spine of cymbiform bract, often keeled,

extending to form a leaf-like point in many species.

Apex obtuse, rounded or acute, mostly coriaceous.

Margins often ciliate to setose (P. burchellii, P. sp. nov.

4, P. pendiila).

EIGURE 2. — Passerina vulgaris,

Bredenkamp 944. A, inflores-

cence; B, leaf exposing adax-

ial surface; C, bract exposing

abaxial surface; D, flower

enveloped by bract; E, fruit

enveloped by hypanthium,

with fragmentation at neck

base; F, achene; G, seed with

black tegmen and white

spots. A, X 10; B-G, x 20.

Scale bars: 10 mm.

Key to taxa based on bract characters (Figures 3; 4; Table 5)

la Bracts hairy outside P. conwsa (Figure 3A), P. sp. nov. 3 (Figure 3B, C)

lb Bracts glabrous outside:

2a Bracts shorter than 4.5 mm:

3a Bracts rhombic in outline:

4a Bracts dark green when fresh, dark brown in dried specimens, coriaceous; membranous wings absent . . .

P. hurchellii (Figure 3D)

4b Bracts blue-green (glaucous) In fresh and dried specimens, softly coriaceous; membranous wings present

P. pendida (Figure 3E)

Bothalia 31,2 (2001)

219

3b Bracts variously shaped, but not rhombic in outline:

5a Membranous wings present:

6a Wings present as membranous rims along lamina margins:

7a Lamina convex, ovate to obovate, thinly coriaceous, obscurely ribbed P. montana (Figure 3F)

7b Lamina helmet-shaped, widely obovate, thinly chartaceous, smooth P. sp. nov. 2 (Figure 3G)

6b Wings variously shaped, but not as membranous rims along lamina margins:

8a Wings widely ovate, evenly concave, main vein forming small, obtuse apex P. paleacea (Figure 3H)

8b Wings oblate or obovate, often bullate:

9a Main vein extended into leaf-like point P. galpinii (Figure 31)

9b Main vein shortly extended into subacute apex P. sp. nov. 1 (Figure 3J)

5b Membranous wings absent:

10a Bracts oblong in outline, apex obtuse p. ericoides (Figure 3K)

10b Bracts widely ovate or obovate to widely obovate in outline, apex various:

1 la Bracts widely ovate with distinct, short acute point, glaucous, coriaceous; lamina with 2 or 3 shallow

folds p. rigida (Figure 3L)

1 lb Bracts obovate to widely obovate, point obtuse to acute, thickly coriaceous; lamina ribbed and reticu-

lately veined P. glome rata subsp. glomerata (Figure 4A), P. glomerata subsp. nov. 1 (Figure 4B)

2b Bracts longer than 4.5 mm:

12a Bracts lanceolate and glaucous P. drakenshergensi.s (Figure 4C)

12b Bracts variously shaped and coloured, but not lanceolate and glaucous:

13a Main vein shortly extended into an acute apex; lamina rhombic to obtrullate, distinctly angled. 4- or 5-

ribbed p, vulgaris (Figure 4L)

13b Main vein extending beyond lamina into a leaf-like point:

14a Leaf-like point obtuse at apex; lamina closely 2-ribbed at margins P. ohtusifolia (Figure 4D)

14b Leaf-like point not obtuse; lamina extending into a membranous margin:

15a Adaxial (inner) surface of bracts basally to centrally setose or tomentose over entire length of midrib;

wings glabrous:

16a Midrib and leaf-like point stout and strongly developed, apex acute P. paludosa (Figure 4E)

16b Midrib shortly extended or forming a straight or filiform, leaf-like point:

17a Midrib shortly extended, wings ovate-acuminate, gradually narrowing to a point

P.filifonnis subsp. nov. (Figure 4F)

17b Midrib extended, forming a straight or filiform, leaf-like point:

18a Midrib extended into a filiform, slightly falcate point; wings widely obovate, narrowing abruptly

into a leaf-like point P.filifonnis subsp. filiformis (Figure 4G)

18b Midrib extended into a straight point or slightly incurved point; wings ovate with margins hairy

in upper half, or obtrullate, narrowing abruptly into a leaf-like point . , . . F. sp. nov. 4 (Figure 4H, I)

15b Adaxial (inner) surface of bracts completely villous;

19a Midrib extended, leaf-like point falcate; wings ± 4-ribbed P. falcifolia (Figure 4J)

19b Midrib shortly extended into a short point, apex acute; wings ± 5-ribbed P. rubra (Figure 4K)

Generic description of floral morphology

Flowers actinomorphic, bisexual, hypogynous. Floral

envelope membranous during pollination and yellowish

in P. rigida, P. paleacea, P. sp. nov. 1 and P. sp. nov. 2,

slightly succulent and greenish in P. ericoides, mostly

yellow-pink in all other species, dehydrated after shed-

ding of pollen, becoming papyraceous or coriaceous,

yellow-pink tones turning red, (4.0-)5.3-7.3{-8.4) mm

long. Pedicel very short or absent. Receptacle very short.

Hypanthium a membranous to coriaceous cylindric tube;

indumentum variable in density, trichomes nonglandular,

uniseriate, often spiralled, whitish, density of indumen-

tum at ovary ranging from glabrous to tomentose or

strigose; neck (narrowed tube between apex of ovary and

sepals) (0.3-)0.6-2.6(-3.0) mm long, density of indu-

mentum ranging from glabrous to tomentose on outside,

inside often hairy, abscission tissue not macroscopically

discernible, articulation plane absent, after fruiting frag-

mentation of neck base caused by dehydration and torsi-

fication of tissue, shedding sepals and androecium

(Figure 2D, E; Table 8) in most species. Sepals petaloid

(Figures IE; 2A, D); lobes 4, imbricate in bud (Figure

8C, D), flexed in flower, often setose with up to 5 long

trichomes on outer surface and glabrous to tomentose on

inner surface; outer lobes cymbiform or concave; inner

lobes oblong, elliptic or obovate. Corolla absent.

Petaloid scales absent. Androecium dimorphic diploste-

monous, inserted at rim of hypanthium, filaments of

antipetalous whorl (0.4-)0.7-1.2(-1.5) mm long, anti-

sepalous whorl ( 1.2-)1.4-2.2(-2.4) mm long; anthers

(0.5-)0.7-0.9(-l.l) X (0.2-)0.3-0.4(-0.7) mm, subbasi-

fixed, 2-thecous and 4-locular (Figure 9A), extrorse.

Disc absent. Ovary superior, ( 1 .6-)2.0-2.5(-2.7) x

(0.5-)0.6-1.4(-1.7) mm. bicarpellate during embryonic

stage (Bunniger 1972), pseudomonomerous (Heinig

195 1 ) at maturity, placentation parietal, uniloculate, with

1 pendulous ovule laterally attached near top of ovary;

style separating laterally from top of ovary, maintaining

lateral position in hypanthium neck, reaching beyond

hypanthium rim; stigma ± globose, mop-like or penicil-

late (wind-pollination). Fruit enveloped by persistent,

loosely arranged hypanthium fragmented at neck base or,

in some species, fragmenting over widest circumference

of fruit, shedding fragmented hypanthium, sepals and

androecium (Figures ID; 2E), in P. ericoides (Figure IF)

and P. rigida a fleshy 1-seeded berry, 5.3 x 4.0 mm, in

all other species (Figure 2F; Tables 6, 7) an achene, peri-

carp membranous and dry, 2.5 x 1 .2 mm. Seed: tegmen

black and shiny, often with white spots, broadly fusiform

with outgrowths at both micropylar and funicular ends

(Figures IG; 2G), 2.2(-2.9) x 1.2(-1.6) mm.

Generic description of floral anatomy (as seen in

transverse section)

Receptacle base: vascular tissue arranged in a central

stele from which 8 traces are derived in a single whorl

(Figure 5A). Receptacle at apical position: carpellary

bundles arranged in continuous central cylinder; 8 vas-

220

Bothalia 31,2 (2001)

FIGURE 3. — Camera lucida drawings of abaxial, lateral and adaxial views of bracts of Passerina taxa, arranged according to sequence in key. A,

P. comosa. Andreae 1288: B-C, P. sp. nov. 3, Esterhuysen 10734, 28006; D, P. burchellii. Bolus 687; E, P. pendula, Bredenkamp 908; F,

P. montana. Bredenkamp 893; G, P. sp. nov. 2, Esterhuysen 26859; H, P. paleacea, Bredenkamp 960; I, P. galpinii, Bredenkamp 946; J,

P. sp. nov. 1, Oliver 9281; K, P. ericoides. Bredenkamp 962; L, P. rigida, Bredenkamp 1013. A-L, x 20. Scale bar: 5 mm.

cular bundles, fused sepal and stamen traces occupying

peripheral position (Figure 5B); cells at perifery of cor-

tex arranged in rows (abscission tissue), differentiation

of inner epidermis of hypanthium and outer epidermis of

ovary wall, separating hypanthium from ovary wall

(Figure 5C, D). Hypanthium irregularly lobed or scal-

loped; outer and inner epidermis variously hairy; cuticle

sometimes strongly developed; 8 vascular bundles

stretching over entire length (Figure 5E, F). Calyx with

4 imbricate lobes developing at hypanthium rim, each

containing 3 vascular bundles; epidermal and hypoder-

mal layers containing large amounts of pigment; spongy

parenchyma aerenchymatic (Figure 8C, D; Table 6).

Androecium; each of the 4 fused commisural sepal and

antipetalous stamen bundles (cs-pst) split into 2 sepal

lateral bundles (si) and I antipetalous stamen bundle

(pst), resulting in the first whorl of 4 antipetalous sta-

mens (situated slightly lower in the hypanthium) and

each sepal containing 3 vascular bundles (Figure 8A,

B); each of the 4 fused sepal midrib and antisepalous

stamen bundles (s-sst) split into a sepal midrib bundle (s)

and an antisepalous stamen bundle (sst), forming the

second whorl of 4 antisepalous stamens (Figure 8B, C);

anthers extrorse, with wall of locule comprising epider-

mis and endothecium only, periclinal walls of epidermis

thin and folded inwards, cell wall thickenings of

endothecium ± stellate, with rib-like extensions directed

towards epidermis (Figure 9B), partitions between

loculi withered and ruptured (Figure 9C), accompanied

by final rupturing of outer walls of thecae (Figure 9D).

Ovary ha.se: wall independent of hypanthium or loosely

adhering to hypanthium at distal side away from placen-

ta (Figure 5E, E; Table 6); outer and inner epidermal

walls strongly developed, containing ample amounts of

tanniniferous substances (Eigure 5E) or less sturdy

(Figure 5G); mesophyll of densely arranged parenchy-

Bothalia 31,2 (2001)

221

FIGURE 4. — Camera lucida drawings of abaxial, lateral and adaxial views of bracts of Passerina taxa, arranged according to sequence in key.

A. P. glomerata subsp. glomerata. Bredenkamp 973; B, P. glomerata subsp. nov., Esterhuysen 28587; C, P. drakensbergensis. Bredenkamp

1012; D, P. obtusifolia, Bredenkamp 919; E, P. paludosa. Jangle 156; F, P . fdifonnis subsp, nov., Schlechter 5125; G, P.fdifonnis subsp.

filiformis, Bredenkamp 1039; H, P. sp. nov. 4 from Knysna, Gillen 4537; I, P. sp. nov. 4, Bredenkamp 1327; J, P.falcifolia. Bredenkamp

917; K, P. rubra. Bredenkamp 914; L, P. vulgaris, Bredenkamp 944. A-L, x 20. Scale bar: 5 mm.

TABLE 3. — Comparison of inflorescences and bracts in Passerimi. Sequence of taxa according to key based on bract characters (see text)

222

Bothalia3l,2 (2001)

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223

TABLE 6. — Distinction between P. ericoides and P. vulgaris based on floral anatomy, seen in t/s (Figures 5; 7; 10)

ma cells (Figure 5E, G) or aerenchymatic (Figure 5F,

H); vascular bundles arranged in a single ring-like whorl

(Figure 5B), median and dorsal carpellary bundles sep-

arating from stele (Figure 5C, D), vascular bundles dif-

ferentiating into dorsal carpellary bundle as well as

median and commissural carpellary bundles (Figure 5E,

G), or dorsal carpellary bundle absent or poorly defined

(Figure 5F, H); funiculus ventral (Figures 5; 6); ovule

bitegmic, outer integument (oi) consisting of outer epi-

dermis (oeoi), mesophyll and inner epidermis (ieoi),

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dermis (oeii), mesophyll and tanniniferous inner epider-

mis (ieii) (Figures 5G, H; 6A, B), crassinucellate;

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5G, H), Ovary at median position: ovule wall, integu-

ments and nucellus remaining the same; embryo sac

with 2 cells of functional macrospore (Figure 6A. B),

Ovary' at apical position: placental vascular bundles and

funiculus strongly developed; ovule anatropous, posi-

tion ventrally epitropous; obturator of elongate cells

extending from base of style to micropyle (Figures 6C,

D; 7A, B); outer integument horseshoe-shaped, opening

close to funiculus facing placenta (Figures 6C; 7A, B);

micropyle formed by inner integument, initially facing

upwards (Figures 6C, D; 7A), but close to base of style,

incurved towards placenta (Figure 7B), Style four-lobed

or elliptic, stylar channel well developed (Figure 7C, D),

lined by conducting tissue; vascular bundles 4, 1 per

lobe (Figure 1C) or inconspicuously arranged in a row

along the long axis of elliptic style (Figure 7D; Table 6),

Stigma reaching beyond hypanthium rim, penicillate,

ramified into numerous simple papillae, dispersed

between filament bases (Figure 8A-C), Seed exotegmic,

outer integument undergoing atrophy; outer epidermis

of inner integument (oeii) lengthening and becoming

palisade-like, mesophyll consisting of 2 or 3 layers of

parenchyma, inner epidermis of inner integument (ieii)

tanniniferous (Figure lOA, B); tegmen black, lignified,

still portraying palisade origin (Figure IOC, D); outer

layer of nucellus ornate with cellulose thickenings,

nucellus 3-5 cell layers thick; endosperm formation

nuclear (Davis 1966), but later becoming cellular

throughout (Figure lOD), absorbed by cotyledons con-

taining no starch but copious amounts of oil.

Floral morphology at species level

Floral morphological characters and taxonomically

important fruit characters are summarized in Table 7, and

specialized hypanthium and sepal characters in Table 8,

All these are associated with specific leaf anatomical

characters (Bredenkamp & Van Wyk 2001).

DISCUSSION

Inflorescences

Weberling (1989) regards polytelic synflorescences as

dominant within the Thymelaeaceae. He found monotelic

synflorescences in the Gonystyloideae, a relatively primi-

tive group, as well as certain genera of the Thymelaeoideae

and Aquilarioideae. In the Gnidieae, it was found in

Laclinaea L. (= Cryptadenia Meisn.) (Beyers & Van der

Walt 1995; Beyers 1997), a genus endemic to the Cape

Floristic Region (Beyers 1992). Weberling (1989) never-

theless concluded that it appears impossible to draw any

taxonomic conclusions from the existence of monotelic

synflorescences within these taxa. Passerina is character-

ized by polytelic synflorescences. Most species have multi-

flowered main and co-florescences, and a reduction of

florescences to single and subcapitulate spikes is clearly

shown (Table 5).

Bracts

In their descriptions of the Thymelaeaceae, Domke

(1934) reports the presence or absence of bracts and bracte-

oles, sometimes involucral, and Peterson (1978) mentions

that deciduous or persistent bracts are often present. In

Passerina, single flowers are always enveloped by persis-

tent bracts. In the present study, this constant taxonomic

character has been employed in a key for application in

herbarium and field work (Figures 3, 4; Table 5).

Floral morphology and anatomy at generic level

Receptacle

Historically the interpretation of the receptacle in the

Thymelaeaceae has been controversial. Tables 1 and 2

Fruit Floral envelope (hypanthium and sepals) | Ovary | Filaments Anthers

224

Bothalia 31,2 (2001)

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Bothalia 31.2 (2001)

225

FIGURE 5. — LM photographs of P. ericoides. Bredenkamp 956, with fleshy fruit, and P. vulgaris, Bredenkamp 951, with dry fruit, showing structure of

receptacle and base of ovary. A-D. t/s at different levels of receptacle. A-C. P. vulgaris: A, departure of eight traces from central stele; B, carpel-

lary bundles arranged in ring, fused stamen and sepal traces in peripheral position; C, differentiation of cortex tissue into inner epidennis of hypan-

thium and outer epidermis of ovary wall, separation of dorsal carpellary bundle. D, comparison to similar stage in P. ericoides. E-H, P's at base

of ovary showing vasculation of ovary wall, funiculus and chalaza of pendulous ovule: E. P. ericoides. ovary wall independent of hypanthium,

carpellary bundles in a single whorl; F. P. vulgaris, young bud, ovary wall loosely adhering to hypanthium distally; G. P. ericoides, one cell of

functional macrospore; H, comparison to similar stage in P. vulgaris, reduction of dorsal carpellary bundle, c, carpellary bundle; cc. commissur-

al carpellary bundles; cs-pst, fused commissural sepal and antipetalous stamen bundles; d, area of tissue differentiation; dc, dorsal carpellary bun-

dle; f, funiculus; fm, functional macrospore; h, hypanthium; ieh. inner epidermis of hypanthium; ieow, inner epidermis of ovary wall; ii, inner

integument; me, median carpellary bundle; n, nucellus; o. ovule; oeow, outer epidermis of ovary wall; oi. outer integument; ow, ovary wall; p,

placenta; s-sst. fused sepal midrib and antisepalous stamen bundle; t. trichome; vh. vascular bundle of hypanthium. Scale bars: A-H. 100 pm.

226

Bothalia 31,2 (2001)

FIGURE 6. — LM photographs of P. ericoides, Bredeiikamp 956. with fleshy fruit, and P. vulgaris, Bredenkamp 951, with dry fruit, showing

structure of ovary and ovule. A, B, t/s in median position of ovary: A, P. ericoides, embryo sac and two cells of functional macrospore;

B, higher magnification of similar stage in P. vulgaris. C, D, t/s at top of ovary: C, P. ericoides. placenta, obturator and micropyle

originating from inner integument; D, comparison of similar stage in P. vulgaris. Abbreviations as in Figure 5. es, embryo sac; ieii,

inner epidermis of inner integument; ieoi, inner epidermis of outer integument; m, mesophyll; mi, micropyle; ob, obturator; oeii, outer

epidermis of inner integument; oeoi, outer epidermis of outer integument. Scale bars: B, 10 pm; A, C, D, 100 pm.

show that Meisner (1857) regards the floral envelope as

perigynous and hypocrateriform, implying a cup-shaped

receptacle or hypanthium. Gilg (1891, 1894) describes

the lloral envelope as a cylindric receptacle which is

articulate in the upper half, and Endlicher ( 1837, 1847),

Leandri (1930) and Dahlgren & Thorne (1984) regard

the floral arrangement as perigynous. The present study

indicates that the receptacle (in t/s) is very short (Figure

5A, B) and definitely not cup-shaped. This is evident

from the arrangement of peripheral cortex cells in rows,

followed by differentiation into the inner epidermis of

the hypanthium and the outer epidermis of the ovary wall

(Figure 5D), finally by the .separation of the hypanthium

(including the vascular bundles differentiated from the

stele) from the ovary wall and the pre.sence of trichomes

in the space subsequently formed (Figure 5D-F).

Hypanthium and androecial position

Owing to reduction of the receptacle, the hypanthium

in Passerina is here interpreted as being formed by the

fused calyx and androecium only. The vascular tissue of

the hypanthium constitutes the fused sepal and stamen

traces (Heinig 1951), separating from the central stele in

a single whorl and forming a peripheral ring of eight vas-

cular bundles (Figure 5A, B), which persist throughout

the length of the hypanthium. A similar pattern of fusion

and distribution of vascular tissue has been reported by

Heinig (1951) and for the genus Lacimaea (= Cryptade-

nia) by Beyers ( 1992) & Beyers & Van der Walt ( 1995).

In Passerina the central stele differentiates into carpellary

bundles after the separation of the fused sepal and stamen

traces (Figure 5C-F). It can therefore be concluded that

Bothalia3I.2(2001)

227

FIGURE 7. — LM photographs of P. ehcoides. Bredenkamp 956, with fleshy fruit, and P. vulgaris. Bredenkamp 951, with dry fruit, showing

structure of micropyle and style. A. P. ericoides, obturator, pollen tube penetrating micropyle, opening of outer integument towards

micropyle. B, similar stage in P. vulgaris, style departing laterally. C. D, t/s of style. C. four-lobed style in P. ericoides. D, elliptic style

in P. vulgaris. Abbreviations as in Figure 5. ct. conducting tissue; mi. micropyle; ob. obturator; pt, pollen tube; st, style. Scale bars: C.

D, 10 pm; A. B. 100 pm.

the segments of the floral envelope and the androecium

arise below the gynoecium, the floral arrangement being

hypogynous and the ovary superior (Weberling 1989).

Identity of floral envelope

From Wikstrom (1818) to Takhtajan (1997) (Tables

1; 2) the floral envelope in Thymelaeaceae and, in many

cases Passerina, was variously interpreted as an in-

fundibular corolla, hypocrateriform hypanthium, infun-

dibular perigone, perianth, cylindric receptacle, floral

tube formed by coalescence of four external whorls,

perianth tube, or a hypanthium. Heinig (1951) supports

the interpretation of Leandri (1930) and Domke (1934)

of the floral tube as appendicular in origin, composed

of the fused bases of the sepals and adherent stamen

filaments, also pointing out that the sepal is with few

exceptions a three-trace organ. Bunniger (1972) is of the

opinion that the hypanthiums in families of the Myrtales

and Thymelaeales have a similar origin. Our results

show eight vascular bundles running along the length of

the hypanthium and separating into sepal and stamen

bundles, each sepal lobe eventually with three vascular

bundles (Figure 8 A, B). We regard the floral envelope as

a hypanthium (fused calyx and androecium), differen-

tiating into four petaloid sepals and a diplostemonous

androecium, arising from the hypanthium rim at the

separation of the sepals. A very short receptacle (Heinig

1951), which does not contribute to the hypanthium,

indicates a reduction in tissue and a possible advanced

state. This is indeed the case in Passerina.

A study of petaloid scales in Thymelaeaceae has

added further evidence to the interpretation of the floral

envelope. These scales have been regarded by various

authors as perigynous scales or glands, perigynous nec-

taries, petals or staminodes, petaloid scales and petaloid

appendages (Tables 1; 2). Heinig (1951) is convinced

that the morphology and vascularization of the petaloid

scales resemble that of stipules, an opinion shared by

Rao & Dahlgren ( 1969) on the floral anatomy and rela-

tionships of the Oliniaceae. In their floral description of

Olinia, Dahlgren & Van Wyk (1988) consider the

petaloid scales as true petals. Heinig (1951) concludes

that the Thymelaeaceae is apetalous. In Passerina there

are no petaloid scales or corolla (Tables 1; 2). Our

results have shown the separation of antipetalous as well

as antisepalous stamens, but petaloid scales or even ves-

tiges of them were not observed. We therefore regard

Passerina as truly apetalous. The complete reduction of

the corolla or the absence of petaloid scales can be

regarded as an advanced state in the Thymelaeaceae; it

could also be interpreted as part of the anemophilous

syndrome displayed by the genus. Based on both the

pattern of the vascular tissue and the absence of petaloid

scales, we consider the floral envelope in Passerina as a

hypanthium consisting of the fused calyx and androecium,

differentiating into four sepals and the diplostemonous

stamens.

TABLE 8. — Specialized characters of the floral envelope in Passerina, correlated with leaf anatomy

228

Bothalia 31,2 (2001)

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Bothalia 31,2 (2001)

229

FIGURE 8. — LM photographs of P. ericoides. Bredenkamp 956. with fleshy fruit, and P. vulgaris. Bredenkauip 951, with dry fruit, showing

structure of androecium. A-D. serial t/s of sepals and two anther whorls. A-C, P. ericoides: A. each of fused commissural sepal and

antipetalous stamen bundles splitting into two sepal laterals and one antlpetalous stamen bundle; B, fused sepal midrib and antisepa-

lous stamen bundle, splitting into sepal midrib and antisepalous stamen bundle; C. final stage in separation of antisepalous stamen bun-

dles. D. P. vulgaris, t/s through both anther whorls. Abbreviations as in Figure 5. pst. antipetalous stamen bundle; s, sepal midrib bun-

dle; si. sepal lateral bundle; sm. stigma; sp, sepal; sst. antisepalous stamen bundle. Scale bars; A-D, 100 pm.

Fragmentation of hypanthinm

The fruit in Passerina is enveloped by a persistent,

loosely attached hypanthium. Bentham & Hooker (1880),

Gilg (1891, 1894), Domke (1934) and Melchior (1964)

mention that the hypanthium is articulated above the ovary.

We found no definite articulation point in the hypanthium

neck (narrowed part of hypanthium between apex of ovary

and sepals) in Passerina, possibly because of the absence

of receptacle tissue and the appendicular namre of the

hypanthium. The base of the neck fragmented as a result of

desiccation and torsification of cells (Figure 2E). In some

species, fragmentation of the hypanthium takes place over

the widest circumference of the fruit, shedding the frag-

mented distal part of the hypanthium, sepals and androeci-

um (Figure ID). A strong association was found between

flowers with a short hypanthium neck and fragmentation of

the hypanthium over the widest part of the fruit, and also

between flowers with a long hypanthium neck and frag-

mentation at the base of the neck (Table 8).

A plane of circumscission, dividing the floral tube into

a basal and upper portion, is clearly illustrated in Gnidia

and Struthiola (Peterson 1978) and Lachnaea (Beyers

1992; Beyers & Van der Walt 1995). We hypothesize that

the plane of circumscission possibly indicates a differ-

ence in tissue composition between the basal and upper

portions of the hypanthium and that this articulation can

be of morphological importance in the Thymelaeaceae.

The basal portion of the floral tube below the plane of cir-

cumscission possibly indicates the inclusion of receptacle

tissue in the hypanthium, whereas the upper portion con-

sists of calyx and androecium tissue only (accepting the

apetalous state). An alternative interpretation, offered by

one of the referees of this paper, regards the vasculature

as a prerequisite to decide whether one is dealing with a

hypanthium (appendicular in origin) or a receptacle (axial

in origin). The significance of an articulation indicating a

distinction between parts of the hypanthium of different

derivation, should be further investigated.

Androecium

In his de.scription of Thymelaeaceae, Peterson (1978)

describes the stamens as twice as many or as many as the

sepals (rarely reduced to two or one), in one or two whorls,

the outer whorl antisepalous. In the subfamily Aqui-

larioideae (Heinig 1951), stamens are of approximately

230

Bothalia 31,2 (2001)

FIGURE 9. — LM photographs of P. ericoides, Bredenkamp 956 and P. vulgaris, Bredenkamp 951, showing structure of anther loculae in t/s: A,

P. vulgaris, 4-locular anther; B, P. ericoides, cell wall thickenings of endothecium rib-like; C, P. vulgaris, rupturing of partition between

loculi; D, P. ericoides, rupturing of outer walls of thecae, extrorse. e, epidermis; en, endothecium; po, pollen. Scale bars: B, C, 10 pm;

A, D. 100 pm.

equal length and the traces supplying them separate from

the sepal traces, practically in a single whorl. The

Thymelaeoideae is characterized by stamens arising as

two distinct whorls at two distinct levels in the floral tube.

The dimorphic diplostemonous nature of stamens in

Passerimi (Thymelaeoideae), in which the antipetalous

stamens are shorter than the antisepalous ones, has been

confinried by our observations (Figure 8A-C). According

to Heinig (1951) evolution within the androecium has

been from polystemony to diplostemony to dimorphic

diplostemony, indicating the advanced state of the androe-

cium in Passerina. Peterson (1978) describes the anthers

as usually introrse, rarely extrorse. Species of Passerina

have large, exserted, extrorse anthers, clearly an adapta-

tion to the anemophilous syndrome of the genus.

Separation of fused sepal and stamen traces

Heinig ( 1951 ) is of the opinion that a foreshortening

of the floral axis has resulted in a fusion of the calyx and

androecium and that progressive stages of adnation can

be ob.served in the family. In all species the antipetalous

stamen traces are fused to the commisural sepal traces

and the antisepalous stamen traces to the sepal midrib

traces at their point of origin from the stele. In the

Aquilarioideae, in Octolepis dinklagei, they become

separated in the receptacle or, in other cases, low down

in the floral tube. Except for Gnidia splendens (- Lasio-

siphon splendens), in which the separation of the

antipetalous and anti.sepalous stamen traces takes place

in the receptacle, Thymelaeoideae is characterized by

separation higher up in the floral tube, e.g. at the top of

the ovary in Dirca occidentalis and below the origin of

the sepals in Gnidia suhidata (Heinig 1951). In Passe-

rina the separation of the antipetalous stamen traces

takes place at the origin of the sepals (Figure 8 A, B) and

separation of the antisepalous traces slightly higher up

(Figure 8C), indicating what appears to be a phylogeneti-

cally advanced tendency.

Pollen

Pollen grains of most members of Thymelaeaceae are

monads, spheroid and pantoporate, characterized by a

typical croton pattern, comprising rings of more or less

trihedral sexine units mounted on an underlying reticulum

of circular muri (Bredenkamp & Van Wyk 1996). In

Passerina, the basal reticulum, as in the typical croton

pattern, is no longer discernible as it is replaced by a sec-

ondary reticulum derived from fused sexine. The supra-

tectal subunits are fused completely to form a continuous

reticulum which replaces the underlying reticulum. The

reticulum in Passerina is therefore secondary in origin

and considered phylogenetically advanced. This modifi-

cation of the crotonoid pattern is probably also of func-

tional significance as pollen in Passerina is adapted to

anemophily.

Disc

In Aquilarioideae a hypogynous disc is generally

absent, but is almost always present in Thymelaeoideae

Bothalia 31,2 (2001)

231

FIGURE 10. — LM photographs of P. ericoides. Bredenkamp 956. with fleshy fruit, and P. vulgaris, Bredenkamp 951. with dry fruit, showing

structure of young fruit and developing seed in Us. A, B, P. ericoides: A. pericarp fleshy, atrophy of outer integument; B, higher mag-

nification. outer epidermis of inner integument palisade-like, mesophyll, inner epidermis of inner integument tanniniferous. C. D,

P. vulgaris: C. palisade-like exotegmen discemable in tegmen. D. tegmen black and lignified, crassinucellate. endosperm initially

nuclear. Abbreviations as in Figure 5. ens. endosperm; es. embryo sac; ieii. inner epidermis of inner integument; m, mesophyll; oeii,

outer epidermis of inner integument; pc. pericarp; te, tegmen. Scale bars; A-D. 100 pm.

(Heinig 1951; Peterson 1978). Possibly because of a

reduction of tissue, no disc was observed in Passerina, a

state confirmed by Bunniger (1972).

Ovary

The ovary in Passerina was described as unilocular

up to the time of Domke (1934) (Table 1 ). The pseudo-

monomerous state is mentioned by Eckardt (1937),

Melchior (1964), Dahlgren & Thorne (1984) and

Cronquist (1988) (Table 2), while most authors agree

that ovules are solitary and pendulous. According to

Takhtajan (1997) the ovary in Thymelaeoideae consists

of two carpels, it is monolocular and the ovule is solitary.

Heinig (1951) is convinced that within the carpellary

whorl a reduction series may be observed, ranging from

four- or more-carpellate members in the Aquilarioideae

to the two-carpellate members of the Thymelaeoideae in

which one carpel is markedly contracted, thus a

pseudomonomerous ovary. According to Heinig (1951)

the ovules have been reduced to one per locule in the

entire family. In Passerina, at the base of the ovary, the

dorsal and median carpellary bundles initially separate

from the central stele (Figure 5C, D). After differentia-

tion has taken place, the dorsal, median and commissur-

al carpellary bundles (Figure 5E) can be distinguished.

Because of a redistribution of carpellary bundles, the sin-

gle dorsal carpellary bundle is arranged at the opposite

side of the horseshoe-shaped median and commissural

carpellary bundles (Figure 5G). In Dirca palustris.

232

Bothalia 31,2 (2001)

Heinig (1951) has illustrated the authenticity of the

pseudomonomerous ovary by the presence of two dorsal

carpellary bundles, one in the fertile carpel and one in the

second, reduced, sterile carpel positioned between two

groups of commissural carpellary bundles. Bunniger

(1972) showed the presence of two carpels in the flower

primordia of P. filiformis. In the present study, which

included the young bud stage of P. vulgaris (Figure 5F),

no indication of a second carpel or a second dorsal

carpellary bundle was found, possibly because of reduc-

tion and fusion processes, which had already taken place

in the formation of the young ovary, consequently we

consider the ovary of Passerina as pseudomonomerous.

Ovule type and position

Domke (1934) describes the ovule in Thymelaeaceae

as pendulous, anatropous, with a ventral funiculus, ex-

ceptionally hemi-anatropous or orthotropous, indicating

a phylogenetic tendency. Our study clearly indicates a

pendulous, anatropous ovule in Passerina (Figure 6C,

D). The funiculus is ventral and has been sectioned from

the base of the ovule (Figures 5E-H; 6A) to the point of

attachment with the placenta (Figure 6C, D). Close to the

embryo sac the micropyle is a trilete opening formed by

the inner integument (Figures 6C, D; 7A), facing

upward. Towards the micropyle, the outer and inner

integuments become horseshoe-shaped (in t/s), resulting

in the micropyle being directed towards the elongated

obturator cells, located at the base of the style (Figure

7B). Based on these results, we agree with Dahlgren

(1975b), who regards the ovule as pendulous and

epitropous.

Obturator

Davis (1966) defines an obturator as any structure asso-

ciated with directing the growing pollen tube towards the

micropyle, but elongated cells extending from the base of

the style to the micropyle are considered exclusive to the

Thymelaeaceae. In Passerina such elongated obturator

cells can be clearly seen at the level of the placenta, at the

departure of the funiculus, touching on the inner integu-

ment (Figures 6C; 7A) and finally these cells extend from

the base of the style entering the micropyle (Figure 7B).

Fruit

Most authors (Tables 1; 2) agree that the fruit in

Thymelaeaceae is indehiscent. In Passerina, Wikstrom

(1818), Meisner (1857) and Endlicher (1837, 1847) con-

sider the fruit as a nut or a nutlet. Domke (1934) concludes

that the fruit of Dais, Gnidia, Lachnaea (= Cryptadenia)

and many taxa of Passerina can be defined as an achene,

and that of P. ericoides as a bemy. Dahlgren (1975b) con-

siders the fruit of Thymelaeaceae as a nut or drupe, Peter-

son (1978) regards it as a berry, a nut, a drupe or a loculi-

cidal capsule and Takhtajan (1997) describes them as nut-

like, baccate or dmpaceous. A relevant family character,

that the outer integument of the ovule disintegrates and the

inner integument becomes palisade-like and hardens to

form a seed coat or tegmen (Figure lOA-D), is illustrated

by the present study. Structurally the dry, membranous fruit

in Passerina conforms to an achene (Spuyt 1994) and the

reduction in pericarp tissue can probably be considered as

a specialized adaptation. A reduction in tissue from a drupe

to a membranous 1 -seeded berry or an achene can be illus-

trated in Thymelaeaceae and therefore the fruit in

Passerina could be considered advanced within the family.

We agree with Domke ( 1934) that P. ericoides (Figures IF;

lOA) and P. rigida are characterized by a fleshy 1 -seeded

berry, while all the other species have an achene (Figure

2F; Table 7). The achene remains enveloped in the rem-

nants of the papyraceous hypanthium, nestled adaxially in

the tomentum of the concave, persistent cymbiform bracts.

Seed

The existing confusion concerning the state of the

tegmen in Passerina is a reflection of the total lack of

information of this aspect in descriptions of the group by

various authors (Tables 1; 2). Meisner (1857) describes a

crustaceous pericarp and Domke (1934) a black, crusta-

ceous testa. The ovule in Passerina is bitegmic (Figures

6; 7; 10). Corner (1976) refers to outer integument (oi)

and inner integument (ii), the product of the outer integu-

ment becoming the testa and that of the inner integu-

ment, the tegmen. In Passerina the outer integument dis-

integrates, whereas the inner integument remains (Figure

lOA), its outer epidermis becoming palisade-like (Figure

lOA, B). Hence, the seeds of Passerina are exotegmic

with a palisade, a state common to the family (Corner

1976). In Passerina the tegmen is black and lignified,

and in t/s, still portraying its origin from the palisade-like

epidermis (Figure IOC, D).

Floral and fruit morphology at species level (Tables 1; 8)

In the present study, specific results in leaf anatomy

indicating the arrangement of taxa in Passerina

(Bredenkamp & Van Wyk 2001), are associated with flo-

ral morphological structures as well as fruit and seed

types. In Table 7, leaf structural types A and especially

B3 are associated with four species that have smaller,

yellow, membraneous flowers (up to 5.3 mm long). Leaf

structural types B4, B5, B6, C and D are associated with

most species having larger, yellow-pink, papyraceous

flowers (up to 8.4 mm long). The same tendency in the

length of the neck, and the size of the inner sepals is

shown in Table 8.

Fruit

Eleshy fruit in P. ericoides (Eigure ID, F) and P. rigida

is possibly correlated with the moist maritime climate of

the coastal habitat of these species; it is possibly also an

adaptation to bird dispersal. Both species occur in the

Western Cape, and the range of P. rigida extends along

the coast to St Lucia. P. ericoides has red berries and

P. rigida has yellow benies. All other species are char-

acterized by achenes and are adapted to drier habitats,

from mountainous areas along the Great Escarpment to

the arid Karoo. In Passerina, each achene is enveloped

by papyraceous remnants of the fragmented hypanthium

and enclosed within the tomentum of an enlarged bract

(Eigure 2D, E), which often takes on a rounded shape

and turns yellow, red or brownish.

The fruits of Passerina clearly illustrate the phenome-

non of transfer of function from protection to dispersal

Bothalia 31,2 (2001)

233

(Stebbins 1974). In P. ericoides and P. rigida, with fleshy

fruit, the pericarp has the double function of protecting

the ovules during early stages of development and disper-

sal. The mature fruit enlarges beyond the bracts and is

protected by the coriaceous pericarp, while the patent

bract does not have a protective function (Figure ID). The

pericarp of the fruit is yellow or red when it is ready for

dispersal, probably by birds or small mammals (Richards

1986), and the dispersal unit is the berry. In all other

species which are characterized by achenes, the protec-

tion of the ovule is apparently transferred from the peri-

carp to the persistent bract. The bract enlarges around the

achene, protecting it in the woolly tomentum of the con-

cave adaxial surface (Figure 2D). The mature fruit is

often still enveloped by the reddish, papyraceous rem-

nants of the hypanthium. Both P. montana and P. glom-

erata are characterized by subcapitulate inflorescences,

with proliferating growth more common in the latter

species. In P. glomerata (growing in the arid Karoo) the

accompanying bract turns yellow and becomes more

patent when the fruit is mature; the achene is shed after

abscission. The yellow colour is associated with senes-

cence of the bracts and these structures are eventually

shed, leaving conspicuous bract scars on the remaining,

often terminal, woolly inflorescence axis. The unit of dis-

persal, in this case, is the achene, assisted by the patent

senescing bract. The achene falls to the ground where it

could either be dispersed by ants or small mammals or

germinate under favourable conditions. In P. montana

(growing along the Great Escarpment), the margin of the

fruiting bract turns red and it becomes more patent,

exposing the achene enveloped in the beak-like, reddish,

papyraceous hypanthium, which fragments at the neck

base. Perhaps birds, attracted by the red colour (Richards

1986), feed on the exposed achenes. Leafless, terminal,

woolly branchlets, with terminal scars are a conspicuous

feature of the plant after fruiting and it therefore also

seems possible that the subterminal capitulum with sever-

al achenes is broken off as a unit. In this case the disper-

sal unit could be the achene or the achene accompanied

by the bract or perhaps even the entire subcapitulum.

There is a need for further observations on seed dispersal

in the field to test some of these suggestions.

Filament length

For a comparison of filament length, it is easier to

measure the antisepalous filaments as they are ± twice

the length of the antipetalous ones. Both P. sp. nov. 1 and

P. sp. nov. 2 have short antisepalous filaments of ± 1.2

mm, corresponding to their small flowers. P. palndosa,

P. fdiformis and P. falcifolia have exserted stamens

because of their long (2. 1-2.4 mm) filaments.

Anthers

Conspicuous differences in anther size have been

noticed among the species; this has also been reported by

Thoday (1924). In relation to flower size, most species

have large exserted anthers between 0.7(-0.9) x

0.3(-0.7) mm, possibly an anemophilous adaptation. P.

ericoides has ± elliptic anthers (Figure 1 ), whereas those

of P. drakensbergensis are narrowly oblong, 0.9 x 0.3

mm. The largest anthers of 1.0 x 0.5 mm are found in P.

obtusifolia and P. falcifolia.

Floral envelope

The hypanthium and sepals in P. ericoides are char-

acterized by their coriaceous (almost fleshy) appearance

and dull green to pinkish colour. The floral envelope in

P. rigida, P. paleacea, P. sp. nov. 1 and P. sp. nov. 2 is

pale yellowish and quite membranous. P. pendida is dis-

tinguished by a pinkish floral envelope, with a membra-

nous texture. In all the other species the floral envelope

is yellow-pink at anthesis, with a papyraceous texture.

After pollination these flowers turn red and the hypan-

thium and sepals become thinly papyraceous and dry.

For practical purposes the total length of the floral

envelope indicates flower size, and its taxonomic impor-

tance is clearly illustrated by the general increase in size

from species 1-20 (Table 7). P. rigida, P. paleacea and

P. sp. nov. 2 are characterized by small flowers, the

length of the floral envelope 4.0^. 6 mm. In most other

species it ranges from 5. 3-7. 3 mm long. P. rubra and P.

falcifolia are characterized by large flowers, the floral

envelope being 8.4 mm long.

Specialized characters (Table 8)

a. Fragmentation of hypanthium after fruiting

During the fruiting phase, the persistent hypanthium

fragments over the broadest part of the ovary, in eight of

the 20 species. Except in P. obtusifolia, the hypanthium

of all these species is characterized by a short neck of

O. 3-1.0 mm. Anatomically their leaves also correlate

with leaf structural type B (Bredenkamp & Van Wyk

2001 ). When fragmentation occurs at the neck base, the

hypanthium usually has a long neck length of mostly

1. 3-3.0 mm, and such species are associated with leaf

structural types A, B, C and D.

b. Indumentum of hypanthium

In certain species of the genus Lachnaea (Beyers 1992;

Beyers & Van der Walt 1995), different trichome types are

found below and above the articulation plane in the hypan-

thium. This state could possibly also be present in other

genera of the Thymelaeaceae. In Passerina the trichome

type remains constant over the length of the hypanthium,

possibly because there is no articulation plane in the

hypanthium. As Passerina is distributed over a wide range

of habitats, the density of the indumentum has been impor-

tant in the distinction of certain species (Table 8). P. eri-

coides is distinguished by the strigose indumentum over

the length of the hypanthium, whereas the indumentum of

the neck is strigose in P. palndosa. A completely glabrous

hypanthium is characteristic of P. paleacea. In 12 of the

species, the hypanthium surrounding the ovary is

glabrous, scantily tomentose or tomentose at the apex,

with the neck scantily tomentose or tomentose. In P. sp.

nov. 1 , P. sp. nov. 2 and P. filiformis subsp. filifonnis the

hypanthium is tomentose over its entire length. In

P. galpinii the whole of the hypanthium is pubescent,

whereas in P. rubra only the neck is pubescent and the rest

of the hypanthium is glabrous.

A strong correlation was found between the indumen-

tum of the lower hypanthium and of the bract. There is a

234

Bothalia31,2 (2001)

tendency for species characterized by a glabrous hypan-

thium base (Table 8) to have a protective bract with a

very hairy adaxial surface (Table 5), whereas species

with a hairy hypanthium base have a less hairy to almost

glabrous adaxial bract surface. When the hypanthium

neck is not covered by a bract, it tends to be hairy in

varying degrees. Only P. paleacea has a completely

glabrous hypanthium. In this species the entire hypanthi-

um is completely covered by the hairy adaxial surface of

the bract because of the very short hypanthium neck.

This tendency shows that the function of protection of

the ovule is partly transferred from the hypanthium to the

hairy bract (Stebbins 1974).

c. Indumentum of sepals

Thoday (1924) uses the character ‘outer sepals beard-

ed behind the apex’ in his key to distinguish between

species. The abaxial surfaces of the outer and inner lobes

of seven species are setose at the apex only and the adax-

ial surfaces range from glabrous to variously hairy (Table

8). In f. comosa abaxial surfaces of both outer and inner

lobes are tomentose and adaxial surfaces are glabrous,

P. drakensbergensis is similar except for the tomentose

adaxial surface of the inner lobes. All sepals are com-

pletely glabrous in P. paleacea. The indumentum of the

sepals varies infraspecifically and should, however, be

used with discretion to distinguish between groups of

species.

d. Size and shape of sepals

The size of the inner sepals is of taxonomic impor-

tance (Table 8). P. sp. nov. 1, P. ericoides, P. rigida, P.

paleacea and P. sp. nov. 2 are characterized by shorter

and broader inner sepals, varying from widely ovate,

obovate, widely obovate to subrotund. All other species

have longer, narrower inner sepals, the shape varying,

with one exception, from narrowly oblong, oblong,

oblong-elliptic, elliptic, to obovate. P. drakensbergensis

is distinguished by lanceolate sepals arranged in the

shape of a cross.

Less important taxonomic characters

Ovary size (Table 7) has been considered less impor-

tant, as it is difficult to measure all ovaries at the same

developmental phase. In Passerina the size of the ovary

increases markedly after anthesis and the enlarged ovary,

enveloped by the persistent hypanthium, can already be

observed in older flowers, rapidly followed by matura-

tion of the fruit.

Taxonomic relationships

Up to 1930, priority was given to the definition of

subordinal or subfamilial taxa in Thymelaeaceae, based

mainly on floral morphology (Table I ). Applying both

morphological and anatomical evidence, Domke (1934)

proposed a satisfactory subfamilial classification system

and envisages a phylogenetic relationship between the

Thymelaeaceae, Malvaceae and Euphorbiaceae. Modern

techniques have enabled taxonomists to find relation-

ships between families and to arrange them into higher

hierarchies, with ranks such as superorders or subclass-

es. Thus Dahlgren (1980) placed the Thymelaeaceae in

the superorder Malviflorae (= Dilleniiflorae), Cronquist

(1981, 1988) placed it in the subclass Rosidae, Thorne

(1992a, 1992b) in the superorder Malvanae and

Takhtajan (1997) in the subclass Dilleniidae (Table 2).

Heinig (1951) discussed the relationships of the

Thymelaeaceae with the Myrtales, Saxifragaceae,

Lythraceae, Gonystylaceae and Malvales on the basis of

floral morphology and concluded that a polyphyletic ori-

gin of the Thymelaeaceae could be sought in both the

Flacourtiaceae and Tiliaceae.

Cronquist (1968, 1981, 1988) considered the

Thymelaeaceae as completely at home in the Myrtales

(Rosidae) on account of the strongly perigynous

polypetalous to apetalous flowers, internal phloem, ves-

tured pits and obturator. However, he admited a possible

relationship with other families, based on the

pseudomonomerous ovary and crotonoid pollen. Dahl-

gren (1975 a, b) placed the superorder Thymelaeanae

between the Dillenianae (Dilleniales, Cistales, Malvales,

Urticales, Euphorbiales) and the Myrtanae. Within the

superorder Malviflorae, Dahlgren (1980) recognized a

close affinity between the Malvales and Euphorbiales,

and indicated a strong relationship with the Urticales and

the Thymelaeales, but a phylogenetic affinity between

the Malviflorae and Myrtiflorae was not supported. The

inclusion of the Thymelaeaceae in the Myrtales was

reviewed by Dahlgren & Thome (1984). They argued

that anatomically, most members of the family possess

Myrtalean characters. On the other hand, embryological,

chemical and palynological evidence strongly indicates

an affinity with the Malviflorae. Thorne (1992a) accept-

ed the superorder Malvanae, but included the Thyme-

laeaceae in the order Euphorbiales. Takhtajan (1969)

considered the Thymelaeales to have a common origin

with the Euphorbiales and Malvales, all arising from a

Elacourtiaceae-type ancestor, and Takhtajan (1997)

placed the Thymelaeales in the superorder Euphor-

bianae, with the Gonystylaceae as the only other family

in the order.

Palynological evidence indicates that the very distinc-

tive pollen in Thymelaeaceae is totally different from

that of any Myrtales and similar to that of most

Euphorbiaceae. Archangelsky (1971) concluded that

both the Euphorbiales and the Thymelaeales belong to

the subclass Dilleniidae and originated from ancestral

lines of the Dilleniales -> Violales Malvales. In a

paper on palynology of Passerina, Bredenkamp & Van

Wyk (1996) supported Dahlgren (1980) in placing the

Thymelaeales in the Malviflorae (= Dilleniiflorae).

In Passerina, the structure of the integuments sur-

rounding the ovule provides taxonomically useful

embryological evidence. We have shown the disintigra-

tion of the outer integument and the differentiation of the

inner integument into a palisade-like outer epidermis, a

mesophyl layer and an inner epidermis (Figure lOA, B).

It is also clear that the mechanical part of the seed coat is

derived from the palisade-like outer epidermis, hence it

is an exotegmen (Figure IOC, D). Corner (1976) distin-

guished the Euphorbiales-Malvales-Thymelaeales-

Tiliales (Malvalean complex) on the basis of the exo-

Bothalia 31.2 (2001)

235

tegmic palisade. He dismissed the derivation of the

Malvalean complex from the Dilleniales (endotestal

seeds) or Violales and looked towards the Myristicaceae

(Magnoliales-Ranales) for the origin of the Malvalean

seed.

Recent evidence from molecular phylogeny (Table 3)

should be interpreted in context with the evidence from

other botanical fields. The primary focus of The

Angiosperm Phylogeny Group (APG 1998) is on orders,

with a secondary focus on families of flowering plants.

Above the ordinal level, ranks are defined as subgroups,

clades or supraordinal subgroups. Magallon et al. (1999)

attempted to compare these groups to the existing systems

mentioned in the preceding paragraphs. Both APG ( 1998)

and Magallon et al. (1999) recognized the Eudicots, a

group characterized by tricolpate pollen, as well as the

supraordinal group Core Eudicots, supported by pen-

tamerous and isomerous flowers. Thymelaeaceae is desig-

nated to the Rosid clade by most authors. APG (1998)

placed the Thymelaeaceae in the subgroup Eurosids II and

order Malvales, whereas all the other authors placed it in

the Expanded Malvales. The Malvalean relationship of the

Thymelaeaceae seems to be strongly supported by molec-

ular phylogeny, as well as floral morphology, anatomy,

embryology and palynology.

The Euphorbiales-Malvales-Thymelaeales relationship

indicated by embryology and palynology is, however,

not supported by molecular data. APG (1998) placed the

Euphorbiaceae in the order Malpighiales in the subgroup

Eurosids I and Magallon et al. (1999) placed it in the

Core Rosid Clade. Cronquist (1968, 1981, 1988) was

convinced that if the Thymelaeaceae is not placed in the

Myrtales, it would stand next to it. Conti et al. (1996),

APG (1998), and Alverson et al. (1999) all regarded the

Myrtales as a sister group of the Malvales or the Expand-

ed Malvales to which the Thymelaeaceae is designated.

Speculations on phylogeny

Within Thymelaeaceae, both Domke (1934) and

Heinig (1951) agreed that the subfamily Thymelaeoideae

is phylogenetically more advanced than the Aquilari-

oideae. On the basis of the advanced pollen, Bredenkamp

& Van Wyk ( 1996) raised the subtribe Passerininae to the

tribe Passerineae, a decision supported by the present

study. Although many of the following advanced charac-

ter states are present in other genera of the Thyme-

laeoideae, these advanced character states are all found

together in Passerina: receptacle reduced to a ± lenticu-

lar structure; departure of the fused sepal and stamen

bundles before carpellary bundles; hypogynous floral

arrangement; petal-like floral envelope comprising a

hypanthium (fused calyx and androecium), differentiat-

ing into four sepals and a diplostemonous androecium;

separation of stamen bundles high up in hypanthium, at

formation of sepals; exserted, extrorse anthers; anemo-

philous habit; secondary reticulum of pollen; complete

absence of petals or petaloid scales; asymmetric devel-

opment of the style; superior, pseudomonomerous, uni-

locular ovary; asymmetric attachment of ovule at top of

ovary; ventrally epitropous ovule; distinctive obturator;

bitegmic ovule with exotegmic palisade; fruit a 1 -seeded

berry or an achene; seed with lignified, black exotegmen.

Considering all the characters mentioned, Passerina is

considered highly advanced in relation to other genera in

Thymelaeoideae.

Systematic value

Family level

The exotegmic palisade and the distinctive obturator

are regarded as family characters. They form the basis of

Comer’s (1976) Euphorbiales-Malvales-Thymelaeales-

Tiliales complex.

Subfamily level

The Thymelaeoideae is distinguished on the basis of

the calyx tube (hypanthium in the present study),

diplostemous androecium and pseudomonomerous ovary

(Domke 1934; Heinig 1951).

Tribal level

On the basis of the secondary reticulum, unique to the

pollen of Passerina, Bredenkamp & Van Wyk (1996)

raised the subtribe Passerininae to the tribe Passerineae.

Genus level

The present study indicates the exserted, extrorse

anthers and the anemophilous habit as unique to Passe-

rina.

Species level

Characters useful at species level are summarized in

Tables 7 and 8.

CONCLUSIONS

The evidence on floral morphology not only con-

firmed the identity of 20 species and four subspecies, but

also proved significant in the taxonomy of the genus. The

status of the following taxa is confirmed by the present

floral morphological study: P. burcliellii Thoday,

P. comosa C.H.Wright, P. drakensbergensis Hilliard &

B.L.Burtt, P. ericoides L., P. falcifolia C.H.Wright, P.fd-

iformis L. subsp. fdiformis, P. fdiformis L. subsp. nov.,

P. galpinii C.H.Wright, P. glomerata Thunb. subsp.

glomerata, P. glomerata Thunb. subsp. nov., P. montana

Thoday, P. obtusifolia Thoday, P. paleacea Wikstr.,

P. paludosa Thoday, P. pendula Eckl. & Zeyh., P. rigida

Wikstr., P. rubra C.H.Wright, P. vulgaris, P. sp. nov. 1,

P. sp. nov. 2, P. sp. nov. 3 and P. sp. nov. 4.

For almost three centuries evidence from floral mor-

phology has been basic to plant taxonomy and applied at

all hierachical levels. Our research on the flowers in

Passerina has produced new morphological and anatomi-

cal evidence, especially as Heinig’s classical study of floral

morphology in Thymelaeaceae (1951) did not include

Passerina. The present study has succeeded in resolving

the floral morphology in Passerina, as many mistakes

have been perpetuated by previous authors. We conclude

that the flower in Passerina is a phylogenetically

advanced structure and consider the genus advanced

236

Bothalia 31,2 (2001)

within the Thymelaeoideae. Possible taxonomic relation-

ships of the Thymelaeaceae with the Malvales are strong-

ly supported by the presentation of floral morphological,

anatomical, embryological and palynological information

to which this study has also contributed. Finally, all the

above-mentioned evidence serves to form a firm taxo-

nomic basis for future comparative studies, especially in

the expanding molecular field.

ACKNOWLEDGEMENTS

The authors wish to thank Dr E. Steyn for assistance

with the LM, Ms G. Condy for the line drawings and Ms

A. Romanowski for the developing and printing of the

photographs.

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Bothalia 31,2: 237-240 (2001 )

Miscellaneous notes

ALOACEAE

ARE OVULES AND SEEDS IN LOMATOPHYLLUM WIELD. (ALOE SECT. LOMATOPHYLLUM SENSE AUCT.) ANATROPOUS AND

EXARILLATE?

INTRODUCTION

The seeds of all aloes are enveloped by an extra tissue

that forms an additional covering to the seed coat

(Dahlgren et al. 1985). Known as an aril and sometimes

referred to as a third integument (Joshi 1937; Danilova et

al. 1995), this tissue arises early in the development of

the aloe ovule as a cup-shaped outgrowth of the placen-

ta (Schnarf & Wunderlich 1939) or base of the funicle

(McNaughton & Robertson 1974). The presence of the

broad, impeding structure developing so early and close

to the base of the ovule has a profound influence on the

ultimate shape of the ovule: instead of turning through

180° to reach a completely upside-down (anatropous)

position, the ovule primordium only completes a 90° turn

and becomes hemitropous (Steyn & Smith 1998: fig.

2A). Morphologically, mature ovules of aloes therefore

show two very distinctive characters, namely hemitropy

and broad aril primordia (Steyn & Smith 1998).

In fertilized ovules the aril immediately starts grow-

ing around the developing seed to form a succulent mem-

brane which completely covers the hemitropous seed

body. The succulent tissue of the aril may have adaptive

value by acting as a handy, nearby reservoir for supply-

ing water to the vital, inner ovular tissues in times of

water stress. Having grown around the developing seed,

the aril possibly keeps the tender, immature seed in a

moist atmosphere. As seed development progresses, the

aril tissue extends laterally to fill the spaces among the

seeds in the locules of the capsule (E.M.A. Steyn pers.

obs.). In this way, the wings of aloe seeds are formed.

The shape and size of the wings may be moulded by the

available space in the developing locule (Werker 1997).

In some aloes, for instance Aloe asperifolia A. Berger, A.

claviflora Burch., A. falcata Baker and A. variegata L.

the wings can be large, frilly structures, while in others

the wings are less elaborate or sometimes absent

(Kamstra 1968; Reynolds 1982). At seed maturity the

succulent aril tissue dries out and becomes papery when

the cartilaginous capsule splits open. The winged seeds

are then released to be dispersed by wind.

It has been reasoned that character states may be lost

during evolution of a taxon if they become functionally

superfluous, e.g. an aril will easily become rudimentary

if the fruit becomes indehiscent (Dahlgren 1991). The

question therefore arises whether this has happened in

about 20 species of aloe look-alikes occurring on

Madagascar and the neighbouring islands of Aldabra,

Reunion, Mauritius and Rodrigues. Unlike all other

members of the Aloaceae, these species do not bear cap-

sular fruit, but produce fleshy-fruited, colourful berries

containing wingless black seed (Rowley 1996). Although

some of these ‘berried aloes’ (Rowley 1996) occur in dry

mountainous situations or on dry, hilly banks (Baker

1877), they generally favour a warm, humid forest cli-

mate and rely on animals (birds?) for the dispersal of

their seed (Rauh 1993, 1998). If the wingless seed in the

berries of Lomatophyllum have lost their arils, one would

then expect reports of anatropy in the berried aloes —

suppression of aril development would allow curvature

of the ovule to the anatropous position.

During a preliminary evaluation of literature dealing

with ovule morphology in Asphodelaceae (Steyn &

Smith 1997) we noticed that Schill (1973) described the

ovules of Lomatophyllum Willd. as anatropous, although

the presence (or absence) of an aril was not mentioned in

this 1973 report. However, subsequent research (Steyn &

Smith 1998) on ovule structure in the genera of the

Aloaceae did not include Lomatophyllum ovules, since

suitable material was not available at the time. In addi-

tion to rectifying this omission, the present paper also

includes a report on seed development and seed coat

structure of L. purpureum (Lam.) TDurand & Schinz.

MATERIAL AND METHODS

Open flowers and berries were collected in the nurse-

ries of the National Botanical Institute, Pretoria from

cultivated plants of Lomatophyllum purpureum (Acces-

sion No. GPTA 3 1155). The plants were propagated from

seed, originally collected by Dr Wendy Strahm on

Mauritius in 1988. For the present study, ovaries, devel-

oping berries and ripe seed were dissected from the flow-

ers or fruit and fixed for 24 h in a 0.1 M cacodylate

buffered solution (pH 7.4) containing 4% formaldehyde

and 2.5% gluteraldehyde. By following the methods of

Feder & O’Brien (1968), ovules and seeds were dehy-

drated in an alcohol series and impregnated with glycol

methacrylate (GMA). The ovules and seed were subse-

quently imbedded in GMA and sectioned in a sagittal

plane at a thickness of 2-3 pm on a Jung RM 2045

microtome. Median sagittal sections were stained with

the periodic acid/Schiff reaction and counterstained with

toluidine blue O (pH 4.4) by using the protocols of

O’Brien & McCully ( 1981 ).

RESULTS

Orientation and structure of ovules

The sessile, trilocular ovary of Lomatophyllum pur-

pureum contains many ovules. Each locule holds several

ovules alternately arranged in two longitudinal rows and

obliquely placed on axile placentae. The ovules are pleu-

rotropous, i.e. the micropyles are turned outwards, facing

the nearest septum (Figure 1 ). In each ovule the micropy-

lar-chalazal (i.e. longitudinal) axis of the ovule is perpen-

dicular to the longitudinal axis of the short funicle (Figure

238

Bothalia 31,2 (2001)

FIGURE 1 . — Mature ovule of Loma-

tophyllwn purpiireum in sa-

gittal view, seen in an oblique

section of ovary: a, aril; c,

central cell nucleus; f, filiform

apparatus of synergid; h, hypo-

stase in chalazal nucellus; i,

inner integument forming

endostome; o, outer integu-

ment. Direction of longitudi-

nal axis of short funicle indi-

cated by arrow. Scale bar: 100

pm.

1 ). Thus, during early development and curvature towards

the anatropous position, the ovules of L purpureum have

only turned through 90 ° to become hemitropous.

At the base of the ovule, the aril forms a conspicuous,

broad and many-layered podium for the virtually sessile,

bitegmic ovule (Figure 1). The outer integument is three-

layered and slightly shorter than the two-layered inner

integument, so that the latter alone forms the micropyle

canal. Around the endostome the inner integument com-

prises more than two layers and here the cells have

increased in size. The embryo sac is relatively large, hav-

ing penetrated into the greater part of the micropylar

nucellus, and only the nucellus epidermis has remained.

The micropylar part of the embryo sac contains an egg

cell and two prominent synergids with conspicuous, stri-

ated and strongly PAS-positive filiform apparatus. A

large central cell nucleus, situated directly above three

ephemeral antipodes, occupies the chalazal part of the

embryo sac. The base of the embryo sac is sunken into

the chalazal nucellar tissue. The nucellus cells surround-

ing the base have developed into a prominent hypostase

consisting of thick-walled, dark-staining cells which pos-

sibly prevented further penetration of the embryo sac

into the chalazal nucellus.

Development and structure of aril and seed coat

After fertilization the aril rapidly expands to enfold

the developing seed (Figure 2A). At this stage, the aril

tissue comprises several layers of thin-walled cells with

living protoplasts (Figure 2B). Except for the notably

large cells of the outer epidermis and scattered idioblasts,

the cells of the aril are small and densely packed (Figure

FIGURE 2. — .Seed development in Loinatophylliim purpureum. A, median sagittal section of young seed; B, structure of developing aril and seed

coat in sagittal section: a. aril; b, enlarging embryo sac; d. outer epidermis of testa; e, epidermis of nucellus; g, inner epidermis of testa; j,

outer epidermis oftegmen; k, inner epidermis of tegmen; n. chalazal nucellus; s, developing testa; t, developing tegmen; u, funicle; w, fruit

wall. Position of micropyle indicated by curved arrow. Scale bars: A, 500 pm: B, 100 pm.

Bothalia 31.2 (2001)

239

FIGURE 3. — Mature seed structure in Lomatophyllum purpureum. A, ripe seed in longitudinal section showing: a, aril: m, large straight embryo

in endosperm; n, some remains of nucellus; u. funicle. Position of micropyle indicated by curved arrow. B, structure of aril and seed coat

in ripe seed: curved arrow, outer epidermis of aril; straight arrow, flattened cell layers of testa and tegmen; g, outer epidermis of testa filled

with phytomelan; p, endosperm. Scale bars: A, 500 pm; B, 100 pm.

2A, B). During enlargement of the embryo sac and the

concurrent increase in circumference of the seed, the aril

possibly keeps pace by marginal growth, i.e. cell divi-

sions in the overlapping tips. Later, when the endosperm

becomes fdled with reserves and seed ripening occurs,

the thin-walled aril tissue collapses. Only the outer epi-

dermis remains as a thin layer of tangentially stretched

cells covering the seed (Figure 3 A, B).

During the enlargement of the growing seed the layers

of both the tegmen (inner integument) and testa (outer

integument) are discernible. The tegmen comprises two

to three layers of thin-walled and closely packed cells,

easily discernible from the adjacent cells of the nucellus

epidermis and testa (Figure 2A, B ). At this stage, the testa

consists of six to eight cell layers. The small, inner epi-

dermal cells have large nuclei and are still in an actively

dividing stage. The outer epidermis cells are noticeably

larger than the cells of the other layers and later become

filled with dark-staining deposits of phytomelan.

Eventually, when seed ripening occurs, the structure of

the outer epidermis cells breaks down, but the black pig-

ment remains as a melanin crust covering the collapsed

cell layers of the testa and tegmen (Figure 3B).

The hemitropous nature of the ovule is retained dur-

ing seed development, although some curvature towards

the ventral side occurs in the distal part of the funicle

(Figure 2A). The ripe seed contains some remains of the

nucellar tissue (perisperm) in the chalazal region. The

linear embryo is straight and occupies more than three-

quarters of the length of the seed (Figure 3A).

DISCUSSION AND CONCLUSIONS

Our results on L. purpureum strongly suggest that

Schill’s (1973) conception of anatropy in the ovules of

Lomatophyllym was erroneous, and that ovule curvature in

Lomatophyllum is arrested in the hemitropous stage of the

ovule primordium by the presence of a broad aril at the base

of the funicle. A re-evaluation of Schill’s data strengthens

our view; his own micrograph (Schill 1973; 277, fig. 3),

depicting a transverse section of the ovary of L citreum

Guillaumin. clearly shows hemitropous ovules above the

aril primordia in the locules. This micrograph was not

intended to illustrate ovule curvature, but to show that the

dorsal walls of the locules in the fmit of Lomatophyllum

remain intact and do not open by median splits like those of

Aloe viquieri H.Perrier (Schill 1973; fig. 4).

During seed formation the growing seed of L. pur-

pureum is eventually enfolded by arillate tissue. Although

elaborate wings do not develop from this tissue as is often

the case in Aloe, the remains of the aril can still be iden-

tified in the ripe, hemitropous seed. Internal ovular struc-

ture in L. purpureum also conforms to that of Aloe and the

other genera of the Aloaceae (Astroloba, Chortolirion,

Gasteria, Haworthia and Poellnitzia) investigated by

Steyn & Smith (1998). Among the characters shared, are

a prominent hypostase, a conspicuous central cell lying

directly above the ephemeral antipodes and synergids

containing striated, well-developed filiform apparatus

that are strongly PAS-positive.

The results obtained in this investigation lend support

for the proposal of Rowley (1996) to subsume Lomato-

phyllum under Aloe, but this may be an oversimplifica-

tion of a rather intricate taxonomy (Rauh 1998). The

genus remains controversial as evidenced by Newton

(1998) who expressed the view that the species in the

genus do not necessarily form a natural infra-generic

group within Aloe.

REFERENCES

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tion of the flowering plants and ferns of those islands: 374.

Reeve, London.

DAHLGREN. G. 1991. Steps toward a natural system of the dicotyle-

dons: embryological characters. Aliso: 107-165.

DAHLGREN, R.M.T., CLIFFORD, H.T. & YEO. P.F. 1985. The fami-

lies of the monocotyledons: structure, evolution and taxonomy:

179-182. Springer Verlag. Berlin.

DANILOVA, M.F., NEMIROVICH-DANCHENKO. E.N.. KOMAR.

G.A. & LODKINA, M.M. 1995. The seed structure of mono-

cotyledons. In RJ. Rudall, RJ. Cribb. D.F. Cutler & C.J. Hum-

240

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phries. Monocotyledons: systematics and evolution'. 461^72,

Royal Botanic Gardens, Kew.

FEDER, N. & O’BRIEN, T.R 1968. Plant microtechnique: some princi-

ples and new methods. American Journal of Botany 55: 123-142.

JOSHl. A.C. 1937. Megasporogenesis in Aloe vera Linn. Journal of the

Indian Botanical Society 15: 297-300.

KAMSTRA. M."W. 1968. Aloe seeds. Aloe 6,2: 8-10, 23-27.

MCNAUGHTON, J.E. & ROBERTSON, B.L. 1974. Megasporogene-

sis and megagametogenesis in Aloe africana Mill. Journal of

South African Botany 40: 75-79.

NEWTON, L.E. 1998. A new species of Aloe (Aloaceae) on Pemba,

with comments on section Lomatophyllum. Cactus and

Succulent Journal 70: 27-31.

O’BRIEN, T.P. & McCULLY, M.E. 1981. The study of plant structure:

principles and selected methods'. 4.19-4.22, 6.81-6.88.

Termacarpi, Melbourne.

RAUH, W. 1993. Lomatophyllum belavenokense Rauh & Gerold spec,

nov., a new species from south-eastern Madagascar. Excelsa 16:

1-6.

RAUH, W. 1998. Three new species of Lomatophyllum and one new

Aloe from Madagascar. Bradleva 16: 92-100.

REYNOLDS, G.W. 1982. The aloes of South Africa, edn 4. Balkema,

Cape Town.

ROWLEY, G.D. 1996. The berried aloes: Aloe section Lomatophyllum.

Excelsa 17: 59-62.

SCHILL, R. 1973. Studien zur systematische Stellung der Gattung

Lomatophyllum Willd. — ein Vergleich mit Aloe Tourn.

(Liliaceae). Beitrdge zur Biologic der Pflanzen 49: 273-289.

SCHNARF, K. & WUNDERLICH, R. 1939. Zur vergleichenden

Embryologie der Liliaceae-Asphodeloideae. Flora 133:

297-327.

STEYN, E.M.A. & SMITH, G.F. 1997. Ovule structure in Trachyandra

saltii (Asphodelaceae). South African Journal of Botany 63:

223-226.

STEYN, E.M.A. & SMITH, G.F. 1998. Ovule orientation, curvature

and internal structure in the Aloaceae. South African Journal of

Botany 64: 192-197.

WERKER, E. 1997. Seed anatomy. Handbuch der Pflanzenanatomie,

special issue 10,3: 283. Bomtraeger, Berlin.

E.M.A. STEYN* and G.F. SMITH*

* National Botanical Institute, Private Bag XI 01, 0001 Pretoria.

MS. received: 2000-11-27.

Bothalia31,2: 241-244 (2001)

OBITUARY

HEINRICH JOHANN WILHELM GIESS (1910-2000)

Willi Giess (Figure 1 ), as he was known to most, was

bom on 21 Febmary 1910 in Frankfurt am Main,

Germany. He was the elder of the two sons of Wilhelm

and Lilly Giess. He completed his schooling at the

‘Adlerflycht Realschule’ in Frankfurt in 1925. The Giess

family then emigrated to South West Africa in 1926

(Kolberg & Strohbach 2000; Gunn & Codd 1981).

Willi Giess started working as a volunteer on farms at

the tender age of 16 until he enrolled for a diploma in

agriculture at the Neudamm Agricultural College near

Windhoek from 1928 to 1929 (Kolberg & Strohbach

2000). He was one of the first three students that were

registered at Neudamm. During his subsequent career he

often re-visited Neudamm as a lecturer to try and impart

his knowledge of local rangelands, as well as a love for

the local flora, to budding young farmers.

In 1931 Willi Giess contracted Malta fever and, as

was quite common in those days, was sent to Germany to

recuperate. Tme to his nature, he was not satisfied with

FIGURE 1. — Heinrich Johann Wilhelm Giess (1910-2000).

spending time resting, but enrolled at the Animal

Breeding Institute (Herdbuch-Abteilung der Karakul-

schafzucht, Tierzuchtinstitut der Universitat Halle). He

completed his training as Technical Adviser for Karakul

Breeding in 1933 and returned to South West Africa

where he applied his newly gained knowledge as manag-

er of a karakul farm. After four years, Willi Giess pur-

chased his own farm, Dornfontein-Sud, between

Dordabis and Leonardville in the east-central part of the

country (Kolberg & Strohbach 2000).

Soon World War II followed and Willi Giess, togeth-

er with a number of others who later were to become

well-known botanists, was interned at Andalusia (now

Jan Kempdorp, South Africa) for six years. The

‘Andalusia Camp University’ was promptly established

and under the guidance of the late Prof. O.H. Volk, Giess

studied plant taxonomy. A number of plants were col-

lected from the area in the camp that was accessible to

internees, identified and drawn (Figure 2). He con-

tributed significantly to the booklet Bestimmungs-

schliissel fiir siidwesl-afrikanische Grasgattungen by

producing engravings on tomato-box wood for the illus-

tration of the key (Glen & Perold 2000). He was allowed

to return to his farm in December 1946 after having col-

lected plants in the Cape with the University of

Stellenbosch as his base.

In 1953, initiated by Prof. Dr H. Walter of the Uni-

versity of Hohenheim, Germany, the Director of

Agriculture, Dr J.S. Watt, agreed to establish a botanical

section within the directorate. Prof. Walter donated over

2000 herbarium specimens and Willi Giess agreed to

temporarily work on the development of a herbarium.

For four years he travelled weekly from his farm

Bergland, about 50 km south of Windhoek, into town. In

1957 the SWA Administration took Giess into permanent

employment as curator of what was then the SWA

Herbarium (Giess 1978). He held this post until his

retirement in 1975, but continued to work in the herbari-

um until 1980 under the new curator, the late Dr Mike

Muller (Gunn & Codd 1981). From April 1985 to

January 1986 he was again acting curator while his suc-

cessor completed his Ph.D. degree.

Willi Giess made an important contribution to the col-

lection of what is now the National Herbarium of

Namibia (WIND). Of the ± 73 000 specimens in the pre-

sent-day collection, about 18 750 were collected by him,

often together with others like O. Volk, D. van Vuuren,

B. de Winter and H. Merxmiiller. These specimens were

always collected in triplicate, often more. Anyone who

has ever collected for a herbarium will appreciate the

amount of work involved in such a collection. Even

today, the Giess specimens, with their carefully typed or

hand-written data labels, are often the best, if not the

only ones of a certain species, to be found in the collec-

tion of the National Herbarium.

242

Bothalia 31,2 (2001)

FIGURE 2. — Ipomoea bolusiana

painted by W. Giess in 1942.

During his collecting trips, Giess covered almost the

entire country (Figure 3). Important collections, which

have to a large extent not been repeated in years following

his expeditions, are those made in the Kaukauveld (later

known as Bushmanland), Andara area on the Okavango

River, Brandberg area, Luderitz District (Aus,

Zebrafontein, Namuskluft, Witputz), Farm Ameib in the

Erongo Mountains and the Kaokoveld. Plant collecting

trips to the Kaokoveld in the late 1950s must have been

quite a feat, considering that even to this day, with modem

vehicles, equipment, maps, GPSs and improved roads, a

trip to the Kaokoveld is considered somewhat of an adven-

ture! The collected specimens of Merxmiiller and Giess

were often the basis for taxon descriptions in the

Prodromus einer Flora von SUdwestafrika, the last volume

of which was completed in 1971, and to date is still the

only reference work for many Namibian plant groups.

Other collections that made a considerable difference

to botanical knowledge in Namibia were his many index

cards of indigenous plant names. When Willi Giess

moved from Windhoek to Swakopmund, just two months

before his death, these valuable boxes of cards were

donated to the National Botanical Research Institute

where they will be incorporated into a proposed publica-

tion of local common names for plants.

• Up to 100 specimens collected per grid

s Between 101 and 300 specimens collected per grid

▲ Between 301 and 600 specim ens collected per grid

FIGURE 3. — Quarter-degree grid

squares in Namibia where W.

Giess collected herbarium

specimens.

Bothalia3l.2 (2001)

243

Willi Giess first published the journal Dinteria (named

in honour of Prof. Kurt Dinter, Namibia’s first official

botanist) under the auspices of the South West Africa

Scientific Society in November 1968. He remained editor

until 1991 (vol. 21). The journal aimed at making contri-

butions to the flora and vegetation of Namibia. In vol. 4

(1971), Giess first published his Preliminary vegetation

map of South West A frica, which was re-printed twice — in

1978 and 1998, and is still the only approximation to a veg-

etation map of Namibia to this day (Kolberg & Strohbach

2000). Despite his very busy schedule, he managed to pub-

lish several papers (see end of this obituary; Giess 1989),

ranging from popular to scientific articles, including the

description of a number of new plant species: Aloe argen-

ticauda Merxm. & Giess, A. dewinteri Giess, A. narniben-

sis Giess, Aristida dewinteri Giess, Spliaeranthus wattii

Giess ex Merxm., Tridentea marientalesis (Nel) L.C.Leach

subsp. albipilosa (Giess) L.C.Leach and Zygophyllum

schreiberianum Merxm. & Giess.

Whilst in retirement, Giess utilized his very compre-

hensive private literature collection to compile the

Bibliography of South West African Botany, published by

the SWA Scientific Society in 1989.

The pioneering work of Willi Giess did not go unno-

ticed in the international botanical world. He was hon-

oured on several occasions: in July 1964, he became a

Correspondent of the Naturhistorisches Museum in

Vienna; in 1968, the Swedish Royal Academy of Sciences

in Stockholm awarded him the Great Linee Medal in

Silver; on the 200th anniversary of the birth of Alexander

von Humboldt. Willi Giess received a commemorative

issue on the life of Von Humboldt from the consul of the

Federal Republic of Germany to honour his achievements

in the field of botany; the Ba\ arian Academy of Science

honoured him with the Bene Merenti Medal for his con-

tributions to the Prodromus einer Flora von Sudwest-

afrika; the South African Association of Botanists award-

ed him a Certificate of Merit in 1979; in 1980, Giess

received the Academy Medal in Gold from the South

African Academy of Science and Arts; and in 1998, the

Windhoek Municipality honoured Willi Giess by naming

a street after him.

A number of plant species were named in honour of

Giess, bearing testament to his collecting efforts and the

assistance he gave to numerous scientists the world over;

Isoetes giessii Launert, Aizoon giessii Friedrich. Salsola

giessii Botsch., Crassula ausensis Hutch, subsp. giessii

(Friedrich) Toelken, Indigofera giessii A.Schreib.,

Zygophyllum giessii (Merxm.) A.Schreib., Euphorbia

giessii L.C.Leach, Commiphora giessii J.J.A.van der

Walt. Heliotropiurn giessii Friedr.-Holzh., Jarnesbritte-

nia giessii Hilliard, Petalidium giessii RG.Mey., Erio-

cephalus giessii M.A.N. Muller, Senecio giessii Merxm.,

Lachenalia giessii W.F.Barker and Stipagrostis giessii

Kers.

His keen interest and an eye for the unknown, meant

that he often also collected insects associated with

plants. In this way, a termite, Okavangoterrnes giessii

(Coaton 1971), and a beetle, Zophosis (Protodactylus)

giessii (Koch 1962), were also named after him since he

collected the first specimens. The former is a monotyp-

ic genus and is known from only a single site east of

Rundu. The beetle is the most widespread of the

Protodactylus subgenus and occurs along the outer mar-

gin of the Namib Desert from Gobabeb to Rosh Pinah.

I had the pleasure of working with Willi Giess for

merely one year and was absolutely amazed by the

wealth of his knowledge. For every species, locality or

collector, he had some anecdote to tell or some memories

to share. I remember going on a short field trip with him

to collect material for Mike Muller, who was struggling

through his Ph.D. in Stellenbosch. While passing his for-

mer farm Bergland, he showed me a fair-sized hill, which

he had de-bushed manually in the early 1950s. To this

day, the vegetation on this hill is visibly different from

those nearby. Every little hill or farm we passed he would

have something to say, like: ‘Oh yes. Aloe viridiflora

occurs in these koppies. I collected it there in 19 ’or

‘When old so-and-so had this farm, the grazing condi-

tions looked much better!’ or ‘We spent three days under

this tree in 19.... when our car broke down and we had to

wait for the Government Garage to send a mechanic.’ I

continued to be amazed by this extraordinary person

when, about 20 years after retirement, he could still iden-

tify just about any plant you presented to him!

It was also a great inspiration to see Willi Giess and

his wife Brigitte (formerly Bleissner) work together and

support each other in all aspects of life. If one of them

forgot where something was written or to be found, the

other would remember. From 1963 to 1980 Brigitte

Giess assisted her husband in the herbarium and accom-

panied him on his many collecting trips on a voluntary

basis. Together with Volk they collected a large number

of specimens in the early 1960s. Sadly his partner and

soul-mate pre-deceased him in 1999.

Willi Giess died in Swakopmund on 28 September

2000. It is with sadness that we say farewell to our friend,

colleague and mentor. Namibian botany would have been

much the poorer without his invaluable pioneering work.

ACKNOWLEDGEMENTS

I gratefully acknowledge the information received

from Dr E. von Koenen. Patricia Craven is thanked for

producing the map, extracting information on Giess col-

lections from the WIND database and making some

valuable suggestions for inclusion in this text. The use of

data from the National Botanical Institute, Pretoria, is

gratefully acknowledged. My appreciation also to

Eugene Marais for providing information on the insects

mentioned in this article. Many thanks to Gillian Maggs-

Kdlling, who reviewed drafts of this paper.

REFERENCES

GIESS. W. 1978. 25 Jahre Landesherbarium Stidwestafrika. Mitteilungen

der SWA Wissenschaftlichen Gesellschaft XIX/2: 3-5.

GIESS. W. 1989. Bibliography of South West African Botany. SWA

Scientific Society, Windhoek.

GLEN, H.F. & PEROLD, S.M. 2000. Obituary; Otto Heinrich Volk

(1903-2000). Bothalia 30: 215-224.

GUNN. M. & CODD, L.E. 1981. Botanical exploration of southern

Africa: 400. Balkema, Cape Town.

KOLBERG. H. & STROHBACH, B. 2000. Obituary: Heinrich Johann

Wilhelm Giess (1910-2000). Dinteria 26: 21-24.

244

Bothalia 31.2 (2001 )

PUBLICATIONS BY W. GIESS

EISEB. E.. GIESS, W & HAACKE, W.H.G. 1991 . A preliminary list of

Khoekhoe (Nama/Damara) plant names. Dinteria 21: 17-30.

GAFF. D.F. & GIESS, W. 1985. Drought resistance in water plants in

rock pools of southern Africa. Dinteria 18: 17-36.

GIESS, W. 1962a. Some notes on the vegetation of the Namib Desert

with a list of plants collected in the area visited by the Carp-

Transvaal Museum Expedition during May 1959. Cimbebasia

2:1-35.

-1962b. Wunder unserer Flora — Wonderwereld van ons plantegroei —

Wonders of our flora. Kleine Reihe 5: 1-34. Afrika Verlag. Der

Kreis, Windhoek.

-1962c. Herbarium und Planzensammeln, aus der Praxis fur die Praxis.

Wissenschaflliche Forschimg in Siidwestafrika 1 : 38^3. SWA

Scientific Society. Windhoek.

-1965. Die Vegetationsverhiiltnisse (des Schwarzen Nossobgebietes).

In H.W. Stengel. Der Schwarze Nossob. Wissenschaftliche For-

schimg in Siidwestafrika 4: 45^9. SWA Scientific Society,

Windhoek, (translated into English and Afrikaans)

-1966a. 'Veldkost' in Siidwestafrika. Journal of South West Africa

Scientific Society 20; 59-68.

-1966b. Hoodia pan'iflora N.E.Br., eine fiir Siidwestafrika neue Hoodia-

Art. Mitteilungen der Botanischen Staatssammlimg, Miinchen

6: 239-243.

-1966c. Plantkundige versamelingsritte in Suidwes-Afrika. SWA Annual

1966: 21-25.

-1966d. The ‘Veldkost’ of South West Africa. SWA Annual 1966:

105-107.

-1968a. Kurt Dinter. Dinteria 1: 4-7.

-1968b. A short report on the vegetation of the Namib coastal area from

Swakopmund to Cape Frio. Dinteria 1: 13-29.

-1968c. Die Gattung Rhigozum Burch, und ihre Arten in Siidwest-

afrika. Dinteria 1: 31-51.

-1969a. Die Verbreitung von Lindernia intrepidus (Dinter) Oberm.

(Chamaegigas intrepidus Dinter) in Siidwestafrika. Dinteria 2:

23-27.

-1969b. Prodromus einer Flora von Siidwestafrika. Dinteria 2: 29-36.

-1969c. Welwitschia mirabilis Hook. fil. Dinteria 3: 1-55.

-1970a. Eine neue Aloe aus der Namib {Aloe namibensis Giess).

Mitteilungen der Botanischen Staatssammlimg, Miinchen 8:

123-126.

-1970b. Ein Beitrag zur Flora des Etoscha Nationalparks. Dinteria 5:

19-55.

-1970c. Moringa ovalifolia, die boom van die sprokieswoud in die

Nasionale Etosha Wildtuin. Dinteria 5: 57, 58.

-1970d. Die Verbreitung von Moringa ovalifolia Dinter & Berger in

Siidwestafika. Dinteria 5: 59-64.

-197()e. Pflanzenwunder in Vley, Pfannen und Wassertiiinplen im

trockenen Siidwestafrika. SWA Annual 1970: 181-183.

-1971a. Gramineae, a new species of Aristida from South West Africa.

(Aristida dewinteri Giess). Bothalia 10: 365, 366.

-1971b. A preliminary vegetation map of South West Africa. (Text in

English. German & Afrikaans). Dinteria 4: 1-1 14.

-1972a. Vegetation types. In W.G.H. Coaton & J.D. Sheasby,

Preliminary report on a survey of the termites (Isoptera) of

South West Africa. Cimbebasia Memoir 2: 15.

-1972b. Der Botanische Garten auf der Farm Lichtenstein bei Wind-

hoek im Jahre 1922. Dinteria 7: 13-15.

-1973. A new species of Aloe from South West Africa. (Aloe dewinteri

Giess). Bothalia 11: 120-122.

-1974a. Zwei Fahrten zur Jensenobotrva lossowiana Herre. Dinteria

10: 3-12.

-1974b. South West Africa: Flora. Standard Encyclopedia of South Africa

10: 149-150. Nasou, Cape Town.

-1974c. Beobachtungen zur Siidwester Flora. Dinteria 10: 35^4.

-1974d. Eine neue Stapelie aus Siidwestafrika. (Stapelia albipilosa

Giess). Mitteilungen der Botanischen Staatssammlimg, Miinchen

11: 349-352.

-1976. A preliminary vegetation map of South West Africa. Boissiera

24: 651.

-1978a. 25 Jahre Landesherbarium Siidwestafrika. Mitteilungen der

SW4 Wissenschaftlichen GeselLschaft XlX/2: 3-5.

-1978b. Die boom van die sprokieswoud in die Nasionale Etosha

Wildtuin. Staatsmuseum Windhoek 9: 10-12.

-1978c. Ein schbner botanischer Fund. (Drosera indica). Mitteilungen

der SWA Wissenschaftlichen GeselLschaft XIX/2: 6.

-1979. The genus Criniim in South West Africa. The Indigenous Bulb

Growers Association of South Africa 28: 1-6.

-1981a. Pflanzen der Namib. SWA Annual 1981 : 34— 41 .

-1981b. Eine botanische Namibfahrt. SWA Annual 1981: 81-86.

-1981c. Die in der Zentralen Namib von Stidwestafrika/Namibia fes-

gestellten Pflanzenarten und ihre Biotope. Dinteria 15: 13-72.

-1982a. Weitere Neunachweise zur Flora des Brandberges. Dinteria

16: 7-9.

-1982b. Zur Verbreitung des Tabaks in Siidwestafrika (Nicotiana afri-

cana Merxm.). Dinteria 16: 11-20.

-1984a. Die Pflanzenwelt des GroBen Gamsbergs. Journal of South

West Africa Scientific Society XXXVlll : 29-41 .

-1984b. Short note on Brachystelma blepharanthera Huber. Dinteria

17:81,82.

-1989a. Einiges zu unserer Flechtenflora. Dinteria 20: 30-32.

-1989b. Bibliography of South West African Botany. 236. SWA Scien-

tific Society, Windhoek.

GIESS, W. & SNYMAN, J.W. 1986. The naming and utilisation of

plant life by the Zu’hoasi Bushmen of the Kau-Kauveld. In R.

Vossen & K. Keuthmann. Contemporary studies on Khoisan I

& II. In Honour of Oswin Kohler on the occasion of his 75th

birthday: 237-346. Buske, Hamburg.

GIESS, W. & TINLEY, K.L. 1968. South West Africa (with vegetation

map of SWA). Conservation of vegetation in Africa south of the

Sahara. Acta phytogeographica suecica 54: 250-253.

MERXMULLER, H. & GIESS, W. 1974. Aloe pachygaster Dinter und

eine damit verwechselte neue Art. (Aloe argenticauda).

Mitteilungen der Botanischen Staatssammlimg, Miinchen 11;

437-444.

ROBINSON, E.R. & GIESS, W. 1974. Report on the plants noted in

the course of a trip from Liideritz Bay to Spencer Bay, January

10-21, 1974. Dinteria 10: 13-17.

HERTA KOLBERG*

* National Plant Genetic Resources Centre, National Botanical Research

Institute, Private Bag 13184, Windhoek, Namibia.

Bothalia31.2: 245 (2001)

Book Review

MEDICINAL. POISONOUS. AND EDIBLE PLANTS IN NAMIBIA

by EBERHARD VON KOENEN, translation of German edn 2. Klaus

Hess Publishers, Windhoek-Gottingen. 2001. Pp. 335. 128 full-page

black-and-white drawings. 6 figures (maps and graphs). A4 size, hard

cover. ISBN Namibia 99916-747-4-8. ISBN Germany 3-9804518-7-9.

Price: ZAR 350.

'The rigid barriers that existed between conventional and alterna-

tive medicine are beginning to open up. The awareness is growing that

the greatest benefit for the sick lies not in opposing but in combining

the different methods of healing. Experience at my Windhoek surgery

has shown the versatility and effectiveness of natural remedies in close

collaboration with orthodox medicine. Many pharmacies which in the

past stocked only pharmaceutical medicines, today keep large supplies

of homeopathic and phytotherapeutic remedies'. Observations similar

to these, made in the preface, are discussed in an article in the New

Scientist of 26 May (Anon. 2001 ): ‘Billions a year are spent worldwide

on herbal medicines. In the US the use of complementary and alterna-

tive medicine has doubled in the past decade. In Britain almost half the

adult population admits to using an alternative therapy'. In southern

Africa more than 80 % of the population subscribe to traditional medi-

cal care. According to a recent estimate, some 20 million people in

South Africa alone consume almost 20 000 tonnes of plant material per

year during 90 million incidents of use (Mander 1998). Witness, there-

fore. the growing number of publications dealing with plants and their

power to heal, to feed and to kill. A forthcoming publication by the

National Botanical Institute (Arnold et al. in prep.) cites 16 major

works dealing with African ethnomedicinal plants which have appeared

during the last 10 to 12 years alone.

The predecessor of the present work first appeared in 1 977 under the

title Heil- und Giftpflanzen in Siidwestafrika. It has since gone through

several reprints and a second edition in German. The present book is a

translation of this latest edition. Binding, paper and printing are still of

a similar exceptionally high standard as in the first edition, and text and

illustrations have been considerably expanded. The introductory chapter.

The Natural Environment, presents an overview of the climate, geology,

physiography and vegetation of Namibia, illustrated by maps, graphs

and diagrams. Under the heading. The Biotopes, ten striking full-page

pencil drawings of Namibian landscapes by the author are included,

from barren wind-swept dune wastes and wide drainage lines support-

ing isolated specimens of Welwitschia in the Namib. to mopane savan-

na and the 'Fairy forest' along the Omaruru River.

Among the healers, and sometimes mischief-makers, as the author

puts it. he distinguishes three main groups: the traditional healers, the

soothsayers or clairvoyants and the black magicians. The soothsayer

must have a disposition for spiritual perception, he tosses bones or

sticks and draws conclusions from their position and thus finds the

causes of diseases, disasters or inexplicable deaths. The black magician

is an evil-doer; he sends disaster, lingering illness and death to others.

The traditional healer uses many medicinal herbs, physical healing

techniques and various utensils in order to invoke metaphysical forces

to neutralise the symptoms of a disease. The essential role of the tradi-

tional healer is today recognised by Public Health Services in many

parts of the world, including Namibia and southern Africa at large.

Dr von Koenen has acquired his knowledge of the healing, feeding

and harmful potential of plants primarily in Namibia, from traditional

healers and herb experts in all regions of the country. This is reflected

in the many common names of plants in 13 of the languages spoken in

Namibia. In the words of the author: ‘In receiving and handing over

this knowledge. I have also assumed the responsibility of pointing out

incessantly that the healing plants are meant to serve the ill and not the

greed for profit of unwelcome moneymakers’. Here he may have

thought of species such as Harpagophytum procwnbens. the Devil’s

claw or Grapple plant, here referred to as Wool spider: some 40 years

ago a dreaded weed; today, due to its extreme popularity as a healing

plant, especially in Central Europe, occasional to rare, except in more

remote or protected areas. And there are numerous much rarer species

which are seriously threatened by plant collectors. Von Koenen adds

another caveat; ‘It should not be expected that the whole range of

medicinal herbs can be recommended for the pharmaceutical process.

Interest should initially be focused on those plants which are used by

different healers to treat more or less the same syndromes.’

The 600 species dealt with — two fungi, five ferns, one gymnosperm

and 592 flowering plants-are numbered and arranged in alphabetical

sequence with an indication of the family to which they belong. The 49

families of vascular plants represented are also shown on a separate

alphabetical list ending with the two groups to which the two fungi

belong. This is followed by a list of families in ‘botanical’ (± Englerian)

order with all species of a given family arranged in alphabetical

sequence. An alphabetical list of all species dealt with precedes a listing

of the southern African distribution of all species in terms of countries

or provinces. A diagram describing the degree square system and a map

of Namibia explain the method according to which the distribution of

species is indicated. Abbreviations used for the 13 ethnic groups or lan-

guages cited as source for the vernacular names are given, and the per-

sons responsible for the naming and the descriptions of the species are

mentioned. The bibliography comprises 45 literature sources.

The treatment of each species begins with the number assigned to

it in the work. and. in the case of trees, with the National Tree Number.

They are followed by the scientific name and family, and a list of the

common names. Eor some of the more common plants, such as Euclea

divinorum, the Wild ebony, or Pterocarpus angolensis, the Transvaal

teak, vernacular names in ten different languages are listed, and for

most other plants in at least three or more. The plant descriptions are

brief and image-forming and the translators have attempted to follow

the free, sometimes almost florid style of the author. The discussion of

the importance of the plant is usually much longer than the descrip-

tions. consisting, on average, of 200 words or more. The source of the

information is indicated — no. 42 (Watt & Breyer-Brandwijk, The

medicinal and poisonous plants of southern and eastern Africa, edn 2.

1962) is the most commonly cited reference. The species treatments

end with an indication of the distribution in Namibia.

One hundred and seventeen species are illustrated on full-plate

drawings by the author. 40 more than in the first German edition. The

new drawings can easily be distinguished: they are not signed and dated

by the author and shading on them is not brought about by stippling, as

in the old drawings, but by stippling combined with grey toning. In con-

trast to the old drawings they therefore had to be screened before they

could be printed. The grey tones often run over the lines, and the new

drawings therefore tend to lack the clear outlines of the old ones. All in

all. 1 prefer the ones from edition 1 . In most cases the artist has caught

enough diagnostic features to make the species recognisable. Soft

leaves, however, often look leathery, and cylindrical organs may look

rather flat. The drawing of Diospyros lycioides subsp. lycioides should

rather have been omitted. The work ends with a list of vernacular

names, a subject index excluding plant names, acknowledgements,

subscriber list and a short curriculum vitae of the author.

Let me conclude with a brief quote from a typical conversation

between the author and a traditional healer: '1 am doing this work not for

my personal profit but in the interest of our country and our fellow coun-

trymen. It must be done today, for the old. who stiU have the knowledge, will

take it with them into the grave. And when you ask the young, you will

mostly just get the reply: a pill is more convenient. But if some day the

urgent question is asked about the medicinal plants which grow in our

country — and the day will come — and there is nobody left who knows,

this book, as the outcome of our work, will provide the answer’.

REFERENCES

ANON. 2001. Hype, hope and healing. New Scientist 2292: 28-30.

ARNOLD, T., PRENTICE, C. A., HAWKER. L.C., SNYMAN. E.E.,

TOMALIN, M., CROUCH, N.R. & POTTAS-BIRCHER, C. in

prep. Medicinal and magical plants of southern Africa: an anno-

tated checklist. Strelitzia.

MANDER, M. 1998. The marketing of medicinal plants in South

Africa: a case study in KwaZulu-Natal. FAO, Rome.

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

and poisonous plants of southern and eastern Africa, edn 2.

Livingstone, Edinburgh & London.

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Bothalia 31,2: 247-263 (2001)

National Botanical Institute South Africa:

administration and research staff 31 March 2001,

publications 1 April 2000-31 March 2001

Compiler: B.A. Momberg

CHIEF DIRECTORATE (ADIR)

CAPE TOWN— PEARSON HOUSE

Huntley, Prof. B.J. M.Sc. Chief Executive Officer

Adams, Ms M.A. Typist/Receptionist

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MARKETING AND COMMUNICATION SUBDIRECTORATE (AMAR)

CAPE TOWN

Moeng, S. M.A. (Languages). Deputy Director

Davis, G.W. Ph.D. Assistant Director. Programme Development/Fundraising

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HUMAN RESOURCES DIRECTORATE (APER)

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Tlali, P.L.N. B.Iuris Lib. Director. Human Resources

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Haupt, Mrs C.S. Administration Aid II. Guest house

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Van Zyl, J.M. M.Econ.dnd. Psych.). Assistant Director: Organizational Development & Training

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Du Toit, Ms R. H.E.D. Senior Training Officer Williams, Mrs L.R. Dip. Sec. Senior Personnel Officer II

Fredericks, C.H. Courier II

248

Bothalia 31,2 (2001)

ADMINISTRATION AND HUMAN RESOURCES— PRETORIA (RPTA)

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Africa, S. B.Tech. Internal Auditor

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Cassiem, Ms S. Senior Accounts Clerk III. Staff benefits

& salaries

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Willis, C.K. M.Sc. (Cons. Biol.) — Director: Gardens and Horticultural Services

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Woodward, Ms Y.J. Senior Administration Clerk III. Secretary

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Britz, R.M. Curator: Fowveld NBG Oliver, I.B. Curator: Karoo NBG

Turner, Ms S.F. Curator: Witwatersrand NBG Tarr, B.B. Curator: Natal NBG

Heilgendorff, J.P. Curator: Pretoria NBG

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Control Inspector of Works

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Peck, W.I. Artisan’s Assistant II. Building maintenance

Koyana, Ms Z. Senior Accounts Clerk I. Salaries

Mcontsi, Ms N. Accounts Clerk II. Finance

Mirkin, Ms Y.A. Senior Secretary II

Paulse, Mrs D.W.S. Senior Accounts Clerk III

September, Ms M. Senior Accounts Clerk I. Debtors

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HAROLD PORTER NBG— BETTY’S BAY (GHPG)

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Arendse, Ms M. Groundsman II. Gate Jacobs, L.H. Groundsman II. Plant sales

Bezuidenhout, Mrs H.M. Senior Administration Clerk III October, Ms R.P. Groundsman II. Plant records

Forrester, Ms J.A. N.T.C.IIKHort.). Chief Technician. Hor- Samuels, Ms D.C. Cleaner II

ticulture Van Wyk, A.B. Groundsman I. Index nursery

Harper, C.C. Driver

Bothalia 31.2 (2001)

KAROO NBG— WORCESTER (GKAR)

Oliver, I.B. N.Dip.(Hort.)(PRA). Control Technician

249

Ashworth, Mrs E.H. Senior Administration Clerk III Viljoen, D.M. N.Dip.tHort.). Chief Technician. Collec-

Mpeke, Ms E.N. Cleaner II tions

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KIRSTENBOSCH NBG— CAPE TOWN (GKBC)

Le Roux, PH. N.Dip.(Hort.), N. Dip. (Parks & Recr.). Deputy Director; Garden. Curator.

Head; Agricultural Support Services

Goldschmidt. S.M. B.A.(Soc.)(Hons.). Assistant Director. Personnel Management

Morkel, A.T. N. Dip. (Nat. Cons.). Control Technician. Estate Manager

Hitchcock. A.N. N.H.Dip.(Hort.). Control Technician. Supervisor; Nursery

Jaques, R.F. Chief Technician. New plant introductions

Notten, Ms A.L. B.Sc., N.Dip.(Hort.). Chief Technician. Supervisor; Seed room

Trautman, C.E. Artisan. Supervisor; Workshop

Adams, TD. N.Dip.(Hort.). Technician. Greenhouse Supervisor

Arends, Ms S.J. Administration Clerk. Plant records

Duncan, G.D. N.Dip.(Hort.). Chief Technician. Bulbs

Engelbrecht, F. Storeman II

Engelbrecht, Mrs L.D. Technician. Plant records

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Jamieson, Mrs H.G. N. Dip. (Parks & Rec.). Chief Tech-

nician. Restio/Asparagus (part time)

Jodamus, Ms N.L. N.Dip.(Hort.). Senior Technician; An-

nuals

Kamalie, Ms S. Typist II. Receptionist

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Ericas

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Outreach Programme

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Herbaceous collections (part time)

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cian. Succulents

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production

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N. Dip. (Parks & Rec. Admin.), N.T.C.IIKHort.).

Control Technician. Horticulture

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III

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Johnson, Ms I. B.Sc. Senior Technician Van der Merwe, Mrs M.E.H. Senior Administration Clerk

Nonjinge, S.H.B. N.T.C.IIKHort.). Chief Technician III

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FREE STATE NBG— BLOEMFONTEIN (GFSG)

Mogale, A.O. N.Dip.(Hort.). Control Technician

Eysele, Mrs J.P. Senior Administration Clerk III

Harris, Ms S. N.Dip.(Hort.). Technician

Lumley, M.J. Chief Scientific Officer. Nursery

Raditlhare, Mrs E.M. Administration Aid II

Thaele, Mrs M.E. Administration Aid II

250

Bothalia 31,2 (2001)

PRETORIA NBG (GPTA)

Heilgendorff, J.P. H.N.Dip.(Hort.). Control Technician

Baloyi, K.J. Senior Auxiliary Services Officer II. Garden

records

Baloyi, M.S. Dip.(EBM), Dip.(PTM), Dip.(Payroll Admin.).

Senior Administration Clerk I. Personnel records

Creighton, Ms D.D. Administration Clerk III

Eyssell, Ms A. B.Sc.(Hons)(Agric.). Senior Technician.

Production and sales nursery

Keyter, B.A. Senior Security Officer I

Klapwijk, N.A. N.Dip.(Hort.), N. Dip. (Plant Prod.),

N. Dip. (Diesel Fitting). Control Technician. Plan-

ning and development. Index nursery. New Plant

Company

Law, C. N.Dip.(Hort.). Senior Technician

Mariri, Ms M.A. Administration Aid II. Cleaner

Matshika, S.P Groundsman II. Cook

Modisha, M.D. Groundsman II. Cleaner

Swartz, Ms P.P. M.Sc. Chief Horticulturist. Scientific

and horticultural curation of living collections of

succulents, orchids, medicinal and rare and endan-

gered plants; garden planning and development;

Madagascan plants; tour groups and visitors

Venter, W.A. N.T.C.II. Senior General Foreman. Main-

tenance

WITWATERSRAND NBG— ROODEPOORT (GWIT)

Turner, Ms S.L. B. Sc. (Hons), N.Dip.(Hort.). Control Technician

Aubrey, Mrs A.E. B.Tech.(Hort.). Senior Horticulturist.

Plant records, interpretation, information

Hankey, A.I. N.Dip.(Hort.). Specialist Horticulturist. Gar-

den, estate, collections, nursery

Head, Mrs S.E. Dip. (Typing). Administration Officer

Manjati, Mrs N.L. Senior Administration Clerk I

Mmola, Mrs B.E. Administration Aid II. Cleaner

Mutshinyalo, T.T. N.Dip.(Hort.) Horticulturist. Nursery,

garden

Ndzondo, Mrs P.G. Administration Aid II. Cleaner

Tebeile, Ms Z.M. Senior Administration Clerk I. Recep-

tionist

RESEARCH DIRECTORATE (RDIR)

PRETORIA

Smith, Prof G.F. Ph.D., F.L.S. Director: Research

Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director: Ecology and Conservation (Cape Town)

Wolfson, Mrs M.M. Ph.D. Deputy Director: Education and Research Support

Klopper, Ms R. M.Sc. Technician (contract worker) worker)

Marais, Mrs A.C. Senior Secretary IV Steyn, Dr E.M.A. D.Sc. Principal Scientist. Embryology,

Meyer, Mrs N.L. B. Sc. (Hons) Technician (contract anatomy, taxonomy

PLANT SYSTEMATICS SUBDIRECTORATE

PRETORIA

Smith, Prof. G.F. Ph.D., F.L.S. Systematics of succulents and rosulate, petaloid monocots

Arnold, T.H. Head: Data Management (Pretoria)

Crouch, N.R. Head: Ethnobotany Unit (Durban)

Koekemoer, Miss M. Curator: National Herbarium (Pretoria)

Leistner, O.A. D.Sc. F.L.S. Scientist (contract worker)

Rourke, Dr J.P. Curator: Compton Herbarium (Cape Town)

Siebert, S.J. Regional Project Co-ordinator: SABONET (Pretoria)

Williams, Ms R. Curator: Natal Herbarium (Durban)

COMPTON HERBARIUM— CAPE TOWN (RHEC)

Rourke, J.P. Ph.D., F.M.L.S., F.R.S.S.Af Assistant Director. Systematics of

southern African Proteaceae, Stilbaceae

Baatjes, Ms A. Data Capturer (SABONET contract wor- Curator: Collections. Taxonomy of the Gnidieae

ker) (Thymelaeaceae)

Beyers, Mrs J.B.P. Ph.D. Principal Scientist. Assistant Chesselet, Ms P.C.M. M.Sc. Chief Scientific Officer

Bothalia 31.2 (2001)

251

Conrad. Ms C. M.Sc. Scientist. Molecular systematics

laboratory (SABONET contract worker)

Cupido, C.N. M.Sc. (Hons). Chief Scientific Officer

Cupido, Mrs C.S. Auxiliary Services Officer II. Tech-

nical Assistant

Engelbrecht, Ms M. (SABONET contract worker)

Foster, Mrs S.E. Senior Secretary IV

Kurzweil, H. Ph.D. Specialist Scientist. Systematics of

southern African terrestrial orchids

Leith, Mrs J. Senior Administration Clerk III

Manning, J.C. Ph.D. Specialist Scientist. Systematics of

Iridaceae and Orchidaceae; anatomy

Marinus, Ms E.D.A. Principal Auxiliary Services Offi-

cer. Herbarium Assistant

Oliver, E.G.H. Ph.D. Principal Scientist. Taxonomy of

the Ericoideae (Ericaceae)

Oliver, Mrs I.M. (contract worker)

Parker, Ms F. B. Sc. (Hons). (SABONET contract worker)

Paterson-Jones, Mrs D.A. (nee Snijman) Ph.D. Principal

Scientist. Systematics of Amaryllidaceae; cladistics

Reeves, Ms G. Ph.D. Molecular systematics (contract

worker)

Roux, J.P. N.T.C.IIKHort.), F.L.S., Ph.D. Principal

Scientist. Systematics of Pteridophyta

Williams, Mrs V.J. Data Capturer (SABONET contract

worker)

NATAL HERBARIUM— DURBAN (RHED)

Williams, Ms R. B. Sc. (Hons), H.D.E. Chief Scientific Officer

Govender, D. M.Sc. Student

Govender, Mrs N. B. Sc. (Hons) Chironia systematics

(SABONET contract worker)

Hlongwane, Mrs N.C. Administration Aid II. Cleaner &

messenger

Ngwenya, A.M. Senior Scientific Officer. Herbarium As-

sistant. Plant identification and information, Zulu

Plant Names Project

Mazibuko, J.V.G. Auxiliary Services Officer II. Herba-

rium Assistant

Noble, Mrs H-E. Senior Administration Clerk III

Ntuli, Mrs N. Data Capturer (SABONET contract worker)

Singh, Ms Y. M.Sc., H.E.D. Senior Scientist. Taxonomy

of Araceae, Hypoxis

ETHNOBOTANY UNIT— DURBAN (RETH)

Crouch, N.R. Ph.D. Principal Scientist. Ethnobotany of southern African flora

NATIONAL HERBARIUM— PRETORIA (RHEN)