Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 39,1

May 2009

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE PRETORIA

Obtainable from the South African National Biodiversity Institute (SANBI), Private Bag X 1 0 1 , 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 South African National Biodiversity 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, (c) to Vols 26-30, and (d) to Vols 31-37 (2001—

2007) 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 South African National Biodiverity 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 1-49 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, syn-

onymy, 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 palaeofiora on a pattern comparable to that of the Flora of southern Africa. Much of the information is pre-

sented in the form of tables and photographic plates depicting fossil populations. Now available:

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

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), 1983, by J.M. & H.M.

Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, 1985, by J.M. & H.M. Anderson.

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

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

Anderson (ed.).

Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications,

2003, by J.M. Anderson & H.M. Anderson. Strelitzia 15.

Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology, 2007, by

J.M. Anderson, H.M. Anderson & C.J. Cleal. Strelitzia 20.

Molteno ferns: Late Triassic biodiversity in southern Africa, 2008, by H.M. Anderson & J.M. Anderson.

Strelitzia 2 1 .

SANBI BIODIVERSITY SERIES

A series of occasional reports on projects, technologies, workshops, symposia and other activities initated by or

executed in partnership with SANBI.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 39,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

national

biodiversity

institute

S A N B I

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

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

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

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

Carter Holmes, Mrs S. Royal Botanic Gardens, Kew, UK.

Coates Palgrave, Mrs M. 9 Blue Kerry, 30 Steppes Rd, P.O. Chisipite, Harare, Zimbabwe.

Dold, A. Albany Museum, Grahamstown, RSA.

Edwards, Dr T.J. 81 Diane Crescent, Croydon, Victoria, 3136 Australia.

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

Hartmann, Dr H.E.K. Institut fur Allgemeine Botanik, Hamburg, Germany.

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

Lavranos, J. Apartado Postal 243, 8100 Louie, Portugal.

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

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

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

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

Van Wyk, Prof. A.E. University of Pretoria, RSA.

Venter, Prof. H.J.T. University of Free State, Bloemfontein, RSA.

Walsh, Dr N.G. Royal Botanic Gardens, Victoria, Australia.

Welman, Ms W.G. South African National Biodiversity Institute, Pretoria, RSA.

Winter, PJ.D. South African National Biodiversity Institute, Pretoria, RSA.

Xiang, Dr Q.-Y. Gardner Hall, North Carolina State University, Raleigh, USA.

Date of publication of Bothalia 38,2: 14 October 2008.

CONTENTS

Bothalia 39,1

1 . New species of Moraea (Iridaceae: Iridoideae), with range extensions and miscellaneous notes for southern

African species. P. GOLDBLATT and J.C. MANNING 1

2. Three new species of Diascia (Scrophulariaceae) from the Western Cape, South Africa. K.E. STEINER 11

3. Aloe in Angola (Asphodelaceae: Alooideae). R.R. KLOPPER, S. MATOS, E. FIGUEIREDO and G.F.

SMITH . 19

4. Three new species of Gladiolus (Iridaceae) from South Africa, a major range extension for G. rubellus and

taxonomic notes for the genus in southern and tropical Africa. J.C. MANNING and P. GOLDBLATT 37

5. A revision of Fumariaceae (Fumarioideae) in southern Africa, including naturalized taxa. J.C. MANNING,

P. GOLDBLATT and F. FOREST 47

6. Two new species of Nemesia (Scrophulariaceae) from arid areas of the Northern Cape, South Africa. K.E.

STEINER 67

7. Taxonomy and phylogeny of two subgroups of Pelargonium section Otidia (Geraniaceae). 1. The

Pelargonium camosum complex. M. BECKER and F. ALBERS 73

8. A review of the genus Curtisia (Curtisiaceae). E. YU. YEMBATUROVA, B-E. VAN WYK and P.M.

TILNEY 87

9. Notes on African plants:

Asphodelaceae: Alooideae. Aloe hahnii, a new species in the section Pictae, in the Soutpansberg Centre

of Endemism, Limpopo Province, South Africa. R.R. KLOPPER and G.F. SMITH 98

Asphodelaceae: Alooideae. Formalizing the synonymy of Bulbine triebneri. R.R. KLOPPER and

G.F. SMITH 100

Campanulaceae. Theilera robusta, the correct name for Theilera capensis. C.N. CUPIDO 97

Ebenaceae. Typification and a new status in Diospyros. M. JORDAAN 101

Thymelaeaceae. The true identity of Synaptolepis kirkii. M. JORDAAN and C.L. BREDENKAMP. ... 104

10. Closing bodies in the capsular fruits of Ruschioideae (Aizoaceae) — a review. H. KURZWEIL and P.

BURGOYNE 107

11. Obituary: Robert Baily Drummond (1924-2008). M. COATES PALGRAVE 117

12. Book review 121

New combinations, name, species, subspecies and status in Bothalia 39,1 (2009)

Aloe hahnii Gideon F.Sm. & R.R.KIopper, sp. nov., 98

Cysticapnos vesicaria (L.) Fedde subsp. namaquensis J.C. Manning & Goldblatt, subsp. nov., 60

Diascia appendiculata K.E. Steiner, sp. nov., 15

Diascia collina K.E. Steiner, sp. nov., 1 1

Diascia pusilla K.E. Steiner, sp. nov., 13

Diospyros natalensis (Harv.) Brenan subsp. nummularia (Brenan) Jordaan, stat. nov., 102

Discocapnos mundii Cham. & Schltdl. [as ‘ mundtii\ subsp. dregei (Harv.) J.C. Manning & Goldblatt, stat. nov., 51

Gladiolus dolichosiphon Goldbatt & J.C. Manning, sp. nov., 37

Gladiolus karooicus Goldblatt & J.C. Manning, sp. nov., 39

Gladiolus reginae Goldblatt & J.C. Manning, sp. nov., 41

Gladiolus sulculatus Goldblatt, nom. nov., 45

Moraea longipes Goldblatt & J.C. Manning, sp. nov., 5

Moraea pearsonii Goldblatt & J.C. Manning, sp. nov., 2

Moraea tanquana Goldblatt & J.C. Manning, sp. nov., 4

Nemesia aurantia K.E. Steiner, sp. nov., 70

Nemesia suaveolens K.E. Steiner, sp. nov., 67

Pelargonium carnosum (. L .) L Her. subsp. ferulaceum ( Cav :) M.Becker & F. Albers, comb, nov., 73

Theilera robusta (A. DC.) C.N.Cupido, comb, nov., 97

IV

Bothalia 39,1: 1-10 (2009)

New species of Moraea (Iridaceae: Iridoideae), with range extensions

and miscellaneous notes for southern African species

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

Keywords: Iridaceae, Iridoideae, Moraea Mill., new species, southern Africa, taxonomy

ABSTRACT

Three new species are described in the largely sub-Saharan genus Moraea Mill. (± 200 spp.), all from its centre of diver-

sity in the winter rainfall region of southern Africa. Moraea pearsonii, from Hottentotskloof near Ceres in Western Cape,

flowers in late November and December when its leaves are ± dry, and has small, pale lilac, stellate flowers with the style

branches each divided to the base into filiform arms. Moraea tanquana, from the Tankwa River Basin in Northern Cape,

resembles the southern Namaqualand M. deserticola but has broad, plane leaves, short anthers exserted from a shallower

floral cup and a short style. In section Acaules, M. longipes from Namaqualand stands out in its early flowering habit, a stem

consisting of a single long intemode reaching well above the ground, short style and unusually long anthers. Moraea jarmilae

described from Ox Bow, Lesotho in 2002, is conspecific with M. albicuspa and is reduced to synonymy. Significant range

extensions are reported for M. elsiae, M. falcifolia, M. pseudospicata , M. spathulata, M. tricolor, M. vegeta, M. verecunda, M.

vespertina and M. vlokii. A yellow-flowered morph, local in the Perdebont Valley of the Little Karoo, is reported for the first

time in typically blue- to violet-flowered M. bipartita , as well as the occurrence of a hybrid swarm, rare in Moraea , between

M. bipartita and M. polyanthos.

INTRODUCTION

The old world and largely sub-Saharan genus Moraea

Mill. (Iridaceae: Iridoideae) comprises some 200 species

of cormous geophytes. Although florally diverse, Moraea

is recognized in Iridoideae by a bifacial, channelled (rarely

terete) leaf blade and corms of a single intemode derived

from a lateral bud. Most species have iris-like flowers with

flattened, petaloid style branches, with filaments at least

partially united. Other floral types make exact definition

of the genus difficult. Moraea is most diverse in southern

Africa and has a marked concentration of species in the

winter rainfall region of western South Africa and adja-

cent southwestern Namibia, the likely area of origin of

the genus (Goldblatt et al. 2002). Here we describe three

new species. Moraea pearsonii from Hottentotskloof,

near Karoopoort in the Ceres District, was re-collected in

2007 for the first time since its discovery 99 years ago,

when H.H.W. Pearson gathered fragmentary material in

November 1908. A late flowering species, it blooms in

November and late December, and the small, stellate flow-

ers have unusual, filiform style branches each divided to

the base. This style conformation appears to have evolved

independently four times in the genus. Moraea tanquana

from the Tankwa National Park, is allied to M. deserticola

Goldblatt from southern Namaqualand and M. speciosa

(L. Bolus) Goldblatt from the Western Karoo, and shares

with these species, cup-like, blue to mauve flowers with

subequal tepals. Although superficially resembling M.

deserticola, it differs from that species in the short anthers

exserted from the floral cup and short style dividing oppo-

site the anther bases so that the style branches emerge

between the anthers. In section Acaules, M. longipes from

Namaqualand stands out in its early flowering, aerial stem

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

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

blatt@mobot.org.

** Compton Herbarium, South African National Biodiversity Insti-

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

sanbi.org.

MS. received: 2008-05-27.

consisting of a single long intemode reaching well above

the ground, short style and unusually long anthers. Of

these new species all but M. longipes are currently known

from a single locality but we suspect that collecting nearby

at the appropriate times of the year, will show them all to

have wider ranges.

We take this opportunity to reduce Moraea jarmilae

J.J.Halda, described in 2002 from Ox Bow in Lesotho,

to synonymy in M. albicuspa Goldblatt, and to report

significant range extensions for several southern African

species. Populations of the southwestern Cape species M

elsiae Goldblatt and M. tricolor Andrews have been dis-

covered in the vicinity of Stilbaai in the southern Cape,

range extensions of 100 km or more from their next

nearest stations. The Stilbaai populations of M. tricolor

also exhibit a shift in flowering time and habitat, unprec-

edented in Moraea. The widespread eastern southern

African M. spathulata (L.f.) Klatt is now known from

the higher mountains of the Great Karoo, a surprising

record for a plant of otherwise mesic coastal and mon-

tane grassland habitats. A population that we refer to M.

vespertina Goldblatt & J.C. Manning, a species until now

known from two localities on the Bokkeveld Plateau

west of Calvinia, has been discovered in the Tankwa

National Park, some 200 km to the south. Although dif-

fering in some respects from the Bokkeveld populations,

notably their slightly larger flowers and drier habitat, the

Tankwa plants seem best included here for the moment.

We also report range extensions for Moraea falcifo-

lia Klatt, M. pseudospicata Goldblatt, M. spathulata, M.

vegeta L ., M. verecunda Goldblatt and M. vlokii Gold-

blatt, the last-named species known until now from

a small portion of the Swartberg. For M. tricolor, we

report for the first time, capsule and seed shape, and

compare these characters with closely related M. ciliata

(L.f.) Ker Gawl. in which seeds and capsules are unusu-

ally variable but appear to be correlated with flower col-

our. For M. bipartita L.Bolus we document the existence

of yellow-flowered populations in an otherwise blue- to

2

Bothalia 39,1 (2009)

violet-flowered species, and discuss the presence of a

hybrid swarm between M. bipartita and closely allied

M. polyanthos L.f. Lastly, we discuss a likely new spe-

cies from Namibia, too inadequately known to formally

describe and name.

NEW SPECIES AND TAXONOMIC ADJUSTMENTS

1. Moraea pearsonii Goldblatt & J.C. Manning,

sp. nov.

Plantae 200-450 mm altae ex cormo globoso 20-30

mm diam. profunde sub terrain infosso, tunicis ex fibris

nigris constantibus, folio producto solitario lineari-

canaliculato marginibus involutis 2 3( — 4) mm lato ad

1 m longo, caule erecto ramoso ramis ultimis rhipidia

sessilia ferentibus, spathis siccis, floribus pallide lila-

cinis exteme pallidis bubalinis inodoris, tepalis ± 14 mm

longis, extemis 4. 5-5.0 mm latis intemis ± 4 mm latis

unguibus 1.0-1. 5 mm longis, filamentis liberis ad basem

contiguis 3 .5^4.0 mm longis suberectis malvinis, anth-

eris 4. 5-5.0 mm longis flavis, stylo filiformi malvino ±

2.5 mm longo, ramis styli ad basem furcatis in brachia

filiformia duo 3. 5^1.0 mm longa productis, capsula doli-

iformi 4—5 x 3^1 mm.

TYPE. — Western Cape, 3319 (Worcester): Farm

Vrede, dry wash west of junction between Touwsrivier

and Ceres-Sutherland roads, (-BA), 15 December 2007,

Manning 3128 (NBG, holo.; K, MO, iso.).

Plants 200-450 mm high. Corm globose, 20-30 mm

diam., deeply buried; tunics of coarse black fibres, drawn

into finely fibrous neck up to 150 mm long. Stem erect,

usually with \-A suberect branches at upper nodes, dull

purplish where exposed; branches with (l-)4-6 sessile

lateral flower clusters. Foliage leaf solitary, basal, much

longer than stem, trailing, linear and channelled but

margins inrolled when dry and thus appearing terete, 2-

3(^1) mm wide and up to 1 m long, partially or entirely

dry at flowering, leathery; cauline leaves bract-like and

entirely sheathing, dry and papery, attenuate margins

united in lower half. Spathes dry and papery at flower-

ing, pale buff or lightly flushed with purple, attenuate,

inner 20-33 mm long, outer ± as long as inner. Flow-

ers pale lilac flushed buff on reverse; tepals with small,

transversely oblong, yellow nectar guides at limb bases,

unscented, shortly clawed, claws erect, 1.0-1. 5 mm long,

apressed to base of filaments, limbs spreading or slightly

reflexed, oblong-elliptical, outer broader, 13 x 4. 5-5.0

mm, inner 13x4 mm. Stamens free; filaments contigu-

ous at base, 3. 5^4.0 mm long, suberect, mauve; anthers

erect, 4.5-5. 0 mm long, yellow, curving inwards distally

at anthesis. Ovary ovoid, 2. 5-3.0 mm long; style erect,

filiform, ± 2.5 mm long, mauve, branches spreading to

ascending between anthers, divided to base into two fili-

form arms 3. 5-4.0 mm long. Capsules barrel-shaped, 4—

5 x 3^1 mm. Seeds angled by pressure, ± 1.5 mm diam.,

reddish brown, testa surface rugulose. Flowering time :

late November to late December; flowers opening at ±

1 7:30 and wilting at ± 20:30. Figure 1 .

Distribution and ecology : Moraea pearsonii is cur-

rently known only from the roadside between Hottentots-

kloof Farm and Karoopoort (Figure 2), almost midway

between the towns of Ceres and Touws River. Plants

grow in sandy alluvium over shale in washes at the foot

of the Baviaansberg, the southernmost extremity of the

Swartruggens, in open shrubland transitional between

renosterveld and arid fynbos.

Flowering in Moraea is typically in the wet season,

which is summer in eastern, southern and tropical Africa,

and late winter and spring in western southern Africa and

southwestern Namibia. A significant number of species,

however, exhibit a shift in their flowering to the dry sea-

son, although they tend to produce their leaves in the wet

season when conditions are optimal for vegetative growth.

Examples of such species include the eastern African M.

stricta Baker, which blooms in August to October but pro-

duces its leaf in November, and M. pseudospicata from

southwestern South Africa, which flowers from Decem-

ber to February when the leaves are dry and brown, only

producing new foliage leaves in May. M. pearsonii has a

similar ecology, and the rather leathery leaves are mostly

dry by early summer when flowering takes place. The pale

lilac flowers open in the late afternoon between 1 7:30—

1 8:00, and last for just three hours before withering at sun-

set. The lack of floral scent, absence of evident nectar, and

the prominently positioned anthers, suggest that M. pear-

sonii is adapted for pollination by pollen-collecting bees.

Diagnosis and relationships'. Moraea pearsonii has

a single, long, trailing leaf up to 1 m long, linear and

channelled when fresh but with the margins involute

on drying so that the leaf then appears to be terete. The

corm, deeply buried among the rocky alluvium up to

20 cm below the surface and thus difficult to extract,

has a tunic of coarse, black fibres. The branched inflor-

escences bear numerous sessile lateral flower clusters

of pale lilac, stellate flowers, 20-25 mm in diameter.

These open exceptionally late in the day, and until the

tepals unfold at ± 17:30 the plant is virtually invisible

in the dry, light brown vegetation. Structurally the flow-

ers resemble those of the Hexaglottis group of Moraea

(Goldblatt & Manning 2000), with the style branches

each divided to the base into thread-like arms extend-

ing between the stamens. In addition, the tepals are sub-

equal, with short claws and spreading limbs, and free fila-

ments. The Hexaglottis- type flower is typical of several

species of Moraea , notably among the six, pale yellow-

flowered species of section Hexaglottis (Vent.) Gold-

blatt, previously recognized as the genus Hexaglottis

Vent, until merged in Moraea (Goldblatt 1987, 1998). It

is also known in the yellow-flowered M. nana (L. Bolus)

Goldblatt & J.C. Manning of section Tubiflora Gold-

blatt; and in the taxonomically isolated, blue-flowered

Namibian M. hexaglottis Goldblatt of section Moraea

(Goldblatt 1986a). Based on vegetative morphology, we

conclude that M. pearsonii is not allied to any of these

species but belongs in section Polyanthes Goldblatt, and

more precisely with the blue- to lilac- or mauve-flowered

M. crispa alliance of western South Africa. Within this

group, it is perhaps most closely allied to M. pseudospi-

cata, which has a similar branching pattern with sessile

rhipidia, similar corm tunics of coarse black fibres, and

it also flowers late in the season when the leaves are ±

dry (Goldblatt 1986a). The style branches of this species

are more typical of Moraea, being flattened and forked

only at the tips and the filaments are united basally.

Bothalia 39,1 (2009)

3

FIGURE 1 . — Moraea pearsonii , Man-

ning 3128 (NBG): A, corm and

flowering stem; B, flower with

two tepals removed; C, inner

(left) and outer (right) tepals;

D, style branches; E, capsules;

F, seed. Scale bar: A, E, F, 10

mm; B, C, 5 mm; D, 2.5 mm.

Artist: John Manning.

History : the species was first collected by H.H.W. Pear-

son, Professor of Botany at the South African College

(later the University of Cape Town) in late November

1908 during an expedition from Cape Town to Namibia.

Pearson collected just the inflorescence branches of a sin-

gle plant, in bloom close to the hottest time of the year.

Although the floral spathes were dry at the time of col-

lection, the rhipidia (flower clusters) bore flowers that

had wilted that day or the previous one. His notes on

the sheet (Pearson 4810 ) indicate that the flowers were

4

Bothalia 39,1 (2009)

FIGURE 2. — Known distribution of Moraea deserticola, A; M. pear-

sonii. A; M. tanquana , •; M. vesper! ina, O; and M. longipes ,

blue, and examination of the withered flowers revealed

that they had free filaments and slender style arms, thus

unlike most Moraea species, which have the filaments

united below and flattened style arms. More than that

could not be determined from the available material and

it remained impossible to identify the plant or even to

draw up a satisfactory description. In December 2007,

we mounted an expedition to try and locate the species,

collected northeast of Ceres, between Hottentotskloof

and Karoopoort. A small population of plants matching

Pearson’s original collection were located in just this

area and confirmed that they represented an undescribed

species.

Additional specimen examined

WESTERN CAPE. — 3319 (Worcester): roadside between Hotten-

tots Kloof and Karoo Poort, (-BA), 29 November 1908, Pearson 4810

(K).

2. Moraea tanquana Goldblatt & J.C. Manning,

sp. nov.

Plantae ad 0.5 m altae ex cormo subgloboso, tunicis

ex fibris crassis atrobrunneis usque nigris constantibus,

foliis 3 infimo basali lineari canaliculato 2-5 mm lato

superioribus caulinibus, caule usque 3-ramoso, spathis

herbaceis ad apicem siccis attenuatis interiore ± 30 mm

longa, floribus pallide caeruleis vel malvinis cum cupula

flava, tepalis exterioribus obovatis ± 25 mm longis lim-

bis 15-16 x 12 mm, interioribus ± 22 mm longis, fila-

mentis ± 10 mm longis in columnam connatis, antheris

contiguis ad anthesin 5 mm longis postea ± 3 mm longis

luteis, ramis styli ± 1 mm longis antheris obtectis, lobis

stigmatis bifidis sine cristis.

TYPE. — Northern Cape, 3219 (Wuppertal): Tankwa

National Park, small koppie east of Leeuberg, 404 m, (—

BB), 4 August 2006, Steyn 872 (NBG, holo.; PRE, iso.).

Plants up to 0.5 m. Corm subglobose, ± 20 mm diam.;

tunics of coarse, dark brown to blackish fibres. Stem up

to 3-branched from upper nodes; branches subtended

by sheathing, attenuate, bract-like cauline leaves. Cata-

phylls pale and membranous. Leaves 3, lowermost basal,

linear and channelled, 2-5 mm wide, ± three quarters

as long as stem, upper 2 leaves cauline, ± as long as the

basal. Spathes green but dry and membranous apically

with brownish cusps; inner ± 30 mm long, outer ± half

as long as inner. Flowers pale blue or mauve with a yel-

low cup, and small, rounded, yellow nectar guides on all

tepals at mouth of cup, tepal claws minutely puberulous

at base, 8-9 mm long, ascending and forming wide cup,

limbs spreading horizontally; outer tepals obovate, ±

25 mm long, limb 15-16 x 12 mm, inner tepals slightly

shorter, ± 22 mm long. Stamens with filaments ± 10

mm long, reaching to mouth of floral cup or exserted

± 1 mm, united, column cylindrical, minutely puberu-

lous at base; anthers erect, contiguous, 4-5 mm long at

anthesis but shrinking to 3 mm after dehiscence, yellow,

apressed to and concealing style. Ovary ovoid, 4—5 mm

long, exserted from spathes, uniformly pale green; style

branching at top of filament column, branches ± 1 mm

long, concealed by anthers; stigma lobes bifid, crests

lacking. Capsules and seeds unknown. Flowering time'.

August, probably also in late July. Figure 3.

Distribution and ecology : Moraea tanquana is known

from a single hill in the Tankwa Karoo National Park

east of Leeuberg (Figure 2). The colony was found on a

dolerite outcrop and comprises numerous plants.

Diagnosis and relationships'. Moraea tanquana is

most likely allied to M. deserticola from the Knersvlakte

and the more widespread M. speciosa of the Western

Karoo (Goldblatt 1986b). All three have upright stems

bearing multiple leaves, branched stems, cup-shaped,

pale blue to mauve flowers with nearly equal tepals, con-

tiguous anthers carried on a slender, cylindrical filament

column and style branches lacking crests. They differ in

several vegetative and floral characteristics (Table 1).

M. speciosa, most robust of the three, has several, broad

foliage leaves up to 40 mm wide, tepals 35-45 mm

long, anthers (8-) 12-1 7 mm long, and the style divides

between the middle and apex of the anthers; when fully

extended the style branch tips typically exceed the

anthers. Smaller M. deserticola has narrow foliage leaves

2-3 mm wide, tepals 30-36 mm long, anthers 6. 0-6. 5

mm long (shrinking after anthesis to 5 mm) that remain

contiguous around the style. The style itself divides at or

just beyond the anther tips into short branches ± 1 mm

long that are carried above the anthers. In contrast, M.

tanquana has foliage leaves up to 5 mm wide, tepals ±

22 mm long, short anthers, 4—5 mm long, that exceed the

style? and enclose them. The style divides opposite the

lower third of the anthers so that the stigmatic tips of the

short style branches, ± 1 mm long, emerge from between

the middle of the ± contiguous anthers. The flower of M.

tanquana most closely resembles that of M. deserticola

in general aspect, but its shorter anthers are held beyond

the floral cup, whereas the anthers are longer in M

deserticola and their bases are retained within the floral

cup.

Vegetatively Moraea deserticola also differs from

M. tanquana in its ± membranous spathes, the longer

inner spathe 3CM10 mm long, whereas in M. tanquana

the spathes are green with dry attenuate tips and the

Bothalia 39,1 (2009)

5

FIGURE 3. — Moraea tanquana, Steyn 872 (NBG): A, corm and flower-

ing stem; B, stamens and style; C, style branches. Scale bar: A,

10 mm; B, 4 mm; C, 2.5 mm. Artist: John Manning.

inner spathe is ± 30 mm long. The ovary is conspicu-

ously veined with dark red in M. deserticola thus

unlike the uniformly green ovary of M. tanquana.

Edaphically the two also differ: M. deserticola occurs

in the Knersvlakte to the northwest and favours light,

loamy clay surrounding limestone outcrops, quite dif-

ferent from the dolerite outcrops and heavy clay soils

in which M. tanquana grows.

3. Moraea longipes Goldblatt & J.C. Manning, sp.

nov.

Plantae 100-180 mm altae, caule eramoso supra ter-

rain bene extenso, cormo globoso 15-20 mm diam.,

tunicis ex fibris crassis pallidis constantibus obtecto,

foliis 2 (spathis extends exclusis) suboppositis rectis

45-85 mm longis canaliculatis marginibus laevigatis vel

parce ciliatis, rhipidium floribus nonnullis; spathis 50-

70 mm longis, subaequalibus vel spatha externa paulo

maiore, floribus pallide flavis vel ± albis tepalis intemis

cupreis, tepalis extemis ± 30 mm longis unguibus ± 10

mm longis munitis, limbis ± patentibus, tepalis intemis

± 28 mm longis, filamentis 7-9 mm longis basin versus

per ± 1.5 mm connatis, antheris 7. 7-9.0 mm longis, stylo

± 1 mm longo, ramis styli 13-16 mm longis, cristis 8-10

longis, anguste triangularibus.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Nama-

qualand, 6 km NW of Bitterfontein on Kotzesrus road,

in stony granitic ground, (-AA), 1 June 2008, Manning

31 72 (NBG, holo.; MO, iso.).

Plants 100-180 mm high, base surrounded by a well-

developed fibrous collar. Corm globose, 15-20 mm

diam., tunics of ± straw-coloured, coarse fibres often

with thickened vertical ridges. Stem unbranched, con-

sisting of single intemode extending from corm to well

above ground level. Leaves 2 (excluding opposed leaf-

like spathes enclosing flowers), subopposite, inserted at

base of spathes, suberect, straight, 45-85 mm long, chan-

nelled, paler green adaxially; margins smooth or sparsely

ciliate. Rhipidium several-flowered; spathes subequal, or

outer slightly larger, clasping inner for ± half its length,

50-70 mm long, inner with broad, transparent membra-

nous margins. Flowers pale yellow or translucent white

with pale copper inner tepals, margins of outer tepals and

on reverse of tepals; nectar guide yellow speckled with

black dots; tepals unequal, outer ± 30 mm long, claw

suberect, ±10 mm long, hairy in midline, limb ± 20 x

19 mm, laxly spreading, hairy toward base, inner tepals

± 28 x 2-4 mm, claws suberect, ± 9 mm long, limbs

spreading. Stamens with filaments suberect, 7-9 mm

long, united basally for ±1.5 mm; anthers apressed to

style branches, 7.5-9 mm long, reaching base of stigma

lobes. Ovary elongate, fertile part ovoid, 5-9 mm long,

usually at least partly included; style dividing ± 1 mm

above base, branches ascending, 13-16 mm long; stigma

lobe rounded; crests erect, narrowly triangular, outer

margins slightly and unevenly serrated, 8-10 mm long,

2.5-3 mm wide at base. Capsules and seeds unknown.

Flowering time : late May to early July. Figure 4.

Distribution and habitat : Namaqualand, from Spring-

bok in the north to the Gifberg Flats in the south; on gra-

nitic or sandy gravel flats (Figure 2).

Diagnosis and relationships', a member of section

Acaules, Moraea longipes has the general aspect of M.

ciliata and M. tricolor except that the stem is not sub-

terranean at flowering but extends up to 180 mm above

the ground. Like most species of the section, the ovary

is retracted to the base of the inflorescence spathes after

6 Bothalia 39,1 (2009)

TABLE 1. — Comparison of flowers of Moraea tanquana, M. deserticola and M. speciosa. All measurements were taken from fresh material.

Anthers measured before anthesis

flowering. The flowers are most like those of M. ciliata

but have slightly broader style crests, triangular with the

base 2. 5-3.0 mm wide (± linear in M. ciliata ) and the

filaments are united basally for ±1.5 mm, about one fifth

of their length. In other species of the section, the fila-

ments are united for one third to half their length. The

anthers of M. longipes are also the longest in the section,

measuring 7. 5-9.0 mm. A unique feature in the section

is the collar of fibres around the underground part of the

stem, this alone making M. longipes easily recognized

even without flowers. The species flowers early in the

season, May to early July, and grows in places where

M. ciliata has been found in bloom six to eight weeks

later. Flower colour in M. longipes is either pale, watery

yellow or translucent white, then with the edges of the

outer tepal limbs and the inner tepal limbs a bright cop-

per colour and with the tepals flushed deep copper on

the reverse. Like other members of section Acaules, the

flowers are fugaceous and last just one day.

Examining the ovary of Moraea longipes closely, we

noted that there is no line of abscission at its base as in

most other species of Moraea. After examining other

members of the section, we found they too lack this

abscission line, which elsewhere in the genus is the point

at which unfertilized ovaries are abscised. We tentatively

suggest that the so-called pedicel of the flower in section

Acaules may actually be part of the ovary and that the

flowers then lack a true pedicel. In freshly open flowers

of M. ciliata and M. tricolor , the ovary stalk is hollow

and contracts in faded flowers, then becomes closely

wrinkled as the tissue collapses on itself, then draws the

ovule-containing part of the ovary into the lower part of

the spathes. Anatomical comparison of true pedicels of

Moraea species with those of the ovary stalk of section

Acaules may solve this question.

Additional specimens examined

NORTHERN CAPE. — 2917 (Springbok): Sanagas, near Spring-

bok, (-DC), 4 June 1980, Dryfhout 2788 (NBG). 3117 (Lepelfontein):

Namaqualand, 15 km SE of Kotzesrus, Farm Biesiesfontein, in stony

granitic gravel, (-BB), I June 2008, Manning 3173 (NBG).

WESTERN CAPE. — 3018 (Vanrhynsdorp): Matsikamma, (-DB),

16 June 1983, Snijman 714 (NBG).

4. Moraea albicuspa Goldblatt in Annals of the

Missouri Botanical Garden 64: 230 (1973). Type: South

Africa, [KwaZulu-Natal], Drakensberg, source of the

Tina River, March 1904, Galpin 6846 (BOL, holo.!;

PRE!, SAM!, iso.).

M. jarmilae J.J.Halda: 69, fig. 79 (2002). Type: Lesotho, Drakens-

berg in the vicinity of Ox Bow, ±3 100 m, 10 March 1989, J.J. & J.

Haldovi s.n. PR4710 (PR, holo.!), syn. nov.

Moraea jarmilae was described for plants from Ox

Bow in Lesotho that had unusual fringed edges to the

outer tepals, which were otherwise ± lanceolate in shape.

The inner tepals were described as tricuspidate and

greenish (Halda 2002) and the species was likened to

the Mpumalanga and northern KwaZulu-Natal species,

M. pubiflora, with the notable exception of the fringed

tepal limbs, but M. pubiflora has the outer tepals velvety

on the reverse, a feature not noted for M. jarmilae. The

type specimen, however, clearly shows the inner tepals

to be linear and apically undivided. Tepals of this shape

are known among Drakensberg Moraea species only in

M. albicuspa, which has white flowers, the outer tepal

claws with a linear, yellow nectar guide. We are confi-

dent that M. jarmilae represents plants with abnormally

formed outer tepal limbs, possibly due to foraging by an

insect while in bud. We accordingly reduce the species

to synonymy in M. albicuspa, which has previously been

collected in Lesotho from Sehlabathebe, southeast of Ox

Bow, and is known from Giant’s Castle in the central

high Drakensberg of KwaZulu-Natal southwards along

the escarpment to Engcobo in Eastern Cape.

RANGE EXTENSIONS AND MISCELLANEOUS NOTES

Moraea bipartita L. Bolus

This widespread species extends from the Swellendam

District in the west through the Little Karoo and part of

the southern Cape to Uitenhage in the east (Goldblatt

1986a). Over much of its range, plants have blue to vio-

let flowers of the standard Moraea- type, thus with larger

outer tepals bearing nectar guides at the base of the limb,

filaments united for half their length, and a style divid-

ing at the apex of the filament column into three peta-

loid branches that terminate in prominent, paired, erect

crests. The stigmatic lobe lies at the base of the crests on

the abaxial surface of the style branch, and in M. bipar-

tita is bilobed, as in many species of the genus. The spe-

cies is distinguished among its allies in section Polvan-

thes by the presence of several (at least three) channelled

foliage leaves, a branched stem, and relatively small

flowers. Plants with yellow flowers discovered by ecolo-

gist J. Vlok in the Perdebont Valley of the Little Karoo

near the northern end of Robinson Pass south of Oudts-

hoom, seemed worth investigating in view of the other-

Bothalia 39,1 (2009)

7

FIGURE 4. — Moraea longipes, Manning 3172 (NBG): A, corm. B,

flowering stem and flowers. Scale bar: A, 10 mm; B, 4 mm.

Artist: John Manning.

wise universal blue to violet flower colour not only in

M. bipartita but in almost all members of section Poly-

anthes ( M. polyanthos L.f. itself may have blue-mauve

or white flowers). The yellow-flowered plants resembled

typical M. bipartita so exactly when compared with typi-

cal blue-flowered plants found elsewhere the same day,

that we were forced to conclude that colour was the only

difference between these plants, growing on the Farm

Saffaanrivier ( Goldblatt et al. 12955 ) and typical M

bipartita. Moraea bipartita then, may rarely have yellow

flowers, a striking exception for section Polyanthes.

At the Saffaanrivier site, yellow-flowered Moraea

bipartita grew together with closely related M. polyan-

thos, a species distinguished from M. bipartita by flow-

ers with subequal tepals, all with small nectar guides,

and reduced style branches as wide as the anthers and

without vestigial style crests. Hybrids between the two

species were common and represent one of the few

examples of a hybrid swarm in Moraea. A range of

intermediate flower forms were represented among the

hybrids, the most common of which had yellow or grey

tepals with blue style branches, narrower than normal

with smaller crests ( Goldblatt et al. 12955A).

WESTERN CAPE. — 3321 (Ladismith): northern foot of the Out-

eniqua Mountains in Perdebont Valley, Farm Salfaanrivier, (-CC), 6

September 2007, Goldblatt, Vlok & Porter 12955 (NBG). Hybrids with

M. polyanthos : loc. cit., Goldblatt, Vlok & Porter 12955A (NBG).

Moraea elsiae Goldblatt (1986a)

Moraea elsiae is one of several species of subge-

nus Visciramosa, an alliance distinguished by woody,

rather than fibrous corm tunics, multiple foliage leaves,

branched stems always viscid below the nodes, and

flowers with filaments free but contiguous around the

style (elsewhere in the genus the filaments are entirely

connate or united in the lower half). Most species of the

subgenus have typical Moraea- type flowers with outer

tepals larger than the inner and with nectar guides at

the base of the limbs, and flattened style branches bear-

ing paired, petaloid crests. M. elsiae is an exception in

having narrow style branches with vestigial crests and

subequal tepals, all of which have nectar guides. The

recorded range of M. elsiae is from the Cape Peninsula

to the Potberg near Cape Infanta. The presence of the

species has now been established in the Pauline Bohnen

Reserve at Stilbaai, a range extension of some 100 km

eastward.

WESTERN CAPE. — 3421 (Riversdale): Stilbaai, Pauline Bohnen

Reserve, (-AD), 20 October 2000, De V. Pienaar & Pienaar PB570

(NBG).

Moraea falcifolia Klatt

One of the most widespread of the winter rainfall

Moraea species, M. falcifolia has been recorded from

southern Namibia through Namaqualand south to the

Breede River Valley and the western Little Karoo, as

well as from Bushmanland and the Upper Karoo as far

east as Kimberley. In Goldblatt’s (1976, 1986a) revi-

sions of the genus, no records were known from Eastern

Cape or the central and eastern Little Karoo. Collections

have now been made from the Little Karoo near Sebras-

fontein in the northern foothills of the Outeniqua Moun-

tains, in the Long Kloof, and from near Alexandria, thus

8

Bothalia 39,1 (2009)

enlarging its already wide range significantly. Plants

from these sites do not differ in any significant way from

known collections.

WESTERN CAPE. — 3322 (Oudtshoom): low northern foothills

of the Outeniqua Mountains near Sebrasfontein, (-CC), 13 July 1986,

Vlok 1518 (MO).

EASTERN CAPE.— 3323 (Willowmore): Onder Kouga, off Long

Kloof, (-DD), 28 September 1975, Bayliss BS7098 (PR E). 3326 (Gra-

hamstown): Salem- Alexandria road, Bushmans River Gorge, Long-

ford Grange Farm, (-CB), 26 August 1995, Dole! & Cocks 1771 (GRA;

MO, photo.).

Moraea pseudospicata Goldblatt ( 1986b)

When described, this late summer- and autumn-flow-

ering species was known with certainty only from the

Nieuwoudtville Wildflower Reserve and from Loken-

burg, 30 km south of Nieuwoudtville. In March 1997,

the species was found on the Nieuwoudtville Sports

Ground and in March 2000 we found it to be common on

Glenlyon Farm, now the Hantam National Botanic Gar-

den, on soils derived from Dwyka tillite. Then in 2003

we discovered large numbers of what appeared to be M.

pseudospicata in fruit on the Hantamsberg at Calvinia in

areas that had been burned two years earlier. While flow-

ers are not known for the Hantamsberg plants, they have

the sessile lateral inflorescences, single linear leaf with

tightly inrolled margins, and small, globose capsules that

are diagnostic for M. pseudospicata. The species now

seems to be common north and south of Nieuwoudtville

in a belt along the western end of the Bokkeveld Plateau

as well as inland on the Hantamsberg. The reasons for its

apparent rarity are not only that it flowers in February,

March and April when little collecting is done in this hot

and summer-dry part of the Western Karoo, but also that

the flowers only open late in the afternoon, after 16:00

and are until then virtually invisible except to the spe-

cialist collector. We suspect that M. pseudospicata has

an even wider range than is currently documented.

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville Sports

Ground, (-AC), 31 March 1997, Van Rooyen, Steyn & De Villiers 411

(NBG, PRU); Glenlyon Farm, renosterveld near reserve at entrance to

farm property, (-AC), 13 March 2000, Goldblatt 11305 (MO); Glen-

lyon Farm, renosterveld on road to Camel Koppie, (-AC), 13 March

2000, Goldblatt 11305 (MO); Hantamsberg slopes, (-BD), 4 Septem-

ber 2002 (late fr.), Goldblatt & Porter 12163 (MO, NBG, PRE).

Moraea spathulata (L.f.) Klatt

A widespread, eastern southern African species, M

spathulata extends from George in the southern Cape

through eastern South Africa and Swaziland to eastern

Zimbabwe, and is typically found in well-watered, mon-

tane habitats except at the extreme southern end of its

range between George and Humansdorp where it occurs

close to the coast. Exploration in the high mountains

fringing the Great Karoo by E.G.H. Oliver in the 1970s

and later by Ralph Clarke in 2005-2007, have resulted

in the documenting of the species on the Koudeveld-

berge near Graaff-Reinet. Plants from the mountains of

the Karoo differ in no way from those in the south at

Knysna and Humansdorp, nor from those in the Drak-

ensberg, despite the drier, and in winter, colder habitat.

They constitute a remarkable range extension.

EASTERN CAPE. — 3224 (Graaff-Reinet): Koudeveldberge SE of

Doombosch, (-AA), frequent scattered clumps on summit plateau and

north slopes with low tussock grass, 6 November 1974, Oliver 5221

(NBG). 3225 (Somerset East): Sneeuberg above Suurkloof, Asante

Sana Private Game Reserve, 1 800 m, (-AC), 6 December 2005,

Clarke & Coombs 155 (GRA).

Moraea tricolor Andrews

Until now, this southwestern Cape species has been

recorded from Darling, the Malmesbury and Tulbagh

Districts, historically from the Cape Peninsula (where it

is now extinct), and locally in the Caledon District as far

east as Napier (Goldblatt 1986a). A member of section

Acaules Goldblatt (5 spp.), it has the acaulescent habit

characteristic of the group, with flowers borne above the

ground and then withdrawn by contractions of the stalk of

the ovary to lie within the spathes and near ground level

where the fruits mature. Within the section, M. tricolor

is recognized by the broad, triangular style crests, 8-10

mm long and 4—5 mm wide at the base, usually with the

filaments united for half their length, and often glabres-

cent foliage. Flower colour in M. tricolor ranges from

pale yellow to pink, brick-red, violet or deep mauve-pink,

always with prominent deep yellow nectar guides edged

in darker pigment, often red or purple, at the base of the

outer tepal limbs, hence the specific epithet tricolor. Col-

lections from Stilbaai in the southern Cape ( Naude s.n.),

where four separate populations are known (J. Naude

pers. comm. 2008), represents a range extension of some

150 km. These plants are also remarkable for flowering

in May, whereas other populations of M. tricolor typi-

cally bloom in August and September, although plants

have been collected in flower near Hopefield in June.

The habitat of the Stilbaai populations, which grow in the

well-drained sandy ground of coastal dunes, is unusual

for the species, which typically favours seasonally moist

to inundated, poorly drained sandy or clay flats. Despite

the ecological shift and change in flowering time of the

Stilbaai populations, we find no character of any taxo-

nomic significance that permits their recognition. For the

present we record the range extension and note the shift

in ecology and flowering time.

Fruiting plants from Stilbaai ( Goldblatt , Manning

& Naude 11199, MO, NBG) have capsules 9-12 x ± 6

mm, and light brown, angular to subglobose seeds, ±1.4

x 2 mm, with reticulate sculpturing. Capsules from one

population of southwestern Cape Moraea tricolor, not

recorded before, are 7-11 mm long and the seeds are

also angular to subglobose with reticulate sculpturing

and 1 x ± 1.3 mm ( Goldblatt 11572, MO). These seeds

match those of most other species of the section except

for small differences in size and are probably the plesio-

morphic type for the section.

We also include a minor range extension for Moraea

tricolor from Langebaan on Saldanha Bay, the northern-

most record for the species ( Goldblatt & Porter 13066

(fl.), MO, NBG, 13210 (fr.), MO, NBG). Plants at this

site have violet tepals with an orange rather than yellow

nectar guide, colouring not before reported for the spe-

cies. The capsules are ± 10 mm long, typical for the spe-

cies.

Capsules and seeds of populations of Moraea ciliata

(e.g., Goldblatt & Porter 12710, MO, 12691, MO) with

yellow or white flowers, are similar in size and shape to

those of M. tricolor, the seeds rather more angular and

Bothalia 39,1 (2009)

9

often 5-sided, obscurely reticulate and ± 1 x 1.5 mm.

Curiously, blue-flowered populations of the species that

we have examined (e.g. Snijman & Perry 2142 , NBG, in

flower) have larger, globose seeds with less pronounced

sculpturing and 2 x 2-3 mm, and are borne in elongate

capsules 18-20 mm long ( Goldblatt et al. 12863 , MO).

The variation in capsule and seed shape and size in col-

our morphs of M. ciliata , suggests differentiation among

populations of the species associated not only with peri-

anth colour but also capsules and seeds, and perhaps

other features. Insufficient collections with capsules and

fully developed seeds make more detailed exploration of

this question impossible at this time.

WESTERN CAPE. — 3318 (Cape Town): Langebaan, wet site in

granite outcrop opposite post office, (— AA), 3 September 2008, Gold-

blatt & Porter 13066 (MO, NBG), 28 September 2008, Goldblatt &

Porter 13201 (fr.) (MO, NBG). 3421 (Riversdale): Stilbaai, Geelkrans

Nature Reserve, (-AD), May 2006, Naude s.n. (NBG), 5 September

2007 (fr.), Goldblatt , Manning & Naude 12950 (MO, NBG); Stilbaai,

Panorama Circle, 29 May 1980, P. Bohnen 7542 (NBG); Stilbaai,

grounds of The Gem Nursery, road from Stilbaai to Jongensfontein, 24

May 2003, U. de V. Pienaar & A. Pienaar 378 (NBG).

Moraea vegeta L.

Locally common on the Cape Peninsula and recorded

as far north as Darling and Hermon in the greater Malm-

esbury District, and as far east as Swellendam (Goldblatt

1986a), Moraea vegeta is a fairly inconspicuous species,

with watery yellow or dull purple flowers. It is readily

recognized by the several channelled, glaucous leaves

and nodding, globose capsules with soft walls that show

the outlines of the seeds. A new record from the Kobee

Valley, some 250 km to the north is an unexpected and

remarkable range extension. In the Kobee Valley, plants

grow in sheltered kloofs and gullies in open bush domi-

nated by wild olive ( Olea europaea subsp. africana ) on a

west-facing slope above the valley floor. This is the only

record of the species from the northwestern centre of the

Cape flora region.

WESTERN CAPE. — 3119 (Calvinia): Kobee Valley, gully on west-

facing slopes above valley floor in light bush, (-CA), 1 September

2001, Goldblatt & Porter 11801 (NBG).

Moraea verecunda Goldblatt

Although currently known from a few sites in the

immediate vicinity of Nieuwoudtville (Goldblatt 1986a),

specimens collected from south of the Bokkeveld Moun-

tains, on Uitkyk Pass descending into Biedouw Valley,

appear to represent this species. The collection ( Stirton

11507) has sessile lateral inflorescences, a solitary, nar-

row basal leaf, and small violet flowers with tepals 10-

12 mm long, and partially fused filaments. No capsules

are fully developed but the ovaries are fusiform and

one immature capsule is beaked. These features accord

exactly with M. verecunda and we tentatively treat this

as a range extension for the species. A photograph (R.

MacFarlane pers. comm. 2008) taken on the mountain

slopes ± 3 km NW of Wuppertal on the road to Biedouw

Valley on 24 September 2006, also appears to be M.

verecunda. Plants should be checked at these new locali-

ties, which represent the first records of the species in

Western Cape and 80 and 100 km from Nieuwoudtville.

WESTERN CAPE. — 3219 (Wuppertal): 11 km from Pakhuis Pass

to Biedouw Valley, descending down Uitkyk Pass, (-AA), 1 1 Novem-

ber 1986, Stirton 11507 (NBG).

Moraea vespertina Goldblatt & J.C. Manning

Described in 2000, based on plants from the dolerite

hills east of Nieuwoudtville in Northern Cape, Moraea

vespertina was distinguished in subgenus Visciramosa

Goldblatt by its several foliage leaves and relatively

large, white flowers opening in the late afternoon and

fading shortly after sunset. Subsequently, a second popu-

lation was reported from Matjiesfontein Farm, lying to

the east, between Nieuwoudtville and Calvinia ( Manning

s.n.). The flowers of this population are slightly larger

(the outer tepals are ± 45 mm long, filaments 10 mm,

anthers 6-7 mm) (Table 2) and on fading become faintly

suffused with grey blue. In 2006 and 2007 we collected

plants resembling M. vespertina in the Tankwa National

Park, but with larger, pale grey-blue flowers. At first we

considered that they represented a new species, distin-

guished from M. vespertina by the larger perianth, blue-

grey coloration, crescent-shaped nectar guide, purple

markings and veins on the outer tepal claws and other

floral details (Table 2), notably the longer filaments,

anthers and style crests. The Tankwa population grows

among dolerite boulders on south-trending slopes. Dif-

ferences between the populations are relatively small and

at present we prefer to expand the circumscription of M.

vespertina rather than to recognize a new taxon. We sus-

pect that more populations of this species remain to be

discovered between the Bokkeveld Plateau sites and the

Tankwa National Park, over 120 km to the south (Figure

TABLE 2. — Comparison of Nieuwoudtville and Tankwa National Park populations of Moraea vespertina. All measurements were taken from fresh

material

10

Bothalia 39,1 (2009)

2) and if so, will no doubt cast more light on questions

about the taxonomic status of the southern, larger-flow-

ered population.

The flowers of the Tankwa population opened at

17:00-17:30 and closed at ± 17:30. When fully open,

as the sun began to set, they produced a strong odour of

stocks (a heavy, sweet, clove scent). As night fell, we

observed settling moths (not captured) visiting the flow-

ers. The strong scent, pale flower colour and timing of

anthesis make it all but certain that the Tankwa popula-

tion is adapted for pollination by settling moths, as was

previously reported for the Nieuwoudtville plants (Gold-

blatt & Manning 2000).

In a curious aside, we report that the corms of Moraea

vespertina are boiled in goat’s milk and eaten locally

in the Calvinia District as a delicacy (I. Coetzee pers.

comm. 2007). This leads us to think there are more sites

for the species, but unfortunately not yet recorded.

NORTHERN CAPE. — 3119 (Calvinia): Farm Matjiesfontein

between Calvinia and Nieuwoudtville, (-AD), 20 October 2007, Man-

ning s.n. (NBG). 3220 (Sutherland): Tankwa National Park, east end

of Elandsberg, (-AA), 16 September 2006, Manning 3060 (NBG);

Tankwa National Park, kloof along Maansedam road to Elandsberg,

(-AA), 10 September 2007, Goldblatt & Porter 12978 (MO, NBG).

Moraea vlokii Goldblatt

When described (Goldblatt 1992), Moraea vlokii

was known from two collections in the Swartberg near

Gamkakloof at fairly high elevations, one at ± 985 m

and the other at ± 490 m. We found a third population

in 1997 near Montagu, which establishes a wider range

for the species and, though still in the Little Karoo, at

a lower elevation, below 300 m and some 150 km dis-

tant. The species is vegetatively very like the widespread

M. gawleri Spreng. in corm tunic structure and the unu-

sual, short inflorescence spathes, although it has a sin-

gle foliage leaf (M gawleri normally has two or three

foliage leaves), and Goldblatt postulated that the two

were immediately allied. DNA sequence analysis using a

single sample of each species shows the two to be sister

taxa (Goldblatt et al. unpublished data).

WESTERN CAPE. — 3320 (Montagu): below Ouberg Pass to Mon-

tagu, rock outcrop, ± 260 m, (-CC), 27 September 1997, Goldblatt &

Manning 10764A (MO, NBG).

Moraea sp.

A collection from the summit of the Aurusberg in

southwestern Namibia, Williamson & Hamer 4564,

NBG, found on 2 November 1992, appears to repre-

sent a new species. The flowers, said to be yellow, were

destroyed by insects before we received the specimen

but it is worthwhile describing the plants for future refer-

ence and to establish its existence. Plants appear to grow

in tufts in rock crevices on steep south slopes, and have

corms ± 1 2 mm in diameter and are covered by medium-

textured, dark fibres, much like those illustrated for M.

tanquana (Figure 3). The stems, about 60 mm long, are

unbranched, trailing and bear several, channelled leaves,

up to 1 mm wide. The blades are somewhat twisted,

have undulate margins and like the stem, they are trail-

ing, but are much longer than the stem. The spathes are

almost equal, the inner 13-14 mm long and the outer ±

12 mm long. Nothing remains of the flowers or buds,

but apart from the colour, were said to be Spiloxene- like,

this with at least partially free filaments and thread-like

style branches. We speculate that the flowers were of the

Hexaglottis- type as illustrated in Figure 1.

ACKNOWLEDGEMENTS

Support for this study by grants 7316-02, 7799-05 and

8248-07 from the National Geographic Society is grate-

fully acknowledged. We thank Elizabeth Parker and Len-

don Porter for assistance and companionship in the field;

Jan Vlok for alerting us to the existence of yellow-flow-

ered Moraea bipartita and M. tricolor at Stilbaai; Ralph

Clarke for alerting us to the presence of M. spathulata in

the Karoo highlands; Tony Dold for providing informa-

tion about the occurrence of M. falcifolia near Alexan-

dria; Janet Naude for guiding us to populations of M. tri-

color, Roger MacFarlane for the Wuppertal record of M.

verecunda; and Sharon Bodine for help with herbarium

searches. Roy Gereau kindly checked the Latin descrip-

tions. Collecting permits were provided by the nature

conservation authorities of Western Cape and Northern

Cape Provinces, South Africa.

REFERENCES

GOLDBLATT, R 1973. Contributions to the knowledge of Moraea (Iri-

daceae) in the summer rainfall region of South Africa. Annals of

the Missouri Botanical Garden 60: 204—259.

GOLDBLATT, P. 1 976. Evolution, cytology and subgeneric classifica-

tion in Moraea (Iridaceae). Annals of the Missouri Botanical

Garden 63: 1-23.

GOLDBLATT, P. 1986a. The moraeas of southern Africa. Annals of

Kirstenbosch Botanic Gardens 14: 1-224.

GOLDBLATT, P. 1986b. Convergent evolution of the Homeria flower

type in six new species of Moraea (Iridaceae) in southern Africa.

Annals of the Missouri Botanical Garden 73: 102-116.

GOLDBLATT, P. 1987. Systematics of the southern African genus

Hexaglottis (Iridaceae-Iridoideae). Annals of the Missouri

Botanical Garden 74: 542-569.

GOLDBLATT, P. 1992. New species, chromosome cytology and notes on

the southern African Iridaceae-Irideae: Moraea , Roggeveldia and

Homeria. South African Journal of Botany 58: 209-214.

GOLDBLATT, P. 1998. Reduction of Bamardiella, Galaxia, Gynandr-

iris, Hexaglottis, Homeria and Roggeveldia in Moraea (Iridaceae:

Irideae). Novon 8: 371-377.

GOLDBLATT, P. & MANNING, J.C. 2000. New species of Moraea (Iri-

daceae-Iridoideae) from southern Africa. Novon 10: 14—22.

GOLDBLATT, P, SAVOLAINEN, V., PORTEOUS, 0., SOSTARIC,

I., POWELL, M„ REEVES, G., MANNING, J.C., BARRA-

CLOUGH, T.G. & CHASE, M.W. 2002. Radiation in the Cape

flora and the phylogeny of peacock irises Moraea (Iridaceae)

based on four plastid DNA regions. Molecular Phylogenetics

and Evolution 25: 341-360.

HALDA, J.J. 2002. Moraea jarmilae J.J.Halda, spec. nov. Acta Musei

Richnoviensis sect. Natur 9,1 : 69, fig. 79.

Bothalia 39,1: 11-17 (2009)

Three new species of Diascia (Scrophulariaceae) from the Western

Cape, South Africa

K.E. STEINER*

Keywords: Diascia Link & Otto, new species, oil-collecting bees, oil-secreting trichomes, Scrophulariaceae, South Africa, Western Cape

ABSTRACT

Three new annual species of Diascia Link & Otto are described from the Western Cape Province of South Africa. D.

collina is characterized by greyish magenta flowers with two divergent yellow sacs containing oil-secreting trichomes. It

is restricted to granite outcrops in the vicinity of Saldanha Bay, from the West Coast National Park and Langebaan north to

Vredenburg. D. pusilla is closely related to D. collina, but differs from that species in having smaller flowers with shorter,

± parallel sacs, and posticous filaments that lack a protuberance where they bend sharply backwards towards the upper lip.

It occurs in grey to whitish sands usually near seasonally moist or wet areas. It has not been found more than 35 km from

the coast and ranges from Modderrivier, south of Darling, north to Lambert’s Bay. D. appendiculata is related to D. diffusa

(Thunb.) Benth. and is characterized by having small, mainly reddish lilac to greyish magenta flowers, two shallow depres-

sions in the corolla tube at the base of the upper lip, and posticous filaments with sterile appendages. It is known from only

six localities in the general vicinity of Citrusdal and occurs in fynbos vegetation on lower mountain slopes or flats, in loose

alluvial sands derived from Table Mountain Sandstone.

INTRODUCTION

Diascia Link & Otto is a genus of ± 72 species of

annual and perennial herbs endemic to southern Africa.

Two sections have been recognized, section Racemosae

with 27 species and section Diascia with about 45 spe-

cies (Hilliard & Burtt 1984; Steiner unpubl.). Section

Racemosae was revised by Hilliard & Burtt (1984)

and three additional taxa were described more recently

(Steiner 1989, 1999). Section Diascia has not been

revised since Hiem’s (1904) treatment in Flora capensis,

although many new species have been described in recent

years as part of a revisionary study (Steiner 1992a, b, c, d

1995). Section Diascia consists solely of annual species,

whereas section Racemosae is mostly perennial (81 %).

Three additional new species in Section Diascia from

the Western Cape are described here. All descriptions are

based on living material collected from the field. Flower

colours are based on the Methuen handbook of colour

(Komerup & Wanscher 1984). Chromosome counts of

these species were reported by Steiner (1996).

Diascia collina K.E. Steiner, sp. nov., D. pusillae

K.E. Steiner proxima, sed differt corolla grandiore, sac-

culis corollae grandioribus divergentibus non parallelis,

et filamentis posticis protuberatione instructis.

TYPE. — Western Cape, 3318 (Cape Town): Postberg

Nature Reserve, Vlaeberg loop road, picnic and view

site, ± 200 m, (-AA), 14 Sept. 1988, Steiner 1816 (NBG,

holo.; BOL, E, K, MO, PRE, US, iso.).

Annual herb, rosulate, glabrous, simple or branching

from base. Stems decumbent, up to 340 mm long, angu-

lar, up to 6-sided, ribs 2 or more, sides up to 2 mm wide.

Leaves simple, alternate, opposite or whorled, petiolate;

lamina ovate to obovate, 4— 33(-60) * 3-11 (-13) mm,

apex rounded to acute or apiculate, base attenuate; mar-

* Department of Botany, California Academy of Sciences, 55 Music

Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA.

MS. received: 2008-05-29.

gins lobed to divided, lobes or divisions oblong-ovate to

triangular, entire, opposite or subopposite, occasionally

alternate, apices rounded to acute or apiculate; petioles

up to 37 mm long; cauline leaves progressively smaller

upwards. Flowers axillary, 1 or 2 open per stem, faintly

sweet-scented, nodding in early bud stage; flowering pedi-

cels 20-65 mm long, ascending, dorsiventrally flattened

especially where attached to flower, recurving in fruit

except for upward curving apical portion. Calyx lobes

5, spreading, lanceolate, ± equal, 3. 2-3. 6 x 1.4— 1.8 mm,

acuminate, the two lower sepals slightly reflexed; mar-

gins white-ciliate. Corolla bilabiate, 5-lobed, limb 13.3—

23.0 x 14.3-26.0 mm; lobes broadly oblong-obovate,

falciform, outer sides longer than inner sides, 4.4-7. 1 x

5.2-5. 7(-7. 9) mm, apices rounded, bases oblique; lateral

lobes broadly obovate, emarginate, 5.4— 7.1 x 5. 2-6. 8

(-8.3) mm, sides ± equal; lower lobe obcordate, 6. 2-9. 7 x

5.4-8. 1(-1 1.4); upper lobes greyish magenta (14D6) with

deep magenta veins or lines at base; other lobes similar in

colour but without veining, all with scattered, dark pur-

ple, peltate glandular trichomes, especially on inner sur-

face near base; tube shallowly cupped, deep magenta; bi-

saccate, sacs oblong-ovate, rounded, 4—5 x 2. 0-2. 5 mm,

mostly yellow, widely diverging, oil-secreting glandular

trichomes within; central, stamen-bearing boss oblique,

anticous portion 1.1-1. 5 mm high, deep magenta, pos-

ticous portion 0.3-0. 6 mm high, yellow. Stamens 4,

erect, partly hidden; anticous filaments (twisted at base

and appearing posticous) falciform, ± 3.4— 3.6 mm long,

bases strongly curved, pubescent, trichomes clavate, pur-

ple; posticous filaments geniculate, thickened, 1. 9-2.0

mm long, bend with protuberance ± 1 mm from base,

pubescent, trichomes clavate, purple; anthers ± 0.50-

0.80 mm, strongly cohering, grey; pollen orange (fading

to yellow in pressed specimens). Ovary oblong-ovoid,

laterally compressed contrary to septum, 1.6-1. 7 x 1

mm; style ± 1. 6-2.0 mm long, reddish purple, curving

forward in distal third; stigma capitate, surrounded by

anthers; ovules ± 35-45. Capsule ovoid to oblong-ovoid,

5. 1-8.1 x 3. 3-5. 8 mm, exceeding sepals at maturity, base

oblique. Seeds reniform, 1.0-1. 2 mm long, dorsal sur-

face ridged, ventral surface with an oblong keyhole-like

12

Bothalia 39,1 (2009)

FIGURE 1. — Diascia collina, Steiner 2219 (NBG). A, habit. B- D, flower: B, C, front and rear views; D, side view partially cut away. E, calyx; F, pistil; G,

capsule. H, I, seed: H, ventral view; I, side view. Scale bars: A, 10 mm; B, C, 4 mm; D, E, 1 mm; F, 3 mm, G, 2 mm; H, I, 0.5 mm. Artist: Ellaphie

Ward-Hilhorst.

opening formed by extensions of seed coat, long sides of

opening bearing a reniform perforation; embryo curved.

Chromosome no.: 2n = 18. Flowering time: August-Sep-

tember. Figure 1.

Diagnostic features: Diascia collina is most similar

to D. pusilla, but it differs from that species in having a

larger corolla (13.3-23.0 x 14.3-26.0 mm vs 9.1-13.5

x 9.0-14.3 mm), longer corolla sacs (4-5 mm vs 2.2-

Bothalia 39,1 (2009)

13

2.5 mm long) and a protuberance from the posticous fila-

ments where they bend backwards (Figure 1). D. collina

is also similar to D. capensis (L.) Britten, but differs from

that species in having stamens that are ± half the size

and backwards-bending, rather than forward-arching. D.

collina also has a shorter style (1. 3-2.0 mm vs 3. 5-5. 2

mm) that is less curved, and corolla sacs that are strongly

divergent, not ± parallel like those of D. capensis.

Etymology : the name refers to the hills of granite

where the species occurs.

Distribution and habitat : Diascia collina is known

only from the Postberg section of the West Coast

National Park, from undeveloped areas in and around

the town of Langebaan directly across the lagoon from

Postberg, and from the granite outcrops on the south-

ern edge of Vredenburg (Figure 2). It ranges in eleva-

tion from near sea level to about 200 m. In Postberg,

D. collina is fairly common around the Uitkyk pic-

nic area on Vlaeberg ridge overlooking Langebaan

Lagoon. It has also been seen near the entrance to the

SADF restricted area at the northwest end of the Post-

berg Reserve. D. collina occurs under and around

medium to large shrubs and can be considered endemic

to Saldanha Granite Strandveld vegetation (Mucina &

Rutherford 2006). In Langebaan and Vredenburg, this

habitat is quickly disappearing due to residential expan-

sion.

Pollination and breeding system : based on observa-

tions of cultivated plants, Diascia collina is self-incom-

patible and, at Postberg, it is pollinated by two species

of oil-collecting bees, Rediviva peringueyi Friese and

R. aurata Whitehead & Steiner (Melittidae) (Whitehead

& Steiner 2001). These bees use the specially modified

setae on their forelegs to collect oil from the paired yel-

low corolla sacs. The pollen is deposited on the frons or

face of the pollinating bees.

Other specimens examined

WESTERN CAPE. — 3217 (Saldanha): Vredenburg. new housing

development at Witklip near old granite quarry (S32° 55.228’ E17°

58.700’), ± 150 m, (-DD), 16 Sept. 2004, Steiner 4101 (CAS); ibid.,

2 Sept. 2005, Steiner 4121 (CAS, K, NBG. PRE, US). 3318 (Cape

Town): Postberg Nature Reserve, Vlaeberg loop road, picnic and view

site, (-AA), 5 Sept. 1990, Steiner 2219 (NBG). Langebaan Hill, sandy

slopes, (-AA), 24 Aug. 1995, Goldblatt & Manning 10280 (NBG);

ibid., 25 Sept. 1995, Goldblatt & Manning 10321 (NBG); Langebaan.

rocky outcrop near town, (-AA), 23 Aug. 1998, Goldblatt & Manning

10994 (NBG); Langebaan hills, above town, in vacant lot opposite 82

Sunbird Lane (S 33°06.172’ E 18°02.574’), ±18 m, (-AA), 30 Aug.

2001, Steiner 3697 (NBG, CAS); Langebaan, between day care centre

and Pikkieland fun park, (S33°05.461' E18°02.207’), ± 5 m, (-AA), 6

Sept. 2002, Steiner 3870 (NBG, CAS).

Diascia pusilla K.E. Steiner, sp. nov., D. collinae

K.E. Steiner proxima, sed differt corolla breviore, sac-

culis corollae parallelis non divergentibus et filamentis

posticis sine protuberationibus.

TYPE. — Western Cape, 3218 (Clanwilliam): Farm

Droogerivier, road 365, 8.6 km N of turnoff to Alexan-

dershoek, ± 200 m west of road, (-BC), 16 Sept. 1988,

Steiner 1819 (NBG, holo.; K, MO, PRE, iso.).

Annual herb, rosulate, glabrous, simple or branch-

ing from base. Stems decumbent, up to 150 mm long,

FIGURE 2. — Known distributions of Diascia collina, •; D. pusilla, ▲;

andZ). appendiculata, ★.

angular, up to 6-sided, ribs 2 or more, sides up to 2 mm

wide. Leaves simple, opposite or alternate, petiolate;

lamina obovate to elliptic, 5-28 x 2-9 mm, apex acute

to apiculate, base attenuate; margins lobed to divided,

lobes or divisions ovate to triangular, entire, acute to

apiculate; petioles up to ± 12 mm long; cauline leaves

progressively smaller upwards. Flowers axillary, 1

to 3 open per branch, unscented; pedicels 17-23 mm

long, ascending, dorsiventrally flattened especially

where attached to flower, recurved in fruit except for

upward curving apical portion. Calyx lobes 5, spread-

ing, lanceolate, ± equal, 2. 3-2. 9 x 1.0-1. 2 mm, acumi-

nate; margins white-ciliate. Corolla bilabiate, 5-lobed,

limb 9.1-13.5 x 9.0-14.3 mm; lobes broadly ovate;

upper lobes 2. 6-3. 5 x 2. 9-4.0 mm, outer sides longer

than inner sides, apices rounded to emarginate, bases

oblique; lateral lobes 2. 5-3. 5 x 3. 0-3. 5 mm, sides ±

equal, apices rounded; lower lobe 3. 3-5.0 x 3. 2-5. 8

mm, emarginate to obcordate; upper lobes greyish

magenta (14D6) on inner surface, pinkish white (13A2)

on reverse side, with deep magenta (14E8) lines at

base; other lobes similar in colour but lacking lines,

sparsely glandular puberulous with dark violet, glan-

dular trichomes, especially on inner surface near base;

tube shallowly cupped, deep magenta and yellow, bi-

saccate, sacs ovate in outline, 2. 2-2. 5 x 1.2-1 .4 mm,

projecting downward and diverging slightly at tips,

yellow, oil-secreting glandular trichomes within; cen-

tral stamen-bearing boss oblique, anticous portion 1 .2-

1.9 mm high, deep magenta (14E8), posticous portion

0.3-0. 5 mm high, yellow. Stamens 4, erect, partly hid-

den; anticous filaments (twisted at base and appearing

posticous) falciform, 3. 0-3. 2 mm long, bases strongly

curved, sparsely pubescent, trichomes clavate; posti-

cous filaments geniculate, thickened, 2. 0-2. 5 mm long,

bend pubescent, trichomes clavate, dark violet, apical

portions, below anthers, bent forward without enlarge-

ment; all filaments greyish magenta (13E6) except

14

Bothalia 39,1 (2009)

FIGURE 3. — Diascia pusilla, Steiner 1819 (NBG). A, habit. B -D, flower: B, C, front and rear views; D, side view partially cut away. E, calyx; F,

pistil; G, capsule. H, I, seed: H, ventral view; I, side view. Scale bars: A, 10 mm; B, C, 5 mm; D, 2 mm; E, 3 mm; F, 1 mm, G, 2 mm; H, I,

0.5 mm. Artist: Ellaphie Ward-Hilhorst.

just below anthers; anthers 0.3-0. 5 mm long, strongly

cohering, grey; pollen yellow to orange. Ovary oblong-

ovoid, laterally compressed contrary to septum, 1 .2-1.5

x 0. 9-1.0 mm; style 1.3-1. 5 mm long, curving for-

ward at tip; stigma subcapitate, surrounded by anthers;

ovules ± 23-38. Capsule falciform ovoid, 6.0-7. 1 x

3. 5-4. 6 mm, ± twice as long as calyx at maturity. Seeds

reniform, 0.9-1. 2 mm long, dorsal surface ridged,

ventral surface with an oblong, keyhole-like opening

formed by extensions of seed coat, long sides of open-

Bothalia 39,1 (2009)

15

ing bearing a reniform perforation; embryo curved.

Chromosome no.: 2n = 18. Flowering time: August-

September. Figure 3.

Diagnostic features: Diascia pusilla differs from its

nearest relative, D. collina, in corolla size, shape of the

posticous filaments, size and shape of the corolla sacs,

and habitat. The difference in flower size between the

two taxa is not simply a function of plant vigour, since

small plants of D. collina at Postberg, with only a few

leaves, have much larger flowers than robust plants of D.

pusilla, with many large leaves and long thick stems, at

Droogerivier. Corolla limb length of D. pusilla reaches

13.5 mm, but averages 11.5 mm, whereas limb length of

D. collina ranges from 13.3 to 23 mm, but averages 17

mm. The corolla sacs of D. pusilla are about half the size

of D. collina sacs (2.1 mm vs 4.4 mm long) and are par-

allel or only slightly divergent near the tips. They are not

strongly divergent like those of D. collina. The shapes of

the filaments also differ between these two species. D.

pusilla lacks the protuberance on each posticous filament

that is present in D. collina.

Etymology: the name refers to the small size of the

flowers.

Distribution and habitat: Diascia pusilla is known

from a narrow strip along the Cape west coast, from the

Farm Modderrivier (Mud River) (southwest of Darling),

north to Lambert’s Bay and as far east as Droogerivier

(± 4.5 km SE of Sandberg). It occurs no more than 35

km from the sea and ranges in elevation from sea level

to nearly 100 m (Figure 2). On the farms Droogerivier

and Suurfontein (near Lambert’s Bay), it occurs in or

near riverine or vlei systems, often in poorly drained,

seasonally wet sands, but in other localities, the habitat

is drier. In all cases, it occurs in loose white to greyish

sands. Southwest of Darling it has been reported from

short fynbos in deep sand ( Hugo 2427). It does not occur

in areas with granitic outcrops, typical of D. collina.

Pollination and breeding system: like most Diascia

species, D. pusilla has oil-secreting glands in its corolla

sacs. Unlike D. collina , this species is facultatively auto-

gamous and, therefore, does not need visits by oil-col-

lecting bees to set seed. At the type locality, and at least

one other locality, D. pusilla occurs and flowers concur-

rently with Hemimeris racemosa (Scrophulariaceae),

another oil-secreting species. Since at both these sites

oil-collecting bees have been caught on H. racemosa,

it is probable that these same bees visit D. pusilla. At

the type locality, Droogerivier, and at Kersefontein,

D. pusilla occurs with Rediviva parva (Whitehead &

Steiner 2001).

Other specimens examined

WESTERN CAPE. — 3218 (Clanwilliam): Farm Suurfontein

(±13 km E of Lambert’s Bay), 6.8 km E of turnoff to Doringbaai

(S32°07.209' E18°25.710’), ± 50 m, (-AB), 4 Sept. 2001, Steiner 3718

(CAS, NBG); Farm Wagendrift, 12.1 km S of Lambert’s Bay, ± 5 m,

(- AB), 12 Sept. 1991, Steiner 2386 (NBG); Elandsbaai-Redelinghuys

rd, 8.9 km NW of Redelinghuys turnoff, ± 40 m, (-AD), 14 Sept. 1984.

Steiner 775 (NBG); ibid., 19 Sept. 1988, Steiner 1822 (NBG); Farm

Droogerivier, Road 365, 8.6 km N of turnoff to Alexandershoek, ± 200

m west of road, (-BC), 14 Aug. 1987, Steiner 1570 (NBG); ibid., 19

Aug. 1993, Steiner 2663 (NBG); Langebaanweg, (-CC), 13 Sept. 1991,

Goldblatt & Manning 9207 (NBG); Farm Kersefontein, 5.9 km N of

Berg River, ± 20 m, (-CD), 17 Sept. 1991, Whitehead 1 (NBG); Farm

Doomfontein A, ± 30 m, (-CD), 9 Sept. 1994, Steiner 2886 (NBG);

Farm Suurfontein, 0.7 km W of Sauer Post Office, ± 30 m, (-DC),

10 Sept. 1990, Steiner 2226 (NBG). 3318 (Cape Town): Farm Baar-

huis (NW of Darling), 0.5 km N of entrance to Farm Skilpadfontein,

(S33° 14.769’ E18° 18.334’), ± 65 m, (-AB), 16 Sept. 2005, Steiner

4168 (CAS, NBG); Farm Modderrivier, southwest of Darling along

new National Road, ± 80 m, (-AD), July 1980, Hugo 2427 (PRE).

Diascia appendiculata K.E. Steiner, sp. nov., D.

diffusae (Thunb.) Benth. proxima, sed differt floribus

brevioribus, sacculis corollae destituti, staminibus erec-

tis nec patentibus et filamentis glabratis.

TYPE. — Western Cape, 3218 (Clanwilliam): Grey’s

Pass (Modderfontein), 4.2 km N of turnoff to Paleisheu-

wel, ± 290 m, (-DB), 9 Sept. 1989, Steiner 1978 (NBG,

holo.; CAS, K, MO, PRE, US, iso.).

Annual herb, rosulate, glabrous, simple or branch-

ing from base. Stems decumbent, up to 220 mm

long, angular, up to 6-sided, ribs 2 or more, sides up

to 2 mm wide. Leaves simple, alternate, opposite or

whorled, petiolate, erect or spreading; lamina mostly

oblong, but also ovate to obovate, 9-35 x 3-9 mm,

apex acute to rounded, base attenuate; margins sinu-

ate, pinnatipartite or pinnatisect, lobes up to ± 5 mm

long, ovate, obovate, oblong or deltoid, apices rounded

to acute; petioles up to ± 15 mm long; cauline leaves

progressively smaller upwards. Flowers axillary, 1 or

2 open flowers per stem, nodding in bud, long pedi-

celate; pedicels 17-53 mm long, ascending, dorsiv-

entrally flattened especially where attached to flower,

elongating and spreading at right angles to stem in

fruit, with an abrupt downward curve 3—4 mm from

base of developing capsule. Calyx lobes 5, spreading,

lanceolate, acuminate, margins white-ciliate, upper

3 segments ± equal, 2.4-3. 1 x 1.0-1. 6 mm, lower 2

segments slightly wider. Corolla bilabiate, 5-lobed,

limb 7.4-14.3 x 7.7-14.8 mm; upper lobes ovate to

obovate, 2.0— 4.7 x 2. 5-5. 2 mm, outer sides longer

than inner sides, apices rounded to emarginate, bases

oblique; lateral lobes oblong-ovate, 2. 5^4. 6 x 2. 8^4. 6

mm, sides ± equal in length, apices rounded to emar-

ginate, lower lobe obcordate, 3. 1-5.0 x 3. 6-6. 4 mm,

all lobes reddish lilac (14C4) to greyish magenta

(14D6) on inner surface and violet- white to purplish

white on reverse side, with scattered black or clear

glandular trichomes on both sides; tube shallowly

cupped, dark ruby to violet-brown, very shallowly bi-

saccate, sacs or depressions 0. 3-1.0 x 1.0-1. 6 mm,

0.5-0. 7 mm deep, yellow, oil-secreting trichomes

clustered within; central stamen-bearing boss oblique,

sparsely glandular pubescent, anticous portion 0. 5-1.0

mm high, ruby, posticous portion ± 2.0 mm high, yel-

low. Stamens 4, erect from the boss; filaments ruby,

usually glabrous, occasionally covered with dark pur-

ple, clavate trichomes; anticous filaments (twisted at

the base and appearing posticous) ± straight, 2. 1-2.5

mm long, base strongly curved; posticous filaments

± 2.3 mm long, with sterile appendages, ± 0.8 mm

from base, appendages 0. 1-0.8 mm long, sometimes

reduced in length to a small nub, filament above bend

± 1.5 mm long; anthers 0.3 mm long, strongly coher-

ing, greenish yellow; pollen orange. Ovary ovoid, lat-

erally compressed contrary to septum, 1.2-1. 5 x 0.8-

1.1 mm, falciform; ovules 27—47; style ± 1.2 mm long,

16

Bothalia 39,1 (2009)

FIGURE 4. — Diascia appendiculata , Steiner 1978 (NBG). A, habit. B-D, flower: B, C, front and rear views; D, side view partially cut away. E,

pistil; F, capsule; G, calyx. FI, I, seed: H, ventral view; I, side view. Scale bars: A, 10 mm; B, C, 3 mm; D, E, 1 mm; F, 1 mm; G, 4 mm; H,

I, 1 mm. Artist: Ellaphie Ward-Hilhorst.

deflected upwards, ± straight or curved forward near

tip; stigma capitate, surrounded by anthers or emerging

slightly. Capsule falciform ovoid, (5.0-)6.4— 7.5 x (2.5-)

4.0-5. 0 mm, about twice as long as calyx at maturity, base

oblique, often resting on soil surface during development

with pedicel ascending just before dehiscence. Seeds reni-

form, 1.3-1. 5 mm long, dorsal surface with parallel ridges,

ventral surface with oblong keyhole-like opening formed

Bothalia 39,1 (2009)

17

by extensions of seed coat, long sides of opening bearing

reniform perforation; embryo curved. Chromosome no. : 2n

= 18. Flowering time: August-September.

Diagnostic features : Diascia appendiculata is most

closely allied to D. diffusa. Both species have posticous

filaments with sterile appendages, but the stamens are

erect in D. appendiculata and projecting forward in D.

diffusa. Furthermore, the filaments in D. appendiculata

are usually glabrous, whereas those of D. diffusa have

clavate trichomes. Both species also have two localized

patches of oil-secreting trichomes, but in D. diffusa they

are clustered in two short, but distinct, spurs (at the base

of the upper corolla lip), whereas in D. appendiculata

they are present in two shallow, yellow depressions that

may or may not be visible on the outside of the corolla

as a slight swelling of the tube.

Etymology: the name refers to the filament appendages.

Distribution and habitat: Diascia appendiculata is

known from only six localities in the general vicinity

of Citrusdal (Figure 2). It occurs between elevations of

100 to 300 m in fynbos vegetation on lower mountain

slopes or flats in loose alluvial sands derived from Table

Mountain Sandstone. In five of the localities it occurred

on first year bums, while in the other locality it was col-

lected from a roadside area next to cultivated land at the

northeastern base of the Piketberg. On Grey’s Pass, it

was most abundant in the first season after fire, but was

also observed in the second and third years (1990, 1991)

after fire. It could not be found in the fifth and sixth

years. The stimulation of germination in response to fire

is also found in other Diascia species such as D. elon-

gata Benth. and D. maculata K.E. Steiner.

Pollination and breeding system: Diascia appendicu-

lata is facultatively autogamous. However, because it

secretes floral oil, it is probably visited and cross-pol-

linated, at least occasionally, by small oil-collecting

Rediviva bees (Melittidae) such as R. parva Whitehead

& Steiner, R. intermixta (Cockerell) or R. aurata White-

head & Steiner (Whitehead & Steiner 2001).

Other specimens examined

WESTERN CAPE. — 3218 (Clanwilliam): Farm Swartboskraal,

6.7 km N of turnoff to Citrusdal on Paleisheuwel-Sandberg road, 270

m, (-BC), 22 Aug. 1991, Steiner 2334 (NBG); ibid, 1.5 km S of farm

entrance, 218 m, (-BC), 31 Aug. 2004, Steiner 4082 (NBG); Farm

Krieberg, 7.2 km E of old Clanwilliam-Citrusdal road on road to Alge-

ria, ± 290 m, (-BD), 16 Sept. 1989, Steiner 2009 (NBG); ibid., 24

Sept. 1989, Steiner 2025 (NBG); N7, 1.6 km N of turnoff to Citrusdal,

± 280 m, (-DB), 10 Aug. 1998, Steiner 3287 (NBG); Farm Kanarie-

berg. Road 366, 5.7 km S of junction with Road 365 to Lambert’s Bay,

± 110 m, ( DB), 21 Sept. 1984, Steiner 776 (NBG). 3219 (Wuppertal):

Farm Moddervlei, 13.6 km south of Citrusdal on road to Keerom, ±

210 m, (-CA), 6 Sept. 1991, Steiner 2357 (NBG); Farm Kammelks-

vlei, 19.6 km south of Citrusdal on road to Keerom, ± 250 m, (-CC), 6

Sept. 1991, Steiner 2362 (NBG).

ACKNOWLEDGEMENTS

I thank Western Cape Nature Conservation and the

Western Cape National Park (formerly Postberg Wildflower

Reserve) for permission to collect in areas under their juris-

diction. I thank SANBI for the loan of specimens from

NBG and PRE and for permission to use the illustrations

by E. Ward-Hilhorst. I also thank the Compton Herbarium

for the use of their facilities and S. Smithies and T. Arnold

for helpful comments on the manuscript.

REFERENCES

HIERN, W.P. 1904. Scrophulariaceae. In W.T. Thiselton-Dyer, Flora

capensis 4,2: 121-420. Reeve, London.

HILLIARD, O.M. & BURTT, B.L. 1984. A revision of Diascia section

Racemosae. Journal of South African Botany 50: 269-340.

KORNERUP, A. & WANSCHER, J.H. 1 984. Methuen handbook of col-

our. Fletcher, Norwich.

MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of

South Africa, Lesotho and Swaziland. Strelitzia 19. South Afri-

can National Biodiversity Institute, Pretoria.

STEINER, K.E. 1989. A new species of Diascia (Scrophulariaceae)

from the southern Drakensberg. South African Journal of Botany

55: 250-253.

STEINER, K.E. 1992a. A new Diascia species (Scrophulariaceae) from

the Richtersveld, South Africa. South African Journal Botany

58: 36-38.

STEINER, K.E. 1992b. Two new Diascia species (Scrophulariaceae)

from the Little Karoo. South African Journal of Botany 58:

39-47.

STEINER, K.E. 1992c. Two new Diascia (Scrophulariaceae) species

from the Nieuwoudtville area, western Cape. South African

Journal of Botany 58: 202—206.

STEINER, K.E. 1992d. Three new species of Diascia (Scrophulariace-

ae) from the western Cape. Bothalia 22: 13-18.

STEINER, K.E. 1995. Three new Diascia species from arid areas of the

Western Cape, South Africa. South African Journal of Botany

61:72-79.

STEINER, K.E. 1996. Chromosome numbers and relationships in

tribe Hemimerideae (Scrophulariaceae). Systematic Botany 21:

63-76.

STEINER, K.E. 1999. A new species of Diascia (Scrophulariaceae)

from the Eastern Cape (South Africa), with notes on other mem-

bers of the genus in that region. South African Journal of Botany

65:223-231.

WHITEHEAD, V.B. & STEINER, K.E. 2001. Oil-collecting bees of

the winter rainfall area of South Africa (Melittidae, Rediviva).

Annals of the South African Museum 108: 143-277.

■

■

Bothalia 39,1: 19-35 (2009)

Aloe in Angola (Asphodelaceae: Alooideae)

R.R. KLOPPER*, S. MATOS**, E. FIGUEIREDO*** and G.F. SMITH*+

Keywords: Aloe L., Angola, Asphodelaceae, flora

ABSTRACT

Botanical exploration of Angola was virtually impossible during the almost three-decade-long civil war. With more areas

becoming accessible, there is, however, a revived interest in the flora of this country. A total of 27 members of the genus Aloe

L. have been recorded for Angola. It is not unlikely that new taxa will be discovered, and that the distribution ranges of oth-

ers will be expanded now that botanical exploration in Angola has resumed. This manuscript provides a complete taxonomic

treatment of the known Aloe taxa in Angola. It includes, amongst other information, identification keys, descriptions and

distribution maps.

INTRODUCTION

The Republic of Angola covers an area of ±

1 246 700 km2 in southwest-central Africa. Its west-

ern boundary is 1 650 km along the Atlantic Ocean and

it is bordered by Namibia in the south, the Democratic

Republic of Congo in the north and northeast, and Zam-

bia in the east. The detached province of Cabinda has

a border with the Republic of Congo in the north and

the Democratic Republic of the Congo in the southeast

(http://www.angola.org) (Figure 1).

The geography of Angola is extremely varied. The flat

coastal part has a few shallow bays and is bordered by a

sparsely vegetated coastal plain that extends inland for

48-165 km. This coastal belt is separated from the central

plateau by an intermediate mountain belt of irregular ter-

races, running mostly parallel to the coast. Water is more

abundant in these mountain chains and the vegetation is

therefore lush. The central plateau has an altitude of 1 200-

1 800 m and consists of rolling plains and low hills with

scanty vegetation. The plateau falls away in the east to the

basins of the Congo and Zambezi Rivers and merges with

the barren, sandy Namib Desert in the south. Several small

rivers arise in the mountain belt and drain westward to the

sea. The largest of these are the Cuanza and Cunene Riv-

ers. From the plateau, the Cuango and other rivers flow

northwards to join the Casai River, one of the largest tribu-

taries of the Congo River. Rivers in the south of the coun-

try either belong to the Zambezi River system or, like the

Okavango, drain to Lake Ngami in Botswana (http://www.

biocrawler.com/encyclopedia/Geography_of_Angola).

Angola is situated in a subtropical zone, but owing to

several factors, the climate of the country is not typical of

such areas. Angola’s climate is influenced by the cold Ben-

guela Current along the southern part of the coast, the high-

lands in the interior and the Namib Desert in the southwest.

* Biosystematics Research and Biodiversity Collections Division,

South African National Biodiversity Institute, Private Bag X101, 0001

Pretoria. E-mail: Klopper@sanbi.org.

** Institute de Investigate) Cientifica Tropical, Trav. Conde da Ribeira

9, 1300-142 Lisboa, Portugal. E-mail: SusanaMatos78@gmail.com.

*** H.G.W.J. Schweickerdt Herbarium, Department of Botany, Uni-

versity of Pretoria, 0002 Pretoria. E-mail: EstrelaFigueiredo@hotmail.

com.

“Acocks Chair, H.G.W.J. Schweickerdt Herbarium, Department of Bot-

any, University of Pretoria, 0002 Pretoria. E-mail: SmithG@sanbi.org.

MS. received: 2008-09-04.

The country has two distinct seasons: the rainy season

from October to May, with average coastal temperatures

of around 21°C and the dryer season with lower average

coastal temperatures of around 16°C and mist or Cacimbo

from June to September. The heaviest rains occur in April

and are accompanied by violent storms. Rainfall along the

coast is high and gradually decreases from 800 mm in the

north to 50 mm in the south. The interior can be divided

into three zones: the North, with very heavy rains and high

temperatures; the Central Plateau, a dry area with average

temperatures; and the South with temperature fluctuations

due to the proximity of the Kalahari Desert and the influ-

ence of tropical air currents (http://www.biocrawler.com/

encyclopedia/GeographyofAngola).

The vegetation of Angola is predominantly Zambezian

and falls mainly within the Tropical Grassland (Savanna)

zone. Six of White’s phytochoria are represented in the

country (White 1983). Humid tropical rainforest occurs

in the north of the country and the arid Namib Desert

and Karoo-Namib shrubland occur in a narrow coastal

strip in the southwest. Elsewhere the vegetation consists

mostly of miombo woodland, dry evergreen forest, dry

deciduous forest, grassland and savanna with Afromon-

tane formations in the highlands (Airy Shaw 1947; Costa

et al. 2004). In a recent classification of terrestrial ecore-

gions based on ecological features, climate, and plant

and animal communities, 15 regions are represented in

Angola (World Wildlife Fund 2001).

The geography and unique climatic characteristics of

the area provide Angola with a rich biodiversity. How-

ever, the flora of the country remains poorly known, a

situation recently addressed by Figueiredo & Smith

(2008). Although the first botanical specimens from

Angola were collected towards the end of the 18th cen-

tury and various explorations were carried out during

the 18th, 19th and 20th centuries, the Angolan Civil War

(1975-2002) prevented the country from being properly

surveyed for almost three decades. After the end of the

civil war, some areas are still inaccessible owing to the

threat of landmines (Costa et al. 2004). Even before the

war, the poor condition of some roads prevented explor-

ers from reaching many isolated habitats in this country.

This situation still prevails to some extent today. With

certain areas of the country becoming increasingly acces-

sible, a new interest in the botanical wealth of Angola is

surfacing.

20

Bothalia 39,1 (2009)

FIGURE 1. — Map of Angola showing the provinces.

In 2006, a project on the flora of Angola, Flora of

Angola Online (FLAN) was initiated in the South Afri-

can National Biodiversity Institute (SANBI), with the

collaboration of the Institute de Investigaipao Cientlfica

Tropical in Lisbon, Portugal, and the Institute de Investi-

gagao Agronomica in Angola. The project had the objec-

tive of producing a compilation of plant names with

associated specimen data, with the collaboration of 32

botanists. As a result of this joint effort, a comprehen-

sive list of the vascular plants of Angola was compiled

(Figueiredo & Smith 2008).

The results of that project (Figueiredo & Smith 2008;

Figueiredo et al. in press) provide the following data for

the plant diversity of the country: a total vascular flora of

7 296 taxa, consisting of 6 961 species (of which 6 735

are indigenous) and 335 infraspecific taxa (of which 331

are indigenous), belonging to 250 families; 997 species

and 72 infraspecific taxa are endemic.

This manuscript provides a complete treatment of

the known Aloe L. taxa in Angola. It is the culmination

of research done for the Angolan Flora Project and also

forms part of the Aloes of the World Project, which is

funded by the Andrew W. Mellon Foundation. Its objec-

tives are to compile as much information as possible on

representatives of the genus Aloe and to make this data

available to a broad range of stakeholders through the

internet as part of the African Plants Initiative of Aluka

(Smith et al. 2008a, b).

MATERIALS AND METHODS

Geo-referencing of specimens was undertaken for

all the specimens examined. The co-ordinates of the

collecting locality of each specimen were determined

using the Angolan map collection kept at the Institute

de Investiga9ao Cientifica Tropical. Distribution maps

are based on the type specimens and those cited under

Additional specimens examined. These specimens are

mainly housed in LISC and PRE, but also in BM, BR,

E, G, K, LISU, M and MO (acronyms as in Holmgren et

al. 1990). Specimens not housed at LISC, LISU or PRE

were viewed on the Aluka website (http://www.aluka.

org) of the African Plants Initiative. Specimens housed

at B were viewed on the virtual herbarium website of the

Herbarium Barolinense (Ropert 2000). Further distribu-

tion records for taxa with a distribution range extend-

ing outside Angola, were obtained from specimens cited

in treatments of the genus in Flora zambesiaca (Carter

2001) and Flora of tropical East Africa (Carter 1994), as

well as specimens housed at PRE.

This treatment includes a summary of the discovery

of aloes in Angola, as well as identification keys to the

taxa using either field or herbarium characters. For each

taxon a description, based on the abbreviated description

template of the Aloes of the World Project (Smith et al.

2008a, b), is provided. Other information that is supplied

for each taxon includes the protologue citation, type

specimen information, diagnostic characters and speci-

mens investigated, as well as notes on habitat, flowering

time, distribution and endemism (indicated by E in front

of the taxon name). Each taxon is further accompanied

by a distribution map.

HISTORY OF ALOE DISCOVERY IN ANGOLA

The known Aloe taxa in Angola currently number 27,

of which at least 16 (59 %) are endemic to the country.

Only five of the 27 Angolan aloes (A. bulbicaulis Chris-

tian, A. christianii Reynolds, A. dinteri A.Berger, A.

hereroensis Engl, and A. nuttii Baker) do not have their

type localities within Angola. The other 22 Aloe taxa

occurring in the country were described from material

collected during plant collecting surveys or expeditions

to Angola from the mid- 1800s up to as recently as 1973.

Important collectors of Aloe in Angola were Dr Friedrich

Welwitsch (1853-1861), Lieut. Wissmann & P. Pogge

during their expedition through Angola and the Demo-

cratic Republic of Congo (1881-1882), Hugo Baum dur-

ing the Kunene-Zambezi Expedition (1899-1900), John

Gossweiler (1903-1944), Edgar Milne-Redhead (1938),

Eduardo J. Mendes (1955-1956 and 1959-1960), Gil-

bert W. Reynolds (1959), Larry C. Leach & I. C. Cannell

during the early 1970s and Baptista de Sousa (1973).

Botanical exploration and the possible discovery of new

Aloe taxa ceased with the advent of the 27-year-long

Angolan Civil War. As a result of vast areas of the coun-

try remaining unexplored, it is likely that more taxa will

be found once detailed botanical surveys of these areas

are resumed.

The first Aloe specimens from Angola were collected

by Welwitsch during 1853-1861. From these collections

Baker (1878) described six new species, of which only

A. platyphylla Baker was later reduced to synonymy

under A. zebrina Baker, and A. angolensis Baker is

sometimes considered to be either a hybrid between A.

zebrina and A. littoralis Baker or to be a synonym of

the latter species (Glen & Hardy 2000; Carter 2001).

A further species, A. venenosa Engl., was described by

Engler (1893) from material collected by Wissmann &

Pogge during their expedition through Angola and the

Democratic Republic of Congo in 1881-1882. This spe-

cies has not been collected since and remains insuffi-

ciently known. During the Kunene-Zambezi Expedition

Bothalia 39,1 (2009)

21

of 1899 to 1900, Baum collected Aloe specimens from

which three new species were described by Engler &

Gilg (1903). However, only A. metallica Engl. & Gilg

is still regarded as a valid species, whereas A. brunneo-

punctata Engl. & Gilg and A. baumii Engl. & Gilg have

been reduced to synonymy under A. nuttii Baker and A.

zebrina, respectively.

In 1903, Gossweiler collected material that was later

described by Berger (1906) as Aloe paedogona A. Berger.

Christian (1936) described A. bulbicaulis Christian from

material collected by Porter in Zambia, but Reynolds

(1966) later established that it is a later synonym of A.

paedogona. The latter species is, furthermore, consid-

ered by some (Reynolds 1966; Keay 1968) as a syno-

nym of A. buettneri A. Berger, although more recent

views (Carter 1994) consider the three as separate taxa.

Milne-Redhead collected a further Aloe in 1938 that was

subsequently described by Christian (1940a) as A. milne-

redheadii Christian.

Reynolds visited Angola during June and July of

1959, but could not reach certain areas in the northeast

and southeast of the country due to the inaccessibility

of these areas. During his journey he established that

A. hereroensis Engl., from Namibia and the Northern

Cape Province of South Africa, also occurs in Angola.

An additional six species were described by Reynolds

(1960, 1961, 1962, 1964) after his Angolan travels, all of

which are still upheld.

During the early 1970s, Leach & Cannell collected

Aloe material in Angola from which Leach (1971,

1974) described five new species and a new variety of

A. andongensis Baker. At the same time Leach (1974)

also described a further species of which the material

was collected and given to him by Baptista de Sousa in

1973. All seven of these taxa are still considered cur-

rent, although A. esculenta L.C. Leach is sometimes con-

sidered to be synonymous with A. angolensis (Glen &

Hardy 2000).

IDENTIFICATION KEYS

Two identification keys are presented below. The first

uses field characters for the identification of living plants,

whereas the second uses mostly leaf and inflorescence

characters (flowers, pedicels and floral bracts) and is

aimed at identifying herbarium specimens. Aloe venenosa

is not included in the identification keys owing to lack of

information for this insufficiently known species.

Key using field characters

1 a Acaulescent or with very short stem:

2a Leaf bases markedly enlarged below ground to form bulb-like swelling:

3a Leaves with marginal teeth ± 1 mm long, densely crowded; floral bracts ovate-acuminate A. bulbicaulis

3b Leaves with marginal teeth ± 3—4 mm long, 5—40 mm apart; floral bracts linear-lanceolate A. paedogona

2b Leaf bases not markedly enlarged to form bulb-like swelling:

4a Flowers vertically disposed (secund) when open:

5a Inflorescence up to 1 m high; racemes subdense; flowers scarlet with a bloom, ± 40 mm long A. guerrae

5b Inflorescence 2.20-2.75 m high; racemes lax; flowers dull reddish purple, 28-33 mm long A. procera

4b Flowers horizontally or pendulously disposed when open:

6a Racemes capitate:

7a Leaves with copious small, white, circular spots near base on lower surface; leaf margin with small crowded teeth that are some-

times almost serrate A. grata

7b Leaves obscurely lineate, with few to many whitish spots scattered or arranged in transverse bands on lower surface; leaf margin

with pungent, red-brown teeth A. hereroensis var. hereroensis

6b Racemes acuminate:

8a Leaves not marked with whitish spots on either surface:

9a Inflorescence ± 0.9 m long; flowers yellow, 20-25 mm long A. angolensis

9b Inflorescence longer than 1 m; flowers reddish pink, longer than 30 mm:

1 0a Flower buds at first covered by densely imbricate, long, white, prominently nerved bracts A. metallica

10b Flower buds not covered by dense floral bracts A. christianii

8b Leaves marked with whitish spots on one or both surfaces:

11a Leaves trifarious A. dinteri

lib Leaves not trifarious:

12a Leaves with prickles along median line of lower leaf surface A. esculenta

12b Leaves without prickles along median line of lower leaf surface:

13a Leaves copiously white-spotted on upper surface, usually obscurely spotted on lower surface; marginal teeth 4-7 mm

long A. zebrina

13b Leaves densely white-spotted on both surfaces, especially on lower surface; marginal teeth 2-3 mm long ... A. milne-redheadii

lb Stems erect, procumbent, ascending or pendent:

14a Stems erect:

15a Leaves grass-like A. nuttii

1 5b Leaves not grass-like:

16a Stem usually shorter than 1 m:

17a Stem 0.5-1 m long; leaves obscurely lineate, rarely with few spots near base A. scorpioides

17b Stem up to 0.3 m long; leaves with spots arranged in wavy transverse bands A. lepida

16b Stem usually longer than 1 .5 m:

1 8a Stems without persistent dried leaves A. rupicola

1 8b Stems with persistent dried leaves:

1 9a Stem unbranched A. littoralis

1 9b Stem branched A. palmiformis

14b Stems procumbent, ascending or pendent:

20a Stems pendent:

21a Stems without persistent dried leaves; leaves obscurely lineate, without spots A. mendesii

21b Stems with persistent dried leaves; leaves obscurely lineate with few small, whitish, H-shaped spots; spots more numerous and in

transverse bands on lower surface z A. inamara

22

Bothalia 39,1 (2009)

20b Stems procumbent, ascending:

22a Stems slender, branched, forming thickets:

23a Stems without persistent dried leaves A. catengiana

23b Stems with persistent dried leaves A. gossweileri

22b Stems branched, but not slender, not forming thickets:

24a Inflorescence 0.5-0. 9 m high, unbranched or branched once, with lax racemes A. vallaris

24b Inflorescence 0. 3-0.4 m high, branched, with dense racemes:

25a Leaves sometimes sparsely spotted on lower surface, with many crowded spots near base, 200-250 x 60-70 mm, with mar-

ginal teeth 2-3 mm long, 5-7 mm apart; ovary pale green A. andongensis var. andongensis

25b Leaves more copiously white-spotted with spots tending to be arranged in wavy transverse bands, smaller and narrower than

above, with marginal teeth smaller and more crowded; ovary brownish orange A. andongensis var. repens

Key using herbarium characters

la Floral bracts longer than pedicels at anthesis:

2a Flowers 20-24 mm long; outer perianth segments free for < 10 mm; leaves 40-50 mm wide A. angolensis

2b Flowers longer than 25 mm; outer perianth segments free for > 10 mm; leaves wider than 60 mm:

3a Floral buds and open flowers secund:

4a Racemes ± 200 mm long; flowers ± 40 mm long, ± 8 mm across ovary; floral bracts 6-8 mm long; leaves 60-70 mm wide, with

marginal teeth 4—5 mm long A. gnerrae

4b Racemes 250-400 mm long; flowers 28-33 mm long, 5-6 mm across ovary; floral bracts 5-6 mm long; leaves 80-95 mm wide,

with marginal teeth 1 .5-3.5 mm long A. procera

3b Floral buds and open flowers not secund:

5a Leaves with blackish brown marginal teeth and similar spines along median line of lower leaf surface; pedicels 5-6 mm long;

floral bracts 20-27 mm long A. esculenta

5b Leaves with reddish brown marginal teeth and no spines along median line of lower leaf surface; pedicels longer than 6 mm; floral

bracts shorter than 20 mm:

6a Leaves up to 600 x 100-130 mm. with marginal teeth 3-4 mm long; inflorescence much-branched and rebranched; floral bracts

12-18 mm long A. littoralis

6b Leaves 250-400 x 70-90 mm, with marginal teeth 2-3 mm long; inflorescence sparsely branched; floral bracts 1 8-20 mm

long A. metallica

lb Floral bracts shorter than or equal to pedicels at anthesis:

7a Floral bracts about as long as pedicels at anthesis:

8a Floral bracts almost equal to or slightly shorter than pedicels at anthesis; leaves < 50 mm wide:

9a Inflorescence up to 0.15 m long, simple or 1- or 2-branched, descending at base and then curving upwards; racemes rather dense;

flowers 21-28 mm long, well constricted above ovary; outer perianth segments free for 8.5-10.0 mm A. scorpioides

9b Inflorescence 0.5-0. 6 m long, simple or 1-branched, oblique or suberect; flowers 20-25 mm long, only slightly constricted above

ovary; outer perianth segments free for 4. 5-6.0 mm A. vallaris

8b Floral bracts almost equal to pedicels at anthesis; leaves > 50 mm wide, usually > 60 mm wide:

10a Leaves 100-120 mm wide, unspotted; flowers 35^15 mm long, not constricted above ovary A. christianii

10b Leaves < 80 mm wide, spotted on both surfaces; flowers up to 35 mm long, abruptly constricted above ovary:

11a Leaf margin narrow, white, with minute white teeth, ± 0.5 mm long, 1-2 mm apart, edge of keel white cartilaginous with similar

teeth A. dinteri

lib Leaf margin cartilaginous, with stout, pungent, red-brown teeth, 4-7 mm long, 10-15 mm apart, no spines on median line ... A. zebrina

7b Floral bracts markedly shorter than pedicels at anthesis:

12a Outer perianth segments free for ± !4 or > 'A its length:

13a Pedicel > 25 mm long at anthesis:

14a Flowers 25-33 mm long; outer perianth segments free for 14—16 mm A. hereroensis var. hereroensis

14b Flowers 35-42 mm long; outer perianth segments free almost to base, or for % of its length A. nuttii

13b Pedicel < 20 mm long at anthesis:

15a Flowers ± 42 mm long A. nipicola

15b Flowers ± 25 mm long:

16a Pedicels 18-20 mm long; floral bracts ± 12 mm long A. mendesii

16b Pedicels 14-18 mm long; floral bracts 5-8 mm long:

17a Leaves 200-250 x 60-70 mm, with marginal teeth 2-3 mm long, 5-7 mm apart A. andongensis var. andongensis

1 7b Leaves smaller and narrower than above, with marginal teeth smaller and more crowded A. andongensis var. repens

12b Outer perianth segments free for < 'A its length:

1 8a Leaves with marginal teeth up to 1 mm long:

19a Flowers 35^10 mm long; pedicels ± 20 mm long; leaves ± 150 mm wide; acaulescent plant with leaf bases enlarging below

ground to form bulb-like swelling A. bulbicaulis

19b Flowers 26-29 mm long; pedicels 22-27 mm long; leaves 40-50 mm wide; plants with branched stem, growing pendent on

cliff faces A. inamara

18b Leaves with marginal teeth > 2 mm long:

20a Floral bracts 15-25 mm long; pedicels 25-30 mm long; acaulescent plants with leaf bases enlarging below ground to form bulb-

like swelling A. paedogona

20b Floral bracts < 7 mm; pedicels usually < 20 mm; shrubs or if acaulescent, then leaf bases not enlarging below ground to form

bulb-like swelling:

21a Floral bracts up to 3 mm long:

22a Leaves 70-80 mm wide; flowers 25-28 mm long A. grata

22b Leaves ± 50 mm wide; flowers ± 30 mm long:

23a Flowers subsecund when open; pedicels ± 10 mm long; leaves with marginal teeth spaced ± 15 mm apart A. gossweileri

23b Flowers nodding to pendulous, not subsecund when open; pedicels 13-15 mm long; leaves with marginal teeth spaced ±

10 mm apart A. palmiformis

2 1 b Floral bracts 5 mm or longer:

24a Leaves ± 35 mm wide; racemes up to 160 mm long; pedicels ± 10 mm long A. catengiana

24b Leaves > 60 mm wide; racemes 200 mm or longer; pedicels longer than 13 mm:

25a Leaves with marginal teeth 3-7 mm long; flowers 25-29 mm long, ± 5.5 mm across ovary; outer perianth segments free

for 5-6 mm A. lepida

25b Leaves with marginal teeth 2-3 mm long; flowers 28-35 mm long, ± 8 mm across ovary; outer perianth segments free for

±10mm A. milne-redheadii

Bothalia 39,1 (2009)

23

TAXONOMY

EA. andongensis Baker var. andongensis in Trans-

actions of the Linnean Society of London 1: 263 (1878).

Type: Angola, Pungo Andongo, Welwitsch 3729 (BM,

holo.!; K!, LISC!, LISU!, iso.).

Branched shrub. Stem short or 0.3-0. 6 m high, branched,

ascending, sometimes becoming decumbent, with persistent

dried leaves. Leaves rosulate at branch apices, varying from

spreading and slightly recurved to suberectly spreading and

compact, dull grey-green, upper surface mostly without

spots, sometimes sparsely spotted, lower surface usually

with many crowded spots near base, lanceolate-attenuate,

200-250 x 60-70 mm; margin slightly cartilaginous, with

brownish teeth 2-3 mm long, 5-7 mm apart; leaf exudate

crusty when dry. Inflorescence 0. 3-0.4 m high, erect, 2-

or 3-branched. Raceme subcapitate to cylindrical-acumi-

nate, 60-120 mm long, dense. Floral bracts 5-8 x 3 mm.

Pedicels 14-18 mm long. Flowers : perianth pale orange-

scarlet, paler at tips, 25 mm long, 5-6 mm across ovary,

narrowed above ovary, widening towards mouth giving a

clavate appearance, cylindric and very slightly decurved;

outer segments free for 17 mm. Stamens exserted up to 1

mm. Style exserted up to 1-2 mm. Flowering time : January

to April.

Diagnostic characters : upper leaf surface mostly

without spots, sometimes sparsely spotted, lower surface

usually with many crowded spots near the base. Buds all

spread somewhat horizontally or slightly deflexed. Inflor-

escence subdense, subcapitate, with flowers lacking a

basal swelling.

Relationships with other species'. Reynolds (1966) placed

Aloe andongensis in his Group 19: Plants of shrubby

growth. It seems to be closely related to A. lepida

L.C.Leach and also shows a strong link in vegetative char-

acters with A. squarrosa Baker from Socotra (Leach 1974).

Habitat, exposed rocky places.

Distribution : endemic to Angola (Cuanza Sul, Huambo,

Malange) (Figure 2).

Illustrations'. Reynolds: 347 (1966).

Additional specimens examined

ANGOLA. — 0915: Pungo Andongo, (-DA), 22 March 1973,

Bamps, Martins & Silva 4246 (LISC, PRE), 31 March 1967, Barbosa

11332 (LISC), 29 December 1911, Gossweiler 5445 (LISC), 14 May

1960, Reynolds 9385 (BM, PRE). 1014: Cuanza Sul, Gabela, rocha

Chitandero, na Ro?a Afficana de CAD A, (-CD), 16 March 1967, Teix-

eira 11203 (LISC). 1015: Benguela Province, 14 miles [22.5 km] S of

Quibala, (-CC), 18 June 1960, Reynolds 9366 (BM). 1115: Cuanza Sul,

4 miles [6.5 km] NW of Cassongue, (-CC), 26 May 1960, Reynolds

9335 (BM, PRE). 1215: Huambo, Alto Hama, Iumbo, Rio Queve, (-

BA), 6 April 1966, Correia 3638 (LISC).

EA. andongensis Baker var. repens L.C.Leach in

Journal of South African Botany 40: 115 (1974). Type:

Angola, Cuanza Sul, 11 km E of Gabela, 8 June 1973,

Leach & Canned 13950 (LISC, holo.!; BM!, BR!, K!,

PRE!, SRGH, iso.).

Branched shrub, forming large spreading clumps.

Stem up to 0.6 m high, branched freely from base and

above, prostrate, with persistent dried leaves. Leaves

rosulate at branch apices, widely spreading, dull grey-

green, upper surface mostly without spots, sometimes

sparsely spotted, lower surface usually with copious

crowded spots near base, spots tend to be arranged in

wavy transverse bands, lanceolate-attenuate, smaller

and narrower than typical variety; margin slightly carti-

laginous, with brownish teeth smaller and more crowded

than typical variety; exudate crusty when dry. Inflores-

cence 0.3-0. 4 m high, erect, 2- or 3-branched. Raceme

subcapitate to cylindrical-acuminate, 60-120 mm long,

dense. Floral bracts 5-8 x 3 mm. Pedicels 14-18 mm

long. Flowers', perianth pale orange-scarlet, paler at tips,

25 mm long, 5-6 mm across ovary, narrowed above

ovary, widening towards mouth giving a clavate appear-

ance, cylindric and very slightly decurved; outer seg-

ments free for 17 mm. Stamens exserted up to 1 mm.

Style exserted up to 1-2 mm. Flowering time : February

to April.

Diagnostic characters', prostrate habit. Stems up to 0.6 m

long, branching freely from the base and above, thus form-

ing large spreading clumps. Stems foliate for greatest part

of their length. Leaves much smaller, narrower, and more

widely spreading than typical variety of species, also more

copiously white-spotted with spots tending to be arranged

in wavy transverse bands, with smaller, more crowded

marginal teeth. Buds all spread somewhat horizontally or

slightly deflexed. Inflorescence subdense, subcapitate, with

flowers lacking a basal swelling. Ovary brownish orange.

Relationships with other species : see comments under

Aloe andongensis subsp. andongensis.

Habitat : slopes of rounded granite hills.

Distribution : endemic to Angola (Cuanza Sul) (Figure 3).

Illustrations'. Leach: 116 (1974).

Additional specimens examined

ANGOLA. — 1114: Cuanza Sul, Santa Comba-Amboiva, bac du

Queve, (-DB), 28 March 1973, Bamps & Martins 4301 (LISC). 1115:

Cuanza Sul, Santa Comba, proximo de Santa Comba, (-AC), 10 March

1965, Santos 1412 (LISC).

EA. angolensis Baker in Transactions of the Lin-

nean Society of London 1: 263 (1878). Type: Angola,

Barra do Bengo, between Quisoma and Cacuaco at

Mutolo in District Quicuxe, 1858, Welwitsch 3728 (BM,

holo.!; G!, K!, LISC!, LISU!, iso.).

FIGURE 2. — Distribution of Aloe andongensis var. andongensis, ■;

A. angolensis, ▲; and A. bulbicaulis, •.

24

Stem very short or up to 0.7 m high. Leaves densely

rosulate, suberect, glaucous, not spotted below, very

fleshy, lanceolate-ensiform, 600 x 40-50 mm; margin

with teeth, 2 mm long, 15-20 mm apart. Inflorescence

0.9 m high, erect, simple or up to 3-branched. Raceme

cylindrical, slightly acuminate, 100 mm long, dense.

Floral bracts 1 0 mm long. Pedicels 3-6 mm long. Flow-

ers'. perianth sulphur-yellow, 20-24 mm long; outer

segments fused to beyond middle (free for less than 10

mm). Stamens and style scarcely exserted. Flowering

time : unknown.

Diagnostic characters', thick leaves and branched

inflorescence with dense racemes of sulphur-yellow

flowers.

Relationships with other species: Reynolds (1966) stated

that the affinities of this species are uncertain, but noted

that the thick leaves and type of inflorescence suggest a

link with the Aloe littoralis- complex. Reynolds (unpub-

lished notes at PRE) further speculated that A. angolen-

sis might be a natural hybrid between A. littoralis and

A. zebrina Baker. This view is upheld by Glen & Hardy

(2000), although they regarded it as conspecific with A.

esculenta L.C. Leach. Carter (2001) considered it as a

synonym of A. littoralis.

Habitat : low hills facing the Bengo River valley, not far

from the sea.

Distribution : endemic to Angola (Bengo) (Figure 2).

Notes : this species has not been found again since Wel-

witsch collected the type specimen. A specimen col-

lected on steep limestone slopes facing the sea between

the Dande River Mouth and the road to Caxito, north of

Luanda [ Barbosa & Santos 10833 (LISC)] is considered

to possibly be Aloe angolensis. However, this specimen

was not in flower and had dry, dehisced capsules. Until

such time as flowering plants are found at this locality,

it is not possible to say if this specimen belongs to A.

angolensis or not (Reynolds 1966; Reynolds, unpub-

lished notes at LISC).

A. bulbicaulis Christian in The Flowering Plants

of South Africa 16: t. 630 (1936). Type: Zambia, West-

FIGURE 3. — Distribution of Aloe andongensis var. repens , ▲ ; A. ca-

tengiana. B; and A. chrislianii , •.

Bothalia 39,1 (2009)

em Province, Misundu Siding, Porter cult. Christian

PRE20587 (PRE, holo.!).

A. buettneri auct., sensu Reynolds (1966), p.p. ref. loc. in DRC,

Malawi & Zambia.

Acaulescent, up to 0.5 m high, with leaf bases

enlarged below ground to form bulb-like swelling;

rosettes solitary. Leaves rosulate, deciduous, spreading,

bright green, longitudinally striate, slightly fleshy, sur-

face smooth, ovate-lanceolate, up to 500 mm long, 150

mm wide at middle; margin white, cartilaginous, with

densely crowded, fairly evenly-spaced, whitish teeth, 1

mm long, 1-5 mm apart. Inflorescence up to 0.6 m high,

erect, 3- or 4-branched. Raceme cylindrical, 100-200

mm long, lax below, more dense above. Floral bracts 8-

15 x 5-8 mm. Pedicels ± 20 mm long. Flowers : perianth

pale yellow to pinkish or brownish yellow with darker

nerves, 35^-0 mm long, 8-10 mm across ovary, slightly

constricted above ovary, widening towards wide-open

mouth, cylindrical; outer segments free for one-third

(12-13 mm). Stamens scarcely exserted. Style slightly

exserted. Flowering time: February.

Diagnostic characters: underground bulb. Leaves up

to 500 mm long, 150 mm wide in middle, with densely

crowded teeth of 1 mm long. Floral bracts ovate-acumi-

nate, 8-15 mm long.

Relationships with other species: Reynolds (1966) consid-

ered Aloe bulbicaulis to be conspecific with A. buettneri

together with A. paedogona. Carter (1994) stated that the

three taxa are morphologically clearly distinct and also

separated geographically and therefore does not agree with

Reynold’s opinion that the three taxa are conspecific.

Habitat: seasonally wet grassland in open woodland.

Distribution: Eastern Angola (Moxico), southeastern

Democratic Republic of the Congo (Katanga), northern

Malawi, northwestern Mozambique, southwestern Tan-

zania, northern Zambia (Figure 2).

Illustration: Lane: 17 (2004).

Notes: cited in Carter (1994) to occur in western Angola.

This is clearly a mistake and should be eastern Angola. It

has been collected in western Zambia, at Matonchi Farm

[Milne-Redhead 2903A (K-PRE, photo.); Christian 868

(PRE)] very close to the Angolan border.

A. catengiana Reynolds in Kirkia 1: 160 (1961).

Type: Angola, Benguela District, at Catengue, 60 miles

[96 km] SE of Benguela, 2 August 1960, Reynolds 9307

(PRE, holo.!; K, iso.!).

Shrub, forming dense, tangled thickets of 1-2 m

across. Stem 1. 5-2.0 m long, simple or branching low

down, slender, ascending, divergent or sprawling, dried

leaves not persistent. Leaves rather laxly rosulate on api-

cal 0.3 m of stem, spreading to deflexed near base, pale

yellowish grey-green, with copious, very pale green len-

ticular spots on both surfaces, more numerous towards

base, narrowly lanceolate-attenuate, ± 300 x ± 35 mm;

sheath lineate, 15-20 mm long; margin with firm, pale,

reddish brown-tipped teeth, 3 mm long, 8-10 mm apart.

Inflorescence 0.4 m high, erect or suberect, slender,

divaricately about 6-branched. Raceme cylindrical-acu-

minate, terminal raceme longest, 160 mm long, erect,

lateral racemes shorter, oblique, rather lax, flowers sub-

Bothalia 39,1 (2009)

25

secund on lateral racemes. Floral bracts 5x3 mm. Pedi-

cels 10 mm long. Flowers : perianth dull scarlet, 28 mm

long, 7 mm across ovary, slightly narrowed above ovary,

slightly enlarging towards mouth, cylindric, slightly

decurved; outer segments free for 10 mm. Stamens

exserted 1-2 mm. Style exserted up to 2 mm. Flowering

time : March to July.

Diagnostic characters', forms dense, tangled thick-

ets. Stems slender. Leaves peculiar pale yellowish grey-

green, usually copiously spotted on both surfaces. Inflor-

escence slender and divaricately branched. Racemes

vary from terminal erect with flowers evenly distributed

around axis, to oblique with flowers somewhat secund.

Relationships with other species', its closest ally is Aloe

palmiformis (Reynolds 1961).

Habitat', hot, arid bush country and quartzitic sandstone

cliffs.

Distribution : western Angola (Benguela, Namibe),

northern Namibia (Figure 3).

Illustration'. Reynolds: 373 (1966).

Notes: this species was previously only known from the

type locality and considered to be endemic to Angola.

However, it has recently been discovered in the Kaokoveld

in Northern Namibia [ E . van Jaarsveld 18805 (WIND)].

A. christianii Reynolds in Journal of South Afri-

can Botany 2: 171 (1936a). Type: Zimbabwe, Ewanrigg,

E of Harare, Acturus, May-June 1936, Reynolds 1885

(PRE, holo.!; K!, SAM!, SRGH, iso.).

Acaulescent; rosettes usually solitary, up to 1 m

across, or in small groups, rarely suckering. Stem devel-

ops in older plants, up to 1 m long, erect or decumbent,

with persistent dried leaf bases. Leaves densely rosulate,

erectly spreading, dull green above, dull bluish green

below, tinged pinkish in dry conditions, unspotted but

often obscurely lineate, lanceolate-attenuate, 300-600

mm long, 100-120 mm wide at base; margin cartilagi-

nous, with pungent, pinkish to pale brown, brown-tipped

teeth, 2-5 mm long, 10-20 mm apart. Inflorescence

2-3 m high, erect, compactly 6-10-branched, lower

branches sometimes 1- or 2-rebranched. Raceme cylin-

dric-acuminate, 150-300 mm long, terminal raceme

longest, lateral racemes shorter, dense. Floral bracts

8-18 x 3-4 mm. Pedicels 8-20 mm long. Flowers', peri-

anth bright coral-pink with a bloom and obscure nerves,

lighter and brownish at tips, 35-45 mm long, 8-10 mm

across ovary, not narrowed above ovary, cylindric-trigo-

nous, straight; outer segments free for 15 mm. Stamens

exserted 3^1 mm. Style exserted 4—5 mm. Flowering

time: April to July.

Diagnostic characters', rosette large. Inflorescence

tall and branched. Floral bracts small, up to 18 mm long,

pedicels short, 8-20 mm long. Flowers cylindric-trigo-

nous, with thick, fleshy perianth and outer segments free

for only 15 mm.

Relationships with other species : Aloe christianii is

closely related to A. pretoriensis Pole Evans from north-

ern South Africa and Zimbabwe (Reynolds 1966).

Habitat: usually in shade in woodland, sometimes in tall

grassland, never on rocks.

Distribution : eastern Angola (Moxico), southern and

eastern Democratic Republic of the Congo, Malawi,

northern Mozambique, eastern Tanzania, Zambia, north-

ern Zimbabwe (Figure 3).

Illustrations'. Reynolds: 187, 188 (1966); Lane: 32 (2004).

Notes: this species has not yet been collected in Angola,

but it was reportedly seen in Angola near the Zambian

border west of Matonchi by Mr Edgar Milne-Redhead

(Reynolds 1966; Carter 2001).

A. dinteri A. Berger in Dinter, Neue und wenig

bekannte Pflanzen Deutsch-Sudwest-Afrikas: 14 (1814).

Type: Namibia, Outjo, Dinter 2791a (SAM, holo.!).

Acaulescent, 0.26 m high; rosettes solitary. Leaves tri-

farious, spreading to almost horizontal, chocolate-brown

or deep brownish green, copiously spotted with crowded

narrowly elongated white spots arranged ± in broken trans-

verse bands, with small white teeth on white cartilaginous

edge of keel, narrowly lanceolate-acuminate, plicate-cari-

nate, 200-300 mm long, 50-80 mm wide at base; margin

narrow, white, with minute white teeth, ± 0.5 mm long,

1-2 mm apart. Inflorescence 0.5-0.85 m high, erect, 3-8-

branched. Raceme cylindric-acuminate, 150-200 mm long,

terminal raceme the longest, lax. Floral bracts 7-12 x ± 2

mm. Pedicels 10-15 mm long. Flowers', perianth pale rose-

pink with bluish bloom, pale to almost white at mouth,

28-30 mm long, ± 6.5 mm across ovary, abruptly narrowed

above ovary, enlarging towards mouth, slightly decurved;

outer segments free for 5-10 mm. Stamens included or

exserted up to 1 mm. Style exserted up to 1 mm. Flowering

time: January to March.

Diagnostic characters', acutely folded, trifarious,

chocolate-brown leaves of up to 300 mm long, with mar-

gins finely toothed. Inflorescence 3-8-branched and up

to 0.85 m high. Floral bracts 3-nerved.

Relationships with other species'. Aloe dinteri, together

with its close relatives, A. sladeniana Pole Evans from

central Namibia and A. variegata L. from southern, west-

ern and central South Africa and southern Namibia, forms

the Section Serrulatae Salm-Dyck (Glen & Hardy 2000).

Habitat', usually wedged firmly in cracks in limestone in

areas of very low summer rainfall. Sometimes on gran-

ite, in bushveld near edge of Namib Desert.

FIGURE 4. — Distribution of Aloe dinteri , •; A. gossweileri, ■; and

A. grata, ▲ .

26

Bothalia 39,1 (2009)

Distribution : southwestern Angola (Namibe, Cunene),

northern Namibia (Figure 4).

Illustrations'. Reynolds: t. 637 (1936b); Reynolds: 211

(1950); Rothmann: 58, 59 (2004).

Note : this species was previously considered to be

endemic to northern Namibia and occurs near the

Cunene River west of Ruacana. Due to its frequent proxi-

mity to the river, it was believed likely to occur north of

the Cunene River in Angola (Hardy 1992). In fact, a few

plants were reportedly seen near Namibe (Mofamedes)

on low stony ridges and in stony desert areas (Downs

1970). However, no herbarium specimens could be

found to confirm this distribution. Clair Bell (pers.

comm.) saw this species north of the Cunene River in

Angola in 1996. A plant collected during this expedition

is currently growing in the Botanical Society Conserva-

tory at Kirstenbosch National Botanical Gardens, Cape

Town (SW Angola, July 1996, Clair Bell 1120/96). How-

ever, opinions are divided on the correct identification of

this taxon in Angola. According to John Lavranos (pers.

comm.) a plant collected inland from Namibe by Dr

Philip Downs in 1971 suckered freely and did not have

the solitary rosettes characteristic of Aloe dinteri. Fur-

thermore, it bore leaves of no longer than 50-70 mm and

consistently produced simple inflorescences. Lavranos,

therefore, considers it to be closer to A. sladeniana.

A. esculenta L.C. Leach in Journal of South Afri-

can Botany 37: 249 (1971). Type: Angola, near Pereira

d’E9a, 7 August 1967, Leach & Canned 13818 (PRE,

holo.!; BM, K!, LISC!, SRGH, iso.).

Acaulescent or sometimes with short, thick, often

decumbent stem, up to 0.4 m long; rosettes suckering to

form dense clumps. Leaves condensed rosulate, erectly

spreading or recurved, greyish green, with pinkish brown

tinge in dry conditions, densely white-spotted on both sur-

faces, spots more copious on lower surface, arranged in

irregular transverse bands, pungent blackish brown spines

along median line, lanceolate, up to 500 mm long, 80 mm

wide at base; margin with pungent, blackish brown teeth,

3-5 mm long, 10-20 mm apart; leaf exudate not bitter.

Inflorescence up to 2 m high, erect, 3-5 -branched, lower

branches sometimes sparsely rebranched. Raceme cylin-

drical-acuminate, 300-500 mm long, lax. Floral bracts

20-27 x 10-11 mm. Pedicels 5-6 mm long. Flowers : peri-

anth pink with cream-coloured stripes, becoming yellow-

ish when mature, 28-30 mm long, ± 6 mm across ovary,

enlarging towards middle, distinctly subclavate; outer seg-

ments free for 15-18 mm. Stamens exserted up to 6 mm.

Style exserted up to 8 mm. Flowering time : July to August.

Diagnostic characters', spotted aloe. Acaulescent

plants forming clumps, frequently of shrubby habit.

Leaves copiously white-spotted, with large spots

arranged in transverse bands. Strong blackish brown

spines along median line on underside of leaf. Flowers

distinctly subclavate; stigma exserted up to 8 mm.

Relationships with other species', at first sight this species

looks very much like a stemless form of Aloe littoralis ,

to which it is closely allied. Leach (1971) stated that it

may well be considered as an ecologically separated sub-

species of A. littoralis. Glen & Hardy (2000) regarded it

as conspecific with A. angolensis, with the latter name

taking priority according to the rules of the International

Code of Botanical Nomenclature (McNeill 2006).

Habitat', sandy soils of flats of inland drainage areas.

Distribution : southern Angola (Cunene, Huila), north-

western Botswana, northern Namibia, southwestern Zam-

bia (Figure 5).

Illustrations'. Rothmann: 62, 63 (2004).

Additional specimens examined

ANGOLA.— 1515: Huila, (-AC), Carrisso & Sousa 193 (LISC).

1614: Huila, Royadas, (-DB), 8 July 1970, Menezes 3384 (K, LISC,

PRE, SRGH); Cunene, Cuamato, Rofadas, entre Dimba e Humbe, (—

DB), 20 July 1970, Santos & Barroso 2761 (LISC, PRE). 1715: Baixo

Cunene, Cuamato, Rofadas, a 34 km para a Missao do Cuamato, (-

AA), 1 July 1970, Santos & Barroso 2695 (LISC, PRE); Huila, Ro?a-

das, Cuamato, (-AA), 4 July 1970, Menezes 3379 (LISC, PRE); Baixo

Cunene, Cuanhama, Pereira d’Eya, entre Namacunde e Chiede, ( -BB).

9 July 1970, Santos & Barroso 2717 ( LISC, PRE).

EA. gossweileri Reynolds in Journal of South

African Botany 28: 205 (1962). Type: Angola, Cuanza

Sul District, Seles, 7 miles [11 km] SE of Vila Nova de

Seles, 15 July 1961, Reynolds 9760 (PRE, holo.!; K!;

LISC!, iso.).

Thicket-forming shrub. Stem 1.0-1. 5 m long, branch-

ing from ground level only, ascending or divergent,

without persistent dried leaves. Leaves subdensely rosu-

late at branch apices, spreading to slightly recurved,

green, mostly without spots, lanceolate-attenuate, ± 300

mm long, 50 mm wide at base; margin with pale deltoid

teeth, 3—4 mm long, 15 mm apart. Inflorescence 0.4-0. 5

m high, erect, pyramidal, divaricately 6-8-branched.

Raceme cylindric-acuminate, 100-150 mm long, termi-

nal raceme the longest, dense, flowers subsecund when

open. Floral bracts 3x2 mm. Pedicels 10 mm long.

Flowers', perianth scarlet, paler at tips, 30 mm long, 6

mm across ovary, very slightly narrowed above ovary,

slightly enlarging towards trigonal mouth; outer seg-

ments free for 10-12 mm. Stamens exserted 1-2 mm.

Style exserted 2-3 mm. Flowering time'. March.

Diagnostic characters : forms thickets. Stems branched

at ground level only, 1.0-1. 5 m long. Inflorescence divari-

cately branched, pyramidal with almost horizontal racemes

with subsecund flowers.

Relationships with other species: its closest ally appears

to be Aloe palmiformis (Reynolds 1966).

Habitat: rocky hills.

Distribution: endemic to Angola (Benguela, Cuanza Sul)

(Figure 4).

Illustrations: Reynolds: 372 (1966).

Additional specimens examined

ANGOLA.— 1114: Vila Nova de Seles, (-AD), March 1941, Gos-

sweiler 13313 (LISC, para.). 1214: Benguela, Bocoio, a caminho da

povoa^ao da Chicuma, andados ± 40 km da povoafao de Chila, (-AA),

24 May 1973, Raimundo, Matos & Figueira 1415 (LISC).

EA. grata Reynolds in Journal of South African

Botany 26: 87 (1960). Type: Angola, Bie District, 30

miles [48 km] S of Chinguar, on Chimbango Hill, 19

June 1959, Reynolds 9246 (PRE, holo.!; K!; LUA, iso.).

Acaulescent or with short stem, suckering to form

dense groups of rather compact rosettes. Leaves densely

rosulate, erect to erectly spreading, upper surface green,

Bothalia 39,1 (2009)

27

tinged reddish brown, without spots, lower surface paler

glaucous green, with many crowded, pale green, circular,

1 mm spots in lower quarter, lanceolate-attenuate, 200-

250 x 70-80 mm; margin sinuate-dentate, sometimes

almost serrate, with teeth 2-3 mm long, 5-8 mm apart;

leaf exudate drying pale yellow. Inflorescence 0.7-0. 9

m high, erect, simple in young plants, up to 3-branched

in older plants. Raceme capitate or subcapitate, 80-100

mm long, fairly dense. Floral bracts 2 x 1.5 mm. Pedi-

cels ± 20 mm long. Flowers : perianth scarlet, 25-28

mm long, 6 mm across ovary, slightly narrowed above

ovary, enlarging towards middle, slightly narrowing at

mouth, trigonous; outer segments free for 7 mm. Sta-

mens exserted 1-2 mm. Style exserted 2-3 mm. Flower-

ing time : June.

Diagnostic characters: leaves with small crowded mar-

ginal teeth that are sometimes almost serrate, lower surface

with copious white, small, circular spots near base. Inflor-

escence simple in young plants, forked in older plants, 2-

or 3-branched in largest specimens. Racemes capitate or

almost so. Perianth scarcely trigonously indented above

ovary. Floral bracts 2 mm long, 1 -nerved.

Relationships with other species: the nearest ally to this

species seems to be Aloe mzimbana Christian from cen-

tral and southeastern tropical Africa (Reynolds 1966).

Habitat: hillsides, mostly on rocks.

Distribution: endemic to Angola (Benguela, Bie, Huambo)

(Figure 4).

Illustration: Reynolds: 117 (1950).

Additional specimen examined

ANGOLA. — 1215: Benguela, Serra do Moco, (-AC), 3 June 1940,

Gossweiler 12597 (LISC).

EA. guerrae Reynolds in Journal of South African

Botany 26: 85 (1960). Type: Angola, Bie District, E of

General Machado, 18 June 1959, Reynolds 9218 (PRE,

holo.!; K!, LUAI, iso.).

Acaulescent or with very short stem; rosettes single,

usually slightly tilted to one side. Leaves densely rosu-

late, suberectly spreading to spreading with slightly

recurved apical portion, dull green, obscurely lineate on

upper surface, grey-green with no markings on lower

surface, lanceolate-attenuate, ± 400 mm long, 60-70

mm wide at base; margin with pungent, pale brown or

reddish brown teeth, 4—5 mm long, 10-15 mm apart;

leaf exudate drying yellow. Inflorescence 0.9-1 m high,

erect, divaricately 8-10-branched, lower branches some-

times rebranched. Raceme cylindrical, ± 200 mm long,

oblique to almost horizontal, dense, buds and flow-

ers secund, almost erect. Floral bracts 6-8 x 4 mm.

Pedicels ± 5 mm long. Flowers: perianth scarlet with a

bloom, 40 mm long, 8 mm across ovary, scarcely nar-

rowed above ovary, cylindric-trigonous towards slightly

upturned mouth, straight or slightly curved; outer seg-

ments free for 10-12 mm. Stamens exserted up to 2 mm.

Style exserted up to 3 mm. Flowering time: May to June.

Diagnostic characters: inflorescence divaricately branched

with oblique to subhorizontal racemes of secund flowers.

Relationships with other species: its nearest ally is Aloe

secundiflora Engl, from eastern and northeastern Africa

(Reynolds 1966).

FIGURE 5. — Distribution of Aloe esculenta, •; A. guerrae, ®; A. ina-

mara , ▲; and A. vallaris , ■

Habitat: grassland with scattered bushes.

Distribution: endemic to Angola (Benguela, Huambo)

(Figure 5).

Illustration: Reynolds: 229 (1966).

Additional specimens examined

ANGOLA. — 1214: Balombo, 23 miles [37 km] E of Monte Belo,

6 miles [9.6 km] W of Balombo, (-BC), 3 July 1959, Reynolds 9325

(PRE). 1215: Benguela, Nova Lisboa, (-AA), 9 May 1937, Exell &

Mendonga 1677 (LISC); Benguela, Calupiango, (-AD), 31 May 1940,

Gossweiler 12600 (LISC); Bie Province, Posto do Cunje, near Silva

Porto, (-BD), 1 August 1955, Reynolds 6885 (PRE).

A. hereroensis Engl, in Botanische Jahrbiicher 10:

2 (1888), var. hereroensis. Type: Namibia, Usakos, 28

May 1886, Marloth 1438 (B, holo.!; NBG!, PRE!, iso.).

Acaulescent; rosettes solitary or suckering to form

small clumps. Stem sometimes develops in older plants,

up to 1 m long, simple or branched, procumbent, with

persistent dried leaves. Leaves densely rosulate, arcuate

erect, greyish green or often bronzed, obscurely line-

ate, usually without spots on upper surface, few to many

irregularly scattered single or double H-shaped whitish

spots on lower surface, ovate-lanceolate, 300-400 mm

long, 60-90 mm wide at base; margin slightly cartilagi-

nous, with pungent, reddish brown, spreading, sometimes

bifid teeth, 3^1 mm long, 8-12 mm apart. Inflorescences

up to 1 m high, erect, 4— 8-branched, lower branches

rebranched. Raceme corymbose-capitate, 60-80 mm long,

very dense. Floral bracts 20-25 x 3^J mm. Pedicels

30-50 mm long. Flowers: perianth scarlet, sometimes

orange, 25-35 mm long, 8-9 mm across ovary, widen-

ing slightly towards middle, narrowing slightly towards

upturned mouth, cylindric-trigonous; outer segments free

for 14-16 mm. Stamens exserted 2-4 mm. Style exserted

up to 5 mm. Flowering time: June to September.

Diagnostic characters: leaves obscurely lineate with

spotted lower surface. Much branched inflorescence with

short, broad racemes. Flowers distinctly shaped, with

upturned mouth, red or sometimes orange.

Relationships with other species: Aloe hereroensis is

included in Section Asperifoliae (A. Berger) Glen &

28

Bothalia 39,1 (2009)

D.S. Hardy and is closely allied to A. viridiflora Giess

from central Namibia (Glen & Hardy 2000).

Habitat : arid, stony desert conditions with very low rain-

fall. Usually grows on quartzite, but has been found on

dolomite in the Kalahari. Confined to summer rainfall area.

Distribution : southwestern Angola (Huambo, Namibe),

Namibia, South Africa (Northern Cape, Free State) (Fig-

ure 6).

Illustrations : Reynolds: 101 (1966); Rothmann: 66, 67

(2004).

Additional specimens examined

ANGOLA. — 1215: Huambo, entre Alto Hama e Aguas Quentes, (-

BA), 20 August 1967, Silva 2104 (LISC). 1512: Mofamedes, andados

30 km de Mofamedes para Dois Irmaos, ( -AB), 2 May 1960, Mendes

3898 (LISC); Benguela Province, 20 miles [32 km] NE of Mofamedes,

(-AB), 28 June 1959, Reynolds 9283 (PRE); Mofamedes, Caraculo, a

± 25 km para Mofamedes, (-AB), 7 May 1962, Santos 1007 (LISC,

SRGH); Mofamedes, Reserva de Mofamedes junto ao limite NE, (—

DA), 5 March 1969, Teixeira 12871 (LISC); Mofamedes, Reserva de

Mofamedes, (-DA), 11 April 1969, Teixeira 12942 (LISC).

EA. inamara L.C. Leach in Journal of South Afri-

can Botany 37: 259 (1971). Type: Angola, Cuanza Sul

District, S of Novo Redondo, at mouth of Quicombo

River, 11 October 1970, Leach & Cannell 14608 (LISC,

holo.!; PRE, iso.!).

Plants hanging on cliff faces. Stem up to 2 m long,

pendent, branching at base and more sparsely above,

forming dense mats, without persistent dried leaves.

Leaves rosulate on branch apices, widely spreading,

curved with apices pointing downwards, pale yellow-

ish green, turning brown when exposed, obscurely line-

ate with few small, whitish, H-shaped spots, more spots

in transverse bands on lower surface, falcate, 450-600

x 40-50 mm; margin whitish or faint pink, with whit-

ish, often brown-tipped teeth, 0.3-1 mm long, 4-20 mm

apart; leaf exudate yellow, not bitter. Inflorescence 0.4—

0.55 m long, pendulous with tip upturned, 4— 6-branched.

Raceme shortly cylindrical-conical to almost subcapitate,

terminal raceme ±75 mm long, lateral racemes shorter,

arcuate-ascending, lax. Floral bracts 7. 5-9.0 x ± 3 mm.

Pedicels 22-27 mm long. Flowers : perianth dull red with

greenish tips, becoming somewhat yellowish with age,

26-29 mm long, 8 mm across ovary, abruptly narrowed

above ovary, enlarging towards middle, narrowing again

at mouth, slightly decurved, basally truncate and inflated;

outer segments free for 6. 5-8.0 mm. Stamens not or only

rarely slightly exserted. Style scarcely or not exserted.

Flowering time : apparently an extended period including

October.

Diagnostic characters', leaves obscurely lineate with

H-shaped spots. Racemes short, sometimes almost sub-

capitate. Pedicels 22-27 mm long. Flowers dull red with

much inflated truncate base and genitalia scarcely or not

exserted.

Relationships with other species', its nearest ally appears

to be Aloe swynnertonii Rendle from eastern south tropi-

cal Africa (Leach 1971).

Habitat : almost vertical cliff faces.

Distribution', endemic to Angola (Cuanza Sul) (Figure 5).

Illustrations'. Leach: 261, 263, 264 (1971).

Notes', this species is only known from the type locality.

EA. lepida L.C. Leach in Journal of South African

Botany 40: 102 (1974). Type: Angola, Huambo Dis-

trict, Morro de Sume, ± 27 km SSE of Nova Lisboa, 15

February 1973, Baptista de Sousa s.n. in Leach 14538A

(LISC, holo.!; SRGH, iso.).

Low, much-branched shrub, up to 0.3 m high. Stem

branching at base, stout, erect, without persistent dried

leaves. Leaves densely rosulate at branch apices, widely

spreading, rigidly strongly recurved, bright to dark yel-

lowish deep green, conspicuous but irregular white spots

in wavy transverse bands, spots smaller and more numer-

ous on lower surface, broadly ovate-attenuate, 200-280

x 75-90 mm; margin with pungent, brown-tipped teeth

with whitish base, 3-7 mm long, 6-12 mm apart; exudate

crusty when dry. Inflorescence 0.3-0. 5 m high, erect, 1-

or 2-branched. Raceme cylindrical-acuminate, ± 200 mm

long, lax. Floral bracts 6-7 x 3. 0-3. 5 mm. Pedicels 15-20

mm long. Flowers : perianth pale orange-scarlet, some-

what yellowish striped, 25-29 mm long, ± 5.5 mm across

ovary, narrowed above ovary, enlarging towards wide open

mouth, cylindric, slightly curved; outer segments free for

5-6 mm. Stamens not exserted. Style only occasionally

very shortly exserted. Flowering time : February to May.

Diagnostic characters', leaves dark yellowish green,

strongly recurved, conspicuously marked with irregular

transverse wavy bands of whitish spots, armed with large

marginal teeth. Inflorescence with lax, cylindric-acumi-

nate racemes, with buds quickly nodding. Flower slender

with outer segments free for 5-6 mm, with mouth widely

open, stamens included and stigma only occasionally

very shortly exserted.

Relationships with other species : Aloe lepida seems to

be closely related to A. andongensis and also shows a

strong link in vegetative characters with A. squarrosa

Baker from Socotra (Leach 1974).

Habitat : rocky slopes in shade of trees.

Distribution : endemic to Angola (Huambo) (Figure 6).

Illustrations'. Leach: 103, 104, 105 (1974).

Additional specimen examined

ANGOLA. — 1315: Huambo District, Morro de Sume, ± 27 km

SSE of Nova Lisboa, (-BA), 14 May 1973, Leach, Canned & De

Sousa 14538 (BM, PRE).

FIGURE 6. — Distribution of Aloe hereroensis var. hereroensis, •;

A. lepida, A ; A. mendesii, ■; and A. metallica, ®.

Bothalia 39,1 (2009)

29

A. littoralis Baker in Transactions of the Linnean

Society of London 1: 263 (1878). Type: Angola, near

Luanda, Barra do Bengo, 1854, Welwitsch 3727 (BM,

holo.!; K!, LISU!, iso.-PRE, photo.).

A. rubrolutea Schinz: 39 (1896). Types: Namibia, Rehoboth, Fleck

497a ; Namibia, IKuisib, Fleck 47 2\ Botswana, Olifantskloof, Fleck

263 (Z, syn.).

A. schinzii Baker: 459 (1898). Type: Botswana, Olifantskloof, April

1888, Schinz 42 (K, holo.!).

Solitary, arborescent plant 2-4 m high. Stem un-

branched, erect, with persistent dried leaves. Leaves

densely rosulate, erectly spreading to slightly recurved,

pale greyish green with reddish tinge in dry conditions,

without spots when mature, few spots on young plants,

lanceolate, ± 600 mm long, 100-130 mm wide at base;

margin cartilaginous, yellow, with pungent, red-brown

teeth, 3-4 mm long, 10-15 mm apart; leaf exudate dry-

ing yellow. Inflorescence 1-2 m high, erect, up to 10-

branched, lower branches usually rebranched. Racemes

narrowly cylindrical-acuminate, 300-600 mm long, lax.

Floral bracts 12-18 x 5-6 mm. Pedicels 6-10 mm long.

Flowers : perianth pinkish red, yellowish towards tips

when mature, with bloom, 27-34 mm long, 6 mm across

ovary, enlarging very slightly towards middle, cylindri-

cal and almost straight; outer segments free for 15-17

mm. Stamens exserted 1-2 mm. Style exserted 2-3 mm.

Flowering time : July to February, varying according to

locality and rainfall.

Diagnostic characters : caulescent, simple-stemmed,

solitary plants. Leaves without spots when mature, few

spots on young plants. No spines along median line of leaf.

Inflorescence branched from low down. Flowers cylindri-

cal, stigma seldom if ever exserted more than 3 mm.

Relationships with other species : closely allied to Aloe

esculenta (see notes under latter species). A. littoralis is

included in Section Pachydendron (Haw.) Salm-Dyck

(Glen & Hardy 2000).

Habitat : usually grows on rocky outcrops in mixed open

woodland and grassland. Also on calcrete or sand. Sum-

mer rain and very dry, warm to cool winters.

Distribution : Angola (Bengo, Benguela, Cuanza Norte,

Cuanza Sul, Huambo, Huila, Luanda, Lunda Norte,

Malange, Moxico, Namibe), Botswana, western Mozam-

bique, Namibia, South Africa (Limpopo), southern Zim-

babwe (Figure 7).

Illustrations'. Reynolds: 317, 318 (1966); Rothmann: 68,

69 (2004).

Additional specimens examined

ANGOLA. — 0720: Arredores de Dundo, (-BD), April 1953, S.c.

135 (LISC). 0813: Luanda, (-CB), Exell & Mendonga 37 (LISC);

Luanda, proximo da Corimba, (-CC), 29 August 1961, Santos 433

(LISC); Luanda, near Cacuaco, 10 miles [16 km] NE of Luanda,

(-CD), 12 July 1959, Reynolds 9402 (PRE); Luanda, Viana, Vale do

Bengo, (-CD), 23 April 1966, Teixeira 10326 (LISC). 0914: Luanda.

Cambambe-Dondo, (-CB), 18 August 1931. Gossweiler 9633

(LISC); Cuanza Norte. Dondo, Rio Cuanza, (-CB), March 1938,

Gossweiler 11963 (LISC); Cuanza Norte, Dondo, (-CB), Gossweiler

12613 (LISC). 0915: Malange, Cacuso, proximo do salto do Cavalo,

(-DB), 15 July 1970, Raimundo 327 (LISC); Malange, na picada que

vai da Aldeia Formosa para o salto do Cavalo, (-DB), 15 July 1970,

Raimundo. Matos & Figueira 327 (LISC). 1013: Foz de Cuvo, (-DD),

30 March 1973. Bamps & Martins 4342 (LISC). 1113: Cuanza Sul,

Gabela, picada para o Pau do Capador e Gungo a 43 km de Novo

FIGURE 7. — Distribution of Aloe littoralis, •; and A. milne-redheadii,

©.

Redondo, (-DB), 19 April 1969, Teixeira 11430 (LISC). 1121: Lago

Calundo, (-CB), January 1955, Machado 358 (LISC). 1214: Lobito,

9 miles [14.5 km] W of Bocoio (Sousa Lara), 49 miles [79 km] E of

Lobito, (-AC), June 1959, Reynolds 9317 (PRE). 1215: Benguela,

Chicala-Calenga, near Sete River, (-CD), 19 June 1940, Gossweiler

12598 (LISC). 1313: Benguela, Catengue, (-BA), 29 July 1940, Goss-

weiler 12619 (LISC). 1314: Benguela, pres de Caimbambo, (-AA), 17

February 1974, Dechamps, Murta & Silva 1095 (LISC). 1413: Serra da

Chela, Vila Arriaga, Bibala, (-CD), 12 August 1941, Gossweiler 13297

(LISC); Huila, Mofamedes, Bibala, Cacanda, (-CD), 7 May 1960,

Mendes 4003 (LISC); Sa da Bandeira. Mapunda, (-CD), 16 June 1963,

Santos 1126 (LISC, LISU), Huila, Sa da Bandeira, Mapunda, (-CD),

16 June 1963, Santos 1128 (LISC, LISU); Mofamedes, Vila Arriaga,

Montipa, (-CD), Teixeira & Santos 3855 (SRGH). 1414: Huila, at Vila

Paiva Couceiro (Quipuno), 28 miles [45 km] E of Sa da Bandeira, (-

DC), 25 June 1959, Reynolds 9267 (PRE). 1512: Mofamedes, 30 km

a ocidente do Pico do Azevedo, (-AD), 9 March 1969, Teixeira 12888

(LISC). 1513: Huila, 11 km do Jau para Bata Bata, (-AB), 30 February

1961, Barbosa 9554 (PRE); Serra da Chela, Tchivinguiro, Humpata,

(-AB), 6 October 1941, Gossweiler 13346 (LISC). No grid: Cascata

do Rio Cuango Muque, 16 July 1954, Machado 200 (LISC).

EA. mendesii Reynolds in Journal of South Afri-

can Botany 30: 31 (1964). Type: Angola, Huila District,

Humpata, Tundavala escarpment, 4 July 1963, Santos &

Henriques 1131 (LISC, holo.!; LISU!, LUAI, PRE!, iso.).

Plants growing pendent on vertical cliff faces. Stem

usually unbranched, up to 1 m long, pendent, without

persistent dried leaves. Leaves rosulate at stem apex,

hanging downwards, green, obscurely lineate, without

spots, ensiform, falcately decurved, 500 x 70-80 mm;

margin narrow cartilaginous edge, with blunt, cartilagi-

nous teeth, 1-2 mm long, 10-15 mm apart. Inflorescence

up to 0.6 m long, pendent, 3- or 4-branched. Racemes

cylindrical-acuminate, 100 mm long, arcuate-ascending,

dense; buds hidden by imbricate bracts. Floral bracts 12

x 5 mm. Pedicels 18-20 mm. Flowers', perianth scar-

let, 25 mm long, 4 mm across ovary, enlarging towards

mouth, narrowing just below mouth, cylindric, slightly

ventricose; outer segments free for 20 mm. Stamens

exserted 2-3 mm. Style exserted up to 3 mm. Flowering

time'. April to July.

Diagnostic characters', grows hanging down on cliffs.

Leaves broad, unspotted. Branched inflorescence pen-

dent, with only racemes arcuate-ascending. Flowers

scarlet. Floral bracts and pedicels relatively long.

30

Bothalia 39,1 (2009)

Relationships with other species : appears to be closely

allied to Aloe veseyi Reynolds from Zambia (Reynolds

1966).

Habitat', vertical cliff faces.

Distribution : endemic to southwestern Angola (Huila,

Namibe) (Figure 6).

Illustrations'. Verdoom: t. 1764 (1964); Reynolds: 170,

171 (1966).

Notes: this species is sometimes wrongly recorded as

occurring in northwestern Namibia (Newton 2001).

The species in Namibia with which it is mistaken is the

Kaokoland endemic A. corallina I.Verd.

Additional specimens examined

ANGOLA. — 1413: Huila, Sa da Bandeira, Tundavala, (-CD),

October 1967, Leach 14015 (PRE); Huila, Lubango, Humpata, Buraco

do Bimbe, (-CD), 22 April 1960, Mendes 3815 (LISC, para.); Huila,

Lubango, Escarpa da Tundavala (serra da Chela), (-CD), 20 May 1964,

Menezes 1133 (LISC, PRE); Huila, Sa da Bandeira, Humpata, Bimbe,

(-CD), 3 May 1963, Santos 1109 (LISC, para.).

EA. metallic;! Engl. & Gilg in Warburg, Kunene-

Zambesi Expedition: 191 (1903). Type: Angola, Bie Dis-

trict, near Cuchi, above Capulo, 4 May 1900, Baum 891

(B, holo.!-LISC, photo.!).

Acaulescent or stem short; rosettes single. Leaves

densely rosulate, erectly spreading, bluish grey with

metallic sheen (sometimes lost, especially in cultiva-

tion), without spots or markings, broadly lanceolate-atte-

naute, 250-400 mm long, 70-90 mm wide at base; mar-

gin slightly reddish brown, homy, with pungent, reddish

brown teeth, 2-3 mm long, 10-20 mm apart. Inflores-

cence up to 1 .2 m high, erect, sparsely branched. Racemes

narrowly cylindrical-acuminate, terminal raceme 350

mm long, dense; buds hidden by large, densely imbricate

bracts. Floral bracts 1 8-20 x 8 mm. Pedicels 8 mm long.

Flowers', perianth reddish pink, 32 mm long, 7 mm across

ovary, enlarging slightly from middle towards mouth,

slightly compressed-trigonous; outer segments free for 13

mm. Stamens exserted up to 2 mm. Style exserted up to 3

mm. Flowering time : April to July.

Diagnostic characters: bracts long, white, promi-

nently nerved. Apical buds of racemes at first covered by

densely imbricate bracts.

Relationships with other species: Reynolds (1966)

placed Aloe metal lica in his Group 9. It keys out with

A. massawana Reynolds (now a synonym of A. eumas-

sawana S. Carter & M.G. Gilbert) from eastern Africa, A.

vacil/ans Forssk. and A. officinalis Forssk. from Yemen.

Habitat: top of rocky gorge, between high sandstone rocks.

Distribution: endemic to Angola (Cuando-Cubango,

Luanda) (Figure 6).

Illustration: Reynolds: 152 (1966).

Additional specimens examined

ANGOLA. — 0713: Luanda, estradada Corimba, (-CC), 7 March 1971,

Henriques 1329 (LISC, LISU). 1416: Cuchi River Gorge, (-DD), Leach

13850 (PRE); Menongue, Cuchi, Caquima, Rio Cuchi, (-DD), 3 April

1960, Mendes 3444 (LISC), 16 July 1964, Reynolds 10104 (LISC, PRE).

A. milne-redheadii Christian in Journal of South

African Botany 6: 177 (1940a). Type: Angola, Moxico

District, between River Zambezi and River Lusavo, Milne-

Redhead 4253 ex Christian 926 (SRGH, holo.; K, iso.).

Acaulescent or with short stem, suckering to form

dense clumps. Leaves densely rosulate, erect and curv-

ing inward, greyish green to brownish green, becoming

slightly reddish in dry conditions, densely white-spotted

on both surfaces, spots more copious on lower surface

and arranged in transverse bands, ovate-lanceolate, 200-

300 mm long, 60-80 mm wide at base; margin cartilagi-

nous, with minute, brown-tipped teeth, 2-3 mm long,

10-15 mm apart. Inflorescence 0.6-0. 9 m high, erect,

1-7-branched. Raceme narrowly cylindrical-acumi-

nate, 200-250 mm long, dense. Floral bracts 5-6 x 2-3

mm. Pedicels 13-18 mm long. Flowers: perianth bright

coral-red, 28-35 mm long, ± 8 mm across ovary, slightly

narrowed above ovary, curved and widening towards

mouth, cylindrical-trigonous; outer segments free for 10

mm. Stamens exserted 0-1 mm. Style exserted 1-2 mm.

Flowering time: June to July.

Diagnostic characters: racemes short, rather dense.

Pedicels long. Leaves copiously spotted with spots in

transverse bands. Plants forming clumps.

Relationships with other species: its nearest ally is Aloe

bukobana Reynolds from Rwanda, Burundi and Tanza-

nia (Reynolds 1966).

Habitat: ridges of hills.

Distribution: eastern Angola (Moxico), northwestern

Zambia (Figure 7).

Illustration: Reynolds: 110 (1966).

Additional specimen examined

ANGOLA. — 1123: Moxico District, between River Zambezi and

River Lusavo, (-DA), material from type plant, 7 July 1941, Verdoorn

PRE29568 (PRE).

A. nuttii Baker in Hooker’s, leones plantarum:

t. 2513 (1897). Type: Zambia, Fwambo, South of Lake

Tanganyika, 1896, Nutt s.n.; Zambia, Fwambo, 1894,

Carson 29 (K, syn.!-PRE, photo.!).

A. brunneo-punctata Engl. & Gilg: 189 (1903). Type: Angola,

Longa, oberh. Minnesera, 2 February 1900, Baum 698 (B, holo.!; BR!,

E!, K!, M!, iso.-PRE, photo.!).

A. corbisieri De Wild.: 29 (1921). Type: DRC, Katanga, Elisabeth-

ville, Welgelegen, 1912, Corbisier 623 (BR, syn.!); DRC, Lukafu, Feb-

ruary 1900, Verdick 417 (BR, syn.!).

A. mketiensis Christian: t. 785 (1940b). Type: Tanzania, Inringa

District, Sao Highlands, north of Mketi, 17 June 1938, Pole Evans c£

Evens 795 (PRE24803) (PRE, holo.!).

Grass aloe, growing singly or with 2 or 3 stems, some-

times up to 12 and more tufted stems. Stems very short or

up to 0.2 m long, erect. Leaves rosulate, erectly spread-

ing, sometimes deflexed at about middle, green, upper

surface sometimes obscurely lineate, usually with few

pale spots near base, lower surface usually copiously

spotted near base, spots occasionally spinulescent, grass-

like and subfleshy, linear, 400-500 mm long, up to 40

mm at dilated base, abruptly narrowed to 15-20 mm,

tapering towards apex; margin very narrow, white, with

densely crowded, minute, white, soft, cartilaginous teeth,

up to 1 mm long. Inflorescence 0.6-0. 8 m high, erect,

unbranched. Raceme cylindrical-acuminate, 150-200

mm long, dense; buds entirely covered by large imbri-

cate bracts. Floral bracts 15-25 x 10-20 mm. Pedicels

25-35 mm long. Flowers: perianth coral-pink to orange-

red with green tips, 35^42 mm long, 7-9 mm across

Bothalia 39,1 (2009)

31

ovary, not narrowed above ovary, cylindrical-trigonous,

base tapering into pedicel; outer segments free almost

to base or for 3A of length. Stamens not exserted. Style

exserted 0-1 mm. Flowering time : January to March,

depending on locality and rainfall.

Diagnostic characters', leaves grass-like, rosulate. Flow-

ers salmon-pink.

Relationships with other species : its nearest ally is Aloe

buchananii Baker from Malawi (Reynolds 1966).

Habitat', montane grassland, often on rocky slopes.

Distribution : southeastern Angola (Cuando-Cubango),

southern Democratic Republic of the Congo, Malawi,

southwestern Tanzania, northern Zambia (Figure 8).

Illustrations'. Christian: t. 762 (1940c); Reynolds: 33, 34

(1966); Lane: 9(2004).

Additional specimens examined

ANGOLA. — 1417: Menongue, Vila Serpa Pinto, vale do Rio

Cabumbe, (-DB), 15 February 1960, Mendes 2575 (LISC). 1517:

Menongue, andados 50 km de Caiundo para Vila Serpa Pinto, (-BC),

4 February 1960, Mendes 2354 (LISC). 1519: Bie, Menongue, Rio

Cuito, Vale do ribeiro Sobi, (-AC), 13 March 1906, Gossweiler 4213

(LISC). 1718: Menongue, entre Longa e Vila Serpa Pinto, vale do

Luassingua, (-CA), 22 March 1960, Mendes 3243 (LISC). No grid:

Longa, oberh. Minnesera, 2 February 1900, Baum 698 (BM. BR. E, K,

M-PRE, photo.); Moxico, between Kangeshi River and Kaboli River,

20 January 1938, Milne-Redhead 4260 (K- PRE. photo.).

A. paedogona A. Berger in Journal of Botany, Brit-

ish and Foreign 44: 57 (1906). Type: Angola, Malange,

June 1903, Gossweiler 946 (BM, holo.!; K, iso.!).

Acaulescent, with leaf bases enlarged below ground

to form bulb-like swelling; rosettes single, rarely suck-

ering to form small groups. Leaves rosulate, usually

deciduous, erectly spreading to slightly recurved, green,

obscurely lineate, sometimes with few scattered whit-

ish spots, surface smooth, ensiform, 450 x 50-60 mm;

margin whitish, cartilaginous, with firm unevenly spaced

teeth, 3 mm long, 5 — 40 mm apart. Inflorescence up to 2

m high, erect, 3-5-branched. Racemes cylindrical-coni-

cal to subcapitate, 70 mm long, dense. Floral bracts

15-25 x 4—6 mm. Pedicels 25-30 mm long. Flowers'.

FIGURE 8. — Distribution of Aloenuttii, ®; A. paedogona, 9; A. palmi-

formis, ■: and A. procera. ▲.

perianth yellow-green, ± 35 mm long, 9-11 mm across

ovary, constricted above ovary, forming globose basal

swelling, enlarging towards middle, narrowing towards

mouth; outer segments free only at tips. Stamens not

exserted. Style exserted up to 2 mm. Flowering time :

September to April.

Diagnostic characters : underground bulb. Leaves ±

450 x 50-60 mm, with firm teeth of ± 3 mm long. Unu-

sual shape of flowers with globose basal swelling. Floral

bracts linear-lanceolate, long-acuminate, 15-25 mm long.

Relationships with other species: Reynolds (1966) consid-

ered Aloe paedogona to be conspecific with A. buettneri

together with A. bulbicaulis. Carter (1994) stated that the

three taxa are morphologically clearly distinct and also

separated geographically and therefore does not agree with

Reynold’s opinion that the three taxa are conspecific.

Habitat', floodplains, woodland, savanna and grassland.

Distribution : Angola (Bie, Huambo, FJuila, Malange),

northern Namibia (Figure 8).

Illustrations', as Aloe buettneri in Rothmann: 46, 47 (2004).

Additional specimens examined

ANGOLA. — 0915: Malange, near Mated, 20 miles W of Malange,

(-BD), 10 April 1960, Reynolds 9384 (PRE). 0916: Malange, type

locality, (-CB), July 1959, Reynolds 9388 (PRE). 1117: Malange,

(-DA), 1903, Almeida s.n. (LISC); Malange, Capunda, Mulundo,

Reserva da Palanca Preta, picada do Luasso, (-BC), 5 August 1965,

Correia 3013 (LISC); Malange, Capunda, Mulundo, Reserva da Pal-

anca Preta, (-BC), 6 August 1965, Henriques 575 (LISC, LISU), 19

August 1965, Menezes 1984 (LISC). 1215: Nova Lisboa, 24 miles

[38.5 km] E of Balombo, 134 miles [215.5 km] E of Lobito on road

to Nova Lisboa, (-AC), 31 March 1960, Reynolds 9328 (PRE). 1216:

Bie Province, 4 miles [6.5 km] E of Chinguar, about midway between

Silva Porto and Nova Lisboa, (-CB), 31 January 1960, Reynolds 9234

(PRE). 1217: Bie, between Coemba and Cuanza River, (-CB), Exell

& Mendonfa 1752 (LISC). 1315: Caconda, Cunene River, Cadinco-

cololo, (-CC), 27 July 1905, Gossweiler 1792 (LISC). 1415: Gangue-

las, Dongo, ribeiro Cussaba, (-DA), 1 August 1905, Gossweiler 1807

(LISC). 1416: Huila, Ganguelas, Vila Artur de Paiva, (-AD), 25 Janu-

ary 1960. Mendes 2175 (LISC).

EA. palmiformis Baker in Transactions of the Lin-

nean Society of London 1: 263 (1878). Type: Angola,

Huila District, Morro de Lopolo, April 1860, Welwitsch

3726 (BM, holo.!; K!, LISU!, iso.).

Shrubby plants. Stem very slender, 1.0-1. 5 m long,

erect, sparsely branched mostly at base, with persistent

dried leaves. Leaves rosulate at branch apices, arcuate-

ascending-recurved or spreading-recurved, dull green

with reddish tinge, lower surface with many small,

crowded, pale green almost white spots towards base,

narrowly lanceolate-attenuate, up to 300 x 50 mm;

sheath lineate, 1 0 mm long; margin with prominent, pun-

gent, pale brown teeth, 4—5 mm long, 10 mm apart; leaf

exudate crusty when dry. Inflorescence 0.4— 0.5 m high,

erect, up to 4-branched, sometimes simple in young

plants. Raceme cylindrical, slightly acuminate, 100-150

mm long, lax. Floral bracts 2-3 x 2 mm. Pedicels 13-15

mm long. Flowers : perianth rose-scarlet, 30 mm long,

5.5 mm across ovary, slightly enlarging towards mouth,

cylindrical-trigonous; outer segments free for 10 mm.

Stamens exserted up to 1 mm. Style exserted up to 2 mm.

Flowering time : April to June.

Diagnostic characters', stems very slender, usually

sparsely branched. Leaves sometimes unspotted on both

32

Bothalia 39,1 (2009)

sides, usually copiously spotted on lower surface near

base. Marginal teeth prominent. Inflorescence usually 2-

or 3-branched, 0.4-0. 5 m high.

Relationships with other species : it is closely allied to

Aloe gossweileri (Reynolds 1966).

Habitat : among standstone rocks in woodland.

Distribution : endemic to Angola (Huambo, Huila, Namibe)

(Figure 8).

Illustrations'. Reynolds: 354, 355 (1966).

Additional specimens examined

ANGOLA. — 1215: Benguela, Capanga, Veva, (-CB), 17 June

1940, Gossweiler 12599 (LISC). 1413: Huila, Lubango, Tundavala,

ao km 16, (-CD), 27 April 1971, Borges 119 (LISC, PRE); Huila,

Lubango, Humpata, pr. do Perimetro Florestal, (-CD), 15 April 1960,

Mendes 3604 (LISC); Huila, Sa da Bandeira, 7 miles [11 km] NE of

Humpata, 8 miles [13 km] SW of Sa da Bandeira, (-CD), 29 June

1959, Reynolds 9292 (BM, PRE); Huila, Sa da Bandeira, Humpata,

proximo de Bimbe, (-CD), 3 May 1963, Santos 1110 (LISC, LISU);

Huila, Sa da Bandeira, Humpata, a 6 km de Sa da Bandeira, (-CD), 3

May 1963, Santos 1112 (LISC, LISU). 1513: Morro de Lopolo, near

Huila village, 16 km SE of Sa da Bandeira, (-AA), 27 May 1965, Rey-

nolds 9288 (LISC, PRE); Huila, Serra do Lopolo, pr. Cascata, (-BA),

20 May 1966, Correia 3824 (LISC).

EA. procera L.C. Leach in Journal of South African

Botany 40: 117 (1974). Type: Angola, Cuanza Sul Dis-

trict, ± 50 km N of Quibala, 12 July 1972, Leach & Can-

ned 14617 (USC, holo.!; BM, LISC!, SRGH, iso.).

Solitary rosettes, erect. Stem short, up to 0.25 m,

sometimes acaulescent. Leaves densely rosulate, erectly

spreading, very slightly recurved at apex, pale green,

obscurely lineate, without spots, ovate-attenuate, apical

portion drying out, up to 550 mm long including withered

apical portion, 80-95 mm wide; margin narrow, cartilagi-

nous, pale yellow, with orange-brown tipped teeth, 1.5-

3.5 mm long, 10-18 mm apart. Inflorescence 2.2-2.75 m

high, erect, 9-12-branched, lower branches rebranched.

Raceme cylindrical, 250-400 mm long, oblique, lax;

buds and flowers secund. Floral bracts 5-6 x ± 4 mm.

Pedicels 1.5-5 mm long. Flowers : perianth dull reddish

purple, 28-33 mm long, 5-6 mm across ovary, not or

only slightly narrowed above ovary, slightly widening

at mouth, fairly straight, cylindric-trigonous; outer seg-

ments free for 9-1 1 mm. Stamens slightly exserted. Style

exserted up to 6 mm. Flowering time : July to August.

Diagnostic characters', leaves pale green, 80-95 mm

wide, apical portion dry and withered. Inflorescence

remarkably tall, up to almost 3 m, branched above mid-

dle, with long laxly flowered racemes of small purple

flowers, subtended by small few-nerved bracts.

Relationships with other species : this species belongs

to the group containing Aloe chris tianii from south and

central tropical Africa, A. pretoriensis from northern

South Africa and Zimbabwe and A. luapulana L.C. Leach

from Zambia, but has its closest affinity with A. guerrae

(Leach 1974).

Habitat', tall grass in deciduous woodland on steep hillside.

Distribution : endemic to Angola (Cuanza Sul) (Figure 8).

Illustration: Leach: 119 (1974).

Additional specimen examined

ANGOLA.— 1014: Cuanza Sul, 50 km N of Quibala, (-BD), 6

August 1972, Leach & Connell 14617B (PRE).

EA. rupicola Reynolds in Journal of South African

Botany 26: 89 (1960). Type: Angola, Bie District, Chim-

bango, rocky hills 3 miles [5 km] S of Chinguar, 19 June

1959, Reynolds 9243 (PRE, holo.!; K!, LUAI, iso.).

Arborescent plant, 2-3 m (sometimes up to 5 m) high.

Stem erect, simple or branched at base, 100-120 mm

diam., without persistent dried leaves. Leaves densely

rosulate, suberectly spreading to spreading, green,

obscurely lineate on upper surface, more prominently

lineate on lower surface towards base of leaf, lanceolate,

apical portion usually drying out, 300-350 mm long

plus 100 mm dried apical portion, 60 mm wide; margin

with pungent, reddish brown teeth, 4—5 mm long, 10

mm apart; leaf exudate drying pale yellow. Inflorescence

0.7-0. 9 m high, erect, 3-5-branched. Racemes cylindri-

cal, 150-180 mm long, dense. Floral bracts ±9x5 mm.

Pedicels ±12 mm long. Flowers: perianth orange-scar-

let, 42 mm long, 7 mm across ovary, slightly enlarged

above ovary towards mouth, cylindrical-trigonous,

slightly clavate; outer segments free for 21 mm. Stamens

exserted 2-3 mm. Style exserted up to 3 mm. Flowering

time: June to July.

Diagnostic characters: stems averaging 2-3 m,

mostly simple or branched from the base. Old dry leaf

remains not persistent. Leaves obscurely lineate on

upper surface, more prominently lineate on lower sur-

face, especially in lower half. Perianth slightly clavate.

Relationships with other species: its closest ally appears

to be Aloe volkensii Engl, from eastern Africa (Reynolds

1966).

Habitat: rocky hills.

Distribution: endemic to Angola (Bie) (Figure 9).

Illustrations: Reynolds: 323, 324 (1966).

Notes: this species is only known from the type locality.

EA. scorpioides L. C. Leach in Journal of South Afri-

can Botany 40: 106 (1974). Type: Angola, Mofamedes

District, near Humbia, towards base of W escarpment

of Serra de Chela, April 1973, Leach & Canned 14654

(LISC, holo.!; BM!, BR, K!, LUA, LUAI, M!, MO!,

PRE, SRGH, ZSS, iso.).

Bothalia 39,1 (2009)

33

Low scrambling shrub, up to 0.5 m high, rarely up to

1 m. Stem branching at base and above, slightly diver-

gent, usually widely spreading, only rarely erect, with

persistent dried leaves. Leaves rosulate at branch apices,

more cauline dispersed below, spreading or somewhat

recurved, yellowish green, upper surface without spots,

lower surface darker, obscurely lineate, rarely with few

spots near base, very narrowly ovate-acuminate, up to

300 x 25-35 mm; sheath striate, 10-20 mm long; mar-

gin with pungent, yellowish or brownish tipped teeth,

somewhat forward-hooked, 2-3 mm long, 10-15 mm

apart; leaf exudate crusty when dry. Inflorescence ±0.15

m long, descending at base and curving upwards, sim-

ple or 1- or 2-branched. Raceme narrowly conical or

cylindrical-acuminate, 110-250 mm long, laterals usu-

ally shorter, dense. Floral bracts ± 6.5 x up to 3.5 mm.

Pedicels 6-10 mm long. Flowers : perianth scarlet, yel-

low-striped with green at base, 21-28 mm long, ± 7 mm

across ovary, narrowed above ovary, enlarging towards

wide-open mouth, cylindric-trigonous; outer segments

free for 8.5-10.0 mm. Stamens exserted. Style exserted

up to 2.5 mm. Flowering time : April to May.

Diagnostic characters : plants scrambling and shrubby

in habit. Leaves yellow-green, almost invariably immacu-

late, with small marginal teeth. Inflorescence with very

slender peduncle, which is at first usually descending,

then arcuately-ascending, with the rather dense, narrow,

acuminate racemes held erect. Buds suberect. Flowers

curved. Bracts large, prominently nerved, orange-brown.

Relationships with other species : its closest relative

seems to be Aloe palmiformis Baker (Leach 1974).

Habitat, rocky slopes, often in shade of woodland.

Distribution: endemic to Angola (Huila, Namibe) (Fig-

ure 9).

Illustrations: Leach: 107, 108 (1974).

Additional specimen examined

ANGOLA. — 1413: Mofamedes, western slopes of the Serra da

Chela, ± 14 miles [22.5 km] W of Sa da Bandeira, (-CB), 21 April

1972, Reynolds 9275 (LISC).

EA. vallaris L.C. Leach in Journal of South African

Botany 40: 111 (1974). Type: Angola, Huila District,

Serra da Chela escarpment, cliffs W of Tchivinguiro, 23

April 1973, Leach & Canned 14651 (LISC, holo.!; BM!,

BR!, K!, LISC!, LUAI, M!, MO!, PRE!, SRGH, WIND,

ZSS, iso.).

Shrubby plant, 0.3-0. 5 m high. Stem branched from

base; rosettes tilted to one side, greyish or greenish blue

to bluish green. Leaves rosulate at branch apices, more

widely spaced below, few small, oval or round, whitish

spots near base, more spots on lower surface, narrowly

ovate-attenuate, 220-340 x 40-50 mm; sheath copiously

spotted; margin narrow, yellowish, with pungent, yel-

lowish, orange- or brown-tipped teeth, 2. 0-2. 5 mm long,

10-12 mm apart; leaf exudate frothy, drying to opaque,

crystalline, yellow crust. Inflorescence 0.5-0. 6 m,

oblique or suberect, simple or divaricately 1 -branched.

Racemes narrowly elongate cylindrical-acuminate, ter-

minal raceme 250 — 450 mm long, lateral raceme 1 70—

300 mm long, oblique, lax. Floral bracts up to 4.5 x 2.5

mm. Pedicels 4. 0-4. 5 mm long. Flowers: perianth bright

scarlet, somewhat purplish at apex, mouth becoming

yellowish at maturity, 20-25 mm long, ± 5 mm across

ovary, narrowed slightly above ovary, enlarging slightly

towards open mouth, slightly curved, cylindric; outer

segments free for 4. 5-6.0 mm. Stamens and style not or

scarcely exserted up to 1 mm. Flowering time: April.

Diagnostic characters: very shrubby plant of rela-

tively dwarf stature, with stem branched from base.

Rosettes of grey-blue to blue-green leaves tilted to one

side. Inflorescence simple or 1 -branched from low down,

with narrow acuminate racemes. Short, bright scarlet

flowers with outer segments free for only 4. 5-6.0 mm

carried on relatively short pedicels.

Relationships with other species: it appears to be most

closely allied to Aloe gossweileri and A. catengiana

(Leach 1974).

Habitat: cliffs.

Distribution: endemic to Angola (Namibe) (Figure 5).

Illustrations: Leach: 113 (1974).

Notes: this species is only known from the type locality.

EA. venenosa Engl, (insufficiently known sp.) in

Botanische Jahrbucher 15: 471 (1893). Type: Angola,

Lunda, between Quimbundu and Nyangwe, 20 May

1882, P. Pogge 1460 (B, holo.-LISC, photo.!).

Growth form unknown. Leaves 350 x 70 mm; mar-

gins with teeth, 6-7 mm long. Inflorescence copiously

paniculate. Raceme 250-350 mm long. Pedicels 25 mm

long. Flowers: perianth pale red, 27-30 mm long. Flow-

ering time: May.

Relationships with other species: unknown.

Distribution: endemic to Angola (Lunda Norte or Lunda

Sul).

Notes: this species has not been located since the type

specimen was collected by Pogge in May 1882. The true

identity of this species, therefore, remains a mystery.

Furthermore, the type locality is extremely vague: Quim-

bundu is in northeastern Angola, whereas Nyangwe is

in the eastern Democratic Republic of the Congo. It is

therefore possible that this species was not collected in

Angola, but in the Democratic Republic of the Congo.

A. zebrina Baker in Transactions of the Linnean

Society of London 1: 264 (1878). Type: Angola, Loanda

District, Barra do Bengo, Quicuxe towards Cacuaco, July

1854, Welwitsch 3721 designated by Reynolds (1966)

(LISU, lecto.!; BM!, G!, K!, isolecto.-LISC, photo.!).

A. platyphylla Baker: 264 (1878). Type: Angola, Pungo Andongo,

1879, Welwitsch 3722 (K, lecto.!; BM, G!, LISU!, isolecto.).

A. constricta Baker: 168 (1880). Type: Mozambique, near Sena, 8

April 1860, Kirk 34 (K, holo.!).

A. lugardiana Baker: 135 (1901). Type: Botswana, Botletle River,

30 June 1897, Lugard 2 (K, holo.!).

A. baumii Engl. & Gilg: 191 (1903). Type: Angola, Chirumbu, 14

October 1899, Baum 275 (B, holo.; E, iso.!).

A. bamangwatensis Schonland: 122 (1904). Type: Botswana, Pala-

pye Road, March 1904, Schonland 1656 (GRA, holo.; PRE, iso.!).

Acaulescent; rosettes sometimes solitary or usually

suckering to form groups. Leaves densely rosulate, spread-

ing, dull green, upper surface with transverse bands of

conspicuous, whitish, oblong spots, lower surface usually

obscurely or copiously spotted, lanceolate, usually dried

34

Bothalia 39,1 (2009)

and twisted at apex, 150-350 mm long, 60-70 mm wide

at base; margin with stout, pungent, red-brown teeth,

4—7 mm long, 10-15 mm apart; leaf exudate yellowish,

drying purplish or orange. Inflorescence 0.75-2 m high,

erect, 4— 12-branched, lower branches often rebranching.

Racemes narrowly cylindrical-acuminate, 300-400 mm

long, very lax. Floral bracts 6-15 x 2-3 mm. Pedicels

6-15 mm long. Flowers : perianth dull red to pinkish

red or coral-coloured with paler segment margins, 25-

35 mm long, basally inflated to 7-9 mm across ovary,

abruptly constricted just above ovary, widening towards

mouth, slightly decurved, cylindrical-trigonous; outer

segments free for 7-11 mm. Stamens exserted 1-2 mm.

Style exserted up to 2 mm. Flowering time : variable,

(November) February to April (May), or June to July,

depending on the locality.

Diagnostic characters', spotted aloe. Upper leaf sur-

face always conspicuously spotted, spotting on lower sur-

face varies. Very laxly flowered racemes, 300-400 mm

long; pedicels 6-15 mm long; perianth averaging 30 mm.

Relationships with other species'. Aloe zebrina belongs

to the Section Pictae Salm-Dyck or maculate aloes (Glen

& Hardy 2000).

Habitat', variety of veld types and soil, mostly grassland

and thickets on dry hills.

Distribution : Angola (Bengo, Cuando-Cubango, Cuanza

Sul, Cunene, Huambo, Huila, Luanda, Lunda Sul,

Malange, Moxico), Botswana, western Mozambique,

Malawi, northern Namibia, South Africa (North-West),

Zambia, Zimbabwe (Figure 10).

Illustrations'. Reynolds: 90 (1966); Rothmann: 94, 95

(2004).

Additional specimens examined

ANGOLA. — 0813: Luanda, entre a foz do Dande e a estrada de

Caxito, (-AD), 18 July 1964, Barbosa 10831, 10832 (LISC); Luanda,

Quicuxe, (-BD), Welwitsch 3720, 3724 & 3725 (K, LISU); Luanda,

Caxito, praia de S. Tiago a caminho do Dande, (-CB), 8 May 1958,

Monteiro, Santos & Murta 119 (LISC, LISU); Luanda, near Cac-

uaco, 10 miles [16 km] NE of Luanda, (-CD), 12 July 1959, Reynolds

9406 (PRE); Luanda, Icola e Bengo, near Dande River, (-DA), June

1944, Gossweiler 13299 (LISC). 0817: Malange, Cambo, Montalegre,

(-CC), 27 May 1948, Rocha 75 (LISC). 0915: Cuanza Norte, Catete,

(-BB), 1930, Gossweiler 9210 (LISC); Pungo Andongo, (-DA), 1879,

FIGURE 1 0. — Distribution of Aloe zebrina.

Welwitsch 3722 (G, K). 0920: Lunda, near Vila Henrique de Carvalho,

(-CA), 22 April 1937, Exell & Mendonga 921 (LISC); Lunda, Dala,

(-CA), 25 April 1937, Exell & Mendonga 1134 (LISC). 1013: Cuanza

Sul, Porto Amboim, estrada S. Filipe-Porto Amboim, a 10 km de S.

Filipe, (-DB), 27 April 1967, Teixeira 11553 (LISC). 1215: Huambo,

Nova Lisboa, Chianga, (-DB), 1 June 1971, Fernando 4 (LISC). 1320:

Moxico, (-BA), Exell & Mendonga 1662 (LISC). 1413: Huila, Sa da

Bandeira, Hoque, (-DB), 2 June 1966, Henriques 1022 (LISC, LISU);

Huila, Lubango, Sa da Bandeira, arredores, proximo da escola do

Marques, (-DC), 7 March 1972, Couto 192 (LISC); Huila, Sa da Ban-

deira, Vio, (-DC), 4 May 1965, Henriques 378 (LISC, LISU). 1417:

Menongue, entre Cuchi e Vila Serpa Pinto, vale do Luassenha, (-CA),

4 April 1960, Mendes 3460 (LISC). 1418: Cuito-Cuanavale, andados

40 km de Longa para Cuango, (-BD), 18 March 1960, Mendes 3175

(LISC). 1513: Huila, Humpata, Tchivinguiro, (-AB), 24 April 1972,

Menezes 4087 (LISC); Huila, Lubango, ao km 17 da estrada para a

Chibia, (-BA), 24 April 1965, Menezes 1590 (LISC); Huila, proximo

da Missao Catolica, (-BA), 10 February 1956, Santos 182 (LISC).

1515: Ganguelas, entre os Rios Cubango e Cunene, Rio Oxo, (-BA),

4 August 1905, Gossweiler 1834a (LISC). 1516: Menongue, Cuito,

vale de Sobi, (-BD), 15 March 1906, Gossweiler 3704 (LISC). 1519:

Cuando-Cubango, Cuito-Cuanavale, sede, (-AA), July 1967, Pereira

s.n. (LISC). 1614: Royadas, Centro de Estudos do Cunene, (-DD), 27

March 1970, Silva 3151 (LISC). 1714: top of Ruacana Falls, on Ango-

lan side, (-AC), Rycroft 2435 (NBG). No grid: Moxico, S of Luhanda

River, 15 January 1938, Milne-Redhead 4146 (K-PRE, photo.).

ACKNOWLEDGEMENTS

The authors would like to thank: Ms Hester Steyn,

Data Management Unit, SANBI, for the distribution

maps; Mr Steve Cafferty, for providing digital images

of some BM specimens; two referees for suggesting

improvements to the manuscript.

REFERENCES

AIRY SHAW, H.K. 1947. The vegetation of Angola. The Journal of

Ecology 35: 23 — 48.

BAKER, J.G. 1878. Report on the Liliaeeae, Iridaceae, Hypoxidaceae

and Haemodoraceae of Welwitsch’s Angolan herbarium. Trans-

actions of the Linnean Society of London 1 : 245-273.

BAKER, J.G. 1 880. A synopsis of Aloineae and Yuccoideae. Journal of

the Linnean Society, Botany 18: 148-241.

BAKER, J.G. 1897. Aloe nuttii. In Hooker's leones Plantarum 6,1: t. 2513.

BAKER, J.G. 1898. Aloe. In W.T. Thiselton-Dyer, Flora of tropical

Africa 7: 454-469. Reeve, London.

BAKER, J.G. 1901. Diagnoses Africanae 13. Kew Bulletin 1901: 1 19—

138.

BERGER, A. 1906. A new aloe from Angola. Journal of Botany, British

and Foreign 44: 57, 58.

CARTER, S. 1994. Aloaceae. In R.M. Polhill, Flora of tropical East

Africa. Balkema, Rotterdam.

CARTER, S. 2001. Aloaceae. In G.V. Pope, Flora zambesiaca 12,3:

48-98.

CHRISTIAN, H.B. 1936. Aloe bulbicaulis. The Flowering Plants of

South Africa 16: t. 630.

CHRISTIAN, H.B. 1 940a. Some new Aloe species from central and east

tropical Africa. Journal of South African Botany 6: 1 77-190.

CHRISTIAN, H.B. 1940b. Aloe mketiensis. The Flowering Plants of

South Africa 20: t. 785.

CHRISTIAN, H.B. 1940c. Aloe nuttii. The Flowering Plants of South

Africa 20: t. 762.

COSTA,' E„ MARTINS, T. & MONTEIRO, F. 2004. A checklist of Ango-

lan grasses — Checklist das Poaceae de Angola. Southern African

Botanical Diversity Network Report No. 28. SABONET, Pretoria.

DE WILDEMAN, E.A.J. 1921. Contribution a l 'etude de la flore du

Katanga. Reynaert, Bruxelles.

DINTER, K. 1914. Neue und wenig bekannte Pflanzen Deutsch-Sud-

west-Afrikas. The author, Okahandja.

DOWNS, PE. 1970. The road to Angola. The National Cactus and Suc-

culent Journal 25: 37-40.

ENGLER, H.G.A. 1888. Plantae Marlothiariae; ein Beitrag zur Kennt-

niss der Flora Sud-afrikas. I. Teil: Monokotyledoneae und Diko-

tyledoneae archichlamydeae. Botanische Jahrbucher 10: 1-50.

Bothalia 39,1 (2009)

35

ENGLER, H.G.A. 1893. Liliaceaea africanae. Botanische Jahrbucher

15: 467—479.

ENGLER, H.G.A. & GILG, E.R 1903. Liliaceae. In O. Warburg, Kunene-

Zambesi Expedition. Verlag des Kolonial-Wirtshaftlichen Komitees,

Berlin.

FIGUEIREDO, E. & SMITH, G.F. 2008. Plants of Angola/Plantas de

Angola. Strelitzia 22. South African National Biodiversity Insti-

tute, Pretoria.

FIGUEIREDO, E., SMITH, G.F. & CESAR, J. in press. The flora of

Angola: first record of diversity and endemism. Taxon.

GLEN, H.F. & HARDY, D.S. 2000. Aloaceae (First part): Aloe. In G.

Germishuizen, Flora of southern Africa, Vol. 5, Part 1 , Fascicle

1: 1-167. National Botanical Institute, Pretoria.

HARDY, D.S. 1992. Kaokoland — Africa’s last wilderness. Aloe 29:

46-49.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index Herbariorum, Part 1: The Herbaria of the World, edn 8.

New York Botanical Gardens, New York.

KEAY, R.W.J. 1968. Liliaceae: Aloe. In F.N. Hepper, Flora of West

tropical Africa 3,1: 90-92. Whitefriars Press, London.

LANE, S.S. 2004. The aloes of Malawi. Umdaus Press, Hatfield, Pretoria.

LEACH. L.C. 1971. Two new species of Aloe (Liliaceae) from south

tropical Africa. Journal of South African Botany 37: 249-266.

LEACH, L.C. 1974. Notes on the aloes of South tropical Africa with

four new species and a new variety. Journal of South African

Botany 40: 101-122.

MCNEILL, J., BARRIE, F.R.. BURDET, H.M., DEMOULIN, V.,

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

PRADO, J., SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. (eds). 2006. International Code of Botanical

Nomenclature (Vienna Code) adopted by the Seventeenth Inter-

national Botanical Congress Vienna, Austria, July 2005. Regnum

Vegetabile 146: 1-568. Gantner Verlag, Liechtenstein.

NEWTON, L.E. 2001. Aloe. In U. Eggli, Illustrated handbook of suc-

culent plants: Monocotyledons: 103-186. Springer, Berlin.

REYNOLDS, G.W. 1936a. Notes on a new aloe from Rhodesia and a

new aloe from the Transvaal. Journal of South African Botany

2: 171-175.

REYNOLDS, G.W. 1936b . Aloe dinteri. The Flowering Plants of South

Africa 16: t. 637.

REYNOLDS, G.W. 1950. The aloes of South Africa. Aloes of South

Africa Book Fund, Johannesburg.

REYNOLDS, G.W. 1960. Notes on the aloes of Angola, with descrip-

tions of three new species. Journal of South African Botany 26:

81-92.

REYNOLDS, G.W. 1961. Anew aloe from Angola. Kirkia 1: 160, 161.

REYNOLDS, G.W. 1962. A new aloe from Angola. Journal of South

African Botany 28: 205-207.

REYNOLDS, G.W. 1964. A new aloe from Angola. Journal of South

African Botany 30:31,32.

REYNOLDS, G.W. 1 966. The aloes of tropical Africa and Madagascar.

Aloes Book Fund, Mbabane.

ROPERT, D. 2000-( continuously updated). Digital specimen images

at the Herbarium Berolinense. Published on the Internet http://

ww2.bgbm.org/herbarium/default.cfrn [accessed August 2008].

ROTHMANN, S. 2004. Aloes, aristrocats of the Namibian flora. ST

Promotions, Swakopmund.

SCHINZ, H. 1896. Die Pflanzenwelt Deutsch-Sudwest-Afrikas. Bulle-

tin de l 'Herbier Boissier 4, App. 3: 1-57.

SCHONLAND, S. 1904. On some new and some little known species

of South African plants-II. Records of the Albany Museum 1:

114-124.

SMITH, G.F., WALTERS, M., KLOPPER, R.R. & CROUCH, N.R.

2008a. Aloes of the World Workshop. Reaching consensus on

content for a scholarly electronic resource on Aloe, 19-22

November 2007. Final Report (30 April 2008). Unpublished

workshop report.

SMITH, G.F., WALTERS, M„ KLOPPER, R.R. & CROUCH, N.R.

2008b. Aloes of the World: African Plants Initiative. An interna-

tional web-based collaboration to promote scholarly research on

Aloe L. Bradleya 26: 121-128.

VERDOORN, I.C. 1964 . Aloe mendesii. The Flowering Plants of Africa

45: t. 1764.

WHITE, F. 1983. The vegetation of Africa, a descriptive memoir to accom-

pany the UNESCO/AETFAT/UNSO vegetation map of Africa.

Natural Resources Research 20. UNESCO, Paris.

WORLD WILDLIFE FUND. 2001. Terrestrial ecoregions of the world.

Available at http://www.nationalgeographic.com/wildworld/ter-

restrial.html [accessed July 2008],

.

■

rl

Bothalia 39,1: 37^5 (2009)

Three new species of Gladiolus (Iridaceae) from South Africa, a major

range extension for G. rubellus and taxonomic notes for the genus in

southern and tropical Africa

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

Keywords: Gladiolus L., Iridaceae, Namibia, new species. South Africa, Tanzania, taxonomy

ABSTRACT

Three new species of Gladiolus L. are described from South Africa. G. dolichosiphon is the second known member of

series Blandus from the mountains of the Little Karoo in Western Cape, and is distinguished from other members of the long-

tubed, pink-flowered G. carneus complex by its 5 or 6 linear leaves, creamy pink to salmon flowers with a tube 30-50 mm

long and longer than the dorsal tepal, and its late summer flowering. G. karooicus from the Klein Roggeveld and the northern

foothills of the Witteberg, is a spring-flowering species allied to G. permeabilis but has bright, canary-yellow flowers with

the lower part of the lower tepals involute and conspicuously auriculate. G. reginae is an edaphic endemic of the Sekhuk-

huneland Centre of Floristic Endemism in Mpumalanga, and flowers in autumn. It is evidently a glabrous member of section

Densiflorus series Scabridus, distinguished by its long-tubed flowers, streaked with red on the lower tepals and blotched with

red in the throat. Anomalously, however, it has the tubular inner bracts and large capsules diagnostic of section Ophiolyza

series Oppositiflorus. A re-examination of the morphology suggests that series Scabridus is better placed in section Ophiolyza

and a slightly revised classification of Gladiolus in southern Africa is proposed. We also propose the replacement name G.

sulculatus for the Tanzanian species, G. sulcatus Goldblatt, a later homonym of G. sulcatus Lam. Finally, a recent sighting

of what appears to be G. rubellus from northern Namibia constitutes the first record of this species in the country and a major

range extension from its previous known occurrence in southeastern Botswana.

INTRODUCTION

Gladiolus L., now including some 262 species, is the

largest genus of Iridaceae subfamily Crocoideae. It is also

the largest genus of Iridaceae in Africa, where 168 species

are now known from southern Africa, 82 species from tropi-

cal Africa, eight in Madagascar, and a scant 10 in Eurasia.

The genus is taxonomically well researched, and recent

monographs exist for Madagascar (Goldblatt 1989), tropi-

cal Africa-Arabia (Goldblatt 1996), and southern Africa

(Goldblatt & Manning 1998). New species continue to be

discovered, however, and two have since been added to the

genus from southern Africa (Manning et al. 1 999), another

from the Democratic Republic of Congo (Geerinck 2001)

and two more have been recognized from Zimbabwe and

Mozambique (Goldblatt 2008). Here we describe a further

three species from South Africa. Gladiolus reginae Gold-

blatt & J.C. Manning from northwestern Mpumalanga was

discovered during a botanical survey of the Steelpoort

River Valley, which transects the Sekhukhuneland Cen-

tre of Floristic Endemism (Van Wyk & Smith 2001), a

region that is still poorly explored botanically. This inter-

esting species is evidently a member of series Scabridus

(currently placed somewhat uncomfortably at the end of

section Densiflorus in the classification of the southern

African species) but it displays the critical characteristics

of section Ophiolyza. As a result, we have reassessed the

taxonomic position of series Scabridus within the genus.

A second species, G. karooicus Goldblatt & J.C. Manning

of section Hebea, from the Western Karoo, was discovered

in the spring of 2006, a year of unusually ample rainfall in

this semi-arid winter rainfall zone. A third new species, G.

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

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

den, P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2008-06-02.

dolichosiphon Goldblatt & J.C. Manning, from the moun-

tains of the Little Karoo in Western Cape, was collected in

early 2008, although a previously unplaced collection was

made two decades earlier.

In addition to describing these new species, we take

this opportunity to provide a new name for the Tanzanian

Gladiolus sulcatus Goldblatt. After examining the check-

list of Iridaceae being prepared for the World checklist of

selected plant families (R. Govaerts pers. comm. 2007),

we have learned that the name is a later homonym, and

therefore illegitimate. Finally, a recent collecting trip to

northwestern Namibia produced what is evidently a new

record for that country of G. rubellus Goldblatt, previ-

ously thought to be endemic to southeastern Botswana.

Gladiolus dolichosiphon Goldblatt & J.C. Manning,

sp. nov. (section Blandus)

Plantae 200^100 mm altae, cormo subgloboso 10-15

mm diam., tunicis extemis aetate in fibras molles solutis,

cataphyllis supra terrain purpurascentibus obscure viridi-

bus vel albis maculatis, foliis 6 vel 7, inferioribus 3

basalibus grandioribus laminibus linearibus ad anguste

ensiformibus 1 .5 — 4.0(— 6.5) mm latis marginibus hya-

linis foliis caulinibus sine laminis, caule simplici vel

1-ramoso, spica ad basem inflexa inclinata 4- vel 5-(ad

7-)flora, ramis 1- vel 2-floris, bractea externa subacuta

vel infime attenuata 20— 30(— 35) mm, interna subacuta ad

apicem indivisa, floribus pallide cremeis vel salmoneis

inodoris, tepalis inferioribus maculis pallidis medianis

circumscriptio atrorubro praeditis, tubo perianthii cylin-

drico (30-)40-55 mm longo, tepalis inaequalibus lanceo-

latis dorsalibus 30^10 x 15-17 mm prorsum supra stam-

ina arcuatis, lateralibus superioribus 25-28 x 12-14 mm,

inferioribus tribus prorsum extensis basi ± 2 mm conna-

tis, 20-25 x 4—5 mm, filamentis 16-20 mm longis, anth-

eris 8-9 mm longis, stylo arcuato ramis 8-9 mm longis.

38

Bothalia 39,1 (2009)

FIGURE 1. — Gladiolus dolichosi-

phon, Schutte & Vlok 941.

Photographer: Jan Vlok.

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

south of track between Rooiberg Pass and Bailey’s Peak,

(-DB), 7-23 February 2008, A.L. Schutte & J. Vlok 941

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

Plants 200—400 mm. Corm subglobose, 10-15 mm

diam.; outer tunics decaying into soft fibres, pale grey-

ish brown. Stem inclined outward above basal leaves,

unbranched or with a small lateral branch from axil of

one or both uppermost leaves, 1-2 mm diam. below

main spike. Catapylls flushed purple above ground and

obscurely mottled with green or white. Leaves 6 or 7,

lower 3 basal and largest, reaching or shortly exceeding

spike; blades linear to narrowly sword-shaped, 1.5— 4. 0(-

6.5) mm wide, midrib slightly thickened and margins

hyaline; upper 2 cauline leaves without blades, some-

times subtended by a lateral branch each, margins open

to base. Spike flexed at base, inclined, weakly flexuose,

4- or 5(-7)-flowered, subsecund, lateral branches 1- or 2-

flowered; bracts foliose, outer subacute or lower attenu-

ate, 20-30(-35) mm, green flushed purple distally, inner

slightly shorter or up to two thirds as long, subacute

and not forked apically. Flowers pale creamy or salmon

pink, lower three tepals each with a pale lozenge-shaped

median mark outlined in dark red, unscented; perianth

tube cylindric, expanded in upper 10 mm, straight or

arched distally, (30-)40-55 mm long; tepals unequal,

lanceolate, dorsal largest, 30—40 x 15-17 mm, arching

forward over stamens and curved upwards distally, upper

laterals 25-28 x 12-14 mm, extending forward and curv-

ing outward in distal third to half, lower three tepals

basally fused for ± 2 mm, narrowly lanceolate, 20-25 x

4—5 mm, in profile shorter than upper tepals. Filaments

16-20 mm long, exserted 10-14 mm from tube; anthers

8-9 mm long, purple with cream-coloured pollen.

Ovary ovoid, ± 5 mm long; style arching beneath dorsal

tepal, dividing just before or just beyond anther apices,

branches 5-6 mm long. Capsule and seeds unknown.

Flowering time : late January to mid-February. Figure 1.

Distribution and ecology, known from two collections

from the mountains around Ladismith in the western Lit-

tle Karoo (Figure 2), where it has been recorded from

the top of the Klein Swartberg at an altitude of 1 900 m

and from the Rooiberg at over 1 300 m. Plants occur on

cooler south-facing slopes in seasonally wet situations,

on rocky outcrops or along kloofs. On both occasions

Gladiolus dolichosiphon was collected on trips that had

been undertaken in response to fires that had occurred the

preceding summer and while it, like many Gladiolus spe-

cies, evidently flowers well after fire, it is probably not a

true pyrophile.

The long-tubed, pink flowers with red markings on

the lower tepals, are similar to several other Gladiolus

species, as well as other Iridaceae, that are adapted to

FIGURE 2. — Known distribution of Gladiolus dolichosiphon , O; G.

karoo icus, •.

Bothalia 39,1 (2009)

39

pollination by long-proboscid flies in the families Taba-

nidae and/or Nemestrinidae (Goldblatt & Manning 1999)

and there is no doubt that G. dolichosiphon is likewise

adapted to this pollination system.

Gladiolus dolichosiphon was first collected in Febru-

ary 1986 on the Klein Swartberg by ecologist Jan Vlok

while checking on the condition of plants of Protea pru-

inosa after a fire the previous year. This puzzling col-

lection could not be identified, however, and was not

included by Goldblatt & Manning (1998) in their revi-

sion of the genus in southern Africa. In February 2008,

Jan Vlok, accompanied by AnneLise Vlok and Willem

Wagenaar from CapeNature, encountered a population

of Gladiolus in flower on the Rooiberg during a biodi-

versity survey of the mountain. Suspecting that it was of

interest, they collected ample material and photographed

the flowers. From this it was clear that the plants repre-

sented an undescribed species and further investigation

in the herbarium revealed the earlier collection from the

Klein Swartberg, which is undoubtedly the same species.

Diagnosis and relationships: the basal fan of nar-

rowly lanceolate leaves and spike of relatively large,

long-tubed, pink flowers with median lozenge-shaped

markings on the lower three tepals, place Gladiolus doli-

chosiphon among the species of section Blandus series

Blandus. Centred on the variable Gladiolus carneus

Delaroche, series Blandus currently includes 13 species

endemic to the mountains of the southwestern Cape,

making G. dolichosiphon the fourteenth member of the

alliance and one of just two that are known from the

inland mountains of the Little Karoo. The other Little

Karoo species, G. aquamontanus Goldblatt & Vlok, is

another discovery of Jan Vlok’s and this hydrophyte is

restricted to the Great Swartberg, where it grows in per-

ennial streams and on wet cliffs.

The relationships between the nine members of the

cream- or pink-flowered Gladiolus carneus complex are

poorly understood. Members of the alliance are distin-

guished primarily on the relative lengths of the perianth

tube and tepals, the proportions of the upper and lower

tepals, and on flowering time and geographical distribu-

tion (Goldblatt & Manning 1998) (Table 1).

In its narrow, linear leaves and creamy pink or pale

salmon flowers with the tube longer than the dorsal tepal,

and the lower tepals with median, lozenge-shaped mark-

ings, Gladiolus dolichosiphon has a marked similarity to

G. bilineatus, a local endemic to the southern foothills of

the Langeberg, where it occurs mainly in a narrow band

along the renosterveld-fynbos transition, and flowers in

autumn, in March and April. G. bilineatus has just three

leaves, usually only the lower one or two with a well-

developed blade, 6-8 mm wide, and flowers with a peri-

anth tube 50-70 mm long. G. dolichosiphon, from the

interior Little Karoo mountains on seasonally moist slopes

at high altitude in fynbos, flowers in late summer, in Feb-

ruary. Plants have 6 or 7 linear leaves with blades mostly

less than 5 mm wide, and a perianth tube 30-55 mm long.

These differences in foliage, flower morphology, ecology

and distribution are all significant in the context of species

delimitation in the G. carneus complex.

Etymology. Greek dolicho, long; tubus, tube, referring

to the long perianth tube.

Other material seen

WESTERN CAPE. — 3321 (Ladismith): top of Klein Swartberg

Mountains, next to Besemfontein track, (-AD), 3 February 1986, Vlok

1407 (NBG).

Gladiolus karooicus Goldblatt & J.C. Manning,

sp. nov. (section Hebea)

Plantae 150-500 mm altae, cormo conico 25 x 15

mm, foliis usitate 4 laminis linearibus ± 2 mm latis, folio

caulino vaginato, caule simplici vel 1-2-ramoso, spica

2- ad 5-flora, bracteis 20-25 mm longis, floribus bilabia-

tis flavis perodoratis, tubo perianthii subcylindrico ± 12

mm longo, tepali inaequalibus, dorsalibus ± 25 x 10 mm

arcuatis, tepalis lateralibus superioribus erectis, tepalis

lateralibus inferioribus 12 x 4 mm in dimidio inferiore

marginibus incurvis tubum calcariformem formantibus,

filamentis ± 15 mm longis 10 mm ex tubo exsertis, anth-

eris ± 8 mm longis, stylo arcuato ramis ± 2.5 mm longis.

TABLE 1. — Selected characteristics of members of Gladiolus carneus complex. Distribution data follows centres of endemism in Goldblatt & Man-

ning (2000)

40

Bothalia 39,1 (2009)

TYPE. — Northern Cape, 3320 (Sutherland): Klein

Roggeveld, Komsberg Pass, along seasonal stream in

damp gully, in shaly gravel, (-DA), 8 September 2006,

Goldblatt & Porter 12804 (NBG, holo.; K, MO, PRE,

iso.).

Plants 150-500 mm high. Corm conical, 25 x 15 mm;

outer corm tunics brown and papery, decaying below

into fine to medium-textured fibres. Stem suberect, or

inclined, flexed outward above sheath of third leaf, sim-

ple or with one or occasionally two branches. Cataphylls

pale and membranous, uppermost reaching shortly above

ground and then dull purple. Leaves usually 4, more if

plants branched, lower 3 with expanded blades, lower 2

longest and reaching at least to base of spike or shortly

exceeding it; blades linear, ± 2 mm wide, firm-textured,

slightly twisted, midrib moderately thickened, margins

evidently not thickened; cauline leaf or leaves short and

largely sheathing. Spike inclined ± flexuose, 2-5-flow-

ered; bracts grey-green, sometimes flushed purplish

above, often dry and pale near apices, outer 20-25 mm

long, inner two thirds to almost as long as outer, acute,

not forked at tip. Flowers yellow, dorsal and upper later-

als flushed and veined grey-purple outside, lower later-

als deep yellow in proximal half and turning dull purple

on fading, sweetly violet-scented; perianth tube subcy-

lindric, slightly wider near apex, ±12 mm long; tepals

unequal, all narrowed below into claws, lanceolate,

dorsal largest, ± 25 x 10 mm, arched and hooded over

stamens, tip curving upward, upper laterals erect, distal

halves arching over dorsal tepal, adaxial margins often

touching one another, ± 25 x 7 mm, windowed between

lower halves of dorsal and upper lateral tepals, lower 3

tepals joined to upper laterals for ± 2 mm and to each

other for ± 3 mm, lower laterals with erect claws ±1.5

mm long, limbs horizontal, ±12x4 mm, expanded dis-

tally, margins of proximal half involute to form tube

extending backward as auricles, lower median ±15x8

mm long, arching downward in distal third. Filaments

unilateral and contiguous, arched under dorsal tepal, ±

1 5 mm long, exserted 1 0 mm from tube; anthers ± 8 mm

long, parallel and contiguous, light purple; pollen whit-

ish. Ovary oblong, ± 5 mm long; style arching over sta-

mens, dividing opposite upper third of anthers, branches

±2.5 mm long, not or barely exceeding anthers. Cap-

sules obovoid and rounded apically or ellipsoid, 15-20

mm long. Seeds ovate, 5-7 x 4-5 mm, broadly winged,

translucent golden brown, with dark seed body ± 2 mm

diam. Flowering time : mid-August to mid-September.

Figure 3.

Distribution and ecology. Gladiolus karooicus is

restricted to the Klein Roggeveld and nearby foothills of

the Witteberg (Figure 2). The type collection was made

in the Komsberg, the scarp separating the high Rog-

geveld from the Klein Roggeveld but the species has also

been recorded from the northern foothills of the Witte-

berg at Memorial, west of Matjiesfontein, and from near

Prince Albert and the foot of the Great Swartberg. It is

mostly encountered in damp gullies and along seasonal

streams in gravelly, decomposed shale and tillite, among

grass clumps and low shrubs. G. karooicus occurs sym-

patrically with the common and widespread G. venustus,

also section Hebea, which favours drier, more exposed

sites. The bright yellow flowers, moderately long peri-

FIGURE 3. — Gladiolus karooicus, Goldblatt & Porter 12804 (NBG).

A, flowering plant; B, flower front view; C, half flower; D, cap-

sules and seed. Scale bar: 1 0 mm. Artist: John Manning.

anth tube, and sweet floral scent suggest that the species

is pollinated by long-tongued bees.

In a remarkable coincidence, Gladiolus karooicus was

independently discovered by three different parties in the

spring of 2006. This was a year of unusually good rains

for the western Karoo and it is likely that this attracted

botanical collectors to the region as well as encouraging

good flowering in the species.

Diagnosis and relationships : Gladiolus karooicus has

the unusually narrow, linear leaves without thickened

margins, clawed tepals, and distinctive dark seed body

characteristic of section Hebea (Goldblatt & Manning

1998) and there is no doubt that it should be placed here.

Bothalia 39,1 (2009)

41

Within section Hebea, the soft-textured corm tunics and

ellipsoid capsules closely resemble those of members

of series Permeabilis and on this basis we conclude that

G. karooicus is probably immediately allied to the small

group of species that includes G. permeabilis, G. stella-

tus and G. wilsonii, comprising the members of the alli-

ance that occur in the winter rainfall zone of southern

Africa. G. karooicus is distinctive in series Permeabilis

on account of its bright yellow flowers with geniculate

lower tepals that are involute and tubular in the basal

half of the blades, with conspicuous auricles above the

claw. In G. permeabilis and G. wilsonii the lower tepals

are ± recurved and although slightly canaliculate, they

are evenly narrowed towards the claw and not at all

auriculate. In addition, the flower colour is generally dull

brownish or lilac, or white. G. stellatus has very distinc-

tive, stellate, actinomorphic flowers.

The curiously constructed flowers of G. karooicus

are remarkably similar to those of G. venustus (series

Deserticola, also section Hebea) in their sharply flexed,

auriculate lower tepals but the members of this species

group share derived woody, clawed corm tunics, a scal-

loped flower spike, and distinctive, squat, barrel-shaped,

apically retuse capsules. In addition, the seed body in

series Deserticola is brown and not black. The floral

similarities between G. karooicus and G. venustus are

thus evidently the result of convergence, possibly the

result of pollinator-driven selection. When not in fruit,

the two species are separated by the differences in their

corm tunics, which are soft in G. karooicus and woody

in G. venustus, and by differences in the orientation of

the upper lateral tepals. In G. venustus the upper later-

als are suberect from the base and not significantly fused

to the lower tepals, whereas in G. karooicus the upper

laterals are fused to the lower tepals for ± 2 mm and thus

horizontal in this basal portion, but abruptly suberect in

the distal, free portion, resulting in the presence of the

characteristic window between the upper lateral tepals

and the lower tepals.

Etymology, karooicus, from the karoo, the semi-arid

and arid interior part of South Africa.

Other material seen

WESTERN CAPE. — 3320 (Montagu): Matjiesfontein, Memorial

Siding, among monuments, (-AB), 1 September 2006, J.C. McMaster

s.n. (NBG, photo.), 1 November 2006 (fruit), Manning 3171 (NBG);

Komsberg Pass, approximately halfway up pass, (-DB), 22 September

2007, V.R. Clark & C. Kelly 102 (GRA, NBG). 3322 (Oudtshoom):

Prince Albert, hill ENE of village on road to Platberg, (-AA), 31

August 2006, J.P. Roux s.n. (NBG, photo.).

Gladiolus reginae Goldblatt & J.C. Manning, sp.

nov. (section Ophiolyza)

Plantae 0.4— 1.5 m altae, cormo subgloboso ± 20 mm

diam., tunicis papyraceis vel aetate aliquantum fibrosis,

foliis 8 vel 9 inferioribus 6 vel 7 basalibus grandioribus

laminis anguste ensiformibus (3— )5— 1 0 mm latis, caule

simplici vel 1-ramoso, spica erecta 7- ad 16-flora dis-

ticha floribus suboppositis vel oppositis, bracteis molli-

bus pallide viridibus siccentibus pallide stramineis post

anthesin bractea, externa (20-)22-32 mm longa interna

ad apicem minute furcata marginibus connatis in dimidio

inferiore, floribus pallide cameis, tepalis inferioribus tri-

bus lateraliter atrorubro-suffusis in tertia parte basali, in

dimidio basali striatis, inodoris, tubo perianthii obliquiter

infundibuliformi 33-35 mm longo, tepalis inaequalibus

superioribus tribus grandioribus ad basem erectis recur-

vatis distaliter lanceolato-attenuatis marginibus leviter

undulatis, tepalo dorsali 33 x 11-12 mm, superioribus

lateralibus 32-34 x 10-12 mm, inferioribus tribus per

1 mm connatis 29-30 x 9-10 mm, filamentis 17 mm

longis ex tubo 7 mm exsertis, antheris 8-9 mm longis

purpureis, stylo arcuato ramis 4 mm longis.

TYPE. — Mpumalanga, 2430 (Pilgrim’s Rest): Kenne-

dy’s Vale, Two Rivers Mine, hills west of bridge over

Klein Dwarsrivier, (-CC), 26 March 2008, Manning &

Valente 3156 (NBG, holo.; K, MO, NBG, PRE, iso.).

Plants 0.4-1. 5 m. Corm subglobose, ± 20 mm diam.;

outer tunics papery, becoming irregularly broken and

somewhat fibrous with age. Stem erect or inclined out-

ward above basal leaves, unbranched or occasionally

with a branch from axil of uppermost stem leaf, 2-3 mm

diam. below spike. Catapylls brownish above ground.

Leaves 8 or 9, lower 6 or 7 basal and largest, reaching to

base of spike or above; blades narrowly sword-shaped,

(3— )5— 1 0 mm wide, relatively soft-textured with midrib

raised but other veins and margins not thickened, slightly

twisted in distal half, remaining 2 or 3 leaves cauline and

smaller, uppermost largely or entirely sheathing, margins

open to base. Spike erect, 7-16-flowered, flowers in two

ranks, subopposite and 60-90° apart, or opposite; bracts

soft-textured, pale green becoming pale straw-coloured

and dry shortly after anthesis, outer (20-)22-32 mm

long, inner slightly shorter or ± as long, minutely forked

apically, margins fused in basal half and thus tubular

below. Flowers pale flesh-pink, tepal sutures and tube

flushed deeper salmon, lower three tepals flushed deep

red laterally in basal third, basal half streaked with three

main and two minor longitudinal lines, upper three tepals

flushed deep red at base and filaments deep red in basal

third, unscented; perianth tube obliquely funnel-shaped,

slightly arched distally, 33-35 mm long, basal cylindri-

cal portion 25-27 mm long; tepals unequal, upper three

larger, erect below, recurved in distal half, lanceolate-

attenuate, margins slightly undulate, dorsal tepal hori-

zontal in basal half and deeply concave, 33 x 1 1-12 mm,

upper laterals 32-34 x 10—12 mm, lower three tepals

joined for an additional 1 mm, lower laterals shortest,

28-29 x 9-10 mm, lower medial 30 x 9-10 mm. Fi la-

ments 17 mm long, exserted 7 mm from tube; anthers

8-9 mm long, purple; pollen cream-coloured. Ovary

ovoid, ± 6 mm long; style arching beneath dorsal tepal,

dividing at or beyond anther tips, branches 4 mm long.

Capsule obovoid, 30 x 12 mm, 3-lobed and retuse api-

cally. Seeds oval to oblong, 8-10 x 4—6 mm, ± evenly

winged or wing not developed on one side, rich brown,

seed body relatively large, ± 3 mm diam. Flowering

time: mid-March to mid-April. Figure 4.

Distribution and ecology. Gladiolus reginae is cur-

rently known only from two populations along the lower

slopes of the Dwarsrivier Mountains (Figure 5), one on

the Two Rivers Platinum Mine concession and the sec-

ond on Steelpoort Park, some 5 km to the west. Plants

grow in open woodland, partially shaded by shrubs and

trees, and are largely restricted to rocky outcrops, where

the corms are wedged in pockets of humus between the

Bothalia 39,1 (2009)

FIGURE 4. — Gladiolus regime, Man-

ning & Valente 3156 (NBG).

A, corm and flowering stem; B,

inner bract; C, half flower; D,

capsule and seeds. Scale bar: 1 0

mm. Artist: John Manning.

rocks. Geologically the substrate has been identified as

the igneous rock gabbro-norite (G. Deall pers. comm.),

which contains slightly higher concentrations of heavy

metals than granite, thus weathering into heavy metal-

enriched soils (Van Wyk & Smith 2001). G. reginae

may thus be considered another of the several edaphic

endemic species of Gladiolus that have been identified

in Iridaceae (Goldblatt & Manning 1996, 1998).

The Dwarsrivier is a tributary of the Steelpoort River,

which bisects the Sekhukhuneland Centre of Floristic

Endemism identified by Van Wyk & Smith (2001). This

mountainous region is relatively poorly known botani-

cally but its rocks hold large reserves of chrome and plati-

num-group metals, and the area supports a rich flora of

local edaphic endemics. Gladiolus reginae is one of two

Gladiolus species endemic to the Sekhukhuneland Centre.

The first to be described, G. sekukuniensis P.J.D. Winter,

is restricted to alkaline calcretes (Manning et al. 1999).

Although over 100 species of plants are estimated to be

endemic or near-endemic to the Sekhukhuneland Centre

(Van Wyk & Smith 2001), the endemic Iridaceae thus far

known include just these two species of Gladiolus. As far

as is known, Gladiolus is the only genus of Iridaceae in

which species have evolved that are tolerant of heavy met-

als (Goldblatt & Manning 1996, 1998).

The long-tubed, unscented, pale pink flowers of Gladi-

olus reginae are evidently adapted to pollination by

long-proboscid flies, and its co-occurrence with the labi-

ate shrub Orthosiphon tubiformis, which is pollinated by

Stenobasipteron wiedmannii (Nemestrinidae) (Goldblatt

Bothalia 39,1 (2009)

43

FIGURE 5. — Known distribution of Gladiolus reginae, •.

& Manning 1999) makes it all but certain that G. reginae

is a member of this pollination guild. Intriguingly, the

species exudes minute droplets of dilute photosynthate

from the tips of the inner and outer bracts. These attract

numerous ants, which crawl over the inflorescence axis

among the flowers. We noted a similar strategy in G.

pole-evansii J.Verd. (Goldblatt & Manning 1998) and

speculated that ants may play a defensive role in deter-

ring predators. This phenomenon had not been noted in

any other species of Gladiolus.

The species was brought to our attention by Graham

Deall, who encountered a population near Steelpoort dur-

ing a botanical survey of the area in February 2006. In

the same season a second population was photographed

in the nearby Twin Rivers Platinum Mine concession by

geologist Louise Fouche. A visit to the Steelpoort locality

in February 2007 after a very dry summer was unsuccess-

ful and the corms had evidently failed to sprout. Good

flowering plants were, however, found on a second visit

to the Twin Rivers Platinum Mine site in March 2008,

following good summer rains. Although plants appear to

be locally common where they occur, the extent of the

species distribution is evidently very limited, making it

vulnerable to increased transformation of the habitat.

Diagnosis and relationships', despite the lack of

pubescence on the stems or leaves, numerous features

point to a relationship with members of series Scabridus.

These include the tall stature of the plants and their nar-

row, relatively soft-textured leaves without visibly thick-

ened margins, papery outer corm tunics, ± erect spike

with 2-ranked, moderately sized, pink flowers, thin-tex-

tured, pale bracts, and large, obovoid capsules. Its habi-

tat, in hot, dry savanna, is also consistent with this rela-

tionship. Within the series, Gladiolus reginae is florally

most similar to G. scabridus M.P.de Vos from northern

KwaZulu-Natal but the glabrous stem and leaves, and the

pale salmon or flesh-pink flowers with a deep red cen-

tre, including the base of the filaments, points to a closer

relationship with G. pavonia Goldblatt & J.C.Manning

from northeastern Mpumalanga. G. reginae differs from

G. pavonia in lacking the characteristic stolons of the

latter species, in its longer floral tube, 33-35 mm vs. ±

16 mm, and in its attenuate rather than acute tepals.

In general appearance, particularly the long-tubed

pink flowers with tubular inner bracts, Gladiolus regi-

nae resembles G. dolomiticus Oberm., another savanna

species from northern South Africa. This dolomite

endemic, however, is covered with a velvety pubescence

on leaves and stem, the leaf blades are distinctly fibrotic,

have thickened margins, the flowers have a shorter tube,

1 8-27 mm long, and the lower tepals are marked with a

median pale blotch rather than with red streaks.

Etymology. Latin reginae, queen, after the legendary

Queen of Sheba (Saba), who controlled rich gold mines

in eastern Africa, in a reference to both the type locality

of the species on the site of a platinum mine and to the

beauty of its flowers.

Other material seen

MPUMALANGA. — 2430 (Pilgrim’s Rest): Steelpoort Park, (-CC),

March 2006 (photo.), April 2006 (fruit), G. Deall s.n. (NBG).

IDENTIFICATION KEYS

The following replacement couplets are provided for insertion in the relevant keys to the species published in

Gladiolus in southern Africa (Goldblatt & Manning 1998).

Key to section Blandus (page 52)

13. Perianth tube longer than the dorsal sepal, (30-)40-100 mm long

14. Tepals attenuate and strongly undulate; lower tepals slightly shorter than the upper G. undulatus

14'. Tepals obtuse to acute, not attenuate and weakly undulate; lower tepals usually about two-thirds as long as the upper

15. Leaves four or five; flowers cream to yellowish with prominent spear-shaped markings in red on the lower tepals; flowering October

to November; West coast mountains and flats G. angustus

15'. Leaves three to seven; flowers salmon with red linear markings on the lower tepals; flowering February to April; southern Cape and

Little Karoo

44

Bothalia 39,1 (2009)

1 5a. Leaves three, the blades 6-8 mm wide; perianth tube 50-70 mm long; flowering March and April, southern foothills of the Lange-

berg G. bilineatus

15a'. Leaves six or seven, the blades 1.5-6. 5 mm wide; perianth tube 30-55 mm long; flowering February, Little Karoo mountains G. dolichosiphon

13’. Perianth tube shorter than to ± as long as the dorsal tepal, (15-)20-50 mm long

Key to section Hebea (page 55)

29'. Perianth tube 9-15 mm long; dorsal tepal 16-33 mm long

29a. Flowers bright yellow; lower lateral tepals with lower half of blade involute-tubular and auricuate G. karooicus

29a'. Flowers whitish to dull grey, purple or brownish yellow; lower lateral tepals with lower half of blade canaliculated and clawed but not

auriculate G. permeablilis

Key to sections Densiflorus and Ophiolyza (page 50)

60'. Leaves narrow and usually exceeding the spikes, in a lax fan usually arising some distance above the ground; leaf blades always with

a prominent midrib and sometimes secondary veins also developed

60a. Plants glabrous; perianth tube 33-35 mm long; lower tepals streaked with red and base of filaments flushed dull red G. reginae

60a'. Plants glabrous or pubescent; perianth tube 10-27 mm long; lower tepals not streaked and filaments concolorous 7

CLASSIFICATION

The discovery of Gladiolus reginae, a member of

series Scabridus (see above), highlights several morpho-

logical inconsistencies in the current placement of the

series in section Densiflorus Goldblatt & J.C. Manning.

These are examined in detail here.

The current classification of Gladiolus in southern

Africa recognizes seven sections, diagnosed princi-

pally by the shape, number and insertion of the leaves

on the stem, the shape and number of the flowers, and

the size of the capsules (Goldblatt & Manning 1998).

Sections Densiflorus and Ophiolyza (Klatt) Goldblatt &

J.C. Manning are characterized by the possession of sev-

eral basally inserted, lanceolate leaves arranged in a fan,

and flowers without distinct spade- or lozenge-shaped

markings on the lower tepals. Both are predominantly

or entirely restricted to the summer rainfall region. Sec-

tion Densiflorus is recognized by its many-flowered,

subsecund spike and small capsules, less than 10 mm

long, and section Ophiolyza by its generally large stat-

ure and especially by flowers with sharply recurved

lower tepals.

There is little doubt that Gladiolus reginae is cor-

rectly placed in series Scabridus, despite the lack of

the pubescence on leaves and/or stem that characterizes

many members of the series. The large stature of the

plants, their slender, soft-textured leaves without thick-

ened margins, and the 2-ranked spike of moderately

large, pink flowers with the lower tepals marked with

red longitudinal lines are all characteristic of the series.

Series Scabridus is currently included in section Densi-

florus but its placement at the end of the section and just

before section Ophiolyza indicates its somewhat inter-

mediate character.

The possibility that the series is incorrectly placed

in section Densiflorus is highlighted by the observation

that Gladiolus reginae shares several highly distinc-

tive features with G. pole-evansii, a member of section

Ophiolyza. The most unusual of these is the fusion of the

margins of the inner bracts in the basal portion such that

they are tubular below. This characteristic was identified

as diagnostic of series Oppositiflorus by Goldblatt &

Manning (1989). Its occurrence in G. reginae must thus

be interpreted either as a homoplasy, or as evidence for

a closer alliance between the species (and by inference

the entire section Scabridus ) and series Oppositiflorus.

The shared occurrence of other unusual features in G.

reginae and G. pole-evansii, however, makes the latter

interpretation more likely. These include the exudation

of droplets from the tips of the bracts and bracteoles (a

character unknown elsewhere in the genus), the suboppo-

site or opposite arrangement of the flowers in the spike

(a second diagnostic characteristic of series Oppositi-

florus and found also among several other members of

series Scabr-idus but anomalous in section Densiflorus),

the recurving lower tepals (diagnostic of section Ophio-

lyza) and the large, obovoid capsules (characteristic

of most members of series Scabridus and anomalous

in section Densiflorus). Coupled with this is the shared

development of velvety pubescence in several members

of both series Scabridus and Oppositiflorus and its other-

wise total absence in section Densiflorus.

All the evidence thus indicates that series Scabridus is

misplaced in section Densiflorus and is actually closely

allied to series Oppositiflorus of section Ophiolyza.

We accordingly move the series to that section. The

exact relationship between series Scabridus and series

Oppositiflorus is less easily determined, however, and

we retain the two as separate pending further study.

Until then it is least disruptive to merely transfer series

Scabridus from the end of section Densiflorus to the

beginning of section Ophiolyza, a move that will not

require rearrangement of the sequence of species in the

current classification (Table 2).

TABLE 2. — Summary of revised classification of Gladiolus in southern

Africa

Gladiolus L.

Section Densiflorus Goldblatt & J.C. Manning [series not listed]

Section Ophiolyza (Klatt) Goldblatt & J.C. Manning

Series Scabridus

Series Oppositiflorus

Series Ecklonii

Series Ophiolyza

Section Blandus (Baker) Goldblatt [series not listed]

Section Linearifolius (M.P.de Vos) Goldblatt & J.C. Manning [series

not listed]

Section Heterocolon O.Kuntze [series not listed]

Section Hebea (Pers.) Benth. & Hook, [series not listed]

Section Homoglossum (Salisb.) Goldblatt & J.C. Manning [series not

listed] ___

Bothalia 39,1 (2009)

45

NEW NAME

Gladiolus sulculatus Goldblatt, nom nov., pro

G. sulcatus Goldblatt, Gladiolus in tropical Africa: 1 1 3

(1996), nom. illeg. non G. sulcatus Lam.: 119 (1791) [=

Babiana hirsuta (Lam.) Goldblatt & J.C. Manning],

The name Gladiolus sulcatus Goldblatt (1996) is

a later homonym for G. sulcatus Lam. (March 1791),

now Babiana hirsuta (Lam.) Goldblatt & J.C. Manning

(= Babiana thunbergii Ker Gawl.) (Goldblatt & Man-

ning 2006). We rename the species G. sulculatus , thus

preserving the reference in the specific epithet to the

finely ribbed leaves of this rare species of central Tan-

zania. Although G. sulcatus Lam. is a superfluous name

for Antholyza hirsutus Lam., it is nevertheless valid

(McNeill et al. 2006 Art. 52.3). Possibly Lamarck (1791)

provided the new epithet in Gladiolus because he was

aware of the impending publication of G. hirsutus Jacq.,

dated imprecisely as Tate’ 1791.

RANGE EXTENSION

Gladiolus rubellus Goldblatt, a member of section

Heterocolon , is readily recognized by its small, scarlet

flowers with bright yellow markings on the lower lat-

eral tepals, hooded dorsal tepal, and linear leaves with

thickened margins and midrib, the margins raised at

right angles to the blade (Goldblatt 1993; Goldblatt &

Manning 1998). It is currently known only from a small

region of southeastern Botswana, between the towns of

Lobatse and Molepolole. Plants grow in stony ground

in dry savanna and flower in late summer and autumn,

between January and March.

On a recent field trip to northern Namibia by staff

of the South African National Biodiversity Institute, a

species of Gladiolus was photographed in the Baynes

Mountains in the northwest of the country that represents

a species not before recorded for the country. It is evi-

dently close to G. rubellus, although the flowers may dif-

fer slightly in the relative proportions of the lower tepals,

possibly less intense red colouring of the perianth, and

leaf margins that are thickened rather than raised at right

angles to the blade surface. It is difficult to assess these

characters more accurately from a single photograph but

there is no doubt as to the importance of this record as

just three species of Gladiolus are currently known from

the summer rainfall part of Namibia, G. dalenii van

Geel, G. permeabilis subsp. edulis (Burch, ex Ker Gawl.)

Oberm. and G. saccatus (Klatt) Goldblatt & M.P.de Vos,

all of them widespread in southern Africa. Pending the

collection of additional material, we include the Baynes

Mountain population in G. rubellus, thereby extending

the range of the species some 1 300 km to the northwest

and rendering it no longer endemic to Botswana.

Material examined

NAMIBIA. — 1712 (Posto Velho): Cunene Region, Baynes Moun-

tains, Omavanda Mountains, open savanna on gentle SE slope, (-BB),

25-30 April 2008, G. Nicolson s.n. (NBG, photo.).

ACKNOWLEDGEMENTS

Support for this study from Elizabeth Parker, and by

grant 7799-05 from the National Geographic Society, are

gratefully acknowledged. Collecting permits were pro-

vided by the Nature Conservation authorities of Northern

Cape and Western Cape, South Africa. We thank Lendon

Porter and Elizabeth Parker for their assistance and com-

panionship in the field; Graham Deall for collecting fruit-

ing material of Gladiolus reginae and for guiding us to

the species; AnneLise and Jan Vlok for collecting mate-

rial of Gladiolus dolichosiphon; and Greg Nicolson for

alerting us to the existence of G. rubellus in Namibia.

REFERENCES

GEERINCK, D. 2001. Nouvelle espece: Gladiolus goldblattianus Geer-

inck ainsi que les deux combinations nouvelles: G. erectiflorus

var. verdickii, G. gregarius var. anguslifolius pour la famille des

Iridaceae en Afrique centrale. Taxonomania 4: 2, 3.

GOLDBLATT, P. 1989. Systematics of Gladiolus L. (Iridaceae-Ixioi-

deae) in Madagascar. Adansonia 3: 235-255.

GOLDBLATT, P. 1 993 . Iridaceae. In G. V. Pope, Flora zambesiaca 1 2, 4:

1-106. Flora Zambesiaca Managing Committee, London.

GOLDBLATT, P. 1996. Gladiolus in tropical Africa. Timber Press,

Portland, Oregon.

GOLDBLATT, P. 2008. Additions to Gladiolus (Iridaceae) in the Flora

zambesiaca region: the new species G. metallicola and the rein-

statement of G. mosambicensis. Novon 18: 164—167.

GOLDBLATT, P. & MANNING, J.C. 1996. Two new edaphic endem-

ic species and taxonomic changes in Gladiolus (Iridaceae) of

southern Africa, and notes on Iridaceae restricted to unusual sub-

strates. Annals of the Missouri Botanical Garden 86: 758-774.

GOLDBLATT, P. & MANNING, J.C. 1998. Gladiolus in southern Afri-

ca. Femwood Press, Cape Town.

GOLDBLATT, P. & MANNING, J.C. 1999. The long-proboscid fly pol-

lination system in Gladiolus (Iridaceae). Annals of the Missouri

Botanical Garden 86: 758-774.

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

of the Cape flora of South Africa. Strelizia 9.

GOLDBLATT, P. & MANNING, J.C. 2006. Systematics of the southern

African genus Babiana (Iridaceae). Strelitzia 18.

JACQUIN, N.J. VON 1791. Collecteana botanicam 4. Wappler, Vienna.

LAMARCK, J.B.A.P. DE 1791 . Tableau encyclopedique et methodique

1 . Panckoucke, Paris.

LEWIS, G. J. 1 972. A revision of the South African species of Gladiolus.

Journal of South African Botany, Suppl. vol. 10.

MANNING, J.C., GOLDBLATT, P. & WINTER, P.D.J. 1999. Two new

species of Gladiolus (Iridaceae: Ixioideae) from southern Africa

and notes on long-proboscid fly pollination in the genus. Both-

alia 29:217-223.

McNEILL, J., BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,

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

PRADO, J., SILVA, P.C., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. (eds). 2006. International Code of Botanical

Nomenclature (Vienna Code) adopted by the 1 7th International

Botanical Congress, Vienna, Austria, July 2005. Regnum Vegeta-

bile 146. Gantner, Ruggell.

VAN WYK, A.E. & SMITH, G.F. 200 1 . Regions offloristic endemism in

southern Africa. A review with emphasis on succulents. Umdaus

Press, Hatfield, Pretoria.

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Bothalia 39,1: 47-65 (2009)

A revision of Fumariaceae (Fumarioideae) in southern Africa, includ-

ing naturalized taxa

J.C. MANNING*, P. GOLDBLATT** and F. FOREST***

Keywords: Cysticapnos Mill., Discocapnos Cham. & Schltdl., Fumaria L., Fumariaceae, southern Africa, taxonomy, Trigonocapnos Schltr.

ABSTRACT

The native and naturalized species of Fumariaceae in southern Africa are reviewed, and keys and full descriptions are

provided. All relevant regional synonyms are given and the indigenous species are illustrated. Three native genera with four

species are recognized. The previously taxonomically unplaced genus Cysticapnos Mill. (3 spp.) is included with two other

monotypic native South African genera, Discocapnos Cham. & Schltdl. and Trigonocapnos Schltr., in an enlarged circum-

scription of subtribe Discocapninae, which is morphologically defined by tendrilliferous leaves and mostly sessile racemes

with the lowermost flower basal. Two subspecies are recognized in Cysticapnos vesicaria (E.Mey. ex Bemh.) Liden, subsp.

vesicaria with fewer, smaller flowers and subsp. namaquensis J.C. Manning & Goldblatt for plants from Namaqualand

with more numerous, often larger flowers. C. parviflora Liden appears to be nothing more than a dwarf-fruited form of C.

vesicaria , in which heterocarpy has long been known. C. prninosa (E.Mey. ex Bemh.) Liden is recorded for the first time to

be a short-lived perennial and not an annual, thus unique in the tribe Fumarieae. Discocapnos mundii var. dregei Harv. from

the southern Cape is treated as subsp. dregei (Harv.) J.C. Manning & Goldblatt on account of its slightly smaller fruits and

significant geographical disjunction from subsp. mundii. Three introduced species are included, Fumaria capreolata L., F.

muralis Sond. ex W.D.Koch subsp. muralis and F. parviflora Lam. var. parviflora, although only the last two appear to be

truly naturalized.

INTRODUCTION

Fumariaceae are a well-circumscribed family, sister

to a monophyletic Papaveraceae sensu stricto (APG

2003). Although included in a broader circumscription

of Papaveraceae by some authors, e.g. Bremer et al.

(2003), the two groups represent reciprocally mono-

phyletic clades that are optionally retained as separate

families (APG 2003). We adopt the latter option, fol-

lowing Liden (1993) as the most recent complete clas-

sification of Fumariaceae. Treating Fumariaceae in

Papaveraceae as subfamily Fumarioideae has nomen-

clatural implications on the rank of lower-order taxa

that have yet to be fully implemented.

Fumariaceae are readily distinguished from Papav-

eraceae by the colourless or yellow, watery or trans-

lucent (not milky) exudate, strongly zygomorphic or

bilaterally symmetric flowers with small sepals, and

petals that are not crumpled in bud. In addition, one

or both of the outer petals are spurred basally, the sta-

mens are usually connate into two bundles, and the

bicarpellate, syncarpous ovary ripens into a capsule

or nut (Bremer et al. 2003; Liden 1993).

Fumariaceae are primarily north temperate, com-

prising 17 genera and ± 530 species. The family is

taxonomically relatively well understood (Liden

1993), and comprises the two subfamilies Fumari-

oideae (DC.) Endl. and Hypecoideae (Dum.) Prantl

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

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

den, P.O. Box 299, St. Louis, Missouri 63166, USA.

*** Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Sur-

rey, TW9 3DS, UK.

MS. received: 2008-08-06.

& Kundig, the latter consisting of the single genus

Hypecoum L. from the Mediterranean and Asia. Sub-

family Fumarioideae includes the tribe Corydaleae

Rchb., characterized by many-seeded capsules con-

taining seeds with a conspicuous elaiosome, and the

tribe Fumarieae, mostly with nuts or few-seeded cap-

sules, usually a caducous style, and seeds lacking an

elaiosome. The tribe Fumarieae is in turn divided into

the three subtribes Sarcocapninae Liden, Fumariinae,

and Discocapninae Liden.

The family is poorly represented in sub-Saharan

Africa with just two species known from tropical

Africa (Lucas 1962), Fumaria abyssinica Hamm,

and Corydalis mildbraedii Fedde. Both are widely

distributed from Ethiopia to Kenya and Tanzania,

and represent large genera that are primarily north-

ern hemisphere in distribution. This contrasts sharply

with the situation in southern Africa, which is home

to five species in three genera, all endemic to the sub-

region. Two of these genera, Discocapnos Cham. &

Schltdl. and Trigonocapnos Schltr., are monotypic and

together comprise subtribe Discocapninae Liden but

the third genus Cysticapnos Miller (3 spp.) is not cur-

rently placed at tribal level (Liden 1986, 1993). The

subtribe Discocapninae is characterized by tendrillif-

erous leaves and single-seeded, pubescent achenes.

Cysticapnos has similar tendrilliferous leaves but a

many-seeded, dehiscent, capsular fruit. DNA studies

(Forest & Manning in prep.) indicate that the three

southern African genera, Cysticapnos, Discocapnos

and Trigonocapnos constitute a monophyletic lineage

within Fumarieae, and all three genera are thus most

appropriately accommodated in an enlarged Disco-

capninae. The subtribe is defined morphologically by

its tendrilliferous leaves, found elsewhere in the fam-

ily only in the Mediterranean genus Ceratocapnos

Dur. (tribe Fumarieae subtribe Sarcocapninae) and in

48

Dactylicapnos Wall, (tribe Corydaleae Rchb.) from

the Himalayas. Several of the species also have ses-

sile racemes (the lowermost flower arising at the base

of the peduncle), whereas other genera of Fumarieae

typically have distinctly pedunculate racemes (the

lowermost flower inserted well above the base of the

peduncle). Anatomy of the seed coat is essentially

identical in Cysticapnos, Discocapnos and Trigono-

capnos (Fukuhara & Liden 1995a) and also in sev-

eral other genera of Fumarieae and Corydaleae, thus

offering no additional synapomorphies for the sub-

tribe. Of potentially greater interest is the observation

that Discocapnos and Trigonocapnos share an unu-

sual vasculature of the fruit wall in which the funicu-

lar supply is ultimately derived from the ventral and

dorsal bundles rather than from the lateral bundles

as in other fumarioids (Fukuhara 1995; Fukuhara &

Liden 1995b). Unfortunately the pericarp vasculature

of Cysticapnos has not been studied and it is there-

fore not known which pattern it follows.

The molecular analysis (Forest & Manning in

prep.) retrieves Discocapnos as sister to Trigono-

capnos plus a monophyletic Cysticapnos, with the

entire clade sister to Fumaria L. This relationship

supports the interpretation that the nutlet is plesio-

morphic for the subtribe Discocapninae and that the

capsules of Cysticapnos represent an autapomor-

phy for the genus. This topology offers two logical

alternatives for circumscribing genera in the subtribe

Discocapnineae. The first retains three separate gen-

era as circumscribed by Liden (1986, 1993), whereas

the second includes Discocapnos and Trigonocapnos

within an enlarged Cysticapnos. The latter treatment

is not consistent with the current circumscription of

genera in the family (Liden 1993), in which fruit type

is extensively used as the basis for separating related

genera, resulting in constellations of oligotypic gen-

era clustered around the two large genera Corydalis

DC. and Fumaria. The merits of this approach may

be debated but at this stage it is unjustifiable to treat

the southern African genera in a radically different

way in the face of the molecular evidence that they

are indeed monophyletic as currently construed. The

capsular fruits of Cysticapnos represent a significant

evolutionary innovation that in the current classifi-

cation justifies its retention as a separate genus. Any

changes to generic circumscriptions in the tribe Dis-

cocapninae are best done in the context of a reap-

praisal of the entire family. The southern African

clade is adequately identified as the subtribe Disco-

Bothalia 39,1 (2009)

capninae for purposes of biogeographic or phyloge-

netic investigation.

The first southern African species of Fumariaceae to

be described were placed either in the genus Corydalis

(capsular-fruited species) or Fumaria (those with nut-

lets). The varied and distinctive fruits of several of the

southern African taxa, however, soon led to their taxo-

nomic separation into distinct genera, beginning with the

genus Cysticapnos (Miller 1754), erected for the blad-

der-fruited species that was until then known as Cory-

dalis vesicaria L. The genus Phacocapnos Bemh. was

later described (Bemhardi 1838) for the remaining cap-

sular-fruited species with compressed fruits but is now

included within Cysticapnos (Liden 1986). The two nut-

let-fruited species remain respectively in the monotypic

genera Discocapnos (Chamisso & Schlechtendel 1826)

and Trigonocapnos (Schlechter 1 899).

The southern African Fumariaceae were first reviewed

by Harvey (1894) for Flora capensis and later more thor-

oughly by Hutchinson (1921) as part of his treatise on the

biogeography of the family. The most recent, and only

modem revision of the southern African species, is the

worldwide study of the family by Liden (1986), in which

the current generic concepts and classification were

established, and the application of the names of the spe-

cies was fixed. A milestone in the study of the family, this

treatment is nomenclaturally incomplete and is also based

on examination of very limited material of the southern

African taxa, essentially the types. With more extensive

material available, it is evident that two of the species

recognized there, viz. Cysticapnos grandiflora sensu

Liden and C. parviflora Liden, cannot be upheld. In addi-

tion, the patterns of geographical variation shown in both

C. vesicaria and Discocapnos mundii, support the recog-

nition of subspecies in each. We provide here a complete,

illustrated account of the native species of southern Afri-

can Fumariaceae, based on extensive field and herbarium

study. Three introduced species of Fumaria that have

become ± naturalized are also included, following the

taxonomy of Liden (1986).

MATERIALS AND METHODS

This study is based on an examination of living

plants in the field and of specimens in the following

herbaria: BOL, K, MO, NBG, NU, PRE, TCD (acro-

nyms as in Holmgren et al. 1990). The abbreviation of

authors’ names follows Brummitt & Powell (1992).

Key to genera of Fumariaceae in southern Africa

la Leaves not tendrilliferous; racemes pedunculate; style twice or more as long as ovary, caducous; fruiting pedicels usually suberect,

thickened; fruit a subglobose nutlet with woody endocarp subtribe Fumariinae: Fumaria (introduced species)

lb Terminal leaflets of some or all leaves developed into tendrils; racemes pedunculate or sessile (lowermost flower basal); style

± as long as ovary or shorter, usually persistent; fruiting pedicels recurved, not thickened; fruit a capsule or achene, without

woody endocarp subtribe Discocapninae:

2a Fruit a several-seeded capsule; spur on upper petal ± as long as wide or shorter; style less than half as long as ovary, persistent

3. Cysticapnos

2b Fruit a 1 -seeded, pubescent or scabridulous achene; spur on upper petal ± twice as long as wide; style ± as long as ovary, persistent or

caducous:

3a Flowers and fruit with pedicels ± 1 .5 mm long; achene disciform, sparsely puberulous; style persistent 1 . Discocapnos

3b Flowers and fruit with pedicels 3^1 mm long; achene obovate-trigonous, densely scabridulous; style caducous 2. Trigonocapnos

Bothalia 39,1 (2009)

49

I. Subtribe Discocapninae Liden, Opera Botanica

88: 104 (1986), emend. J.C. Manning & Goldblatt. Type

genus: Discocapnos Cham. & Schltdl.

Scandent annuals or rarely perennial (one species

only) with bipinnate, tendrilliferous leaves. Racemes

pedunculate or sessile (lowermost flower basal). Flow-

ers pink with purple tip, upper petal spurred, inner pet-

als apically keeled or winged. Style usually persistent

(rarely deciduous); stigma with two papillae. Fruit either

a many-seeded capsule or a one-seeded achene, without

a woody endocarp.

Three genera: Cysticapnos, Discocapnos and Trigono-

capnos. Southern Africa.

1. Discocapnos Cham. & Schltdl., Linnaea 1: 569

(1826); Harv.: 18 (1894); Hutch.: 114 (1921); Liden: 104

(1986), 316 (1993). Fumaria sect. Discocapnos (Cham.

& Schltdl.) Prantl in Prantl & Kiindig: 145 (1891). Type

species: D. mundii Cham. & Schltdl.

Brittle, semi-succulent, glabrous, climbing annual;

stems quadrate, with obscure unicellular papillae along

angles; sap watery, yellow. Leaves alternate, bipinnately

compound, primary divisions alternate, ultimate leaflets

obovate and 3-partite; terminal leaflets of all or only

upper leaves transformed into tendrils. Inflorescence

pedunculate, terminal but leaf-opposed through rapid

growth of axillary bud, racemose; bracts scale-like, peta-

loid, irregularly toothed. Flowers zygomorphic, bilabi-

ate, pink, unscented. Sepals 2, lateral, much shorter than

petals, scale-like, petaloid, irregularly toothed. Petals 4

in 2 series, outer petals larger, naviculate-spathulate, lat-

erally winged apically, upper spurred at base, inner pet-

als apically connate, clawed, limb inflated with dorsal

crest, claw basally adnate to upper petal. Stamens 6, dia-

delphous in abaxial and adaxial bundles of 3; filaments

of each bundle fused, lanceolate, membranous, clasping

ovary, upper filament cluster adnate to margins of dor-

sal petal basally to form small chamber, with nectary

decurrent on dorsal petal into spur. Ovary suborbicular,

dorsoventrally compressed with peripheral rim, bearing

unicellular papillae; ovule solitary, lateral; style short,

± as long as ovary, flexed upwards apically, persistent;

stigma compressed, 2-lobed. Fruit pendulous, dorsov-

entrally compressed, suborbicular with peripheral wing,

pubescent, indehiscent and dropping entire, papery. Seed

solitary, lenticular, glossy black, colliculate, without

elaiosome.

One sp., South Africa, southwestern and southern Cape.

1.1. Discocapnos mundii Cham. & Schltdl. [as

‘ mundtii ’] in Linnaea 1: 569 (1826); Harv.: 18 (1894);

Hutch.: 114 (1921). Liden: 104 (1986). Fumaria mun-

dtii (Cham. & Schltdl.) Prantl & Kiindig: 145 (1891).

Type: South Africa, Cape of Good Hope, without precise

locality, August [without year], Mund & Maire s.n. [B-

WILLD, holo.!].

Climbing annual up to 1 m, with yellow sap. Leaves

bipinnately compound, rachis flexuouse, ultimate seg-

ments broadly obovate, 2- or 3-lobed almost to base,

lobes elliptical to obovate, terminal segments trans-

formed into tendrils. Inflorescence up to 100 mm long in

fruit, 8-15-flowered; pedicels erect, apically recurved,

± 1.5 mm long; bracts erect, clasping pedicel, i 1 x 0.5

mm, irregularly toothed. Flowers pale pink with dark

reddish tips. Sepals peltate, ovate, ± 1.0 x 1.5 mm, irreg-

ularly toothed. Petals: outer petals naviculate-spathulate,

4. 5-5.0 x 1.0-1. 5 mm, apically short-winged, wings pat-

ent, ± 0.5 mm wide, upper spurred at base, spur 2. 5-3.0

mm long; inner petals ± 4 mm long, claw ±1.5 mm long,

adnate to upper petal in lower 1 mm, blade inflated,

ellipsoid, 2.5 x 1 mm, with fleshy dorsal crest ± 1mm

wide. Stamens: filament bundles lanceolate, 2. 5-3.0 x 1

mm, basal nectary decurrent up back of upper petal spur

to near apex, vestigial; anthers ± 0.25 mm long, yellow.

Ovary discus-shaped, 2x2 mm, green, minutely papil-

late; style maroon, flexed sharply upwards apically, ±1.5

mm long, persistent. Fruit pendent, discus-shaped with

peripheral, radially costate wing, (3-)4— 5 mm diam.,

pubescent with unicellular trichomes longest over seed-

body, papery, indehiscent and dropping entire, brownish

black when mature. Seed lenticular, (3— )4 — 5 mm diam.,

glossy black, colliculate. Flowering time : (August) Octo-

ber-December. Figure 1 .

Distribution and ecology, apparently a rare species,

with a discontinuous distribution along the cooler, south-

ern foothills of the coastal mountains of the southwest-

ern and southern Cape (Figure 2). The species has been

recorded from the Cape Peninsula and adjacent moun-

tains in the southwest, and then some 300 km to the east

in the Outeniqua and Tsitsikamma Mountain ranges. Dis-

cocapnos mundii is restricted to moist, loamy or humic

soils on the edge of forest and bush clumps, usually in

sheltered valleys or drainage lines, where plants scram-

ble among bushes and grasses. Its distribution tracks

the scattered occurrence of affotemperate forests in the

southwestern and southern Cape but it is curious that the

species has not been recorded from forest patches along

the southern slopes of the intervening Riviersonderend

or Langeberg ranges. The plants are, however, incon-

spicuous and easily overlooked and suitable habitats in

these mountain ranges should be investigated in search

of additional populations linking the two known areas of

occurrence.

Diagnosis and relationships : Discocapnos mundii

is recognized by its distinctive, discus-shaped samaras

containing a solitary seed with a colliculate testa. The

seeds of the other southern African species of Fumar-

iaceae have scalariform-colliculate surface sculpturing.

The one-seeded, indehiscent fruits with pubescent peri-

carp are shared with Trigonocapnos lichtensteinii but the

trichomes in D. mundii , with their uniquely sculptured

cuticular ornamentation (Liden 1986), are already evi-

dent at anthesis rather than developing after fertilization,

as in T. lichtensteinii , possibly indicating that the peri-

carp vestiture is independently derived in the two spe-

cies. Other similarities between the two include the flex-

uous leaf rachis with the primary divisions of the blade

alternate, unlike those of C. cracca and C. vesicaria,

which are often opposite, the strongly spurred dorsal

petal, and the relatively rudimentary wings on the lower

petal. These characters, however, are also shared with

Fumaria and are thus evidently ancestral conditions and

thus not indicative of a sister-species relationship.

History, the species has an uncomplicated history,

presumably on account of its rarity and distinctive fruits.

50

Bothalia 39,1 (2009)

FIGURE 1 .- Dicocapnos mundii subsp. mundir. A, portion of stem; B, lower leaf; C, flower, front and lateral views; D, lower outer petal; E, inner

petals; F, androecium and section through spur; G, gynoecium, dorsal and lateral views; H, fruit; I, seed. Scale bar: A, B, 10 mm, L U,

mm; H, 1.6 mm; I, 1 .2 mm. Artist: John Manning.

Bothalia 39,1 (2009)

51

FIGURE 2. — Known distribution of Discocapnos mundii subsp. mun-

dii, •; D. mundii subsp. dregei, A; and Trigonocapnos lichten-

steinii, O.

It was first collected by the German pharmacist Johannes

Mund, who had been despatched to the Cape in 1816 as

official plant collector by the Prussian government (Gunn

& Codd 1981). His fruiting collection of Discocapnos

mundii was made in the vicinity of Cape Town, probably

soon after his arrival. Mund met Chamisso in Cape Town

in 1818 (Gunn & Codd 1981), who subsequently named

the species for him (Chamisso & Schlechtendel 1826)

in the new genus Discocapnos on account of its unusual

disc-like fruit. It has been recognized under this name

ever since. The original spelling of the epithet as mundtii

should be treated as an orthographic error and corrected

to mundii (McNeil et al. 2006: Art. 60.7) since his letters

to Hooker are signed L. Mund (Gunn & Codd 1981).

Key to subspecies

la Fruits mostly 4—5 mm long subsp. mundii

lb Fruits ± 3 mm long subsp. dregei

1.1a. subsp. mundii

Fruits (3.5)4— 5 mm diam., with peripheral wing 0.8-

1 .0 mm wide.

Distribution : restricted to the southwestern Cape and

mostly recorded from the Cape Peninsula, where it is

best known from the lower slopes of Table Mountain

between Devil’s Peak and Constantiaberg, with addi-

tional collections from the northern Hottentots Holland

Mountains around Jonkershoek (Figure 2).

1.1b. subsp. dregei (Harv.) J.C. Manning & Gold-

blatt, stat. nov.

D. mundii [as mundtii ] var. dregei Harv. in Flora capensis 1: 18

(1894). D. dregei (Harv.) Hutch.: 114 (1921). Type: South Africa,

[Western Cape], Knysna, Bosch River, between the bush and the river,

October 1839, Drege s.n. (TCD, holo.!; K, S, iso.!).

Fumaria scandens E.Mey.: 125, 186 (1844), nom. nud.

Fruits 3. 0-3. 5 mm diam., with peripheral wing 0.5-

0.8 mm wide.

Distribution : known from three collections along the

southern foothills of the Outeniqua and Tsitsikamma

Mountains, between Knysna and Humansdorp (Figure 2).

Distinguished from the typical subspecies by its

slightly smaller fruits, 3. 0-3. 5 mm diam., with narrower

wing less than 1 mm wide.

History, this taxon is based on a collection made near

Knysna by the German botanical collector Johann Drege

in October 1839. Ernst Meyer, who wrote the botanical

accounts of Drege ’s collections, intended to describe

the species as Fumaria scandens, and this name appears

on the TCD sheet but was never published beyond its

appearance in Drege’s Zwei pflanzengeographische

Dokumente (1844). The smaller fruits of these plants

compared with those of the typical forms from the Cape

Peninsula led Harvey (1894) to distinguish them as the

variety dregei, which he further defined by its more

floriferous racemes. The taxon was subsequently recog-

nized as a distinct species by the British botanist John

Hutchinson (1921) in his account of the family. Further

collections of the typical form show that there are no

differences in floral characters between the two sets of

populations. The difference in fruit size did not impress

Liden (1986), who treated them as comprising a single

entity, and indeed there are apparently no significant

floral or vegetative differences between the two sets of

populations. A subsequent collection of var. dregei from

Clarkson ( Thode s.n. BOL) confirms that the eastern

populations are consistently smaller-fruited but the size

difference between the two sets of populations is slight,

and a plant from Jonkershoek ( Levyns 8511 BOL) bears

occasional fruits of 3.5 mm diam. among the more typi-

cal larger fruits of 4 mm diam. The slight but consistent

difference in average size of the fruits between the west-

ern and eastern populations, combined with the signifi-

cant disjunction between them, supports taxonomic rec-

ognition at some level, and we treat them as subspecies.

2. Trigonocapnos Schltr. in Botanische Jahrbiicher

27: 131 (1899); Hutch.: 113 (1921); Liden: 105 (1986),

316 (1993). Type species: T. curvipes Schltr. [= T. lich-

tensteinii (Cham. & Schltdl.) Liden],

Brittle, semi-succulent, glabrous, climbing annual;

stems quadrate, with obscure unicellular papillae along

angles; sap watery, yellow. Leaves alternate, bipinnately

compound, primary divisions alternate; ultimate leaflets

obovate and 3-partite; terminal leaflets of all or only

upper leaves transformed into tendrils. Inflorescence

shortly pedunculate or sessile, terminal but leaf-opposed

through rapid growth of axillary bud, racemose; bracts

scale-like, petaloid, irregularly toothed. Flowers zygo-

morphic, bilabiate, pink, unscented. Sepals 2, lateral,

much shorter than petals, scale-like, petaloid, irregu-

larly toothed. Petals 4 in 2 series, outer petals larger,

naviculate-spathulate, laterally winged apically, upper

spurred at base, inner petals apically connate, clawed,

limb inflated with dorsal crest, claw basally adnate to

upper petal. Stamens 6, diadelphous in abaxial and adax-

ial bundles of 3; filaments of each bundle fused, lanceo-

late, membranous, clasping ovary, upper filament cluster

adnate to margins of dorsal petal basally to form small

chamber, with vestigial nectary decurrent on dorsal petal

into spur. Ovary obovoid with peripheral rim, concave

52

Bothalia 39,1 (2009)

abaxially, bearing unicellular papillae; ovule solitary,

sub-basal; style short, ± as long as ovary, flexed upwards

apically, deciduous; stigma compressed, 2-lobed. Fruit

pendulous, dorsally crested with lateral rims folded

downwards, thus abaxially conduplicate, puberulous,

indehiscent and dropping entire, leathery. Seed solitary,

narrowly obovoid, yellowish brown, scalariform-collicu-

late, without elaiosome.

One sp., South Africa, northwestern parts of winter rain-

fall region.

2.1. Trigonocapnos lichtensteinii (Cham. &

Schltdl.) Liden in Opera Botanica 88: 105 (1986).

Fumaria lichtensteinii Cham. & Schltdl.: 568 (1826).

Type: South Africa, [Western Cape], Cape of Good

Hope, without precise locality or date, Lichtenstein s.n.

( B-WILLD12938 , holo.!).

T. curvipes Schltr. : 131 (1899); Hutch.: 113 (1921). Type: South

Africa, [Western Cape], Clanwilliam, 21 August 1896, R. Schlechter

8585 (BOL, lecto.!, here designated; S, iso.).

F. capreolata (3? burchellii DC: 130 (1824), name without rank.

Type: South Africa, Karoo [Roggeveld], 1 August 1811 , Burchell 1298

(not located at B, BM, K or P).

F. eckloniana Sond. in Harv.: 18 (1894), nom. nud.

F. micrantha Licht. ms. Lichtenstein s.n. (B-WILLD12938).

Climbing annual up to 1 m, with yellow sap. Leaves

bipinnately compound, rachis flexuous, ultimate segments

broadly obovate, 2- or 3-lobed to base, lobes elliptical, ter-

minal segments transformed into tendrils. Inflorescence up

to 70 mm long in fruit, 20^10-flowered; pedicels suberect,

apically recurved, 3^1 mm long; bracts erect, clasping pedi-

cel, 1.0-1. 5 x 0.5 mm, irregularly toothed. Flowers pale

pink with dark purple tips. Sepals peltate, ovate, ± 0.5 mm

long, irregularly toothed. Petals', outer petals naviculate-

spathulate, 2. 5-3.0 x 0. 5-1.0 mm, apically short-winged,

wings patent, ± 0.5 mm wide, upper spurred at base, spur

2. 0-2. 5 mm long; inner petals ± 3 mm long, claw ± 1.5 mm

long, adnate to upper petal in lower 1 mm, blade inflated,

ellipsoid, 1 x 0.75 mm, with fleshy dorsal crest ± 0.75

mm wide. Stamens : filament bundles oblong, 1. 5-2.0 x 1

mm, basal nectary decurrent up back of upper petal spur to

near apex, vestigial; anthers ± 0. 1 mm long, yellow. Ovary

obovoid with peripheral rim, concave beneath, 1 x 0.5 mm,

green, colliculate; style translucent, flexed sharply upwards

apically, ± 1 mm long, basally constricted, deciduous. Fruit

pendent, obovoid-trigonous, dorsally crested with lateral

rims folded downwards, thus abaxially conduplicate, 3 mm

long, 1.5 mm high and 1.0 mm wide, puberulous, indehis-

cent and dropping entire, brown when mature, leathery.

Seed narrowly obovoid, 2 x 1 mm, yellowish brown, sca-

lariform-colliculate. Flowering time : August-September.

Figure 3.

Distribution and ecology'. Trigonocapnos lichtensteinii

occurs along the Bokkeveld-Matsikamma Escarpment

southwards through the northern Cedarberg to Clanwil-

liam, and inland from the foot of the Hantamsberg at

Calvinia along the Roggeveld Escarpment as far south as

the Koedoesberg at the northwestern edge of the Klein

Roggeveld (Figure 2). Plants grow in sheltered, season-

ally moist situations, along drainage lines or at the base

of rocky outcrops in fine-grained clay soils, where they

clamber among the surrounding shrubbery.

Diagnosis and relationships: the tricostate, puberu-

lous achenes are diagnostic for the species, as is the nar-

rowly obovoid seed. The style is basally constricted and

deciduous, and in this respect Trigonocapnos lichten-

steinii more closely resembles species of Fumaria rather

than the indigenous southern Africa taxa of Fumariaceae,

in which the style is persistent. The curiously shaped

fruit is the result of progressive downward curvature of

the margins of the ventrally concave, disc-shaped ovary

during ripening of the fruit. In its puberulous, single-

seeded, indehiscent fruit, T. lichtensteinii most resem-

bles Discocapnos mundii, but in that species the fruit is

disciform and less obviously puberulous, and contains a

rounded, lenticular seed. The trichomes on the fruit of T.

lichtensteinii, unlike those of D. mundii, are lacking in

cuticular ornamentation (Liden 1986) and only develop

after anthesis, suggesting that pubescence in the two

species may have been independently derived. Other

similarities between the two species, notably their flex-

uous leaves with alternate primary segments, and their

strongly spurred flowers with rudimentary apical wings

on the outer petals, appear to be ancestral and thus not

indicative of a sister-species relationship.

History : the species was first collected sometime in

the early years of the nineteenth century by Martin Hein-

rich Lichtenstein, a German physician and naturalist who

accompanied the incumbent Governor of the Cape of Good

Hope, J.W. Janssens, as family physician on his arrival in

Cape Town at the end of 1802 (Gunn & Codd 1981). Over

the next three years, Lichtenstein made three journeys into

the interior of the country, all of which passed through the

Klein Roggeveld, where he almost certainly collected Tri-

gonocapnos lichtensteinii. Although Lichtenstein intended

to describe his collection under the name Fumaria micran-

tha, this manuscript name remained unpublished and the

species was only formally described more than two dec-

ades later, when it was named for him by the German bota-

nists Ludolf von Chamisso and Diederich von Schlechten-

dal (1826).

The next collection of the species was also made dur-

ing a journey up to the Roggeveld Escarpment, this time

by the English explorer William Burchell on 1 August

1811, who noted in his Travels in the interior of south-

ern Africa of 1822 (Burchell 1953: 174) that among

the 14 specimens collected that day ‘was a Fumitory so

exceedingly like an English species, as hardly to be dis-

tinguished from it’. This opinion was evidently shared

by Alphonse de Candolle (1824), who distinguished

Burchell’s plant only as a form of the European Fumaria

capreolata , basing his identification on the recurved

pedicels that characterize the latter species. Although

seen by de Candolle in Burchell’s herbarium, we have

been unable to locate this specimen but the description

of the cirrhose leaves and relatively long, recurved fruit-

ing pedicels (de Candolle 1824: 130) can only apply to

Trigonocapnos lichtensteini among the South African

taxa and this, combined with the locality, make its iden-

tification certain. The species remained poorly known

and Harvey (1894: 18) in his account of the family for

Flora capensis, following the advice of Sonder, included

it in the European species F. muralis, as yet another of

the multiplicity of ‘book species’ of this variable taxon

that so aggravated him.

Bothalia 39,1 (2009)

53

FIGURE 3. — Trigonocapnos lichtensteinii : A, portion of stem; B, flower, front and lateral views; C, lower outer petal; D, inner petals; E. gyn-

oecium, dorsal, lateral and partial ventral views, style removed; F, immature fruit, front view; G, mature fruit, front, lateral and three-quarter

views; H, seed. Scale bar: A, 10 mm; B-E, G, 1 mm; F, 0.5 mm; H, 0.25 mm. Artist: John Manning.

It is little wonder, therefore, that it was subsequently

re-described as a distinct genus by the German botanist

Rudolph Schlechter (1899) from a collection that he had

made a few years earlier in the Pakhuis Mountains. This

is the name under which the species remained known

until Liden (1986) recognized that Chamisso and Sch-

lechtendal’s name took priority and accordingly effected

its transfer to Trigonocapnos. The species is now rela-

tively well known from the Bokkeveld and Pakhuis

Mountains but has only been sporadically collected from

the Roggeveld and Klein Roggeveld Escarpments.

3. Cysticapnos Mill., The gardener’s dictionary

(1754); Harv.: 16 (1894); Hutch.: 110 (1921); Fedde:

286 (1924); Liden: 105 (1986), 316 (1993). Type spe-

cies: C. vesicaria (L.) Fedde.

Phacocapnos Bemh.: 664 (1838); Hutch.: 109 (1921).

Corydalis sect. Phacocapnos (Bemh.) Prantl in Prantl &

Ktindig: 144 (1889); Harv.: 16 (1894). Type species: P.

cracca (Cham. & Schltdl.) Bemh.

Brittle, semi-succulent, glabrous, climbing annuals

with slender taproot or rarely (C. pruinosa ) perennial

with fleshy taproot; stems quadrate, with obscure unicel-

lular papillae along angles; sap watery, clear or yellow.

Leaves alternate, pinnately or bipinnately compound,

primary divisions alternate or opposite; ultimate leaflets

elliptical or obovate-cuneate and deeply 3-lobed; termi-

nal leaflets of all or only upper leaves transformed into

tendrils. Inflorescence pedunculate or sessile, terminal

but leaf-opposed through rapid growth of axillary bud,

54

Bothalia 39,1 (2009)

racemose; bracts scale-like, petaloid, irregularly toothed.

Flowers zygomorphic, bilabiate, pink, unscented. Sepals

2, lateral, much shorter than petals, scale-like, peta-

loid, irregularly toothed. Petals 4 in 2 series, outer pet-

als larger, naviculate-spathulate, laterally expanded or

winged apically, upper saccate or short-spurred at base,

inner petals apically connate, clawed, limb inflated

with dorsal crest, claw basally adnate to upper petal.

Stamens 6, diadelphous in abaxial and adaxial bundles

of 3; filaments of each bundle fused, lanceolate, mem-

branous, clasping ovary, upper filament cluster adnate

to margins of dorsal petal basally to form small cham-

ber, with nectary basal or decurrent on dorsal petal into

spur, pulvinate. Ovary ovoid, sometimes dorsoventrally

compressed, glabrous; ovules several in 2-several series

on lateral placentas; style short, straight or geniculate;

stigma compressed, 2-lobed. Fruit pendulous, capsular,

papery, sometimes inflated and bladder-like, circumfer-

entially partially or completely dehiscent. Seeds several,

lenticular, glossy black, testa obscurely scalariform-col-

liculate or almost smooth.

Three spp., temperate South Africa, southern Namibia

and Lesotho, winter rainfall region and along eastern

escarpment to Mpumalanga.

Key to species

la Perennial with fleshy taproot; leaves finely divided, carrot-like;

racemes pedunculate, mostly 14— 35-flowered; inner petal

crests papillate; nectary adnate to lower half of spur; capsules

lanceolate-quadrate, transversely flexed 3.2 C. pruinosa

lb Annuals with slender taproot; leaves not finely divided,

Aquilegia- like, ultimate segments elliptic; racemes ses-

sile (lowermost flower basal), l-15(-20)-flowered; inner

petal crests smooth; nectary at base of staminal sheath, not

adnate to spur; capsules not transversely flexed:

2a Raceme 4— 1 5(-20)-flowered; ± 1 mm long, pedicels shorter

than bracts; outer petal wings small, ± 1 mm wide, not

extending apically; capsule flattened, lanceolate, 5-12

mm long; style straight; sap yellow 3.1 C. cracca

2b Raceme (l)2-7-flowered; pedicels 2-8 mm long, longer

than bracts; outer petal wings large, 2-4 mm wide, encir-

cling petal apices; capsule ovoid, usually inflated, 20-25

mm long, rarely dwarfed, 4-10 mm long; style sharply

sigmoid at base; sap clear 3.3 C. vesicaria

3.1. Cysticapnos cracca (Cham. & Schltdl.) Liden

in Opera Botanica 88: 108 (1986). Corydalis cracca

Cham. & Schltdl.: 567 (1826); Harv.: 17 (1894). Phaco-

capnos cracca (Cham. & Schltdl.) Bemh.: 664 (1838);

Hutch.: 110 (1921). Type: South Africa, [Western Cape],

Cape of Good Hope, without precise locality or date,

Bergius s.n. (S, holo.).

Phacocapnos dregeanus Bemh.: 664 (1838). Type: South Africa,

[Western Cape], Enon, White River, 11 November 1829, Drege 7586

(MO, holo.!; PRE, iso.!).

Corydalis laevigata E.Mey.: 95, 125 (1844), nom. nud.

Climbing annual up to 1 m, with yellow sap. Leaves

bipinnately compound, ultimate segments broadly obo-

vate, 2- or 3-lobed almost to base, lobes elliptical to

obovate, terminal segments transformed into tendrils.

Inflorescence sessile with lowest flower basal, up to 80

mm long in fruit, 4— 1 5(-20)-flowered; pedicels suberect

and slightly curved outwards, ± 1 mm long; bracts

recurved, lanceolate, 1. 5-2.0 x 0.5-0.75 mm, irregularly

toothed. Flowers pale pink with dark reddish tips. Sepals

ovate, 1.0-1. 5 x 0.5-0.75 mm, irregularly toothed. Pet-

als: outer petals naviculate-spathulate, 6.0-6. 5 x 1. 5-2.0

mm, apically winged, wings reflexed, ± 1 mm wide,

upper saccate at base, sac ± 1 mm deep; inner petals 5-6

mm long, claw ± 3 mm long, adnate to upper petal in

lower 1 mm, blade inflated, ellipsoid, 2-3 x 1 mm, with

fleshy dorsal crest ± 1 mm wide. Stamens: filament bun-

dles lanceolate, 4. 0-4. 5 x 1 mm, upper bundle adnate

to upper petal along margins in lower 3 mm, nectary

basal, pulvinate; anthers ± 0.25 mm long, yellow. Ovary

dorsoventrally flattened, lanceolate, 4x2 mm, green,

sparsely and obscurely papillate; style straight or slightly

curved, ± 1 mm long; ovules biseriate, ± 8 per placenta.

Fruit pendent, dorsoventrally compressed, lanceolate,

(5— )9— 12 x 2-3 mm, dehiscing from base to ± halfway,

remaining attached to replum (placenta) in distal half,

several-seeded. Seeds lenticular, 0.6-0. 8 mm diam.,

glossy black, obscurely scalariform-colliculate. 2n = ±

32 (Liden 1986). Flowering time: (August) September to

October. Figure 4.

Distribution and habitat: occurs among bushes in sea-

sonally moist, sheltered situations in coastal and inland

scrub up to ± 1 000 m, from the Bokkeveld Escarpment

through the southwestern and southern Cape and into

the Eastern Cape as far as Port Elizabeth, Port Alfred

and Grahamstown (Figure 5). The species usually grows

in fine-grained clay or loamy, granite-derived soils but

sometimes in neutral sands, often on south-facing slopes

that are shaded in the afternoon. On nutrient-poor sand-

stone soils plants have been recorded as favouring richer,

nitrified hyrax middens.

Diagnosis and relationships: Cysticapnos cracca is

recognized by its lanceolate, blade-like capsules, 10-12

x 2-3 mm, containing numerous seeds in two periph-

eral rows. The' pericarp dehisces from the base, with

the two halves peeling away but remaining attached in

the distal half. In general form and dehiscence they are

very similar to those of C. pruinosa, but the latter has

larger, ± quadrate capsules, 15-17 mm long, that are dis-

tinctly flexed upwards in the distal half, with obscurely

scalloped margins. The racemes in C. cracca are almost

invariably sessile, with flowers smaller than those of C.

pruinosa (6. 0-6. 5 mm vs 7-8 mm) and generally fewer

per raceme (mostly up to 15 vs more than 15). Vegeta-

tively C. cracca is readily distinguished from C. pru-

inosa by its annual habit, less finely divided leaves, and

yellow vs clear sap.

Cysticapnos cracca is most likely to be confused

with C. vesicaria, with which it co-occurs in parts of

the southwestern Cape. Although typically smaller and

more compact than C. vesicaria , C. cracca is otherwise

almost indistinguishable in foliage, both species hav-

ing broadly obovate, deeply 2- or 3-lobed ultimate leaf

segments. The sap of C. cracca, however, is distinctly

yellow, staining the skin bright orange, whereas that of

C. vesicaria is clear. The two species are readily distin-

guished in flower and fruit, C. cracca having smaller,

less prominently winged flowers and flattened, non-

inflated capsules. Although C. cracca is typically found

in heavier, clay soils, with C. vesicaria favouring deep

coastal sands and gritty granite-derived soils, the two

species have been recorded growing together in several

places throughout their area of sympatry.

Bothalia 39,1 (2009)

55

FIGURE 4. — Cysticapnos cracca : A, flower, front and lateral views; B, lower outer petal; C, inner petals; D, gynoecium, lateral view; E, mature

fruit, lateral view before dehiscence and dorsal view after dehiscence; F, seed. Scale bar: A-C, 10 mm; E, 2.5 mm; F, 0.4 mm. Artist: John

Manning.

History ; Cysticapnos cracca was first collected by

the German apothecary Carl Bergius, who arrived at the

Cape in 1815. He almost certainly gathered the speci-

mens in the immediate surrounds of Cape Town itself,

and as he died here a few years later, at the beginning

of 1818 (Gunn & Codd 1981), his collection must date

from this three-year period. The species was described

several years later in the genus Corydalis by the Ger-

man botanists Ludolf von Chamisso and Diederich von

Schlechtendal (1826) but was later transferred by Bem-

hardi (1838) to his new genus Phacocapnos, which he

distinguished from Corydalis primarily on the lack of

an aril or strophiole on the seed. This treatment was not

followed by Harvey (1894) but was adopted by later

botanists. At the same time Bemhardi (1838) described

a later collection from Enon, north of Port Elizabeth,

made by the German collector Johann Drege in 1828

as the new species P. dregeanus on the basis of its acute

rather than obtuse fruits. As subsequently concluded by

Harvey, this purported difference is not significant.

3.2. Cysticapnos pruinosa (E.Mey. ex Bernh.)

Liden in Opera Botanica 88: 106 (1986). Phacocapnos

pruinosa [as pruinosus ] E.Mey. ex Bernh.: 664 (1838);

Hutch.: 110 (1921). Corydalis pruinosa (E.Mey. ex

Bernh.) Harv.: 17 (1894). Type: South Africa, [Eastern

Cape], 3027 (Lady Grey): Witberg [Witteberge], (-DA),

4000-5000' [1 200—1 500 m], November without year

[1832], Drege 3846 133 (I, a) (MO, holo.!; B!, BOL!,

PRE!, S, iso.).

Tufted perennial with fleshy taproot, and climbing

stems up to 1 m, with clear sap. Leaves bipinnately com-

pound, ultimate segments broadly obovate, 2- or 3-lobed

almost to base; terminal segments of upper leaves trans-

formed into tendrils. Inflorescence pedunculate, up to

160 mm long in fruit, 14— 3 5 -flowered, sometimes bear-

ing 1 or 2 reduced leaves below; pedicels suberect and

slightly curved outwards, 1-2 mm long; bracts recurved,

lanceolate, 1. 5-2.0 x 0.5-0.75 mm, irregularly toothed.

Flowers pale pink with dark reddish green gibbae.

Sepals ovate, 2 x 1.5 mm, irregularly toothed, basally

auriculate. Petals : outer petals naviculate-spathulate,

prominently clawed, 7-8 x 0. 8-1.0 mm, apically winged

and 3-4 mm wide, wings patent, ±1.5 mm wide, upper

saccate at base, sac 3^4 mm deep; inner petals 5-6 mm

long, claw ± 2 mm long, adnate to upper petal in lower 1

mm, blade inflated, ellipsoid, 3^4 x 1.5 mm, with papil-

late dorsal crest ± 1 mm wide. Stamens : filament bundles

lanceolate, 3. 5^4.0 x 1.2 mm, upper bundle adnate to

J*

16° 18° 20° 22° 24° 26°

FIGURE 5. — Known distribution of Cysticapnos cracca.

56

Bothalia 39,1 (2009)

upper petal along margins in lower 1 mm, nectary elon-

gate-pulvinate, decurrent up lower half of spur; anthers

± 0.25 mm long, yellow. Ovary dorsoventrally flattened,

lanceolate-quadrate, 4 x 1.2 mm, green; style straight,

suberect, ± 0.7 mm long; ovules biseriate, ± 9 per pla-

centa. Fruit pendent, dorsoventrally compressed, lan-

ceolate-quadrate, (1 2-) 15-25 x 6-7 mm, flexed upwards

in distal half, margins obscurely scalloped-dentate at

maturity, dehiscing from base to ± halfway, remaining

attached to replum (placenta) in distal half, many-seeded.

Seeds lenticular, 1.0-1 .5 mm diam., glossy black, scalari-

form-colliculate peripherally. 2n = ± 32 (Liden 1986).

Flowering time : December to February. Figure 6.

Distribution and ecology, a montane and subalpine

species, distributed along the eastern escarpment of South

Africa between 1 200 and 3 000 m, from the Koueveld-

berge west of Graaff-Reinet northwards through the

Drakensberg Mountains of Eastern Cape and KwaZulu-

Natal to Harrismith and westwards across the higher-

lying parts of Lesotho to the Leribe Plateau (Figure 7),

with isolated northern outliers along the eastern escarp-

ment in Mpumalanga around Wakkerstroom and Ermelo

and on the Mauchsberg near Lydenburg. Cysticapnos

pruinosa typically grows in gritty, basaltic soils, scram-

bling among bushes and grasses on mountain slopes and

in scree, in rocky boulder beds in drainage lines, or along

mountain streams. It is the only member of the tribe Dis-

cocapninae to occur in the summer rainfall region.

The species has previously been regarded as an annual

but excavation of several individuals confirms that it is

perennial. Seedlings form the characteristic fleshy tap-

root within their first year but appear to flower only from

the second season. Older plants develop a taproot sev-

eral millimetres in diameter, and produce a crown with

multiple growing points, from which new shoots arise.

These, like the seedlings, seem not to flower in their first

season, accumulating old leaf bases along the subterra-

nean portion. Plants die down in winter to the crown and

resprout in the spring.

The species is used medicinally by the Sotho, among

whom it is known colloquially as Musa pelo oa noka

(the river comforter), literally ‘the one who puts the

heart (of the river) right’. It is taken as a relief for sor-

row, especially in times of bereavement ( Dieterlen 873).

Diagnosis and relationships'. Cysticapnos pruinosa

is distinguished by its perennial habit with a fleshy tap-

root, and by its finely divided, almost carrot-like leaves.

Other species of Cysticapnos are annuals with a slender,

almost fibrous taproot and broader, ± elliptical ultimate

leaflet segments. The elongate, floriferous racemes of C.

pruinosa produce up to 35 relatively large flowers, 7-8

mm long, with papillate rather than smooth dorsal crests

on the lateral petals, and lanceolate-quadrate fruits with

a distinctive transverse flexure. The clear, rather than

yellow, sap is shared with C. cracca and with species of

Fumaria.

History: one of several South African Fumariaceae

collected by the German plant hunter Johan Drege, the

species is based on his collection from the mountains

around Lady Grey in Eastern Cape. Although the col-

lection is undated as to year, his itinerary indicates that

he was in this area in late 1832 and early 1833 (Gunn

& Codd 1981). Ernst Meyer, the German botanist who

dealt with Drege ’s collections, intended describing the

species in the genus Corydalis but his name remained

unpublished and it was left to Bemhardi to formally

describe the species in his new genus Phacocapnos,

which he distinguished from Corydalis by the lack of an

aril or strophiole on the seeds. The Bemhardi herbarium

formed the basis for the Missouri Botanical Garden her-

barium and the specimen there, which is marked as part

of the Bemhardi Herbarium, is thus appropriately recog-

nized as the holotype.

3.3. Cysticapnos vesicaria (L.) Fedde in Reper-

torium specierum novamm regni vegetabilis 19: 287

(1924); Liden: 106 (1986). Corydalis vesicaria (L.)

Pers.: 269 (1806). Fumaria vesicaria L.: 701 (1753).

Type: South Africa, without locality or date, LINN881.16

(LINN, holo.!).

C. cirrhosa Moench: 52 (1794), nom. illeg. superfl. a Fumaria vesi-

caria L.

Climbing annual up to 2 m, with clear sap. Leaves

bipinnately compound, ultimate segments broadly obo-

vate, 2- or 3-lobed almost to base, lobes narrowly ellip-

tical to obovate; terminal segments transformed into

tendrils. Inflorescence sessile (lowest flower basal), 10-

60(-80) mm long, (l)2-7-flowered; pedicels suberect,

curved outwards apically, 2-8 mm long, elongating to 25

mm in fruit; bracts spreading, obovate-cuneate, 2. 0-2. 5 x

1.0-1. 5 mm, apically toothed. Flowers pale pink, upper

petal with dark reddish mark at base of wing, inner pet-

als translucent. Sepals sagittate, 1— 2(— 3) x 0.5-1.0(-2.0)

mm, irregularly toothed. Petals: outer petals naviculate-

spathulate, 7— 9(— 1 3) x 2-3 mm, winged in outer half,

wings reflexed, 2^1(-5) mm wide, upper petal rounded

at base; ‘inner petals 6-7 mm long, claw 1. 5-2.0 mm

long, adnate to upper petal in lower 1 mm, blade ellip-

soid, inflated, 4.5-5 .0 x 1 mm, with inflated dorsal crest

1. 5-2.0 mm wide. Stamens: filament bundles broadly

lanceolate, 3. 5^4.0 xl.5 mm, upper bundle adnate to

upper petal along margins in lower 1 mm, nectary basal,

pulvinate; anthers ± 0.25 mm long, yellow. Ovary ovoid,

3 x 1.5 mm, green; style geniculate-sigmoid, ± 0.5 mm

long; ovules 4-seriate, 30-40 per placenta. Fruit pendu-

lous on elongated pedicel, heterocarpic, ovoid, usually

inflated and (10-)20-25 x (5-) 15-20 mm, sometimes

compressed and not inflated, then 4-10 x 3-5 mm, exo-

carp splitting longitudinally for entire length, mesocarp

usually aerenchymatous, endocarp remaining attached to

exocarp by radiating threads, rupturing irregularly, many-

seeded, rarely mesocarp not developing and fruits then

much smaller, 5-10 x 4—5 mm. Seeds lenticular, 1.0-1. 5

x 0.8-1. 3 mm diam,, excavated on faces, glossy black,

obscurely colliculate and almost smooth. 2n = 28, ± 30

(Liden 1986). Flowering time: mainly August to early

October but as early as June in the Little Karoo. Figure 8.

Distribution and ecology: common and widely dis-

tributed from southern Namibia through Namaqualand

and the southwestern Cape to the Little Karoo around

Oudtshoom but apparently absent from the Knersvlakte

(Figure 9). The species usually occurs in coarse-grained

soils, mainly coastal sands in fynbos and coastal scrub or

gritty, granitic soils in renosterveld.

Bothalia 39,1 (2009)

57

FIGURE 6. — Cysticapnos pruinosa : A, portion of flowering stem; B, base of plant; C, infructescence; D, flower, front and lateral views; E, section

through spur; F, lower outer petal; G, inner petals; FI, gynoecium, lateral and three-quarter views; I, seed. Scale bar: A-C, 10 mm; D-G, 1 mm;

H, 0.5 mm. Artist: John Manning.

58

Bothalia 39,1 (2009)

FIGURE 7. — Known distribution of Cysticapnos pruinosa.

Diagnosis and relationships', although similar to Cysti-

capnos cracca in foliage, C. vesicaria is distinguished

from this and other species by the basally geniculate style

and by its ± inflated, bladder-like fruit, mostly 20-25 x

15-20 mm. This unique fruit results from the develop-

ment of a thick, highly aerenchymatous, spongy mesocarp

between the papery exo- and endocarps. Even at flowering

the ovary wall is clearly 3 -layered. At maturity the exo-

carp splits longitudinally but remains attached to the endo-

carp by the radiating, thread-like vascular bundles, and the

seeds are released through irregular rupturing of the endo-

carp. In some fruits, however, the mesocarp fails to expand

and the capsules remain small and compressed, measuring

just 4-10 x 3-5 mm. The seeds in these small capsules are

quite normal and the same plant may produce both dwarf

and normal fruits. These flattened capsules are strikingly

similar to those of C. cracca and C. pruinosa. The clear,

watery sap is shared with C. pruinosa.

Heterocarpy in Cysticapnos vesicaria was first noticed

and reported by Fedde (1924), and later by Hilliard &

Burtt, who recorded that ‘small fruits, not inflated but

with ripe seeds, [are] sometimes present on [the] same

raceme [as normal inflated fruits]’ ( Hillard & Burtt

13025, NU). Dwarf, flattened fruits have been recorded

throughout the range of the species and are a regular fea-

ture of the species. They have no taxonomic significance

although they have caused confusion in the past. Cory-

dalis hurmannii Eckl. & Zeyh. ex Harv. was based on

plants with such dwarf fruits, and a cultivated plant that

produced a single dwarf fruit formed the basis of Cysti-

capnos parviflora Liden.

The species is typically few-flowered, with 1-4 flow-

ers borne on short racemes up to 45 mm long. The flow-

ers are mostly 7-8 mm long, with the upper and lower

petal wings 2-3 mm wide. Populations from the northern

part of the range, in Namaqualand and the Richtersveld,

however, have up to seven, mostly larger flowers borne

in racemes 20-80 mm long. In these plants the outer pet-

als may reach 10-12 mm long with apical crests 3-5 mm

wide. These northern forms are treated here as subsp.

namaquensis.

History: the first of the southern African Fumariaceae

to be named, Cysticapnos vesicaria was described in the

genus Fumaria by Carl Linnaeus (1753). The identity of

the original collector is not known but the species was

soon in cultivation in Europe and became known to later

botanists under the name Cysticapnos africana , based on

an illustration of the very distinctive fruit reproduced in

Gaertner’s (1791) De fructibus et seminibus plantarum.

It was only in the early decades of the twentieth cen-

tury that Fedde (1924) resuscitated the use of the earlier

name, C. vesicaria, in an article in which he first iden-

tified the existence of marked heterocarpy in the taxon.

This led him to suspect that Harvey’s (1894) Corydalis

burmannii was nothing more than a smaller-fruited form

of C. vesicaria, an opinion that has now been vindicated

by further observations of plants in the field. This small-

fruited form was still recognized by Liden (1986) under

the illegitimate name Cysticapnos grandiflora. The vari-

ous forms of C. vesicaria distinguished by Fedde (1924)

on the basis of the size of the leaf segments and flowers

represent nothing more than the normal variation in leaf

and floral development evident even within individuals

in the same population.

Plants thought to have originated from Alexandria in

Egypt were cultivated in Europe under the unpublished

name Cysticapnos alexandrina. Although thought by

Fedde (1924) to have been validated by Don, this is not

Jn fact the case as Don (1831) indicated quite clearly his

opinion that the plants in question were not distinct from

C. africana , and the name thus remains invalid (McNeil

et al. 2006: Art 34:1).

Key to subspecies

la Raceme 1 0—20(^45) mm long, 1 ^-flowered; outer petals 7—

9 mm long with wings 2-3 mm wide; plants from south-

western and southern Cape subsp. vesicaria

lb Raceme (20-)30-80 mm long, 3-7-flowered; outer pet-

als 9-13 mm long with wings 3-5 mm wide; plants from

Namaqualand and southern Namibia subsp. namaquensis

3.3a. subsp. vesicaria

Cysticapnos africana Gaertner, De fructibus et seminibus plantarum

2: 161 (1791); Harv.: 16 (1894); Hutch.: 110 (1921). Type: Gaertner:

161, t. 115(1791), icono.!

Corydalis burmanii Eckl. & Zeyh. ex Harv.: 17 (1894). Phaco-

capnos burmanii (Eckl. & Zeyh. ex Harv.) Hutch.: 110 (1921). Type:

South Africa, [Western Cape], Saldanha Bay and near Brackfontein,

Clanwilliam, Ecklon & Zeyher 23 (SAM, lecto.!, designated by Gold-

blatt & Manning (2000); B!, MO!, iso.).

Cysticapnos vesicaria forma brevilobus Fedde: 287 (1924), syn.

nov. Type: South Africa, [Western Cape], Stellenbosch and Swellen-

dam, without date, Ecklon 21 (B, lecto.!, here designated).

Cysticapnos vesicaria forma longilobus Fedde: 287 (1924), syn.

nov. Type: South Africa, [Western Cape], Brakdam, August 1897, R.

Schlechter 11131 (B, lecto.!, here designated; MO, iso.!).

Cysticapnos vesicaria forma latilobus Fedde ms. (without collector

or date, B).

Bothalia 39,1 (2009)

FIGURE 8. — Cysticapnos vesicaria subsp. vesicaria : A, portion of flowering stem; B, base of plant; C, flower, lateral view; D, lower outer petal; E,

inner petals; F, gynoecium, lateral view; G, detail of nectary; H, infructescence (nearside lower leaf segment omitted); I, half normal fruit; J,

dwarf fruit; K, seed. Scale bar: A, B, H-J, 10 mm; C-G, 1 mm; K, 0.5 mm. Artist: John Manning.

60

Bothalia 39,1 (2009)

Cysticapnos alexandrina Link & Otto, nom. nud.

Cysticapnos grandiflora E.Mey.: 95 (1844), nom. nud.

Cysticapnos grandiflora [Bemh.] sensu Liden: 106 (1986), nom.

nud. [cited as C. grandiflora Bernh. in Linnaea 12 (as ‘13’) by Liden:

664 (1986), but not mentioned therein].

Raceme 10-20(^45) mm long, l-3(4)-flowered. Outer

petals 7-9 mm long with wings 2-3 mm wide.

Distribution and ecology, widespread through the

southwestern and southern Cape, from the northern

Bokkeveld Plateau southwards along the west coast and

the inland mountains of the southwestern Cape as far

east as Still Bay on the coast and Oudtshoom in the Lit-

tle Karoo (Figure 9). Plants are most commonly found

in sandy soils in fynbos and thicket, especially along the

coast, but inland populations occur on finer-grained clay

or limestone soils in renosterveld.

The typical subspecies is characterized by short, 1-4-

flowered racemes and relatively small flowers, 8-10 mm

long, borne close to the stem and thus relatively incon-

spicuous.

3.3b. subsp. namaquensis J.C. Manning & Gold-

blatt, subsp. nov.

Cysticapnos parviflora Liden: 106 (1986), syn. nov. Type: South

Africa, Northern Cape, 3018 (Kamiesberg): Farm Kamagap, 10 km NE

of Bitterfontein on Kliprand road, (-CC), 10 September 1974, Norden-

stam & Lundgren 1803 (S, holo.!).

Racemus (20-)3(M10(-80) mm longus, 3-5(-7)-flora,

alis petali extemis 3-5 mm latis.

TYPE. — Northern Cape, 3018 (Kamiesberg): Lang-

kloof, renosterveld near river, 735 m, (-CA), 7 Septem-

ber 2006, Snijman 2081 (NBG, holo.; K, iso.).

Raceme (20-)3(M10(-80) mm long, 3-5(-7)-flowered.

Outer petals 9-13 mm long with wings 3-5 mm wide.

Distribution and ecology, widespread through the

higher-lying parts ofNamaqualand, from Nuwerus north-

wards into the Richtersveld, and into southern Namibia

on the Numaissspitze at the southern edge of the Huib

Hock Plateau (Figure 9). Plants grow in sandy or gritty

soils, often along seasonal washes and watercourses.

This subspecies is distinguished from subsp. vesicaria

by the mostly longer, more floriferous racemes bearing

3-7 flowers. Populations between Garies and Spring-

bok have especially large flowers, 10-15 mm long, and

constitute a particularly attractive form of the species in

which the racemes of showy flowers project conspicu-

ously above the stems that twine through the upper twigs

of supporting shrubs. The collections from southern

Namibia, in contrast, have the small flowers of subsp.

vesicaria but their longer, more floriferous racemes are

typical of subsp. namaquensis.

A cultivated plant grown in Europe in 1974 from seed

collected between Bitterfontein and Kliprand produced

unusually small flowers and a single dwarf fruit, and was

described as the new species Cysticapnos parviflora by

Liden but is evidently nothing more than a poorly devel-

oped individual of this subspecies. The wild collection

has the relatively many-flowered racemes and large flow-

FIGURE 9. — Known distribution of Cysticapnos vesicaria subsp. vesi-

caria, • ; and C. vesicaria subsp. namaquensis , O.

ers typical of subsp. namquensis with the petal wings up

to 7 mm wide, compared to 2 mm in the cultivated mate-

rial (Liden 1986). The seeds of these cultivated plants,

which measured 0.8 mm in diameter, were contrasted

against the larger seeds 1.3 mm in diameter of C. grandi-

flora (Liden 1986) but seed size is variable in C. vesicaria

and can range from 0.8-1. 5 mm in diameter within a sin-

gle collection (e.g. Buys 474, Rosch 547). This apparent

difference in seed size is thus not significant. No fur-

ther material matching the cultivated plants, from which

much of the description and illustrations were based, has

been collected despite the fact that the type locality is on

a well-travelled road through Namaqualand.

History, although mostly quite distinct from the

southern forms of the species in the wild, the larger-

flowered northern populations have not been recognized

taxonomically until now.

II. Subtribe Fumariinae

Scandent annuals (rarely perennial) with 2^1-pinnatc

leaves, lacking tendrils. Racemes pedunculate. Flow-

ers white to pink, or yellow, upper petal spurred. Style

deciduous; stigma with two papillae. Fruit a hard-walled

nutlet, one-seeded, indehiscent, usually with two apical

germination pores.

Four genera: Cryptocapnos, Fumaria, Fumariola and

Rupicapnos. Mainly Mediterranean to central Asia, also

North and East Africa, and the Himalayas.

4. Fumaria L., Species plantarum: 699 (1753).

Type species: Fumaria L.

Brittle, semisucculent, glabrous, erect or diffuse

annuals; stems quadrate, with obscure unicellular papil-

lae along angles; sap watery, clear. Leaves alternate,

2^1-pinnately compound, primary divisions alternate

or opposite; ultimate leaflets linear to obovate, without

tendrils but sometimes petiolule and rachis prehensile.

Inflorescence sessile or pedunculate, terminal but leaf-

Bothalia 39,1 (2009)

61

opposed through rapid growth of axillary bud, racemose,

pedicels often thickening in fruit; bracts scale-like, peta-

loid, irregularly toothed. Flowers zygomorphic, bilabi-

ate, pink, unscented. Sepals 2, lateral, much shorter than

petals, scale-like, petaloid, often irregularly toothed. Pet-

als 4 in 2 series; outer petals larger, naviculate-spathu-

late, laterally winged apically, upper spurred at base,

inner petals apically connate, clawed, limb inflated with

dorsal crest, claw basally adnate to upper petal. Stamens

6, diadelphous in abaxial and adaxial bundles of 3; fila-

ments of each bundle fused, lanceolate, membranous,

clasping ovary, upper filament cluster adnate to margins

of dorsal petal basally to form small chamber, with nec-

tary decurrent on dorsal petal into spur. Ovary subglo-

bose; ovules 1 or 2; style longer than ovary, ± slightly

upcurved or flexed upwards apically, deciduous; stigma

compressed, 2-lobed. Fruit ± erect or pendulous, subglo-

bose or obreniform, smooth or rugulose, indehiscent and

dropping entire, exocarp papery, endocarp hard and bony

with paired apical germination pores. Seed solitary, len-

ticular, thin-walled, without elaiosome.

± 50 spp., Mediterranean to India and east tropical

Africa, mainly North Africa and Spain.

Key to naturalized species

la Peduncle longer than raceme; pedicels recurved, 3-5 mm

long; sepals large, 4—6 mm long 4.1/1 capreolata

lb Peduncle shorter than raceme; pedicels stiffly suberect or

spreading, sometimes recurving slightly in fruit, 1.5-2. 5

mm; sepals small, 0. 5-3.0 mm long:

2a Leaves not as finely divided, parsley-like, ultimate seg-

ments oblanceolate-obovate (rarely ± linear); flowers

pink and purple, 7-10 mm long, including spur; bracts

1.0-1. 5 mm long, half to three-quarters as long as pedi-

cels 4.2 F. muralis

2b Leaves finely divided, almost dill-like, ultimate segments

linear; flowers white turning pink, and purple, ± 5 mm

long, including spur; bracts 2-3 mm long, as long as or

longer than pedicels 4.3 F. parviflora

4.1. Fumaria capreolata L., Species plantarum:

701 (1753); Sell: 256 (1964); Liden: 67 (1986); Walsh

& Norton: 407 (2007). Type: France, prope Olbyam Gal-

liae Narbonense ( UPS-LINN74560 , holo.).

Sprawling or scandent, branching annual up to 1 m.

Leaves bipinnately compound, rachis sometimes flexu-

ous, ultimate segments obovate, dissected almost to

base, lobes elliptical, apiculate-aristate. Inflorescence

pedunculate, up to 80 mm long in fruit; raceme shorter

than peduncle, 10-25(-35)-flowered; pedicels recurved,

3-5 mm long, apically thickened in fruit; bracts patent,

not clasping pedicel, linear-lanceolate, 2. 5^1.0 x 0.5-

0.8 mm. Flowers creamy white or pale pink with dark

reddish tips. Sepals peltate, ovate, 4—6 x 2.5 — 4.0 mm,

irregularly toothed. Petals : outer petals naviculate, upper

spathulate and apically short-winged, wings reflexed, ±

0.5 mm wide, spurred at base, spur compressed saccate,

2. 5-3. 5 mm long, lower linear with rudimentary apical

wings, 8-9 x 1.0-1. 5 mm; inner petals 7-9 mm long,

claw ± 2 mm long, adnate to upper petal in lower 1 mm,

blade inflated, narrowly oblong, 5-7 x 1 mm, with fleshy

dorsal crest ± 0.5 mm wide. Stamens : filament bundles

lanceolate-attenuate, 5-6 x 1 mm; anthers ± 0.25 mm

long, yellow. Ovary subglobose, 1.5 mm diam., green;

style slightly curved, 4—5 mm long, green, deciduous.

Fruit globose, 2. 0-2. 5 mm diam., indehiscent and drop-

FIGURE 10. — Known distribution in southern Africa of Fumaria capre-

olata and F. muralis, ® ; F. muralis, •; and F. parviflora, ■.

ping entire, brown, exocarp membranous, finely tubercu-

late or almost smooth, endocarp woody with small hour-

glass-shaped apical pit. Seed not seen. Flowering time :

September to October.

Distribution and ecology, native to the Mediter-

ranean and southwestern Europe but widely natural-

ized in warmer temperate countries, including southern

Australia. Known in South Africa from a single collec-

tion made near Somerset West in Western Cape in 1952

( Parker 4818) (Figure 10), it is so far at most an occa-

sional adventive here.

4.2. Fumaria muralis Sond. ex W.D.J.Koch, Syn-

opsis florae germanicae et helveticae, edn 2, part 3: 1017

(1845); Sell: 257 (1964); Liden: 78 (1986); Walsh &

Norton 408 (2007). Lectotype designated by Walsh: 496

(1992): Germany, Hamburg in muris, July 1844, Sonder

s.n. {MEL1 584466, lecto.).

4.2a. subsp. muralis

F. officinalis var. grandflora DC.: 135 (1824). Type: South Africa,

without precise locality or collector (G-BU, holo.).

F. officinalis var. capensis Harv.: 18 (1894), syn. nov. Type: South

Africa, [Western Cape], Witzenberg, without date, Ecklon & Zeyher 25

(TCD, holo.; PRE, SAM, iso.!).

Sprawling or scandent, branching annual up to 0.5

m, with clear sap. Leaves bipinnately compound; rachis

sometimes flexuous; ultimate segments obovate, dis-

sected almost to base, lobes elliptical (rarely ± linear),

apiculate-aristate, terminal segment rarely without a

blade. Inflorescence pedunculate, up to 30 mm long

in fruit, 3-12-flowered; pedicels suberect or spread-

ing, 2. 0-2. 5 mm long, thickened, firm and sometimes

slightly recurved in fruit; bracts erect, clasping pedicel,

1.0-1. 5 x 0.5 mm, irregularly toothed. Flowers pale to

deep pink with dark reddish tips. Sepals peltate, ovate,

2-3 x 1. 5-2.0 mm, irregularly toothed. Petals : outer pet-

als naviculate, 5-7 x 1.0-1. 5 mm, upper petal spathulate

and apically short-winged, wings reflexed, ± 0.5 mm

wide, spurred at base, spur compressed saccate, 2. 0-3. 5

mm long, lower petal linear with rudimentary apical

wings; inner petals 5-7 mm long, claw ± 1.5 mm long,

adnate to upper petal in lower 1 mm, blade inflated, nar-

62

Bothalia 39,1 (2009)

rowly oblong, 4-5 x 1 mm, with fleshy dorsal crest ± 0.5

mm wide. Stamens : filament bundles lanceolate-attenu-

ate, 5-6 x 1 mm; anthers ± 0.25 mm long, yellow. Ovary

subglobose, 1.5 mm diam., green; style ± straight or

slightly curved, 4-5 mm long, green, deciduous. Fruit

suberect, globose, 2. 0-2. 5 mm diam., indehiscent and

dropping entire, brown, exocarp membranous, finely

tuberculate or ± smooth, endocarp woody with small

hourglass-shaped apical pit. Seed depressed-subglobose,

1. 5-2.0 mm diam., testa membranous, containing scat-

tered, elongate orange deposits. Flowering time : mainly

August to October, later or earlier in gardens. Figure 11.

Distribution and ecology, native to Europe but intro-

duced elsewhere, including Australia and New Zea-

land. The species is widely distributed in South Africa

as a ruderal through the wetter, more temperate parts of

both winter and summer rainfall regions (Figure 10). It

is especially common in the southwestern and south-

ern Cape but has also been recorded from the higher-

lying Kamiesberg in central Namaqualand and into the

Eastern Cape, in KwaZulu-Natal between Durban and

Pietermaritzburg, and in Gauteng, with isolated collec-

tions from Kimberley, Bloemfontein and Phalaborwa.

It is found along the margins of cultivated lands and

in fallow fields, in waste places, and as a weed in gar-

dens. Fumaria muralis is essentially cleistogamous, the

anthers dehiscing before the petals separate, and the style

abscising from the ovary by an thesis.

The species appears to have been first collected in

South Africa during the latter part of the eighteenth cen-

tury by Carl Thunberg, who observed (Thunberg 1823)

that it had become widely established in the southwest-

ern Cape, both in gardens and outside of them. This very

early appearance of Fumaria muralis makes it one of the

first exotic plant species to naturalize in South Africa.

Other early collections from the southwestern Cape,

made during the first half of the eighteenth century,

include those by Christian Ecklon & Carl Zeyher ( Eck -

Ion & Zeyher 25) and by Johann Drege ( PRE24456 ).

Collections from the end of the century by Henry George

Flanagan and Alice Pegler record its occurrence in the

Eastern Cape, and it had reached central Namaqualand,

KwaZulu-Natal and Gauteng by the early decades of the

twentieth century.

It is almost certain that Fumaria muralis was an acci-

dental introduction to South Africa, probably among

wheat seed brought from Europe. Species of Fumaria

are common weeds of cereal crops in Australia and

wheat was already cultivated widely in the southwest-

ern Cape by the time that F. muralis was first collected

in the region by Thunberg (Thunberg 1823). This mode

of introduction would also explain its rapid spread

through the country. The alternative, that it was a delib-

erate introduction, is not supported by any evidence. The

small flowers of F. muralis make it useless as an orna-

mental and although the closely allied F. officinalis is

used medicinally in Europe (Launert 1981), the absence

of any mention of the species in Pappe’s early treatise on

South African medicinal plants (Pappe 1868) (and which

included other introductions) or in the later compendium

by Watt & Breyer-Brandwijk (1932), suggests that it was

not introduced for medicinal purposes.

Early collections of the species from southern Africa

were identified as forms of Fumaria officinalis L. (at that

time F. muralis had still to be described), and even until

recently F. muralis was often treated as a synonym of F.

officinalis. True F. officinalis is characterized by more

pronouncedly spathulate lower petals, and nutlets that

are broader than long, apically truncate or emarginate

and often obreniform in shape (Sell 1964; Liden 1986).

When pressed, green fruits of F. muralis may split open

at the apex and thus appear to be obreniform, and this

has resulted in some collections being misidentified as

F officinalis. Although an uncommon introduction in

Australia, F. officinalis does not yet appear to have been

recorded in South Africa.

Nomenclatural note : Harvey (1894) treated the South

African material as Fumaria officinalis var. capensis Harv.,

based on Ecklon & Zeyher 25, and although his citation

of the earlier name F. capreolata p. burchellii DC. (1824)

would render his name superfluous were de Candolle’s epi-

thet to be accepted at varietal level, we interpret de Can-

dolle’s hesitant citation of the rank of his name as F. capre-

olata p? burchellii to be sufficient grounds for considering

the name rankless and thus without priority at varietal rank

(McNeil et al. 2006: Art. 52.1).

4.3. Fumaria parviflora Lam., Encyclopedie metho-

dique. Botanique 2: 567 (1788); Sell: 258 (1964); Liden:

88 (1986); Walsh & Norton: 410 (2007). Type: In cult.

Paris, of Mediterranean origin, Lamarck (P-LAM, holo.).

4.3a. var. parviflora

Sprawling, branching annual up to 0.5 m. Leaves ter-

nately compound; primary divisions alternate or oppo-

site, ultimate segments finely dissected to base, lobes lin-

ear, apiculate-aristate. Inflorescence up to 30 mm long in

fruit, (7-)10-15(-22)-flowered; pedicels suberect, 1.5—

2.0 mm long, thickened and firm in fruit; bracts erect,

clasping pedicel, 2. 0-3.0 x 0.5 mm, irregularly toothed.

Flowers white, turning pink with age, with dark reddish

tips. Sepals cordate, broadly ovate, 0. 5-1.0 x 0.5-0. 7

mm, irregularly toothed. Petals', outer petals naviculate,

3^4 x ± 1 mm, upper petal spathulate and apically short-

winged, wings patent, ±0.5 mm wide, spurred at base,

spur compressed saccate, 1.0-1. 5 mm long, lower petal

linear with rudimentary apical wings; inner petals 3^1

mm long, claw ±1.5 mm long, adnate to upper petal in

lower 1 mm, blade inflated, narrowly oblong, ±2.5 x 1

mm, with fleshy dorsal crest ± 0.5 mm wide. Stamens :

filament bundles lanceolate-attenuate, 2. 5-3.0 x 1 mm;

anthers ±0.1 mm long, yellow. Ovary subglobose, ± 1

mm diam., green; style flexed sharply upwards apically,

± 2 mm long, green, deciduous; ovule 1, lateral. Fruit

suberect, globose, ± 2 mm diam, indehiscent and drop-

ping entire, brown, exocarp membranous, finely rugose-

tuberculate, endocarp woody with small hourglass-

shaped apical pit at stylar end. Seed not seen. Flowering

time'. August to February.

Distribution and ecology, native to the Mediterranean

but widely naturalized elsewhere, including Australia,

where it is scattered but uncommon. In South Africa the

species appears to be naturalized only in the drier west-

ern parts of the Northern Cape and Free State, where it

has been recorded from Hopetown and Kimberley (Fig-

Bothalia 39,1 (2009)

63

FIGURE 1 1 . — Fumaria muralis : A, portion of stem; B, detached lower leaf; C, flower, three-quarter and lateral views; D, lower outer petal; E, inner

petals; F, gynoecium, lateral view; G, detail of nectary; H, fruit; I, endocarp, front and dorsal views; J, seed. Scale bar: A, B, 10 mm; C-J, 1

mm. Artist: John Manning.

ure 10), typically along irrigation ditches or channels,

where it is regarded as a troublesome weed, forming

dense stands in wet places and choking the canals. The

species is evidently a recent introduction to the country,

and no records from earlier than the latter decades of the

twentieth century have been seen.

Fumaria parviflora, like F. muralis , is essentially

cleistogamous, the anthers dehiscing before the pet-

als separate, and the style abscising from the ovary by

anthesis.

OTHER SPECIMENS EXAMINED

The taxa are indicated in brackets by the number

assigned to them in the text, followed by the herbarium

acronym.

Abbot 4775, 5509 (3.2) MO, NH. Acocks 10477 (4.2) PRE; 14883,

16960, 17782 (2) PRE; 14950 (3.3b) PRE; 20433 (3.2) PRE; 20653,

21571, 22723 (3.1) PRE. Adamson 1936 (3.3a) BOL; 2103 (la) BOL.

Andreae 390 (3.3a) NBG. Axelson 477 (3.3a) NBG.

Baker 6 (4.2) PRE. Balkwill, Manning & Cadman 1145 (3.2) MO, NU,

PRE. Barker 1360 (3.3b) NBG; 5929 (3.3a) NBG. Baur ex Marloth

11457 (3.1) PRE. Bayer 227 (3.1) NBG; 240 (4.2) NBG. Bayliss 87

(3.2) MO. Behr 481 (4.2) PRE. Bester 2700 (3.2) NH; 3953 (3.2) PRE.

Bohnen 4762 (4.2) NBG, PRE. Bolus 1003 (3.1) BOL; 4697 (la) BOL,

NBG; 2702, 8052, 12596 (3.3a) BOL; 8920 (2) PRE. Bond 509 (3.3a)

NBG. Boot 132, 135 (4.2) PRE. Borchardt PRE463 (3.1) PRE. Bosen-

berg & Rutherford 75 (3.3a) NBG. Botes 397 (3.1) NBG. Boucher 528

(3.1) NBG, PRE; 6579 (3.3a) NBG. Braun 2224 (3.2) PRE. Bremer

281 (3.3a) PRE. Brooke 3 (3.3a) BOL. Brynard 255 (4.2) PRE. Burg-

ers 2114 (4.2) NBG; 2933 (3.3a) NBG. Burrows 6393 (3.2) PRE.

Burtt-Davy 10811 (4.2) PRE. Buys 474 (3.3a) NBG.

Cairns PRE54105 (4.2) NBG, PRE. Castelnau PRE24458 (4.2) P,

PRE. Cloete & Haselau 283 (2) NBG; 285 (3.3a) NBG. Coleman 312

(3.2) PRE. Compton 2688, 24323 (2) NBG; 3508 (3.1) BOL; 6808,

64

Bothalia 39,1 (2009)

17216 (3.3b) NBG; 21516 (3.2) NBG, PRE; 22296 (la) NBG. Crosby

660 (3.3b) PRE. Cruz 54 (3.3b) NBG; 126 (4.2) NBG.

Dahlstrand 2112 (3.3a) MO, NBG, PRE; 2453 (4.2) PRE. De la Bats.n.

(3.1) NBG. De Vos 1461 (3.1) NBG. Devenish 513, 1203 (3.2) PRE.

Dieterlen 873 (3.2) PRE, SAM. Dobay 65 (3.1) NBG; 66 (3.3a) NBG.

Dive 104 (3.2) PRE. Dr 'ege 3847 (3.1 ) BOL; PRE24456 (4.2) P, PRE.

Ecklon 25 (4.2) PRE. Eliovson 35 (3.3b) PRE. Esterhuysen 231, 11226

(la) PRE; 564, 11227 (3.1) PRE; 5539 (2) BOL; 5825 (3.1) NBG;

11848 (la) BOL, PRE. Evrard 8934 (3.3b) PRE.

Fabian 1244 (3.2) PRE. Fellingham 236 (4.2) NBG, PRE. Flanagan

806 (4.2) PRE, SAM. Fourcade 398 (4.2) NBG; 506 (3.1) BOL, MO;

904 (lb) BOL. Fugler 101 (4.2) NBG, PRE.

Galpin 1803 (3.2) BOL; 6569 (3.2) BOL, NH, SAM; 8229 (4.2) PRE.

Giess 13070 (3.3b) PRE. Giffen 710 (4.2) PRE. Gill 64 (4.2) PRE. Gil-

lett 547, 1080 (3.1) NBG; 3380 (la) NBG, PRE. Glass 21 (4.2) NBG.

Goldblatt 1980 (3.1) MO; 2334 (3.3a) MO, PRE; 5793 (2) PRE. Gold-

blatt & Manning 10520, 10554 (2) NBG; 10577 (2) BOL, MO, PRE;

13016 (3.1) MO, NBG, PRE. Goldblatt & Porter 12396 (3.3a) NBG;

12421 (2) MO, NBG. Greene 1091 (3.2) NH. Grobler 38 (3.3b) PRE.

Gubb KMG11151 (4.3) PRE. Guillarmod 597 , 1079 (3.2) PRE. Guthrie

2538 (3.1) NBG.

Hahndiek 15 (3.3a) NBG. Hall 4464 (2) NBG, PRE; s.n. (3.3b) NBG.

Hanekom 688 (4.2) NBG, PRE; 1189 (3.3a) NBG, PRE. Harris 30

(4.2) PRE. Harvey s.n. (la) PRE, TCD. Haynes 1110 (la) NBG, PRE;

1110A (3.1) PRE. Heinecken 103 (4.2) PRE. Helme 2615 (3.2) NBG.

Herre s.n. (3.3b) NBG. Heyns s.n. (3.3a) NBG. Hilliard & Burtt 6670

(3.2) MO, NU; 8869, 12192 (3.2) NU; 13025 (3.3a) NU, PRE; 14968

(3.2) NU, PRE. Hilton-Taylor 1185 (3.3b) NBG. Hugo 442 (3.3a)

NBG; 542, 2432, 2959 (3.3a) NBG, PRE. Hutchinson 263 (3.3a) BOL,

PRE; 780 (2) K, PRE.

Jacobsz 2013 (3.2) NBG.

Kemper 1PC621 (3.1) NBG. Killick 2277, 4390 (3.2) PRE. Kinges

1761 (4.2) PRE. Kok & Pienaar 1214 (4.2) PRE. Kroon 38 (4.2) PRE.

Kruger 513 (4.2) NBG, PRE.

Le Roux 2426 (3.3a) PRE; 2483 (3.3a) NBG, PRE; Le Roux & Ramsey

365 (3.3a) NBG, PRE. Leipoldt 20752 (2) BOL. Levyns 3217, 11320

(3.3a) BOL; 3453, 8511 (1.1a) BOL; 4065 (3.3b) BOL; 50572 (3.3a)

BOL, PRE. Lewis 1905, 1906 (3.3b) SAM; 19829 (3.1) BOL. Linder

2196 (3.3a) BOL. Loubser 3457 (3.1) NBG. Low 925 (4.2) NBG, PRE;

976 (4.2) NBG; 927a (3.3a) NBG, PRE; 2799 (3.3a) NBG.

MacOwan 1247 (3.1) BOL; 1713 (3.2) SAM; SAM14025 (3.1) SAM.

Maguire 288 (3.3b) NBG; 1887 (2) NBG; 1916 (3.3a) NBG. Man-

ning 3017 (4.2) NBG. Manning, Hilliard & Burtt 16020 (3.2) E, NU.

Marais 1352 (3.2) BOL, PRE. Marloth 237 (3.3a) PRE; 570 (4.2) PRE;

3612 (la) PRE; 6773 (3.3b) NBG, PRE; 12448 (3.3b) PRE. Marsh 370

(3.3a) NBG, PRE; 399 (3.1) NBG, PRE. Matthews 901 (3.2) NBG.

Mauve 2861 (4.2) PRE; 4544 (3.3a) PRE. Medley-Wood 4597, 5237

(3.2) NH. Meyer 3735 (4.2) PRE. Moffet 71 (4.2) NBG, PRE; 191

(3.3a) NBG, PRE. Mogg 1055, 11688 (4.2) PRE. Montgomery 48 (3.1)

NBG. Moriarty 723 (3.3a) NBG. Muir 5 (3.3a) PRE; 6 (4.2) PRE;

2921 (3.1) PRE.

Nelson 11622 (4.2) PRE. Newnham Brothers PRE54096 (4.2) PRE.

Nicolson 2102 (4.2) PRE.

O’Callaghan, Fellingham & Van Wyk 2 (4.2) NBG, PRE. Oliver 3716

(4.2) NBG, PRE; 4350 (4.2) NBG. Oliver, Tolken & Venter 475, 643

(3.3b) NBG, PRE; Olivier 90 (3.3a) NBG, PRE; 781 (4.2) NBG.

Parker 4235 (4.2) NBG; 4818 (4.1) NBG. Pearson 172, 6603 (4.2) NBG.

Pegler 519 (4.2) PRE. Perry & Snijman 2289 (3.3a) NBG, PRE; 2290 (2)

NBG. Phillips 1390 (4.2) SAM; 7616 (3.1) SAM; s.n. (3.3a) NBG. Phil-

lipson 717, 1449 (3.2) PRE; 717 (3.2) MO. Phillipson & Hutchings 167

(3.2) K, MO, PRE. Pienaar 1155, 1166 (3.3b) PRE. Pillans 2859, 6921

(3.1) BOL. Potts 5133 (3.2) BOL, PRE; 2817 (4.2) PRE. Pretorius 270

(2) NBG. Purcell SAM89465 (4.2) SAM.

Rennie s.n. (3.2) NU. Roberts 3439 (3.2) PRE. Retief 651 (4.2) PRE.

Rodin 1475 (3.3b) MO, PRE. Rogers 3484 (3 .1) MO . Rosch 547 (3.3a)

NBG; 644 (2) NBG. Rosch & le Roux 696 (3.3b) PRE; 1452 (4.2)

PRE. Rourke 577 (3.3a) MO, NBG; 1114 (3.3a) MO, NBG, PRE. Roux

Grootfontein 132 (4.3) PRE. Rubin 420 (4.2) PRE.

Schelpe 239 (3.3b) BOL; 506 (3.2) NU, PRE; 1443 (3.2) NU. Schlech-

ter 4596 (3.1) PRE; 4941 (3.3a) NBG, PRE; 10863 (2) MO; 10869 (2)

BOL, PRE; 11060 (3.3b) BOL, MO; 11131 (3.3b) PRE. Schmidt 383

(3.3a) PRE. Schmitz 8352 (4.2) PRE; 9312 (3.2) PRE. Schonland 3636

(4.2) PRE. Schweickerdt 2569 (3.3b) PRE. Shearing 1295 (4.2) PRE.

Sidey 2242 (3.1) MO. Sikhakhane & Williams 390 (3.2) NH, PRE.

Smith 3027 (3.3a) PRE. Smuts s.n. PRE59122 (3.3a) PRE. Staples

217 { 3.2) PRE. Steiner 529 (3.2) MO, NBG; 748 (2) NBG; 768 (3.3a)

NBG; 861 (3.2) NBG; 921, 3657 (3.3b) NBG. Stewart 1903 (3.2) MO,

NU. Steyn 14, 95 (4.2) PRE; 572 (3.3a) NBG; 592 (2) PRE. Strey 2823

(3.3a) PRE; 11013, 3909 (4.2) PRE. Stokoe SAM67430 (3.3a) SAM.

Strid & Strid 37430, 38077 (3.3a) PRE. Symons 14553 (3.2) PRE.

Taylor 1553 (3.1) SAM; 2718 (3.3a) NBG; 3951, 8150 (4.2) NBG;

4103, 11827 (3.1) NBG; 4104, 7411 (3.3a) NBG, PRE; 7410, 7421

(3.1) NBG, PRE; 11788 (2) NBG, PRE; 11801 (3.3a) MO, NBG,

PRE. Theron 1089 (4.2) PRE. Thode 5441, 5442, 6132 (3.1) NBG;

6133, 6134 (1.1a) NBG; 8267 (3.2) NBG; 9281 (3.3a) NBG; A473,

A1140 (3.2) PRE; s.n./A738 (1.1b) BOL, NH, PRE; A1878 (3.1) PRE;

A1958 (3.3a) NBG, PRE; A1959 (4.2) NBG, PRE; STE5670 (3.2) NH.

Thompson 394, 1034 (3.3b) NBG, PRE; 1875 (3.1) NBG. Thompson

& Le Roux 49 (3.3b) MO, NBG, PRE. Thorne SAM48867 (3.3b) SAM.

Trauseld 365 (3.2) NU. Tyson 498, 1728 (3.2) SAM; 640 (3.3a) NBG,

PRE, SAM.

Van Breda 43, 80 (3.3a) PRE. Van der Merwe 866 (4.2) PRE; 1746

(3.3a) NBG; 2557 (3.3a) PRE. Van der Walt 157 (3.3b) NBG, PRE;

s.n. (3.3a) NBG. Van Jaarveld 1389 (3.3b) PRE. Van Niekerk 814 (3.1)

BOL. Van Rooyen 2254 (3.3a) PRE. Van Wyk 717 (4.2) NBG, PRE;

1244 (3.3a) NBG, PRE; 1937 (3.1) NBG, PRE; 1937A (3.3a) NBG;

6305 (3.3b) PRE. Van Wyk & Abbott 12058 (3.2) PRE. Van Wyk, Win-

ter & Tilney 3458 (3.1) PRE. Van Zyl 3122 (3.3a) NBG, PRE. Venter

9537 (3.3b) PRE. Verdoorn s.n. PRE54147 (3.1) PRE. Victor 211 (4.2)

PRE. Viviers 602 (3.3a) NBG, PRE.

Walgate 295 (3.3a) NBG. Walters 2004 (4.2) NBG. Ward 606 (4.2)

PRE. West 630 (3.2) PRE. Wiese 6 (4.2) NBG. Williams 3320 (4.2)

NBG. Williamson 3562, 3916 (3.3b) NBG. Wilman 1475 (3.3b) BOL.

Winkler 92 (4.2) NBG. Wolley Dod (3.1) BOL; 114 (1.1a) BOL. Wright

1798 (3.2) NU.

Zantovska 117 (4.2) PRE. Zeitsman & Zeitsman 1003 (3.3b) PRE; 1101

(2) PRE?; 1458 (3.2) PRE. Zeyher 4936 (3.1) BOL, SAM; PRE54116

(4.2) PRE; SAM14029 (3.3a) SAM. Zeyher, Preiss & Krauss

SAM14028 (3.3a) SAM.

ACKNOWLEDGEMENTS

We are grateful to the curators and staff of the various

herbaria consulted and especially those of the Berlin and

Stockholm herbaria for making loans of the types in their

collections available to us for study. We also thank Roy

Gereau and Nick Turland for their help on nomenclature.

REFERENCES

ANGIOSPERM PHYLOGENY GROUP (APG). 2003. An update of

the Angiosperm Phylogeny Group classification for the orders

and families of flowering plants: APG II. Botanical Journal of

the Linnean Society 141: 399—436.

BERNHARDI, J.J. 1838. Nachtragliche Bemerkungen iiber Papavera-

ceen und Fumariaceen. Linnaea 12: 651-668.

BREMER, K„ BREMER, B. & THULIN, M. 2003. Introduction to phy-

logeny and systematics of flowering plants. Symbolae Botanicae

Upsalienses 33: 2. Department of Systematic Botany, Uppsala.

BRUMMITT, R.K. & POWELL, C.E. 1992. Authors of plant names.

Royal Botanic Gardens, Kew.

BURCHELL, W. J. 1 822. Travels in the interior of southern Africa. 2 vols.

MacLebose, Glasgow. Reprinted 1953 by Batchworth Press, Lon-

don.

CHAMISSO, L.A. VON & SCHECHTENDAL, D.F.L. 1826. Fumar-

iaceae. Linnaea 1: 556-570.

DE CANDOLLE, A.-P. 1 824. Fumariaceae. Prodromus systematis regni

vegetabilis 1: 125-130. Treutel & Wiirtz, Paris.

DON, G. 1831.4 general system of gardening and botany, vol. 1 . Riv-

ington, London.

FEDDE, F.K.G. 1924. Cysticapnos vesicarius (L.) heterokarp. Reperto-

rium specierum novarum regni vegetabilis 19: 285-288.

FOREST, F. & MANNING, J.C. in prep. A molecular phylogeny of the

southern African Fumariaceae.

Bothalia 39,1 (2009)

65

FUKUHARA, T. 1995. Vascular pattern in the fruit of Trigonocapnos

and Discocapnos (Papaveraceae-Fumaroideae). International

Journal of Plant Science 156: 547-554.

FUKUHARA, T. & LIDEN, M. 1995a. Seed-coat anatomy in Fumari-

caeae-Fumaroideae. Botanical Journal of the Linnean Society

119: 323-365.

FUKUHARA, T. & LIDEN, M. 1995b. Pericarp anatomy in Fumari-

caeae-Fumaroideae. Botanische Jahrbiicher 117: 499-530.

GAERTNER, J. 1791. De fructibus et seminibus plantarum, vol 2.

Schramm, Tubingen.

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

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

Institute, Cape Town & Missouri Botanical Garden, St Loius.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

HARVEY, W.H. 1894. Fumariacae. In W.H. Harvey & O.W. Sonder,

Flora capensis 1: 15-19. Reeve, Kent.

HOLMGREN, P.K., HOLMGREN, N.H. & BARTLETT, L.C. 1990.

Index herbariorum , edn 8. Regnum vegetabile 120. New York

Botanical Gardens, Bronx, New York.

HUTCHINSON, J. 1921. The genera of Fumariaceae and their distribu-

tion. Bulletin of Miscellaneous Information, Royal Botanic Gar-

dens, Kew 1921: 97-115.

KOCH, W.D. 1845. Synopsis florae germanicae et helveticae, edn 2,

part 3. Gebhardt & Resiland, Leipzig.

LAMARCK, J.B.A.P.M. DE. 1788. Encyclopedic methodique. Bota-

nique 2. Panckoucke, Paris.

LAUNERT, E. 1981. Edible and medicinal plants of Britain and north-

ern Europe. Hamlyn, London.

LIDEN, M. 1986. Synopsis of Fumarioideae (Papaveraceae) with a

monograph of the tribe Fumarieae. Opera Botanica 88: 1-133.

LIDEN, M. 1993. Fumariaceae. In K. Kubitzki, J.G. Rohwer & V. Bit-

trich. The families and genera of vascular plants, vol. II. Flower-

ing plants-dicotyledons: 310-318. Springer, Berlin.

LINNAEUS, C. 1753. Species plantarum. Salvius, Stockholm.

LUCAS, G.L. 1962. Fumariaceae. In C.E. Hubbard & E. Milne-Red-

head. Flora of tropical East Africa. Crown Agents for Oversea

Governments and Administrations, London.

McNEILL, J„ BARRIE, F.R., BURDET, H.M., DEMOULIN, V.,

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

PRADO, J., SILVA, P.C., SKOG, J.E., WIERSMA, I.H. & TUR-

LAND, N.J. 2006. International Code of Botanical Nomencla-

ture (Vienna Code) adopted by the 17th International Botanical

Congress, Vienna, Austria, July 2005. Regnum vegetabile 146.

Koeltz Scientific Books, Konigstein.

MEYER, E. 1844. In J.F. Drege, Zwei pflanzengeographische Doku-

mente. Leipzig.

MILLER, P. 1754. Cysticapnos. The gardener’s dictionary, abridge-

ment 4, vol. 3. Rivington, London.

MOENCH, C. 1794. Methodus. Facsimile edn. Koeningstein-Taunus.

PAPPE, L. 1868. Florae capensis medicae. Brittain, Cape Town.

PERSOON, C.H. 1806. Synopsis plantarum, vol. 2. Cramerum, Paris.

PRANTL, K. & KUNDIG, J. 1889. Papaveraceae. In A. Engler &

K. Prantl, Die naturlichen Pflanzenfamilien, vol. 3: 130-144.

Engelmann, Leipzig.

PRANTL, K. & KUNDIG, J. 1891. Papaveraceae. In A. Engler & K.

Prantl, Die naturlichen Pflanzenfamilien, vol. 3: 145. Engel-

mann, Leipzig.

SCHLECHTER, R. 1899. Plantae schlechterianae novae vel minus cog-

nitae describuntur II. Botanische Jahrbiicher 27 : 86-220.

SELL, P.D. 1964. Fumaria L. In T.G. Turin, V.H. Heywood, N.A. Burg-

es, D.H. Valentine, S.M. Walters & D.A. Webb, Flora europaea,

vol. 1: 255-258. Cambridge University Press.

THUNBERG, C.P. 1823. Flora capensis. Cottae, Stuttgart.

WALSH, N.G. 1992. Lectotypification of Fumaria muralis Sond. ex

Koch (Fumariaceae). Muelleria 7: 495, 496.

WALSH, N.G. & NORTON, W. 2007. Fumaria. Flora of Australia 2:

405-412.

WATT, J.M. & BREYER-BRANDWIJK, M.B. 1933. The medicinal and

poisonous plants of southern Africa. Livingstone, Edinburgh.

■

'

Bothalia 39, 1:67-72 (2009)

Two new species of Nemesia (Scrophulariaceae) from arid areas of

the Northern Cape, South Africa

K.E. STEINER*

Keywords: Namaqualand, Nemesia Vent., new species, Scrophulariaceae, South Africa, Tanqua Karroo

ABSTRACT

Two new annual species of Nemesia Vent, are described from southern Africa. Nemesia suaveolens is characterized by

magenta and yellow flowers. It differs from the closely related N. euryceras by having a lower lip that is yellow rather than

white with pale violet margins, an upper lip with a conspicuous yellow rectangular patch just above the corolla opening, a

spur that is ± equal to the length of the lower lip, not half the length, and a hypochile that is yellow rather than dark violet.

This new species is known only from the arid Tanqua Karoo east of the Cedarberg Mountains. N. aurantia is characterized

by orange saccate flowers with a brown and orange bearded palate. It is closest to N. versicolor, but differs from that species

by its orange corolla, the absence of a spur, and its bearded palate with brown and orange trichomes. It is known from a single

locality adjacent to the Swart Doring River in Namaqualand between Nuwerus and Garies.

INTRODUCTION

Nemesia Vent, is a genus of ± 62 species of annual

and perennial herbs endemic to southern Africa (Steiner

1994). It is over a hundred years since the last revision

of the genus (Hiem 1 904) and many new species have

come to light in the intervening years (Steiner 1989,

1994, 2006). A partial revision for species occurring

chiefly in KwaZulu-Natal was published by Hilliard &

Burtt (1986), but most of the new species that remain

to be described occur in the Cape Floristic Region and

southern Namibia, where ± 75 % of the species occur.

The purpose of this paper is to describe two new species

from arid areas in the Northern Cape Province of South

Africa. Descriptions are based primarily on living mate-

rial collected in the field and maintained in cultivation.

Nemesia suaveolens K.E. Steiner, sp. nov., N. eury-

cerate Schltr. proxima, sed differt lobo inferiore flavido

nec bianco et violaceomarginato, hypochilo corolla aureo

non atroviolaceo, macula rectangulari citrina conspicua

basi labio superioro.

TYPE. — Northern Cape, 3219 (Wuppertal): Tanqua

Karoo, 1.9 km N of entrance to Stompiesfontein (11.1

km N of Elandsvlei turnoff), S32° 14.392' E19° 41.463',

330 m, (-BA), 1 Sept. 2007, Steiner 4286 (NBG, holo.;

CAS, iso.).

Annual herb up to 320 mm tall, simple or branching

from base; stems glandular pilose, rectangular in cross

section, comers ridged, sides up to 1.8 mm wide, lat-

eral stems up to 160 mm long. Leaves simple, opposite,

mostly sessile to shortly petiolate; lamina lanceolate to

ovate or elliptical, 4—22 x 2-12 mm, sparsely glandular

pilose, apex rounded to acute; base rounded to cuneate;

margins entire to shallowly and sparsely dentate; peti-

oles 0-12 mm long, glandular pilose. Flowers axillary

or in lax, terminal racemes, pleasantly spicy scented;

racemes up to 270 mm long; bracts alternate, sessile,

* Department of Botany, California Academy of Sciences, 55 Music

Concourse Drive, Golden Gate Park, San Francisco 94118, USA.

MS. received: 2008-07-22.

cordate, reflexed, glandular pilose, lowermost leaf-like

up to 12.5 x 8 mm, uppermost reduced to ± 3 x 3 mm,

apex acute, base cordate; margins entire; pedicels 4—12

mm long, ascending, glandular pilose. Calyx lobes 5,

lanceolate to ovate, acute, spreading, densely glandular

pilose, central upper lobe lanceolate, 3. 0-3. 9 x 0.9-1. 2

mm, upper lateral lobes lanceolate, 2. 8-3. 7 x 0.8-1. 3

mm, lower lateral lobes ovate, 2. 8-3. 5 x 1.0-1. 6 mm.

Corolla bilabiate, 11.6-21.8 x 10.5-10.9 mm, upper

lip 4-lobed, 2 inner lobes oblong to obovate, 6. 4-6. 7

x 2.6-3. 1 mm, apices rounded to acute, bases strongly

oblique, 2 outer lobes spreading to revolute, oblong,

3. 1-5.1 x 3.2^4. 1 mm, apices rounded to emargin-

ate, bases strongly oblique; upper lip deep magenta at

base (greyish magenta reverse) surrounding a bright

golden yellow rectangular patch, 1.2-1 .6 x 1.7-1. 9 mm,

just above corolla opening, becoming greyish white

on lobes distally, lower lip widely obovate, 6. 0-8. 2 x

6-8 mm, apex emarginate, light yellow with yellowish

white reverse, base with a raised palate; palate convex,

oblong, cristate, 3. 1-4.6 x 4. 7-6.0 mm, divided by a

central groove into 2 raised yellow bosses, bosses gla-

brous or with minute trichomes distally, densely pilose

with conspicuous pale yellow or white trichomes basally

around corolla opening; hypochile (floor of corolla

tube) ± 3.9 — 4. 1 mm long, central boss low, densely

pilose, trichomes white to pale yellow; sides and upper

inside surface of corolla tube greyish with magenta to

purple lines, base of tube with a narrow spur, 3. 5-4. 8

x 0.7-0. 8 mm, pale yellow to whitish, deflexed and ±

straight or curving forward slightly in distal third, out-

side sparsely glandular pubescent, spur opening flanked

by 2 greenish yellow to orange patches. Stamens 4, usu-

ally white, purple flecked distally, lying in a shallow

depression (2. 5-2. 9 x 0. 8-1.0 mm) in upper inside sur-

face of corolla tube; filaments of anticous pair (twisted

into posticous position) 2.7-2. 8 mm long, sigmoid,

± straight in middle, glabrous or with a few glandular

trichomes below middle; posticous filaments ± straight

except at base, 0.8-1 .0 mm long, glandular pubescent;

anthers 0. 9-1.0 mm long, each pair strongly coherent.

Ovary oblong-ovoid, 1.2-1. 6 x 1.0-1. 6 mm, laterally

compressed; style oblong, ±0.8 mm long, compressed

68

Bothalia 39,1 (2009)

FIGURE 1 . Nemesia suaveolens, Steiner 4288 (CAS, NBG). A, habit; B, calyx. C-F, corolla: C, front view; D, side view; E, rear view; F, side

view partially cut away. G, stamens, anticous left, posticous right; FI, pistil; I, capsule; J, seed. Scale bars: A, 10 mm; B, 3 mm; C-F, 2 mm;

G, H, 1 mm; I, 2.5 mm; J, 1 mm. Artists: A-H, John Manning; I, Sarah Adler; and J, Nicole Bollinger.

Bothalia 39,1 (2009)

69

contrary to ovary, lying between anther pairs and curv-

ing slightly away from corolla opening; stigma cres-

cent-shaped, 0.1 x 0.5 mm. Capsules ovate to oblong

in outline, 3.9-10.6 x 3.6— 6.5 mm, laterally compressed

contrary to septum, apex emarginate to bilobed, lobes

rounded. Seeds winged, ovate to widely ovate, ± 1.5 x

1.0- 1.5 mm, light brown, verruculate, wing membranous

with numerous parallel, brownish veins. Flowering time:

(May-)July-September. Figures 1; 2A, B.

Diagnostic features : Nemesia suaveolens is easily

recognized by its yellow and magenta flowers with a ±

straight spur and a prominent yellow rectangular spot

(nectar guide) at the base of the upper corolla lip. It is

most closely related to N. euryceras, but differs from

that species by having a lower lip that is yellow rather

than white with pale violet margins, an upper lip with

a conspicuous yellow rectangular patch, just above the

corolla opening, a spur that is ± equal to the length of

the lower lip, not half the length of the lower lip, and

a hypochile that is yellow rather than dark violet. Sch-

lechter (1899) and Hiem (1904) describe the lower lip

of N. euryceras as sulphur yellow, but this is apparently

based on a single flower on the type specimen ( Schlech -

ter 8126 at K) that became yellowish from drying. The

other flowers on the same plant do not look yellowish,

but rather are consistent with the author’s observation at

the type locality {Steiner 3686 at NBG) of a lower lip

that is white with pale violet margins. Other collections

in PRE from the type locality and nearby areas also give

FIGURE 2. — Nemesia flowers: A, B, N. suaveolens: front and side views; C, D, N. aurantia: front and side views.

70

Bothalia 39,1 (2009)

FIGURE 3. — Known distribution ofN. suaveolens , •; and N. aurantia.

flower colour as white and purple, mauve and white, or

pink (e.g. Grant 4763; Hall 3726; Le Roux 2167).

Nemesia suaveolens often grows in close proxim-

ity to N. karroensis Bond and has been confused with

that species. It is similar in coloration to N. karroensis,

especially when both are pressed and dried, but differs

in having a short, straight, downwardly oriented spur

instead of a long (8.9-12.3 mm), backwardly project-

ing one that is usually strongly recurved in its distal half

(Bond 1940).

Etymology, the name refers to the pleasant spicy fra-

grance of the flowers.

Distribution and habitat. Nemesia suaveolens is

known from a very limited area of the central Tanqua

Karoo at elevations between 320 and 445 m (Figure 3).

The Tanqua is an arid desert plain situated in the rain

shadow of the Cedarberg Mountains that receives only

50 to 70 mm of rainfall in an average year, mostly fall-

ing in late autumn or early winter. N. suaveolens is an

annual that germinates in response to these rains, but in

years of no appreciable rainfall, plants remain dormant

as seeds.

Breeding systems : in cultivation, where insects are

excluded, Nemesia suaveolens does not set seed. This

suggests that, like many Nemesia species, this species is

self-incompatible. Although pollinators were not encoun-

tered on flowers in the field, the presence of capsules sug-

gests that plants were successfully pollinated. The spurs

of N. suaveolens do not secrete nectar, so pollinating

insects only obtain pollen as a reward.

Other specimens examined

NORTHERN CAPE. — 3219 (Wuppertal): Tanqua Karoo, Ceres-

Calvinia road (R355), 13.4 km N of turnoff to Elandsvlei, S32° 13.487'

El 9° 42.086’, ± 320 m, (-BA), 23 Aug. 2004, Steiner 4069 (CAS);

Tanqua Karoo, 13.4 km N of Elandsvlei turnoff (4.2 km N of entrance

to Stompiesfontein), S32° 13.279' E19° 42.195', 324 m, (-BA), Steiner

4288 (CAS, NBG); Tanqua Karoo, Ceres-Calvinia road (R355), 13.9

km N of turnoff to Elandsvlei, S32° 13.311' E19° 42.189', ± 320 m, (-

BA), 23 Aug. 2004, Steiner 4070 (CAS); Tanqua Karoo, Stompiesfon-

tein, (-BA), 26 July 1941, Bond 1181 (NBG); Tanqua National Park,

Bo Stompiesfontein and Varsfontein, S32° 12' 56" E19° 43' 02.1"), 368

m, (-BA), 25 July 2006, Rosch 415 (NBG); 4.8 km NNE of Kom-

mando Drift, ± 375 m, (-BC), 30 Aug. 1957, Acocks 19477 (PRE);

near Eendjies Kraal [Uintjieskraal] on road to Hottentots Kloof, (-BD),

28 Sept. 1929, Grant 4909 (PRE); Tanqua Karoo, 6.1 km S of Papkuil

on Ceres-Calvinia road, [± 442 m], (-DA), 17 May 1983, Snijman

1274 (NBG); Coega Kamma, 70 miles [112 km] from Ceres (-DA),

25 Aug. 1968, Stayner s.n. (NBG). 3220 (Sutherland): south of Tan-

qua National Park on Middlepos-Ceres road, after Platfontein turnoff,

Farm No. 10, 390 m, (-AC), 11 July 2006, Rosch 353 (NBG).

Nemesia aurantia K.E. Steiner, sp. nov., N. ver-

sicolori Drege proxima, sed differt floribus aurantis,

corolla saccato, non calcarato, palato barbato brunneo et

aurantio.

TYPE. — Northern Cape, 3018 (Kamiesberg); Farm

Stinkfontein, ± 160 m, (-CC), 20 Aug. 2001, Steiner

3640 (NBG, holo.; CAS, iso.).

Annual herb up to 410 mm tall, simple or branching;

stems glandular pilose, rectangular in cross section, cor-

ners ridged, sides up to 2 mm wide, lateral stems up to

265 mm long. Leaves simple, opposite, glabrous, apices

acute, bases cuneate to truncate, margins sparsely den-

tate; basal leaves ovate to elliptical, 10.8-18.4 x 3. 1-8.4

mm; petioles up to 7 mm long; upper leaves sessile,

lanceolate to linear, 9.0-39.6 x 1.3-9. 4 mm, glabrous.

Inflorescence terminal, racemose, up to 190 mm long,

or flowers axillary; bracts alternate, sessile, reflexed,

narrowly lanceolate to deltoid, 1.1 -2. 4 x 0.9 mm, gla-

brous above, sparsely glandular puberulent below, apex

acute, base truncate; margins entire; pedicels ± 8-24 mm

long, ascending, glandular pilose. Calyx lobes 5, spread-

ing, acute, densely glandular pilose, upper and lateral

lobes oblong, ± 2.5 x 0.8 mm, lower lobes shorter, lan-

ceolate, ± 2.1 x 0.8 mm. Corolla bilabiate, 13.1-18.8 x

1 1 .1-16.0 mm, upper lip 4-lobed, 2 inner lobes oblong to

obovate, 5. 1-6.2 x 1.9-3. 2 mm, apices rounded to acute,

bases strongly oblique, 2 outer lobes spreading, oblong,

4. 7-6. 5 x 3.4^1. 1 mm, apices emarginate, bases strongly

oblique; all lobes orange distally and pale yellow with

brownish streaks at base, lower lip widely obcordate,

9.2-12.3 x 10.5-16.0 mm, deep orange, basal portion

inflated into a dark brown, convex palate, 4. 1-8.0 x

4. 5-5. 2 mm long; bosses 2, oblong, divergent, orange,

0.5-1. 7 x 1.3-1. 4 mm, densely villous, trichome stalks

pale yellow or brown, translucent, simple or branched

below head, heads capitate or discoid; hypochile (floor

of corolla tube) ± 3.7-4. 1 x 4 mm, ± flat, streaked with

reddish brown, sparsely to densely pilose, especially

on inner lateral walls, outer surface glandular pilose;

upper inside surface of corolla tube pale yellow with

brown lines, base shallowly saccate, 0. 7-1.0 x 4. 7-4.9

mm, pale yellow, lower outside portion pilose. Stamens

4, lying in a shallow depression in upper inside surface

of corolla tube; filaments of anticous pair (twisted into

posticous position) ± 2.2 mm long, sigmoid, ± straight

in middle, glabrous; posticous filaments ± 1.5 mm long,

straight except at base, glabrous; anthers ± 1.1-1. 2 mm

long, each pair strongly coherent, opening downward.

Ovary oblong-ovoid, ± 1.0-1. 2 x 0. 9-1.0 mm, later-

Bothalia 39,1 (2009)

71

FIGURE 4. — Nemesia aurantia , Steiner 3640 (CAS, NBG). A, habit; B, calyx. C-F, corolla views: C, front, D, rear; E, corolla intact; F, corolla

partially cut away. G, H, stamens: G, anticous, H, posticous. I, pistil. J, K, seed: K, with outer coat and wing removed. L, mature capsule.

Scale bars: A, 20 mm: B, E, F, L, 2 mm; C, D, 5 mm; G, H, J, K, 0.5 mm; I, 1 mm. Artists: A, Inge Oliver^ C-K, Nicole Bollinger.

ally compressed; style ± 0.5 mm, deflected away from

corolla opening, oblong, truncate, lying between anther

pairs; stigma deltoid ± 0.3 x 0.5 mm. Capsules ovate

to oblong in outline, ± 3. 8-8. 3 x 4. 5-7.0 mm, com-

pressed contrary to septum, apex emarginate to bilobed,

lobes acute to rounded. Seeds winged, ovate to widely

ovate, 1. 7-2.4 x 1. 3-2.1 mm, outer layer of testa white,

lacey, verruculate, enveloping inner seed at maturity,

wing membranous with numerous parallel white veins,

internal portion of seed narrowly elliptical, dark brown,

1.0-1. 2 mm long, surface alveolate. Flowering time:

August-September. Figures 2C, D; 4.

72

Bothalia 39,1 (2009)

Diagnostic features : Nemesia aurantia is most closely

related to N. versicolor, but has orange rather than blue

or yellow flowers, lacks a conspicuous spur, and has a

bilobed palate on the lower lip that is less prominent.

The only other saccate Nemesia species with orange

flowers is N. strumosa Benth. Plants of N. aurantia are

typically shorter than N. strumosa (up to 410 mm vs up

to 730 mm) with smaller flowers (corolla limb 13-19

mm vs 17-35 mm long). The sac of the corolla in N.

aurantia is less than half the length, on average, of N.

strumosa and the raised palate and divergent bosses on

the lower lip of N. aurantia are absent in N. strumosa.

Etymology, the name refers to the bright orange flow-

ers.

Distribution and habitat'. Nemesia aurantia is known

only from the Northern Cape, just north of its boundary

with the Western Cape, ±31 km S of Garies. It occurs

with Arctotis fastuosa in a very localized area of loose

sand near the Swart Doring River on the Farm Stinkfon-

tein (Figure 2).

Conservation status', until other populations have

been located, this species should be considered rare.

The intensity of sheep grazing on the Farm Stinkfontein

is currently unknown, but may represent a threat to the

long-term survival of this population.

Pollination and breeding systems : nothing is known

about the pollination biology of Nemesia aurantia, but

because of its open saccate flowers, bright orange col-

our, and densely pubescent palate, it may be pollinated

by small monkey beetles. Based on the absence of cap-

sule formation in cultivation, N. aurantia is probably

self-incompatible.

Other specimens examined

NORTHERN CAPE. — 3018 (Kamiesberg): Farm Stinkfontein, 950

feet [± 300 m], (-CC), 21 Sept. 1929, Grant & Theiler 4778 (BOL, K).

Locality uncertain: Namaqualand, along road, Garies to O’Okiep, Aug.

1925, Marloth 6742 (NBG).

ACKNOWLEDGEMENTS

I thank Northern and Western Cape Nature Conserva-

tion authorities for permission to collect in their respec-

tive areas. I also thank John Manning, Inge Oliver1,

Nicole Bolinger, and Sarah Adler for their contributions

to the illustrations. I am grateful to Dee Snijman and the

Compton Herbarium staff for help with localities and

for the use of their facilities. I also thank SANBI for

Nemesia loans from NBG and PRE.

REFERENCES

BOND, P. 1940. Plantae Novae Afficanae, Nemesia karroensis. Journal

of South African Botany 6: 65-67 .

HIERN, W.P. 1904. Scrophulariaceae. In W.T. Thiselton-Dyer, Flora

capensis 4,2: 121^420. Reeve, London.

HILLIARD, O.M. & BURTT, B.L. 1986. Notes on some plants of

southern Africa chiefly from Natal: XIII. Notes from the Royal

Botanic Garden Edinburgh 43: 345 — 405 .

SCHLECHTER, R. 1899. Plantae Schlechterianae novae vel minus

cognitae describuntur. II. Botanische Jahrbucher 27: 175.

STEINER, K.E. 1989. A new perennial Nemesia (Scrophulariaceae)

from the western Cape. South African Journal of Botany 55:

405^408.

STEINER, K.E. 1994. A new Nemesia (Scrophulariaceae) from the

interior of the southern Cape, South Africa. South African Jour-

nal of Botany 60: 211-213.

STEINER, K.E. 2006. Two new species of Nemesia (Scrophulariaceae)

from southern Africa. Bothalia 36: 39^44.

Bothalia 39,1: 73-85 (2009)

Taxonomy and phylogeny of two subgroups of Pelargonium section

Otidia (Geraniaceae). 1. The Pelargonium carnosum complex

M. BECKER* and F. ALBERS*

Keywords: amplified fragment length polymorphism (AFLP), distribution, Geraniaceae, hybridization, morphology. Pelargonium carnosum

(L.)L'Her., phylogeny. South Africa, taxonomy, winter rainfall area

ABSTRACT

This contribution deals with the taxonomy and phylogeny of the Pelargonium carnosum complex, a group of closely

related taxa of Pelargonium L’Her. section Otidia (Sweet) DC. (Geraniaceae) that is distributed in the winter rainfall area

of South Africa. According to molecular analyses via AFLP, P. adriaanii M. Becker & F.Albers, P. carnosum (L.) L’Her., P.

ferulaceum (Cav.) Willd. and P. polycephalum (E.Mey. ex Harv.) R.Knuth form a monophyletic clade. Although hybridization

may occur between the taxa, three are assigned to specific rank. The fourth taxon, P. ferulaceum is recognized as a subspe-

cies of P. carnosum. As is implied from the occurrence of morphological intermediates and partly from molecular evidence,

hybridization does not only occur among the taxa in this complex but also involves species closely related to this group (P.

parvifiorum J.C.Wendl., P. laxum (Sweet) G.Don, P. dasyphyllum R.Knuth). For the taxa in the P. carnosum complex, distri-

bution areas are delineated and diagnostic features that have until now remained obscure, are outlined.

INTRODUCTION

The genus Pelargonium L’Her. comprises ± 280-290

species which are subdivided into 16 sections (Bakker et

al. 2004). The infrageneric classification relies on molecu-

lar evidence, on differences in chromosome sizes and basic

chromosome numbers, and on geographical distribution

and growth forms. Within the major group characterized by

small chromosomes, section Otidia (Sweet) DC. belongs to

a subgroup showing xerophytic growth. Within this ‘xero-

phytic clade’, it is part of the so-called winter rainfall clade

(Bakker et al. 2004). Section Otidia comprises 25 taxa,

some of them distinctive and isolated, others closely related

and hardly distinguishable.

The Pelargonium carnosum (L.) L’Her. complex

includes several taxa that are linked by intermediates.

Morphological intermediates are usually assigned to

introgression. The close relationship of these taxa was

recognized early in the taxonomic history of Pelargo-

nium. Harvey (1860), when describing P. polycephalum

(E.Mey. ex Harv.) R.Knuth as P. ferulaceum var. poly-

cephalum., stressed its strong resemblance to P. camo-

sum. Dyer (1953) reduced P. ferulaceum (Cav.) Willd.

and P. polycephalum to the rank of a variety under P.

carnosum, which can be taken as the year of inception

of the P. carnosum complex. Becker & Albers (2005a)

added P. adriaanii M. Becker & F.Albers, thus increasing

the number of taxa in this complex to four: P. adriaanii,

P. carnosum , P. ferulaceum and P. polycephalum.

Vorster (1990) placed the closely related Pelargonium

parvifiorum J.C.Wendl. in the P. carnosum complex as

well. However, as P. parvifiorum comprises several taxa

that are characterized by a distinctive floral structure, we

prefer to treat it as a separate taxon and with further taxa

as a separate subgroup {P. parvifiorum complex, Becker

& Albers in press a).

* Institut fur Botanik, Schlossgarten 3, D-48149 Munster, Germany.

E-mail: Prof. Dr F. Albers, albersf@uni-muenster.de; Dr M. Becker,

m.becker@massey.ac.nz.

MS. received: 2007-05-30.

A phylogenetic tree that results from extensive

molecular analyses is presented. Polymorphic mark-

ers have been detected via AFLP (amplified fragment

length polymorphism, Vos et al. 1995), a method that

requires no previous knowledge of DNA sequences

and provides a large amount of reliable and repeatable

bands. AFLP markers are generated from the entire spec-

trum of genomic DNA including fast evolving regions,

leading to a high resolution at the subspecies and even

population level in phylogenetic analyses. We discarded

sequence analyses based on nuclear ITS and plastid tm-

LF regions, as the differences between studied taxa were

too marginal.

MATERIAL AND METHODS

Plant material

Specimens of the living collection of Munster Botani-

cal Garden (Table 1) were included in the molecular

analysis. Voucher specimens were deposited in MSUN.

For delimiting the distribution ranges of the taxa, 224

herbarium specimens were examined from the following

herbaria: BM, BOL, K, MSUN, NBG and PRE— acro-

nyms as in Holmgren et al. (1990).

DNA extraction and AFLP analysis

Genomic DNA was extracted from ± 500 mg of

fresh leaf tissue per plant following the CTAB pro-

cedure described by Doyle & Doyle (1987) and modi-

fied by Bakker et al. (1998). AFLP (amplified fragment

length polymorphism) analyses were performed using

the protocol of Vos et al. (1995), with minor modifi-

cations (Marschalek 2003). DNA was restricted with

enzymes EcoRI (rarely cutting) and Msel (frequently

cutting). Single strands of EcoRI and Msel adapter

were 3’-CATCTGACGCATGGTTAA-5’, 5’-CTCGTA-

GACTGCGTACC-3 ’ and 3 ’ -TACTC AGGACTC AT-5 ’ ,

5’-GACGATGAGTCCTGAG-3’, respectively. Nine com-

binations of primers based on three selective bases (EcoRI -

AAC, -AGG, -ATA and Msel -CAA, -CAG, -CCG, -CGA,

74

Bothalia 39,1 (2009)

TABLE 1. — Localities, collectors’ names and numbers of specimens in Pelargonium carnosum complex and further species of sect. Otidia from

South Africa (RS A) and Namibia selected for AFLP analyses. Grid references given per quarter-degree square. STEU = Stellenbosch Uni-

versity Botanical Garden

-CTA) were chosen for the second selective PCR

amplification. EcoRI primer was fluorescence labeled

(IRDye™ 800 infrared dye, Licor). Electrophoresis of

AFLP fragments was done on 7 % (w/w) polyacrylamide

gels (250 x 0.2 mm) on a one-dye model 4200 Licor

DNA automatic sequencer.

Cladistic analysis

Each AFLP fragment was counted as a separate puta-

tive locus and scored as present (1) or absent (0) for

each sample. Only polymorphic bands that could be read

unambiguously on each gel image were used for data

analysis.

Phylogenetic analyses were performed with PAUP

Version 4.0b 10 (Swofford 2002), using neighbour join-

ing (NJ; Restriction-site distances: Upholt) and a maxi-

mum parsimony criterion. For the latter, starting trees

were generated by stepwise addition, swapping on best

tree only in case of multiple trees. One thousand random

addition replicates were chosen. The heuristic search

for best topologies used TBR branch swapping. Support

for clades in both distance and parsimony analyses was

measured using the non-parametric bootstrap method

(Felsenstein 1985; 10 000 replicates).

RESULTS

The taxa in the P. carnosum complex: etymology’ and

taxonomic history

The oldest mention of Pelargonium carnosum is found

on a herbarium sheet dating back to 1724 (BM649367, Fig-

ure 1A). The brief diagnosis reads: ‘'Geranium africanum

frutescens, Chelidonii folio; petalis florum angustis, albi-

dis; carnoso caudice ’. The herbarium specimen originates

from a plant that arrived at Chelsea Physic Garden in Lon-

don in the same year. The collector and place of origin of

this specimen are not known. A first detailed description

of P. carnosum was provided by Dillenius in 1732, who

described the species under the phrase name ‘ Geranium

Afric. carnosum, petalis angustis albicantibus ' — a pre-Lin-

Bothalia 39,1 (2009)

75

FIGURE 1. — A, B, Pelargonium camosum subsp. camosum, BM649367 : A, one of the earliest herbarium specimens; B, lectotype; C, lectotype of

P. camosum subsp. ferulaceum. D-H, habit: D, P. adriaanii; E, P. camosum subsp. camosum; F, G, P. polycephalum; H, P. camosum subsp.

ferulaceum. D, E, G, H, plant’s natural setting; F, in Worcester Botanical Garden. Scale bars: 100 mm.

76

Bothalia 39,1 (2009)

nean name not valid according to the rules of the Interna-

tional Code of Botanical Nomenclature (ICBN). Although

Dillenius did not give any indication as to the origin of

the material examined by him, it can be assumed from

the wording of the phrase name that he had indeed seen

that particular herbarium specimen from 1724. In 1755

the species became the first member of (the later) section

Otidia to be described by Linnaeus which he placed under

the genus Geranium as G. carnosum, as he did not distin-

guish the genus Pelargonium. Linnaeus referred to Dille-

nius’s illustration as the iconotype that accompanied the

phrase name of Geranium carnosum (Figure IB). Linnae-

us’s specific epithet, carnosus, the Latin word for fleshy or

succulent, refers to the succulent stem as a presumed cha-

racteristic feature — but nonetheless fairly common within

section Otidia — which he chose to separate P. carnosum

from P. gibbosum (sect. Polyactium, with swollen nodes

of the stem). In 1789, L’Heritier placed the species under

Pelargonium.

The first diagnosis of Pelargonium ferulaceum was

published by J. Burman in 1738 under the phrase name

‘ Pelargonium foliis ferulaceis, multifidis, flore rubello' .

The first part of the description refers to the deeply incised

delicate leaves that resemble those of the genus Ferula

(Apiaceae). The illustration by Burman is regarded as the

iconotype. Burman’s son (1759) was the first to adopt the

Linnaean binary nomenclature and, hence, has traditionally

been accepted as the author of this species name. Though

he referred to his father’s illustration (Figure 1C), he obvi-

ously described a member of section Polyactium : ‘dwarf

plant lacking stems, a simple stalk rising from the centre

of the root-bearing long-tubed flowers as found in Gera-

nium triste ’. Due to this error, we propose to drop Bur-

man filius as author of P. ferulaceum. Instead, Cavanilles

(1787) should be credited as the author who took up a part

of J. Burman’s diagnosis when describing ‘ Geranium feru-

laceum\ Willdenow (1800) placed this species under the

genus Pelargonium.

Pelargonium polycephalum was introduced by Meyer

(1843) who referred to the numerous capitate pseudo-

umbels as the character of distinction in this species {poly

is the Greek word for many, and cephalium is the Greek

word for head). As Meyer failed to publish a proper diag-

nosis, Harvey (1860) is credited as author of this species’

name. A specimen collected by Drege was designated

as the holotype. Harvey described the species under the

name P. ferulaceum var. polycephalum with P. carnosum

as the most closely related taxon. In 1912 Knuth raised

the taxon to specific rank as P. polycephalum.

The last species described in this complex was Pelar-

gonium adriaanii M. Becker & F.Albers which was pub-

lished by the present authors in 2005 in honour of the

late J.J. Adriaan van der Walt for his valuable contribu-

tions in the genus.

Key to taxa

la Pedicel longer than hypanthium P. adriaanii

lb Pedicel shorter than hypanthium:

2a Compact pseudo-umbel with ±10 flowers P. polycephalum

2b Loose pseudo-umbel with 4-6 flowers:

3a Leaves pinnately incised P. carnosum subsp. carnosum

3b Leaves bipinnately divided, pinnae petiolate

P. carnosum subsp. ferulaceum

Pelargonium carnosum and P. ferulaceum are treated

as subspecies, due to their close relationship and the fre-

quent occurrence of hybrids (see below). With P. carno-

sum being the older name of the two, the correct names

are P. carnosum subsp. carnosum and P. carnosum

subsp. ferulaceum (Cav.) M. Becker & F.Albers, comb,

nov. Pelargonium adriaanii and P. polycephalum are still

treated as separate species.

1 . Pelargonium adriaanii M.Becker & F.Albers in

Botanische Jahrbiicher 126,2: 153 (2005a). Type: North-

ern Cape, 2916 (Port Nolloth), (-BB), 17 Sept 2003,

Becker & Albers 4235 (MSUN, holo.!).

2a. Pelargonium carnosum {L.) L’Her. in Aiton,

Hortus kewensis 2: 421 (1789). Geranium carnosum

L.: 20 (1755). Otidia carnosa (L.) Sweet: t. 98 (1822).

Geraniospermum carnosum (L.) Kuntze: 94 (1891). Lec-

totype: Geranium africanum carnosum Dill.: 153, t. 127,

fig. 154(1732).

Pelargonium sisonifolium Baker, t. 28 (1869). Lectotype: ‘Cape’

[icono., Baker in Saunders, Refugium Botanicum: t. 28 (1869)].

2b. Pelargonium carnosum subsp. ferulaceum

(Cav.) M.Becker & F.Albers, comb. nov.

Geranium ferulaceum Cav., Quarta Dissertatio botanica: 265, t.

110, fig. 2 (1787). Pelargonium ferulaceum (Cav.) Willd.: 687 (1800).

Otidia ferulacea (Cav.) Eckl. & Zeyh.: 69 (1835-1837). Geraniosper-

mum ferulaceum (Cav.) Kuntze: 94 (1891). Lectotype: Pelargonium

foliis ferulaceis Burm.: 93, t. 36, fig. 1 (1738).

Pelargonium mammulosum J.C.Wendl.: 77 (1810). Otidia burman-

niana Eckl. & Zeyh., non DC.: 69 (1835-1837). Pelargonium burman-

nianum Steud.: 284 (1841). Lectotype: ‘Vorgebirge der guten Hoff-

nung’ [icono., Wendland, Collectio plantarum 2: t. 70 (1810)].

3. Pelargonium polycephalum ( Harv .) R. Knuth

ex E.Mey. in Drege, Zwei pflanzengeographische Doku-

mente: 209 (1843-1844); Engler: 372 (1912). P. feru-

laceum var. polycephalum Harv.: 279 (1859-1860).

Type: Northern Cape, ‘Kous-Silverfontein’, Drege 3244

(according to Knuth 1912), nomen nudum by E. Meyer:

209(1843-1844).

Morphological characters of section Otidia

In addition to the commonly occurring succulent

stems and pinnate leaves, the set of characters defining

section Otidia also includes short-spurred flowers.

The typical auricles borne at the base of the poste-

rior petals are restricted to this section, although similar

petal structures are found in certain species of sections

Campylia, Hoarea, Ligularia and Pelargonium (Struck

1997). Stamens that curl upwards at the end of the stam-

inate phase are restricted to section Otidia and a couple

of species in section Pelargonium (Struck 1997).

Pelargonium alternans J.C.Wendl. which hitherto has

also been placed in section Otidia , differs in various cha-

racters (Becker & Albers in press b). A large genetic

gap between this species and the rest of the section has

been established (Bakker et al. 2004; Becker & Albers

in press c), and P. alternans is excluded from the section

in the strict sense (= Otidia s.str.) in the present account.

We refer to Otidia s.l. in order to indicate traditional cir-

cumscription of the section, i.e. including P. alternans.

Bothalia 39,1 (2009)

77

The taxa in the Pelargonium carnosum complex

do not differ much from the other members of section

Otidia. However, most taxa in the P carnosum com-

plex and the closely related P. parviflorum complex

(Becker & Albers in press a) are distinguished by a char-

acter which is otherwise absent in the section (except

in P. altemans ): the pedicel is much shorter than in the

remaining Otidia species. The ratio between the lengths

of hypanthium and pedicel — a valuable characteristic in

Pelargonium (Miller 1996) — varies between 2 and 10

within both complexes, but between 0.1 and 1.0 in the

remaining members of Otidia s.str. However, P. adriaa-

nii does not fit into this pattern: in this species the pedi-

cel is much longer than the hypanthium. Therefore, no

phenotypic character clearly delineates the P. carnosum

complex. The taxa in the P. parviflorum complex differ

from those in the P. carnosum complex in possessing

tiny, mostly yellowish petals.

Diagnostic features in the P. carnosum complex (Table 2)

Pelargonium adriaanii (Figure ID) and P. poly-

cephalum (Figure IF, G) can develop into large plants of

1 m across that possess stems of 50 mm in diameter. In

P. carnosum subsp. carnosum (Figure IE) the stems are

much thinner and in subsp . ferulaceum (Figure 1H) they

remain shorter.

In Pelargonium adriaanii and P. carnosum subsp.

carnosum the lateral roots form series of small tubers;

P. carnosum subsp. ferulaceum and P. polycephalum

exhibit a simple root system without any thickenings.

Apart from their pinnate shape, the leaves do not

share a further character. They differ in the degree of

succulence and density of the indumentum (Figure

2A). In Pelargonium carnosum subsp. carnosum and P.

adriaanii , the leaves are rather herbaceous and densely

covered with partially very long hairs; in P. carnosum

subsp. ferulaceum and P. polycephalum, the leaves

are slightly succulent and covered with microscopi-

cal papillae. Conversely, Harvey in his description of

P. polycephalum, maintained that the leaves are much

more hairy in this species than in P. carnosum s.str.,

although the reverse is correct according to our obser-

vations. Leaf shapes differ clearly in the four taxa: in

P. carnosum subsp. carnosum and P. polycephalum.

the leaves are mostly pinnately incised, but bipinnately

divided in P. carnosum subsp. ferulaceum and in P.

adriaanii. In P. carnosum subsp. ferulaceum almost all

pinnae have a petiolule, but only the first pair of pin-

nae in the remaining three taxa have a petiolule. Peti-

oles tend to persist in section Otidia and old leaf bases

cover the stems as acute or blunt, ‘thorny’ fragments.

Only in P. carnosum subsp. carnosum are the petioles

usually shed with the rest of the leaves, giving the

stems a smooth overall outline.

As to floral structure, the four taxa in the complex

can be divided into two groups: 1, Pelargonium adri-

aanii; and 2, P. carnosum subsp. carnosum, P. car-

nosum subsp. ferulaceum and P. polycephalum. The

pedicel is exceptionally long in P. adriaanii (Figure

2F) compared to those of all other taxa in the P. carno-

sum complex and the pseudo-umbels of the first taxon

appear fairly loose. In contrast, the pseudo-umbels of

P. carnosum subsp. carnosum (Figure 2B), P. carnosum

subsp. ferulaceum (Figure 2C, E) and P. polycephalum

(Figure 2G) appear rather compact due to the very short

pedicels. The capitate shape of the pseudo-umbels in P.

polycephalum results from the shortened hypanthia and

a higher number of flowers per partial inflorescence. P.

carnosum subsp. carnosum, P. carnosum subsp. ferula-

ceum and P. polycephalum exhibit long and narrow pet-

als. The flowers are most commonly white in all three

taxa sometimes suffused with pink and bearing dark

pink markings or stripes on all five petals (Figure 2D,

E). Since there is no detectable pattern in the distribu-

tion of these variations in floral pigmentation, the three

taxa can be separated as a subgroup with respect to

flower structure. Conversely, P. adriaanii exhibits the

longest and broadest petals within section Otidia (Fig-

ure 2F). With the exception of their purple markings,

the petals are pure white; no additional flower colour

occurs.

A comparison of early and later states of inflorescence

development reveals important distinctions (Figure 3).

Pelargonium carnosum subsp. carnosum and P. adri-

aanii exhibit a loosely branched inflorescence with 4-5

nodes and each node generates a single or a branched

peduncle (simplified in Figure 3). Altogether the inflor-

escence is composed of 6-12 pseudo-umbels with 4-6

TABLE 2. — Morphological differences in taxa of Pelargonium carnosum complex

78

Bothalia 39,1 (2009)

FIGURE 2. — Leaves and flowers of taxa in Pelargonium carnosum complex. A, leaves, from left to right: P. carnosum subsp. carnosum ( STEU1815 ),

P. carnosum subsp. ferulaceum (A&B4118), P. polycephalum ( A&B4232 ), P. adriaanii (A&B4237). B-G, flowers: B, P. carnosum subsp.

carnosum (STEU1815)\ D, P. carnosum subsp. carnosum x P carnosum subsp .ferulaceum (A2622). C, E, P. carnosum subsp .ferulaceum'.

C, (. 4&B4II8); E, (A&B4435). F, P adriaanii ( A&B4237 ); G, P polycephalum ( A&B4232 ). Scale bars: 10 mm.

Bothalia 39,1 (2009)

79

onset of flowering — * later in the year

9 flowering pseudo-umbel X shed pseudo-umbel

FIGURE 3. — Inflorescence branching patterns in P. camosum complex.

The inflorescence depicted to the left is the basal condition from

which all taxa start at different times of the year. The drawings

to the right (which are drawn to a smaller scale than the drawing

to the left) represent three distinct inflorescence types at different

times later in the year.

flowers each. The inflorescence of P camosum subsp.

femlaceum fits the overall pattern, but continues to pro-

duce further pseudo-umbels after the older flowers have

already faded. The youngest pseudo-umbels are formed

successively a few mm higher and higher; this leads to

a delicately branched structure that resembles a panicle

of grasses when not in flower. Individual inflorescences

may last several months in P. camosum subsp. ferula-

ceum. Due to its great vitality it may even produce veg-

etative shoots from the lower half of its inflorescence

after flowering has ceased (not represented).

Pelargonium polycephalum likewise differs from P.

camosum subsp. camosum and P. adriaanii by a larger

number of pseudo-umbels. However, in contrast to P.

camosum subsp. ferulaceum, these do not emerge suc-

cessively at the tip of the continuously growing pedun-

cle, but develop in large numbers from already visible

buds. Each of the 4 or 5 nodes of the principal axis may

produce 10-15 pseudo-umbels with about 10 flowers

each, which may add up to some 500 flowers per inflor-

escence within a relatively short period of time. Com-

pared to P. camosum subsp. ferulaceum , the life span of

an inflorescence is short in P. polycephalum-. the period

between full flowering and fruiting and immediate with-

ering of the inflorescence will last a few weeks only.

Hence, fully developed inflorescences are easily assigned

to the respective taxa, but not young inflorescences, as

these all represent variations of the same basic structural

type.

Distribution and habitat

The four taxa in the Pelargonium carnosum complex

occur within clearly circumscribed distribution areas

which do not overlap much (Figure 4). P. adriaanii

from the Northern Cape is geographically well isolated:

there is no record of any other member of section Otidia

within a radius of 50 km. Only a single plant resem-

bling P. parviflorum is known from Kleinsee south of

Port Nolloth ( Drijfhout 2842 sub STEU2979). With the

exception of a single disputed specimen, P. adriaanii has

always been collected near the coast at altitudes of up to

50 m (Becker & Albers 2005a). P. adriaanii lodges at

the fringes of the Gariep Centre, which is a major centre

of endemism (Van Wyk & Smith 2001) bordering on the

southern Namib where the species receives less than 100

mm annual rainfall (Figure 4).

Pelargonium camosum subsp. camosum is exclu-

sively found in the Western Cape within an area that

stretches from the Atlantic coast to the chain of the Ced-

erberg-Swartruggens-Hexrivier Mountains at altitudes

of up to 200 m. Receiving an annual rainfall of more

than 300 mm, this region exhibits transitions to fynbos

vegetation. The most typical specimens of P. carnosum

subsp. carnosum are restricted to the Sandveld right at

the coast. The two remainders in the P. carnosum com-

plex occur further inland, receiving 150-250 mm of

annual precipitation. The area of P. carnosum subsp.

femlaceum is largely situated in the Western Cape and

stretches into the Northern Cape. The range of subsp.

ferulaceum borders on the area of Pelargonium carno-

sum subsp. carnosum , but instead of proceeding to the

Atlantic coast, stretches in the opposite direction towards

the shrubland of the Great and Little Karoo. The subspe-

cies occurs mostly at altitudes of 600-1 000 and in the

vicinity of Vanrhynsdorp, merely at 100 m. P. polyceph-

alum is largely restricted to the Namaqualand Hills in the

Northern Cape Province at altitudes of up to 1 200 m.

Flowering phenology

As to the onset of flowering, there is a general dine in

section Otidia running from north to south: taxa from the

north (Namibia) exhibit an earlier blooming than their

relatives from the south (Western Cape). This is par-

ticularly true for Pelargonium ceratophyllum L’Her., P.

klinghardtense R.Knuth and P. paniculatum Jacq. which

already flower in early August. Although flowering starts

at the same time in P. laxum (Sweet) G.Don from the

Eastern Cape (receiving rainfall during summer), this

species is subject to different environmental and climatic

conditions. In most members of the P. carnosum com-

plex, flowering starts not earlier than mid-September (P.

adriaanii, P. carnosum subsp. femlaceum) or early Octo-

ber (P. carnosum subsp. carnosum). The same is true for

the taxa of the closely related P. parviflorum complex.

With the exception of subsp. karasbergense (Becker &

Albers in press a) which, similar to P. laxum occurs in

80

Bothalia 39,1 (2009)

* P. adriaanii

■ P. carnosum subsp. carnosum

• P. carnosum subsp. ferulaceum

O P. polycephalum

(■> Hybrid 1

® Hybrid 2

FIGURE 4. — Distribution of taxa in

P. carnosum complex accord-

ing to records from herbarium

and living specimens. Dark

grey shading = winter rain-

fall area; light grey shading =

annual rainfall area.

the transitional zone between the winter and summer

rainfall areas, flowering commences only from mid-Sep-

tember onwards in all subgroups of the latter complex —

even including the taxa from Namibia. As was deduced

from the collecting dates of dried specimens bearing

flowers, P. polycephalum from the Namaqualand Rocky

Hills is the only taxon in both complexes exhibiting a

relatively early blooming. This species flowers from

August to September. Hence, there is little overlap in the

timing of the reproductive seasons of P. polycephalum

and the remaining species in the P. carnosum complex

and in the P. parviflorum complex. Onset and end of the

flowering period may vary according to rainfall patterns;

nevertheless, the sequence of ‘early’ and Tate’ blooming

taxa is maintained.

Morphological evidence of hybridization

Although the four taxa in the Pelargonium carnosum

complex are clearly distinguished, the existence of inter-

mediates is unmistakable. Interbreeding has repeatedly

been recorded from areas where different taxa in the P.

carnosum complex occur in close proximity.

Hybrids within the complex

The region west of the Cederberg (Vredendal-

Clanwilliam) harbours hybrids between the two sub-

species of Pelargonium carnosum which exhibit leaf

features of both parental taxa ( A&B4373 ). Plants from

this area show the same type of indumentum as found in

typical P. carnosum and pinnae as stalked and narrow as

found in P. carnosum subsp. ferulaceum.

Plants from the coastal region near Papendorp and

Doringbaai mostly exhibit pink petals ( A&B4389 ,

A&B4397). In this region, not less than three genotypes

appear to intergrade, as the specific characters of all

three taxa were found to combine in plants growing next

to each other. These plants develop leaves as observed in

P. carnosum subsp. carnosum , the distinct inflorescence

of P polycephalum and the long pedicels of P. adriaanii.

Hybridridzation involving taxa outside this complex

The closest relatives of the Pelargonium carnosum

complex are found in the P. parviflorum complex. Struc-

tural intermediates occur in several regions which point

to large-scale interbreeding between both complexes.

The resulting cluster of hybrids are discussed elsewhere

in more detail (Becker & Albers in press a) and are only

briefly characterized in the present account. In this clus-

ter of hybrids, floral structure is conspicuously varied

exhibiting a range of petal colours and shapes.

The closest relatives of both complexes combined are

Pelargonium laxum (Sweet) G.Don and P. dasyphyllum

R.Knuth (Becker et al. 2008). Both species are clearly

distinguished on account of a set of structural charac-

teristics. P. laxum possesses posterior petals which are

sharply reflexed from bases at nearly 1 80° and unusually

long stamens. P. dasyphyllum exhibits a cushion-shaped

growth habit and relatively small leaves. In both spe-

cies, plants recorded from the geographical fringes also

exhibit characters of the P. carnosum complex.

One record pertains to a plant of remote resemblance

to Pelargonium dasyphyllum (. A&B4286) which we

tracked down in the midst of a population of typical

members of that species. In this plant, the stem is unusu-

ally thickened and leaves are exceedingly large. The

wide range of different petal shapes of individual speci-

mens of P dasyphyllum even includes the petal structure

found in typical P. carnosum flowers.

Molecular analysis (AFLP)

The phylogram (Figure 5) results from a neigh-

bour joining (NJ) analysis using 416 AFLP markers.

Non-parametric bootstrap values (BVs) are indicated.

A maximum parsimony analysis was also undertaken

(not shown) employing 193 informative characters. The

major groupings of taxa were congruent between both

analyses.

Bothalia 39,1 (2009)

81

P allernans \

P. ceratophyllum

P klinghardtense

| P. crithmifolium

' P paniculatum

P. dasyphyllum

P. laxum

P. brevipetalum

P. parvifiorum s.l

subsp. rotundipetalum

subsp. tuberculum

subsp. parvifiorum

ubsp. karasbergense

other species of

O lidia s.1.

'r P parvifiorum

s.str.

A&B 4405

j A&B 4120

_fl A&B 4118

611

1 A&B 4435

— A&B 4444

A&B 4373

A&B 4426

STEU 2401

STEU 1815

- A&B 4419

A&B 4430

A&B 4389

A&B 4397

A&B 4232

- A&B 4313

A&B 4298

A&B 4238

A&B 4236

A&B 4237

Hybrid

(P cam q-sovt? x p. par/Mofum)

P. carnosum

subsp ferulaceum

Hybrid 1

P. carnosum

ijbap carnosum

Hybrid 2

P. polycephalum

P. adriaanii

2

3

o

cn

C

3

o

o

3

■o

ct>

X

10 changes

FIGURE 5. — Cladistic analysis of

taxa in P. carnosum complex

based on AFLP patterns; phy-

logenetic tree reconstructed

by neighbour joining analy-

sis using 416 AFLP markers.

Bootstrap values derived from

a maximum parsimony analy-

sis employing 193 informative

characters are indicated for

clades that are congruent to

the strict consensus tree. Col-

lections that were unequivo-

cally grouped under defined

phenotypes of the complex

are highlighted with dark grey

shading.

There are three questions with regard to the Pelargo-

nium carnosum complex: 1, is this complex a monophy-

letic group?; 2, are the distinct phenotypes reflected in

distinct genotypes?; and 3, can the presumed existence

of structural intermediates be verified?

Comparison with other members of section Otidia s.l.

( P. altemans, P. ceratophyllum , P. crithmifolium Sm., P.

dasyphyllum , P. klinghardtense, P. laxum and P. panic-

ulatum) yields indications as to the monophyly of the

complex. The study also involved the subspecies of the

closely related P. parvifiorum complex (Becker & Albers

2005b, in press a).

In the cladogram (Figure 5) the four taxa of the Pel-

argonium carnosum complex emerge as a single mono-

phyletic group (clade B) with the P. parvifiorum com-

plex (clade A) as sister group. The clade that comprises

clade A and clade B is well supported by a BV of 95.

With less good support (BV 79), P. dasyphyllum and

P. laxum share a clade with the group that comprises

clades A and B. Despite the many morphological signs

of interbreeding between the two complexes, the typi-

cal phenotypes of P. adriaanii, P. polycephalum and

subspecies of P. carnosum emerge as monophyletic

groups (highlighted in dark grey in Figure 5). After

excluding the samples of the supposed hybrids from

the analysis, BVs rise to 95-100 in support of these

groups (not represented in Figure 5). The cladogram

seems to corroborate the presumed interbreeding pro-

cesses among several taxa in the P. carnosum complex.

AFLP data are also consistent with the notion of phe-

notypic intergrades among the two subspecies of Pelar-

gonium carnosum (Hybrid 1: A&B4444, A&B4373,

A&B4426, STEU2401 ) and among P. carnosum subsp. car-

82

Bothalia 39,1 (2009)

nosum/P. polycephalumlP. adriaanii (Hybrid 2: A&B4389,

A&B4397). However, possible interbreeding processes

also involve taxa in the P. parviflorum complex.

In terms of the grouping of taxa, one would expect

both subspecies of Pelargonium carnosum to share one

clade. However, subsp . ferulaceum appears in the clado-

gram as a sister group to clade B2b, which contains not

only subsp. carnosum but also P. po/ycephalum and P.

adriaanii. This may be seen as a conflict of molecular

data and proposed taxonomy, which is based on mor-

phology. On the other hand, this sister group relation-

ship to the remaining taxa of the P carnosum complex

might reflect the ongoing gene flow between P. carno-

sum subsp. ferulaceum and the P. parviflorum complex,

which has been predicted after evaluation of morpholog-

ical traits.

Hybrid 1 comprises four plants in the cladogram that

are supposed to result from gene flow between both sub-

species of Pelargonium carnosum. As subsp. ferulaceum

and subsp. carnosum do not share a clade, it is no sur-

prise that the same is true for their hybrids.

The distribution areas of several taxa of the two com-

plexes meet in the southern part of the Knersvlakte. In

this area particularly, the two subspecies of P. carnosum

and P. parviflorum s.l. appear to interbreed. Collection

A&B4405 from Clanwilliam shows a phenotype similar

to the intermediates of the two subspecies of P. carno-

sum ( A&B4373 , Vredendal population), but it is only one

selected from an array of plants that intergrades to the P.

parviflorum complex (Becker & Albers in press a).

DISCUSSION

Original descriptions

The distribution area of Pelargonium carnosum was

assumed to be fairly large (Van der Walt 1977; Vorster

1990), since the delineation of P. carnosum subsp. car-

nosum was blurred, due to the occurrence of numerous

intermediates. Furthermore, even taxa foreign to this

complex were uncritically grouped under P. carnosum as

well. However, when taxonomically treated in a stricter

sense, P. carnosum is actually restricted to a compara-

tively small geographical area (Figure 4). This taxonomi-

cal concept is in line with Dillenius’s original description

of 1732 and the available herbarium records dating back

to the first half of the 1 8th century both with respect to

the morphological characteristics as well as to their geo-

graphical distribution.

With regard to the circumscription of Pelargonium

carnosum subsp. ferulaceum, contradictions in the ear-

lier descriptions (Burman f. 1759; Cavanilles 1787; Lin-

naeus 1826) led to a similar state of taxonomic limbo.

This becomes obvious when comparing leaf characters

given for lP. ferulaceum' and lP. carnosum’’, respec-

tively (degree of division or separation of the leaf blade,

degree of hairiness and succulence). Burman ’s iconotype

of P. ferulaceum contains two variants of the same taxon

which differ in leaf shape and (according to the diagno-

sis) in petal colour. The significance and reliability of

this information can be questioned in view of Burman’s

erroneous representation of the number of petals: the

author believed both of his specimens to carry a single

posterior petal and three or four anterior petals, which is

not found in the genus Pelargonium.

So it happens that the view adhered to in the present

account of ‘ Pelargonium ferulaceum' not being synony-

mous to ‘ P. carnosum' is based to a lesser degree on the

original description (with the exception of leaf morphol-

ogy) than on the extensive study of herbarium speci-

mens, observations in the field and a molecular analysis

of populations sampled from their natural habitats.

While the precise identity of the specimens on which

'Pelargonium ferulaceum' were based remains obscure,

the diagnosis of P. polycephalum is fairly straightfor-

ward. Although we could not examine the holotype itself

( Drege 3244), we were able to study a comparable speci-

men ( Drege 1033) which clearly showed the diagnostic

features given in the original description for P. poly-

cephalum'. capitate pseudo-umbels and a panicle-like

inflorescence. The notion of a conspicuous indumentum

shown on the leaves remains incomprehensible. All dried

and live specimens seen by us showed almost glabrous

leaves.

Morphology and ecological function in a geographical

context

Many structural differences between the taxa in the

complex can be explained as adaptations to prevailing

environmental conditions. With regard to leaf shape,

two basic traits can be distinguished which correlate to

the geographical distance from the coast. Plants occur-

ring close to the sea exhibit virtually non-succulent and

densely hairy leaves, whereas taxa from further inland

are characterized by succulent and (to the naked eye)

glabrous leaves. Another character shaped by habitat

factors is the underground organs of the plants. Pelargo-

nium adriaanii and P. carnosum subsp. carnosum occur-

ring in Sandveld habitats near the Atlantic coast pos-

sess lateral roots with thickened sections, whereas the

two taxa occurring further inland, P. carnosum subsp.

ferulaceum and P. polycephalum, mostly on rugged soil,

exhibit fibrous root systems lacking tubers.

Since the taxa in the Pelargonium carnosum com-

plex are adapted to different climatic, edaphic and topo-

graphical factors and are associated with certain habitats,

one could view them as mere ecotypes of a single taxon.

However, this view is contradicted by the observation

that P. adriaanii and P. polycephalum maintain their typi-

cal growth habit even under cultivation, and do not seem

to hybridize on a large scale, which speaks in favour of

treating them as separate species. While intermediates

between P. adriaanii, P. polycephalum and P. carnosum

were rarely recorded, the situation is different in "P. car-

nosum' and ‘ P. ferulaceum' . In view of their extremely

different habitats (Sandveld and Little Karoo) on the one

hand and their relatively subtle structural differences on

the other hand, it seems appropriate to treat them as sub-

species. This view is supported by the occurrence of a

range of phenotypic intermediates which is correlated to

the distance from the sea and the immediate degree of

aridity, respectively.

Bothalia 39,1 (2009)

83

Molecular analysis and its taxonomic application

The results of the molecular analyses (AFLP) are

mostly consistent with the hypotheses based on morpho-

logical observations. The four taxa in the Pelargonium

carnosum complex emerge as a monophyletic group

within section Otidia (Figure 5). P. carnosum subsp. car-

nosum and P. carnosum subsp. ferulaceum were treated

as a single taxon by many authors (Van der Walt 1977;

Vorster 1990), since neither their geographical range nor

their morphological features were sufficiently known.

Our molecular study confirms their close relationship but

also provides evidence in favour of the existence of two

distinct genotypes.

The close relationship of Pelargonium adriaanii and

P. polycephalum as revealed by the AFLP analysis is sur-

prising as there are profound differences in floral struc-

ture between P. adriaanii and P. polycephalum. Preced-

ing the description of P adriaanii, a short pedicel was

among the diagnostic features given for the circumscrip-

tion of the complex. The constricted pseudo-umbels of

P. polycephalum versus the loose pseudo-umbels of P.

adriaanii represent the extremes in this feature.

If the analysis is exclusively restricted to typical

samples in the various taxa, high values are yielded in

the maximum parsimony analysis and increased lengths

of branches in the phylogenetic tree (not represented in

Figure 5). The inclusion of all morphological intermedi-

ates demonstrates that interbreeding is the rule within the

Pelargonium carnosum complex on the one hand (Figure

5) and among the P. carnosum complex and the closely

related P. parviflorum complex on the other hand (Becker

& Albers in press a).

Finally, the question arises: which taxonomic rank

is appropriate in the Pelargonium carnosum complex?

Following the biological species concept (Dobzhansky

1937), a species is a reproductive group; interbreeding

among species is prevented by various isolation mecha-

nisms. These include, in simple terms, mechanical, tem-

poral, habitat-related and genetic barriers to interbreed-

ing (Avise 2004). Mechanical isolating mechanisms, as

brought about in the flower of different subgroups of

Pelargonium alternans (Becker & Albers in press b), are

absent in the P. carnosum complex. Despite differences

in petal size and in lengths of pedicels, the taxa show

basically the same floral structure.

In contrast, a seasonal isolation resulting from the

development of different flowering periods is found in

Pelargonium polycephalum. Nevertheless, a slight over-

lap of this species’ reproductive season with that of P.

carnosum subsp. carnosum results in the occurrence of

hybrids in the region of Papendorp-Doringbaai (3118

CA and CC). Interbreeding between P. polycephalum

and the cluster of hybrids in the P. parviflorum complex

could also be expected due to their generally sympat-

ric distribution and insufficiently separated flowering

seasons. However, the occurrence of rudimentary and,

consequently, sterile anthers in many ‘ P. polycephalum ’

plants speak against consecutive interbreeding among P.

polycephalum and P. parviflorum beyond the F 1 genera-

tion (Becker & Albers in press a).

A habitat-related or geographical isolation is particu-

larly found in Pelargonium adriaanii. While the distri-

bution areas of the remainders in the complex border

on each other, the range of P. adriaanii is situated fairly

isolated in the region around Port Nolloth — as far as this

can be assessed in view of the fact that we were unable

to enter the local diamond mining area. From rare occur-

rences of P. adriaanii- like features near Papendorp, the

existence of a gene substitution along the stretch of the

Sandveld is presumed.

Intermediates between Pelargonium carnosum and

P. polycephalum or P. adriaanii are on record, but such

intermediates occur markedly less often than intermedi-

ates among the subspecies of P. carnosum. Since numer-

ous morphological characteristics support the distinct-

ness of their gene pools and as there are no indications of

interbreeding processes going beyond the FI generation,

P. polycephalum and P. adriaanii continue to qualify as

separate species. The position is completely different

in the subspecies of ‘'P. carnosum ’ and ‘P. ferulaceum' .

Large-scale hybridization as in the region of VredendaL

Clanwilliam demonstrate the absence of effective repro-

ductive isolation mechanisms; P. ferulaceum is therefore

reduced to the rank of subspecies. There is evidence that

P carnosum subsp. ferulaceum is connected to P. parvi-

florum subsp. parviflorum via a full range of intergrading

forms in the area of geographical overlap. In Becker &

Albers (in press a) we will provide material that none-

theless supports the treatment of P. parviflorum and near

related taxa as a distinct complex.

A recent breakdown of reproductive barriers result-

ing from human action as has been presumed by Vorster

(1990) cannot be excluded, but such an assumption is

not needed to explain the large number of interbreeding

taxa. A shift in species distribution ranges due to recent

climatic changes may have played a role.

As long as there is no artificial transport of propa-

gules, the spread of settlements, agriculture, overgrazing

and diamond mining could also isolate small populations

from each other, as this would render large intermittent

areas unsuitable to support the natural plant life. Nev-

ertheless, man will hardly contribute to increase biotic

diversity but rather destroy genetic diversity of the local

flora as well as its natural habitats. Pelargonium adriaa-

nii is currently threatened with extinction unless the dia-

mond companies denounce their rights to fully exploit

the diamond fields within the species’ range.

The massive impact of human activities on the veg-

etation within the Cape floristic region is uncontested.

Today the species-rich flora and fauna is protected

within an increasing number of national parks and nature

reserves. However, the Cape floristic region stands out in

its wealth of endemics which often show a very localized

distribution outside of protective areas. In order to pro-

tect even this species from extinction, centres of diver-

sity have to be identified and plant diversity has to be

put on record through taxonomic contributions. A further

article on Pelargonium sect. Otidia will elucidate the

unexpected radiation within the P. parviflorum complex,

which hitherto has been treated as a single taxon.

84

Bothalia 39,1 (2009)

SPECIMENS EXAMINED

Acocks 14847 (2b) K, PRE. Albers 2262 , 2492 (2a) MSUN; 4174 ,

4175, 4176 (2b) MSUN. Albers & Becker 4235, 4236 (1) MSUN;

4184, 4185, 4186, 4382, 4383, 4384, 4385, 4410, 4411, 4412, 4413,

4414 (hybr. 1) MSUN; 4388, 4390, 4392, 4393, 4395, 4416, 4420,

4421, 4422, 4423, 4427, 4428, 4430, 4431, 4432 (2a) MSUN; 4461

(2b) MSUN; 4232, 4301, 4302, 4318, 4319, 4396, 4398 (3) MSUN.

Albers, Becker & Strauch 4083, 4091, 4095, 4112, 4116, 4117, 4118,

4119, 4120, 4121, 4122, 4123 (2b) MSUN; 4081, 4084 (3) MSUN.

Barks.n. (2a) K 74365. Baver 1651 (3) NBG. Bolus 6531 (1) K, PRE;

s.n. (2a) BOL107101, K74361; 13047 (2b) PRE; 446 = Herb. Bolus

6531 (3) BM, BOL, K. Boucher 2972 (2a) NBG; 2871 (2b) NBG,

PRE. Bremer 363 (2a) PRE. Bultro 123 (hybr. 1) K. Chelsea Phys.

Gdns s.n. (2a) BM649367, 649368.

Compton 8515 (2b) NBG; 6838, 17303 (3) NBG.

Dobay 59/75, 70/75, 79/75 (2b) NBG. Drake s.n. (2a) PRE 56350.

Drege 7481 (2a) PRE; 01033 (3) PRE. Dreyer 39 (1) PRE. Drijfhout

1301 (2b) NBG, 2702 (2b) PRE.

Esterhirysen 1806 (2b) BOL; 22851 (2b) BOL, NBG; s.n. (2b) BOL

107074; 23593 (3) BOL. Euthen 2550 (hybr. 1) NBG.

Fischer 30 (2b) PRE.

Germishuizen 4811, 5311 (1) PRE. Gibby & Crompton 19 (hybr. 1)

BM. Goldblatt 2395 (3) M, NBG, PRE.

Haftrom & Acocks 1986 (hybr. 1) BOL. Hall s.n. sub NBG. 98/68 (2b)

NBG; 3475 (hybr, 2) NBG, PRE; 5260 (hybr. 2) NBG, PRE. Hardy

241 (2b) PRE. Hattingh 572/68 (hybr. 1) NBG. Herb. Regiurn Mona-

cense s.n. (2a) M 36641, 36642, M 36643, 36644, 36645, 36646,

36647. Herre 2892, 7503 (2b) BOL; 2988 (3) BOL, K. Hugo s.n. (1)

NBG 177505, PRE 650792.

Leighton 1243 (3) BOL. Leipoldt s.n. sub N.B.G. 1419/25 (2a) BOL;

19396 (2b) BOL; 3795 (3) BOL. Leistner 707 (2b) PRE; 2553 (3) K.

Liede & Meve 619 (2b) MSUN.

Maguire 986 (3) NBG. Marloth 4026 (2a) PRE; s.n. (2a) PRE 56347;

8213 (2b) PRE; 12261 (3) BOL, NBG, PRE. Morris 5598 (3) K.

O ’ Call . & Steensma 1689 (2a) NBG. O ' Callaghan 650 (hybr. 2) NBG.

Odensen 1480 (hybr. 1 ) BOL.

Pearson 5301 (2b) K; 6487 (3) BM, K. Pienaar 1156 (1) PRE. Pillans

5626 ( 1 ) BOL, K; 6631, 6981 (2a) BOL.

Rechinger A3231 (hybr. 2) M.

Salter 1718 (2a) BM; 6465 (2a) BOL. Schlechter s.n. (1) NBG 177507;

1007 (3) PRE; 8199 (3) BM, K; s.coll. s.n. (2a) BM649375, 649379.

Scully 196 (3) BOL. Stell. Univ. Gdns 2996 (2b) BOL. Stokoe 9541

(2a) PRE.

Taylor 12359 (hybr. 1 ) NBG; 7542 (2b) NBG.

Van der Walt 763 (hybr. 1) MSUN, PRE; 918 (2a) MSUN; 532 (2b)

PRE; 1201 (2b) MSUN; s.n. (2b) PRE572097. Van der Walt & Vorster

1034 (hybr. 1) PRE. Van Jaarsveld s.n. (2b) NBG 1 1 4903; 5704 (hybr.

2) NBG. Van Niekerk s.n. (2b) BOL107029. Vorster 2399, 2404 (2b)

PRE.

Wagener 30 (2b) NBG. Ward 32, s.n. sub STEU3853 (2a) MSUN.

Wilman 2316 ( 1) BOL. Wisura 584 ( 1) NBG; 268 (hybr. 1) NBG; 374,

1708 (2a) NBG; 2105 (2b) NBG.

ACKNOWLEDGEMENTS

We are indebted to the Department of Agriculture,

Land Reform, Environment and Conservation of the

Northern Cape and the Department of Fauna, Flora,

CITES & Hunting of the Western Cape for granting col-

lecting permits. The curators of the herbaria BM, BOL,

K, NBG, and PRE are thanked for loans of numerous

specimens.

REFERENCES

AITON, W. 1789. Hortus kewensis 1, vol. 2: 421. Nicol, London.

AVISE, J.C. 2004. Molecular markers, natural history, and evolution 2.

Sinauer Associates, Sunderland, Massachusetts.

BAKER, J.G. 1869. In W.W. Saunders, Refugium botanicum 1: t. 28.

London.

BARKER, F.T., CULHAM, A., HETTIARACHI, P., TOULOUMENI-

DOU, T. & GIBBY, M. 2004. Phylogeny of Pelargonium (Gera-

niaceae) based on DNA sequences from three genomes. Taxon

53: 17-28.

BARKER, F.T., HELLBRUGGE, D„ CULHAM, A. & GIBBY, M.

1998. Phylogenetic relationships within Pelargonium section

Peristera (Geraniaceae) inferred from nrDNA and cpDNA

sequence comparisons. Plant Systematics and Evolution 211:

273-287.

BECKER, M. & ALBERS, F. 2005a. Pelargonium adriaanii (Gera-

niaceae), a new species from the Northern Cape Province, South

Africa. Botanische Jahrbiicher 126: 153-161.

BECKER, M. & ALBERS, F. 2005b. Was ist Pelargonium parvifloruml

Avonia 23,2: 31-39.

BECKER, M„ SCHAPER, K. & ALBERS, F. 2008. Description of Pel-

argonium keeromsbergense and two subspecies of Pelargonium

laxurn of section Otidia (Geraniaceae). Schumannia 5 : 181-190.

BECKER, M. & ALBERS, F. in press a. Taxonomy and phylogeny of

two subgroups of Pelargonium section Otidia (Geraniaceae). 2.

The Pelargonium parviflorum complex. Bothalia.

BECKER, M. & ALBERS, F. in press b. Pollinator shift and speciation

in Pelargonium alternans (Geraniaceae). Biodiversity & Ecol-

ogy-

BECKER, M. & ALBERS, F. in press c. Phylogeny and speciation in

succulent Geraniaceae (Geraniales). Biodiversity & Ecology.

BURMAN, J. 1738. Rariorum Africanarum plantarum : 93. Boussiere,

Amsterdam.

BURMAN, N.L. 1759. Specimen botanicum de Geraniis: 40. Haak,

Leiden.

CAVANILLES, A.J. 1787. Quarta dissertatio botanica : 265-266, t. 99,

110. Didot, Paris.

DILLENIUS, J.J. 1732. Hortus elthamensis. London.

DOBZHANSKY, T. 1937. Genetics and the origin of species. Columbia

University Press, New York.

DOYLE, J.J. & DOYLE, J.L. 1987. A rapid DNA isolation procedure

for small quantities of fresh leaf tissue. Phytochemical Bulletin

19: 11-15.

DYER, R.A. 1953. Pelargonium carnosum. The Flowering Plants of

Africa 29: t. 1145.

ECKLON, C.F. & ZEYHER, C. 1835-1837. Enumeratio plantarum

Africae australis extratropicae 1-3: 68, 69. Hamburg.

FELSENSTEIN, J. 1 985. Confidence limits on phylogenies: an approach

using the bootstrap. Evolution 39: 783-791.

HARVEY. W.H. 1859-1860. In W.H. Harvey & O.W. Sonder, Flora

capensis: 279. Hodges, Smith, Dublin.

L'HERITIER DE BRUTELLE, C.L. 1789. In W. Aiton, Hortus kewen-

sis 1, vol. 2: 421. Nicol, London.

HOLMGREN, P.K., HOLMGREN, N.H. & BARTLETT, L.C. 1990.

Index herbariorum, edn 8. Regnum Vegetabile 120. New York.

KNUTH, R. 1912. Geraniaceae. In A. Engler, Das Pflanzenreich, vol. 4.

129: 369-372, 587. Engelmann, Leipzig.

KUNTZE, O. 1891. Revisio generum plantarum: 94. Felix, Leipzig.

LINNAEUS, C. 1755. Centuria 1. plantarum: 20. Uppsala.

LINNAEUS, C. 1826. Systema vegetabilium 16: 63. Gottingen.

MARSCHALEK, R. 2003. Marker Assisted Selection for the develop-

ment of intervarietal substitution lines in rapeseed (Brassica

napus L.) and the estimation of QTL effects for glucosinolate

content. Ph.D. thesis, Georg-August-Universitat, Gottingen.

MEYER, E.H.F. 1843-1844. In J.F. Dreg e, Zwei pflanzengeographische

Dokumente: 209.

MILLER, D. 1996. Pelargoniums — a gardener's guide to the species

and their cultivars and hybrids. Batsford, London.

STRUCK, M. 1997. Floral divergence and convergence in the genus

Pelargonium (Geraniaceae) in southern Africa: ecological and

evolutionary considerations. Plant Systematics and Evolution

208: 71-97.

SWEET, R. 1820-1822. Geraniaceae. The natural order of Gerania 1 :

t. 98. London.

SWOFFORD, D.L. 2002. PAUP: Phylogenetic Analysis Using Parsi-

mony and other methods, Version 4.0 beta 10 Version. Computer

program distributed by the Laboratory of Molecular Systemat-

ics, Smithsonian Institution, Washington DC.

Bothalia 39,1 (2009)

85

VAN DER WALT, J.J.A. 1977. Pelargoniums of southern Africa, vol. 1.

Purnell, Cape Town.

VORSTER, P.J. 1990. Taxonomy of the genus Pelargonium : review of

the section Otidia. Proceedings of the International Geraniaceae

Symposium-. 281-294. University of Stellenbosch.

VOS, P, HOGERS, R., BLEEKER, M., REIJANS, M„ VANDELEE,

T„ HORNES, M„ FRIJTERS, A., POT, J„ PELEMAN, J., KUI-

PER, M. & ZABEAU, M. 1995. AFLP: a new technique for

DNA fingerprinting. Nucleic Acids Research 23: 4407^4414.

WENDLAND, J.C. 1810. Collectio plantarum tarn exoticarum, quam

indigenarum, cum delineatione, descriptione culturaque earum

2: 69, 70, 77, 78. Hannover.

WILLDENOW, K.L. 1800. In C. Linne, Species plantarum 4: 686, 687.

Nauck, Berlin.

■

Bothalia 39,1: 87-96 (2009)

A review of the genus Curtisia (Curtisiaceae)

E. YU. YEMBATUROVA* * +, B-E. VAN WYK* • and P. M. TILNEY*

Keywords: anatomy, Comaceae, Curtisiaceae, Curtisia dentata (Burm.f.) C.A.Sm., revision, southern Africa

ABSTRACT

A review of the monotypic southern African endemic genus Curtisia Aiton is presented. Detailed studies of the fruit and

seed structure provided new evidence in support of a close relationship between the family Curtisiaceae and Comaceae.

Comparisons with several other members of the Comales revealed carpological similarities to certain species of Comus s.l .,

sometimes treated as segregate genera Dendrobenthamia Hutch, and Benthamidia Spach. We also provide information on the

history of the assegai tree, Curtisia dentata (Burm.f.) C.A.Sm. and its uses, as well as a formal taxonomic revision, including

nomenclature, typification, detailed description and geographical distribution.

INTRODUCTION

Curtisia Aiton is a monotypic genus traditionally placed

in the family Comaceae. It is of considerable interest

because of the many uses of its timber and bark — but no

recent reviews of the morphology, taxonomy or anatomy

are available. Recent cladistic and molecular systematic

studies have revealed new evidence of relationships at fam-

ily level (Murrell 1993; Xiang et al. 1993; Noshiro & Baas

1998; Xiang & Soltis 1998; Xiang 1999), including support

for a separate family, Curtisiaceae, or a close relationship

with the family Grubbiaceae, even resulting in a proposal

for extending it to also include Grubbia P.J.Bergius (Xiang

et al. 2002). As no taxonomic revision of Curtisia has ever

been published since its original description in 1789, we

present here a formal taxonomic treatment of this inter-

esting southern African endemic genus. A further aim was

to investigate the structure of the fruit and seeds to allow

comparisons with putative relatives.

MATERIALS AND METHODS

Young branches with leaves and fruits at different

stages of maturation were fixed in FAA (for a minimum

of 24 hrs). The voucher specimen is Van Wyk & Yem-

baturova 4218 (JRAU), collected at Diepwalle Forest

(3323CC), Knysna District, Western Cape, South Africa.

Some Curtisia fruits and seeds were also obtained from

the carpological collections of LE and STU (collec-

tion of Felix Hohmann). Suitable portions were treated

according to the method of Feder & O’Brien (1968) for

embedding in glycol methacrylate (GMA). Specimens

from the following herbaria were examined: JRAU,

NBG, PRE and STU. Herbarium acronyms are given

after Holmgren et al. (1990).

A Porter-Bliim ultramicrotome was used to cut trans-

verse sections of the fruits from 3 to 5 pm thick, and

the periodic acid-Schiff/toluidine blue staining method

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

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

+ Present address: Department of Botany, Russian State Agrarian Uni-

versity— K.A. Timiryazev MSKHA, ul. Timiryazevskaya, 49 Moscow

127550, Russian Federation.

• Corresponding author e-mail address: bevanwyk@uj.ac.za; tele-

phone: 027 011 559 2412.

MS. received: 2008-03-28.

(Feder & O’Brien 1968) was applied. Suitable sections

were photographed. Fruits obtained from carpological

collections were rehydrated and then softened by means

of prolonged heating in Strassburger mixture (water,

glycerol and 96 % ethyl alcohol in equal proportions), in

accordance with traditional anatomical procedures (Pro-

zina 1960) and then sectioned either by hand or sledge

microtome. Test-reactions to identify lignification (phloro-

glucine and concentrated HC1), cutin and fatty substances

(Sudan III, IV), starch (I-KI) and protein (biuret test with

5 % copper sulphate and 50 % KOH) were applied to the

sections. The terminology used is illustrated in the fig-

ures.

RESULTS AND DISCUSSION

Vegetative morphology

The plants are tall shrubs or medium-sized trees 8 to

13 m high (Figure 1A) although specimens of up to 20 m

are not uncommon (Coates Palgrave 2002). Young twigs

are densely tomentose; the hairs are rusty brown to dark

brown but become sparser and greyish with age. The bark

is initially brownish and smooth; in older trees it becomes

darker, tinted with grey and rough with square fissures

(Figure ID, E). The leaves are opposite, simple, petiolate

and lack stipules. They are elliptic to ovate-lanceolate,

130-150 x 50-70 mm, leathery, bright to dark green and

glossy above (but sometimes sparsely pubescent, mostly

along the midrib, with simple, straight, thin hairs) and

markedly reddish tomentose below (tending to become

somewhat glabrescent with age). The lamina is entire,

dorsiventral, with its apex broadly tapering to rounded

and abruptly pointed; the base is mostly broadly tapering,

less often cuneate; the margins are usually slightly revo-

lute and are markedly dentate to serrate or almost sinuate.

The venation is pinnate and markedly raised on the abax-

ial surface (Figure 1C). Stomata are anomocytic. Petioles

are up to 30 mm long and rusty tomentose when young

but may become sparsely hairy with age. Bud scales are

brown or rust-brown and very densely pubescent.

Reproductive morphology and anatomy

Floral morphology and vasculature was thoroughly

investigated by Eyde (1967, 1988). The small flowers of

Curtisia are arranged in terminal, rather large (up to 120

Bothalia 39,1 (2009)

mm long) panicles (Figure IB). Sometimes second-order

inflorescences are formed of flowers clustered together,

giving a somewhat capitate appearance. The peduncles

and pedicels are covered with dense yellowish hairs. The

opposite primary branches of the panicles are subtended

by pronounced, markedly pubescent bracts (Cannon

1978). As is common for Comaceae, floral parts are in

fours (visible in Figure IB). The petals are twice as long

as the calyx tube, oblong in shape; the stamens are equal

to petals in length. A characteristic centrally located

vascular bundle in the four-loculed inferior ovary was

described by Eyde (1967) who suggested the removal of

Curtisia from Comaceae on the basis of this feature.

Curtisia fruits (Figure 1C) are usually referred to as

drupes (or ‘drupaceous’) (e.g. Dyer 1975; Leistner 2000)

but this term is imprecise. True drupes are found only in

the subfamily Pmnoideae of the family Rosaceae where

the fruit develops from a monocarpous gynoecium

(Shibakina 1984; Levina 1987). In members of Cor-

nales, as well as many other taxa, the fruits are syncar-

pous (with two or more fused carpels) and often pseu-

domonomerous (with only one of two or more locules

and seeds reaching full development). Therefore, the

term ‘pyrenarium’, proposed by Z. Artyushenko (Arty-

ushenko & Fyodorov 1986) for such fruits as those of

Curtisia , will be used. The term ‘pyrene’ applies to fruits

such as those of Ilex L. and some Araliaceae, where each

seed is surrounded by a lignified endocarp (each called

a pyrene). However, when the endocarp is fused into a

single bony structure around the seeds, the term ‘fused

pyrenes’ or ‘pyrenarium’ is preferable. The pyrenaria

of Curtisia are small (10-15 mm in diameter), globose,

subglobose or rarely ovoid, snow-white and sometimes

with ± persistent calyx teeth (Figure 1C). The outer peri-

carp tissues are fleshy (Figure 2 A) and comprise both

hypanthial exocarp and mesocarp, the latter developed

from both hypanthial and carpellary tissues. The heav-

ily lignified endocarp constitutes a four-loculed stone,

usually with a fully developed seed in each. Only the

very central area, where the vascular bundle is located,

is not lignified (Figure 2F, arrow). The seeds are exotes-

tal (see description of the seed coat later on), with copi-

ous endosperm and a well-developed, small, centrally

located embryo.

The structure of the fruit wall and the seed was stud-

ied in detail. The exocarp is formed by a single layer

of radially elongated cells with heavily thickened and

cutinized walls (Figure 2B). The mesocarp is primarily

parenchymatous. A hypoderm is clearly visible (Fig-

ure 2B, K) as the outermost zone of the mesocarp, with

two to three layers of minute cells which are flattened in

the tangential direction and sometimes have an almost

FIGURE I . Morphology of Curtisia dentata: A, growth form (cultivated tree at Kirstenbosch Botanical Garden); B, inflorescence with flowers

(note densely tomentose vestiture); C, fruits (note white colour) and leaf surface; D, E, bark. Scale bars: A, 1 m; B, 6 mm; C, 10 mm; D, E,

40 mm. Photographs by B-E. van Wyk.

Bothalia 39,1 (2009)

89

FIGURE 2. — Fruit and seed anatomy in c/s: A— I, K, Curtisia dentata; J, L, Dendrobenthamia japonica (Sieb. et Zucc.) Hutch. A, fleshy part of peri-

carp; B, exocarp (note cell wall cutinization) and hypoderm (outermost zone of mesocarp); C, mesocarp sclereids; D, derivatives of mesocarp

vascular bundles and transitional zone between mesocarp and endocarp; E, endocarp sclereids; F, endocarp with seed-bearing locules and

central vascular bundle (arrow); G, seed coat; H, seed coat vascular bundle; I, embryo and endosperm cells; J, K, exomesocarp with scattered

sclereids (note similarity in structure); L, exocarp cells. Scale bars: A, J, K, 0.2 mm; B-E, G-I, L, 0.07 mm; F, 0.7 mm.

tabular arrangement. Most of the mesocarp consists of

loosely arranged, large, thin-walled parenchymatous cells

of varying shape. These cells form up to 16 layers. Small

sclereids (Figure 2C) can be found scattered through-

out the entire mesocarp (solitary or in clusters of two to

four). Also scattered in the mesocarp are the derivatives

of vascular bundles (Figure 2A, D, arrow); they are fairly

large but usually only slightly lignified. The inner zone of

the mesocarp is composed of two or three layers of small,

thin-walled cells which are elongated tangentially.

90

Bothalia 39,1 (2009)

There is a very conspicuous intermediate zone between

the mesocarp and endocarp, formed by five to seven (or

up to nine) layers of small, tangentially elongated cells

filled with brightly coloured tanniniferous substances

(dark zone in Figure 2A, D). These cells possess slightly

thickened walls that are somewhat lignified.

The typically woody endocarp comprises 17 to 20

layers of fairly small sclereids (Figure 2E), mostly ori-

entated parallel to the longitudinal fruit axis, except for

the innermost layer(s), which can be variously orientated

to form a ‘lining’ layer for each locule. Some of the scle-

reids appear to contain tannins.

The seed coat of Curtisia was previously incom-

pletely described as ‘thin and filmy, made of compressed

and partially obliterated cells’ (Trifonova 2000). How-

ever, our study revealed several interesting structural

details. Firstly, the exotesta is composed of fairly large,

tangentially elongated cells, which appear oval or almost

crescent-shaped in cross section (due to concave outer

tangential walls) (Figure 2G). These cells are filled with

tanniniferous substances and have walls consisting of

cellulose. In certain parts of the seed coat, large scleri-

fied vascular bundle derivatives are found (Figure 2H),

surrounded by a few layers of very small, compressed

parenchymatous cells. This type of seed coat can be clas-

sified as ‘exotestal’ because of the pronounced, thick-

walled outer cells.

The meso- and endotestal components of the seed

coat largely consist of compressed and deformed cells

of irregular shape, sometimes only with their walls per-

sisting. The part of the endotesta which lies against the

endosperm is homogenous and the cell walls are indis-

tinguishable and cutinized.

The seed has a massive copious endosperm, with pro-

tein as the main ergastic substance; the minute embryo is

located centrally within the endosperm (Figure 21).

Phylogenetic relationships

The relationships of Curtisia have not yet been clari-

fied. Its position within the Comaceae had been stable

for a long time (Harms 1898; Wangerin 1910; Philipson

1967; Cronquist 1981; Thome 1992) but its isolated posi-

tion within the family was noted — hence the subfamily

Curtisioideae of Harms (1898). Evidence had gradually

accumulated (Adams 1949; Eyde 1967; Yeramyan 1971)

for separating it as a monotypic family Curtisiaceae

Takht. (validated by Takhtajan 1987) but still within

Comales. This followed an earlier trend of splitting the

broad family concept of Harms (1898) by elevating most

of the 15 genera to family status, e.g. Davidia Baill. to

Davidiaceae (Li 1954), Mastixia Blume to Mastixiaceae

(Calestani 1905 — cited by Takhtajan 1987) and Toricel-

lia DC. to Toricelliaceae (Hu 1934 — cited by Takhtajan

1987).

Detailed molecular systematic studies by Xiang and

co-authors (Xiang et al. 1993, 1998, 2002; Xiang &

Soltis 1998; Xiang 1999; Fan & Xiang 2003) showed

that Curtisia is sister to another southern African

endemic genus Grubbia, and that the combined clade is

again sister to the rest of the Comales. A suggestion by

Xiang et al. (2002) to expand the Gmbbiaceae to include

Curtisia (citing similarities such as the geographical

distribution, woody habit, leathery simple leaves with

revolute margins, opposite phyllotaxy, lack of stipules,

hermaphroditic flowers, inferior ovary and copious

endosperm) was not followed in later treatments (e.g.

Angiosperm Phylogeny Group II 2003; Heywood et al.

2007) where the two monotypic families were retained.

According to Xiang et al. (2002), ‘no apparent morpho-

logical synapomorphies can be found for the two genera

at present’ and there are indeed important differences

between them. The distribution of Curtisia extends along

the southern and eastern coast of South Africa north-

wards as far as Mozambique (coastal zone — Da Silva

et al. 2004) and eastern Zimbabwe (Baker 1911-1912;

Eyles 1917), whereas Grubbia is restricted to fynbos.

The large, leathery leaves of Curtisia are coarsely den-

tate or sometimes sinuate and only slightly revolute,

whereas the small, almost ericoid leaves of Grubbia spe-

cies have entire margins which are distinctly revolute.

There are also important morphological differences in

reproductive structures — inflorescences, flowers and

especially the fruits. Xiang et al. (2002) ascribed these

differences to ancient divergence. It is possible that a

more detailed investigation of the anatomy of Grub-

bia fruits and seeds could shed some light on the Curti-

sia-Grubbia relationship, but hitherto this appears to be

based only on molecular evidence.

Carpological research of Comales and allies by Yem-

baturova (2001, 2002), the comparative results of which

are summarized in Table 1 , showed a number of basic

structural traits that appeared to be of diagnostic value.

Curtisia resembles Cornus L. s.l., Nyssa L., Alangium

Lam. and Corokia A.Cunn. most closely, and Davidia

and Mastixia to a lesser extent. All of these taxa have

epigynous flowers (however, this is still question-

able for Davidia), typical pyrenaria, thick, succulent

exomesocarps and well-developed woody endocarps.

Curtisia, however, differs from all other taxa in having

stones with all four locules similar in size and shape,

each containing a fully developed seed. The other taxa

usually have only a single fully developed locule and

seed. There are also differences in some finer structural

details. Alangium and Mastixia, for example, have con-

spicuous secretory structures in their fleshy mesocarp

tissues which are absent in Curtisia. Species of Nyssa

have either no sclereids or far more mesocarp sclereids

than Curtisia. Furthermore, Curtisia is the only genus

with a centrally located vascular bundle in the ovary;

other putative Comaceae relatives are reported to have

transseptal bundles (Eyde 1 967).

As is well known, the Comaceae have a rich fossil

record which includes wood, leaves and especially fruits

(Miki 1956; Eyde & Barghoom 1963; Eyde et al. 1969;

Mai & Walther 1978; Arbuzova 1988; Eyde 1988; Eyde

& Xiang 1990). Most of the fossil structures have been

identified as belonging to the genera Davidia, Mastixia,

Diplopanax Hand.-Mazz., Nyssa, Alangium and Cornus.

Recently, a fossil species of the Eocene London Clay

flora, originally referred to as Leucopogon quadrilocu-

laris Reid & Chandler or Cornus quadrilocularis (Reid

& Chandler) Chandler was shown to belong to Curtisia

(Manchester et al. 2007) on the basis of the endocarp

structure. The fossil endocarps of Cornus quadrilocula-

Bothalia 39,1 (2009)

91

ris are composed of slightly elongated sclereids and pos-

sess four seed-bearing locules and a vascular bundle in

the centre, as in extant Curtisia.

The greatest degree of resemblance to Curtisia in

pericarp structure (Figure 2J, L, with a portion of Cur-

tisia pericarp given in Figure 2K for comparison) is

shown by two representatives of the genus Comus s.l.,

currently segregated into the genera Dendrobenthamia

(Asian dogwoods with compound fruits) and Bentha-

midia (American dogwoods with fruits clustered together

without fusing). These three taxa all have radially elon-

gated exocarp cells with heavily cutinized walls, solitary

or clustered sclereids scattered throughout the parenchy-

matous mesocarp and a homogeneous woody endocarp.

It is noteworthy that scattered sclereids were previously

treated as an adaptation to synzoochorous dispersal in the

relatively large, compound fruits of Dendrobenthamia,

believed to be dispersed by monkeys (Eyde 1985). Cur-

tisia fruits, despite their bitter taste, are also dispersed by

animals such as birds (loeries, rameron pigeons), wild

pigs, monkeys and less often baboons (Watt & Breyer-

Brandwijk 1962). It seems likely that Curtisia fruits

may be oil-yielding, as many white-fruited dogwoods

from Comus s.l. [e.g. Swida alba (L.) Opiz, S. sericea

(L.) Holub and other shrubby species] have fruits rich in

oil (Sozonova 1992). These fruits look white because of

the oil droplets that reflect light. Thus, fruit anatomy has

hereby provided evidence of a possible Curtisia-Comus

affinity and therefore supports Takhtajan’s (1987) treat-

ment of Curtisia as a monotypic family Curtisiaceae

within the order Comales, close to the core family Cor-

naceae. The latter is currently treated as consisting only

of Comus s.l. but it has often been divided into several

segregates (Takhtajan 1987, 1997).

TAXONOMY

Curtisia Aiton, Hortus kewensis 1: 162 (Aug-

Oct. 1789) nom. cons.; Thunb.: 100 (1792); Thunb.: 28

(1794); Thunb;: 141 (1823); Willd.: 687 (1797); Roem.

& Schult.: 294 (1818); Spreng.: 442 (1825); DC.: 12

(1825); Harv.: 143 (1838); Endl.: 799 (1839); Lindl.:

783 (1847); Harv.: 579 (1862); Hook.f.: 949 (1867);

Baill. : 3: 334 (1863), 163 (1879a), 253 (1879b); Harms:

262 (1898); Wangerin: 29 (1910). Type: C.faginea Aiton

[now C. dentata (Burm.f.) C.A.Sm.].

Note: the generic name Curtisia Aiton was conserved against the

earlier homonym Curtisia Schreber, Apr. 1789 (family Rutaceae)

and is listed in the International Code of Botanical Nomenclature

(McNeill et al. 2006). Both these generic names were given in honour

of William Curtis, the founder and for a long time the proprietor of the

famous Curtis’s Botanical Magazine , published since 1786 and known

as Botanical Magazine (Burtt Davy 1932; Marais 1985).

Trees, medium-sized to large, up to 20 m high, or tall

shrubs. Bark brown and smooth when young, dark brown

and square-fissured with age. Young branchlets with red-

dish or rust-brown hairs. Leaves simple, opposite; petiole

up to 20(-30) mm long, pubescent; lamina oblong-elliptic

to ovoid, 25-10 x 25-75 mm, penninerved, leathery, mar-

gins dentate, revolute to sinuate, glabrous on upper surface

and rust-brown pubescent beneath. Inflorescences termi-

nal, paniculate, loose or compact, inflorescence branches

tomentose; bracts conspicuous, linear-lanceolate, rust-

brown tomentose, subtending opposite primary inflores-

cence branches. Flowers minute, sessile or shortly pedicel-

late, scentless, hermaphrodite, epigynous, tomentose. Calyx

tube 4-lobed, adnate to ovary. Petals 4, cream-coloured

or white with a tint of purple, valvate. Androecium didy-

mous; stamens 4, alternating with petals; filaments subu-

late; anthers short. Gynoecium syncarpous; ovary 4-locular

(occasionally 3-locular); ovules pendulous, one per locule.

Fruit globose, snow-white, drupaceous (pyrenarium), stone

TABLE 1. — Comparative carpological traits in genera of Comaceae sensu Harms (1898) [From Yembaturova (2001), translated from Russian]

92

Bothalia 39,1 (2009)

(3)4-locular, (3)4-seeded, with a central vascular bundle.

Seeds exotestal. Endosperm copious. Embryo minute,

centrally located. Flowering time: late October to March.

Fruiting time May to October. Chromosome number n = 1 3

(Goldblatt 1978). Conservation status: Low Risk. Figure 1.

The species of Curtisia

There is only one extant species in the genus (see

generic description given above).

Curtisia dentata (Burm.f) C.A.Sm., in Journal of

South African Forestry Association 20: 50, t. 170 (1951);

Cannon: 635 (1978). Type: South Africa, Sideroxylum

dentatum J.Burm., Rariorum Africanarum plantarum 235,

t. 82 (1738), iconotype (Figure 3).

Sideroxylon dentatum Burm.f.: 6 (1768).

Curtisia faginea Aiton: 162 (1789); Willd.: 687 (1797); Roem. &

Schult.: 294 (1818); Thunb.: 141 (1823); Spreng.: 442 (1825); DC.:

12 (1825); Pappe: 17 (1854); Harv.: 570 (1862); Wangerin: 30 (1910);

Baker f.: 76 (1911-1912); Eyles: 435 (1917); Burtt Davy: 512 (1932);

Steedman: 60 (1933). Type: South Africa, introduced by Francis Mas-

son in 1775 (Banks Herbarium, sheet BM000794113, BM!).

Illustrations: Burm.: t. 82 (1738); Lam.: t. 71 (1823); Sim: t. 77

(1907); Wangerin: 30 (1910); Burtt Davy: 513 (1932); Cannon: t. 170

(1978). There is a beautiful unpublished painting of Curtisia dentata

by Clemenz Heinrich Wehdemann (1762-1835) in the repository of

Natural History Museum (BM). This original watercolour forms part of

the Wehdemann collection entitled ‘Sketches of plants growing about

Plettenburg Bay on the coast of Africa’.

History and uses

The most well-known name for Curtisia is assegai,

assegai-tree (Afrikaans: assegaaiboom) or assegai-

wood ( assegaaihout ). The records for this name date

back to the earliest colonial days, recorded, according

to Smith (1966), by Hermann in 1672 and Burmann in

1692. These plant names apparently were the first cases

of modem spelling of the name. Early writers used such

versions as hazegaiejn boomen hout, assghaij, asseg-

aje and hassagay-hout. As far as the origin of the name

assegai/ assagay is concerned, several versions exist. One

of them is that the name is derived through Portuguese

from the Latin word hast a, meaning a spear (Palmer &

Pitman 1972). Another idea is that it refers to the Arabic

word Azzaghayah, adopted by the Portuguese with vari-

ous spelling modifications and taken over by early Dutch

writers (Smith 1966), evolving to its present-day form

with time but always referring to weapons (spears and

bows) used by the native African people. The ethnobotani-

cal importance of the tree is reflected in the large number

of vernacular names recorded in southern Africa. In Table

2, all these are listed exactly as they were published.

The wood of Curtisia is iron-hard but elastic, strong

and very durable — traditional javelins and spears, with

1.8 m-long tapering shafts, were made from this timber.

Thunberg, in his descriptions of his travels in the Cape

of Good Hope between 1772 and 1775 (Forbes 1986),

repeatedly mentioned Curtisia (as ‘assagay wood’) and

its practical use for ‘assagays’ (javelins that Khoikhoi

people carried with them on their journeys, with which

they ‘defended themselves against their enemies and

wild beasts, and were able to kill them, buffaloes and

other animals’). Later on, many other uses were found.

The fact that assegai wood shrinks less than most

other woods, made it very useful for the early settlers.

It was the preferred timber for the axles and poles for

wagons as well as the spokes, naves and felloes of the

wheels. The highly decorative qualities of Curtisia tim-

ber— resembling mahogany due to developing a reddish

brown, cinnamon tint with age — were appreciated by

craftsmen producing tool handles (hammer handles in

particular), implements of husbandry, furniture and floor

covering (Watt & Breyer-Brandwijk 1962; Smith 1966;

Palmer & Pitman 1972; Van Wyk et al. 2000). The bark

was used for tanning, yielding 2.98 to 14.05 % of tan-

nin; up to 4 % of tannin was yielded by the leaves and

twigs of the plant (Watt & Breyer-Brandwijk 1962; Van

Wyk et al. 1997). Curtisia is also an attractive ornamen-

tal tree, especially for moist gardens.

Curtisia dentata is important in traditional medicine

(Arnold et al. 2002). Zulu people use the bark to treat

diarrhoea and stomach ailments. It serves as an aphro-

disiac (a ‘love charm’ to make a man attractive) and as

a ‘blood purifier’ or ‘strengthener’ (Cunningham 1988;

Hutchings et al. 1996; Van Wyk et al. 1997; Ngwenya et

al. 2003). Since the tannin-rich bark is used for medici-

nal purposes, the debarking and ringbarking of trees

caused significant damage in some localities. The bark

(Figure IE) used to be sold by traditional Zulu herb

gatherers for R30 per 50 kg bag but after becoming

scarce due to overexploitation, it is only included in spe-

cial bark mixtures, such as ‘Special Khubalo’ (Cunning-

ham 1988). Rapid coppicing and the apparent capability

of the bark to regrow, played a very important role in the

survival of this forest tree. Despite a broad phytochemi-

cal survey of the Comaceae by Bate-Smith et al. (1975),

the chemical rationale for the traditional uses is still

poorly understood. It may be assumed, however, that the

high tannin content contributes to the efficacy in treating

diarrhoea and stomach ailments.

Geographic distribution

Curtisia dentata is restricted to southern Africa, pri-

marily along the southern and the eastern coasts of

South Africa and extends into Swaziland, Mozambique

and eastern Zimbabwe (Figure 4). The genus is confined

to montane forests (mainly found on southern and south-

western slopes), at altitudes from sea level to 2 300 m.

SPECIMENS EXAMINED

Anon. 9701 (PRE). AM. W. 1005 (NBG). Acocks 8992 (PRE). Airy

Shaw 3614, 3736 (PRE).

Bakker 281 (PRE). Balkwill 1474 (PRE). Balsinhas 02576 (PRE).

Bayliss 127. 6169 (PRE, 2 sheets); 466, 1328, 1490 (PRE). Bazer 809

(PRE). Bird 1042, 1213, 1241, 1396, 1412 (PRE). Bos 965 (NBG,

PRE); 1314 (NBG). Botha 0007 (PRE). Botha & Cilliers 3636 (PRE).

Boucher G19, 322, 2078 (PRE). Bredenkamp 1511 (PRE). Britten 1333

(PRE). Burgers 1111 (PRE). Burgoyne 1726 (PRE); 6206 (NBG, PRE).

Burtt Davy 1262, 2439, 2442 (PRE).

Cheadle 737 (PRE). Codd 3050 (PRE). Codd & Dyer 9117 (PRE).

Comins 933 (PRE). Compton 28675, 32241 (PRE); 29500 (NBG,

PRE). Curson & Irvine 86 (PRE, 2 sheets).

Dahlstrand 2661 (PRE). Davis 56 (PRE). De Winter 8266A (PRE).

Dlamini s.n. (PRE). Dohse 140 (PRE). Duthie 632 (NBG). Dyer 4850

(PRE).

Eastwood 2430 (PRE). Ecklon & Zeyher 558 (NBG, 2 sheets); 596

(NBG); s.n. (PRE). Edwards 1450, 3311 (PRE).

Bothalia 39,1 (2009)

93

FIGURE 3. — Iconotype of Curtisia den-

tata [Burmann: plate 82 (1738)].

Fisher 1105 (PRE). Flanagan 143 (PRE). Forest Dept. s.n. sub

STEU18857 (PRE).

Galpin 4229, 11612, 13676 (PRE). Gertzner 4387, 4488, 4499 (PRE).

Giffen G849 (PRE). Gillet 72, 1311 (NBG). Glen 2413 (PRE). Goodier

& Phipps 157 (PRE). Grant 3263 (PRE). Guillarmod 9314 (PRE).

Hemm 40, 522 (PRE). Henkel 2545 (PRE). Herman & Prior 322

(PRE). Hoffmann 171 (PRE). Humbert 9675 (PRE).

Jacobsen 1812, 4679 (PRE). Jones & Leach 2531 (PRE).

Kemp 1200 (PRE). Kerfoot 6122 (PRE, 2 sheets; STU). Kerfoot,

Forrester & Gooyer 131 (PRE). Killick 866, 1900 (PRE); s.n. (PRE,

photo!). Kluge 1358 (PRE).

Legal 2442 (PRE). Liebenberg 7955A (PRE).

MacOwan 731 (PRE). Maguire 1438 (NBG). Marloth 524 (PRE);

5246 (NBG). Meyer 1927, 2610 (PRE). Miller S/26 (PRE). Muir 117

(PRE). Mogg 17209, 34609 (PRE). Mohle 416 (PRE). Moll 1789

(PRE). Morze 2010 (PRE). Mullin 16/52 (PRE). Mullins s.n. (PRE, 2

sheets). Muller 487 (PRE). Muller & Scheepers 118 (PRE).

Nicholson 658 (PRE). Nienaber EN162 (PRE).

Obermeyer 890, 1179 (PRE); 2008 (NBG, PRE). Osborne 30 (PRE).

Paterson 2109 (PRE). Pegler 1332, 4606 (PRE); s.n. (NBG). Phillips

s.n. (PRE). Pillans 3664 (PRE). Plantk. Hons. 477 (JRAU). Pole Evans

3619, 15490, s.n. (PRE). Prosser s.n. (PRE).

Raal 226 (PRE). Ranger 290 (PRE). Reid 4063 (PRE). Reyneke 189

(PRE). Rodin 3147, 4104 (PRE). Rogers 23038 (PRE). Rudatis 1776

(PRE).

Scharf 1784 (PRE). Scheepers 832 (PRE). Schutte 29 (JRAU).

Sebothoma 334, 346 (PRE). Sim 2020 (PRE). Smith 40 (PRE). Smuts

11, 98 (PRE); 1094 (PRE, 2 sheets): s.n. (NBG). Smuts & Gillet 2262

(NBG, PRE); 3600 (PRE). Stalmans 335, 1164 (PRE). Stokoe s.n.

(NBG). Stork 61580 (PRE). Story 1665 (PRE). Strey 9424, 10738

(PRE). Symons 504, 15736 (PRE).

Thode A273 (PRE); A844 (NBG, PRE); A845 (PRE, 2 sheets). Thorn-

croft 1005 (PRE). Topper 97 (NBG). Torre & Pereira 12.738 (PRE).

Uys s.n. (PRE).

94

Bothalia 39,1 (2009)

TABLE 2. — Vernacular names for Curtisia dentata. All names given in exactly same form as originally published

Van der Merwe 1309 (PRE). Van der Schijff 4642, 4675, 5121 (PRE).

Van Son s.n. (PRE). Van Warmelo s.n. (PRE). Van Wyk 4110, 7005, 7528

(PRE). Van Wyk & Theron 4564 (PRE). B-E. & C.M. van Wyk 28 (JRAU).

Van Wyk & Yembaturova 4218 (JRAU). Von Ludwig s.n. (STU)

Wager 23012 (PRE). Ward 3422 (PRE). Wells 2996 (PRE). West 3012,

3814 (PRE). Williams 2418 (NBG, PRE).

Zeyher 558, 7699 (PRE).

ACKNOWLEDGEMENTS

The authors offer their thanks to the University of

Johannesburg and German Academic Exchange Service

(DAAD) for funding the research. Also, we are much

indebted to A.R. Magee (University of Johannesburg),

Dr A. I. Konstantinova (Moscow State University), Dr

A. Wiirz (STU), Anne Lise Fourie and Estelle Potgieter

(both of the Mary Gunn Library, PRE) for valuable help

during the preparation of the manuscript. We thank V.

Papworth (BM) for kindly providing us with high resolu-

tion images of type specimens in the Banks Herbarium,

and the curators and staff of PRE and NBG for allowing

us to study their collections.

REFERENCES

ADAMS, J.E. 1949. Studies in the comparative anatomy of the Cor-

naceae. Journal of the Elisha Mitchell Scientific Society 65:

218-244.

AITON, W. 1789. Hortus kewensis 1 : 162. Nichol, London.

ANGIOSPERM PHYLOGENY GROUP II. 2003. An update of the

Angiosperm Phylogeny Group classification for the orders and

families of flowering plants: APG II. Botanical Journal of the

Linnean Society 141: 399-436.

ARBUZOVA, O.A. 1988. K paleokarpologii Nyssa L. [On the pal-

aeocarpology of Nyssa L.] Trudy Leningradskoi molodyozhnoi

konferentsii botanikov. [Proceedings of Leningrad conference of

young botanists] 1: 194—204. [In Russian]

ARNOLD, T.H., PRENTICE, C.A., HAWKER, L.C., SNYMAN, E.E.,

TOMALIN, M., CROUCH, N.R. & POTTAS-BIRCHER, C.

2002. Medicinal and magical plants of southern Africa: an anno-

Bothalia 39,1 (2009)

95

FIGURE 4. — Known distribution of Curtisia dentata, •, in South Afri-

ca, Swaziland, Mozambique and Zimbabwe. Additional locali-

ties given by Von Breitenbach (1995), ▲.

tated checklist. Strelitzia 13; 53. National Botanical Institute,

Pretoria.

ARTYUSHENKO, Z.T. & FYODOROV, A.A. 1986. Atlas po

opisatel'noi morfologii vysshikh rastenii: plod [Atlas on descrip-

tive morphology of higher plants: the fruit]: 110-143. Nauka,

Leningrad. [In Russian]

BAILLON, H.E. 1863. Remarques sur l’organisation florale de queques

Bruniacees et sur les affinites du genre Grubbia. Adansonia 3:

318-335.

BAILLON, H.E. 1879a. Adansonia: recueil d 'observations botaniques

/ redige par le Dr H. Baillon 12: 163. Baillon & Savy, Paris.

BAILLON, H.E. 1879b. Histoire naturelle des plantes 7: 253. Imprime-

rie Nationale, Paris.

BAKER, E.G. 1911-1912. A contribution to the flora of Gazaland.

Journal of the Linnean Society (Botany) 40: 76.

BATE-SMITH, E.C., FERGUSON, I.K., HUTSON, K„ JENSEN, S.R.,

NIELSEN, B.J. & SWAIN, T. 1975. Phytochemical interrela-

tionship in the Comaceae. Biochemical Systematics and Ecology

3: 79-89.

BURMANN, J. 1738. Rariorum Africanarum plantarum: 235, 236.

Boussiere, Amsterdam.

BURMANN, N.L. 1768. Flora indica: cui accredit series zoophytorum

indecorum, nec non prodromus florae capensis: 6. Haak, Leiden,

& Schreuder, Amsterdam.

BURTT DAVY, J. 1932. A manual of the flowering plants and ferns of

the Transvaal with Swaziland, South Africa, part 2: 512, 513.

Longmans, Green, London, New York, Toronto.

CANNON, J.F.M. 1978. Comaceae. Flora zambesiaca 4: 635-638.

Flora Zambesiaca Managing Committee, London.

COATES PALGRAVE, M. 2002. Keith Coates Palgrave Trees of south-

ern Africa, edn. 3: 858, 859. Struik, Cape Town.

CRONQUIST, A. 1981 .An integrated system of classification offlower-

ing plants: 668-670. Columbia University Press, New York.

CUNNINGHAM, A.B. 1988. An investigation of the herbal medicine

trade in Natal/KwaZulu. Investigational Report No. 29. Institute

of National Resources, University of Natal, Pietermaritzburg.

DA SILVA, M.C., IZIDINE, S. & AMUDE, A.B. 2004. A preliminary

checklist of the vascular plants of Mozambique: 49. South Afri-

can Botanical Diversity Network Report No. 30. SABONET,

Pretoria.

DE CANDOLLE, A.P. 1 825. Prodromus systematis naturalis regni veg-

etabilis 2: 12. Treuttel & Wiirtz. Paris.

DE LAMARCK, J.B.A.P. 1823. Recueil de planches de botanique de

I’encyclopedie 1: t. 71. Imprimeur-Libraire, Paris.

DYER, R.A. 1975. The genera of South African flowering plants 1 —

Dicotyledons: 429. Department of Agricultural Technical Ser-

vices, Pretoria.

ENDLICHER, S.L. 1839. Genera plantarum secundum ordines natura-

les disposita: 799. Beck, Vienna.

EYDE, R.H. 1967. The peculiar gynoecial vasculature of Comaceae