Both alia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 41,1

May 2011

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BIODIVERSITY INSTITUTE PRETORIA

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BOTHALIA

Botha/ia 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 palaeoflora 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. I . 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 41,1

Scientific Editors: G. Germishuizen, O.A. Leistner

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

May 2011

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

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

Braun, Ms K.P. P.O. Box 174, Mbabane HI 00, Swaziland.

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

Figueiredo, Dr E. University of Coimbra, 3001-455 Coimbra, Portugal / Nelson Mandela Metropolitan

University, Port Elizabeth, RSA.

Forster, Dr P.L, Queensland Herbarium, Brisbane Botanic Gardens, Australia.

Geerinck, Dr D. Rue Charles Pas 4, B-1160, Brussels, Belgium.

Henderson, Ms L. Agricultural Research Council, Private Bag XI 01, Pretoria, RSA.

Herman, P.P.J. South African National Biodiversity Institute, Pretoria, RSA.

Hutchings, Ms A. P.O. Box 1362, Empangeni, KwaZulu-Natal, RSA.

Isawumi, Dr M.A. Natural History Museum, Obafemi Awolowa University, Ile-ife, Nigeria.

Jongkind, Dr C.C.H. Agricultural University, Wageningen, The Netherlands.

Klopper, Mrs R.R. South African National Biodiversity Institute, Pretoria, RSA.

Lammers, Prof. T.G. University of Wisconsin, Oshkosh, USA.

Leistner, Dr O.A. 1 94 Griselda Rd, Murrayfield, Pretoria, RSA.

Moffett, Prof. R. Omruil, P.O. Box 326, Kestell, Free State, RSA.

Nagalingum, Dr N.S. University of California, Berkeley, USA.

Nicholas, Prof. A., Howard College, University of KwaZulu-Natal, Durban, RSA.

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

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

Robinson, Dr H. Smithsonian Institution, Washington DC, USA.

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

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

Tilney, Prof. PM. Department of Botany, University of Johannesburg, RSA.

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

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

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

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

Windham, Dr M. Duke University, Durham, North Carolina, USA.

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

Yatskievych, Dr G. Missouri Botanical Garden, St Louis, USA.

Date of publication of Bothalia 40,2: 1 Oct. 2010.

CONTENTS

Bothalia 41,1

1 . Systematics and biology of the African genus Ferraria ( Iridaceae: Irideae). P. GOLDBLATT and J.C. MAN-

NING

2. Annotated catalogue of the flowering plants of Sao Tome and Principe. E. FIGUEIREDO, J. PAIVA, T.

STEVART, F. OLIVEIRA and G.F. SMITH

3. Systematics of the southern African genus Ixia (Iridaceae). 3. Sections Hyalis and Morphixia. P. GOLD-

BLATT and J.C. MANNING

4. A conspectus of Combretum (Combretaceae) in southern Africa, with taxonomic and nomenclatural notes

on species and sections. M. JORDAAN, A.E. VAN WYK and O. MAURIN

5. Generic status of Quisqualis (Combretaceae), with notes on the taxonomy and distribution of Q. parviflora.

M. JORDAAN, A.E. VAN WYK and O. MAURIN

6. Notes on African plants:

Agavaceae. Furcraea foetida : an invading alien in South Africa. N.R. CROUCH and G.F.

SMITH

Apocynaceae. Distribution of Ciyptolepis delagoensis (Periplocoideae-Cryptolepideae), a sub-

continental southern African endemic and selection of a neotype. S.P. BESTER and L. JOU-

BERT

Asphodelaceae: Alooideae. Gasteria croucheri subsp . pondoensis, a new cremnophyte from Pon-

doland. South Africa. N.R. CROUCH. G.F. SMITH and D.G.A. STYLES

Asteraceae. The true identity of Oxylaenci acicularis. M. KOEKEMOER

Asteraceae. Vernonia (tribe Vemonieae) and related genera in southern Africa: updates and correc-

tions. P.P.J. HERMAN and N. SWELANKOMO

Bryophyta. New and interesting records of mosses in the Flora of southern Africa area: 5. New

provincial records. J. VAN ROOY

Campanulaceae. A new species of Wahlenbergia from Western Cape, South Africa. C.N. CU-

PIDO

Iridaceae. Tritonia cedarmontana and T. linearifolia (Crocoideae), two new species from the Cape

Floristic Region; T. lineata var. pan’ifolia included in T. gladio/aris; and the correct author

citation for T. strictifolia. J.C. MANNING and P. GOLDBLATT

Marchantiophyta. New liverwort distribution records in South Africa. J. VAN ROOY, N. PHEPHU

and S.M. PEROLD

Molluginaceae. Adenogranvna natans, a remarkable new aquatic species from Western Cape,

South Africa. J.C. MANNING, P. GOLDBLATT and F. FOREST

Pteridophyta-Marsileaceae. Pilularia dracomontana , a new species of pillwort from the KwaZu-

lu-Natal Drakensberg, South Africa. N.R. CROUCH & J. WESLEY-SMITH

Pteridophyta. New pteridophyte records for the flora of Swaziland. N.R. CROUCH and J.E. BUR-

ROWS

Pteridophyta-Sinopteridaceae. A new subspecies of Cheilanthes deltoidea from Gauteng and Lim-

popo, South Africa. R.R. KLOPPER and A.E. VAN WYK

Pteridophyta-Thelypteridaceae. Metathelypteris burrows iorum, a new species from Swaziland

and a first genus record for southern Africa. N.R. CROUCH

7. The medical ethnobotany of Lesotho: a review. A. MOTEETEE and B-E. VAN WYK

8. Miscellaneous notes:

Tribute to Gerrit Germishuizen, editor of Bothalia. G.F. SMITH

1

41

83

135

161

196

199

183

187

176

188

178

171

185

189

201

181

204

193

209

229

New combinations, names, species, statuses, subgenera and subspecies in Bothalia 41,1 (2011)

Adenogramma natans J.C. Manning & Goldblatt , sp. nov., 191

Cheilanthes deltoidea Kunze subsp. silicicola Klopper & A.E.van Wyk , subsp. nov., 204

Combretum psidioides Welw. subsp. grandifolium (F.Hoffm.) Jordaan, stat. nov., 147

Combretum sylvicola O.Maurin, comb, et nom. nov., 168

Ferraria flava Goldblatt & J.C. Manning, sp. nov., 31

Ferraria macrochlamys subsp. kamiesbergensis (M.P.de Vos) Goldblatt & J.C. Manning, comb, et stat. nov., 36

Ferraria macrochlamys subsp. serpentina Goldblatt & J.C. Manning, subsp. nov., 37

Ferraria ornata Goldblatt & J.C. Manning, sp. nov., 24

Ferraria parva Goldblatt & J.C. Manning, sp. nov., 26

Ferraria spithamea (Baker) Goldblatt & J.C. Manning, comb, nov., 15

Gasteria croucheri (Hook.f) Baker subsp. pondoensis N.R. Crouch, Gideon F.Sm. & D.G. A. Styles, subsp. nov., 183

Ixia alata Goldblatt & J.C. Manning, sp. nov., 121

Ixia cedarmontana Goldblatt & J.C. Manning, sp. nov., 126

Ixia dolichosiphon Goldblatt & J.C. Manning, sp. nov., 126

Ixia ecklonii Goldblatt & J.C. Manning, sp. nov., 118

Ixia linderi Goldblatt & J.C. Manning, sp. nov., 97

Ixia linearifolia Goldblatt & J.C. Manning, sp. nov., 121

Ixia longituba N.E.Br. subsp. macrosiphon Goldblatt & J.C. Manning, subsp. nov., 95

Ixia monticola Goldblatt & J.C. Manning, sp. nov., 115

Ixia parva Goldblatt & J.C. Manning, nom. nov., 116

Ixia pavonia Goldblatt & J.C. Manning, sp. nov., 120

Ixia ramulosa (G.J. Lewis) Goldblatt & J.C. Manning, stat. nov., 117

Ixia recondita Goldblatt & J.C. Manning, sp. nov., 103

Ixia saundersiana Goldblatt & J.C. Manning, sp. nov., 1 1 1

Ixia stenophylla Goldblatt & J.C. Manning, nom. et stat. nov., 129

Metathelypteris burrowsiorum N.R. Crouch, sp. nov., 193

Pilularia dracomontana N.R. Crouch & J. Wesley-Smith, sp. nov., 201

Subgen. Glutinosae (M.P.de Vos) Goldblatt & J.C. Manning, subgen. & comb, nov., 11

Tritonia cedarmontana Goldblatt & J.C. Manning, sp. nov., 171

Tritonia linearifolia Goldblatt & J.C. Manning, sp. nov., 173

Wahlenbergia suffruticosa Cupido, sp. nov., 179

IV

Bothalia 41,1: 1-10(2011)

Systematics and biology of the African genus Ferraria (Iridaceae:

Irideae)

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

Keywords: Ferraria Burm. ex Mill., floral biology, Iridaceae, new species, taxonomy, tropical Africa, winter rainfall southern Africa

ABSTRACT

Following field and herbarium investigation of the subequatorial African and mainly western southern African Ferraria

Burm. ex Mill. (Iridaceae: Iridoideae), a genus of cormous geophytes, we recognize 1 8 species, eight more than were included

in the 1979 account of the genus by M.R de Vos. One of these, F. ovata , based on Moraea ovata Thunb. (1800), was only

discovered to be a species of Ferraria in 2001, and three more are the result of our different view of De Vos’s taxonomy. In

tropical Africa, F. gluiinosa is recircumscribed to include only mid- to late summer-flowering plants, usually with a single

basal leaf and with purple to brown flowers often marked with yellow. A second summer-flowering species, F. candelabrum,

includes taller plants with several basal leaves. Spring and early summer-flowering plants lacking foliage leaves and with

yellow flowers from central Africa are referred to F. spithamea or F. welwitschii respectively.

The remaining species are restricted to western southern Africa, an area of winter rainfall and summer drought. We rec-

ognize three new species: F. flava and F. ornata from the sandveld of coastal Namaqualand, and F. parva, which has among

the smallest flowers in the genus and is restricted to the Western Cape coastal plain between Ganzekraal and Langrietvlei near

Hopefield. Ferraria ornata blooms in May and June in response to the first rains of the season. Among the remaining species,

F. uncinata subsp. macrochlamys is now F. macrochlamys and is treated as comprising three subspecies: subsp. macroch-

lamys from central and northern Namaqualand has leaves with thickened, crisped margins; subsp. kamiesbergensis from the

southern Kamiesberg has leaves with unthickened margins and blades curved in one direction; and subsp. serpentina from

gravels and sands of coastal Namaqualand has serpentine leaves, also with unthickened margins. Among the remaining spe-

cies, F. divaricata subsp. arenosa is now treated as a synonym of F. divaricata. Because of our re-interpretation of the type

of F. divaricata, plants which were called F. divaricata subsp. divaricata and subsp. australis are now treated as synonyms

under the name F. variabilis.

Flowers of Ferraria are unique in Iridaceae in having tepal limbs with crisped margins and a style that divides into flat-

tened branches, each deeply forked w'ith finely fringed adaxial margins. Despite relative floral uniformity, the genus displays

a surprising range of discrete pollination systems for so small a genus. Pollinators include Diptera in the families Muscidae,

Calliphoridae, and Sarcophagidae (F. crispa group); anthophorine and honey bees (F. ferrariola)', eumenid wasps (F. di-

varicata., F. macrochamvs, F. variabilis)', and Coleoptera in the families Meloidae and Melyridae (F. uncinata ). Preliminary

phylogenetic analysis using plastid DNA regions shows F. glutinosa to be sister to an unresolved cluster of southern African

species and confirms as plesiomorphic the open branching habit, many-flowered inflorescences and exserted globose capsules

that characterize F. glutinosa and its immediate allies in subgen. Glutinosa.

CONTENTS

Abstract 1

Introduction 1

Generic relationships and geological age 3

Morphology 3

Chromosome cytology 6

Pollination biology 7

History 8

Phylogeny and evolution 9

Systematics 9

Key to species 10

A. Subgen. Glutinosae M.P.de Vos (spp. 1—4) .... 11

B. Subgen. Ferraria (31 spp.) 17

B 1 . Sect. Ferraria M.P.de Vos (spp. 5-10) 17

B2. Sect. Macroscyphae M.P.de Vos (spp. 1 1 — 18) . 25

Series Subdivaricatae (spp. 11, 12) 25

Series Macroscyphae (spp. 13-15) 27

Series Uncinatae (spp. 16-18) 33

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

P.O. Box 299, St. Louis, Missouri 63166, USA.

** Compton Herbarium, South African National Biodiversity Insti-

tute, Private Bag X7, 7735 Claremont, Cape Town/Research Centre for

Plant Growth and Development, School of Biological and Conserva-

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

Bag X01, Scottsville 3209, South Africa.

MS. received: 2010-02-09.

Excluded species 39

Acknowledgements 39

References 39

INTRODUCTION

When last revised by De Vos (1979), Ferraria Burm.

ex Mill. (Iridaceae: Iridoideae) was treated as compris-

ing 10 species: nine native to the southern African win-

ter rainfall zone and a broadly circumscribed F. gluti-

nosa (Baker) Rendle, widespread across the summer

rainfall interior of southern and tropical Africa. Field

work in central and southern Africa in the past 10 years,

often in conjunction with studies of pollination biology

and examination of herbarium collections, has shown

that the taxonomy of the genus needed major adjust-

ment. Some nomenclatural changes have already been

published. F. ovata (Thunb.) Goldblatt & J.C. Manning

was added to the genus (Goldblatt & Manning 2002)

after the rediscovery in the wild in 2001 of the plant

named Moraea ovata by C.P. Thunberg in 1800. Subspe-

cies macrochlamys of F. uncinata Sweet is now recog-

nized as a separate species (Goldblatt & Manning 2004),

and reinterpretation of the type of F. divaricata Sweet

resulted in plants called F. divaricata subsp. divaricata

and subsp. australis M.P.de Vos being referred to a new

species, F. variabilis Goldblatt & J.C. Manning. Subsp.

Bothalia 41,1 (2011)

PLATE I. - Ferraria : A, uncinala (Clanwilliam, Western Cape) (no. 16); B, macrochlamys subsp. kamiesbergensis (Kliprand, Northern Cape)

(no. 17b); C, folios a (near Elands Bay, Western Cape) (no. 6); D, variabilis (Bushmanland, Northern Cape) (no. 14); E, crispa (Langebaan,

Western Cape) (no. 5); F ,ferrariola (Bitterfontein, Northern Cape) (no. 1 1 ); G, glutinosa (northern Namibia) (no. 1 ); H, ornata (near Groen

River Mouth, Northern Cape) (no. 10); I, parva (Vredenburg, Western Cape) (no. 12); i. flava (near Koekenaap, Western Cape) (no. 15).

Bothalia 41,1 (2011)

3

arenosa of F. divaricata is now referred to F. divaricata

(Goldblatt & Manning 2005) and subsp. aurea M.P.de

Vos is most likely a synonym of the new F.flava.

Our examination of herbarium collections (from BM.

BOL, K, LISC, NBG, PRE and SAM), photographs and

drawings of the tropical African Ferraria glutinosa as

circumscribed by De Vos (1979), has convinced us that

there are at least four clearly defined sets of populations

under this name. Mid- to late summer-blooming popula-

tions with brown or dull purple flowers with yellow tepal

margins and one or two poorly developed basal leaves

match the type of F. glutinosa , which is from Angola. We

refer plants with a fan of up to eight basal leaves and yel-

low flowers sometimes marked with brown, also bloom-

ing in mid- to late summer, to F candelabrum (Baker)

Rendle. Among a series of late spring to early summer-

blooming (October to mid-December) populations with

smaller, dull yellow to buff flowers and lacking foliage

leaves on flowering stems, we recognize F. spithamea

(Baker) Goldblatt & J.C. Manning for plants lacking

fringed style branches and tepal limbs with smooth rather

than crisped margins. Plants with tepal limbs with minute

brown, red, purple or dull green spotting and well-devel-

oped fringed style branches represent a fourth species,

the earliest name for which is F welwitschii Baker. We

cannot exclude the possibility that additional species

occur in tropical Africa, but specimens from there have

flowers too poorly preserved to be certain.

These changes render De Vos’s 1979 account of Fer-

raria of limited practical use. We provide a revison of

the genus in which we recognize 18 species, including

three new species, F. flava, F. ornata and F. par\>a, all

from the western coastal belt of South Africa. We also

reduce De Vos’s F. kamiesbergensis to subspecific rank

in F. macrochlamys and describe subsp. serpentina of

that species. After briefly outlining the morphology of

Ferraria and what is known about its phytogeny, we

propose a new subgeneric classification. We also review

the pollination biology of the genus, show how floral

morphology is closely correlated with pollination ecol-

ogy, and conclude with a key and systematic account of

the genus.

GENERIC RELATIONSHIPS AND GEOLOGICAL AGE

Ferraria is a member of the predominantly Old World

tribe Irideae (subfam. Iridoideae), which includes the

northern hemisphere genus Iris L. (± 280 spp.) of Eura-

sia and North America, the largely sub-Saharan African

Moraea Mill. (± 200 spp.), the southern African Bobar-

tia L. (15 spp.), and Dietes Salisb. ex Klatt (6 spp.), also

largely sub-Saharan African, but with one species on Lord

Howe Island in the southern Pacific (Mathew 1980; Gold-

blatt 1990). Additional small genera of the tribe are now

no longer recognized. Flermodactylus Mill. ( 1 sp.), Pard-

anthopsis (Hance) Lenz (1 sp.), and Belamcanda Adans.

(1 sp.) have been shown by molecular study to be nested

in Iris (Tillie et al. 2001 ), whereas Barnardiella Goldblatt

(1 sp.), Galaxia Thunb. (15 spp.), Gynandriris Parlatore

(9 spp.), Hexaglottis Vent. (6 spp.) and Homeria Vent.

(32 spp.) are now known to be nested in Moraea (Gold-

blatt 1998; Goldblatt et al. 2002) and are included in that

genus.

Both morphological analysis and molecular DNA

sequences of plastid DNA regions show that Ferraria is

sister to Moraea (±210 species) (Goldblatt 1990; Reeves

et al. 2001; Goldblatt et al. 2002; Goldblatt et al. 2008)

and this clade is sister to Dietes/Bobartia. Using the

molecular clock and a combined analysis of four chloro-

plast DNA regions, preliminary dating estimates showed

that Ferraria and Moraea diverged in the Miocene, ±

25 mya (Goldblatt et al. 2002). Subsequent dating using

improved molecular clock analysis techniques indicate

that the two genera diverged earlier, ± 30 mya, therefore

in the late Oligocene (Goldblatt et al. 2008). Moraea is

distinguished from Ferraria by its bifacial leaf, single

intemode conns of axillary corm ontogeny, whereas

the leaves of Ferraria are unifacial and isobilateral, the

ancestral type in Iridaceae, its corm consists of multiple

intemodes and is of axial origin (De Vos 1977).

MORPHOLOGY

Corm: unmistakable in Iridaceae, relatively large,

depressed globose to almost discoid, and lacks tunics

when mature. A new corm, produced each season, devel-

ops from the base of the flowering stem (De Vos 1977),

and is therefore regarded as terminal or axial (sensu

Goldblatt et al. 2006) in origin. This ontogenetic pattern

contrasts with development in the immediately related

genus Moraea , corms of which are lateral in origin, aris-

ing from an axillary bud at the base of the flowering

stem (Goldblatt 1976, 1990). Although Ferraria corms

are described as having tunics, the tunics are membra-

nous and short-lived and are rarely present in mature

corms, but in F. flava corms carefully removed from the

sandy ground have a dark, softly fibrous covering.

Although new corms are produced annually, those

of the previous seasons are not completely re-absorbed.

Instead, they persist as hard brown discs attached to

each other, like a string of large beads behind the cur-

rent corm. Anatomically, corms lack an organized stele,

a feature shared with Moraea (De Vos 1977), and as in

that genus, new roots emerge from the base of the ter-

minal bud when growth is renewed at the beginning of

the wet season. This contrasts with the typical pattern in

conn-bearing plants, including Iridaceae subfam. Cro-

coideae (syn. Ixioideae), Colchicaceae and Tecophilae-

aceae, which have an organized stele and in which the

new roots emerge from the base or lower half of the

corm (Goldblatt 1990).

Leaves : the lowennost foliar organs sheath the emer-

gent shoot and lack blades and are cataphylls. There are

usually three present and they are typically pale and mem-

branous. The cataphylls are distinctive in Feiraria ferrar-

iola, where they consistently turn reddish with pale speck-

ling above ground level, and often also in F. crispa and

F. densepunctuata. In F. ornata the uppermost cataphyll

develops red pigmentation above the ground. The true

foliage leaves have a sheathing base and a unifacial, linear

to sword-shaped blade with several to many equal, paral-

lel veins or, in F. crispa, F. foliosa and F. schaeferi, there

is a relatively prominent central vein, the pseudomidrib,

consisting of a pair of large, opposed, vascular traces (De

Vos 1979). Even F. glutinosa has, at least anatomically, a

4

Bothalia 41,1 (2011)

pair of larger vascular traces in the centre of the leaf, but

this is not evident in live plants or pressed specimens.

Foliage leaves are lacking on the flowering shoots of F.

spithamea and F. welwitschii, and leaves are evidently

not normally produced by the flowering stems even after

flowering and fruiting are completed. Non-flowering and

juvenile individuals produce one large, narrowly sword-

shaped leaf and one or two smaller ones on a new shoot

produced from the base of the fruiting stalk as capsules

mature. Ferraria glutinosa may also have a reduced or

vestigial basal leaf, but cauline leaves are always present.

The basal leaves (or the juvenile leaves produced by

immature plants) differ sharply from the cauline leaves

in Ferraria densepunctulata, F. ferrariola, F. ornata and

F. ovata. In the last-named species the short, bifacial,

broadly ovate cauline leaves contrast with the unifacial,

linear basal leaves. In F. ornata , juvenile and non-flow-

ering plants produce 3-5 centric, subterete leaves, round

to oval in cross section and up to 100 mm long, whereas

flowering individuals have shorter falcate-lanceolate and

channelled to concave leaves up to 40 mm long, with

short or vestigial unifacial tips.

In Ferraria foliosa and F. schaeferi the basal leaves are

thickest in the midline and have the central vein internal

to the thickest part of the blade (De Vos 1979). The leaf

blade (the unifacial portion of the leaf) of F. brevifolia is

considerably shorter than the prominent sheaths which

together form a tight fan. Ferraria spithamea lacks uni-

facial foliage leaves except in non-flowering individuals;

flowering plants bear relatively large sheathing leaves at

the base and in axils of the flowering stems, the tips of

which are sometimes extended as short, unifacial blades.

Leaf margins may be undifferentiated or are moder-

ately to prominently thickened and hyaline. The hyaline

appearance is the result of the thickening of the strand

of sclerenchyma, present below the marginal epidermis

in leaves of all species. Thickened margins are particu-

larly prominent in Ferraria macrochlamvs subsp. macro-

chlamys and F. uncinata. In F. uncinata the margins are

also usually moderately to strongly crisped and the mar-

ginal thickenings may be slightly irregularly serrate

to crenulate. The leaf tips are often slightly hooked, the

feature for which F. uncinata was named, but the hook is

only weakly developed and is not restricted to that spe-

cies. In F. macrochlamys subsp. macrochlamys the mar-

ginal thickenings are often slightly and minutely pilose

(visible under lOx magnification) and usually crisped

or at least crenulate. The blades of the basal leaves of F.

macrochlamys subsp. serpentina are slightly curved back

on themselves in loose concertina fashion and the mar-

gins are not thickened. In contrast, the leaves of F. macro-

chlamys subsp. kamiesbergensis all curve gently to the

same side, and in some populations, the curvature is par-

ticularly pronounced, forming almost a half circle.

Leaf blades of the Ferraria glutinosa and F. crispa

groups have a visible central vein but in F. divaricata,

F. ornata, F. variabilis and the F. ferrariola and F. unci-

nata groups, the leaves show no external evidence of an

enlarged central vein and leaf sections show all vascu-

lar traces to be ± the same size in these species (De Vos

1979).

Flowering stem and inflorescence: the stem of all spe-

cies is ± erect, terete and, in well-grown plants, often

branched. There are three distinct growth patterns. In

subgen. Glutinosae (the Ferraria glutinosa group), the

stem bears well-developed branches and the intemodes

are sheathed only near their bases. Most other species,

e.g. F. crispa , F. ferrariola , have short, suberect branches

crowded in the upper half of the stem, and the intemodes

are usually enclosed by large leaf sheaths. The cauline

leaves may be held ± in two opposed, vertical ranks,

or are strongly curved outward in a spirally 2-ranked

arrangement in F. foliosa and F. schaeferi. In subgen. Glu-

tinosae the upper nodes and distal portions of the inter-

nodes are sticky. In a third growth form, typical of a few

species including F. macrochlamys and F. variabilis, the

above-ground part of the stem and internodes are short,

and the few branches produced are usually subequal in

length and crowded close to the ground, giving the plants

a somewhat tufted appearance. Ferraria ornata is acau-

lescent, with the stem entirely underground and the rhi-

pidia are borne at ground level.

The main axis and each branch terminates in a cluster

of flowers, a rhipidium, the basic inflorescence unit of

all Iridaceae subfam. Iridoideae. This is a laterally com-

pressed monochasial cyme with the main axis collapsed

and the flowers enclosed by a pair of opposed, large,

± leaf-like bracts called spathes (Goldblatt 1990). The

flowers are borne serially on short pedicels that elon-

gate at flowering so that the flower bud is raised out of

the sheathing spathes. Within the rhipidium, each flower,

except the first one, is enclosed in a large submembranous

bract, which in Ferraria can usually be seen to have a pair

of weakly defined keels, each often greenish and with a

central vascular trace. In subgen. Glutinosae the rhipidia

usually have at least three and in F. glutinosa up to six

flowers. All members of subgen. Ferraria have just two

flowers in each rhipidium. The inner of the two spathes is

longer than the outer and is partly to entirely sheathed by

the outer spathe. The spathe tips are often slightly hooked

and curved inward, a feature weakly developed in most

species. A feature of several species, notably F. crispa

and F. foliosa, is the elongation of the inner spathe as the

capsules ripen. The enlarging capsule therefore remains

enclosed in protective leafy spathe tissue until the aerial

parts wither and dry.

Perianth: specialized in the family, the Ferraria flower

embodies several derived features. The inferior ovary is

either ovoid and borne on a long pedicel that reaches the

tips of the spathes (subgen. Glutinosae ), or the pedicel

reaches to about the middle of the spathes and the ovary

is ellipsoid (F crispa, F. foliosa, F. ornata, F. ovata), or

fusiform and with a sterile, tubular upper half or beak

(F. schaeferi and sect. Macroscvphae of subgen. Fer-

raria). The ovules are then restricted to the lower part

of the ovary. The six tepals are subequal or those of the

inner whorl are somewhat smaller. The tepals are clawed

and the claws may be relatively short, broad, and slightly

ascending to suberect. The claws typically overlap each

other and together form a wide to narrow floral cup,

whereas the tepal limbs spread horizontally or are slightly

reflexed. In F. ferrariola, F. parva and the F. uncinata

group, the tepal claws are relatively narrow, compara-

tively long, and suberect, and then form a narrow cup or

Bothalia 41,1 (2011)

5

gullet, while the limbs may be distally recurved or the

inner tepal limbs may be fully reflexed. In F. ornata the

lower third of the tepal claws are narrowed with the result

that the floral cup has wide gaps or windows alternating

with the bases of the claws. The tepal limbs are lanceo-

late-attenuate and have tightly crisped margins, except F.

spithamea , in which the margins are plane.

Perianth colour is variable across the genus and

is even, within limits, in species. Colours are typi-

cally relatively dull, and the margins of the tepal limbs

often have a contrasting lighter or darker pigmentation.

Because of intraspecific variation it is hard to general-

ize about colour. Ferraria glutinosa has predominantly

dark brown (chocolate-coloured) to dull purple tepals

(also described as violet), the limbs of which have yel-

low margins and sometimes yellow markings at the

bases of the limbs. F. welwitschii has dull yellow to buff

tepals with tiny, dark red. brown or dull green spots but

the tepal limb margins lack contrasting colour. F macro-

chlamys and the closely allied F. brevifolia consistently

have pale, watery yellow tepal limbs with slightly darker

margins. Populations of these species differ, if at all, in

the colour intensity of the few small spots toward the

limb bases. Ferraria uncinata usually has deep blue-vio-

let tepal limbs, but populations in the north of its range

often have light brown limbs with dark blue spots.

The tepals of Ferraria crispa, F. foliosa , F. ornata

and F. schaeferi are usually irregularly mottled with

dark brown to dull maroon on a paler ground colour.

But at least in F. crispa and F. glutinosa there are indi-

viduals with almost uniformly dark brownish tepals with

pale, almost golden margins (see figure in Manning et

al. 2002: 157). Flowers of F. divaricata are also dark-

coloured, but are not blotched with darker pigment;

instead they have dark brown limbs, described in the

type description as chocolate-brown, with light brown

margins and a pale cup, usually longitudinally streaked

with brownish purple or with a single broad longitudi-

nal streak. The series of populations that De Vos (1979)

called subsp. aurea have flowers with yellow tepals

very faintly marked with contrasting colour. Ferraria

variabilis shows a range of distinctive perianth pat-

terning across its range. In the north, in Namaqualand

and southern Namibia, flowers have tepal limbs with a

darker band toward the base and are pale brown distally

and the margins are the same pale brown. The claws are

pale greenish cream with a narrow darker band running

down the midline. In the western Karoo flower pigmen-

tation is more variable, with shades of brown or yellow

predominating but individual variation is often striking.

In the southern Cape and southern Karoo F. variabi-

lis has greenish yellow or light to middle brown tepal

limbs, usually darker in the midline, and with small or

medium-sized spots in the lower third, or when brown,

then often with spots irregularly scattered across the

limb and the margins sometimes slightly darker col-

oured. The claws are pale, usually with a darker streak in

the midline. Perianth colour can vary as much within a

population as across the entire southern Cape and west-

ern Karoo.

Perhaps most striking in floral pigmentation, Fer-

raria ovata has yellow tepals with dark brown at the

base of the tepal limbs and along the margins and mid-

line (Figure 7). In F. ferrariola the tepal limbs range

from pale yellow to grey-blue or light turquoise usually

with a pale yellow zone toward the base of the outer

tepal limbs bearing small dark dots, while the particu-

larly prominent, feathery style branches are grey-blue to

pale purple. The smaller, less conspicuous inner tepals

are unmarked, often a darker colour, and the limbs are

± reflexed, in contrast to the horizontally extended outer

tepal limbs.

Each tepal claw bears a nectary, which takes the form

of small dark zones or larger shallow depressions cov-

ering one third to almost half the claw. Nectaries are

largest and most conspicuous in the tropical African spe-

cies and in those southern African species with a shal-

low floral cup, as in Ferraria crispa, F. foliosa and F.

schaeferi the nectary is paler than the rest of the claw. In

contrast, the nectaries of F. glutinosa and F. welwitschii

are dark brown and often wider than high. F. densepunc-

tulata and F. ovata have small, pale, bilobed or paired

nectaries above the tepal bases. In F. ornata the nectar-

ies are ± white with violet spots and the distal edges are

raised into an irregularly lobed crest. Smaller nectaries

located close to the tepal base are often not easy to iden-

tify, particularly in species with a deep, narrow floral

cup. In those species with large nectaries, small drop-

lets of intensely sweet nectar, in excess of 50 % sucrose

equivalents, are scattered over the surface, which may

coalesce in a pool in the base of the floral cup when

the air is moist or after light rain. Nectar of remarkably

low sugar concentration, in the range of 5-9 % sucrose

equivalents, accumulates in a pool in the relatively

deep floral cups of F. divaricata and F. variabilis. The

size and position of nectaries is usually constant, but we

have encountered populations of F. variabilis with large

bilobed nectaries forming pockets in the middle of the

outer tepal claws (e.g. Goldblatt & Porter 12977 from

the Tankwa Karoo) although the species usually has

small dark nectaries close to or at the base of the claws

(De Vos 1979).

Androecium and gynoecium : the three stamens have

filaments united for most of their length in a slender,

smooth or rarely puberulous column that reaches to the

apex of the floral cup. The upper 1. 5-3.0 mm of the

filaments are free and diverging. The anthers are either

oblong with conventional, parallel thecae (sects. Glu-

tinosae and Ferraria), or the thecae are held together

only at their tips and diverge from the apex after anthe-

sis, usually becoming widely divergent (sect. Macroscy-

phae). The filaments are inserted in a small pocket in the

connective ± a third of the anther length above the base

or, in sect. Macroscyphae a short distance below the

anther apex.

Pollen is bright orange-red in most species, or occa-

sionally yellow, but Ferraria flava consistently has yel-

low pollen. Pollen of F. welwitschii has been described

as orange-yellow (Milne-Redhead 2664) or is yellow

in photographs we have examined. Anthers shrink a

surprising amount, at least a third and up to half their

length, after pollen is shed. De Vos (1979) described the

pollen grains in some detail: they are always monosul-

cate with reticulate exine but exhibit no features of taxo-

nomic significance within the genus.

6

Bothalia 41,1 (2011)

The ovary is borne on a well-developed (subgen.

Glutinosae) or short pedicel that elongates just before

anthesis, thus raising the flower bud above the sheath-

ing inflorescence spathes. The ovary is either narrowly

ovate-truncate (subgen. Glutinosae) to ± fusiform (sect.

Ferraria except F. schaeferi ) or in all members of sect.

Macroscyphae and F. schaeferi elongate-fusiform,

tapering distally into a slender, sterile tube, the rostrum

or beak. Only the middle portion of the wider, lower

portion of the ovary then contains ovules. The ovary

remains included in the spathes except in subgen. Glu-

tinosae and is then held either just below the apex of the

inner spathe, or when the ovary is beaked, well within

the spathes.

The slender style, held within the filament col-

umn, divides at its apex into three tangentially flattened

branches, each divided for more than half its length into

a pair of conduplicate lobes or arms that curve outward.

The arms consist of an adaxial surface, nearly always

deeply and irregularly fringed, and an abaxial surface that

arches over the anthers and bears the stigma. The fringes

of Ferraria glutinosa can be seen under at least 20x mag-

nification to be minutely papillate, a feature not or barely

developed in other species. In F. spithamea the style

branches lack the fringes that characterize the rest of the

genus.

In subgen. Glutinosae and sect. Ferraria of subgen.

Ferraria, the stigma is minute and located at the apex

of each stylar arm (see illustrations by De Vos 1979). In

sect. Macroscyphae (subgen. Ferraria ) the stigmatic sur-

face consists of an expanded and channelled fold of tissue

below the tip of each style arm. The stigmatic surface is

unusually large in F ferrariola and arches over the adja-

cent anther lobe, partly concealing it.

Capsules and seeds', capsule shape largely mirrors that

of the ovary. Thus, in subgen. Ferraria the capsule is ±

globose-truncate closely resembling that in the related

Bobartia, the less specialized species of Dietes (e.g. D.

bicolor ) and many species of Moraea. In F. crispa , F.

densepunctulata and F. foliosa, which have a fusiform

ovary, capsules are ovoid with a rounded to pointed apex.

In those species with a beaked ovary, the beak persists in

fruit as a firm extension at the tip of the large, ovoid, seed-

bearing portion of the fruit. The capsules are enclosed in

the spathes throughout their development in subgen. Fer-

raria but remain exserted in subgen. Glutinosae as they

ripen.

The relatively large seeds are unique in Irideae in their

size, multifaceted shape and distinctive smooth, glossy

surface. In most species, seeds are five- or six-sided with

the facets separated by raised, somewhat undulate ridges

and the individual facets have a ± plane to slightly undu-

late or somewhat wrinkled surface. These seeds are, to

some extent, variable in size, even in the same capsules,

and ± 2.5-3 .4 mm wide and 3-4 mm at the longest axis.

Seed colour ranges from pale to dark brown, with the

ridges separating the facets often lighter yellow-brown.

The entire seed surface is smooth and glossy without

raised cell outlines. De Vos (1979) noted that F. divari-

cata (as subsp. arenosa ) stands out in having matte, dull

brown, globose seeds with a slightly wrinkled, ± rumi-

nate sculpturing (called reticulate by De Vos) and fove-

ate epidermal cells, which we have confirmed for one

population.

CHROMOSOME CYTOLOGY

The basic chromosome number in Ferraria (Table

I ) is x = l o and diploid numbers range from 2 n = 20

to 60 (Goldblatt 1971; De Vos 1979; Goldblatt & Takei

1997). Most species are paleodiploid but F. crispa and

F. schaeferi have tetraploid and hexaploid populations

(2n = 40 and 60) (Table 1). Largely on the basis of

their tetraploid chromosome number, 2 n = 40, but also

a slightly smaller flower, De Vos (1979) segregated the

northern populations of F. crispa as subsp. nortieri. The

southern and eastern populations of the species are hexa-

ploid, 2 n = 60. The single count for F. glutinosa (from

Namibia) is tetraploid and F. variabilis has diploid and

tetraploid populations, the latter in the southern part of

its range (and were assigned by De Vos to a separate

subsp. australis of F. divaricata). The southern African

F. Jlava, F. ovata, F. ornata, F. par\>a and the tropical F.

candelabrum and F. spithamea are uncounted.

TABLE 1 . — Chromosome numbers of Ferraria. Data from Goldblatt

(1971), De Vos (1979), and Goldblatt & Takei (1997). Ferraria

candelabrum. F. pan’a, F. ornata, F. ovata. F. spithamea and F.

macrochlamys subsp. serpentina are uncounted

Chromosomes are large, as they are in most genera of

subfam. Iridoideae (Goldblatt 1971). The basic karyo-

type consists of nine acrocentric and one metacentric

chromosome pair, reported for Ferraria welwitschii (De

Vos 1979, as F. glutinosa). The southern African species

have, in contrast, a derived karyotype of 10 ± acrocen-

tric chromosomes, some of the smaller pairs tending to

metacentric. Within Iridoideae, the karyotype of F. wel-

witschii with its metacentric chromosome pair, most

closely resembles that of Moraea species that have x =

10, the ancestral base number in that genus. Both Bobar-

tia and Dietes , the two genera immediately related to

the Ferraria-Moraea clade also have x = 10 (Goldblatt

1971; Goldblatt & Takei 1997) and a karytotype with

both aero- and metacentric chromosomes.

Bothalia 41,1 (2011 )

7

POLLINATION BIOLOGY

Somewhat surprisingly for a small genus with super-

ficially similar flowers, pollination of Fen-aria is rela-

tively diverse (Goldblatt et al. 2009). As was long sus-

pected, the flowers of several species are adapted for

pollination by muscid flies, calliphorid flies, flesh flies

and other families of Diptera. Vogel (1954), in his semi-

nal survey of pollination systems in the southern African

flora, regarded Fen-aria as a genus adapted for pollina-

tion by flies. The basis for this contention appears to

have been the reports by Scott Elliot (1891) of visits to

F. crispa (as F. undulata) by two Diptera, the flesh fly

Scathophaga hottentotta (now S. stercoraria ) (Sarco-

phagidae) and the blowfly Chrysomya regalis (Cal-

liphoridae). These insects are evidently attracted to

the strong, molasses or burnt sugar odour of the flow-

ers (sometimes loosely described as putrid or decay-

ing), combined with a dull, mottled perianth coloration,

reminscent of the flowers of Orbea and Stapelia species

(Apocynaceae). Two populations sampled respectively

by R. Raguso (pers. comm. 2009) and R. Kaiser (pers.

comm. 2009) both yielded significant amounts of guai-

acol, a phenylpropionoid, which has a strong molasses

odour. The reward offered is concentrated sugary nectar

retained as tiny droplets on the surface of large perigonal

nectaries within the floral cup.

Ferraria densepunctulata, F. foliosa, F. ovata, and F

schaeferi (all sect. Ferraria) share this pollination sys-

tem, often termed sapromyiophily (Faegri & van der

Pijl 1979) and according to the definition, associated

with a dull-coloured and often mottled perianth, odours

of rotting protein, but without a reward. Clearly, flies

are attracted by the appearance and odour of the flow-

ers but in Ferraria they are rewarded by concentrated

nectar. Four families of Diptera: Calliphoridae (Chryso-

mya), Tachinidae (genus undetermined), Sarcophagidae

( Scathophaga ), and Muscidae (several genera) have now

been recorded visiting flowers of this group of species

(Goldblatt et al. 2009). In fair to warm weather, flowers

of these species are rarely without one or more fly visi-

tors, usually carrying dense loads of bright orange pol-

len covering the dorsal parts of the head and thorax and

sometimes the abdomen. The ± bitter or aminoid scents

produced by the flowers of F. crispa . F. foliosa, and F.

ovata are typical of flowers visited by these flies. In con-

trast, F. densepunctulata and F. schaeferi flowers have

faint, and to the human nose, pleasant floral odours, and

visits by the same suite of fly species are as frequent as

those to F. crispa and F. foliosa.

Not all Ferraria flowers have this floral presenta-

tion and pollination system. Flowers of F. ferrariola are

specialized in the genus in having a narrow floral cup,

17-20 mm deep, a pale blue-grey or yellow perianth

with only the outer tepals bearing nectar guides, nectar

of elevated sugar concentration, and typically a faint

sweet scent recalling violets mixed with a spicy odour,

in some populations recalling vanilla or, in others, bit-

ter almonds. Flowers of some populations appear to

be unscented. The flowers are pollinated only by bees,

including honey bees. Apis mellifera , and long-tongued

anthophorines, including Anthophora praecox (Goldblatt

et al. 2009). Observations on the pollination of other

bee-pollinated Iridaceae show that anthophorine bees of

several species may visit and successfully pollinate spe-

cies of Babiana and Gladiolus (Goldblatt et al. 1998,

2001; Goldblatt & Manning 2006) across their ranges,

and we conclude that different Anthophora species

probably visit F. ferrariola at other sites across its wide

range. Bees climb into the narrow floral cup, evidently

in search of nectar and, in doing so, they passively accu-

mulate loads of orange pollen on the dorsal part of the

thorax. Visits to a second flower result in the deposition

of pollen on the exposed stigmatic surfaces of the style

branches.

Ferraria divaricata and F. variabilis, either have

dull-coloured flowers, either uniformly pale to dark

brown, speckled or banded with purple, brown or green

on a cream to dull yellow background, or in F. macro-

chlamys pale, watery yellow. Floral odour seems to be

absent, or is slightly unpleasant, but indefinable. The

flowers produce large amounts of nectar of exception-

ally low sugar concentration, mostly 3-10 % sucrose

equivalents, which accumulates in a pool in the base

of a wide, fairly deep floral cup. Flowers of both spe-

cies are visited by medium-sized to large potter wasps,

Delta and Allepipona erythrospina (Eumenidae) or

female masarine wasps (Masarinae), Jugurtia koeroe-

gabensis recorded on F. variabilis (as F. divaricata) and

Ceramius on F. macrochlamys subsp. kamiesbergensis

in Namaqualand (Gess 1997, reported as F. divaricata,

now F. variabilis). Other potential insect visitors includ-

ing bees and flies of several families common nearby are

not attracted to these flowers. The eumenid wasps, seen

only infrequently, display a constant foraging pattern,

alighting on flowers of one plant after another. In doing

so they become densely covered in pollen on the dorsal

part of the head and thorax. Microscopic examination

of pollen from captured male and female individuals of

Allepipona and female Delta caffer shows that the only

type of pollen they carry is that of Ferraria. The reason

for visits by wasps is unknown but may simply be for

the water content of the nectar rather than the dissolved

sugar, although the possible presence of amino acids in

the nectar has not been assessed.

One last pollination system in the genus is that

reported in Ferraria uncinata, which is visited only by

small herbivorous beetles, Iselma planidorsus (Meloi-

dae), and an unnamed genus of Melyridae (Goldblatt et

al. 2009). We assume that nectar of moderate sugar con-

centration is the reward for these insects. We are hesitant

to accept these beetles as the legitimate pollinators but

these fairly common flower visitors are almost invari-

ably seen on flowers of F. uncinata.

Pollen carried by flies, bees, and wasps was always

deposited on the dorsal part of an insect’s thorax as

it enters the floral cup in search of nectar. To date, no

insect visitors capable of accomplishing pollen transfer

have been recorded on the small, pale, watery, yellow

flowers of Ferraria brevifolia but we assume the species

has the same pollinator as F. macrochlamys. The flowers

of these species have faintly scented flowers with a nar-

row floral cup containing dilute nectar, 6-12 % sucrose

equivalent. All these species require insect-mediated

pollen transfer and our lack of success in locating pol-

linators remains puzzling.

Bothalia 41,1 (2011)

Nectar sugar chemistry, examined for individuals of

four species shows that Ferraria species secrete hexose-

dominant nectar (Goldblatt et al. 2009). This is consist-

ent with nectar sugars of the sister genus Moraea (Gold-

blatt & Bernhardt 1999) and contrasts with nectar sugars

of Iridaceae subfam. Crocoideae which are sucrose-rich

or sucrose dominant except in a few species adapted to

butterfly or sunbird pollinators (Goldblatt et al. 1999,

2001; Goldblatt & Manning 2006).

HISTORY

The first record of Ferraria in the literature is the illus-

tration of F. crispa in the celebrated volume. Flora sen

de florum cultura (Flora of cultivated plants) by the 17th

century artist, Giovanni Batiste Ferrari (1633). Interest-

ingly the illustration was made from a plant grown in Italy.

Ferrari called it \flos indicus e violaceo fuscus radice tube-

rosa ’ and, unfortunately, did not record the source of his

plant. The genus appears in the botanical literature again

in Robert Morison’s (1680) Plantarum historiae universa-

lis as Gladiolus indicus e violaceo fuscus and in Olof Rud-

beck’s (1701 ) Reliquiae mdbeckianae as Narcissus indicus

flore saturate purpureo but the illustrations in both works

appear merely to have been copied from Ferrari’s original.

Ferraria was formally described by Philip Miller (1759a,

b), who attributed the name to the Dutch botanist, Johan-

nes Burman. Evidently Burman communicated to Miller

his intention to name the genus Ferraria , of course, hon-

ouring Giovani Ferrari. Burman’s own paper was pub-

lished two years later (Burman 1761) and included not

only an extended botanical description, but woodcut

illustrations of two plants, F. crispa , and what he called

F. fimbriata , the identity of which is uncertain. The illus-

trations are in fact the types of the two species and ade-

quate for the identification of F. crispa but not for F. fim-

briata. We agree with De Vos (1979) that the latter is not

F. crispa. Our best guess it that the illustration represents

what is now F. divaricata.

Linnaeus (1759) accepted the genus without cit-

ing Miller or Burman, but listed one species, Ferraria

undata , without description. Later in Species plantarum

edn 2, Linnaeus (1763) admitted one species, F. undu-

lata, thus overlooking Burman’s earlier epithet, F.

crispa , although citing Burman’s paper. Linnaeus’s F.

undulata was based on Miller’s 1759 illustration and

was the name used for the species until the late 20th cen-

tury when Moore (1974) noted that F. crispa has nomen-

clature! priority.

The distinctive and most unusual looking Ferraria

crispa was grown in Paris and Vienna in the later 18th

century and was illustrated by Jacquin (1770) and Re-

doute (1802), amongst others. Thunberg (1787) col-

lected F. crispa himself and realized it was Linneaus’s

F. undulata. However, he transferred the species to

Moraea , which then included such diverse genera as

Bobartia and Aristea (typical Moraea species were

at the time included in Iris). Only two other species of

the genus, F. ferrariola and F. ovata , are known to have

been recorded before 1800. Jacquin described Moraea

ferrariola in 1790, thus following Thunberg ’s circum-

scription of that genus. Jacquin’s description was based

on a plant grown in Vienna, and very likely provided by

Franz Boos and Georg Schol, who collected plants for

Jacquin at the Cape in the years 1786 to 1788 (Gunn &

Codd 1981). Then, in the 1890s, most likely 1893, Fran-

cis Masson, plant collector for the Royal Botanic Gar-

dens, Kew, discovered what we now know to be Fer-

raria ovata in western South Africa. His collection, a

dried specimen lacking preserved flowers or corms, was

described as Moraea ovata by Thunberg (1800).

We owe the restoration of Ferraria to Willdenow

( 1 800) for in edn 4 of Species plantarum he included F.

undulata and F. ferrariola in the genus, as well as two

American species, the superficially similar F. pavonia,

now Tigridia pavonia, and F. ixioides, now Libertia

ixioides. Thunberg’s Moraea ovata was not included in

Willdenow’s account (perhaps because of their con-

temporary publication) and M. ovata remained a puz-

zling species until 1995 when vegetative plants were

collected near the Skilpad Reserve in Namaqualand by

botanist, Annelise le Roux. Flowering specimens were

obtained soon thereafter and the species was found to

have a Ferraria- like flower. M. ovata was transferred

to Ferraria by Goldblatt & Manning (2002). Salisbury

(1796), always a maverick, also recognized Ferraria

but expanded its circumscription to include species of

what are today Moraea and Sisyrinchium as well as Iris

domestica (syn. Belamcanda chinensis). Ker Gawler

(1804, 1827) followed Willdenow in maintaining Fer-

raria but Tigridia pavonia and a few of its American

allies in the tribes Tigridieae and Trimezieae (species

of Cypella, Eleutherine, Gelasine, and Trimezia ) were

sometimes included in Ferraria. Tigridia had actually

been described for Linnaeus fil.’s Ferraria pavonia in

1789 (Jussieu 1789) although the species itself was only

formally transferred to that genus some years later (De

Candolle in Redoute 1802). The separation of Tigridia

from Ferraria, or for that matter Moraea, was not uni-

versally accepted until late in the 19th century (Molseed

1970).

The early 19th century saw the addition of two new

Ferraria species: F. uncinata (Sweet 1826a) and F.

divaricata (Sweet 1827) were both grown in England

from corms or seeds collected by Walter Synnot, land-

drost (magistrate) at Clanwilliam from 1821 to 1825.

Four more species were described in the latter half of the

19th century by J.G. Baker (1878), notably Ferraria glu-

tinosa and its allies, F. candelabrum and F. spithamea,

all from Angola and based on collections by the Austrian

Friedrich Welwitsch in the years 1855-1861, and all were

initially referred to Moraea by Baker who evidently did

not see their resemblance to the southern African Fer-

raria species (Baker 1877, 1896). This error was most

likely due to the condition of the specimens, the flowers

of which are poorly preserved. In 1892, Baker described

another Angolan species, F. welwitschii, this time as

referred to the genus. Based not on herbarium material

but on an illustration of a plant grown in England, the

drawing with some parts painted in watercolour, showed

clearly the hallmarks of Ferraria, fringed style crests,

a floral cup and crisped and undulate tepal limb mar-

gins. It was left to J.B. Rendle (1899) to transfer to Fer-

raria, Baker’s other tropical species. Baker (1876) also

described what is now F. macrochlamys but included it

in Lapeirousia. The species was based on a poor herbar-

Bothalia 41,1 (2011)

9

ium specimen in the Kew Herbarium, originally owned

by William Forsyth and probably collected by William

Paterson in the 1780s, though this is no more than sur-

mise.

Several more species were added in the later 19th

and early 20th centuries but most are synonyms of those

already described. In 1920, Kurt Dinter added Fen-aria

schaeferi based on collections from southern Nanibia

± 1912-1913, unaware that the species had first been

recorded by J.-F. Drege in 1820. G.J. Lewis added Fer-

raria brevifolia and F. foliosa in 1954 and M.P. de Vos, in

her revision of the genus, added F. densepunctulata and

F. kamiesbergensis . De Vos recognized just 10 species

and four subspecies (14 taxa). De Vos unfortunately mis-

understood the tropical African species and included the

four we recognize here in F. glutinosa, following Carter’s

(1963) lead. Our revision reverses this action, and we now

admit 1 8 species and two subspecies (20 taxa) in Ferraria

assigned to two subgenera and three sections.

PHYLOGENY AND EVOLUTION

De Vos (1979) considered the morphologically and

geographically isolated Ferraria glutinosa (in which

she included all four tropical African species we recog-

nize), the least specialized member of the genus in the

light of its open branching growth form, cluster of basal

leaves differentiated from the bract-like cauline leaves,

multi-flowered inflorescences, and globose, truncate

ovary. These features seem intuitively to be ancestral,

recalling species of the related genus Moraea. Molecular

studies of the genera of the Iridaceae using plastid DNA

sequences confirm that Ferraria and Moraea are imme-

diately related (Goldblatt et al. 2002, 2008). The genera

share apomorphic astelic conns that produce roots from

the base of the terminal bud. unique in the Iridaceae,

and also have flowers with the filaments partly united,

whereas other African genera of Irideae and Iris itself,

have free filaments as their ancestral condition.

DNA sequences of 12 Ferraria species, using the

plastid DNA regions trnL-V and rbcL (unpublished

data), confirm our intuitive hypothesis about relation-

ships within the genus. Ferraria glutinosa is sister

to the remaining species in the data set, which did not

include any other tropical African species. The relation-

ships among these species is largely unresolved, leaving

us with the conclusion that F. glutinosa (and presum-

ably the morphologically similar tropical species) are

taxonomically and geographically isolated species that

evolved before the establishment of the present southern

African winter rainfall climate. This in turn is consistent

with the most recent molecular clock estimates of the

divergence of Moraea and Ferraria ± 30 mya (Gold-

blatt et al. 2008), during the later Oligocene, when the

African climate become increasingly dry and seasonal,

but with summer precipitation. This period coincides

with the establishment of circum-Antarctic oceanic cir-

culation and the strengthening of the cold Benguela

current along the southwestern African coast. Although

still under a summer rainfall regime, seasonal aridity is

believed to have caused the reduction of forest in favour

of open grass-dominated habitats (Raven & Axelrod

1974; Coetzee 1993). Such a regime would have encour-

aged the evolution of geophytic plants, which survive

periods of drought using underground storage organs

that enable them to rapidly sprout and flower when suit-

able growing conditions return.

The onset of a winter rainfall regime in south-

ern Africa probably dates from the beginning of the

Pliocene, ± 6 mya, and this is when we assume that a

recent burst of radiation occurred in Ferraria in the

south, associated with summer drought and cool, wet,

winter conditions. We infer that the resulting species we

see today are too recent to have accumulated sufficient

DNA sequence diversity to be successfully resolved in

phylogenetic studies of two chloroplast DNA regions we

utilized for our molecular study.

For the present, except for the position of Fer-

raria glutinosa (and we infer, the other tropical spe-

cies) as sister to the remaining species confirmed by

DNA sequence studies, we are left with morphology as

a guide to the phylogeny of the genus. Outgroup com-

parison reveals two major specializations in the Ferraria

species of the southern African winter rainfall zone: an

ovary with a sterile tubular tip, the beak (and associ-

ated beaked capsule); and divergent anther lobes. The

first of these is present in the species of sect. Macro-

scyphae (subgen. Ferraria ) of De Vos plus F. schaeferi ,

and divergent anther lobes are restricted to that section,

although the expression of the character is delayed in F.

ferrariola and F. parva, in which it is expressed in older,

flowers, after dehiscence of the anther lobes and some-

times only on the second day of anthesis.

We propose a revised infrageneric classification of

Fen-aria based first on the molecular data, supplemented

by the morphological variation evident in the ovary and

anthers. Thus we recognize two subgenera, subgen. Glu-

tinosae, for the tropical species and subgen. Ferraria

for the remaining species, all of winter rainfall southern

Africa. We endorse De Vos’s two sections, Ferraria and

Macroscyphae, the first for species with parallel anther

lobes, broad, spreading tepal claws and a relatively wide

floral cup, and except for F. schaeferi , a subacute cap-

sule apex lacking a beak. Sect. Macroscyphae always

has anther lobes joined only at the tips and usually

widely divergent (rarely sub-parallel), a beaked ovary,

and except in F. divaricata and F. variabilis , suberect

tepal claws that form a narrow floral cup. In sect. Mac-

roscyphae we recognize three informal series, Macro-

scyphae, Subdivaricatae and Uncinatae.

SYSTEMATICS

Ferraria Bunn, ex Mill., Figures of plants, vol. 2:

187, t. 280 (1759a) and The gardeners dictionary, edn 7

(1759b); De Vos: 327-375 (1979). Type species: Fer-

raria foliis lanceolatis ... (= F. crispa Burm.).

Named for the 17th century artist, Giovanni Ferrari,

whose illustrated works were much celebrated by contem-

poraries; his Flora sen de florum cultura (Flora of culti-

vated plants) included the first illustration of Ferraria , F.

crispa , drawn from plants grown in Italy.

Revisionary accounts: De Vos in Journal of South

African Botany 45: 295-375 ( 1979).

10

Bothalia 41,1 (2011)

Small to medium-sized seasonal perennials. Corm

depressed-globose, consisting of several intemodes,

producing roots from terminal bud, lacking tunics when

mature. Leaves several, lower 3 or 4 lacking blades,

thus cataphylls, foliage leaves with isobilateral, sword-

shaped to linear, unifacial blades, sometimes not pro-

duced on flowering stem, basal one or more and larg-

est, progressively smaller above, becoming bract-like at

distal nodes. Flowering stem aerial or largely to entirely

underground, usually branched, branches often short and

crowded distally. Inflorescence a 2- or several-flowered

rhipidium; spathes green, often leafy, enclosing flower

buds, inner always longer than outer, margins free to

base. Flowers radially symmetric, fugaceous or lasting

2 days, often dull-coloured, usually cream-coloured to

buff or brownish, sometimes yellow, rarely blue-violet,

spotted and blotched dark brown, green or purple, with-

out scent or scented with sweet, spicy or rotting odours;

tepals free, clawed, those of outer whorl slightly to sig-

nificantly larger than inner, claws forming a wide or nar-

row cup or shallow basin, limbs spreading to slightly

reflexed, margins crisped, the tips attenuate and twisted,

producing nectar from nectaries at base or middle of

claws, distal edges of nectaries lobed in F. ornata. Sta-

mens symmetrically arranged; filaments united in lower

± two thirds in a smooth column, free and diverging

in upper third or quarter; anthers appressed to style

branches, lobes joined to connective but not to each

other, parallel and ± contiguous or widely diverging,

rarely subparallel but not contiguous. Ovary included

or exserted from spathes, often with sterile beak; style

enclosed by filament column, dividing at column apex

into short branches, each forked into flattened, diverg-

ing, conduplicate arms, each usually prominently

fringed on adaxial margin, rarely fringes vestigial or

absent (F spithamea), stigmatic surface terminal on

style branches or on grooved lobe below tip of abaxial

margin. Capsules globose-truncate, ovoid or ellipsoid,

then often beaked. Seeds angular, mostly 5- or 6-sided,

facets separated by winged angles, facets ± smooth or

wrinkled, in one species ± globose and reticulate. Basic

chromosome number x = 10.

Species: 18, dry parts of central and southwestern

Africa, with a centre along the southern African west

coast and near interior, and four species extending from

interior southern Africa through Namibia to Angola,

Congo, Zambia, Zimbabwe and Malawi in dry grass-

land, generally favouring sandy soils, occasionally rocky

habitats or seasonally wet sites.

Key to species

la Anthers with filament insertion in lower third and lobes parallel and usually contiguous above filament insertion when fully dehisced;

capsules globose- to ovoid-truncate or acute or beaked in one species; perianth with tepal claws usually forming a shallow cup and

bearing prominent nectaries '/3 to '/, as long as claw; stigmas minute, terminal on apices of style branch arms (subgen. Glutinosae and

sect. Ferraria ):

2a Upper intemodes below flower clusters with sticky exudate; ovary ovoid, short, < 10 mm long; capsules ovoid to globose and truncate

(subgen. Glutinosae ):

3a Leaves several (up to 8) in basal fan; rhipidia with outer spathe 7 to ‘/4 as long as inner 2. F. candelabrum

3b Leaves absent at base of flowering stem or 1-4 basal; rhipidia with outer spathe V to slightly more than V, as long as inner:

4a Flowers brown or dark purple, sometimes marked with yellow near tepal limb bases; tepal limbs often with buff to yellow mar-

gins; flowering plants usually with 1—4 basal foliage leaves, or foliage leaves reduced; filament column 8-11 mm long; flower-

ing (Dec.-)Jan.-Mar.(-May) 1 . F. glutinosa

4b Flowers yellow to buff, limbs sometimes speckled with brown, dark red, purple or dull green spots; tepal limb margins not con-

trasting in colour; flowering plants lacking foliage leaves at base and branches subtended by sheathing leaves without blades;

filament column 5. 0-7. 5 mm long; flowering (Sept.-)Oct. to Dec. (-early Jan.):

5a Tepal limbs speckled with dark colour and margins obviously crisped; style branch arms prominently fringed 4 . F. welwitschii

5b Tepal limbs uniformly yellow and margins plane or slightly undulate; style branch arms entire (not fringed) and arching outward

3 . F. spithamea

2b Plants ± acaulescent or stem aerial, then upper intemodes below flower clusters not sticky; ovary ± fusiform, > 10 mm long; capsules

usually ovoid, rounded at apex or with short sterile beak (sect. Ferraria ):

6a Stem aerial, slender with intemodes partly exposed; basal leaves linear to falcate, with blades usually < 5 mm wide:

7a Plants ± acaulescent; tepal claws narrow in lower third, thus forming a windowed cup; nectaries concave, at apex of narrow part of

claw and with raised and lobed to fringed distal edge 10. FI ornata

7b Plants with aerial stems: tepal claws broad and not abruptly narrowed in lower part, cup not windowed; nectaries relatively large,

in centre of tepal claws and without raised distal edges:

8a Cauline leaves lanceolate-attenuate; flowers grey or blue-green with minute brown, maroon or violet speckling; nectaries yel-

lowish green 9. F. densepunctulata

8b Cauline leaves ovate-cucullate; flowers yellow with brown margins and a few conspicuous spots; nectaries cream-coloured

streaked maroon 8. F. ovata

6b Stem stout, mostly covered by leaf bases; leaves sword-shaped to falcate, with blades > 5 mm wide:

9a Ovary with short tapering beak, ± 8 mm long; tepals 22-25(-30) mm long, pale yellow with dark brown margins and blotches,

often coalescing in outer quarter; flowers sweetly scented 1 . F. schaeferi

9b Ovary fusiform, without tapering beak; tepals 25-35 mm long; flowers variously coloured, usually dark maroon or purple with

paler margins or cream-coloured to pale yellow and variously striped and blotched, with an unpleasant, burnt-sugar or putrid scent:

10a Blades of basal leaves with slightly thickened zone in middle and prominent central vein; leaves 2-ranked 5. F. crispa

1 0b Blades of basal leaves much thickened in middle but with weakly developed central vein; leaves spirally 2-ranked 6. F. foliosa

lb Anthers with filament insertion in upper third and lobes held together only at tips, usually widely divergent when fully dehisced, rarely

sub-parallel but not contiguous (not always so in freshly opened flowers); capsules always prominently beaked; perianth with tepal

claws forming deep, narrow or wide cup; stigmas consisting of groove below tips of style branch arms (sect. Macroscyphae ):

1 la Floral cup widening substantially toward apex, rim ± as wide or wider than depth of cup:

12a Flowers small, sweetly, slightly clove-scented; floral cup ± 8 mm deep, ± 7 mm wide at rim; outer tepals 18-22 mm long; filament

column ± 6 mm long 12. F. parva

12b Flower relatively large, unscented or faintly unpleasantly scented; floral cup 12-15 mm deep, 13-20 mm wide at rim; outer tepals

30— J5 mm long; filament column 8-13 mm long:

Bothalia 41,1 (201 1 )

13a Plants usually subacaulescent and stem mostly covered by sheathing parts of leaves; flowers lasting 2 days; seeds (4)5(6)-sided,

facets separated by raised angles, surface smooth 14 . F. variabilis

13b Plants with aerial stem and branching only distally; flowers lasting a single day; seeds ± globose, surface matte 13. F. divaricata

lib Flowers with narrow cup, sides ± erect (rim of cup V, to 2/3 as wide as depth of cup):

14a Stem produced well above ground; basal leaves sharply distinct from cauline leaves and with intemodes partly exposed; flowers

with floral cup 1 7-20 mm deep; outer tepals 30-40 mm long 11 . F. ferrariola

14b Stem not or only shortly extended above ground (unless growing in shade); basal leaves often hardly distinct from cauline leaves

(or from spathes enclosing flower clusters); floral cup 10-12 mm deep; outer tepals 25-35 mm long:

1 5a Basal leaves broad, leathery-succulent, 8-20 mm wide, cauline leaves and spathes different from basal leaves; flowers lasting a

single day, deep yellow; anthers and pollen yellow 15. F. flava

15b Basal leaves hardly distinct from cauline leaves, rarely exceeding 12 mm wide, flowers lasting two days, watery yellow, blue or

shades of beige to cream-coloured dotted with blue spots; anthers and pollen usually orange to ± red:

16a Flowers usually predominantly blue to violet with dull yellow-green to khaki margins, occasionally tepal limbs pale bluish or

cream-coloured, then speckled dark blue; margins of at least lowermost leaves conspicuously thickened, hyaline and partly

crisped and undulate; marginal thickenings smooth 16. FI uncinata

16b Flower pale watery yellow with darker yellow to buff or khaki margins; leaf margins either thickened then lower leaves with

margins plane or serrulate to crenate, or sometimes slightly crisped or unthickened and margins plane; marginal thickenings

when present often shortly velvety:

17a Basal leaves with extended, lanceolate to linear blades longer than sheaths 17. F. macrochlamys

17b Leaves with short, oblique, ovate blades less than half as long as sheaths 18. F. brevifoha

A. Subgen. Glutinosae ( M.P.de Vos) Goldblatt

& J.C. Manning, subgen. & comb. nov. Sect. Gluti-

nosae M.P.de Vos: 329 (1979). Type: Ferraria glutinosa

(Baker) Rendle.

Plants with open, aerial branching system and stem

sticky below nodes. Rhipidia with > 2 and up to 6 flow-

ers each. Flowers with tepal claws forming a wide cup,

bearing prominent nectaries with dry nectar; filaments

inserted in lower third of anthers; anther lobes paral-

lel and contiguous above filament insertion when fully

dehisced. Ovary exserted. ovoid; stigmas minute, termi-

nal on apices of style branch arms. Capsules globose- to

ovoid-truncate.

1 . Ferraria glutinosa (Baker) Rendle , in W.P.Hiem,

Catalogue of the African plants collected by Dr. Frie-

drich in 1853-61. vol. 1,2: 27 (1899); De Vos: 329

(1979); Goldblatt: 11 (1993). Moraea glutinosa Baker:

271 (1878). Type: Angola, Huila, near Lopollo, Ferrao

da Sola, February and April 1860, Welwitsch 1543 (BM,

holo.!; K!, LISU, P!, iso.).

F. bechnanica Baker: 344 (1898). Type: Bechuanaland [Botswana],

Ngamiland, Kalahari Desert near Manumwe, ‘flowers chocolate with

yellow fringe’, 26 Feb. 1897, Lugard 237 (K, holo.!).

Moraea randii Rendle in Rand: 144 (1898). Ferraria randii (Ren-

dle) Rendle: 54 (1905). Type: Southern Rhodesia [Zimbabwe], Bula-

wayo, Jan. 1898, Rand 223 (BM, lecto.! here designated; BR, iso.).

F. viscaria Schinz: 77 (1900). Type: South West Africa [Namibia],

Omupanda, Wulfhorst 45 (Z, holo.).

IMoraea malangensis Baker: 862 (1901). Type: Angola, Malange,

without date, Expedition von Mechow 386 (B, holo., no longer extant,

R. Vogt pers. comm.).

Plants mostly 400-600 mm high. Stem exposed above

sheaths of lower leaves, with 1(2) branches from upper

nodes, branches rarely also branched; distal 5-6 mm

of upper intemodes sticky, often with sand adhering.

Leaves at base solitary or up to 4, sometimes basal leaf

poorly developed; blades linear, straight, 4 — 6(— 1 0) mm

wide, with visible central vein, cauline leaves moder-

ately well developed, often longer than basal leaf, pro-

gressively reduced above and then largely sheathing.

Rhipidia 4— 6-flowered; inner spathe 30-42 mm long,

outer entirely sheathing, usually ± 7 to V3 as long as

inner. Flowers on pedicels slightly longer than spathes,

lasting a single day, usually dark brown to dull purple or

maroon; tepal limbs with pale yellow to gold margins,

sometimes limbs marked or sparsely spotted yellow

near base or with large irregular brown blotches on pale

background; claws brown or yellow streaked with pur-

ple or brown, odourless; tepals diverging, claws forming

wide basin ± 10 mm deep, 15-18 mm wide at rim, limbs

spreading ± horizontally to reflexed; margins crisped;

nectaries dark brown, 1.5 x 2.0 mm, 3-4 mm above

claw bases; outer tepals 23-33 x ± 10 mm, claws usu-

ally slightly longer and ± as wide as limbs; inner tepals

slightly smaller, claws tapering to narrow base. Stamens

with filaments united in a minutely puberulous column

8-11 mm long, free in upper 1. 5-2.0 mm; anther thecae

diverging below, parallel above filament insertion, 5-6

mm long before anthesis, 3-4 mm long after anthesis,

dark brown; pollen orange. Ovary narrowly ovoid, usu-

ally exserted. 5-7 mm long; style branches 2 mm long,

dividing into diverging, prominently fringed arms 2-3

mm long, fringes smooth or minutely papillate, 4-5 mm

long; stigmas terminal on tips of style arms. Capsules

ovoid- to globose-truncate, mostly 12-18 mm long.

Seeds glossy, brown, 4-5 mm at longest axis, ± 5-sided,

facets separated by raised ridges, facet surfaces wrin-

kled. Flowering time : mostly Jan.-Mar., occasionally

later. Figure 1; Plate 1G.

Distribution and biology. Ferraria glutinosa occurs

across a wide belt of summer rainfall southern and cen-

tral Africa; it is relatively common across Botswana and

northern Namibia from where it extends northward to

southern Angola, and westward to the North-West Prov-

ince of South Africa and western Zimbabwe, but with an

isolated record from Malawi (Figure 2). We have seen

no collections from Zambia, where it is likely to occur.

F. glutinosa is usually found in sandy ground with the

corms deeply buried, but sometimes in rocky places.

Several collections specifically mention white sand habi-

tats and others red sand.

Diagnosis and relationships'. Ferraria species in

tropical Africa and summer rainfall southern Africa dif-

fer from all species of the southern African winter rain-

fall zone in their open-branching habit, steins with sticky

exudate below the nodes, rhipidia with several (usu-

ally 4—6) flowers, and subglobose, flat-topped capsules.

The genus in tropical Africa is, nevertheless, unusually

variable as regards plant height, presence or absence of

foliage leaves on the flowering stem, proportions of the

12

Bothalia 41,1 (2011)

FIGURE I . — Ferraria g/utinosa. A, flowering stem; B, flower, side view; C, outer tepal; D, inner tepal; E, staminal column and style,

undehisced anther indicated by broken line; F, anther. Scale bar: A, B, 10 mm; C, D, 5 mm; E, F, 1.5 mm. Artist: John Manning.

Bothalia 41,1 (2011)

13

inner and outer rhipidial spathes, and flower colour and

patterning. It is not altogether surprising then, that 12

species have been described for the area between North-

West Province of South Africa and western Angola,

southern Congo and Zimbabwe. Ferraria glutinosa ,

based on Moraea glutinosa , one of the first four of these

named species, was first collected by Friedrich Wel-

witsch in western Angola in the years 1855-1861, and is

the name used until now for all populations of Ferraria

in tropical Africa (De Vos 1979; Goldblatt 1993).

Three other Welwitsch collections of Ferraria from

Angola formed the basis for Moraea andongensis, M.

candelabrum and M. spithamea, named by J.G. Baker

(1878) at the same time as F. glutinosa. Five more spe-

cies were subsequently described by Baker (1892,

1901), four from Angola and one from Botswana (then

Bechuanaland); one more species was described from

Namibia (Schinz 1900); one from Zimbabwe (Rendle in

Rand 1898); and one from Congo (Bolus 1932a). Five

of these were placed in the synonymy of Ferraria gluti-

nosa by Carter (1963), who suggested that the Namibian

F. viscaria was probably also conspecific. Following

Carter’s lead, De Vos (1979) treated all 12 named spe-

cies from tropical Africa as synonyms of F. glutinosa.

Her taxonomy was uncritically accepted by Goldblatt

(1993) for Flora zambesiaca and by Geerinck (2005) for

Flore d’Afrique Centrale.

We have examined most of the available herbarium

collections of tropical African Ferraria and conclude

that recognition of a single tropical species does not

reflect the extent of the variation among Ferraria popu-

lations there. Most importantly, there are two major veg-

etative types, one with only short sheathing leaves pro-

duced by the flowering stem in plants flowering mainly

in October to mid-December, and another with at least

one and up to eight well-developed basal leaves with

unifacial blades and a relatively prominent central vein

in plants flowering in (December) January to March.

In the light of the variation, which is closely correlated

with flower size and colouring, tepal patterning, relative

proportions of the inner and outer inflorescence spathes,

flowering time, and partly with geography, we propose

an alternative taxonomy, recognizing four species, thus

to a large extent validating Baker’s (1878) original inter-

pretation of Ferraria in tropical Africa.

Ferraria glutinosa , largest-flowered of the tropical

species, is distinguished by the presence of several foli-

age leaves with well-developed blades, at least one of

which is basal, and dark brown to purple flowers. Sev-

eral specimens have the basal leaf virtually vestigial, but

longer cauline leaves (e.g. Evles 8543). Also exceptional

are collections from Gweru [Gwelo] and Lomagundi in

Zimbabwe (Holland s.n.; Rutherford-Smith 593), which

respectively have three or six basal leaves. The inner

inflorescence spathes are (30-)35-42 mm long, with

the outer half or sometimes only one third as long. The

brown to dark purple (also described as brown-violet,

bright mauve or dark maroon) flowers have tepal limbs

edged with yellow, sometimes also marked with yel-

low, and are produced mainly from January to March.

The plants described as F. bechuanica from Botswana

by Baker (1898), Moraea randii by Rendle (in Rand

1898) from Zimbabwe, and F. viscaria from Namibia by

Schinz (1900) are the same species, but we note that the

flowers of the last-named, F. viscaria, were described

as yellow. We also provisionally include F. malangen-

sis (Baker 1901 ) in F. glutinosa because its flowers were

described as violet, but details of the leaves and flower-

ing time, essential for confident identification, are not

known. The type at the Berlin Herbarium is evidently

lost (R. Vogt pers. comm. 2008).

Like the other species of Ferraria from tropical

Africa, F. glutinosa is remarkable in having sticky inter-

nodes, a feature absent in the southern African winter

rainfall species of the genus, but present in a few species

of related genera of the southern African Moraea (only

subgen. Visciramosa) and two species of Bobartia (Strid

1974; Goldblatt 1986). Rand (1898) noted that ants are

trapped in the sticky exudate and suggested that this was

a defence against these insects, which we assume might

consume nectar, the reward for legitimate pollinators.

The fairly open branching pattern of the tropical

species is also quite different from the crowded, short

branches of other species of Ferraria and this, as well

as the ovoid-truncate capsule and several-flowered

rhipidia suggest an ancestral position in the genus. The

morphological indicators of its unspecialized status are

confirmed by molecular analysis using plastid DNA

sequences, which places F. glutinosa as sister to the win-

ter rainfall southern African species of the genus that

were included in a preliminary analysis.

Both the common tropical African Ferraria wel-

witschii and the related but rare F. spithamea are smaller

plants, notable for lacking foliage leaves on flowering

plants. They also have flowers somewhat smaller than

in F. glutinosa, with yellow to buff tepals (in F. wel-

witschii with the limbs marked throughout with small

brown to dark red spots, more densely so on the claws),

and large dark brown nectaries ± in the middle of the

claws (illustrated in Geerinck 2005: plates 9A, B, as F.

glutinosa). The rhipidia typically have the outer spathe

slightly more than half to two thirds as long as the inner.

Both these species bloom mainly in November to mid-

December, but have been recorded in flowering as early

as October.

The corms of Ferraria glutinosa are eaten raw or

roasted by the Kwanyama Ovambo, according to the

14

Bothalia 41,1 (2011)

ethnobotanist Robert Rodin (1985). Except for its value

as an occasional ornamental in the garden or in contain-

ers, we know of no other human use of this or any other

Ferraria species. It is noteworthy in the context of Rod-

in’s statement about the edibility of conus, that Mann-

heimer et at. (2008) report that Ferraria glutinosa is

poisonous to stock, with symptoms including anorexia,

increased heart rate, rapid respiration, diarrhoea and apa-

thy. Post mortem signs are cyanosis, hyperaemia of the

lung, and bloody inflammatory areas of the stomach and

intestine. No southern African species is known to be

toxic and we have seen cattle grazing on F. divaricata ,

but evidently not relishing it, for animals rarely took

more than a mouthful before moving on.

Representative specimens

ANGOLA. — Huila: without precise locality or date, Dekindt 493

(LISC) [specimen incomplete and without base], Huambo: Chianga,

± 1 700 m, 3 Nov. 1962, Teixeira & Andrada 6525 (LISC). Cunene:

Cuvelai, 11 Dec. 1972, Menezes 4270 (LISC).

BOTSWANA. — 1825 (Panda-ma-Tenga): 13 km toward Jol-

lies Pan, flowers yellow, (-DC), 23 Dec. 1996, Bruyns 6912 (NBG).

2033 (Chibabava): near Kwebe Hills, (-CA), Dec. 1898, Lugard 282

(K). 2121 (Ghanzi): Groot Laagte (East) fossil river valley, (-AD), 16

Mar. 1980, Smith 3176 (MO, PRE, SRGH). 2322 (Kang): Phuduhudu,

(-DD), 12 Dec. 1989, Barnard 499 (PRE). 2421 (Tshane): Kalahari

Park, SW of Ritchie’s Pan, white sand savanna, (-AC), Mar. 1979, Van

der Walt 5807 ( PRE).

MALAWI. — Northern: ± 10 km north of Mpherembe, sand at edge

of thicket, brown with yellow centre, 1 180 m, 7 Feb. 1987, La Croix

957 (MO).

NAMIBIA. — 1915 (Okakuejo): Ovamboland, Oshikango, brown-

purple with yellow spots, 1 May 1973, Rodin 9360 (MO, WIND).

1917 (Tsumeb): Tsumeb, 25 Jan. 1911, Dinter 1840 (SAM). 1920

(Tsumkwe): 6 km east of Tsumkwe along Botswana border, white

sand, (-DA), 13 Jan. 1971, Giess. Watt & Snyman 11027 (PRE); 157

miles [251 km] east of Grootfontein, Simkue, (-DA), 16 Jan. 1958,

Stoiy 6156B (PRE, with colour slides). 2017 (Waterberg): Otji-

warongo, turn-off to Waterberg, (-DA), 24 Mar. 1987, Maggs 83

(WIND). 2115 (Karibib): Donkerhuk 91, sandy ground, (-DD), 2 Mar.

1965, Barnard 79 (PRE); Erongo Siding, deep Kalahari sand, (-DB),

22 Apr. (fr.), Bean, Vlok & Viviers 1837 (BOL, MO); Namibrand,

Altenbronn, (-DD), 10 Mar. 1964, Seydel 264 (B). 2117 (Otjosondu):

Waterberg, Quickbom, (-AA), 30 Mar. 1930, Bradfield 146 (PRE).

2118 (Steinhausen): 15 km along Kapps Farm road, Steinhausen to

Windhoek, sandy plain, (-CC), Mar. 1988, Goldhlatt & Manning

8805 (MO). 2216 (Otjimbingwe): dry rocky slopes of Kupferberg

Pass, SW of Windhoek, (-DB), 21 Mar. 1988, Goldblatt & Manning

8843 (MO, WIND). 2217 (Windhoek): Farm Frauenstein, (-AD), 18

Jan. 1976, Giess 13901 (WIND); Goreangab Dam near Windhoek, (-

CA), 21 Feb. 1965, Giess 8395 (NBG); Lichtenstein, (-CC), 20 Jan.

1923, Dinter 4311 (B). 2219 (Sandfontein): east of Gobabis, farm road

between Farm Isabella and Farm Etna, (-AD), 8 Mar. 2002, Bartsch

503 (WIND). 2316 (Nauchas): Khomas, Farm Lichtenstein North,

rock outcrop, (-DD), 3 Mar. 2002, Mannheimer 1732B (WIND). 2317

(Rehoboth): Rchoboth, (-AC), 13 Apr. 191 1, Dinter s.n. (SAM).

ZIMBABWE. — Mazoe, Umvukwes, 5000 ft [I 525 m], Ruorka

Ranch, 17 Dec. 1952, Wild 3911 (MO, K, PRE, SRGH). Lomagundi,

Dyke near Rod Camp Mine, flowers yellow, spotted purple, 22 Feb.

1961, Rutherford-Smith 593 (K). Mangula Township, 17 Dec. 1961,

Jacobsen 1990 (PRE). Plumtree, Jan. 1936, Eyles 8543 (K) [without

basal foliage leaves]. Gwelo, 29 Dec. 1929 (cultivated at Kirsten-

bosch), Holland s.n. (BOL as National Botanic Gardens 92/28, K).

Matobo, Farm Besna Kobila, grassland, Dec. 1955, Miller 3215 (PRE).

NORTHERN CAPE. — 2723 (Kuruman): Kuruman, Esperanza,

(-AD), Apr. 1940, Esterhuysen 2219 (BOL); Kormutsethla, (-BB), 2

Feb. 1934, Cross 1088( PRE).

2. Ferraria candelabrum (Baker) Rendle , Cata-

logue of the African plants collected by Dr. Friedrich

in 1853-61, vol. I, 2: 27 (1899). Moraea candelabrum

Baker: 271 (1878). Type: Angola, Huila, Morro de

Lopollo, Apr. 1860, Welwitsch 1544 (BM, lecto.!, desig-

nated by De Vos, 1 979: 331; K!, LISU, iso.).

? Moraea andongensis Baker: 271 (1878). F. andongensis (Baker)

Rendle: 27 (1899). Type: Angola, Pungo Andongo, Mutollo near

Pedras de Guinga, marshy, low grassland, Jan. (fr. Mar.), Welwitsch

1532 (BM, lecto.!, designated by De Vos: 331 (1979); K!, LISU, iso.).

M. kitambensis Baker: 575 (1898). Type: Angola, Bangala, Kitam-

ba, on Cuango River, in swamp, 1880, Buchner 679 (B, holo., no

longer extant (R. Vogt pers. comm); K, iso!, fragment and drawing of

B specimen).

Plants mostly 600-800 mm high. Stem enclosed

below by overlapping leaf bases, exposed above,

branched from upper nodes, branches sometimes 2(3)

per node, and primary branches themselves often

l(2)-branched; upper nodes and distal parts of inter-

nodes sticky, often with sand adhering. Leaves at base

5-8, ± narrowly sword-shaped-linear, straight, 8-10 mm

wide, with visible central vein; margins not thickened;

cauline leaves well developed, shorter than basal, pro-

gressively reduced above and becoming largely sheath-

ing and bract-like. Rhipidia at least 3- or 4-flowered;

inner spathe 28-32 mm long, outer entirely sheathing,

usually ± 7 (-V4) as long as inner. Flowers on pedicels

slightly longer than spathes, ?lasting a single day, brown

with yellow markings or pale yellow, evidently odour-

less; tepals ascending with claws forming a cup 11-12

mm deep, ± 9 mm wide at rim, limbs ± spreading, ± 1 0

mm long, margins crisped; nectaries not evident; outer

tepals 20-25 mm long, claws 11-13 mm long, inner

tepals slightly smaller, claws tapering to narrow base.

Stamens with filaments united in a column ± 1 1 mm

long, free in upper ± 1 .5 mm; anther thecae ± paral-

lel, ± 3 mm long, shorter after anthesis. Ovaty usually

exserted, narrowly ovoid, ± 3 mm long; style branches

± 1.5 mm long, dividing into diverging, prominently

fringed arms ± 2 mm long; stigmas terminal on tips of

style arms. Capsules globose-truncate, ± 8 mm long.

Seeds unknown. Flowering time : Feb.-Apr., possibly

also in Jan.

Distribution and biology', known to us from just four

collections, none with well-preserved flowers, Ferraria

candelabrum is recorded from central Angola and adja-

cent western Zambia (Figure 3). Plants grow in rocky,

well-drained sites. Collections usually mention hilly

places, but the type of the synonym, Moraea kitamben-

FIGURE 3. — Known distribution of Ferraria spithamea , O; F. cande-

labrum, •.

Bothalia 41,1 (2011)

15

sis, is said to be from a swampy place (which seems

unlikely). The type of Moraea andongensis , provision-

ally assigned here, is from marshy, low grassland.

Diagnosis and relationships : none of the collections

of Ferraria candelabrum has well-preserved flowers

but the colour was described as yellow in the type col-

lection, brownish in one other, and buff with dark brown

stripes and spots in a third. Despite the lack of precise

floral details, the vegetative habit is unique among the

tropical African species. Plants stand up to 800 mm

high, and have a basal fan of several (up to 8) long, rela-

tively soft-textured leaves. The stem is branched repeat-

edly, the branches terminating in unusually slender rhi-

pidia about 30 mm long. Notably the outer of the two

rhipidial sheaths is~V3-'/4 as long^as the inner, giving the

plants a distinctive appearance. Ferraria candelabrum

is no doubt closely allied to the widespread subtropical

African F. glutinosa and was included in that species by

De Vos (1979). The unusually narrow rhipidia and short

outer spathe makes the species easy to recognize. Unlike

F. candelabrum, F. glutinosa has a single or rarely up to

three basal leaves, sometimes poorly developed at flow-

ering, a stem usually with the primary branches them-

selves only occasionally branched and large capsules

12-18 mm long, compared to capsules ± 8 mm long,

known only from the somewhat atypical collection from

Lubango in Angola (De Menezes 1661).

We provisionally place Ferraria andongensis in the

synonymy of F. candelabrum. Although flowering in

January and fruiting in March, plants of the type collec-

tion are much less robust, at most only 300 mm tall, and

have only one basal leaf and a second, well-developed

cauline leaf, both with narrow blades, ± 2.5 mm wide.

The outer inflorescence spathes are short, about a third

as long as the inner, and this as well as the small cap-

sules are consistent with F. candelabrum. Welwitsch, in

his notes, described the flowers as dull sulphur-yellow.

The anthers are short, ± 2 mm long. Little more can be

deduced from the type material.

Representative specimens

ANGOLA. — Huila: Lubango, on plateau of Ponta do Lubango, 1 1

Apr. 1965, De Menezes 1661 (LISC).

Also known from Angola from the types of the species cited above.

ZAMBIA. — Northwestern: Mwinilunga District: Ikelenge, Milomba

Hill, among rocks, 1 440 m, 22 Feb. 1995, Zimba et al. 632 (MO, PRE);

Mwinilunga, base of Kalene Hill, 1 400 m, sandy bank, flowers buff with

dark brown stripes and spots, 22 Feb. 1975, Hooper & Townsend 327 (K).

3. Ferraria spithamea (Baker) Goldblatt &

J.C. Manning, comb. nov.

Moraea spithamea Baker in Transactions of the Linnean Society,

London, ser. 2, Botany 1: 271 (1878). Type: Angola, sandy places

about Humpata and Lopollo, Oct. 1859 (fl. and fr.), Welwitsch 1547

(BM, lecto.!, designated by De Vos: 331 (1979); K, iso.!).

Plants mostly 100-160 mm high. Stem exposed

above sheaths of cauline leaves; upper nodes and distal

parts of intemodes sticky, often with sand adhering, 2

or 3 intemodes long, with a branch at each node, thus

2- or 3 -branched. Leaves of flowering plants absent at

base, borne at aerial nodes, partly sheathing, channelled

throughout, 20^10 mm long; leaves of vegetative plants

unknown. Rhipidia at least 3-flowered; inner spathe

25-30 mm long, outer entirely sheathing, ± half as long

as inner. Flowers on pedicels as long or slightly longer

than spathes, ?lasting a single day, yellow, marked with

brown spots toward base of tepals, tepals ascending,

claws forming a cup, ± 8 mm deep, ± 7.5 mm wide at

rim, limbs spreading, margins slightly undulate (not

crisped); nectaries ± 1 .5 mm diam., in lower third of

outer tepal claws, in middle of inner tepal claws, col-

our unknown; outer tepals ± 20 mm long, inner ± 18

mm long. Stamens with filaments united in column ± 8

mm long, free in upper ± 1 mm; anther thecae parallel

or diverging basally, ± 3 mm long. Ovary oblong to nar-

rowly ovoid, ± 3 mm long, included or exserted 2-3 mm

at anthesis; style branches ± 1.3 mm long, dividing into

two divergent, entire arms ± 2 mm long; stigmas termi-

nal on style arms. Capsules globose-truncate, ± 5 mm

long, exserted from spathes, smooth. Seeds unknown.

Flowering time : Oct.-Dec.

Distribution and biology. Ferraria spithamea is

a narrow endemic of the highlands of southwestern

Angola (Figure 3). Its habitat is described as thorny

thicket in sand, or among rocks.

Diagnosis and relationships', poorly understood,

Ferraria spithamea is known to us from just two col-

lections, the type and one other, both from southwest-

ern Angola. The species is recognized primarily by the

entire, arching style arms, without the fringes character-

istic of other species of the genus, but also by the pale

yellow flowers with the tepal limbs at best undulate,

and not crisped as they are in other species. Plants lack

foliage leaves at flowering and we assume the foliage

leaves are developed later in the season from separate

shoots. The growth form is thus exactly as in the fairly

widespread tropical African F. welwitschii but that spe-

cies differs in the slightly smaller flowers, also yellow

to buff, but with the tepal limbs and distal parts of the

claws liberally scattered with minute dark spots. Nor-

mally a taller plant, F. welwitschii may reach 350 mm,

therefore about twice as tall as F. spithamea and, typical

of Ferraria, it has crisped tepal limb margins and promi-

nently fringed style arms. We have relied in part for

the description of the flowers of F. spithamea on Wel-

witsch’s notes which describe the yellow tepal colour

(flava) marked from the middle to the base with small

dark marks (parvis nigris picta).

The lack of style branch fringing and tepal limbs

without crisped margins are anomalous in Ferraria but

the vegetative form, distally sticky internodes and dis-

tinctive corm, leave us in no doubt that the species is

correctly assigned to the genus.

Representative specimen

ANGOLA. — Huila: Sa de Bandeira, Bata-Bata, 6 Dec. 1961, San-

tos 657 (LISC).

4. Ferraria welwitschii Baker, Handbook of the

Irideae 74 (1892). Type; Angola, locality unknown, cul-

tivated in England (Hort. Saunders), Welwitsch s.n. (K,

holo.! drawing only with tepal painted in watercolour).

? Moraea viscosa R.C. Foster: 48 (1936), as a new name for M.

aurantiaca Baker: 575 (1898), nom. illegit. non A. Dietrich: 485

(1832). Type: Angola, Malange, Oct. 1879, Expedition von Mechow

303 (B, holo.! (fragment), K, drawing!).

16

Bothalia 41,1 (2011)

Moraea aurantiaca Baker: 575 (1898). Type: Angola, Malange,

October 1879, Expedition von Mechow 303 (B, holo.! (fragment), K,

drawing!).

F. hirschbergii L. Bolus: 57 (1932a). Type: Congo, [Shaba], near

Lubumbashi (Elisabethville), von Hirschberg s.n., cultivated at

National Botanic Gardens, Kirstenbosch, 615/29 (BOL, holo.!; K (two

sheets)!, SAM, iso.!).

Plants ( 1 20~) 1 80—350 mm high. Stem exposed above

sheaths of lower leaves; upper nodes and distal parts

of internodes sticky, often with sand adhering. Leaves

of flowering plants sheathing, occasionally with short

blades shorter than sheaths; of vegetative plants (1)2,

narrowly sword-shaped, straight or falcate, 1. 5-3.0 mm

wide, with visible main vein. Rhipidia 4- or 5-flowered;

inner spathe 20-34 mm long, outer entirely sheathing,

mostly V to ± 2/3 as long as inner, 10-17 mm. Flowers

on pedicels ± as long as spathes, lasting a single day,

pale to dull yellow to buff (?also orange) with brown

to dark red or dull green spots on limbs and distal part

of claws, evidently odourless, tepals ascending, claws

forming a cup, 7-9 mm deep, ± 7 mm wide at rim, limbs

spreading to reflexed up to 40°, margins crisped; nec-

taries dark brown, ± 1.5 x 2 mm, ± in centre of claws;

outer tepals 15-20 x ± 5 mm, inner 14-20 mm long.

Stamens with filaments united in a column 5. 0-7. 5 mm

long, free in upper 1.0-1. 5 mm; anther thecae parallel or

diverging basally, 1.5-2. 5 mm long, slightly shorter after

anthesis. Ovary’ usually exserted, oblong to narrowly

ovoid, 2-3 mm long; style branches 1. 5-2.0 mm long,

dividing into diverging, fringed arms; stigmas terminal

on style arms. Capsules globose-truncate, mostly 5-7

mm long, exserted from spathes, smooth or minutely

warty (Mendes 1957). Seeds angular, ± prismatic, 3 nun

long, glossy, brown with pale, raised angles, ± 5-sided,

facet surfaces undulate to wrinkled. Flowering time :

mostly Oct. to early Dec. (early Jan.).

Distribution and biology. Ferraria welwitschii occurs

across a wide belt in central Africa from the Malange

highlands in west-central Angola, across Zambia to

southern Congo and Zimbabwe (Figure 4). Records

indicate that plants grow on the margins of open wood-

land, often in rocky ground and in fairly moist habitats,

including riverbanks and dambos.

FIGURE 4.' Known distribution of Ferraria welwitschii.

Diagnosis and relationships : the type collection of

Ferraria welwitschii is a pencil drawing of a plant cul-

tivated in Britain, with one of the tepals coloured. The

plant evidently stood ± 300 mm high, lacked developed

basal foliage leaves and had flowers with reflexed, dull

yellow tepal limbs covered with fine, dark brown spots.

Friedrich Welwitsch is believed to have collected the

plant but the original locality is not recorded. Plants

matching this collection, although not always as tall,

have been recorded widely across Angola, Zambia,

Congo and parts of Zimbabwe, always flowering early

in the wet season, October to mid-December, and con-

sistently lacking well-developed foliage leaves. Even

those specimens in fruit (e.g. Cruse s.n. from Zambia

collected in late December) lack foliage leaves; other

collections (e.g. Mendes 1957 from Angola in fruit in

January) have short, narrow foliage leaves emerging at

the base, evidently representing late-developing foliage

leaves on a shoot lateral to the main axis. It is clear that

flowering plants do not produce leaves later in the sea-

son but rely on the stems and sheathing leaves for pho-

tosynthesis and production of storage carbohydrates for

the new conns developed after flowering. This pattern

recalls tropical African Gladiolus unguiculatus Baker;

flowering and fruiting specimens of this species have

short or vestigial foliage leaves but plants remain green

even after the capsules have ripened and the seeds are

shed in December (Goldblatt 1996). Corms of both spe-

cies eventually produce new foliage leaves from shoots

lateral to the fruiting axis, after the capsules have rip-

ened. Vegetative specimens of F. spithamea have one

or more well-developed leaves and one or two smaller

leaves in a basal tuft.

Evidently the closely allied Ferraria spithamea from

southwestern Angola has a similar growth form, but the

yellow tepals lack the characteristic small dark spots,

have at best undulate margins (never crisped), and the

style branches lack the feathery fringes of all other Fer-

raria species.

The inclusion of Ferraria welwitschii, and for that

matter F. spithamea, in the larger-flowered F. glutinosa

(e.g. Carter 1963; De Vos 1979; Geerinck 2005), a taller,

more robust plant with brown or purple (to maroon) or

sometimes partly yellow flowers, and often bears one or

more foliage leaves and well-developed cauline leaves

on flowering individuals, now seems mistaken. The

flowers of F. glutinosa have tepal limbs edged in yellow

and bloom mainly from January to March, rarely in late

November or December. Colour photographs in Flore

d’Afrique Centrale (Geerinck 2005) show the flower

form, colour and patterning of F. welwitschii clearly and

they contrast starkly with photographs of F. glutinosa

(e.g. Story 6165B PRE; Mannheimer et al. 2008) and

our illustration of the species (Figure 1). We include F.

hirschbergii in the syonymy of F. welwitschii, the type

of which closely matches that species. We provision-

ally also include a second heterotypic synonym, Moraea

aurantiaca, in F. welwitschii. The type, at the Berlin

Herbarium, is fragmentary, but consists of an apparently

leafless plant without flowers, but with small rounded

capsules. We assume the species was called aurantiaca

because the flowers were orange (or thought to be so)

but the basis for this is not evident to us. Duplicates of

Bothalia 41,1 (2011)

17

the type collection, if they exist, may help establish the

identity of this name. The flowering time, October, and

absence of leaves are consistent with our provisional

assignment to F. welwitschii.

Representative specimens

ANGOLA. — Huila: Ganguelas, Vila Artur de Paiva, banks of

the Cubango, 1 450 m, 4 Jan. 1960, Mendes 1957 (fit), (LISC, MO);

Huambo, outskirts of Nova Lisboa, 1 750 m, 8 Nov. 1970, da Silva

3340 (K, LISC, PRE); Caala to Cuima, 26 Nov. 1959, Stopp 122 (K).

Lubango, Hoque, on road to Dinde, 5 km, 9 Nov. 1962, De Menezes

340 (K, LISC, PRE). Cuanza Sul: Cela-Cassamba, ± 1 350 m, 28 Sept.

1963, Texeira & All 7396 (LISC).

CONGO. — Shaba: Lubumbashi, woodland and savanna, 1 300 m,

6 Dec. 1968, Lewalle s.n. (MO); near Lubumbashi. banks of the Nat-

webo, 25 Oct. 1970, Lisowski 108 (B, BR. K); Lubumbashi. Paturage

de la Karavia, Nov. 1933, Quarre 3633 (BR).

ZAMBIA. — Northwestern: Mwinilunga: Kalenda Ridge, W of

Matonchi Farm, stony ground, 8 Oct. 1937 (fr. 16 Dec. 1937), Milne-

Redhead 2664 (K).

ZAMBIA. — Western: Luano Forest Reserve, Chingola, in granite

rocks, 19 Nov. 1961, Liriley 218 (K, MO, PRE, SRGH).

ZAMBIA. — Copperbelt: Kitwe, dambo margin, 13 Dec. 1959,

Fanshawe 5314 (K, NDO); Mufulira, dambo, 10 Nov. 1947, Cruse 92

(K), late Dec. 1947 (ft.), Cruse s.n. (K); Kawambwa Dist., 900 m, by

M’bereshi River, 2 Dec. 1961, Richards 15485 (K); Mazabuka. Ridge-

way road, 2 Dec. 1931, Central research station 541 (K, PRE). Mum-

bwa Dist., 30 miles [48 km] west of Kafue Hoek pontoon on road to

Mankoya. 21 Nov. 1959, Drummond & Cookson 6734 (K, MO, PRE.

SRGH).

ZIMBABWE. — Bulawayo, 25 Dec., "brown flowers’, Norman R44

(K); Matobo, Paddocks, black land, Dec. 1931. Rattray 440 (K); Farm

Shumbashaba, among rocks, Dec. 1954, Miller 2569 (K); Gokwe,

Charama road fly gate 4 miles [6 km] north of Gokwe, 2 Jan. 1963,

Bingham 393 (PRE).

B. Subgen. Ferraria

Plants acaulescent or with aerial stems with relatively

short branches enclosed by leaf sheaths. Stem not sticky

below nodes. Rhipidia with only 2 flowers each. Flow-

ers with tepal claws widely diverging, forming a shal-

low cup, with large nectaries with non-fluid nectar, or

forming a deep, wide to narrow cup with small nectaries

and with fluid nectar; filaments inserted in lower third or

near apex of anthers; anther lobes parallel and usually

contiguous above filament insertion or lobes not contig-

uous and usually widely diverging when fully dehisced.

Ovary included, ellipsoid, with or without sterile beak;

stigmas minute, terminal on apices of style branch arms,

or larger, on lobes below style arm tips. Capsules ovoid-

oblong and obtuse or with prominent beak.

B1 . Sect. 1 . Ferraria

Flowers with tepal claws widely diverging, forming a

shallow cup. with large nectaries with non-fluid nectar;

filaments inserted in lower third of anthers; anther lobes

parallel and contiguous above filament insertion. Ovary

ellipsoid, sometimes with sterile beak; stigmas minute,

terminal on apices of style branch arms. Capsules ovoid-

oblong and obtuse or with prominent beak.

5. Ferraria crispa Burnt, in Nova acta physico-

medico Academiae Caesareae Leopoldino-Carolinae

germanicae naturae curiosorum 2: 199 (1761); De Vos:

338 (1979). Type: South Africa, without precise local-

ity, illustration in Burman, Nova acta physico-medico

Academiae Caesareae Leopoldino-Carolinae germanicae

naturae curiosorum 2: 199, t. 3, fig. 1 (1761).

F. undulata L.: 1353 (1763), nom. illegit. superfl. pro M. crispa

Burm.f. Moraea undulata (L.) Thunb.: no. 14 (1787). Type: South

Africa, without locality or collector, illustration in Miller: 187, t. 280

(1759a), lecto., here designated.

F. ensiformis Mill. (1768), nom. superfl. pro F. crispa Burm. Type.:

not designated.

F. obtusifolia Sweet: t. 148 (1826b). Type: illustration in Sweet, l.c.

(1826b).

F. major Eckl.: 18 (1827), nom. nud.

F. vandermenvei L. Bolus: 276, fig. D (1932b). Type: South Africa,

[Western Cape], near Swellendam, cultivated in Cape Town, Sept.-Oct.

1932, Van der Meme s.n. BOL20168 ( BOL, holo.!).

F. crispa subsp. nortieri M.P.de Vos: 338 (1979), syn. nov. Type:

South Africa, [Western Cape], heights north of Verlorenvlei, 28 Aug.

1976, De Vos 2366 (NBG, holo.!).

See De Vos (1979: 338) for additional synonymy.

Plants usually robust, (300— )450— 1 500 mm high.

Stem much branched in upper half, sheathed by leaf

bases below; cataphylls and bases of leaf sheaths often

speckled pale on red background. Leaves several, linear

to sword-shaped, (4-)6-12 mm wide, basal leaves long-

est, with visible main vein. Rhipidia 2-flowered; spathes

green with membranous margins, inner 45-65 mm long,

outer ± half as long, usually entirely sheathing. Flow-

ers on pedicels 40-60 mm long, lasting a single day,

variously pale yellow to beige with brown to dull pur-

ple mottling and brown margins to predominantly brown

with pale yellow margins, strongly scented, odour remi-

niscent of caramel or molasses, tepals widely diverging,

forming a shallow bowl 6-8 mm deep, 13-18 mm wide

at mouth, nectary ± 7 length of claw, sometimes slightly

longer, limbs spreading horizontally; outer tepals 28-35

mm long, claws 8-12 mm long, limbs to 25 mm long,

inner tepals slightly shorter and narrower. Stamens with

filaments united in smooth column ± 6 mm long, free

in upper 3 mm; anthers ± 3 mm long, thecae parallel,

shortly apiculate; pollen orange. Ovary spindle-shaped,

not beaked, included in spathes, 12-20 mm long; style

branches ± 4 mm long, forked in upper half, divided into

diverging arms; 4—5 mm long, prominently fringed: stig-

mas terminal on style arms. Capsules ovoid to oblong,

1 5-25 mm long, obtuse or pointed at apex. Seeds angu-

lar, mostly 5-sided, ± 4 mm long, facets slightly wrin-

kled. Flowering time : Aug.-Oct. Plate IE.

Distribution and habitat : Ferraria crispa is best

known as a coastal species and it is common along the

Western Cape coast in rocky sites from Lambert’s Bay

to Hermanus; it also occurs sporadically further east

along the coast at least to Herold’s Bay near George

(Figure 5). Showing a striking ecological shift, it also

grows in montane habitats, sometimes along streams in

deep, coarse sand, or on seasonally moist south-facing

slopes, from Pakhuis Pass in the north through the Wit-

teberg and Swartberg to Georgida in the Baviaanskloof

Mtns of Eastern Cape. Usually fairly robust in stature,

plants of montane populations can reach a remarkable

1.5 m in height, more than has been recorded in any

coastal population of the species. Unlike the coastal pop-

ulations which flower annually, the montane populations

of F. crispa flower well only in late spring following a

wild fire the previous summer.

18

Bothalia 41,1 (2011)

16° 18° 20° 22° 24°

_j i i 1 1 —

FIGURE 5. — Known distribution of Ferraria crispa.

Long known by the later synonym, Ferraria undulata

L., dating from 1763 (e.g. Baker 1892, 1896), the spe-

cies was first described by Johannes Burman in 1761 as

F. crispa. It is the most well-known species of the genus

and the name F. crispa (or F. undulata ) is often care-

lessly applied to other members of the genus. The spe-

cies is best recognized by the habit, with stems produced

well above the ground and bearing numerous branches

crowded in the upper half of the stem, combined with a

flower with a wide, shallow cup and an ovoid capsule

lacking a beak. The sword-shaped lower leaves and the

shorter cauline leaves have a definite main vein and

form two-opposed vertical ranks. The strongly scented

flowers have tepals speckled or blotched dark brown to

dull purple on a cream-coloured to pale yellowish back-

ground, or occasionally predominantly dark brown, but

whatever the dominant colour, the crisped margins are

a pale colour, sometimes almost golden when the tepals

are otherwise brown. More important than their colour

and patterning, which is quite variable, the tepals form a

wide, fairly shallow cup and each claw has a large nec-

tary, often covering almost half the surface. The nectar-

ies produce minute droplets of concentrated nectar scat-

tered over the entire surface.

De Vos (1979) recognized the northern populations

of Ferraria crispa as subsp. nortieri because of their

pale green rather than slightly glaucous leaves, slightly

smaller flowers, the perianth with a cup ± 8 mm deep

and ± 1 0 mm wide at the rim, and the ovary less than 20

mm long. The inner tepals are also slightly narrower, ±

6.5 mm wide versus ± 10 mm wide in the typical sub-

species. The distinction is less clear than De Vos indi-

cated and we prefer not to recognize these northern pop-

ulations formally. Notably, all plants of subsp. nortieri

for which chromosome numbers are available, are tetra-

ploid, 2 n = 40, whereas populations assigned to subsp.

crispa to the south and in the southern Cape mountains

are hexaploid, 2n = 60. The smaller flower dimensions,

and smaller pollen grain size noted by De Vos, seem

directly correlated with the lower ploidy level in these

northern populations.

The ecological range of Ferraria crispa , as noted

above is remarkable. We have looked in vain for differ-

ences between the coastal populations that favour rocky

outcrops and those from montane sandy slopes and in

the absence of any morphological distinction, apart from

greater height in the latter plants, they remain the same

taxon.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): 2 km from Lam-

bert’s Bay to Vredendal, (-AB), 5 Sept. 1976, De Vos 2374 (NBG);

Pakhuis Pass, road to Klein Kliphuis, (-BB), 4 Sept. 1976, De Vos

2368 (NBG). 3317 (Saldanha): Bobbejaankop, Saldanha, (-BB),

2 Sept. 1974, De Vos 2323 (NBG). 3318 (Cape Town): entrance to

West Coast National Park, calcareous slope, (-AA), 4 Aug. 2001,

Goldblatt & Manning 11665 (MO); Dassen Island, (-AC), Jan. 1926

(fr.), Lang sub Marloth 6696 (PRE); Robben Island, (-CC), Walgate

496 (NBG); Spieka near Klipheuwel, (-DA), 10 Sept. 1975, Thomp-

son 2617 (NBG). 3319 (Worcester): Hex River Pass, (-BD), 26 Sept.

1951, Barker 7454 (NBG). 3320 (Montagu): Montagu Baths, (-CC),

Oct. 1921, Page 74 (PRE). 3321 (Ladismith): Swartberg Mtns, road

to Gamkakloof, deep sand along stream, burned last summer, (-BD),

Goldblatt & Porter 11858 (MO, NBG); Gamkaberg Nature Reserve,

(-CB), 15 Aug. 1983, Cattell & Cattell 298 (NBG). 3322 (Oudt-

shoom): Swartberg Mtns, Meiringspoort, (-BC), Loubser sub De Vos

2351 (NBG). 3418 (Simonstown): Strandfontein, (-AB), 18 Sept.

1942, Compton 13705 (NBG). 3419 (Caledon): Kleinmond, (-AC),

De Vos 2360 (NBG); Hermanus, (-AC), Nov. 1921 (fr.), Rogers 22619

(PRE); Femkloof Nature Reserve, (-AD), 2 Oct. 1984, Drewe 154

(MO). 3420 (Bredasdorp): De Hoop, (-AD), 13 Sept. 1979, Burg-

ers 2217 (NBG). 3421 (Riversdale): Stilbaai, hill near tennis courts,

(-AD), 23 Nov. 1978 (fr.), Bohnen 4651 (NBG, PRE); 1 mile [1.6 km]

N of Gouritz River mouth, (BD), 18 Sept. 1968, Mauve 4755 (PRE).

EASTERN CAPE. — 3323 (Willowmore): Georgida, (-AD), Oct.

1930, Fourcade 4411 (NBG).

6. Ferraria foliosa G.J. Lewis in Annals of the

South African Museum 40: 117 (1954); De Vos: 366

(1979). Type: South Africa, [Western Cape], near

Elandsbaai [Elands Bay], 12 Sept. 1955, Lewis 2301

(, SAM60809 , holo.!; SAM (two sheets)!, STE!, PRE!,

iso.).

Plants 300-800 mm high. Stem leafy and much-

branched, branches rotated in clockwise fashion. Leaves :

basal (and juvenile) with linear blades diamond-shaped

in cross section, thickened in midline, ± 8-10 mm wide,

main vein hardly evident below thickened midline;

cauline leaves falcate, channelled below, distally hori-

zontal, main vein often not evident, arranged in a two-

ranked spiral. Rhipidia 2-flowered; inner spathe 48-50

mm long, often prominently inflated, outer 28-30 mm

long, sheathing below, diverging in upper half, hooked

at tips. Flowers on pedicels 17-20 mm long, lasting a

single day, dull maroon to purple or dark brown, usu-

ally with darker purple-maroon blotches, margins pale

brown, claws usually maroon speckled with white,

putrid- or molasses-smelling, tepal claws broad, forming

a wide cup, 7. 5-8.0 mm deep, 13-18 mm wide at rim,

nectaries pale green, heart-shaped, prominent, '/ to V, as

long as claw, limbs ± spreading; outer tepals 33^-0 mm

long, inner tepals 28-35 mm long, claws of both whorls

9-10 mm long. Stamens with filaments united in a col-

umn 8-11 mm long, free in upper 1.5-2. 5 mm; anthers

± 3.5 mm long before anthesis, thecae parallel; pollen

orange. Ovary spindle-shaped, not beaked, included,

15-20 mm long; style branches ± 4.5 mm long, diverg-

ing, with prominently fringed arms; stigmas terminal on

style arms. Capsules ellipsoid, 28-35 mm long, round

Bothalia 41,1 (2011)

19

at apex. Seeds angular, ± 5-sided, ± 3 mm long, facets

slightly wrinkled, light brown, shiny. Flowering time :

late Aug.-Oct. Plate 1C.

Distribution and biology, the vegetatively distinctive

Ferraria foliosa is restricted to dunes and sandy beaches

along the Atlantic coast of Northern and Western Cape

of South Africa (Figure 6). Plants have been recorded

from near Hondeklip Bay in the north to near Velddrif in

the south, where it overlaps the range of F. crispa. The

latter favours rocky sites in contrast to the deep sands,

mostly in strandveld close to the sea shore in which F.

foliosa grows. The dull purple-brown flowers have a

strong unpleasant, molasses-like or putrid odour and

attract numerous flies which successfully accomplish

cross pollination.

Vegetatively, Ferraria foliosa can be recognized

by the long, linear basal leaves much thickened in the

midline, thus diamond-shaped in cross section (see De

Vos 1979), and sometimes dry by flowering time. The

basal leaves differ markedly from the shorter, wider

cauline leaves, the blades of which are also thickened

in the middle and arch outward, ultimately spreading

horizontally. Successive leaves are opposed but col-

lectively the leaves have a spirally two-ranked arrange-

ment. This is also reflected in the short lateral branches,

which are crowded in the upper half of the stem, the suc-

cessive ones rotated slightly in a clockwise direction.

The flowers differ little from those of closely related F.

crispa in their darkly mottled pigmentation, broad tepal

claws with large nectaries, and wide, shallow floral cup.

Whereas the flowers of F. crispa are usually shades of

dull yellow, buff and brown, those of F. foliosa are mot-

tled dull purplish, or brownish maroon on a paler grey-

purple background.

In her key to the species, De Vos (1979) distinguished

Ferraria foliosa and F. schaeferi from F. crispa by the

leaf blades having numerous parallel veins rather than

a single prominent vein. That distinction seems to us to

poorly reflect their morphology, for the blades of F. foli-

16° 18° 20° 22° 24°

_l i l l i —

FIGURE 6. — Known distribution of Ferraria foliosa, O; F. schaeferi ,

osa are so thickened and leathery that no veins are evi-

dent except for the main vein in the centre of the thick-

ened blade. Ferraria foliosa is better distinguished from

F. crispa by the spirally rotated leaves and branches, and

secondarily by the duller flower coloration, the back-

ground tinged with purple and the mottling a maroon

to dull purple in F. foliosa that contrasts with the dark

brownish or dull yellow colouring on a pale cream-col-

oured background, or brown with yellow margins in F.

crispa. The cream-coloured perianth covered with dark

speckles and prominently beaked ovary, immediately

separate F. schaeferi from F. foliosa. The species appears

to have first been collected by Harry Bolus near Veld-

drif in 1892, but that specimen was overlooked by Lewis

and was referred to F. crispa by De Vos in her account of

Ferraria.

Representative specimens

NORTHERN CAPE. — 3017 (Hondeklipbaai): Farm Avontuur, 3

km E of Hondeklipbaai [Hondeklip Bay], (-AD), 25 Aug. 1986, Le

Roux & Lloyd 260 (NBG); sandy slopes above lighthouse at Groen

River Mouth, (-DC), 25 Aug. 2002, Goldblatl & Porter 12113 (MO);

Groen River Mouth, above highwater mark, (-DC), 29 Aug. 1977, De

Vos 2396 (PRE).

WESTERN CAPE. — 3117 (Lepelfontein): Brand-se-Baai, (-BD),

23 Aug. 1993, De Villiers 121 (PRE). 3118 (Vanrhynsdorp): Olifants

River, dunes at river mouth, (-CA), 8 Oct. 1983, O'Callaghan 649

(NBG). 3218 (Clanwilliam): Otterdam, Lambert’s Bay, (-AB), 6 Sept.

1953, Compton 24162 (NBG); sandy slope south of Leipoldtville,

Elandsbaai [Elands Bay] road, (-AD), 15 Sept. 2001, Goldblatt & Por-

ter 11886 (MO, NBG); sandy flats along coast between Nuwedam and

Soutkloof, S of Elandsbaai [Elands Bay], (-AD), 5 Oct. 2004, Gold-

blatt & Porter 12619 (MO, NBG); Elands Bay, sand below Bobbe-

jaankop, (-AD), 28 Aug. 1976, De Vos 2365 (NBG); Dwarskersbos,

strandveld, (-CA), 7 Oct. 2004, Goldblatt & Porter 12636 (NBG);

dunes at mouth of Berg River, (-CC), Oct. 1892, Bolus 6301 (BOL,

NBG, PRE).

7. Ferraria schaeferi Dinter in Repertorium

Novae Species Regni Vegetabilium 16: 339 (1920); De

Vos: 344 (1979). Type: South West Africa [Namibia],

Klinghardt Mtns, Dreikugelberg, 14 Aug. 1913, Schafer

562 (B, neo.!, designated by De Vos 1979: 344).

Plants 200-500 mm high. Stem sheathed by leaf

bases, branching repeatedly in upper axils, branches

rotated in clockwise spiral; with corm bearing long run-

ners from base, each terminating in a small corm. Leaves

with overlapping sheaths; blades thick and leathery,

thickened in midline and main vein usually evident,

arranged in a two-ranked spiral, spreading horizontally

above, 12-20 mm wide. Rhipidia 2-flowered; spathes

firm, leafy in texture, with translucent membranous

upper margins, inner 50-60 mm long, outer somewhat

shorter, sheathing only near base, arching outward in

upper half. Flowers on pedicels 40-50 mm long, lasting

a single day, creamy yellow with small to large brown

spots, margins dark brown, usually sweetly scented (of

violets), tepal claws broad, forming a wide cup, ± 8

mm deep, 12-15 mm wide at rim, with brownish pur-

ple, heart-shaped nectaries in lower half, ± xf as long

as claw; outer tepals ± 25 mm long, inner tepals 22-24

mm long, claws of both whorls ± 8 mm long. Stamens

with filaments united in a column ± 7.5 mm long, free

in upper 2 mm; anther thecae parallel, ± 5 mm long at

anthesis, later shrinking; pollen dull brown to orange.

Ovary spindle-shaped, 20-25 mm long, tapering to a

slender beak, ± 8 mm long, included; style branches ±

20

Bothalia 41,1 (2011)

2 mm long, diverging into prominently fringed arms ±

2 mm long; stigmas terminal on style arms. Capsules ±

ellipsoid, 20-25 mm long, excluding beak. Seeds light

brown, glossy, 5- or 6-angled, facets ± smooth. Flower-

ing time : mid-Jul.-Sept.

Distribution and biology, favouring deep sands,

mainly along the coast, or a short distance inland, Fer-

raria schaeferi extends from near Liideritz Bay in south-

western Namibia in the north, to the coast at Grootmis

in Northern Cape, South Africa in the south (Figure 6).

The type locality, the Klinghardt Mtns of southwesten

Namibia, lies some 40 km inland, which is somewhat

unusual but we have also seen F. schaeferi growing near

Brandkaros east of Alexander Bay, some 30 km inland,

along the Orange River. There is also an early record of

the species at Arrisdrif ( Marloth 12391), even further

inland, so F. schaeferi is not restricted to near-coastal

habitats. The inland stations benefit from coastal fog, the

additional precipitation allowing this otherwise coastal

species to extend into the interior.

Unlike the flowers of the related species, Ferraria

crispa and F. foliosa, those of F. schaeferi sometimes

have, at least to the human nose, a pleasant, sweet scent

reminiscent of violets, but with a slightly sour under-

tone. We have noted in cultivated plants that some have

sweeter-scented flowers at cool temperatures, whereas

others smell putrid both at cooler and warm termpera-

tures. Whatever the scent, the flowers of F. schaeferi are

visited exclusively by a range of small and large fly spe-

cies, mostly of the families Muscidae and Calliphoridae

that forage on the droplets of sweet nectar on the nectaries.

Diagnosis and relationships', with a basic floral mor-

phology very much like that of Ferraria crispa and F.

foliosa , F. schaeferi has widely spreading tepals, the broad

claws of which form a shallow cup, ± 8 mm deep and

some 12 mm wide at the rim. The tepal claws bear large

heart-shaped nectaries, while the limbs have a creamy pale

yellow ground colour speckled with small dark brown

spots, the same colour as the crisped margins. In vegeta-

tive form, Ferraria schaeferi most closely resembles F.

foliosa which also has the leaves and branches arranged

in a two-ranked spiral, and the leaf blades are similarly

thick and leathery with a weakly defined central vein. In

addition to the differently coloured perianth, the ovary

and capsule have a pronounced beak up to 8 mm long,

that readily separates F. schaeferi from F. foliosa. In its

normally sandy habitat, plants spread vegetatively on

long runners produced from the base of the stem that can

extend below the ground a metre or more from the parent

plant, each terminating in a corm.

Although first collected in October 1830 on the

south side of the Orange River by the important early

plant explorer in South Africa, J.F. Drege ( Drege s.n.,

P), Ferraria schaeferi is based on an early 20th cen-

tury collection made about 1912 to 1913 by the medical

doctor, Fritz Schafer, who worked on the Liideritz Bay-

Keetmanshoop railway in the then German colony of

South West Africa (Gunn & Codd 1981), now Namibia.

Schafer made two collections in the Klinghardt Mtns

in the Namib Desert south of Liideritz Bay, and these

specimens formed the basis for Dinter’s description of F.

schaeferi. The broad similarity of the plant to F. crispa

led 19th century botanists to overlook the Drege col-

lection, which lacks flowers, but has the characteristic

beaked capsules that mark F. schaeferi unmistakably.

Representative specimens

NAMIBIA. — 2615 (Liideritz): Kovisberge, (-AA), 8 Sept. 1929,

Dinter 6689 (B); Luderitzbucht, (-AC), Sept. 1917, Knobel s.n.

( SAM12783 ); Kowis Mtns, east of Liideritz, 644 m, (-AA), 15 Aug.

2001, Smook 11394 (PRE); east of Grosse Bucht, wind farm site, (—

CC), 21 Sept. 2001, Mannheimer & Burke 1703 (WIND). 2715

(Bogenfels): south Namib opposite Possession Is., (-AA), 10 Oct.

1976, De Vos 2379 (NBG); Granietberg near Bogenfels, (-AA), Aug.

1911, Schaefer 14 (sub Marloth 5249) (PRE); Buchubergen, (-DD), 9

Aug. 1929, Dinter 6584 (B). 2816 (Oranjemund): rocky outcrop before

Schakel Mtn among quartzite stones, (-BA), 31 Jul. 1977, Muller 752

(WIND).

NORTHERN CAPE. — 2816 (Oranjemund): Beauvallon, sanddune,

(-DA), 22 Aug. 2001, Goldblatt & Porter 11739 (MO); 19 Jul. 2002,

Manning 2746 (NBG). 2817 (Vioolsdrif): stony gravel flats near Arris-

drif, banks of Orange River, (-AC), 31 Aug. 1925, Marloth 12391

(PRE). 2916 (Port Nolloth): 5 km E of Port Nolloth, (-BC), 22 Aug.

2001, Goldblatt & Porter 11734A (MO). 2917 (Springbok): Grootmis,

sanddune, (-CA), Oct. 1977, De Vos 2391 (NBG).

8. Ferraria ovata (Thunb.) Goldblatt &

J.C. Manning in Novon 12: 461 (2002). Moraea ovata

Thunb.: 186 (1800). Type: South Africa, [Northern

Cape,] Namaqualand, near Kooksfontein [as Koksfon-

tein, also Cocksfonteyn], (now Soebatsfontein), without

date, probably in 1793, Masson s.n. UPS-THUNB1225

(UPS-THUNB, holo.l).

Plants often small, (40—) 1 00—200 mm high. Stem

erect, unbranched, lower half partly exposed. Leaves :

basal and juvenile with short, linear blades, up to 45 x

2 mm; cauline entirely bifacial, broadly ovate to rotund,

amplexicaul, 15-18 mm long, spreading, concave; mar-

gins membranous, often reddish. Rhipidia 2-flowered;

spathes leafy, green, often brown at tips, inner 18-20

mm long, outer 12-20 mm long, sheathing in lower half,

arching outward above. Flowers on pedicels 7-12 mm

long, lasting a single day, pale yellow with brown spots

in midline and base of limbs, margins closely crisped,

dark brown, acrid-smelling, tepal claws broad, 7-8

mm long, forming a cup ± 7 mm deep, ± 8 mm wide at

rim, nectaries ± 2 mm long, above base of claws, ± 2 x

2 mm, paired or bilobed, greenish, tepal limbs spread-

ing; outer tepals 17-20 mm long, limbs 8-12 mm long,

inner slightly smaller than outer. Stamens with filaments

united in a column 7-8 mm long, free in upper 1 mm;

anther thecae parallel, ± 2 mm long; pollen orange.

Ovary included, spindle-shaped, ± 6 mm long, without

beak; style branches ± 2 mm long, diverging into promi-

nently fringed arms; stigmas terminal on style arms.

Capsules ovoid (mature capsules and seeds unknown).

Flowering time : Jun.-Jul. Figure 7A-E.

Distribution and biology, found on granitic slopes

and clay-loam flats, Ferraria ovata is restricted to cen-

tral Namaqualand, South Africa, where it has been

recorded from a few isolated sites from Soebatsfontein

in the north to the low hills of the southern Kamiesberg

between Kliprand and Bitterfontein (Figure 8). Plants

typically grow in gritty granitic gravel among rocks that

protect the corms from predation. Like those of other

species of sect. Ferraria, the flowers of F. ovata seem

adapted for pollination by muscid and blowflies.

Bothalia 41,1 (2011)

21

FIGURE 7— A-E, Ferraria ovata : A, whole plant; B, half-flower; C, outer tepal; D, inner tepal; E, anther and style branches. F, Ferraria variabilis:

flower. G, FI, F. densepunctulatcr. G, capsule; H, seed. Scale bar: A, C, D, F, G, 10 mm; B, 5 mm; E, 2 mm; H, 1 .5 mm. Artist: J.C. Manning.

22

Bothalia 41,1 (201 1)

FIGURE 8. — Known distribution of Ferraria densepunctulata, A; F.

ovata, O; F. ornata, •.

Diagnosis and relationships', the parallel anthers,

wide and shallow tepal cup, spindle-shaped ovary with-

out a beak, and stigmas terminal on the style arms, show

that Ferraria ovata belongs in sect. Ferraria. It most

closely resembles the coastal species F. densepunctulata ,

which also has dimorphic basal and cauline leaves, a

sparsely branched or unbranched stem, and the relatively

small bilobed or paired nectaries located well above the

base of the tepal claws. It is one of three winter flower-

ing species in the genus, the others being the coastal spe-

cies, F. densepunctulata and F. ornata.

History, one of the rarest species of the genus, Fer-

raria ovata was first collected in the 1790s, probably

in 1793, by the Royal Botanic Gardens Kew collector,

Francis Masson, and not again for over 200 years. The

species was described as Moraea ovata by Thunberg in

1800, based on Masson’s collection, which was in fruit

and without corms. Plants were rediscovered in 1995 by

botanist, Annelise le Roux, who collected a sterile speci-

men near Soebatsfontein in central Namaqualand. A

later record by P. Desmid and party, also sterile, finally

led us to a site with plants in bloom near Kliprand, east

of Bitterfontein ( Manning 2350), in June 2001, when

it became clear that the plants represented an unknown

species of Ferraria (Goldblatt & Manning 2002).

Representative specimens

NORTHERN CAPE. — 3017 (Hondeklipbaai): Namaqualand, Farm

Doomfontein 464, Steenkamp Kraal, sandy loam slope, (-BA), 2 Sept.

1 995 (sterile), Le Roux 4658 (.IONK).

WESTERN CAPE. — 3018 (Kamiesberg): Farm Gannabos, 25

km from Bitterfontein on road to Kliprand, granite outcrops, (-CD),

10 Aug. 2000 (fr.), Goldblatt & Manning 11373 (MO, NBG), 10 Jun.

2001, Manning 2350 (MO, NBG); Knersvlakte, Farm Steenbokskraal,

(-CD), 20 Aug. 1999 (fr.), National Geographic-IPC Expedition 115

(NBG).

9. Ferraria densepunctulata M.P.de Vos in Jour-

nal of South African Botany 45: 346 (1979). Type: South

Africa, [Western Cape], Elandsbaai [Elands Bay], 14 Jun.

1974, Louhser sub De Vos 2317 ( STE, holo.!; PRE, iso.!).

Plants slender, mostly 120-250 mm high. Stem sev-

eral-branched in upper half, lower half of stem partly

visible; leaf sheaths and cataphylls often flushed red

with pale spotting. Leaves : basal with long compressed-

cylindrical blades, loosely 2-ranked, 3-5 mm wide,

with narrow hyaline to reddish margins; cauline leaves

shorter, spreading outward, tips often hooked; juvenile

leaves in a 2-ranked cluster, up to 6, terete, hollow, dark

green. Rhipidia 2-flowered; spathes green with nar-

row membranous margins either pale or flushed red;

inner 40-50 mm long, outer ± 25 mm long, sheathing

below, diverging and hooked distally. Flowers on pedi-

cels slightly shorter to ± as long as spathes, lasting two

days, pale greenish to grey-blue, outer tepals densely

speckled with small maroon to purple or dark blue spots,

inner tepal limbs maroon to purple or dark blue blotch

in lower V2, faintly spice-scented, tepal claws forming

a shallow cup, ± 9 mm deep, 15-20 mm wide at rim;

nectaries bilobed, above base of outer tepal claws, in

midline of inner tepal claws, tepal limbs spreading hori-

zontally, undulate or only slightly crisped; outer tepals

25-30 x 11-13 mm, inner tepals 20-23 x ± 10 mm,

claws of both whorls ± 10 mm long, inner much nar-

rower than outer. Stamens with filaments united in a col-

umn 8-10 mm long, free in upper ± 2 mm; anther thecae

parallel, ± 4 mm long before anthesis; pollen orange or

dull yellow. Ovary sometimes exserted, spindle-shaped,

12-15 mm long, without obvious beak but without

ovules in upper 3—4 mm; style ± 10 mm long, style

branches ± 3 mm long, divided in upper 7 into diverg-

ing, fringed arms; stigmas terminal on style arms. Cap-

sules ovoid, 15-25 mm long, apex subacute. Seeds pale

glossy brown, ± 3 mm diam., 5-sided with facets slightly

wrinkled. Flowering time : May-Jul. Figure 7G, H.

Distribution and biology. Ferraria densepunctu-

lata grows in rocky or calcareous sandy sites along the

Western Cape coast of South Africa, from Lambert’s

Bay in the north to Langebaan in the south (Figure 8).

At Jacobsbaai near Saldanha, plants grow in limestone

pavement. The spice-scented flowers are pollinated by a

range of short-tongued flies.

Diagnosis and relationships', early flowering and rela-

tively rare, Ferraria densepunctulata was only described

in 1979 by M.R de Vos, and until she began her study

of the genus, the species was known from just three col-

lections, the first made by C.L. Leipoldt in June 1941 in

the hills at Langebaan, south of Saldanha, still the most

southerly record. The species is distinctive in the narrow

basal leaves, oval in cross section, and rather different

from the shorter, broader cauline leaves, and in the unu-

sually coloured flowers, minutely dark blue to purple

spotted on a pale greenish to grey-blue background, but

with the limbs of the inner tepals so densely speckled as

to appear solidly coloured, either maroon or dark slate

blue. The flowers are also unusual in the genus in hav-

ing the tepal limb margins undulate instead of closely

crisped. The small bilobed nectaries are located at the

base of the outer tepals but in the midline of the inner

tepal claws, the lower halves of which are linear and ±

1 mm wide. Like other members of sect. Ferraria , the

stigmas are terminal on the arms of the style branches

and the ovary is not beaked as in all but F. schaeferi of

the section. The flowers are unique in the section in last-

Bothalia 41,1 (201 1 )

23

FIGURE 9. — Ferraria omata, Manning 3174. A, flowering plant; B, fruiting plant; C, outline side view of flower; D, claw of outer tepal;

E, claw of inner tepal; F, staminal columns and style; G, capsule. Scale bar: A— C, G: 10 mm; D, E, 0.5 mm; F, 1.5 mm. Artist: J.C.

Manning.

24

Bothalia 41,1 (201 1 )

ing two days instead of a single day. The leaves of non-

flowering plants are so strikingly different from those of

mature, flowering indivuals that they appear to represent

a different species. Four to six in number, they are held

in a close, 2-ranked fan and the blades are dark green,

terete and hollow.

The early flowering habit of Ferraria densepunc-

tulata is now known to be matched in the genus by F.

ornata and F. ovata, both of which also have the narrow

basal leaves differentiated from the shorter and broader

cauline leaves.

Representative specimens

WESTERN CAPE. — 3217 (Vredenburg): Jacobsbaai, in limestone

pavement, (-DD), Aug. 2007 (fr. ), Goldblatt & Manning 129I4A

(MO); Cape Columbine, hill adjacent to lighthouse, in sand with lime-

stone gravel, (-DD), 8 Sept. 2009, Goldblatt & Porter 13285 (MO,

NBG). 3218 (Clanwilliam): Langdam, near Lambert's Bay, (-AB),

Jun. 1974, De Vos 2298 (NBG); Elands Bay, near school, (-AB),

17 May 1966, Pamphlet t 98 (NBG); Farm St Helenafontein, sandy

ground, (-DB), 11 Sept. 2008 (fr.), Goldblatt & Porter 13111 (MO,

NBG). 3318 (Cape Town): Langebaan, hills behind hotel, (-AA), Jun.

1941, Leipoldt 3844 (BOL).

10. Ferraria ornata Goldblatt & J.C. Manning, sp.

nov.

Plantae acaulescentes usitate 50-80 mm altae, foliis

4 vel 5 amplexicaulibus concavo-lanceolatis ad 40 x 15

mm, lamina folii basalis 5-10 mm longa subtereti ad

cylindrica, spathis coriaceis glaucis marginibus translu-

centibus spatha interna 40-50 mm longa externa 20-30

mm longa extrinsecus arcuata, floribus albis tepalis basi

pall ide flavis, limbis unguibusque brunneis maculatis

inodoris, marginibus undulatis crispisque, nectaribus

bilobis in medio unguium marginibus distalibus cristis

prominentibus, tepalis inaequalibus externis ± 19 x 10

mm unguibus ± 8 mm longis ± 2 mm latis ad basem,

internis ± 13 x 6 mm unguibus ± 6.0 longis ± 1.5 mm

latis ad basem, columna filamentorum ± 8-9 mm longa,

antheris ± 3 mm longis brunneis lobis paralleli, ovario

fusiformi incluso 10-12 mm longo, ramis styli fimbria-

tis.

TYPE. — Northern Cape, 3017 (Hondeklipbaai):

central Namaqualand, sandy slopes 10 km inland from

Groen River mouth. Farm Van Zylsrus, 500 m along dirt

track N of main road, strandveld in deep sand, (-DC), 2

Jun. 2008, Manning 3174 (NBG, holo.; MO, iso.).

Plants acaulescent, congested, 50-80 mm high. Stem

2^1-branched; replacement conns not present at flow-

ering, developing later; cataphylls (usually only upper-

most) turning red above ground. Leaves 4 or 5, amplexi-

caul, lanceolate-concave, up to 40 x 15 mm, glaucous

with purple tips; margins reddish membranous, finely

crisped, basal leaf with blades only 5-10 mm long,

subterete to cylindrical, obtuse-apiculate, 3-4 mm

diam.; non-flowering plants with l-3(-5) leaves; blades

suberect or falcate, subterete or cylindrical, ± quadran-

gular to round in section, 60-80 x 2-4 mm, obtuse-

apiculate, glaucous with purple apex and sheath, sheath

margins and veins abaxially minutely hairy. Rhipidia

2-flowered; spathes leathery, glaucous, with translu-

cent, membranous margins, inner 40-50 mm long, outer

much smaller, 20-30 mm long, sheathing in lower half,

arching outward. Flowers on pedicels 30-40 mm long.

lasting a single day, white with base of tepal limbs yel-

low, limbs finely brown-spotted basally and with fewer,

larger brown spots distally, margins finely crisped, yel-

low and brown, claws streaked with brown, unscented,

with bilobed, concave nectaries in middle of claws,

distal nectary margin prominently crested, pale lilac

with purple streaks, tepal claws narrowed below, form-

ing a windowed cup 7-8 mm deep and ± 10 mm wide

at rim, limbs spreading or slightly reflexed; tepals un-

equal, outer ± 19 x 10 nun with claws ascending, ± 8

mm long, narrowed below and ± 2 mm wide, inner ±

13x6 mm with claws ± 6 mm long, narrow proximal

part ± 1.5 mm wide. Stamens with filaments united in

column 8-9 mm long, free in upper ± 2 mm; anthers

diverging, thecae parallel, ± 3 mm long, brown; pol-

len orange. Ovary included, spindle-shaped, 10-12

mm long; style branches diverging, deeply divided into

diverging, fringed arms; stigmas terminal on style arms.

Capsules broadly ovoid, 14-18 mm long, tapering to

acute tip. Seeds irregularly 5(6)-sided, ± 3 x 2 mm, fac-

ets separated by pale straw-coloured, wavy ridges, facet

surfaces usually ± plane and dark brown. Flowering

time : May to early Jun. Figure 9, Plate 1H.

Distribution and biology. Ferraria ornata is known

from two extended populations, one on slopes above the

lower Groen River in central Namaqualand and the other

west of Koekenaap in southern Namaqualand, some 120

km to the south. It occurs on sandy ground in sandveld

(Figure 8). At its northern site on the northern bank of

the Groen River, ± 1 0 km inland from the Atlantic coast,

plants occur in localized colonies in deep, gritty, sandy

soil in Namaqualand Strandveld vegetation among suc-

culent shrubs, including Othonna cylindrica and O.

coronopifo/ia. The plants appear to be habitat specific

and were not located in patches of Namaqualand Coastal

Fynbos that interdigitate with the strandveld. Plants

flower early in the season, after the first rains, in May

or June, and last a single day, opening ± 08:00 and col-

lapsing ± 16:00. Branches are produced in the upper one

or two leaf axils, which extend the flowering period for

some weeks, possibly into early July in years of favour-

able rainfall. Juvenile and non-flowering plants produce

a tuft of cylindrical leaves quite different from those on

flowering individuals. Replacement corms of flowering

individuals are absent at flowering and develop later as

the capsules ripen.

The species was discovered by Cape Town ecologist,

Rupert Koopman, who collected a fruiting specimen

in spring 2007. The locality was visited in the follow-

ing winter in a year of poor rainfall but we were able to

locate a single flowering individual for description and

illustration.

Diagnosis and relationships', stemless, and with the

tepal claws unusually narrow in the lower part, thus

forming a windowed cup, Ferraria ornata is easily rec-

ognized by its small size, flowering individuals stand-

ing only 50-80 mm high. The flowers are unique in the

fleshy crests bordering the distal margins of the nectar-

ies, which are situated in the middle of the tepal claws,

and which secrete miniscule quantities of sticky, sweet

nectar. The flowers appear scentless to the human nose.

The short, concave cauline leaves of flowering plants

stand in marked contrast to the long, centric leaves of

Bothalia 41,1 (201 1 )

25

non-flowering individuals. Both the early flowering

and unusual, concave cauline leaves of flowering plants

suggest it is most closely allied to F. ovata , which has

a similar growth strategy of early flowering and small,

concave cauline leaves on a taller flowering stem.

Additional specimens

NORTHERN CAPE. — 3017 (Hondeklipbaai): same locality as

type collection, Aug. 2007 (fr. and sterile), Koopman s.n. (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Farm Kommandokraal

west of Koekenaap, sandveld, ± 80 m, (-CA), 9 Sept. 2008 (fr.), Gold-

blatt & Porter 13098 (K, MO, NBG, PRE).

B2. Sect. Macroscyphae Baker in Journal of the

Linnean Society 16: 106 (1877); De Vos: 349 (1979).

Type species: Ferraria divaricata Sweet (lectotype des-

ignated by De Vos 1979: 350).

Flowers with tepal claws forming a deep, wide or

narrow cup containing fluid nectar; filaments inserted

near apex of anthers; anther lobes when fully dehisced

usually widely divergent, rarely parallel but never con-

tiguous. Ovary always with sterile beak; stigmas on

lobes below apices of style branch arms. Capsules with

prominent beak.

Series Subdivaricatae

11. Ferraria ferrariola (Jacq.) Willd., Species

plantarum edn 4, 3: 581 (1800); De Vos: 350 (1979).

Moraea ferrariola Jacq.: (1790). Type: South Africa,

without precise locality or collector, illustration in

Plantarum rariorum horti caesarei schoenbrunnensis 4:

24, t. 450(1804).

F. viridiflora Andrews: 285 (1803) \F. viridis Ker Gawl. (1804)

orth. var.]. Type: South Africa, collected by W. Synnot, without precise

locality, illustration in The botanist’s repository 4: t. 285 (1803); no pre-

served material known.

F. tonga Barnes: 313 (1930). Type: South Africa, [Northern Cape],

Nieuwoudtville, cultivated at Kirstenbosch, Jul. 1930, P. Ross-Frames

s.n. BOL19720 (BOL, holo.!).

Plants slender, mostly 150-250 mm high. Stem

sheathed below by mottled cataphylls and leaf sheaths;

lower intemodes usually partly exposed, usually branched

in upper V . Leaves', basal linear, unifacial, usually slightly

longer than stem, mostly 2-3 mm wide, main vein usu-

ally well delineated; cauline leaves markedly shorter and

broader than basal leaves and resembling spathes. Rhi-

pidia 2-flowered; spathes glaucous or pale green, with

broad transparent margins, inner 45-60(-70) long, often

inflated, outer ± '/ to 2/3 as long, sheathing below, diverg-

ing in upper half and ± hooked at tips. Flowers on pedi-

cels 20-30 mm long, lasting two days, greenish blue,

grey-blue, or pale watery yellow to pale green, usually

with darker spots and streaks on outer tepal limbs, faintly

sweet- or spice-scented, sometimes odourless, tepal claws

forming a narrow, closed cup, 17-20 mm deep, ± 12 mm

wide at mouth; nectaries small, at base of tepals; outer

tepals (30— >35^40 mm long, claws fairly narrow, 17-20

mm long, inner slightly shorter and narrower, limbs of

outer ± spreading, inner usually reflexed. Stamens with fil-

aments united in a column 14—19 mm long, free in upper

1-2 mm; anther thecae initially parallel, ± 4 mm long at

anthesis, later diverging and shrinking to almost 2 mm.

Ovary spindle-shaped, 20-25 mm long, with sterile beak.

(5— )8— 12 mm long; style branches ± 2.5 mm long, divid-

ing into diverging, fringed arms, ± 2.5 mm long; stigmas

irregularly shaped lobes in middle of style arms, arching

over anthers. Capsules ovoid-oblong, 1 4 — 20(— 25) mm

long, beak (5 — )8— 1 2 mm long. Seeds mostly 5-sided, 3.0-

3.5 x 2. 5-3.0 mm, smooth, shiny, brown, facets separated

by prominent, pale wavy ridges, facets light yellow-brown

with surfaces smooth or slightly wrinkled. Flowering time :

Jun.-Aug. Plate IF.

Distribution and biolog\r. usually in rocky sites, the

relatively common Ferraria ferrariola is found on gran-

ite and sandstone slopes, or in sandy ground usually not

far from rock outcrops. Its range extends from the Rich-

tersveld in northern Namaqualand to the Bokkeveld and

Olifants River Mtns as far south as Clanwilliam (Figure

10). The floral cup contains nectar of moderately high

sugar concentration. The scented flowers, which last two

days, appear attractive to bees, the only recorded floral

visitors (Goldblatt et al. 2009). These include honey

bees. Apis mellifera, and Anthophora species, which

visit the flowers to forage for nectar, and during their

visits accomplish pollination.

Diagnosis and relationships'. Ferraria ferrariola is

readily recognized by the linear basal leaves, quite dif-

ferent from the shorter cauline leaves, the stem usually

partly exposed, and the red-flushed and white-speckled

cataphylls and sheath of the lowermost leaf. The large

flowers have a deep and relatively narrow floral cup,

17-20 deep and ± 12 mm wide at the mouth. The flow-

ers are usually shades of turquoise-grey to pale slate-

blue with a pale yellow nectar guide lightly speckled

with darker colour on the limbs of the outer tepals (Man-

ning et al. 2002: 157). The flowers have a faint sweet

odour, often with spicy overtones of almond or cinna-

mon or vanilla, unique in the genus.

The relationships of Ferraria ferrariola are evidently

with sect. Macroscyphae with which it shares the diver-

gent anther lobes and large stigma lobes below the tips

of the style arms, as well as a prominently beaked ovary.

16° 18° 20° 22° 24°

l

FIGURE 10. — Known distribution of Ferraria ferrariola, O; F. parva,

26

Bothalia 41,1 (2011)

The sharp dimorphism between the basal and cauline

leaves does not occur in other species of the section, but

is found in F densepunctulata and F. ovata of sect. Fer-

raries and in F. glutinosa of subgen. Glutinosae. These

species are, however, otherwise very different in having

flowers with the tepal claws forming a wide, shallow

cup, anthers with parallel lobes, small stigmatic surfaces

terminal on the style arms, and an ovary lacking a beak.

In our Western Cape Wildflower Guide (Manning &

Goldblatt 1996: 51) we misdentified as Ferraria fer-

rariola the photograph of a second species, F. parva,

described here as new. Whereas the two have flow-

ers of similar shape, divaricate anthers, and a narrow

floral cup, those of F. pan>a are much smaller, with

a floral cup, ± 6 mm deep, and outer tepals 18-22 mm

long, compared to (30— )35^40 mm in F. ferrariola. The

sheaths and cataphylls of F. parva also lack the red flush

and white speckling so characteristic of F. ferrariola.

Representative specimens

NORTHERN CAPE. — 2817 (Vioolsdrif): Richtersveld, Kodaspiek,

stony clay loam, (-AA), 2 Sept. 1977, Oliver, Toelken & Venter 395

(PRE); Armanshoek, moist gorge, (-AC), Aug. 1995, Williamson

& Williamson 5647 (NBG); stony flats ± 4 km N of Eksteenfontein,

(-CD), 23 Aug. 2001 (fr.), Goldblatt & Porter 11755 (MO). 2917

(Springbok): granite dome W of Springbok, (-DB), 21 Aug. 2002,

Goldblatt & Porter 12085 (MO). 3017 (Hondeklipbaai): Kamieskroon,

(-BB), 24 Jul. 1941, Compton 11110 (NBG); flats east of Karkams on

Tweerivieren road, (-BD), 21 Aug. 2001, Goldblatt & Porter 11723

(MO). 3119 (Calvinia): Nieuwoudtville Wildflower Reserve, dolerite

rocks, (-AC), 11 Aug. 1983, Perry & Snijman 2276 (NBG); Botter-

kloof Pass, (-CA), 12 Aug. 1983, Van Wvk 1453 (NBG); top of Botter-

kloof, (-CC), 28 Jul. 1948, Lewis 1992 (SAM).

WESTERN CAPE.— 3018 (Kamiesberg): ± 9 km N of Bitter-

fontein, (-CC), 20 Aug. 2001, Goldblatt & Porter 11713A (MO).

3118 (Vanrhynsdorp): Holrivier, stony hills, (-AD), 11 Aug. 1970,

Hall 3730 (NBG); Farm KJiphoek, 10 miles [16 km] east of Doring-

baai, (-CC), 12 Aug. 1970, Hall 3743 (NBG). 3218 (Clanwilliam):

Pakhuis Pass, west end, (-BB), 23 Aug. 1966, Barker 10454 (NBG);

Clanwilliam. 350 ft [106 m], (-BB), 22 Aug. 1896, Schlechter 8597

(PRE). 3219 (Wuppertal): between Pakhuis and Botterkloof, 2 km past

Bidouw turn-off, (-AA), 4 Sept. 1976 ( t'r.), De Vos 2369 (NBG).

12. Ferraria parva Goldblatt & J.C. Manning, sp.

nov.

Plantae usitate 80-150 mm altae, foliis unifacialibus,

basalibus 2-3 linearibus 3-5 mm latis, caulinibus lan-

ceolatis 4-6 mm latis, spathis pallidis viridibus interne

45-65 mm longis, externe ± dimidio longiore interne,

floribus atroflavis ad pallide caeruleo-viridibus atro-

maculatis, marginibus crispis bubalinis, tepalis exteri-

oribus 18-22 mm longis, columna filamentorum ± 6 mm

longis, antheris ± 3 mm longis lobis divaricatis.

TYPE. — Western Cape, 3217 ( Vredenburg): sandveld

west of Vredenburg, north side of Saldanha highway,

opposite cemetery, under shrubs and restios, (-DD), 19

Sept. 2007, Goldblatt & Manning 13000 (NBG, holo.;

MO, iso.).

Plants mostly 80-150 mm high, not including basal

leaves. Stem branched near base and in upper part, lower

'/, often partly exposed; cataphylls and sheath of low-

ermost leaf pale. Leaves unifacial, basal 2 or 3 ± linear,

usually exceeding spathes, 3-5 mm wide, upper leaves

subtending branches lanceolate, spreading, shorter and

wider than basal, mostly 4-6 mm wide, margins and

main vein not evident when alive; non-flowering plants

with 2 or 3 linear leaves, oval in cross section, without

differentiated margins or main vein. Rhipidia 2-flow-

ered; spathes pale green, apices slightly hooked, inner

45-65 mm long, often inflated with age, margins trans-

parent, outer spathe ± 7 as long as inner, often diverg-

ing in upper 7V Flowers on pedicels 18-20 mm long,

lasting 2 days, spotted with dark brown in a dull yellow,

light red-brown or pale greenish blue background, mar-

gins crisped, buff to khaki, sweetly scented of a mix of

jasmine and coconut, tepal claws forming a narrow cup

± 8 mm deep, ± 7 mm wide at rim; outer tepals 1 8-22 x

8-10 mm, claws 7-8 mm long with a pouch-like bilobed

nectary in lower midline; inner tepal slightly shorter and

± V as wide, limbs slightly reflexed, inner more so than

outer. Stamens with filaments united in a column ± 6

mm long, free in upper ± 1 .5 mm; anther thecae initially

parallel, later divaricate, ± 3 mm long at anthesis, later ±

2/3 as long. Ovary fusiform, 12-15 mm long, with a ster-

ile beak up to 7 mm long; style branches ± 2 mm long,

forked in upper V2, prominently fringed; stigmas nar-

row, at lateral ends of style branches, arching forward.

Capsules ovoid-oblong, 15-20 mm long, with beak up

to 2-7 mm long. Seeds angular, smooth, shiny, golden

brown, mostly 5-sided, facets smooth or slightly wrin-

kled. Flowering time : late Aug.-mid-Sept. Figure 11,

Plate II.

Distribution and biology. Ferraria parva is a sur-

prising discovery — plants grow close to Saldanha and

Vredenburg, an area believed to be well-explored botan-

ically. Clearly rare, the species is restricted to sand-

veld and limestone fynbos on the Western Cape coast

between Bokbaai and Vredenburg and the Berg River

near Langrietvlei (Figure 10). Plants grow in deep sandy

ground, sometimes over limestone, or in cracks in lime-

stone or calcrete pavement. The flowers last two days

and have a sweet scent reminiscent of a mix of jasmine

and coconut or, less often, a somewhat sour, unpleasant

odour. The floral cup contains moderately sweet nectar

of mean 24.2 % sucrose equivalents. Pollination biology

is unknown but, like its closest relative, F. ferrariola, the

flowers appear adapted for pollination by bees foraging

for nectar. At the type site we noticed plants had been

well pollinated. Unfortunately, many plants in the small

population had been uprooted and the corms eaten, pre-

sumably by porcupines. Apart from the type site, inci-

dentally, the only known locality for Romulea el/iptica

M.P.de Vos, just one other record of the species was

known before 2007, made in 1940 by W.F. Barker ‘on

the road to GanzekraaT (probably the farm southwest of

Darling near Bokbaai). Collecting in the Saldanha area

in 2008 showed the species to be fairly common in the

immediate area of Vredenburg and Saldanha, both on

limestone and in sandy ground. The single flower of the

Ganzekraal collection is poorly preserved but is compa-

rable in size and has the divaricate anthers and general

appearance of F. parva.

Diagnosis and relationships', we first encountered

Ferraria parva in 1995 while taking photographs for

the West Coast Wildflower Guide (Manning & Gold-

blatt 1996) and included therein a photograph of

the species misidentified as F. ferrariola. Later, dur-

ing our study of Ferraria, it became clear that this

Bothalia 41,1 (201 1 )

27

FIGURE 1 1 . — Ferraria parva,

Goldblatt & Manning 13000.

A, flowering plant; B, half

flower; C, outer tepal; D,

inner tepal; E, anther and sin-

gle style branch; F, capsules,

undehisced (left) and dehisced

(right); G, seed. Scale bar: A,

F, 10 mm; B-D, 5 mm; E, G, 2

mm. Artist: J.C. Manning.

was not F. ferrariola, which is a Namaqualand plant

that extends south only as far as Clanwilliam, but

an unknown species. After repeated searches at the

same site, we re-collected the species in Septem-

ber 2007. The flower is similar to F. ferrariola in

basic structure: it has a narrow floral cup and tepals

with slightly reflexed limbs but the flower is less than

half the size of those of F. ferrariola. the tepals just

18-22 mm long, the floral cup ± 8 mm deep and fila-

ment column ± 6 mm long, compared with outer tepals

(30-)35-40 mm long, a floral cup 17-20 mm deep and

a filament column 14-19 mm long of F. ferrariola. The

cataphylls and sheath of the basal leaf of F. parva are

uniformly coloured in contrast to the speckled and usu-

ally red-flushed cataphylls and sheaths of F. ferrariola.

Representative specimens

WESTERN CAPE. — 3217 (Vredenburg): sandveld west of Vreden-

burg, (-DD), 29 Aug. 1995, Goldblatt & Manning 10271 (MO); lime-

stone-topped hill north of Saldanha, (-DD), 13 Sept. 2008, Goldblatt

& Porter 13117 (MO, NBG, PRE). 3218 (Clanwilliam): Eriocephalus-

dominated sandveld on Farm Brakfontein (Nuweland) south of Langriet-

vlei, (-CC), 6 Sept. 2008, Goldblatt & Porter 13077 (K, MO, NBG,

PRE, S); sandveld near Langebaanweg Fossil Park, (-CC), Goldblatt

& Porter 13150 (MO, NBG). 3318 (Cape Town): road to Ganzekraal,

(-?DB), 15 Sept. 1940, Barker 730 (NBG).

Series Macroscyphae

13. Ferraria divaricata Sweet in British flower

garden: t. 192 (1827); De Vos: 354 (1979). Type: South

Africa, without precise locality, collected by W. Synnot,

illustration in British flower garden: t. 192 (1827) (no

preserved material known).

28

Bothalia 41.1 (2011)

F. divaricata subsp. arenosa M.P.de Vos: 358 (1979). Type: South

Africa, [Western Cape], Clanwilliam, Nardouw Pass, Farm de Lille,

Oct. 1 973, Van Breda sub De Vos 2295 (NBG, holo.!; PRE, iso.!).

Plants (250-)350-500 mm high. Stem well devel-

oped, much-branched distally. Leaves sword-shaped

to linear, 6-15 mm wide, often with a slightly raised

main vein, glaucous, margins sometimes slightly

raised, basal leaves longest, cauline leaves progres-

sively shorter above and becoming spathe-like. Rlti-

pidia 2-flowered; spathes glaucous, with broad mem-

branous margins, inner 70-85 mm long, outer ± half as

long as inner, sheathing entirely or in lower two thirds,

then often slightly S-shaped. Flowers on pedicels 6-15

mm long, lasting one day, variously dark chocolate to

golden brown, crisped part of margins concolourous or

paler, claws pale, streaked with fine longitudinal lines

or with a broad median darker centre, forming a cup,

14- 18 mm deep, 15-18 mm wide at rim; outer tepals

(30— )35^44 x 14-16 mm, inner tepals (28-)36-38 x

8-10 mm, claws of both whorls ( 1 3— )1 6—1 9 mm long;

nectaries at base of claws, ± black or pale yellow-green,

1 .3 — 4.0 mm long. Stamens with filaments united in a

column 11-13 mm long, free and arching outward in

upper ± 3 mm; anther thecae usually widely diverging,

initially ± 4 mm long, shrinking after dehiscence; pollen

orange. Ovary fusiform, 33—42 mm long including beak

15- 20 mm long; style branches ± 2 mm long, dividing

into diverging, prominently fringed arms ± 4 mm long;

stigmas on small lobes below tips of style arms, arching

over anthers. Capsules 1 5-22 mm long (excluding beak

15-20 mm long). Seeds globose, ± 3 mm diam., pale

matte brown, surfaces lightly wrinkled in ± ruminate

pattern, epidermal cells foveate. Flowering time : mainly

late Sept, to early Nov.

Distribution and biology'- Ferraria divaricata extends

along the west coast and coastal mountains of North-

ern Cape and Western Cape, South Africa, from near

Komaggas in northern Namaqualand southward to the

Cape Flats (Figure 12). Plants grow in deep sandy soils,

in sandveld or marginal fynbos habitats. The flowers,

FIGURE 12. — Known distribution of Ferraria divaricata , O; F. f/ava.

with a relatively wide, deep floral cup, contain quantities

of dilute nectar, seldom more than 8 % sucrose equiva-

lents. This dilute nectar is remarkable as it is an unlikely

reward for any potential pollinator. Like its relative, F.

variabilis , which has a similar floral cup and dilute nec-

tar, the flowers are visited only by vespid and masarine

wasps (Eumenidae) (Goldblatt & Manning 2006).

Diagnosis and history, the name Ferraria divaricata

has been generally applied to the low-growing, usually

± tufted plant, mostly 100-200 mm tall, now F. vari-

abilis, from southwestern Namibia, the Upper Karoo,

Namaqualand and Western Cape. This species also has

divaricate anthers and pale brown to yellowish tepals

with the limbs either speckled or marked with a dark

brown to blackish purple band of solid colour at the base

and the claws forming a deep floral cup. De Vos (1979)

associated the name F. divaricata with the northerly

populations of F. variabilis , which she divided into four

subspecies.

Examination of the type, a watercolour illustra-

tion in The British flower garden led us (Goldblatt &

Manning 2004) to question De Vos’s interpretation

(Goldblatt & Manning 2005) of subsp. divaricata. The

height, said to be 18 inches (450 mm), the longer basal

leaves with a slightly raised central vein and appear-

ing slightly striated when dry as the leaf veins become

raised above the surrounding leaf tissue and differenti-

ated from the cauline leaves, plus the exposed stem and

uniformly chocolate-brown tepal limbs, seem to us to

accord best with the plant De Vos called F. divaricata

subsp. arenosa. This attribution also seems reasonable

on circumstantial grounds, for plants depicted in the type

illustration were collected (presumably as seed) by Wal-

ter Synnot, magistrate ( landdrost ) at Clanwilliam from

1821 to 1825. Synnot collected plants in the Clanwil-

liam District and nearby (Gunn & Codd 1981), where

subsp. arenosa is found but there is no evidence that he

collected in Namaqualand where subsp. divaricata sensu

De Vos occurs. This generally tufted plant has flowers

with a solid, dark band of colour on the lower half of

the tepal limbs. De Vos’s two tufted subspecies, subsp.

divaricata and subsp. australis , seem best treated as

one species, F. variabilis. The fourth subspecies, subsp.

aurea , is here asociated with the new F. flava.

The vegetative differences outlined above and the

associated different patterns of tepal marking are sub-

stantial and are correlated with differences in the seeds.

Whereas most Ferraria species have brown seeds, com-

pressed by pressure into 5 or 6 smooth, flattened facets

separated by thickened ridges, the seeds of plants called

subsp. arenosa and subsp. aurea are globose with reticu-

late/foveate sculpturing (De Vos 1979). Seeds rarely

vary within a species so that differences in seed mor-

phology usually have strong taxonomic significance.

Another difference between the last two subspecies is

that their flowers last a single day but two days in subsp.

divaricata and subsp. australis.

As here understood, Ferraria divaricata is a plant of

deep sands or stony sandstone slopes, extending from

near Komaggas in northern Namaqualand to the Cape

Flats close to Cape Town. Populations extend inland as

far as Vanrhynsdorp and the northeastern slopes of the

Bothalia 41,1 (2011)

29

Piketberg (60-70 km inland). As in most Ferraria spe-

cies. floral pigmentation patterns and colour are variable.

The tepal limbs are uniform in colour, except for the

margins, and range from light to chocolate-brown with

paler margins, or are dull yellow-brown, then with little

or no colour difference at the margins. The tepal claws

are cream-coloured to palest yellow with fine, dark, lon-

gitudinal streaks and usually a dark median stripe.

The Cape Town botanist, H.M.L. Bolus, annotated

collections of Ferraria divaricata from Langebaan,

north of Cape Town, F. langebaanensis, but did not pub-

lish the name (De Vos 1979).

Representative specimens

NORTHERN CAPE. — 3017 (Hondeklipbaai): 1.2 km east of

Oubees/Wildepaardehoek boundary, 4.7 km E of Springbok-Soebats-

fontein road. (—BA), 15 Oct. 1986, Le Roux & Lloyd 664 (NBG); 38

km from Garies to Wallekraal, (-BC), 4 Sept. 1977, Drijfltout sub De

Vos 2399 (NBG); without precise locality: between Komaggas and

Soebatsfontein, 9 Sept. 1950, Barker 6740 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): 16 km north of Strand-

fontein, front of sand dune, (-CA), 12 Sept. 1975, De Vos 2356 (NBG);

west of Koekenaap, Farm Kommandokraal, (-CA), Sept. 2008, Gold-

blatt & Porter 13188 (NBG); Ratelfontein, Olifants River Mtns, (-DC),

6 Oct. 2004, Goldblatt & Porter 12630 (MO). 3218 (Clanwilliam):

northeastern slopes of the Piketberg, ± 28 km north of town, (-DA),

28 Sept. 2001, Goldblatt & Manning 11931 (MO, NBG). 3318 (Cape

Town): Farm Osfontein east of Saldanha and north of Langebaan, cal-

careous sand and limestone, (-AA), Nov. 1976, Boucher s.n. (PRE);

Silwerstroomstrand, (-CA), 20 Nov. 1974, De Vos 2339 (NBG); Bell-

ville, University of Western Cape campus, (-DC), 10 Nov. 1976, De

Vos 2385 (NBG).

14. Ferraria variabilis Goldblatt & J.C. Manning

in Bothalia 35; 73 (2005). Type: South Africa, [Northern

Cape,] Nieuwoudtville, Klipkoppies, lower slopes, 15

Sept. 1961, Barker 9537 (NBG. holo.!; MO, iso.!).

F. antherosa Ker Gawl.: t. 751 (1804), nom. illegit. superfl. pro F.

viridiflora [as F. viridis] Andrews (= F. ferrariola (Jacq.) Willd.). Type:

South Africa, without precise locality or collector, illustration in Cur-

tis’s Botanical Magazine 19: t. 751 (1804).

F. divaricata subsp. australis M.P.de Vos: 359 (1979). Type: South

Africa, [Western Cape,] Karoo Garden, Whitehill, 17 Sept. 1945,

Compton 17412 (NBG, holo.!).

Plants 60-200(-300) mm high. Stem often branched

just above base, branches crowded and ± equal in length,

often forming small tufts. Leaves sword-shaped to lin-

ear, usually ± as long as stem, sometimes up to twice as

long, (2— )3— 1 5(— 22) mm wide, mostly sub-basal, with-

out visible midrib, often slightly striate, margins often

slightly thickened, rarely obscurely crisped, sheaths usu-

ally overlapping and concealing stem. Rhipidia 2-flow-

ered; spathes glaucous green, inner mostly 60-75 mm

long, outer slightly shorter to ± as V as long, entirely

sheathing or arching outward in distal third. Flowers

on pedicels 10-15 mm long, lasting 2 (rarely 3) days,

predominantly pale to dull yellowish to pale or middle

brown, or dull grey-blue, limbs with solid dark brown

to blackish purple at base, or with scattered dark spots,

margins darker or paler in colour, claws uniformly pale

or with dark longitudinal streaks or with a broad darker

median streak, forming a floral cup 12-15 mm deep,

13-15 mm wide at rim, usually slightly putrid smell-

ing, nectaries usually basal, rarely in pouches in centre

of claws, pale or dark-coloured; outer tepals 3CM10(-

45) x 10-15 mm, claws 10-15 mm long; inner tepals

25 — 40( — 45 ) x 8-10 mm. Stamens with filaments united

in a column 8-13 mm long, free and arching outward

in the upper 2^1 mm; anthers 3. 5-5.0 mm long before

anthesis, shorter after dehiscence, thecae joined only at

apices, usually widely diverging but sometimes almost

parallel. Ovary > fusiform, 15-20 mm long, with a sterile

beak 5-8 mm long; style branches 2-3 mm long, divid-

ing into diverging, prominently fringed arms, ± 4 mm

long; stigmas on small lobes below tips of style arms

and arching over anthers. Capsules ellipsoid, 30-50 mm

long. Seeds rounded, usually angled by pressure, coat

dull and slightly wrinkled. Flowering time ; Aug.-Nov.

Figure 7F, Plate 1 D.

Distribution and biology, most widespread of the

southern African species, Ferraria variabilis extends

from southern Namibia to Oudtshoorn including Bush-

manland and the Great Karoo and as far east as Uping-

ton and Britstown, but is absent from the western coastal

forelands of Western Cape Province (Figure 13). Plants

grow in a variety of habitats including shale flats, gran-

ite outcrops, and deep sands. As described in more detail

above and elsewhere (Goldblatt & Manning 2006; Gold-

blatt et al. 2009), the flowers are adapted for pollination

by mud wasps (Vespidae): the only pollinating insects

recorded on F. variabilis are species of Allepipona and

Delta (Eumeninae), and Jugurtia (Masarinae). A previ-

ous report of pollination of F variablis by the masarine

wasp, Ceramius (Goldblatt et al. 2009), is incorrect: that

record is for F. macrochlamys subsp. kamiesbergensis .

Diagnosis and relationships : Ferraria variabilis

is morphologically fairly coherent in including plants

with relatively few branches crowded near the base, and

rarely higher than 200 mm. Plants thus form low tufts

unless growing under shrubs when they may be taller.

The flowers are fairly large and the broad tepal claws

form a deep, relatively wide bowl in which nectar of low

sugar concentration accumulates in a pool. The spread-

ing tepal limbs vary considerably in colour and marking.

In the north of its range, in Namaqualand and southern

Namibia, flowers have light brown tepal limbs with a

FIGURE 13. — Known distribution of Ferraria variabilis.

30

Bothalia 41,1 (2011)

dark purple longitudinal band toward the base and light

brown margins, while the claws are pale greenish cream

with a narrow darker band running down the midline. The

nectaries in the northern populations are fairly large and

green or less often purple-black. In the western and south-

ern Karoo and the southern Cape, populations have green-

ish yellow or pale to middle brown tepal limbs, usually

darker in the midline, and with small or medium-sized

spots in the lower third or when brown then often with

spots irregularly scattered across the limb and the mar-

gins sometimes slightly darker coloured. The claws are

pale, usually with a darker streak in the midline and have

a small dark nectary at the base or on the inner tepals,

shortly above the base. Perianth colour can vary as much

within a population as across the entire southern Cape and

western Karoo.

Scent is also variable and difficult to describe. In the

protologue of Ferraria antherosa, an illegitimate, super-

fluous name but applicable to F. variabilis, Ker Gawler

(1804) described the scent as reminiscent of green

olives, whereas we have noted both a slightly putrid

smell or a faint, sweet odour in different populations.

Scent characteristics are dependent on temperature and

the condition of the plants, so that little can be said with

certainty except that odours slightly unpleasing to the

human nose are characteristic of the species.

A notable variant of Ferraria variabilis occurs in

the Olifants River Valley and adjacent mountains, rep-

resented by Goldblatt & Porter 12210, and possibly

De Vos 2389 ; leaves are broader and more succulent

than those found elsewhere. The best-developed leaves

are 20-22 mm wide (versus typically 4-12 mm for

most other populations), and the rest of the plant is also

larger, thus the outer spathes are 50-70 mm long and

the inner may reach 85 mm. Plants form circular mats

in stony ground, which suggest spreading by vegetative

means. De Vos (1979: 355) found that her collection

from the Gifberg, was triploid (2 n = 30) and she specu-

lated that it was a hybrid between F. divaricata subsp.

divaricata (i.e. the diploid northern form of F. variabilis

with 2 n = 20) and subsp. australis (the southern tetra-

ploid form with 2 n = 40), both of which occur on the

lowlands below the Gifberg (and have similarly narrow

leaves less than 10 mm wide). Polyploidy and hybrid

vigour seem unlikely to explain the gross differences in

leaf width and thickness of the plants in question. More

likely, they represent a local variant of F. variabilis and

deserve further investigation. Plants from Boklands-

kloof near Lokenburg ( Acocks 19736 ) in the Bokkeveld

Mtns have subspathulate leaves with the blades shorter

than the sheaths and 15-18 mm at the widest point. The

blade margins are also strongly thickened, the thicken-

ings partly wavy to slightly crisped and the apices are

hooked. These plants appear vegetatively very unusual;

unfortunately the flowers are poorly preserved and col-

our was not noted on the label. Revisiting Lokenburg in

2008 we found plants with such leaves but the flowers

were quite typical of F. variabilis.

Some collections of Ferraria variabilis from south-

western Namibia, the Richtersveld and Bushmanland

have long, narrow leaves exceeding the stems and only

2-3 mm wide (e.g. Mittendorf 43; Mostert 1415), so dif-

ferent from the spathes, that the plants appear to repre-

sent another species. Flowers and other features corre-

spond closely to F. variabilis from this, the northwestern

and northeastern extremities of its range.

Populations of Ferraria variabilis from the Tanqua

Basin and nearby (e.g. Goldblatt & Porter 12977, MO,

NBG, PRE) stand out in having the nectaries located

in prominent pouches in the middle of the tepal claws,

rather than at the claw bases. The anthers in the Tanqua

Basin populations and others from nearby (e.g. Jackson

s.n. NBG, from Wuppertal; Compton 17412, NBG, from

Whitehill, incidentally the type of F. divaricata var. aus-

tralis) have the anther lobes held ± parallel, although

joined together only at the tips, as they are in plants

from other parts of the range that have widely diverging

anther lobes.

Histoiy : as discussed above under Ferraria divari-

cata, this name was applied in part by De Vos (1979)

to what we here call F. variabilis. A taller species, F.

divaricata is based on a painting of a plant about 450

mm high, most likely collected in the Clanwilliam Dis-

trict. It has large flowers with dark brown tepal limbs

except for the paler margins, large anthers with widely

divaricate lobes, orange pollen, and long basal leaves

with a grey bloom. That plant corresponds to what De

Vos called F. divaricata subsp. arenosa , which we con-

sidered best treated as a separate species that now bears

the name F. divaricata (Goldblatt & Manning 2005).

The only epithet that can be applied with some degree

of certainty to F. divaricata subsp. divaricata sensu De

Vos is the nomenclaturally illegitimate F. antherosa, a

species based on a plant of uncertain provenance, and

painted from a specimen that flowered in England in

1804. The flowers most closely match those of northern

populations of low-growing, more or less tufted plants

which De Vos distinguished in F divaricata by tepal

markings of solid colour and the claws with large green

nectaries. The name F. antherosa is, however, super-

fluous because Ker Gawler also cited as a synonym F.

viridis Andrews, an orthographic variant of F. viridiflora

(= F. ferrariol a), published in 1803. Ferraria divaricata

subsp. divaricata sensu De Vos was named F. variabilis

by Goldblatt & Manning (2005). The epithet reflects the

variability in tepal colour and marking across its range,

and even within populations. Ferraria atrata Loddiges

(1828) may be this species but the description, accom-

panying the illustration, and a fragment at the Kew Her-

barium that may be the type, are inadequate to identify

the plant. It can even be argued that the species lacks a

description, for the text associated with the painting con-

tains no truly descriptive information. It is impossible

even to determine whether the anther lobes are parallel

or divergent.

Ferraria divaricata subsp. australis was described

by De Vos (1979) for populations that have tepals

speckled and streaked and small dark-coloured nec-

taries at the tepal bases, contrasting with subsp.

divaricata (as she understood it), which has tepals

with solid patterns of colour and larger, pale green-

ish nectaries. We find the distinction less clear in

the field: plants with some tepal streaking or spots

have large nectaries and there seems to be a cline

from the northwest to the southeast of the range of

plants with increasing degrees of tepal spotting and

Bothalia 41,1 (2011)

31

decreasing size of the nectaries. Some northern popu-

lations with tepals marked with solid bands of colour

also have fairly small nectaries. Currently, subsp. austra-

lis is included in the synonymy of F. variabilis.

Representative specimens

NAMIBIA. — 2715 (Bogenfels): Klinghardt Mtns, near ephemeral

pan, (-BC), 22 Sept. 1996, Mannheimer & Mannheimer 271 (PRE,

WIND). 2716 (Witputz): Sperrgebiet, SE side of Auros Mtns, in red

sand, 644 m, (-CB), 11 Aug. 2001, Smook 11318 (WIND); ± 40 km

N of Rosh Pinah, (-DA), 29 Sept. 1983, Goldblatt 7018 (MO); Huib

Hoch Plateau, between Zebrafontein and Witputs, (-DB), 29 Jun.

1989, Oliver 9161 (NBG); Namuskluft, (-DD), MiJtendorf 43 (PRE,

WIND); 12 Sept. 1973, Giess 12900 (PRE). 2816 (Oranjemund):

Luderitz Dist., gravel plain west of rooi duine, (-BA), 1 Oct. 1996,

Mannheimer & Mannheimer 449 (WIND); Schakel Mtn in sand, (—

BA), 1 Aug. 1977, Muller 775 (WIND).

NORTHERN CAPE. — 2816 (Oranjemund): 37 km north of

Lekkersing, red sand, (-DB), 10 Oct., 1991, Germishuizen 5595

(PRE). 2817 (Vioolsdrif): Richtersveld, Rosyntjieberg, dry riverbed, (-

AC), 30 Aug. 1977, Oliver, Tolken & Venter 267 (PRE); 15 km south

of Vioolsdrif, (-DC), 7 Aug. 1976, Giess 14535 (PRE, WIND). 2821

(Upington): Sandveld, Upington, (-AC), 8 Aug. 1961, Mostert 1415

(PRE). 2917 (Springbok): Richtersveld, road to Eksteenfontein, (—

CD), 24 Aug. 1992, Goldblatt & Manning 9312 (MO); 48 km W of

Steinkopf, stony slope, (-BA), 25 Sept. 1974, Goldblatt 2778 (MO).

2918 (Gamoep): Aggenys, Farm Nooisabes, (-AB), 11 Aug. 2000,

Desmet & Opel 2995 (NBG); Goegab Nature Reserve, (-CA), 10 Oct.

1999, Zietsman 3869 (PRE). 2919 (Pofadder): 6 km from Pofadder to

Springbok, (-AB), 21 Sept. 1975, Boucher 2886 (NBG). 2921 (Ken-

hardt): golf course, (-AC), 28 Aug. 1977, De Vos 2395 (NBG). 3019

(Loeriesfontein): 103 km from Brandvlei to Loeriesfontein, (-DC),

Oct. 1975 (fr.), Arnold 905 (PRE). 3023 (Britstown): Britstown, rock

cracks, (-DA), Sept. 1961, Mauve 4133 (PRE). 3119 (Calvinia): Bok-

landskloof, Lokenburg, (-CA), 11 Oct. 1958, Acocks 19736 (PRE);

Agterplaas, west of Calvinia, (-BC), 1 Sept. 1990, Oliver 9615 (NBG).

NORTH-WEST.— 2822 (Glen Lyon): Hay Dist., Langberg, (-DD),

Jul. 1920, Hunter 18 (PRE); Hay Dist., Farm Doring Aar, red sand, (—

DD), 7 Aug. 1936, Acocks 571 (PRE).

WESTERN CAPE. — 3118 (Vanrhynsdorp): top of Gifberg, (—

DB), 25 Sept. 1975, De Vos 2389 (NBG). 3119 (Calvinia): near Nieu-

woudtville, Glen Lyon, rocky hills, (-AC), Goldblatt & Porter 12216

(MO, NBG). 3218 (Clanvvilliam): 12 km N of Clanwilliam on old road

to Bulshoek, (-BB), 15 Sept. 2002, Goldblatt & Porter 12210 (MO);

Clanwilliam, 350 ft [106 m], (-BB), 5 Aug. 1896, Schlechter 964

(PRE). 3219 (Wuppertal): Bidouw Valley, (-AA), 23 Sept. 1952, Mid-

dlemost 1746 (NBG); Farm Jakkalsfontein, 25 km toward Middelpos

from Calvinia-Ceres road, (-BD), 31 Aug. 1982, Snijman 616 (PRE);

Tanqua National Park, Maansedam, 10 Sept. 2007, Goldblatt & Por-

ter 12977 (MO, NBG, PRE). 3220 (Sutherland): ± 2 km N of Matjies-

fontein. Farm Buelhouer, (-BA), 14 Sept. 2004, Snijman 1925 (NBG).

3319 (Worcester): flats east of Prince Alfred’s Hamlet, (-AD). 10 Oct.

1974, Oliver 5062 (MO, NBG, PRE); Doom River, 12 miles [19 km]

N of Villiersdorp. (-CC), 25 Oct. 1962, De Villiers s.n. (NBG). 3320

(Montagu): Tweedside, (-AB), 26 Sept. 1951, Barker 7463 (NBG).

3419 (Caledon): burned stony slopes on western outskirts of Cal-

edon, (-AB), 29 Sept. 2110, Goldblatt & Ndnni 11939 (MO). 3420

(Bredasdorp): 25 km W of Swellendam. slopes of Bromberg, (-AA),

17 Sept. 1978, Goldblatt 4882 (MO).

15. Ferraria flava Goldblatt & J.C. Manning, sp.

nov.

IF. divaricata subsp. aurea M.P.de Vos: 359 (1979). Type: South

Africa, [Western Cape], ± 9 miles [± 13.5 km] north of Lambert’s Bay,

Farm Langdam, 28 Sept. 1973, Van Breda sub M.P de Vos 2297 (STE,

holo.!; PRE, iso., two sheets!).

Plantae ± acaulescentes ad 120 mm altae foliis exclu-

sis, foliis 5-7 ensiformibus ad 300 x 12-20 mm suberec-

tis glaucis ± succulentis, rhipidiis 2-floris, spathis viridi-

bus marginibus transparentibus spatha interna usitate

40-55 mm longa externa vaginanti vel supra divergenti

apice incurvata, floribus flavis limbis tepalorum caer-

uleo-viridibus maculatis marginibus crispis pallidiori-

bus, tepalis exterioribus ± 30 x 10 mm, interioribus ±

12x5 rnm, unguibus ± 15 mm longis, columna filamen-

torum 11-12 mm longa, antheris ± 3.5 mm longis lobis

divaricatis, ramis styli ± 3 mm longis, lobis ramorum

prominenter fimbriatis.

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

veld west of Koekenaap, Farm Kommandokraal, (-AD),

10 Sept. 2008, Goldblatt & Porter 13105 (NBG, holo.;

K, MO, PRE, iso.).

Plants ± acaulescent, up to 120 mm high excluding

leaves. Stem with (2— )4 or 5 primary branches produced

from below or shortly above ground level, each with

several branches, these crowded basally and congested,

together bearing up to 50 rhipidia. Leaves suberect, 5-7,

sword-shaped, up to 300 mm long, lower 3-5 much

exceeding spathes and flowers, mostly 12-20 mm wide,

glaucous, ± succulent, without visible main vein. Rhi-

pidia 2-flowered, numerous, crowded on short, ± hori-

zontal or ascending branches; spathes green, margins

transparent, inner mostly 40-55 mm long, outer ± 2/3

as long as inner, usually entirely sheathing or diverg-

ing above, apex curving inward. Flowers on pedicels

up to 10 mm long, lasting one day, clear yellow, tepal

limbs with minute blue-green spots in lower 7 margins

slightly paler in colour, claws translucent white inside,

forming a floral cup 14-15 mm deep, 8-9 mm wide

at rim, tepal limbs spreading, crisped, usually sweet

vanilla-smelling, nectaries at tepal bases, pale green;

outer tepals ± 30 x l() mm, inner tepal limbs ± 12 x 5

mm, claws of both whorls ± 1 5 mm long. Stamens with

filaments united in a column 11-12 mm long, free and

arching outward in upper ± 1 mm; anthers pale yel-

low with dark purple-brown on the lines of dehiscence,

± 3.5 mm long before anthesis, ± 1.5 mm long after

dehiscence, horizontal, thecae joined only at apices,

widely diverging; pollen pale yellow. Ovary fusiform,

20-25 mm long, with a sterile beak, ± 1 0 mm long, style

branches ± 3 mm long, dividing into diverging, promi-

nently fringed arms, ± 5 mm long; stigmas on small

lobes below tips of style arms and arching over anthers.

Capsules and seeds unknown. Flowering time', early

Sept.-late Oct. Figure 14, Plate 1J.

Distribution and habitat', endemic to the sandveld

of coastal Western Cape and Namaqualand, Ferraria

flava extends from the sandy flat country north of Lam-

bert’s Bay and Klawer to the Groen River and Komag-

gas (Figure 12). Plants grow in deep, moderately fine

sand among sandveld species such as Willdenowia

(Restionaceae) and the small, tree-like, willowy Wibor-

gia obcordata (Fabaceae), and occur most often on the

slopes of low, stabilized and vegetated dunes. It shares

this habitat at some sites with the rare F. ornata which

blooms in May and June and has ripe capsules when F.

flava comes into flower in September. Its pollination

biology is unknown. The flowers last one day, with the

tepals unfolding at ± 9:00 and collapsing at ± 16:00.

They produce small quantities of watery nectar 6-8 %

sucrose equivalents, a feature of F. divaricata and its

immediate allies (Goldblatt et al. 2009). No potential

pollinating visitors have been observed.

32

Bothalia 41,1 (2011)

FIGURE 14. — Ferraria flava, Gold-

blatt & Porter 13105. A, flow-

ering plant; B, flower, side

view; C, inner tepal; D, outer

tepal; E, staminal column and

style; F, anther, undehisced

(above), dehisced (below).

Scale bar: A, B, 10 mm; C, D,

8 mm; E, F, 1.6 mm. Artist:

J.C. Manning.

Diagnosis and relationships'. Ferraria flava is recog-

nized by the yellow, sweetly scented flowers and typi-

cally acaulescent habit with well-grown plants bearing 3

or 4 main branches arising below the ground and bearing

numerous rhipidia set closely together. Combined with

the acaulescent habit are the several broad, firm, almost,

rigid leathery leaves up to 300 x 20 mm. The leaves

recall those of F. divaricata , which has aerial stems and

larger, usually predominantly brown flowers (but some-

times dull yellow) with a wider floral cup. The flowers of

F. flava recall those of F. macrochlamys and its immedi-

ate allies in the narrow floral cup but they are somewhat

larger, with tepals ± 30 x 10 mm and claws ± 15 mm

long, and last a single day, unlike those of the F. macro-

chlamys group, which have somewhat smaller tepals with

shorter claws, 10-14 mm long, and last two days. We

noted that at one site near Koekenaap, F. divaricata , with

tall stems and brown flowers (Goldblatt & Porter 13188

NBG), occurs together with F. flava, but blooms later in

the season, coming into flower there in October.

We provisionally include Ferraria divaricata var.

aurea in synonymy here. The type, from near Lambert’s

Bay, has short, aerial stems but other collections cited by

De Vos (1979) under this name are more or less stemless

Bothalia 41,1 (2011)

33

and do belong here. Flower colour and shape of the type

collection with aerial stems are consistent with F. flava.

We have not been able to locate plants at the type local-

ity and wonder whether subsp. aurea is simply a short-

stemmed variant of F. flava. Living plants from the type

locality need to be examined to determine the true iden-

tity of subsp. aurea.

Representative specimens

NORTHERN CAPE. — 2917 (Springbok): Komaggas, (-DC), 6

Oct. 1934, Herre s.n. (BOL). 3017 (Hondeklipbaai): Farm Hardevlei,

NW of Kotzesrus on road to Groenrivier, sandy flats, bright yellow

with greenish dots in V-shaped pattern, sweetly scented, (-DC), 29

Sept. 1987, Reid 1292 (BOL, MO, PRE).

WESTERN CAPE. — 3117 (Lepelfontein): sandy flats and dunes

west of Lepelfontein, ± 160 m, (-BB), 10 Sept. 2009 (sterile), Gold-

blatt & Porter 13309 (NBG); Brand-se-Baai, sandy slopes inland of

coast, (-DD), 15 Sept. 2008, Goldblatt & Porter 13122 (MO, NBG).

3118 (Vanrhynsdorp): sand dunes 3^1 miles (4. 5-6.0 km] north of

Klawer station, (-DA), 27 Oct. 1944, Leipoldt 4166 (BOL).

Series Uncinatae

16. Ferraria uncinata Sweet in The British flower

garden: t. 161 (1826a); De Vos: 365 (1979), excluding

subsp. macrochlamys. Type: South Africa, without pre-

cise locality, illustration in Sweet l.c. (1826a), originally

collected by W. Synnot (no preserved specimen known).

F. framesii L. Bolus: 123 (1931). Type: South Africa, [Western

Cape], ± 6 miles [± 9 km] south of Clanwilliam, 22 Oct. 1931, Ross

Frames s.n. BOL19928 (BOL, holo.!; SAM, iso.!).

Plants 90-200(-300) mm high. Stem either ± sub-

terranean and then few-branched at base, or stem pro-

duced above ground and branched from upper nodes.

Leaves lanceolate, mostly 8-12 mm wide, sheaths usu-

ally concealing stem, usually without visible main vein,

prominently multi-veined, flat or slightly twisted, apex

often hooked, margins thickened, hyaline, smooth, usu-

ally crisped and undulate at least proximally. Rhipidia

2-flowered; spathes green, usually with hooked tips,

inner mostly 40-60 mm long, outer 26 — 40 mm long (±

V2 to 2/3 as long as inner), usually entirely sheathing.

Flowers on pedicels 12-20 mm long, lasting two days,

usually pale to dark blue-violet, or light brown with blue

speckling, margins crisped, dull yellow-green to khaki,

with a narrow cup, 8-12 mm deep, ± 7 mm wide at rim,

often unscented, faintly foetid-smelling, or scented of

lemon and cinnamon, nectaries minute, at base of tepals;

outer tepals 28-35 * 7-10 mm, limb at least 1.5 times

as long as claw, and up to 3 times as long, often long-

attenuate and coiled, claws 8-12 mm long; inner tepals

28-30 mm long, limbs reflexed, claws slender, 7-12

mm long. Filaments united in a column 7-1 1 mm long,

free in upper ± 1.5 mm; anthers ± 3 mm long, thecae

divergent, shrinking to ± V2 their length after dehis-

cence. Ovaty fusiform, 1 5—28 mm long, with beak 8-12

mm long; style branches ± 1.5 mm long, dividing into

diverging, prominently fringed arms, ± 1.5 mm long;

stigmas terminal on small lobes below tips of style arms

and arching over anthers. Capsules 12-15 mm long,

excluding beak. Seeds angular, 5- or 6-sided, 3^1 mm

long, dark brown, facet surfaces slightly wrinkled. Flow-

ering time: late Aug.-Oct. Plate 1A.

Distribution and biology’’. Ferraria uncinata extends

from the slopes of the Gifberg near Klawer southward.

through the Olifants River Valley and Piketberg to

Malmesbury and Mamre, some 65 km north of Cape

Town (Figure 15). Plants are most often found in sand-

stone outcrops and relatively dry sites, but also some-

times in open ground in coarse, granite-derived gravel or

on sandstone slopes in deep sand.

Diagnosis and relationships’, as circumscribed here,

Ferraria uncinata includes plants with a narrow flor-

al cup 8-12 mm deep, tepals either deep blue to violet

edged with dull yellow-green or in the north of its range,

buff to dull yellow with violet speckles with yellow-

brown margins. The leaves have thickened margins with

at least the edges of the lower leaves crisped and undu-

late. The margin surface itself is smooth. The hooked

tips of the leaves and spathes, the character for which

the species was named ( uncinatus is Latin for hooked),

is frequent, though not constant, not only in F. uncinata

but in several other species of the genus.

De Vos (1959) included the yellow-flowered central

Namaqualand species, Ferraria macrochlamys in F. un-

cinata as subsp. macrochlamys, but we regard this plant

as a separate species (Goldblatt & Manning 2004). It dif-

fers not only in the pale yellow flower colour but the leaf

margins in those populations with thickened margins

resemble those of F. uncinata only superficially. In F.

macrochlamys subsp. macrochlamys the marginal thick-

enings are usually densely ciliate and irregularly ser-

rulate or crenulate, as well as sometimes being slightly

crisped. The tepal limbs of F. uncinata are at least 1.5

times as long as the claws and up to three times as long

in the northern populations, the tepal apices of which are

unusually extended. In F. macrochlamys the tepal limbs

are slightly shorter to ± 1.5 times as long as the claws

and are less prominently attenuate.

The type illustration, of a plant with the tepal limbs

speckled with dark blue dots on a light brown back-

ground, suggests that it was collected between Clanwil-

liam and Bulshoek in the Olifants River Valley. Else-

where across its range, Ferraria uncinata has blue-violet

tepal limbs with somewhat obscure, darker blue spot-

34

Bothalia 41,1 (2011)

ting. The northern populations of F. uncinata differ not

only in colouring, but in the proportions of the flower.

Elsewhere, the floral cup is ± 11-12 mm deep and the

tepal limbs are 16-18 mm long. In the northern popula-

tions, however, the floral cup is 8-9 mm deep and the

tepal limbs are 25-28 mm long, the extended length due

largely to the long, trailing, attenuate tip.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): Olifants River Val-

ley, stony slopes at turn-off to Nardouw, (-BB), 26 Aug. 1957, Lewis

5213 (NBG), 15 Sept. 2002, Goldblatt & Porter 12213 (MO, NBG,

PRE); Clanwilliam, 5 Aug. 1896, Schlechter 8413 (K, MO, PRE, S);

Olifants River Valley, Farm Klawervlei, N-facing slopes at bridge

over Olifants River, (-BD), 2 Sept. 1994, Goldblatt & Manning 9959

(MO); hills N of Aurora near Redelinghuis, (-DA), 14 Sept. 2001,

Goldblatt & Porter 11882 (MO, NBG). 3219 (Wuppertal): Brandewyn

River, (-AA), 26 Aug. 1950, Barker 6572 (NBG); Cedarberg, Algeria

Forest Station, (-AC), 10 Sept. 1997, Van Rooyen et al. 729 (NBG).

3318 (Cape Town): Malmesbury, western edge of town in new devel-

opment, (-BC), 1 Oct. 1998, Goldblatt & Manning 11034 (MO); near

Groenekloof (Mamre), (-AD), Nov. (ft.), Ecklon 305 (K, MO, S). 3319

(Worcester): Mostertshoek, Romans River, (-AC), Sept. 1976, De Vos

2338 (NBG).

17. Ferraria macrochlamys (Baker) Goldblatt

& J.C. Manning in Novon 14: 293 (2004). Lapeirousia

macrochlamys Baker: 338 (1876). F. uncinata subsp.

macrochlamys (Baker) M.P.de Vos: 369 (1979). Type:

South Africa, without precise locality or date, as Herb.

Forsyth (K, holo.!).

Plants mostly 70-100 mm high forming low tufts.

Stem few- to several-branched (rarely simple), branches

crowded close to base; branches and spathes all ± same

length. Leaves narrowly sword-shaped to linear, mostly

(2.0-)4.5-7.0 mm wide, ± straight, or all curving to

same side, or serpentine (loosely wavy in concertina

fashion), tips oblique or hooked, sometimes densely

papillate-ciliate; margins heavily thickened then some-

times velvety, often irregularly serrulate or crenate,

sometimes slightly crisped. Rhipidia 2-flowered; inner

spathes 37-62 mm long, abaxial margins sometimes

crisped, outer 24-50 mm long, usually entirely sheath-

ing, or free distally, often hooked at apex. Flowers on

pedicels 10-23 mm long, lasting two days, pale watery

yellow with slightly darker yellow to light brown mar-

gins, outer tepal limbs often minutely speckled grey-

blue at base, with a narrow cup, 9-13 mm deep, 5-7

mm wide at rim, with a faint, green apple or slightly

sour odour; outer tepals 26-32 mm long, limbs spread-

ing to slightly reflexed, inner tepals 26-28 mm long,

limbs often ultimately ± reflexed, claws of both whorls

10-14 mm long. Stamens with filaments united in a col-

umn 10-12 mm long, free and diverging in upper ± 1

mm; anthers 2. 2-3.0 mm long, thecae divergent; pollen

orange. Ovary fusiform, 1 5-20 mm long with beak 7-15

mm long; style branches 1.0-1. 5 mm long, dividing into

diverging, prominently fringed arms, 1.0-1. 5 mm long;

stigmas on small lobes below tips of style arms and

arching over anthers. Capsules 15-20 mm long exclud-

ing beak. Seeds angular, mostly 5- or 6-sided, ± 2. 7-3.0

mm diam., dark brown, facets slightly wrinkled. Flow-

ering time-, late Aug. to end of Sept., rarely to mid-Oct.

Figures 16-19.

Distribution and biology-, an exclusively Namaqua-

land species, the range of Ferraria macrochlamys

extends from Steinkopf in the north to the southern foot-

hills of the Kamiesberg near Bitterfontein in the south

and west toward the coast (Figure 17). Plants grow on

a variety of soils, subsp. macrochlamys and subsp.

kamiesbergensis in gritty to loamy granite-derived soils,

sometimes in granite outcrops, in vegetation dominated

by succulent-leaved shrubs and subsp. serpentina mostly

in quartzitic sand, usually among rocks. As in other

members of series Uncinatae, the flowers last two days

and often have a slightly unpleasant, sour odour, but

sometimes they appear to have no scent at all. Pollina-

tion biology is known only for subsp. kamiesbergensis ,

the flowers of which are pollinated by the masarine wasp

(Vespidae: Masarinae) Ceramius (Gess 1997, reported as

F. divaricata , now F. variabilis ). So similar are the flow-

ers of all three subspecies that we infer the same pollina-

tion strategy for the other two subspecies.

Diagnosis and relationships : De Vos (1979) treated

Ferraria macrochlamys as a subspecies of the dark

blue-flowered F. uncinata , a plant of the western half of

Western Cape. She considered that the two shared iden-

tically specialized leaves with heavily thickened and

crisped margins and derived flowers with a narrow floral

cup and that they differed only in flower colour, either

violet-blue with dull greenish yellow margins in subsp.

uncinata or pale yellow with yellow to brown margins

in subsp. macrochlamys . The leaf morphology in the two

taxa is, however, not identical: whereas Ferraria unci-

nata has leaf margins crisped and sometimes undulate

with the thickened edges smooth, F. macrochlamys has

more often straight leaf margins but the marginal thick-

enings, when present, are usually irregularly serrate to

crenate, or sometimes crisped, and are also sometimes

densely velvety. Only occasionally do the lower leaf

blades have undulate or crisped margins. Their flow-

ers differ in size as well as colour. The tepal limbs of F.

uncinata exceed the claws by at least 1.5 times and are

usually at least twice and sometimes up to three times

as long, and ( 1 8 — )28— 3 5 mm long (with claws 8-12 mm

long), whereas the tepals limbs of F. macrochlamys are

15-20 mm long and at most half again as long as the

claws, 10-18 mm long, and are less strongly attenuate.

While F. macrochlamys is clearly allied to F. uncinata, it

seems to us most closely related to F. brevifo/ia, a local

endemic of southern Namaqualand. This species has vir-

tually identical flowers to those of F. macrochlamys but

the broad leaf blades are shorter than the sheaths.

We recognize three subspecies in Ferraria macro-

chlamys, based on differences in leaf morphology and

sometimes in habitat. Only subsp. macrochlamys has

leaves with thickened and crisped to undulate margins

and sometimes papillate-ciliate leaf blades. The ± falcate

leaves of subsp. kamiesbergensis are all recurved in the

same direction and have smooth, unthickened margins.

In subsp. serpentina the leaf margins are also unthick-

ened but the blades are serpentine, thus loosely wavy in

concertina fashion.

History-. Ferraria macrochlamys was initially des-

cribed as Lapeirousia macrochlamys by J.G. Baker

(1876). It remained so poorly understood that in 1931

H.M.F. Bolus, who collected plants at Bowesdorp in

central Namaqualand in 1929, described it anew as F.

crispulata. Goldblatt (1972), in his revision of southern

Bothalia 41,1 (201 1 )

35

African Lapeirousia, noted that the type of L. macro-

chlamys was a species of Ferraria akin to F. crispulata,

but it remained to M.P. de Vos in her revision (De Vos

1979) to transfer the species to the genus. The species

was based on a specimen in the Kew Herbarium, osten-

sibly collected by W. Forsyth. It is now understood that

the specimen was part of a collection purchased in 1 835

by George Bentham of the Royal Botanic Gardens,

Kew, from the estate of William Forsyth, son of W.F.

Forsyth (1737-1804). Forsyth, a distinguished Scottish

horticulturist, was one of the founders of the (Royal)

Horticultural Society of London and was also associ-

ated with the Chelsea Physic Garden (1771-1784). The

brief comment about Forsyth by Gunn & Codd (1981)

in their history of plant collecting in southern Africa, is

thus inaccurate with regard to Forsyth (Nelson 2006).

It seems reasonable to suggest that at least some speci-

mens of the Forsyth herbarium were gathered by the

Scottish botanist, James Niven, who collected plants at

the Cape for British patrons and later the Empress Jose-

phine of France. Niven visited Namaqualand in 1799

and travelled to the Kamiesberg through country where

F. macrochlamys grows. Another possibility is that F.

mcicrochlamys was first collected by William Paterson,

who travelled through Namaqualand collecting plant and

animal specimens in 1778 and 1779. Paterson was for a

time associated with W.F. Forsyth.

Key to subspecies

la Leaf margins thickened, hyaline, and sometimes shortly vel-

vety, at least lower leaves with serrulate to crenulate mar-

gins; leaf blades ± straight or laxly undulate; outer spathes

sheathing for most of their length and no more than 2/3 as

long as inner 17a. subsp. macrochlamys

lb Leaf margins hardly, if at all, thickened, and smooth:

2a Leaf blades in upper 7 inclined to curving to same side;

outer spathes not sheathing in distal half and ± as long

as inner; outer tepals 25-35 mm long; filament column ±

12.5 mm long 17b. subsp. kamiesbergensis

2b Leaf blades in upper 7, serpentine, thus loosely folded

back on themselves in concertina fashion; outer spathes

sheathing for most of their length and ± 2/3 as long as

inner; outer tepals 25-30 mm long; filament column

10-11 mm long 17c. subsp. serpentina

17a. subsp. macrochlamys

Fen-aria crispulata L. Bolus: 260 (1931). Type: South Africa,

[Northern Cape], near Bowesdorp, 27 Aug. 1929, L. Bolus s.n.

BOL19162 (BOL, holo.l; BM, Kl, iso.).

Plants with leaves often densely papillate-ciliate;

margins heavily thickened, thickenings smooth or

densely ciliate, often irregularly serrulate or crenate,

sometimes slightly crisped. Rhipidial spathes : inner

45-62 mm long, abaxial margins sometimes crisped,

outer 30-50 mm long, usually entirely sheathing, or

tips free. Flowers with narrow cup 11-13 mm deep, ± 7

mm wide at rim, with faint, green-apple or slightly sour

odour; outer tepals 28-32 mm long, limbs spreading to

slightly reflexed, inner tepals 26-28 mm long, claws of

both whorls (11 — ) 1 2 — 1 4 mm long. Stamens with fila-

ments united in column ± 12 mm long, free and diverg-

ing in upper ± 1 mm; anther thecae ± 3 mm long before

dehiscence. Ovary’ 23-28 mm long with beak 15-18 mm

long. Flowering time', late Aug.-late Sept. Figure 16.

Distribution and habitat', restricted to central Namaqua-

land, subsp. macrochlamys extends from Steinkopf in the

FIGURE 16. — Ferraria macrochlamys subsp. macrochlamys. Gold-

blatt. Manning & Porter 13406 (NBG). Scale bar: 10 mm. Art-

ist: J.C. Manning.

north to Garies in the south and inland to the northern

Kamiesberg (Figure 17). Plants grow in gritty to loamy

granite-derived soils.

Diagnosis', most distinctive of the three subspecies,

subsp. macrochlamys stands out in its spreading leaves

with thickened margins that are crenate or crisped and

sometimes velvety. The flowers often serve to distin-

guish it from subsp. serpentina in their slightly larger

size, but that difference is not consistent, hence unre-

liable. Subsp. macrochlamys is always acaulescent,

whereas subsp. serpentina , in years of ample rainfall,

develops an aerial stem rendering it quite different in

general appearance.

Representative specimens

NORTHERN CAPE.— 2917 (Springbok): 48 km west of Stein-

kopf, stony slope, (-BA), 25 Sept. 1974, Goldblatt 2778 (PRE); 1 mile

[1.6 km] west of Springbok, (-DB), 9 Sept. 1950, Barker 6701 (NBG);

Mesklip, (-DD), Lewis 1035 (SAM). 3017 (Hondeklipbaai): road

to Soebatsfontein, ± 1 km from N7, (-BB), 17 Sept. 2002, Goldblatt

& Porter 12229 (MO); 1 km south of Kamieskroon, (-BB), 29 Sept.

1981, Hugo 2889 (PRE); top of Garies hill along road to Hondeklip-

baai, (-BD), 2 Sept. 2000, Goldblatt & Ncinni 11454 (MO, NBG), 4

Sept. 1945, Compton 17179 (NBG). 3018 (Kamiesberg): Kamiesberg,

slopes north of Leliefontein, burned previous summer, (-AC), 1 7 Sept.

2002, Goldblatt & Porter 12228 (MO).

36

Bothalia 41,1 (2011)

16° 18° 20° 22° 24°

_j i i i i

FIGURE 17. — Known distribution of Ferraria macrochlamys subsp.

macrochlamys, O; subsp. kamiesbergensis, •; subsp. serpenti-

na, A ; and F. brevifolia, ik.

17b. subsp. kamiesbergensis (M.P.de Vos) Gold-

blatt & J.C. Manning, comb, et stat. nov., Ferraria

kamiesbergensis M.P.de Vos: 362 (1979). Type: South

Africa, [Northern Cape], Kamiesberg, Rondefontein, 1

Sept. 1976, Oliver 5970 (NBG, lecto.!, designated by

Goldblatt & Manning, 2004: 292).

Plants forming low tufts, rarely unbranched. Leaves

firm, suberect-curved to falcate-incurved, mostly 70-100

x 3-6 mm, all weakly to strongly curving to same side;

margins plane, barely or not at all thickened. Rhipidial

spathes'. inner 50-65 mm long, outer 50-57 mm long,

sheathing in lower half, arching outward distally. Flow-

ers with tepal claws forming narrow cup mostly 12-14

mm deep, ± 7 mm wide at rim, limbs laxly spreading;

outer tepals 25-35 mm long, 8-10 mm at widest, limbs

13-20 mm long, slightly exceeding claws, inner tepals

25-30 mm long, claws of both whorls 12-14 mm long.

Stamens with filaments united in a column 10.5-12.5

mm long, free and diverging in upper 1 mm; anther

thecae ± 3 mm long, shorter after anthesis. Ovary

18-22 mm long with tubular beak ± 12 mm long; style

branches ± 1 mm long, dividing into diverging, promi-

nently fringed arms, ± 2 mm long. Flowering time :

Sept.-early Oct. Figure 18, Plate IB.

Distribution and biology, subsp. kamiesbergensis is

centred in the southern highlands where it extends from

Rooiberg in the Kamiesberg to Bitterfontein but isolated

populations occur to the east toward Loeriesfontein and

the hills northwest of Calvinia in the western Karoo

(Figure 17). In the Kamiesberg, plants grow in coarse,

gritty soil derived from decomposed granite or in cracks

in granite pavement. Soils favoured by F. kamiesber-

gensis in the western Karoo are not recorded and we

have not seen the species there ourselves. The flowers

last two days and, in the one population we examined

for nectar, produced modest quantities of dilute nectar,

mean 1 0.6 % sucrose equivalents. The one record of pol-

lination in subsp. kamiesbergensis, by Gess ( 1997, origi-

nally reported as F. divaricata ) is by the masarine wasp,

Ceramius (Eumenidae). We have seen no potential pol-

linators on the other subspecies.

Diagnosis and relationships : subsp. kamiesbergensis is

distinctive only in its vegetative morphology. The leaves

are firm and leathery to almost succulent and the blades

are often dark green and in the upper half all curve to the

same side and the margins are plane, smooth and not at

all, or only barely, thickened. Otherwise plants have the

tufted habit of subsp. macrochlamys and flowers virtually

identical to that subspecies. The ranges of subsp. kamies-

bergensis and subsp. macrochlamys do not overlap: pop-

ulations of the latter are mostly from the highlands west

and north of the Kamiesberg and in the Kamiesberg itself

are only known from Leliefontein and nearby, which is

north of the range of subsp. kamiesbergensis.

FIGURE 18. — Ferraria macrochlamys subsp. kamiesbergensis. Gold-

blatt & Porter 13176. Scale bar: 10 mm. Artist: J.C. Manning.

Bothalia 41,1 (2011)

37

Typification of this taxon has proved problematic

(Goldblatt & Manning 2004) as the illustration in the

protologue (De Vos 1979) is not this species but, at least

in leaf, resembles the widespread and common Ferraria

variabilis. Of the three plants on the type sheet only one,

now the lectotype, resembles subsp. kamiesbergensis.

When seen alive, the flowers of the true F. kamiesber-

gensis and F. variabilis are quite different but in speci-

mens with poorly pressed flowers, they are remarkably

similar. Because of lingering confusion over the identity

and circumscription of F. kamiesbergensis (now subsp.

kamiesbergensis ) we list all specimens of the subspecies

that we have seen below, including our own recent col-

lections.

Representative specimens

NORTHERN CAPE. — 3018 (Kamiesberg): Farm Karas, Die Kom,

(-CA), Oct. 1940, Leipoldt 3563 (BOL); Wilgehout Ravine, (-CA),

Sept. 1911, Pearson 6812 (K); Farm Rondefontein, south of Karas,

in fallow field on SE-trending slope, (-CA), 17 Sept. 2002, Goldblatt

& Porter 12223 (MO, NBG); NE slopes of Rooiberg, (-CA), 17 Sept.

1987, Davidse 33353 (MO); southern Kamiesberg, between Farms

Doringkraal and Gemsbokkloof, (-CA), 25 Sept. 2008, Goldblatt &

Porter 13176 (MO, NBG). 3019 (Loeriesfontein): between Kliprand

and Loeriesfontein. Farm Onderste Camdini, (-CD), 11 Sept. 1976,

Thompson 2882 (NBG). 3119 (Calvinia): near Hantams River, 30

miles (± 45 km) NW of Calvinia on Loeriesfontein road, (-BB), 26

Sept. 1952, Lewis 2534 (SAM); 32 miles (± 48 km) from Calvinia to

Loeriesfontein. (-BB), 27 Sept. 1952, Johnson 603 (NBG).

WESTERN CAPE. — 3018 (Kamiesberg): ± 10 km north of Bitter-

fontein. sandy slope, (-CC), 2 Sept. 2002, Goldblatt & Porter 12146

(MO).

17c. subsp. serpentina Goldblatt & J.C. Manning,

subsp. nov.

Ferrariae macrochlamydi subsp. macrochlamydi

similis sed foliis linearibus usitate 3-4 mm latis laminis

undulatis marginibus non incrassatis, spathis interioribus

37-45 mm longis, exterioribus 24-38 mm longis, flo-

ribus cupulo florali 9-11 mm profundo munitis, limbis

tepalorum exteriorum 16-18 mm longis interiorum ± 16

mm longis, ovario 15-28 mm longo.

TYPE. — Northern Cape, NE of Kotzesrus on road to

Garies, 16 Sept. 2001, Goldblatt & Porter 11900 (NBG,

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

Plants like subsp. macrochlamys with branches

crowded close to base or branching above ground, thus

forming rounded plants. Leaves in a loose fan, linear,

(2-)3— 4 mm wide, without a central vein, blades ser-

pentine (loosely wavy in concertina fashion, sometimes

appearing coiled when pressed); margins plane, not

thickened. Rhipidial spathes: inner 37^45 mm long,

outer 24-38 mm long, sheathing in lower two thirds.

Flowers on pedicels 10-15 mm long, with tepal claws

forming a narrow cup, 9-11 mm deep, ± 5 mm wide at

rim; outer tepal limbs 16-18 mm long, tips attenuate

and twisted, inner tepal limbs ± 16 mm long, slightly

reflexed, claws of both whorls 10-12 mm long. Sta-

mens with united filaments in a column 10-11 mm

long, free in upper 1 mm; anther thecae ± 2.2 mm long.

Ovary 15-28 mm long, with a beak 7-18 mm long; style

branches diverging, with style arms ± 1 mm long. Flow-

ering time4, late Aug. and Sept., rarely to mid-Oct. Figure

19.

Distribution and biology : restricted to central south-

ern Namaqualand, subsp. serpentina occurs on the low,

sandy or stony hills inland of the coast between Killians

Pass and Kotzesrus (Figure 17). Plants grow in gritty

soils, either on stony slopes or in deep. red. quartzitic

sand. Like the other subspecies of Ferraria macro-

chlamys, the flowers of subsp. serpentina produce mod-

est quantities of unusually dilute nectar, mean 8.1 %

sucrose equivalents (Goldblatt et al. 2009). Pollination is

unknown.

Diagnosis and relationships : evidently first col-

lected by the American botanist, A.L. Grant in 1929 in

‘Namaqualand' and shortly thereafter by R.H. Compton

near Wallekraal in 1935, these specimens were misun-

derstood by De Vos (1979), who identified them as F.

divaricata subsp. divaricata (now F. variabilis). Later

collections made in 2001 and 2002 east of Kotzesrus,

some 70 km south-southeast of Wallekraal, but in a simi-

lar sandy habitat, made it clear that the flowers of these

plants closely resemble those of what are now F. macro-

chlamys subsp. macrochlamys and subsp. kamiesbergen-

sisdiffering if at all only in their slightly smaller size.

Thus they have a pale yellow perianth and narrow, com-

paratively deep floral cup. Most striking are the leaves,

the narrow blades of which are undulate in loose concer-

tina fashion and have unthickened, plane margins. The

leaf contrasts starkly with the straight leaves of typical F.

macrochlamys, which have strongly thickened, crisped

or crenate margins, and in addition in some populations,

shortly velvety. The flowers of subsp. serpentina have

a floral cup, 9-11 mm deep, ± 5 mm wide at the rim,

outer tepals 25-30 mm long, and filaments 10-11 mm

long, thus slightly smaller than in most populations of

typical F. macrochlamys, which have a cup, 11-13 mm

deep, 6-7 mm wide at the rim. outer tepals 28-32 mm

long, and filaments ± 12 mm long. The inner spathes

are 37^45 mm long, compared to 45-62 mm in subsp.

macrochlamys, and the flowers have pedicels 10-15

mm long (versus 20-23 mm in subsp. macrochlamys),

reflecting the shorter spathes. A particularly narrow-

leaved form of subsp. serpentina has been recorded in

the Komaggas area west of Springbok by Rupert Koop-

man, the blades ± 2 mm wide and when dry, showing the

outline of a raised central vein.

Representative specimens

NORTHERN CAPE. — 2917 (Springbok): near Kommagas, sandy

field, (-CD), 10 Sept. 2007, Koopman s.n. (NBG). 3017 (Hondeklip-

baai): Wallekraal, (-BC), 30 Aug. 1935, Compton 5405 (BOL, NBG);

Farm Horees 441, on road between Killian’s Pass and Wallekraal,

(-BC), 8 Oct. 2003, Le Roux 5124 (NBG); road to Groen River Mouth,

near Farm Nariep, (—DC), 16 Sept. 2008, Goldblatt & Porter 13131

(MO, NBG); ± 6 km NE of Kotzesrus on road to Garies, 573 ft [175

m], (-DD), 28 Aug. 2001, Goldblatt & Porter 11782 (MO), 2 Sept.

2002, Goldblatt & Porter 12141 (MO, NBG, PRE); ± 10 km NE of

Kotzesrus, (-DD), 3 Sept. 2001, Goldblatt c£ Porter 11820 (MO).

WESTERN CAPE. — 3018 (Kamiesberg): Bitterfontein-Garies,

at border of Northern Cape, stony slopes facing Swartdoom River,

(-CC), 24 Sept. 2008, Goldblatt & Porter 13167 (MO, NBG, PRE).

18. Ferraria brevifolia G.J. Lewis in Annals of

the South African Museum 40: 1 19 (1954); De Vos: 369

(1979). Type: South Africa, [Western Cape], 2 miles [3

km] south of Nuwerus, Sept. 1945, Lewis 1374 (SAM,

holo.!).

38

Bothalia 41,1 (201 1 )

FIGURE 19. — Ferraria macro-

chlamys subsp. serpentina,

Goldblatt & Porter 13131 . A,

flowering plant; B, staminal

column and style; C, capsules.

Scale bar: A, C, 10 mm; B, 2.5

mm. Artist: J.C. Manning.

Plants 70—1 20(— 1 80) mm high, usually ± acaulescent.

Stem with 2-A short branches borne close to ground,

all ± equal in length. Leaves overlapping to form a

tight fan, sheaths completely enclosing stem and much

exceeding blades, blades oblong, mostly 7-15 x 5-7(-

10) mm, obtuse or acute with tips often hooked; margins

thickened especially abaxially, thickenings often crenate

to slightly crisped. Rhipidia 2-flowered; inner spathe

mostly 45-55 mm long, outer 35-38 mm long, sheathing

in lower 2/3, arching outward distally. Flowers on pedi-

cels 15-23 mm long, lasting two days, pale to watery

yellow with darker yellow to brown or orange margins,

nectaries small, at base of tepals, claws forming a nar-

row cup, ± 12 mm deep, ± 7 mm wide at rim, faintly

sweet-scented; outer tepals 32-40 x ± 9 mm, attenuate,

tips slightly coiled, claws slender, 10-13 mm long, inner

tepals 30-37 mm long. Stamens with filaments united in

a column ± 10 mm long, free in upper ± I mm; anther

thecae divergent, initially ± 2.5 mm long, ± 1 .5 mm

long after dehiscence; pollen orange. Ovary fusiform,

( 1 5 ) 1 8—25 mm long, with a beak 8-12 mm long; style

branches ± 1.5 mm long, dividing into diverging, promi-

nently fringed arms, ± 1 .5 mm long; stigmas on small

lobes below tips of style arms and arching over anthers.

Capsules ovoid, 15-25 mm long. Seeds angular, irregu-

larly 5- or 6-sided, ± 3 mm diam., brown, shiny. Flower-

ing time : Aug.-Sept.

Distribution and biolog\>\ with a recorded range cov-

ering a linear distance of about 40 km (Figure 17), Fer-

raria brevifolia has one of the narrowest distributions

of any member of the genus. Records are mostly from

the immediate vicinity of Nuwerus, but plants have also

been collected near Bitterfontein to the north — its most

northern station is just 10 km north of the town. Plants

occur on shrub-covered slopes of gritty, granite-derived

ground. The flowers last two days and secrete small

quantities of nectar of the low concentration, ± 6.4 %

sucrose equivalents, typical of the F. uncinata alliance.

There are no reports of pollination in the species.

Diagnosis and relationships', distinctive in Ferraria ,

the leaves of F. brevifolia form a tight, 2-ranked fan and

have elongate sheaths 40-80 mm long, much exceed-

ing the short, ovate, obtuse blades, 7-15 mm long. The

blades, usually less than half as long as the sheaths, have

relatively thick margins and oblique, acute or apiculate,

hooked tips. The pale yellow flowers, in contrast, are

virtually identical to those of F. macrochlamys and have

Bothalia 41,1 (201 1 )

39

a narrow floral cup, ± 12 mm deep, and tepal limbs,

22-28 mm long, with crisped, light brown or rarely

reddish brown margins. The flowers produce a faint,

slightly sweet scent. They can be distinguished from

those of its apparent nearest relative, F macrochlamys,

if at all, by their somewhat longer outer tepals with long,

tapering loosely coiled tips. Although only formally

described in 1954, an early collection of the species, and

possibly the first, Schlechter 11031 , made in 1897, bears

the annotation 'F. namaquensis Schltr.’ in Schlechter’s

hand.

Representative specimens

WESTERN CAPE. — 3018 (Kamiesberg): ± 10 km N of Bitter-

fontein, granite bank, (-CC), 2 Sept. 2000, Goldblatt & Nanni 11450

(MO); hills at Bitterfontein, 1200 ft [365 m], (-CC), 2 Sept. 1897, Sch-

lechter 11031 (BOL, K, MO, PRE, S; B, BM, BR, Z, not seen). 3118

(Vanrhynsdorp): Nuwerus, stony veld, (-AB), 7 Sept. 1945, Barker

3734 (NBG), 6 Aug. 1972 (flowered at Vredendal), Hall 4114 (NBG,

PRE); 2 miles [3 km] south of Nuwerus, (-AB), 6 Sept. 1974, De Vos

2326 ( PRE).

EXCLUDED SPECIES

Ferraria jimbriata Burm. in Nova acta physico-

medico Academiae Caesareae Leopoldino-Carolinae

germanicae naturae curiosorum 2: 201 (1761). Type:

South Africa, without precise locality, illustration in

Burman, loc. cit. : t. 3, fig. 2 (1761).

The woodcut illustration is inadequate to detennine

this plant to species (see History). Our best guess is that

this may represent Ferraria divaricata or alternatively F.

variabilis.

ACKNOWLEDGEMENTS

Support for this study by grants 7103-01. 7316-02,

7799-05 and 8248-07 from the National Geographic

Society is gratefully acknowledged. We extend our grati-

tude to Ingrid Nanni, Greg Nicolson and Lendon Porter

for their assistance and companionship in the field; Felix

Forest for help with the molecular phylogenetic analy-

sis; Rupert Koopman for alerting us to the existence of

Ferraria ornata , and Koos Claasens and Francois Kotze

for helping us establish the range of F. parva. Collect-

ing permits were provided by the Nature Conservation

authorities of Northern Cape and Western Cape Prov-

inces, South Africa. We thank the curators of the fol-

lowing herbaria for loans of types and other specimens

among their collections: B. BM, BOL, K, MO and PRE.

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Bothalia 41,1: 41-82 (2011)

Annotated catalogue of the flowering plants of Sao Tome and Principe

E. FIGUEIREDO*, J. PAIVA**, T. STEVART***, F. OLIVEIRA* and G.F. SMITH+

Keywords: diversity, flowering plants, Sao Tome and Principe

ABSTRACT

A catalogue of the flowering plants of the islands of Sao Tome and Principe (Gulf of Guinea) is presented. Flowering plant

diversity figures are updated to: 135 families (20 introduced), 624 genera (172 introduced), 1 104 species (301 introduced)

plus 15 additional infraspecific taxa. At present 119 taxa ( 107 species and 12 infraspecific taxa) are known to be endemic to

the two islands. The catalogue includes accepted names, synonyms used in recent literature, common names, voucher speci-

mens and information on habit and habitat and on plant uses, particularly medicinally.

INTRODUCTION

The islands Sao Tome and Principe (STP) are part

of an archipelago of four islands situated in the Gulf

of Guinea. These islands resulted from volcanic activ-

ity along a NE-SW line of fractures that extends from

Mounts Kupe and Manenguba in Cameroun, stretching

southward towards St Helena. S.Tome and Principe form

a political entity, whereas the two other islands, Bioko

and Annobon, are part of Equatorial Guinea. Bioko

lies on the continental shelf, and is separated from the

mainland by a stretch of water that is less than 100 m

deep, but the other three islands apparently have an oce-

anic origin and are generally thought to have never been

linked to the continent (e.g. Exell 1944, 1956). Bioko’s

flora is similar to that of the neighbouring continent; it

has a higher diversity of Angiosperms than those of the

other three islands, but lower levels of endemism. Anno-

bon is the furthest away from the mainland. Its flora has

characteristics of that of an oceanic island (Exell 1973).

The first studies of the flora of S.Tome and Principe

were undertaken at the University of Coimbra, Portu-

gal, by J. Henriques (1838-1928), who based his work

on the collections made during the 19th century by A.

Moller (1842-1920) and F. Quintas (birth and death

dates unknown, collected in 1885-1893). In 1932, A.

Exell (1901-1993) visited the four islands in the Gulf

of Guinea as part of a four-month expedition, collect-

ing extensively. Resulting from this collecting trip, he

eventually published the Catalogue of the vascular

* Department of Botany, Nelson Mandela Metropolitan University, RO.

Box 77000, Port Elizabeth 6031 / Centre for Functional Ecology, De-

partment of Life Sciences, University of Coimbra, 3001-455 Coimbra,

Portugal.

** Centre for Functional Ecology, Department of Life Sciences, Uni-

versity of Coimbra, 3001-455 Coimbra, Portugal.

*** Missouri Botanical Garden, Africa & Madagascar Department,

P.O. Box 299, 63166-0299, St Louis, Missouri, USA / Herbarium et

Bibliotheque de Botanique africaine, Universite Libre de Bruxelles -

ULB, 50 Av. F. Roosevelt, CP 169, 1050 Bruxelles. Belgium; National

Botanic Garden of Belgium, Domein van Bouchout, Nieuwelaan 38,

B-1860 Meise, Belgium.

• Herbario Nacional de Sao Tome, Jardin Botanico de Bom Sucesso,

Sao Tome e Principe.

+ Research and Scientific Services, South African National Biodiver-

sity Institute, Private Bag X101, Pretoria 0001 / Acocks Chair, Sch-

weickerdt Herbarium, Department of Botany, University of Pretoria,

Pretoria 0002.

MS. received: 2010-03-17.

plants of S.Tome (Exell 1944), in which 17 new spe-

cies were described and several new records noted. In

the ensuing years, Exell published papers (e.g. Exell

& Rozeira 1958) and a supplement to the catalogue

(Exell 1956), and finally produced a preliminary check-

list, Angiosperms of the islands of the Gulf of Guinea

(Exell 1973). Other relevant botanists who collected on

these islands were A. Chevalier (1873-1956) in 1905,

T. Monod (1902-2000) in 1956 and J. Espirito Santo

(1901-unknown) in the 1960s-70s.

After a period of 20 years, the interest in the flora

of these islands was rekindled through the European-

funded ECOFAC project (www.ecofac.org), as well as

by research on medicinal plants. Extensive collecting

was done over the last 10 years by Santomean, Belgian

and Portuguese collectors. Several studies were pub-

lished on the flora of the islands (see Figueiredo 1994a

for a comprehensive botanical bibliography), but the

only published plant diversity list remains Exell’s pre-

liminary checklist (1973) which was a simple list of

names without vouchers or further information.

Numerous new collections were made over the past

20 years, and it is now possible to improve Exell’s list

considerably. In this new catalogue we include com-

mon names, habit and habitat, medicinal uses and infor-

mation on the ornamental or food value of the taxa.

Recent trends to interlink plant diversity studies, such

as conducting comprehensive surveys of useful plants,

have resulted in a fresh appreciation for the consider-

able value of herbaria as untapped data sources. There

has also been a resurgence in an appreciation for com-

mon names as an important source of information for

plant uses. The reliability of such information increases

exponentially if the names can be linked to specimens,

given that such vouchered anecdotes can be verified by

future researchers. In addition, common names represent

an important, but sadly vanishing, component of the cul-

ture of a country and its peoples. In this paper we create

the critically important link between voucher specimens,

scientific names in current use, common names, and uses

of the plants.

Phytogeography

The island of S.Tome in particular, has floristic ele-

ments that are remarkable in the context of African phy-

42

Bothalia 41,1 (201 1 )

togeography. In this regard, disjunctions of afromontane

taxa are particularly striking. For example, Melchiora

mannii (Oliv.) Kobuski, which is endemic to S.Tome,

is closely related to a species occurring in the eastern

parts of the Democratic Republic of Congo (DRC) and

on the Usambara and Uluguru Mountains. More than

2 000 km separate these two species. Afrocarpus man-

nii (Hook.f.) C.N.Page, an endemic of the summit of

S.Tome, is another well-known example of a wide dis-

junction. Its nearest relative, Afrocarpus usambarensis

(Pilg) C.N.Page, occurs in eastern Africa. The same type

of disjunction is also found in some lowland forest taxa,

such as Lasiodiscus mildbraedii Engl. This species is

distributed in the eastern DRC and, at much lower alti-

tudes, in S.Tome, in remnants of the lowland forest. A

single specimen was also collected in Cameroun.

Narrow afromontane palaeoendemics are probably

remnants of species which were much more widespread

during cooler climatic periods, becoming restricted to

mountains during warmer periods. When the climate

became colder allowing afromontane taxa to extend

to lower altitude, several lowland species became con-

fined to refuges that retained suitable environmental

conditions. They may have survived on the island where

adverse conditions were ameliorated by the effect of a

milder oceanic climate. Exell (1944) suggested that

S.Tome might be of a different ‘botanical age’ in rela-

tion to the other islands in the archipelago, as well as

Mount Cameroun, which would explain the confinement

of several taxa to that island and their absence from the

other mountains on the mainland. He hypothesized that

although these mountains might be of the same geo-

logical age, S.Tome might have been the first to become

available for colonization and it would contain elements

(palaeoendemics) of an older flora which was already

confined to refuges when the other local mountains

became available for colonization.

Vegetation

When the islands of S.Tome and Principe were dis-

covered by Portuguese sailors in 1471-1472, they were

uninhabited and totally covered with dense tropical rain-

forest. The occupation of the islands took place in 1486

and with it the establishment of farms ( rogas ), initially

for the production of sugar cane, later for cocoa and cof-

fee. These had, and still have, a profound impact on the

vegetation of the islands. Deforestation was particularly

high during the 1 9th century, when the lowland forests,

up to an altitude of ± 1 500 m were almost completely

replaced with plantations, with a few original trees kept

for shade. When, at the end of the 16th century, sugar

cane production was stopped, there was a regeneration

of part of the vegetation, giving rise to large patches

of secondary vegetation. Several classifications exist

for the vegetation of the island of S.Tome, but the

most commonly used one is that of Monod (1960) who

divided the island into four regions based on character-

istic species. The littoral region includes dunes [with

Ipomoea pes-caprae (L.) R.Br. subsp. brasiliensis (L.)

Ooststr., Canavalia rosea (Sw.) DC., Cynodon dactvlon

(L.) Pers. and Sporobolus virginicus (L.) Kunth] and

mangroves (with Rhizophora harrisonii Leechm., Acro-

stichum aureum L. and Pandanus thomensis Henriq.).

The next region, extending to an altitude of 800 m

includes the lowland rainforest that was mostly replaced

with cocoa plantations. In the north of the island where

rainfall is much lower, there are anthropogenic savannas

characterized by grasses and legumes. At an altitude of

800-1 400 m, the mountain rainforest is dominated by

Rubiaceae and Euphorbiaceae and many epiphytes, par-

ticularly orchids. Among the numerous endemics char-

acteristic of this area, are several trees such as Trichilia

grandifolia Oliv., Craterispermum montanum Hiem,

Tabernaemontana stenosiphon Stapf, Erythrococca

molleri (Pax) Prain and Discoclaoxylon occidentale

(Miill.Arg.) Pax & K.Hoffm., and the lianas Mussaenda

tenuiflora Benth. var. thomensis G. Taylor, Sabicea exel-

lii G. Taylor and Ecpoma cauliflora (Hiem) N. Halle. The

undergrowth is home to species of Begonia L. and Pep-

eromia Ruiz & Pav., and the very common Psychotria

subobliqua Hiem and Ca/voa crassinoda Hook.f.

Above 1 400 m and up to the summit of the island

(2 024 m), a mist forest is encountered with the endemic

trees Afrocarpus mannii, Melchiora mannii , Psychotria

guerkeana K.Schum., P. nubicola G. Taylor, and Peddiea

thomensis Exell occurring. Under the canopy species

of Begonia and Tristemma Juss., Anthocleista scandens

Hook.f., Pilea manniana Wedd. and P. rivularis Wedd.

are found in numbers. The presence of Erica thomen-

sis (Henriq.) Dorr & E.G.H.Oliv. and Lobelia barnsii

Exell at a higher altitude indicates a tendency towards

a mountain grassland. The area of Lagoa Amelia (at ±

1 430 m) has been well collected and its plant diversity

is well known. For example, this small area of ± 4 km2

has 55 taxa of pteridophytes. The Lagoa (= lake) is the

crater of an ancient volcano. It is now a bog surrounded

by tree ferns (Alsophila manniana (Hook.) R.M.Tryon)

and giant begonias ( Begonia crateris Exell). The bog

itself is populated by ferns such as Nephrolepis cordifo-

lia (L.) C.Presl var. pumicicola (F.Ballard) Hovenkamp

& Miyam. (typical of lava-derived soils), Lycopodiella

cernua (L.) Pic.-Serm. and Lepisorus excavatus (Bory

ex Willd.) Ching.

The vegetation of Principe is similar to that of

S.Tome, with a predominance of representatives of the

families Rubiaceae, Euphorbiaceae and Orchidaceae,

and a scarcity of Fabaceae taxa. Mangroves also occur

there, but savannas are absent. Being an island of much

lower altitude (948 m), the forest resembles the low alti-

tude forest in S.Tome, but the ridges at low altitude tend

to show some characteristics of formations that are only

found at much higher altitude in S.Tome. Submontane

forest is only recorded on the summit of the island, the

Pico do Principe.

Diversity

The flora of S.Tome and Principe has a high diversity

and endemism. This issue has been previously covered

in the literature (e.g. Figueiredo 1994b). The present

work updates the figures of flowering plant diversity

to: 135 families (29 introduced), 624 genera (172 intro-

duced), 1 104 species (301 introduced) plus 15 addi-

tional infraspecific taxa. At present, 119 taxa (107 spe-

cies and 12 infraspecific taxa) are known to be endemic

to the two islands.

Bothalia 41,1 (2011)

43

There is only one native gymnosperm in STP, Afro-

carpus mannii (Hook.) C.N.Page, a species endemic to

S.Tome which is widely planted as an ornamental tree.

However, it is rare in the wild where it occurs only at

high altitudes. The predominant families in the flora of

STP are the Orchidaceae, Rubiaceae and Euphorbiaceae.

Together they account for more than half the number

of endemic taxa. Polystachya Hook. (Orchidaceae) and

Psychotria L. (Rubiaceae) have the highest levels of

endemism. In the remaining families, the endemism of

taxa of the genus Begonia (Begoniaceae) is notewor-

thy, particularly that of the gigantic species B. baccata

Hook.f. and B. crateris, which can reach a height of 4

m. Endemic species of Impatiens L. (Balsaminaceae)

and Calvoa Hook.f. (Melastomataceae) are also note-

worthy for the ornamental value of their flowers. The

pteridophytes are particularly diverse and plentiful in the

flora of S.Tome and Principe, with 157 species and 13

endemics. This group has been recently revised (Figuei-

redo 1998, 2000, 2001, 2002a; Figueiredo & Gascoigne

2001).

MATERIAL AND METHODS

The collections cited in this catalogue are those of

the authors and other recent collectors such as J. Lejoly,

S.P. Carvalho, G. Joffroy and G.C. Matos. Only recent

collections (from 1990 or later) are cited. Most of the

recent collections from the islands are deposited at COI

(Paiva, Madureira & Martins), LISC (Carvalho, Figue-

iredo, Matos) and BRFU (Joffroy, Fejoly, Stevart). The

national herbarium in S.Tome (STPH) has duplicates or

the first set of many of the recent collections. Herbar-

ium abbreviations follow Thiers (continuously updated:

http://sweetgum.nybg.org/ih/).

The nomenclature follows Klopper et al. (2006)

and AFPD (2009). The synonyms cited in the list were

accepted by Exell (1973) or in recent literature.

Common names were mostly obtained from the lit-

erature (Rozeira 1958; Silva 1959; Santo 1969; Roseira

1984; Figueiredo 2002b). Medicinal uses were com-

piled from the literature (Sequeira 1994; Martins 2001;

Madureira 2006; Madureira et al. 2007) and herbarium

specimens.

FIST OF FFOWERING PFANTS

Accepted names are in bold, synonyms in italics. Common names appear in brackets after the taxon and author’s

name(s). Abbreviations/symbols used in the list are: *, introduced; ?, recorded in literature but specimens not known,

or doubts about their origin; §, record requiring confirmation; D, distribution in S.Tome and Principe (STP); E,

edible; E, endemic; H, habit and habitat; M, medicinal (see Appendix); N, notes; O, ornamental; P, Principe; ST,

S.Tome; V, voucher(s).

ACANTHACEAE

(Daniel & Figueiredo 2009)

ACANTHUS L.

montanus (Nees ) T. Anderson (Castanheiro-de-S.Tome, Cumbo-

-muala-ve, Cundu-de-muala-ve)

H: perennial herb with spiny leaves and white flowers, on roadsi-

des and plantations. M: 27; 171. D: STR V: Daniel 11183 (CAS,

STPH); Matos 7522 (BRLU, LISC).

ASYSTASIA Blume

gangetica (L.) T.Anderson ( Fia-mambleble , Fia-qui-sobo, Fia-qui-

-sobd-homem, Home, Muala )

H: perennial herb with white flowers dotted purple, on roadsides, cul-

tivated areas, plantations, ruderal. M: 40; 65; 79; 88; 124; 142. D:

STP. V: Figueiredo 207 (LISC); Matos 7734 (LISC).

BRACHYSTEPHANUS Nees

Eoccidentalis Lindau

H: herb or shrub with white flowers, in montane forest, secondary for-

est and margins of watercourses. D: ST. V: Paiva 208 (COI).

BRILLANTAISIA P.Beauv.

lamium (Nees) Benth. (Fia-de-mina-grande, Fia-fogo)

H: perennial herb with blue flowers, in forest, secondary forest and

plantations, ruderal. D: STP. V: Figueiredo 31 (LISC); Matos 7737

(LISC).

owariensis P.Beauv. ( Mamblembe )

H: herb with blue flowers in secondary forest and on roadsides. D: ST.

V: Figueiredo 213 (LISC).

vogeliana (Nees) Benth.

H: herb with white or blue flowers, in rainforest and secondary forest.

D: ST. V: Figueiredo 32 (K, LISC).

DICLIPTERA Juss.

verticillata (Forssk.) C.Chr.

H: herb with blue, purple flowers, in secondary forest, on roadsides. D:

STP. V: Matos 7575 (BRLU, LISC, STPH).

ELYTRARIA Michx.

marginata Vahl ( Focha-anzolo , Santage-basso-cafe)

H: herb with white or lilac flowers, in secondary forest, cultivated

areas and river margins. M: 16; 51; 59; 96; 173. D: STP. V: Oli-

veira 530 (BRLU, STPH), 1351 (BRLU).

HETERADELPHIA Lindau

Fpaulo\vilhelmia Lindau

H: perennial herb or shrub with purple flowers, in rainforest. D: ST. V:

Matos 7302 (BRLU, LISC).

HYPOESTES Sol. ex R.Br.

sp.

D: ST. N: Hypoestes sp. was recorded by Exell (1944) based on the

collection Don s.n. (BM) and said to be possibly a new species.

JUSTICIA L.

tenella (Nees) T.Anderson

H: herb with pink flowers or white with purple patch on lower lip

(Exell 1944), in secondary forest. D: STP. V: Matos 7393 (BRLU,

LISC, STPH); Oliveira 135/98 (STPH).

Ethomensis Lindau

H: herb. D: ST. N: rare or even extinct plant only known from the type

collection [Souza s.n. (COI)].

NELSONIA R.Br.

smithii Oerst.

H: herb with creeping stem sending up flowering shoots, flowers white

tinged with pink or crimson (Exell 1944), in secondary forest. D:

STP. V: Oliveira 504 (BRLU, LISC), 594 (BRLU, STPH).

44

Bothalia 41,1 (2011 )

PHAULOPSIS Willd.

micrantha (Benth.) C.B. Clarke ( Fia-glavana )

H: herb with white flowers. M: 84; 96; 150. D: ST. V: Oliveira 264

(STPH).

RHINACANTHUS Nees

virens (Nees) Milne-Redh.

H: herb with white flowers in secondary forest. D: P. V: Oliveira

133/98 (BRLU, LISC, STPH).

*RUELLIA L.

*brevifolia (Pohl) Ezcurra ( Mato-bana , Pega-pega)

H: herb or undershrub with red flowers, in secondary forest. D: ST. V:

Figueiredo 30 (LISC).

STENANDRIOPSIS S. Moore

thomensis (Milne-Redh.) Heine

H: herb with pink or blue flowers, in forest. D: STP. V: Lejoly 95/97

(BRLU, LIC, STPH); Oliveira 547 (BRLU, STPH).

*AGAVACEAE

* AGAVE L.

*sisalana Perrine (sisal)

H: succulent shrub. D: ST. V: Paiva 128 (COI).

?*FURCRAEA Vent.

?*foetida (L.) Haw.

N: recorded by Exell (1944) based on the literature; no specimens

known.

AIZOACEAE

SESUVIUM L.

portulacastrum (L.) L.

H: perennial herb occurring in coastal areas. E: leaves. D: ST.

ALANGIACEAE

ALANGIUM Lam.

chinense (Lour.) Harms

H: tree up to 22 m high, with white to yellowish flowers, in secondary

forest. D: ST.

7ALLIACEAE

7ALLIUM L

?*cepa L. (Cebola)

N: cultivated for the bulbs, recorded by Exell (1973) based on Silva

(1958) without specimen citation.

?*sativum L. ( Alho )

N: cultivated for the bulbs, recorded by Exell (1973) based on Silva

(1958) without specimen citation.

AMARANTHACEAE

*ACHYRANTHES L

*aspera L. ( Fia-ponto , Folha-galo, Folha-ponto, Mato-bana, Papue)

H: herb in plantations. M: 14; 41; 67; 60; 89; 90; 168. D: ST. V: Madu-

reira & Martins 86 (COI).

ALTERNANTHERA Forssk.

littoralis P.Beauv. var. maritima (Mart.) Pedersen

maritima (Mart.) A.St.-Hil.

H: succulent prostrate herb, in dunes. D: STP. V: Paiva 266 (COI).

*pungens Kunth ( Piancabla )

H: prostrate herb, on roadsides and cultivated ground. D: STP. V:

Paiva 1051 (COI).

*sessilis (L.) DC. ( Qua-qua-closso )

H: herb, in plantations and on roadsides, ruderal. M: 48; 54; 96; 124.

D: STP. V: Madureira & Martins 295 (COI); Paiva 159 (COI).

AMARANTHUS L.

*caudatus L. (Jimboa)

H: herb, in plantations. E: leaves, seeds. D: ST. V: Oliveira 201

(BRLU).

*dubius Mart, ex Thell.

H: herb, ruderal. E: leaves. D: ST. V: Paiva 263 (COI).

graecizans L. (Chimboa, Gimboa)

H: herb, on roadsides. E: leaves. D: ST. V: Paiva 394 (COI).

*hybridus L. subsp. cruentus (L.) Thell. ( Gimboa )

H: herb, in cultivated fields. E: leaves, seeds. D: ST. V: Paiva 1040

(COI).

*spinosus L. (Gimboa, Jimboa-brava)

H: herb. E: leaves. M: 173. D: STP.

*viridis L. (Belo-vieme, Gimboa-doida)

H: herb, on roadsides. E: leaves. M: 124; 173. D: ST. V: Madureira <£

Martins 282 (COI).

BLUTAPARON Raf.

vermiculare (L.) Mears var. vermiculare

H: perennial herb, in coastal areas. D: ST. V: Matos 7271 (LISC).

CELOSIA L.

*argentea L.

H: herb. E: leaves. O. D: ST.

leptostachya Benth.

H: annual herb, in plantations and on roadsides, ruderal. D: ST. V:

Paiva 160 (COI)

CYATHULA Blume

prostrata (L.) Blume var. pedicellata (C.B. Clarke) Cavaco (Fia-bana,

Mato-bana, Pegacalato)

H: small prostrate herb, ruderal. D: STP. V: Figueiredo 218 (LISC);

Paiva 450 (COI).

DEERINGIA R.Br.

sp.

D: P. N: Exell (1944, 1973) recorded Deeringia amaranthoides (Lam.)

Merr. as an introduced plant in Principe (based on Barter 2009

(K)). He cited D. celosioides R.Br., nom. illegit, as a synonym. In

the AFPD (2009) database, D. celosioides is accepted as a species

endemic to Principe.

*GOMPHRENA L.

*globosa L.

H: annual herb. O. D: ST.

*IRESINE P. Browne

*herbstii Lindl.

H: herb with red leaves, ruderal. D: ST. V: Lejoly 93/601 (BRLU).

PUPALIA Juss.

lappacea (L.) A. Juss. (Pega-lato, Pega-rato)

H: herb, on roadsides and cultivated fields, ruderal. M: 67. D: ST. V:

Madureira & Martins 275 (COI).

AMARYLLIDACEAE

CRINUM L.

jagus (J.Thomps.) Dandy (Cebola-cence, Sabola-cence)

H: bulbous plant with white flowers, in secondary forest and planta-

tions. M: 1 2; 33; 50; 66; 85; 1 86. D: ST. V: Paiva 458 (COI).

*HIPPEASTRUM Herb.

*puniceum (Lam.) Kuntze

H: bulbous plant with pink flowers. O. D: STP.

*HYMENOCALLIS Salisb.

*Iittoralis (Jacq.) Salisb. ( Flor-branca , Sabola-cence)

H: bulbous plant with white flowers. M: 12; 66; 98; 1 14; 191. D: P. V:

Madureira & Martins 648 (COI).

ANACARDIACEAE

*ANACARDIUM L.

*occidentale L. (fruit: Caji't; tree: Cajueiro)

H: tree, cultivated and occurring on roadsides. E: fruits, seeds. M: 2;

22; 23; 26; 37; 60; 67; 1 14; 171. D: STP. V: Paiva 109 (COI).

Bothalia 41,1 (2011)

45

LANNEA A. Rich.

welwitschii (Hiem) Engl. (Mucumbli)

H: tree, on secondary forest (300-1 500 m). M: 22; 57; 87; 109; 124.

D: ST. V: Madureira & Martins 217 (COI).

*MANGIFERA L.

*indica L. (fruit: Manga ; tree: Mangueira)

H: large tree with small red or yellow flowers, cultivated and occurring

on roadsides. E: fruits. M: 14; 27; 37; 40; 57; 67; 98; 177. D: STP.

V: Madureira & Martins 459 (COI); Paiva 821 (COI).

PSEUDOSPONDIAS Engl.

microcarpa (A. Rich.) Engl. (Guegue, Safu-d'obo, Safu-dodo, Zenze,

Zenzem)

H: tree with yellowish white flowers (Exell 1944), in secondary forest

and on roadsides. M: 62; 124; 157. D: STP. V: Madureira & Mar-

tins 512 (COI); Paiva 1915 (COI).

SORINDEIA Thouars

grandifolia Engl. (Gogo, Safii-de-obo )

H: tree, in rainforest and secondary forest. D: ST. V: Matos 7683

(LISC).

juglandifolia (A. Rich.) Planch, ex Oliv.

H: tree, in forest. D: ST. V: Lejoly 98/236 (BRLU).

*SPONDIAS L.

*cvtherea Sonn. (fruit: Cajamanga ; tree: Cajamangueira)

dulcis Sol. ex Parkinson

H: tree with small white flowers. E: fruits. D: STP. V: Paiva 799

(COI).

*mombin L. ( Geg , Gig, Guegue, Guegue)

lutea L.

H: small tree, in hedges. E: fruits. M: 14; 41; 54; 56; 75; 84; 122. D:

STP. V: Paiva 814 (COI).

ANISOPHYLLEACEAE

ANISOPHYLLEA R.Br. ex Sabine

Ecabole Henriq. (Cabole)

H: tree 35^30 m high (Exell 1944). D: ST. N: an undetermined species

of Anisophvllea has also been reported from Principe. V: Matos

7237a (LISC).

ANNONACEAE

ANNONA L.

*cherimola L. (Crimola)

H: small tree. E: fruits. D: ST.

glabra L. (Nona, Nona-concha, Nopa, Nopa-concha )

H: tree. E: fruits. D: ST.

*muricata L. ( Chabe-chabe , Chapo-chapo, Coraqao-da-lndia,

Coraqao-de-preto, Coraqao-preto, Sape-sape, Sapo-sapo)

H: tree. E: fruits. M: 94; 98; 123. D: STP. V: Paiva 1042 (COI).

^reticulata L. (Fruta-conde)

H: tree with reddish flowers, cultivated and occurring in secondary for-

est. E: fruits. D: STP. V: Oliveira 225/98 (BRLU), 1435 (BRLU).

senegalensis Pers.

arenaria Thonn.

H: shrub or small tree. D: ST? N: recorded by Exell (1944, 1973) as A.

arenaria Thonn., based on Don s.n. (BM).

*squamosa L. ( Fruta-pinha , Nona)

H: tree. E: fruits. M: 150. D: ST. V: Madureira & Martins 435 (COI).

?*ASIMINA Adans.

?*triloba (L.) Dunal

D: P? N: a doubtful record from the literature according to Exell

(1944, 1973); no specimens known.

*CANANGA (DC.) Hook.f. & Thomson

*odorata (Lam.) Hook.f. & Thomson (Ilangue-ilangue. Pompeia)

H: tree with strongly fragrant flowers. O. M: 138. D: ST. V: Figueiredo

123 (K).

MONODORA Dunal

brevipes Benth. (lobo)

H: tree, in plantations. D: STP. V: Paiva 822 (COI).

myristica (Gaertn.) Dunal (lobo, lobo, Jobo, Xipobo, Yobo)

H: tree, in plantations. M: 2; 20; 57; 98; 99; 146. D: STP. V: Madu-

reira & Martins 76 (COI), 455 (COI).

POLYALTHIA Blume

oliveri Engl. (Cabore-de-Bom-Sucesso, Inhe-preto, Pau-preto, Preto)

H: tall tree with white flowers, in rainforest. M: 20; 22. D: STP. V:

Madureira & Martins 589 (COI); Oliveira 588 (BRLU).

UVARIA L

ovata (DC.) A. DC. subsp. ovata

sp. sensu auctt. Santo (1970), Exell (1973).

H: shrub with yellow flowers forming thickets, in savanna. D: ST. V:

Espirito Santo 4357 (LISC).

XYLOPIA L.

aethiopica (Dunal) A. Rich. (Cabela, Inhe-bobo, Noe, Pimenta-do-

-Congo, Untue-do-bo)

H: tree ± 20 m high with fragrant white flowers (Exell 1944), in sec-

ondary forest. E: fruits. M: 22; 43; 62. D: STP. V: Joffroy 147

(BRLU); Paiva 1473 (COI).

quintasii Engl. & Diels

H: tree up to 35 m high, in rainforest. D: ST.

staudtii Engl. & Diels (Jnhe-bobe, Inhe-branco, Inhe-ganso, Unue-

-bolina)

africana (Benth.) Oliv.

H: tree 9-30 m high, in plantations, secondary forest and on roadsides.

D: STP. V: Paiva 943 (COI), 1419 (COI).'

APIACEAE

APIUM L

leptophvllum (Pers.) F.Muell. ex Benth.

H: annual herb. D: P.

CENTELLA L

asiatica (L.) Urb. (Fia-d’olha, Folha-de-orelha, Folha-viola,

Olhadato, Ufia-copo)

H: perennial herb with red flowers, creeping and sometimes climbing

(Exell 1944). M: 29; 44; 69; 155; 174. D: STP. V: Figueiredo 115

(K).

*ERYNGIUM L.

*foetidum L. (Celo-sum-zom-maid, Cheiro-de-Joao-Maria, Coentro-

-de-S.Tome, Jamaia, Lamaia, Selo-sum-Joao-Maia, Selo-sum-

-zon-maia)

H: biennial herb. E: leaves. M: 2; 52; 61; 81; 94; 145; 150; 156. D:

STP. V: Paiva 634 (COI), 1149 (COI).

HYDROCOTYLE L.

bonariensis Lam. (Fia-copo, Fid-d'ollta, Home-d'olha)

H: perennial herb. M: 2; 44; 69; 155; 1 74. D: STP. V: Paiva 249 (COI),

1436 (COI).

mannii Hook.f.

H: prostrate herb trailing along the ground (Exell 1944). D: ST. V:

Matos 7428 (BRLU, LISC).

*PETROSELINUM Hill

*crispum (Mill.) A.W.Hill (Salsa)

H: biennial herb. E: leaves. D: ST.

APOCYNACEAE

*ASCLEPIAS L

*curassavica L.

H: herb, in gardens and escaped. O. D: ST. V: Figueiredo 60 (K).

*CASCABELA Raf.

*thevetia (L.) Lippold

Thevetia peruviana (Pers.) K.Schum.

H: shrub, with yellow flowers, ruderal. D: ST. V: Paiva 1297 (COI).

*CATHARANTHUS G.Don

*roseus (L.) G.Don (Labios-de-mulata, Vinca)

H: shrub or perennial herb, with white flowers with a red or pink cen-

tre (Exell 1944), ruderal. M: 118; 171. O. D: ST. V: Paiva 433

(COI).

FUNTUMIA Stapf

africana (Benth.) Stapf (Pau-cadeira, Po-cadela, Po-leite, Po-visgo)

H: tree 45-60 m high (Exell 1944), with yellow-white flowers, in for-

est. M: 56; 115; 176. D: STP. V: Oliveira 88 (BRLU).

46

Bothalia 41,1 (2011)

FUNTUMIA Stapf (cont.)

elastica (P.Preuss) Stapf (Funtumia)

H: tree, with yellow-white flowers, in forest. D: ST.

GONGRONEMA Decne.

latifolium Benth. ( Olage )

H: climber with tuberous base. D: ST.

LANDOLPHIA P.Beauv.

landolphioides (Hallier f.) A.Chev.

dawei Stapf

H: liana with a trunk up to 0.4 m in diameter. E: fruits. D: ST. V: Oli-

veira 51/99 (BRLU).

MARSDENIA R.Br.

'exellii C.Norman

H: liana with yellow flowers, in forest. D: ST. V: Lejoly 98/269

(BRLU).

*NERIUM L.

"“oleander L.

H: shrub. O. D: ST. V: Oliveira 217 (BRLU).

ONCOSTEMMA K.Schum.

cuspidatum K.Schum.

D: ST. N: this monotypic genus has been reduced to synonymy of

Tylophora by Meve & Omlor (2000) and found to be a synonym

of T. oblonga N.E.Br. The new combination Tylophora cuspi-

data (K.Schum.) Meve & Omlor, said to take precedence over T.

oblonga, is illegitimate.

"“PLUMERIA L.

"“rubra L.

H: shrub or tree, very fragrant. O. D: ST.

RAUVOLFIA L.

caffra Sond. (Cata-cuene, Cata-glandje, Caia-grande. Note: not to be

confused with Cata-pequena)

macrophylla Stapf

H: tree, with white flowers and blackish purple fruits, in forest. M: 2;

25; 44; 79; 99; 123; 127; 146; 155; 168; 175. D: ST. V: Madureira

& Martins 16 (COI).

vomitoria Afzel. ( Cata , Cata-manginga, Cata-pequena, Cata-piquina,

Cata-ussubi)

dichotoma K.Schum.

H: shrub or small tree with white flowers and red fruits, in forest and

plantations. M: 60; 79; 99; 103; 123; 175; 184. D: STP. V: Figuei-

redo 256 (LISC).

TABERNAEMONTANA L

*divaricata (L.) R.Br. ex Roem. & Schult.

Ervatamia coronaria (Jacq.) Stapf

E. divaricata (L.) Burkill

H: shrub or small tree with white flowers, in gardens. D: STP. V: Paiva

290 (COI).

pachysiphon Stapf (Cata-glandje, Cata-grande)

H: shrub or small tree, with white or yellowish flowers, in forest. D:

ST. V: Oliveira 1430 (BRLU).

Estenosiphon Stapf (Cato, Cata-d'obo, Cata-piquina, Pau-llrio )

H: small tree with fragrant white flowers, in forest. M: 8; 77; 103. D:

STP. Oliveira 141/98 (LISC); Paiva 76 (COI).

VOACANGA Thouars

africana Stapf (Buie, Cata-contra, Cata-glandje, Cata-grande, Cata-

-quio, Gata-grande, Guibule )

lemosii Philipson

H: small tree, with yellow or white flowers, in forest. M: 8; 103. D:

ST. V: Paiva 1194 (COI).

AQUIFOLIACEAE

ILEX L.

mitis (L.) Radik.

H: tree, with white flowers, in forest. D: ST. V: Oliveira 985 (BRLU).

ARACEAE

*ALOCASIA (Schott) G.Don

*macrorrhiza (L.) G.Don (Coco-veneno, Matabala-veneno)

H: perennial herb with a tuberous rootstock in secondary forest and

plantations. O. M: 5; 133; 174. D: STP. V: Madureira & Martins

449 (COI).

*CALADIUM Vent.

*bicolor (Aiton) Vent. (Coco-floli, Coco-veneno )

H: perennial herb with a tuberous rootstock, in secondary forest and

plantations. O. D: STP. V: Paiva 1200 (COI), 1427 (COI).

"“COLOCASIA Schott

"“esculenta (L.) Schott (Coco, Matabala, Micoco)

H: large herbaceous plant cultivated for the corm (taro). D: STP.

CULCASIA P.Beauv.

angolensis Welw. ex Schott (Pimenta-da-terra)

barombensis N.E.Br.

H: climber, in forest. D: ST.

scandens P.Beauv. ( Home-dum-osso , Homem-de-um-osso-so)

H: epiphyte, climbing herb, in forest. M: 22; 87; 96. D: ST. V: Madu-

reira & Martins 66 (COI).

"“DIEFFENBACHIA Schott

*seguine (Jacq.) Schott

H: herb. O. D: ST. V: Oliveira 216 (BRLU).

*XANTHOSOMA Schott

"“sagittifolium (L.) Schott (Coco, Matabala, Matabala-branca,

Matabala-gema-de-ovo, Matabala-vermelha, Micoco )

maffafa Schott

violaceum Schott

H: large herbaceous plant much cultivated for the corm (coco-yam).

M: 188. D: STP. V: Paiva 761 (COI), 1331 (COI).

ARALIACEAE

"“HEDERA L.

"“helix L. (Hera)

H: climber. O. D: ST. N: recorded by Exell (1944, 1973) but it is pos-

sible that it may be a different species of Hedera.

POLYSCIAS J.R.Forst. & G.Forst.

Equintasii Exell (Guegue-fasso, Veld)

H: tree, in forest. D: STP. V: Oliveira 1253 (BRLU).

SCHEFFLERA J.R.Forst. & G.Forst.

barteri (Seem.) Harms

H: liana, in secondary forest. D: STP. V: Figueiredo 48 (K).

mannii (Hook.f.) Harms

H: small shrub or tree, in forest. D: ST. V: Matos 7513 (LISC).

ARECACEAE

Note: the following palm trees have also been recorded in cultivation:

Roystonea regia (Kunth) O.F.Cook, Butia eriospatha (Mart, ex

Drude) Becc., Caiyota urens L.

BORASSUS L.

aethiopum Mart. ( Palmeira-do-marfim , Palmeira-leque, Ulua)

H: tree up to 20 m high. D: ST.

"“COCOS L.

"“nucifera L. (Coco, Coconjd, Conconja, Coqueiro)

H: tree. E: fruit. D: STP.

ELAEIS Jacq.

guineensis Jacq. (Andim, Dendem, Dendem, Palmeira-andim,

Palmeira-dendim, Palmeira-do-dendem, Pema)

H: tree up to 30 m high. E: fruit (oil). D: STP.

"“PHOENIX L.

"“dactylifera L. (fruit: Tamara ; tree: Tamareira)

H: tree with trunk up to 20 m high. E: fruit. D: ST.

*ARISTOLOCHIACEAE

"“ARISTOLOCHIA L.

"“littoralis Parodi

H: climber, yellow flowers spotted with pink (Exell 1956). O. D: ST.

Bothalia 41,1 (2011)

47

ASPHODELACEAE

*ALOE L.

*humilis (L.) Mill. (Aliba-blabosa, Erva-babosa )

H: scapose perennial herb. M: 39; 51; 54; 49; 60; 72; 77; 93; 123; 124.

D: ST. V; Madureira & Martins 354 (COI).

ASPARAGACEAE

* ASPARAGUS L.

^officinalis L. (Espargo)

N: herb cultivated for the young shoots, recorded by Exell (1973)

based on Silva (1958) without specimen citation.

ASTERACEAE

*ACANTHOSPERMUM Schrank

*hispidum DC. ( Mosquito-dia )

H: annual herb with yellowish green flowers (Exell 1944). D: STP.

ACMELLA Rich, ex Pers.

caulirhiza Delile (Fia-dentche, Fia-dente)

Spilanthes filicaulis (Schumach. & Thonn.) C.D. Adams

H: herb with yellow flowers. M: 132; 175. D: STP. V; Madureira &

Martins 296 (COI); Matos 7491 (LISC).

ADENOSTEMMA J.R.Forst. & G.Forst.

mauritianum DC.

FI: herb with white flowers, in forest and secondary forest. D: ST. V:

Lejoly 94/551 (BRLU, LISC).

perrottetii DC. (Lingua-tela, Tonfonso)

H: herb with white flowers, in forest, plantations and on roadsides. M:

141. D: ST. V; Madureira 646 (COI).

*AGERATUM L.

*convzoides L. subsp. conyzoides (Fia-cuncunha, Fia-male, Fia-

-male-di-que, Fia-male-home, Folha-male)

H: annual herb with blue-purple or white flowers, in secondary for-

est, plantations and on roadsides, ruderal. M: 124; 150. D: STP. V:

Paiva 1385 (COI), 1782 (COI).

AMBROSIA L.

maritima L. ( Atinija , Atlija, Atlimija, Atrimija, Natruja)

H: annual herb or perennial from woody rootstock. M: 29; 77; 115;

150; 184; 189. D: STP. V; Madureira & Martins 93 (COI).

7ASPILIA Thouars

?rudis Oliv. & Hiem subsp. rudis

H: erect, often perennial herb with pale yellow or almost white flow-

ers. D: P. Recorded by Exell (1944) based on a citation in Flora of

tropical Africa of a collection by Barter that he did not see.

BIDENS L.

pilosa L. ( Aliba-giihia , Fia-giihia, Pega-pega)

H: herb with white flowers, in secondary forest, plantations and on

roadsides. M: 29; 46; 69; 88. D: STP. V: Paiva 572 (COI), 705

(COI).

^CHRYSANTHEMUM L.

*indicum L.

H: perennial herb. O. D: ST.

*?CICHORIUM L.

*?intvbus L.

H: perennial herb. E: leaves (var. foliosum), roots (var. sativum). D;

ST. N: a doubtful record based on a specimen collected by Don and

cited by Bentham in Niger flora. Exell (1944) did not see the speci-

men.

CONYZA Less.

attenuata DC. (Folha-male, Lingua-de-vaca, Lingua-tela)

persicifolia (Benth.) Oliv. & Hiem

H: annual herb, in forest. D: ST. V; Matos 7369 (LISC).

*bonariensis (L.) Cronquist

H: annual erect herb. D: STP.

CRASSOCEPHALUM Moench

crepidioides (Benth.) S. Moore

H: annual herb with yellow or mauve flowers, in forest and planta-

tions. D: ST. V: Lejoly 98/211 (BRLU).

montuosum (S. Moore) Milne-Redh.

H; herb, ruderal. D: ST. V: Matos 7438 (LISC).

rubens (Juss. ex Jacq.) S. Moore

H: herb, in secondary forest and plantations. D: ST.

sarcobasis (DC.) S.Moore

H: herb, in disturbed ground. D: ST. V: Matos 7466 (BRLU).

DICHROCEPHALA L’Her. ex DC.

integrifolia (L.f.) Kuntze subsp. integrifolia

H: annual herb with whitish flowers, ruderal. D: ST. V: Lejoly 95/057

(LISC).

ECLIPTA L.

prostrata (L.) L. (Folha-boba, Folha-sapateiro)

H: annual or perennial herb with white flowers. D: STP

*ELEPHANTOPUS L

*mollis Kunth (Fia-budo, Fia-dentche, Folha-budo, Macabali,

Macablali)

H: herb with white flowers, in secondary forest, plantations, and

on roadsides. H: 3; 22; 112; 129; 130; 150; 174; 175. D: STP. V:

Figueiredo 105 ( K).

EMILIA Cass.

?coccinea (Sims) G.Don

N: recorded by Exell (1973) based on Silva (1958) without specimen

citation.

sonchifolia (L.) DC. ex Wight

H; annual herb. E: leaves. D: STP.

§EPALTES Cass.

§*brasiliensis DC.

H; herb. D: STP. N: Exell (1944) cited three collections of this taxon

( Barter 1976, Mann s.n. and Welwitsch 3520). We did not see these

specimens but it is possible that they refer to Litogyne gariepina

(DC.) Anderb., a species widespread from West Africa to South and

East Africa that includes as synonyms the three species of Epaltes

previously cited for tropical Africa. E. brasiliensis has apparently

never been recorded for Africa.

ETHULIA L.f.

conyzoides L.f. subsp. conyzoides

H: annual herb, with white, mauve or purple flowers. D: ST.

*GALINSOGA Ruiz & Pav.

*parviflora Cav.

H: annual herb with white flowers, on disturbed ground and on road-

sides. D: ST. V: Matos 7603 (LISC).

HELICHRV SUM Mill,

foetidum (L.) Moench var. foetidum

H: annual or perennial herb, with yellow flowers, in forest. D: ST. V;

Matos 7371 (LISC).

MICROGLOSSA DC.

pyrifolia (Lam.) Kuntze

H: shrub or scandent shrub, with white, cream-coloured, yellowish or

pinkish flowers, in forest. D; STP. V: Matos 7564 (LISC).

MIKANIA Willd.

chenopodiifolia Willd. (Matucana)

cordata sensu auctt. non (Burm.f.) B.L.Rob.

sp. sensu auct. Exell (1956, 1973)

H: shrub with white flowers. M: 31; 77; 98; 129; 140. D: STP. V:

Figueiredo 257 (LISC).

MIKANIOPSIS Milne-Redh.

paniculata Milne-Redh.

H: perennial climber with orange-yellow flowers. D: ST. V: Matos

7312 (LISC).

PSEUDOGNAPHALIUM Kirp.

luteo-album (L.) Hilliard & B.L.Burtt subsp. luteo-album

Gnaphalium luteo-album L.

H: annual or perennial herb with whitish or yellowish flowers, on

roadsides, ruderal. D: STP. V: Paiva 1064 (COI).

48

Bothalia 41,1 (2011)

SOLANECIO (Sch.Bip.) Walp.

biafrae (Oliv. & Hiem) C.Jeffrey

Crassocephalum biafrae (Oliv. & Hiem) S.Moore

H: perennial, climbing, subsucculent herb with yellow flowers. E:

leaves. D: ST.

SONCHUS L.

oleraceus L. ( Serralha )

H: annual or biennial herb with yellow flowers, in forest, plantations

and on roadsides. E: leaves. D: ST. V: Paiva 33 (COI).

STRUCHIUM P. Browne

sparganophora (L.) Kuntze ( Libo-d’agua , Libd-d'aua )

Sparganophorus sparganophora (L.) C.Jeffrey, nom. illegit.

H: perennial, slightly succulent herb, with whitish or pinkish flowers,

in humid areas. M: 123; 137; 190. D: STP. V: Paiva 1292 (COI).

*SYNEDRELLA Gaertn.

*nodiflora Gaertn. ( Fia-male-dodo , Fia-male-muala, Folha-male-

-doida, Macabali-fia-budo )

H: annual or perennial herb with yellow flowers, in secondary forest,

plantations, and on roadsides, ruderal. M: 27; 51; 52; 81; 94; 113;

123; 156. D: STP. V: Paiva 1006 (COI), 1784 (COI).

*TITHONIA Desf. ex Juss.

*diversifolia (Hemsl.) A. Gray ( Girassol )

H: annual or perennial herb or subshrub up to 2.5 m high, with yellow

flowers, ruderal. M: 123. D: ST. V: Paiva 1281 (COI).

VERNONIA Schreb.

amygdalina Delile (Libo, Libo-da-terra, Libo-de-que, Libo-mucambu,

Limbo, Pau-fede )

H: undershrub, shrub or tree, with white flowers, ruderal. E: leaves. M:

60; 123. D: ST. V: Paiva 1289 (COI).

stellulifera (Benth.) C.Jeffrey ( Fia-bunga )

Triplotaxis stellulifera (Benth.) Hutch.

H: annual herb with mauve to purple flowers, in secondary forest. D:

STP V: Figueiredo 41 (K).

*ZINNIA L.

*peruviana (L.) L.

H: annual herb, with red, pink or purplish flowers. D: ST.

*AVERRHOACEAE

*AVERRHOA L.

*carambola L. (fruit: Carambola\ tree: Caramboleira )

H: shrub or small tree, cultivated. E: fruits. D: ST. V: Paiva 1327

(COI).

AVICENNIACEAE

(Daniel & Figueiredo 2009)

Avicennia L.

germinans (L.) L.

H: shrub or tree up to 17 m high, with pneumatophores, halophyte, in

mangroves. E: fruits. D: ST. V: Figueiredo 93 (K).

BALSAMINACEAE

IMPATIENS L.

*balsamina L. ( Balsamina , Melindres )

tarns iana Exel!

H: herb. O. D: STP. V: Paiva 1583 (COI), 1690 (COI).

' buccinalis Hook.f. ( Camardes )

H: herb or shrub up to 3 m high, with red flowers, in forest or second-

ary forest. D: ST. V: Paiva 1858 (COI).

Emanteroana Exell

H: herb with red flowers in humid and rocky places, in rainforest. D: P

V: Paiva 1726 (COI).

Rthomensis Exell

H: herb up to 1 m high, with red or orange flowers, in humid places, in

forest. D: ST. V: Figueiredo 255 (LISC).

*BASELLACEAE

*BASELLA L.

*alba L. ( Fia-tataluga , Folha-tataruga)

H: annual or perennial succulent herb. E: leaves. M: 173. D: ST. V:

Madureira & Martins 339 (COI).

BEGONIACEAE

BEGONIA L.

ampla Hook.f. ( Fia-boba-d’obo , Fia-boba-glandji )

H: perennial herb, epiphyte, with white flowers, in forest and second-

ary forest. M: 1; 16; 119; 129. D: STP V: Figueiredo 72 (K).

annobonensis A. DC.

H: annual epiphytic succulent herb, in forest. D: STP V: Figueiredo

260 (LISC).

Ebaccata Hook.f. (Fia-boba, Fia-bdba-d'obo, Folha-boba-vermelha)

H: herb, in forest. M: 16; 38; 84; 142. D: ST. V: Figueiredo 39 (K,

LISC).

Ecrateris Exell ( Fia-bdba-d’obo )

H: herb, in forest. D: ST. N: considered by some authors as a synonym

of 5. baccata. V: Lejoly 94/559 (BRLU, LISC).

Efusialata Warb. var. principensis J.J.de Wilde

H: herb. D: P.

Eloranthoides Hook.f. subsp. loranthoides

H: herb. D: STP

Emolleri (C.DC.) Warb.

H: herb. D: ST.

rostrata Welw. ex Hook.f. var. rostrata

H: annual succulent herb. D: ST. V: Figueiredo 161 (LISC).

subalpestris A.Chev.

macambrarensis Exell

H: herb, in forest. D: ST. V: Matos 7552 (BRLU, LISC).

thomeana C.DC.

H: perennial herb, in forest. D: ST. V: Matos 7704 (LISC).

BIGNONIACEAE

*JACARANDA Juss.

*mimosifolia D.Don ( Jacaranda )

H: tree with purple flowers. O. D: ST. N: recorded in the literature as

introduced, there is a single tree known (at Ropa Agostinho Neto).

NEWBOULDIA Seem

laevis (PBeauv.) Seem, ex Bureau (Quime, Quine)

H: shrub or tree, with pink flowers, ruderal. M: 28; 57. D: ST. V: Paiva

674 (C OI).

SPATHODEA PBeauv.

campanulata P.Beauv.

nilotica Seem.

H: tree. O. D: ST.

*TECOMA Juss.

*sp. ( Maioba-bini )

H: vine. O. M: 7; 96; 128; 146; 153. D: ST. V: Madureira & Martins

308 (COI).

*BIXACEAE

*BIXA L.

*orellana L. (Anato, Giclo, Ginclo, Quisafu, Tintureira, Urucu)

H: shrub or small tree with pink flowers. E: seeds. O. M: 22; 27; 63;

67. N: seeds contain a natural orange-red dye. D: STP V: Paiva

493 (COI).

BOMBACACEAE

ADANSONIA L.

digitata L. ( Imbondeiro , Micondo)

H: large tree in savanna. E: leaves, fruits, seeds. D: STP. V: Paiva 1238

(COI).

CEIBA Mill.

pentandra (L.) Gaertn. {Oca)

Bothalia 41,1 (2011)

49

H: large tree in secondary forest. E: seeds. M: 6; 124. D: STP. V:

Oliveira 139 (BRLU).

*PACHIRA Aubl.

*glabra Pasq.

Bombacopsis glabra (Pasq.) A. Robyns

H: tree, in secondary forest and plantations. E: seeds. D: STP. V: Paiva

1237 (COl), 1950 (COI).

BORAGINACEAE

CORDIA L.

*gerascanthus L.

H: tree up to 25 m high, with white flowers, in town. D: ST.

platythyrsa Baker ( Pau-tabaque , Pd-tabaque, Tabaque)

H: tree up to 18 m high, with creamy white flowers, in secondary for-

est. D: ST. V: Lejoly 93/641 (BRLU).

EHRETIA P.Browne

cymosa Thonn. var. zenkeri (Giirke ex Baker & C.H. Wright) Brenan

zenkeri Giirke ex Baker & C.H. Wright

H: shrub or small tree up to 12 m high, with white flowers and yellow

fruits, in forest. D: ST. V: Lejoly 97/344 (BRLU).

Escrobiculata Hiem

H: small tree with whitish flowers. N: probably extinct. D: P.

*HELIOTROPIUM L.

*indicum L. ( Fia-galo , Folha-galo, Galo)

H: annual herb with red flowers, ruderal. M: 41; 51; 57; 70; 77; 89; 90;

112; 183; 184. D: STP. V: Paiva 1290 (COI), 1413 (COI).

BRASSICACEAE

*BRASSICA L.

*juncea (L.) Czem. & Coss. (Mostarda)

H: perennial herb with yellow flowers. E: seed (spice: mostarda). D:

ST.

*CAPSELLA Medik.

*bursa-pastoris (L.) Medik.

H: annual herb, ruderal. D: ST.

CARD AMINE L

africana L.

H: annual herb with white flowers, in forest and secondary forest. D:

ST. V: Lejoly 97/220 (BRLU).

?hirsuta L.

N: a doubtful record in Exell (1973) probably due to a typographic

error; the references cited refer to the occurrence in Bioko and not

in S.Tome.

*DIPLOTAXIS DC.

*tenuisiliqua Delile

H: herb. D: ST. V: Matos 7300 (BRLU).

*LEPIDIUM L.

*didymum L.

Coronopus didymus (L.) Sm.

H: annual or biennial herb. E: leaves. D: ST.

*LOBULARIA Desv.

*maritima (L.) Desv.

H: annual or perennial herb, with white, pink or purple flowers. D: ST.

*RAPHANUS L.

*sativus L. (Nabo-macau)

H: annual or biennial herb with white or pink flowers. E: roots. D: ST.

RORIPPA Scop.

micrantha (Roth) Jonsell

indica sensu auct. non (L.) Hiem

H: annual herb with yellow flowers. D: ST.

*nasturtium-aquaticum (L.) Hayek (Agriao. Fia-guinhom, Fia-

-guiom. Guiom)

H: aquatic or semi-aquatic perennial herb with white flowers. E:

leaves, stems. M: 27; 177. D: ST. V: Madureira & Martins 617

(COI).

*BROMELIACEAE

* ANANAS Mill.

*co mosus (L.) Merr. ( Ananas , Nanas)

H: herb cultivated for fruit. M: 24; 98; 124. D: STP. V: Madureira &

Martins 647 (COI).

BUDDLEJACEAE

LACHNOPYLIS Hochst.

thomcnsis Philipson

Nuxia congesta R.Br. ex Fresen. var. thomensis (Philipson) J. Lewis

H: shrub or small tree, in forest. D: ST. V: Matos 7384 (LISC).

BURMANNIACEAE

GYMNOSIPHON Blume

sp.

H: colourless saprophyte, in forest. D: P. V: Stevart 264 (BRLU).

BURSERACEAE

DACRYODES Vahl

edulis (G.Don) H.J.Lam (fruit: Mubafo, Safit. Safu. Safu-safip tree:

Safueiro, Safuzeiro)

H: tree, in forest and plantations, producing a blue-violet fruit. E:

fruits. M: 185. D: STP. V: Paiva 521 (COI).

SANTIRIA Blume

trimera (Oliv.) Aubrev. ( Balsamo-de-S.Tome , Belambo, Goqui. Oleo-

-melambo, Pau-oleo, Po-oleo, Po-oleo-d'obo)

H: tree, in forest. M: 4; 22; 31; 51; 54; 57; 124; 177; 184;. D: STP. V:

Madureira & Marlins 249 (COI).

*?BUXACEAE

*?BUXUS L.

*?sempervirens L.

N: a doubtful record by Exell (1944, 1973) based on a citation by

Chevalier without specimens.

CACTACEAE

RHIPSALIS Gaertn.

baccifera (J.S.Muell.) Steam subsp. baccifera

H: hanging epiphytic, succulent shrub. D: STP. V: Paiva 1173 (COI).

CAMPANULACEAE

VVAHLENBERGIA Schrad. ex Roth

silenoides Hochst. ex A. Rich.

H: herb. D: ST. V: Matos 7563 (LISC).

*CANNABACEAE

*CANNABIS L

*sativa L. (Liamba)

H: annual herb. E: seeds (oil). D: ST.

*CANNACEAE

*CANNA L.

*indica L. ( Cana-de-jardim , Cana-florifera , Erva-conteira, Salaconta,

Salaconta-blanca, Salaconta-pleta )

H: rhizomatous perennial herb with red flowers. O. M: 45; 67; 69; 79;

98; 128; 153. D: STP. V: Paiva 364 (COI).

CAPPARACEAE

BUCHHOLZIA Engl,

coriacea Engl. ( Cola-congo )

H: tree up to 12 m high, in forest and secondary forest. M: 6; 173; 174.

D: ST. V: Paiva 1271 (COI).

50

Bothalia 41,1 (2011)

CAPPARIS L.

erythrocarpos Isert

H: shrub or climber, in savanna. D: ST. V: Lejoly 93/654 (BRLU).

toinentosa Lam.

H: shrub or climber, in savanna. D: ST. V: Figueiredo 83 (K).

CLEOME L.

gynandra L.

Gynandropsis gynandra (L.) Briq.

H: annual herb. D: ST.

rutidosperma DC. ( Fia-stlela , Mussanfi)

H: annual herb. M: 61; 89. D: STP. V: Madureira & Martins 342

(COi).

*CAPRIFOLIACEAE

*SAMBUCUS L.

*mexicana C.Presl ex DC.

H: shrub or tree with white flowers. D: ST. V: Matos 7520 (BRLU).

*CARICACEAE

*CARICA L

*papaya L. (fruit: Mamom, Papaia\ tree: Mamoeiro, Papaeira)

H: small tree. E: fruits. M: 32; 98; 114; 135; 146. D: STP. V:

Madureira & Martins 29 (COI).

CARYOPHYLLACEAE

DRYMARIA Willd. ex Schult.

cordata (L.) Willd. ex Roem. & Schult. (Chile-branco, Pega-pega,

Tchile-blanco)

H: perennial prostrate herb with white Bowers, in secondary forest,

plantations, and on roadsides, ruderal. M: 36; 53; 57; 174. D: STP.

V: Paiva 1022 (COI).

STELLARIA L.

mannii Hook.f.

H: annual or perennial herb, in forest. D: ST. V: Paiva 84 (COI).

*media (L.) Vill. subsp. media

H: annual herb. D: ST.

*CASUARINACEAE

*CASUARINA L.

*equisetifolia L.

H: tree. D: ST. V: Oliveira 236 (BRLU).

CELASTRACEAE

HELICTONEMA Pierre

velutinum (Afzel.) Pierre ex N. Halle

Hippocratea velutina Afzel.

H: liana with white flowers, in forest. D: ST. V: Lejoly 97/297 (LISC).

MAYTENUS Molina

Emonodii Exell

N: rare species, collected only once. D: ST.

SALACIA L.

pyriforniis (Sabine) Steudel

H: climber, in forest. D: ST.

*CHENOPODIACEAE

♦CHENOPODIUM L.

*album L. subsp. album

H: annual herb. E: leaves. D: ST.

*ambrosioides L. (Erva-de-santa-maria, Fia-madre-matrufo, Fid-

-matluQO, Fia-santa-maia, Folha-santa-maria, Macluso, Mastrugo,

Masturgo, Matlugo, Matlusso, Matrugo, Milongo-de-febres)

II: annual, rarely perennial herb, ruderal. E: leaves, seeds. M: 2; 61;

89; 95; 1 16; 124; 135; 156. D: STP. V: Paiva 1287 (COI).

7CHRYSOBALANACEAE

7NEOCARYA (DC.) Prance ex F. White

?macrophylla (Sabine) Prance ex F. White

Parinari macrophylla Sabine

N: doubtful record (Exell 1944, 1973) based on a collection by Don

which may have originated from the continent.

7PARINARI Aubl.

?curatellifolia Planch, ex Benth.

mobola Oliv.

N: doubtful record (Exell 1973) based on a citation by Silva (1958)

without specimens.

?excelsa Sabine

N: doubtful record (Exell 1944, 1973) based on a collection by Don

which may have originated from the continent.

CLUSIACEAE

GARCINIA L.

*mangostana L. (Mangostao)

H: tree. E: fruits. D: ST. V: Paiva 764 (COI).

?smeathmannii (Planch. & Triana) Oliv.

polyantha Oliv.

N: doubtful record (Exell 1944, 1973) based on a collection by Barter

that may not be from the islands.

MAMMEA L.

africana Sabine (Amargoso, Magloso, Mamdo, Mata-passo, Oba,

Pau-ova, Po-ova)

H: tree, in forest. E: fruits. M: 112. D: STP. V: Madureira & Martins

444 (COI).

PENTADESMA Sabine

butyracea Sabine (Mata-passo, Oba, Po-ova)

H: tree up to 35 m high, in forest. D: ST.

*PLATONIA Mart

*esculenta (Arruda) Oken ( Bacuri , Bacurizeiro)

H: tree. E: fruits. D: ST.

SYMPHONIA L.f.

globulifera L.f. (Oleo-balom, Oleo-barao)

H: tree up to 40 m high, in forest. M: 5; 133. D: STP. V: Matos 7524

(BRLU, LISC).

COMBRETACEAE

COMBRETUM Loell.

*indicum (L.) DeFilipps

Quisqualis indica L.

H: climbing shrub with white, pink or red flowers. D: ST.

paniculatum Vent.

H: liana with red flowers, in secondary forest. D: P. V: Oliveira 206/98

(BRLU).

CONOCARPUS L

erectus L.

H: mangrove shrub or tree. D: ST. V: Oliveira 38 (BRLU).

*TERMINALIA L.

*catappa L. (Amendoeira-da-lndia, Caroceiro, Carogo )

H: tree, in streets and on roadsides. M: 67. D: STP. V: Paiva 286

(COI), 1343 (COI).

COM MELINACEAE

ANEILEMA R.Br.

beniniense (P.Beauv.) Kunth

FI: straggling herb. D: P.

COMMELINA L.

congesta C.B. Clarke ( Bodo-bodo )

H: creeping perennial herb, in forest. M: 75; 142. D: STP. V: Figuei-

redo 37 (K).

diffusa Burnt. f. subsp. diffusa (Bodo-bodo, Capim-d'dgua, Capulo,

Dacala, Fia-bodo-bodo, Mucorroana)

Bothalia 41,1 (2011)

51

H: annual or perennial herb with blue to violet flowers, ruderal. D:

STP. V: Figueiredo 54 (K).

petersii Hassk.

H: straggling or erect perennial herb with blue flowers, in secondary

forest. D: ST. V: Figueiredo 55 (K).

PALISOTA Rchb. ex Endl.

braeteosa C.B. Clarke

H: herb with pinkish white or white flowers, in forest. D: ST.

pedicellata K.Schum. ( Ucuete-macaco , Vaiele)

H: herb, with white flowers, in forest. M: 83; 87; 163. D; STP. V:

Figueiredo 35 (K); Paiva 1918 (COI).

POLLIA Thunb.

condensata C.B. Clarke (Bogoto, Uquete-d'obo)

H: perennial herb with white flowers and blue or violet fruit, in forest.

M; 51. D: STP. V: Figueiredo 100 (K).

ntannii C.B. Clarke

H: straggling herb with white flowers. D: ST.

?*RHOEO Hance

?*spathacea (Sw.) Steam

N: recorded by Exell (1944) based on the literature; no specimens

known.

STANFIELDIELLA Brenan

imperforata (C.B. Clarke) Brenan var. imperforata

H: herb, in forest. D: STP. V: Figueiredo 116 (K).

*TRADESCANTIA L.

*zebrina Heynh. ex Bosse

H: herb with purple flowers. D: STP. V: Paiva 235 (COI).

CONNARACEAE

AGELAEA Sol. ex Planch,

pentagyna (Lam.) Baill.

coccinea Exell

obliqua (P.Beauv.) Baill. var. cordata (G.Schellenb.) Exell

phaeocarpa Exell

reticulata Exell

H: liana, in forest. D: STP. V: Joffroy 139 (BRLU).

rubiginosa Gilg

principensis Exell

H: liana, in forest. D: P.

CNESTIS Juss.

ferruginea Vahl ex DC. ( Mondim-moela, Muandim-mudla )

H: tree, in forest. M: 22; 95. D; STP. V: Madureira & Martins 75

(COI).

CONNARUS L.

africanus Lam. (Corda-ana)

H: shrub or scandent shrub. D: ST.

griffonianus Baill.

H: woody climber, with white flowers. D: P.

ROUREA Aubl.

parv iflora Gilg

H: liana, in forest. D: ST. V: Carvalho 9 (LISC).

CONVOLVULACEAE

*CUSCUTA L.

*campestris Yunck.

H; parasitic herb, in plantations and on roadsides. D: ST. V; Figueiredo

77 (K).

IPOMOEA L

*alba L.

H: liana, with white or pink flowers. D: STP.

*batatas (L.) Lam. (Batata-doce)

H: climber with purple, pink or white flowers, on roadsides. E: tuber-

ous roots. D: STP. V; Oliveira 82 (BRLU).

cairica (L.) Sweet var. cairica

H: climber with white to purple flowers. D: STP.

eriocarpa R.Br.

H: annual herb, white to purple flowers, ruderal. E: leaves. D: ST. V:

Matos 7626 (LISC).

*hederifolia L.

H: annual climbing herb with red flowers. D; STP.

imperati (Vahl) Griseb.

stolonifera J.F.Gmel.

H: on coastal areas. D: ST.

*indica (Burm.f.) Merr.

H: perennial climber with blue flowers, in gardens. D: ST. V: Lejoly

97/275 (BRLU, LISC).

mauritiana Jacq.

H: perennial climber with tuberous roots and flowers reddish purple or

mauve. D: ST. V: Oliveira 1015 (BRLU).

pes-caprae (L.) R.Br. subsp. brasiliensis (L.) Ooststr. ( Fia-tataluga ,

Folha-de-tartaruga)

H: climber, with white or purple flowers, in coastal areas. D: STP. V:

Paiva 1284 (COI).

*quamoclit L.

H: annual or perennial climber, with white, red or pink flowers. D:

STP. V: Paiva 1801 (COI).

rubens Choisy

H: perennial climber, with purple or mauve flowers. D: ST.

*setifera Poir.

H: climber. D: P.

*tiliacea (Willd.) Choisy

H: liana with yellow-orange flowers. D: STP. V: Oliveira 561 (BRLU);

Paiva 1303 (COI).

MERREMIA Dennst.

aegyptia (L.) Urb. (Matazem)

H: annual climber with purple flowers, in coastal areas. M: 86; 98;

122; 124; 128; 179. D: ST. V: Paiva 282 (COI).

umbellata (L.) Hallier f. subsp. umbellata

H: perennial climber with yellow flowers. D: P.

XENOSTEGIA D.F.Austin & Staples

tridentata (L.) D.F.Austin & Staples subsp. angustifolia (Jacq.)

Lejoly & Lisowski

Merremia tridentata (L.) Hallier f. subsp. angustifolia (Jacq.) Ooststr.

H: trailing herb. D: ST.

CRASSULACEAE

KALANCHOE Adans.

crenata (Andrews) Haw. var. crenata ( Fia-damina , Fia-da-mina-que,

Ufia-cabece)

H: succulent herb with yellow, orange or magenta flowers. M; 54; 57;

79; 94; 1 19; 124; 174. D: ST. V: Madureira & Martins 72 (COI).

*pinnata (Lam.) Pers. (Fia-damina, Folha-da-mina)

Bryophyllum pinnatum (L.f.) Oken

H: perennial, succulent plant. M: 37; 54; 57; 79; 94; 119; 124; 167;

174. D: STP. V: Madureira & Martins 139 (COI).

CUCURBITACEAE

CAYAPONIA Silva Manso

africana (Hook.f.) Exell

H: perennial, climbing herb, in forest and secondary forest. D: ST. V:

Figueiredo 249 (LISC).

7CITRULLUS Eckl. & Zeyh.

?Ianatus (Thunb.) Matsum. & Nakai (Melancia)

N: annual climbing herb with yellow flowers cultivated for fruit

(watermelon), recorded doubtfully for Principe (Exell 1 944) with-

out recorded specimens.

COCCINIA Wight & Am.

barteri (Hook.f.) Keay

H: climbing herb, in forest. D: ST. V: Matos 7419 (BRLU, LISC).

CUCUMIS L.

*sativus L. (Pepino)

H: annual climber with yellow flowers. E: fruits, leaves. D: ST.

metuliferus E.Mey ex Naudin

H: annual climbing or trailing herb. E: fruits. D: ST.

DIPLOCYCLOS (Endl.) Post & Kuntze

palmatus (L.) C. Jeffrey (Coedano, Stlofi-d’obo, Ucoedano,

Uquedano)

H; perennial climber. M: 47. D: ST. V; Madureira & Martins 477

(COI).

52

Bothalia 41,1 (2011)

LAGENARIA Ser.

breviflora (Benth.) Roberty (Chimom-coia, Simao-Correia )

H: perennial climber with yellow flowers, in forest, secondary forest

and cultivated ground. M: 5; 22; 53; 87; 150. D: STP. V: Figuei-

redo 203 (LISC).

siceraria (Molina) Standi. (Cabafa, Ocd)

H: annual climber. E: fruits, leaves. D: ST. V: Oliveira 226 (BRLU).

LUFFA Mill.

cylindrica (L.) M.Roem. (Mamalongo)

aegyptiaca Mill.

H: annual herb or perennial climber with yellow flowers, ruderal. E;

young fruits. N: fruits are used as sponges. D: ST. V: Paiva 1298

(COI).

MOMORDICA L

charantia L. var. charantia (Chlofi, Fia-sanzom, Melao-de-Sao-

-Caetano, Sancaetano, Stlofi, Tloufi)

H: annual climber or trailer with yellow flowers, in forest and planta-

tions, ruderal. E: fruits, leaves. M: 57; 60; 79; 95; 112; 124; 136;

150; 158; 183. D: STR V: Figueiredo 201 (LISC).

charantia L. var. abbreviata Ser. (Chcofi, Fia-estelofi)

H: annual climber or trailer. E: fruits. D: STR V: Paiva 1145 (COI).

foetida Schumach.

H: perennial climbing herb with woody rootstock, ruderal. D: ST. V:

Lejoly 95/111 (BRLU).

PEPONIUM Engl.

vogelii (Hook.f.) Engl. ( Batata-pim-pim )

H: climbing or trailing herb with yellow Bowers. M: 16; 87; 178; 183.

D: ST. V: Paiva 1148 (COl).

*SECHIUM Juss.

*edule (Jacq.) Sw. (Chuchu, Pimpinela )

H: climber or trailing herb with cream-coloured flowers, on roadsides.

E: fruits, leaves, roots. D: ST. V: Paiva 106 (COI).

ZEHNERIA Endl.

gilletii (De Wild.) C. Jeffrey

H: climbing herb with slender stems. D: ST.

scabra (L.f.) Sond. subsp. scabra

H: perennial herb, climbing or trailing, near cultivated ground, on

roadsides. D: ST. V: Paiva 1158 (COI).

*CYCLANTHACEAE

*CARLUDOVTCA Ruiz & Pav.

*palmata Ruiz & Pav.

H: tufted perennial palm-like plant. D: ST. V: Paiva 1941b (COI).

CYPERACEAE

ABILDGAARDIA Vahl

ovata (Burm.f.) Krai

Fimbristylis ovata (Burm.f.) Kern

H: perennial herb. D: ST.

CAREX L.

'Teptocladus C.B. Clarke

H: herb, in forest. D: ST. V: Lejoly 94/560 (LISC).

CYPERUS L.

articulatus L.

H: herb. D: P. V: Oliveira 532 (BRLU).

§baronii C.B.CIarke

H: herb. D: ST.

difformis L.

H: annual herb. D: ST.

distans L.f. (Zuya)

H: herb, in forest. D: STP. V: Paiva 80 (COI).

esculentus L.

H: perennial herb. D: STP. V: Paiva 967 (COI).

exaltatus Retz.

H; herb. D: ST.

haspan L.

H: herb. D: ST.

laxus Lam. subsp. buchholzii (Boeckeler) Lye

diffusus Vahl subsp. buchholzii (Boeckeler) Kiik.

H: herb. D: STP. V: Lejoly 94/564 (BRLU).

renschii Boeckeler var. renschii

sylvicola Ridley

H: herb. D: ST.

rotundus L. (Zufa)

H: herb. D: STP.

sphacelatus Rottb.

H: herb, in forest. D: STP. V: Oliveira 521 (BRLU).

ELEOCHAR1S R.Br.

geniculata (L.) Roem. & Schult.

H: herb. D: ST.

FIMBRISTYLIS Vahl

cymosa R.Br.

N: recorded for Principe by Exell (1973) based on a reference by Silva

(1958); no specimens known,

dichotoma (L.) Vahl

H: perennial herb. D: STP.

ferruginea (L.) Vahl

H: perennial herb. D: ST.

FUIRENA Rottb.

umbellata Rottb.

H: perennial herb. D: ST.

HYPOLYTRUM Rich.

'grande (Uittien) Koyama

H: herb. D: STP. V: Matos 7696 (LISC).

heteromorphum Nelmes

H: herb. D: STP. V: Joffroy 198 (BRLU).

KYLLINGA Rottb.

erecta Schumach.

H: herb. D: ST. V: Stevart 167 (BRLU).

nemoralis (J.R.Forst. & G.Forst.) Dandy ex Hutch. & Dalziel

H: perennial herb. D: STP. V: Oliveira 512 (BRLU).

peruviana Lam.

H: herb. D: STP.

pumila Michx.

H: herb. D: STP. V: Matos 7604 (LISC).

MAPANIA Aubl.

Terruginea Ridl. ( Pema-d'obo )

H: herb, in forest. M: 34. D: STP. V: Paiva 1111 (COI), 1490 (COI).

MARISCUS Vahl

cylindristachyus Steud. (Zuga)

alternifolius sensu auct. non Vahl

H: herb, ruderal. D: STP. V: Paiva 573 (COI), 704 (COI).

dubius (Rottb.) Kiik. ex C.E.C.Fisch.

H: herb. D: STP.

flabelliformis Kunth var. flabelliformis

H: herb. D: STP. V: Matos 7453 (LISC).

ligularis (L.) Urb. ( Zuqa )

H: herb. D: STP. V: Matos 7661 (LISC).

longibracteatus Cherm.

H: herb. D: ST.

PYCREUS P.Beauv.

polystachyos (Rottb.) P.Beauv.

H: herb. D: STP. V: Matos 7254 (LISC).

REMIREA Aubl.

maritima Aubl.

H: herb. D: P.

RHYNCHOSPORA Vahl

eorymbosa (L.) Britton

H: perennial herb. D: ST.

SCLERIA P.J.Bergius

naumanniana Boeckeler

H: herb, in forest. D: STP. V: Stevart 113 (BRLU).

TORULINIUM Desv. ex Ham.

odoratum (L.) S.S. Hooper

H: herb. D: STP. V: Paiva 1155 (COl).

Bothalia 41,1 (2011)

53

DICHAPETALACEAE

DICHAPETALUM Thouars

Ebocageanum (Henriq.) Engl. ( Melambo )

H: tree (?). D: ST. N: apparently rare; only two collections have been

recorded.

DILLENIACEAE

*DILLENIA L.

*indica L.

H: shrub or small tree, with white flowers. E: fruits. D: ST. V: Lejoly

95/051 (BRLU, LISC).

TETRACERA L

alnifolia Willd. subsp. alnifolia

H: liana, in forest. D: STP. V: Joffroy 134 (BRLU, LISC).

DIOSCOREACEAE

DIOSCOREA L.

*alata L. ( Gudu , Inhame-branco, Inhame-gudu, Som-Iongo, Sonlongo)

H: climber up to 15 m high, producing one tuber. E: tuber. D: ST.

bulbifera L. (Cara-zambuco, Cuini, lnhame, Inhame-zambuco, Ofd.

Otoni )

H: climber with small woody tuber and axillary bulbils. E: tuber. M; 5.

D: STP. V: Paiva 1013 (COI), 1476 (COI).

cayenensis Lam. (Inhame-amarelo, Inhame-branco, Inhame-gundu,

Ofd )

H: climber producing tubers. E: tuber. M: 152. D: STP. V: Lejoly

98/253 (BRLU); Paiva 1872 (COI).

dumetorum (Kunth) Pax (Bofd, lnhame-bravo, Ofd, Quinin.

Unquiniri)

H: climber with tubers. E: tuber (famine-food). D: ST.

minutiflora Engl. (Inhame-de-Benin)

H: climber with tubers. E: tuber (famine-food). D: ST.

sansibarensis Pax (Cuini)

H: climber with tubers. E: tuber (famine-food). D: ST.

DRACAENACEAE

DRACAENA L

arborea (Willd.) Link ( Pau-sabao , Po-sabom)

H: tree to 20 m high, with creamy white flowers. M: 22; 51; 158; 170;

175. D: STP V: Madureira & Martins 122 (COI).

aubryana Brongn. ex E.Morren ( Fia-fita )

monostachya Baker

H: shrub with white to greenish white flowers. D: P. V: Stevart 241

(BRLU).

laxissima Engl. (Folha-pempe, Jambele)

H: shrub to 2 m high, with greenish white flowers. D: STP. V: Figuei-

redo 62 (K); Paiva 618 (COI).

EBENACEAE

DIOSPYROS L.

ferrea (Willd.) Bakh.

H: tree. E: fruits. D: ST.

ERICACEAE

ERICA L.

Fthomensis (Henriq.) Dorr & E.G.H.Oliv. ( Urze )

H: shrub, in primary forest. D: ST. V: Matos 7370 ( LISC).

^RHODODENDRON L.

*mucronatum (Blume) G.Don

H: shrub. D: ST.

ERYTHROXYLACEAE

ERYTHROXYLUM PBrowne

*coca Lam.

H: shrub. D: ST.

emarginatum Thonn.

H: shrub or small tree, in forest. D: ST. V: Oliveira 1173 (BRLU).

EUPHORBIACEAE

ACALYPHA L.

amentacea Roxb. subsp. wilkesiana (Mull.Arg.) Fosberg

wilkesiana Mull.Arg.

H: shrub. O. D: ST. V: Stevart 164 (BRLU).

ciliata Forssk.

H: annual herb, in secondary forest. D: ST. V; Oliveira 344 (BRLU).

ALCHORNEA Sw.

cordifolia (Schumach. & Thonn.) Mull.Arg. ( Bengue , Bugi-bugi,

Buje-buje, Bungi-bungi )

H: shrub or small tree, erect or climbing, in forest, savanna and sec-

ondary forest. M: 16; 79; 90; 94; 95; 124; 130; 158. D: STP. V:

Figueiredo 238 (LISC).

hirtella Benth.

H: tree or shrub, in forest. D: STP. V: Matos 7706 (LISC).

laxiflora (Benth.) Pax & K.Hoffm.

H; shrub or small tree, in savanna. D: ST. V: Figueiredo 80 (K).

ANTHOSTEMA A.Juss.

aubryanum Baill.

H: tree, in swamps or river margins. D: STP.

ANTIDESMA L

membranaceum Mull.Arg.

H: shrub or tree, in forest. D: ST. V: Matos 7635 (LISC).

vogelianum Mull.Arg.

H: shrub or tree, in forest. D: ST. V: Figueiredo 241 (LISC).

*BREYNIA J.R.Forst. & G.Forst.

*disticha J.R.Forst. & G.Forst. (Filanto, Lagrima-de-Cristo, Pau-flor,

Po-floli)

nivosa (W.G.Sm.) Small

H: shrub, used as hedge plant. O. M: 94; 150. D: ST. V: Paiva 653

(COI).

BRIDELIA Willd.

micrantha (Hochst.) Baill. (Gigo, Muindo, Vundem)

stenocarpa Mull.Arg.

H: tree, in forest. M: 13; 22; 56; 57; 71; 134; 177. D: STP. V:

Madureira & Martins 261 (COI).

CAPERONIA A.St.-Hil.

latifolia Pax

H: herb, by watercourses. D; ST.

CAVACOA J. Leonard

quintasii (Pax & K.Hoffm.) J. Leonard

H: tree up to 20 m high, in forest. D: ST.

*CHAMAESYCE Gray

*hirta (L.) Millsp. (Belo-vleme, Fia-sanope, Folha-centopeia)

H: herb, ruderal. M: 5; 28; 77; 98; 174. D: STP. V: Paiva 271 (COI).

*hyssopifolia (L.) Small

H: annual herb. D: STP.

*prostrata (Aiton) Small (Cento, Centopeia, Fia-cento, Fia-fleminga-

-blanco, F olha-formiga)

H: herb, ruderal. M: 10; 64; 98; 117; 161. D: STP. V: Paiva 291 (COI).

*serpens (Kunth) Small (Fia-flomiga. F olha-formiga)

H: herb, ruderal. D: ST. V: Paiva 122 (COI).

CLEISTANTHUS Hook.f. ex Planch.

libericus N.E.Br. ( Nono , Vilo, Vilo-pleto, Viro-preto)

H: shrub or small tree, in forest. D: ST. V: Matos 7691 (LISC).

*CODIAEUM Rumph. ex A.Juss.

*variegatum (L.) Blume ex A.Juss. var. variegatum

N: ornamental shrub recorded by Exell (1944) based on a literature

citation; no specimens known.

CROTON L.

draconopsis Mull.Arg. (Pau-purga)

H; tree, in forest. D: ST. V: Oliveira 1323 (BRLU).

54

Bothalia 41,1 (201 1)

CROTON L. (cont.)

Estellulifer Hutch. ( Cubango )

H: tree, in forest. M: 6; 9. D: STP. V: Madureira & Martins 483 (COI).

DISCOCLAOXYLON (Mull.Arg.) Pax & K.Hoffm.

EoccidentaIe (Mull.Arg.) Pax & K.Hoffm.

H: tree, in forest. D: STP. V: Joffroy 230 (BRLU); Matos 7502 (L1SC).

DISCOGLYPREMNA Prain

caloneura (Pax) Prain (Pau-branco, Po-blanco )

H: tree, in forest. D: ST.

DRYPETES Vahl

Eglabra (Pax) Hutch. ( Mama-d'obo , Mamon-d'obo)

H: tree in forest. D: ST. V: Matos 7235 (LISC).

Ehenriquesii (Pax) Hutch. (Mamon-d’obo, No-no)

H: tree, in forest. M: 8; 22. D: ST. V: Madureira & Martins 407 (COI).

principum (Mull.Arg.) Hutch.

H: tree, in forest. D: P.

ELAEOPHORBIA Stapf

drupifera (Thonn.) Stapf (Magloso, Margoso, Paga-oe, Paga-olho,

Paga-olho-de-marcaqao, Paga-ue, Paga-ue, Pau-amargoso,

Tapa-oe)

H: tree, in forest edges. M: 13; 177. D: STP. V: Paiva 1169 (COI).

ERYTHROCOCCA Benth.

Ecolumnaris (Mull.Arg.) Prain

H: shrub or small tree, in forest. D: P. V: Oliveira 215/98 (BRLU).

Emolleri (Pax) Prain ( Bugi-bugi , Lutchica)

H: shrub or small tree, in forest. D: ST. V: Figueiredo 252 (LISC).

EUPHORBIA L.

cervicornu Baill.

H: perennial herb. D: ST.

*heterophylla L.

H: herb, on roadsides. D: ST. V: Paiva 108 (COI).

*peplus L.

H: annual herb. D: ST.

*pulcherrima Willd. ex Klotzsch

H: herb, in gardens. D: ST. V: Paiva 830 (COI).

*tirucalli L. (Alvo-sin-fid, Arvore-sem-folha )

H; shrub or tree, used as hedge plant. O. M: 27; 57; 146. D: ST. V:

Madureira & Martins 67 (COI).

EXCOECARIA L.

guineensis (Benth.) Mull.Arg.

H: shrub, in forest. D: P.

GROSSERA Pax

Eelongata Hutch.

H: tree with white flowers, in forest. D: P.

multinervis J. Leonard

H: tree with white flowers, in forest and plantations. D: ST. V: Matos

7648 (LISC).

*HEVEA Aubl.

*brasiliensis (A.Juss.) Mull.Arg. ( Borracha )

H: tree, in plantations. D: ST. V: Paiva 826 (COI).

*guianensis Aubl.

N; recorded by Exell (1944) based on a citation by Henriques without

specimens.

TJATROPHA L.

*curcas L. (Folha-grao, Glom-maluco, Glon, Grao-maluco, Pau-glon,

Pau-pixe, Purgueira )

H: shrub or tree, in plantations and on roadsides. M: 120; 146. D: ST.

V: Paiva 111 (COI).

*multifida L. (Flom-gongo, Glom-congo, Glom-gongo)

H: shrub or tree, used as hedge plant. M: 120; 146. D: STP. V: Paiva

793 (COI).

KLAINEANTHUS Pierre ex Prain

gabonii Pierre

H: tree, in forest. D: ST.

MACARANGA Thouars

monandra Mull.Arg. (Begbe, Begbe, Po-ngamala)

H: shrub or tree, in forest. D: ST.

MAESOBOTRYA Benth.

Eglabrata (Hutch.) Exell

H: shrub or small tree, in forest and secondary forest. D: STP. V:

Oliveira 553 (BRLU), 1391 (BRLU).

*MANIHOT Mill

*esculenta Crantz ( Mandioca )

H: shrub or undershrub with green flowers, on roadsides and cultivated

ground. E: roots, leaves. M: 13; 32; 114. D: ST. V: Paiva 1037

(COI).

*glaziovii Mull.Arg.

H: shrub or tree.O. D: STP.

MANNIOPHYTON Mull.Arg.

fulvum Mull.Arg. (Congo-gloncongo)

H: straggling bush or liana, in forest. D: STP.

MARGARITARIA L.f.

discoidea (Baill.) G.L. Webster var. discoidea ( Pau-ferro , Po-felo,

Sitvi)

Phyllanthus discoideus (Baill.) Mull.Arg.

H: tree, in forest and secondary forest. M: 87; 124; 173. D: STP. V:

Madureira & Martins 467 (COI); Paiva 936 (COI).

NEOBOUTONIA Mull.Arg.

mannii Benth. (Pluga-mato)

H: tree, in forest. M: 146. D: STP. V: Madureira & Martins 485 (COI).

PHYLLANTHUS L.

*amarus Schumach. & Thonn. ( Cafena-tlache , Ucue-clache, Ue-

-tlache)

H: annual herb, ruderal. M: 77; 98; 117; 130; 178; 179; 181. D: STP.

V: Pague 4 (COI); Paiva 1500 (COI).

dinklagei Pax

H: climbing shrub in secondary forest. D: ST. V: Lejoly 97/357

(BRLU, LISC).

gagnioevae Brunei & J.P.Roux

odontadenius Mull.Arg.

H: annual or perennial herb, woody at base, in forest. D: STP. V: Paiva

565 (COI), 1018 (COI).

muellerianus (Kuntze) Exell

H: shrub or climber, occasionally tree, in forest. D: ST. V: Matos 7451

(LISC).

physocarpus Mull.Arg.

H: straggling tree, in forest. N: collected also in Nigeria once. D: P.

tessmannii Hutch.

H: shrub or tree, in forest. D: ST.

urinaria L. (Cafena-tlache, Ucue-clache, Ue-tlache)

H: annual herb,' ruderal. M: 77; 98; 117; 130; 178; 179; 181. N: very

similar to P. amarus and not distinguished from that species by the

local population. D: ST. V: Madureira & Martins 300 (COI).

PROTOMEGABARIA Hutch

macrophylla (Pax) Hutch.

H: tree, in forest. D: STP. V: Oliveira 154/98 (BRLU).

PSEUDAGROSTISTACHYS Pax & K.Hoffm.

africana (Mull.Arg.) Pax & K.Hoffm. subsp. africana (Cacau-de-obo,

Dumo-branco)

H: shrub or tree, with white flowers, in forest. D: ST. V: Matos 7578

(LISC).

*RICINUS L.

*communis L. (Mamanododo, Mamona, Mamono, Mamono,

Mamonon, Ricino, Zague-zague )

H: shrub, ruderal. M: 39; 69; 146; 174. D: STP. V: Paiva 801 (COI).

SCLEROCROTON Hochst.

cornutus (Pax) Kruijt & Roebers

Sapium cornutum Pax

H: climbing shrub or tree, in forest. D: ST. V: Oliveira 7 (BRLU).

SHIRAKIOPSIS Esser

elliptica (Hochst.) Esser ( Pau-maria )

Sapium ellipticuin (Hochst.) Pax

H: shrub or tree, in secondary forest. D: ST. V: Lejoly 97/201 (BRLU,

LISC).

TETRORCHIDIUM Poepp.

didymostemon (Baill.) Pax & K.Hoffm. (Branco, Pau-branco, Pau-

-mole, Po-blanco, Pd-moli)

H: tree, in secondary forest. D: ST. V: Oliveira 1546 (BRLU).

Bothalia 41,1 (2011)

55

THECACORIS A.Juss.

Emanniana (Miill.Arg.) Mull.Arg. (Pau-figado)

H: tree, ± 12 m high, in forest. D: ST.

Emembranacea Pax

H: shrub, in secondary forest. D: ST. V: Oliveira 62/98 (LISC).

stenopetala (Mull.Arg.) Mull.Arg.

H: small tree up to 6 m high with greenish white flowers, in forest. D:

P.

TRAGIA L.

tenuifolia Benth.

H: climbing undershrub, in forest. D: ST.

UAPACA Baill.

guineensis Mull.Arg. ( Mangne-d'obo , Nespera-de-bosque, Nespla-

- d'obo )

H: tree, in forest and secondary forest. M: 3; 22; 61; 124. D: STP. V:

Madureira & Martins 171 (COI).

*?VERNICIA Lour.

*?cordata (Thunb.) Airy Shaw ( Aleurite )

Aleurites cordata (Thunb.) R.Br. ex Steud.

N: recorded by Exell (1973) based on a citation by Silva (1958) with-

out specimens.

FABACEAE

ABRUS Adans.

precatorius L.

H: climbing herb with pink to purple flowers and black and scarlet

seeds. D: ST. V: Matos 7320 (LISC).

ACACIA Mill.

*farnesiana (L.) Willd. ( Bana-muala )

H: shrub or tree with yellow flowers. D: ST.

kamerunensis Gand. (Piam-fogo)

H: liana or climbing shrub, in forest. M: 122; 165. D: ST. V:

Madureira & Martins 211 (COI).

*nilotica (L.) Willd. ex Delile subsp. indica (Benth.) Brenan

H: tree. D: ST.

pentagona (Schumach.) Hook.f.

H: liana, in forest. D: P.

*ADENANTHERA L.

*pavonina L. ( Acacia-coral)

H: tree with small yellowish flowers and scarlet lens-shaped seeds. D:

ST. V: Matos 7681 (BRLU, LISC).

AESCHYNOMENE L.

indica L.

H: annual or perennial herb. D: STP.

*ALBIZIA Durazz.

*lebbeck (L.) Benth.

H: tree, with purple flowers. D: ST. V: Figueiredo 90 (K).

*procera (Roxb.) Benth.

H: tree. D: ST.

ALYSICARPUS Desv.

vaginalis (L.) DC. var. vaginalis

H: perennial herb with pink flowers. D: ST.

*ARACHIS L.

*hypogaea L. ( Amendoim , Ginguba, Gumbo)

H: annual herb with yellow flowers. E: seeds. M: 27; 103; 154. D: ST.

V: Madureira & Martins s.n. (COI).

BAPHIA Afzel. ex Lodd.

nitida Lodd.

H: shrub or tree, in rainforest and coastal areas. D: ST. V: Oliveira 529

(BRLU); Stevart 166 (BRLU).

*BAUHIMA L

*monandra Kurz

H: tree with pink flowers. D: ST. V: Figueiredo 86 (K).

*purpurea L.

Perlebia purpurea (L.) A. Schmitz

H: shrub with purple flowers. D: ST. V: Paiva 832 (COI).

*tomentosa L.

Pauletia tomentosa (L.) A. Schmitz

H: shrub or tree with large yellow flowers. D: ST.

CAESALPINIA L

bonduc (L.) Roxb. (Hido-hido, ldo-ido)

H: shrub or tree. M: 27; 170. D: ST. V: Madureira & Martins 448

(COI).

*pulcherrima (L.) Sw.

H: tree with red and yellow flowers. O. D: STP.

*CAJANUS DC.

*cajan (L.) Millsp. ( Feijao-congo , Fezom-congo, Uandru )

H: perennial shrub with yellow flowers. E: seeds. M: 106. D: STP. V:

Madureira & Martins 331 (COI).

*CALOPOGONIUM Desv.

*mucunoides Desv.

H: annual or perennial climber with blue flowers. D: ST.

CANAVALIA DC.

*ensiformis (L.) DC.

H: annual climber with white flowers. E: young pods or immature

seeds. D: ST.

rosea (Sw.) DC. (Fia-aganga)

maritima Thouars

H: herb with blue flowers, in coastal areas. D: ST. V: Paiva 980 (COI).

CASSIA L. emend. Gaertn.

*alata L.

H: shrub ± 4 m high with yellow flowers. D: ST.

*flstula L. (Canafistula, Canapistula. Cassia-oficinal)

H: deciduous or semi-evergreen tree up to 15 m high, with scented,

pale or vivid yellow flowers. M: 79; 87; 98; 103; 114; 119. D: ST.

V: Madureira & Martins 326 (COI).

*javanica L.

H: tree ± 6 m high, with purple to pink flowers. D: ST.

mannii Oliv.

sieberiana sensu auct. non DC.

H: tree with white or pink flowers, in forest. D: STP.

*obtusifolia L

Senna obtusifolia (L.) Irwin & Bameby

H: annual or perennial herb or undershrub, possibly introduced. D: ST.

*occidenta!is L. (Fedegoso, Maioba. Maiopa, Manhanoca,

Munhanoca)

H: herb or undershrub with yellow flowers, on roadsides. M: 19; 27;

53; 57; 77; 98; 99; 103; 114; 135; 158. D: STP. V: Paiva 1067

(COI).

podocarpa Guill. & Perr. (Fia-zaia, Folha-zaia )

H: shrub with yellow flowers. M: 68; 146; 178. D: ST. V; Madureira &

Martins 309 (COI).

senna L. ( Maioba )

angustifolia Vahl

H: shrub with yellow flowers. D: ST. V: Paiva 1288 (COI).

septemtrionalis Viv.

floribunda sensu auct. non Cav.

laevigata Willd.

Senna septemtrionalis (Viv.) H.S.Irwin & Bameby

H: undershrub with yellow flowers, on roadsides. D: ST. V: Figueiredo

112 (K).

*siamea Lam.

H: tree with yellow flowers. D: ST.

*sophera L. ( Maioba-beni )

H: shrub with yellow flowers. D: STP.

*spectabilis DC.

H: tree with yellow flowers. D: ST. V: Paiva 254 (COI).

*CENTROSEMA Benth.

*plumieri (Pers.) Benth.

H: climbing shrub. D: ST. V: Oliveira 913 (BRLU).

*pubescens Benth.

H: climber with purple flowers, on roadsides. M: 146; 186. D: ST. V:

Paiva 972 (COI).

*?CERCIS L.

*?siliquastrum L. ( Olaia )

N: recorded by Exell (1973) based on a citation by Silva (1958) with-

out specimens.

56

Bothalia 41,1 (2011)

CHAMAECRISTA Moench

pratensis (R.Vig.) Du Puy

Cassia mimosoides L.

H: herb with yellow flowers. D: ST. V: Lejoly 93/653 (BRLU).

CLITORIA L.

falcata Lam.

nibiginosa Juss. ex Pers.

H: climber with white flowers streaked with mauve or red. M: 64. D:

ST. V: Paiva 1198 (COI).

ternatea L. var. ternatea

H: perennial climbing herb. M: 64; 119. D: ST. V: Madureira & Mar-

tins 351 (COY).

7COLOPHOSPERMUM J.Kirk ex J.Leonard

?mopane (J.Kirk ex Benth.) J.Kirk ex J.Leonard

N: doubtfully recorded by Exell (1973) based on a citation by Silva

(1958) without specimens.

CROTALARIA L.

doniana Baker

H: perennial herb. D: ST.

ochroleuca G.Don

H: annual herb. D: ST.

pallida Aiton var. obovata (G.Don) Polhill

mucronata Desv.

H: annual or perennial herb or shrub. D: STP.

retusa L. var. retusa (Maleboque. Malimboque, Marimboque)

H: herb with yellow and pink flower, ruderal. M: 146. D: STP. V:

Paiva 978 (COI).

trichotoma Bojer

zanzibarica Benth.

H: annual or perennial herb. D: ST.

CYNOMETRA L

mannii Oliv. (Quebla-machado. Zungo. Znngii)

H: tree up to 20 m high, in forest. M: 33; 146; 186. D: ST. V: Fignei-

redo 245 (LISC).

DALBERGIA L.f.

ecastaphyllum (L.) Taub. ( Maja , Popian)

H: shrub or tree, in mangroves and coastal bush. D: STP. V: Paiva

1168 (COI).

7DANIELLIA Benn.

Joblonga Oliv.

N: very large tree, reported by Exell (1944) based on Chevalier 14227

(S.Tome, Porto Alegre), a collection not seen by Exell nor by us.

*DELONIX Raf.

*regia (Bojer ex Hook.) Raf.

H: tree with red flowers. O. D: ST. V: Paiva 800 (COI).

*DESMANTHUS Willd.

*virgatus (L.) Willd. (Fia-foguete, Pau-foguete, Pd-fuguete)

H: undershrub with white or yellow flowers, on roadsides, ruderal. M:

5; 133. D: STP. V: Figueiredo 91 (K).

DESMODIUM Desv.

adscendens (Sw.) DC. var. adscendens ( Fia-vintem . Pega-pega)

H: perennial herb with pink flowers. M: 3; 11; 174. D: STP. V:

Madureira & Martins 142 (COI).

gangeticum (L.) DC.

II: perennial herb or shrub. D: STP. V: Oliveira 1198 (BRLU).

*incanum DC. (Pega-pega, Vala-case )

canum (J.F.Gmel) Schinz & Thell.

H: perennial herb with purple flowers, on roadsides and in plantations.

M: 61. D: STP. V: Figueiredo 45 (K); Paiva 1416 (COI).

*procumbens (Mill.) Hitchc.

H: herb. D: ST. V: Matos 7324 (BRLU).

ramosissimum G.Don ( Pega-pega )

H: shrub with white and mauve flowers. M: 61; 112; 158; 168; 183.

D: STP. V: Madureira & Martins 287 (COI); Stevart 160 (BRLU).

repandum (Vahl) DC.

H: perennial herb or shrub with orange red flowers, in secondary forest

and plantations. D: ST. V: Figueiredo 109 (K).

salicifolium (Poir.) DC.

H: perennial herb or shrub. D: ST. V: Stevart 16 (BRLU).

*tortuosum (Sw.) DC.

H: perennial herb or undershrub with purple flowers, on roadsides. D:

ST. V: Paiva 409 (COI).

triflorum (L.) DC.

H: annual or perennial herb. D: STP.

velutinum (Willd.) DC. subsp. velutinum (Valacasei)

H: woody perennial herb or shrub, in savanna. D: STP. V; Matos 732 7

(LISC).

DIALIUM L.

guineense Willd. ( Salamba )

H: tree. D: STP. V: Matos 7686 (LISC).

DIOCLEA Kunth

reflexa Hook.f. (Coda-ipe. Codo-plego, Corda-iple)

H: climbing shrub. D: STP. V: Matos 7276 (LISC).

ERYTHRINA L

?caffra Thunb.

N: doubtful record of a South African species (Exell 1944) based on a

literature citation; no specimens known,

droogmansiana De Wild. & T.Durand

H: shrub or tree up to 20 m high, in forest. D: ST.

*fusca Lour.

H: tree. D: ST. V: Paiva 150 (COI).

*mitis Jacq.

umbrosa Kunth

N: recorded by Exell (1973) based on a citation by Silva (1958); no

specimens known.

*poeppigiana (Walp.) O.F.Cook (Litrina)

H: tree, in plantations and secondary forest. M: 58; 191. D: ST. V:

Figueiredo 118 (K).

variegata L. ( Litrina )

indica Lam.

H: tree. M: 58; 191. D: STP. V: Figueiredo 234 (LISC).

*velutina Willd.

N: recorded by Exell (1944, 1973) and by Silva (1958); no specimens

known.

FAIDHERBIA A.Chev.

albida (Delile) A.Chev.

H: tree up to 18 m high with yellow flowers. D: ST.

*HAEMATOXYLUM L

*campechianum L.

H: tree. D: ST.

INDIGOFERA L.

astragalina DC.

H: annual herb. D: ST.

colutea (Burm.f.) Merr. var. colutea

H: annual semi-woody herb. D: ST.

hirsuta L. var. hirsuta

H: annual semi-woody herb. D: ST.

spicata Forssk. var. spicata (Banco)

H: perennial herb. M: 99; 126. D: ST. V: Madureira & Martins 286

(COI).

*suffruticosa Mill. (Ago)

H: undershrub, possibly introduced. D: ST. V: Lejoly 97/294 (LISC).

tinctoria L. var. tinctoria (Ago. Ago-mato, Anileiro)

H: shrub. N: one of the original plants used to produce indigo dye. D:

STP. V: Matos 7321 (LISC).

trita L.f. var. subulata (Vahl ex Poir.) Ali

H: woody herb. D: ST. V: Matos 7231 (LISC).

*INGA Mill.

*edulis Mart. (Cajareiro, Cajaseiro. Ingareiro. Ingaseiro)

H: tree. E: fruits. D: STP. V: Figueiredo 53 (K).

*LEUCAENA Benth.

*leucocephala (Lam.) de Wit (Leucena. Pau-foguete)

H: shrub or tree with white flowers, ruderal. D: STP. V: Figueiredo 92

(K).

LONCHOCARPUS Kunth

sericeus (Poir.) Kunth (Colema, Colima. Cohna)

H: shrub or small tree with lilac flowers, in savanna. M: 146; 175. D:

STP. V: Madureira & Martins 487 (COI).

Bothalia 41,1 (2011)

57

MACHAERIUM Pers.

lunatum (L.f.) Ducke

Drepanocarpus lunatus (L.f.) G.Mey.

H: shrub with purple flowers, in coastal areas. D: STP.

MILLETTIA Wight & Am.

barteri (Benth.) Dunn ( Colima-doido )

H: shrub or liana. M: 147; 167. D: ST. V: Madureira & Martins 423

(COI).

griffoniana Baill.

Lonchocarpus griffoniamts (Baill.) Dunn

H: tree up to 10 m high, with reddish purple flowers. D: ST.

thonningii (Schumach. & Thonn.) Baker ( Colima , Colima-fria,

Colma)

H: tree up to 18 m high, with reddish purple flowers. D: ST.

*MIMOSA L.

*polydactvla Humb. & Bonpl. ex Willd. (Fia-malicia, Fissope, Mafi-

-sope)

H: perennial herb with white flowers. D: STP. V: Paiva 713 (COI).

*pudica L. var. hispida Brenan ( Fia-malicha , Fia-malichia)

H: annual or perennial herb. M: 167; 170. D: ST. V: Paiva 156 (COI).

MUCUNA Adans.

flagellipes Hook.f.

H: liana with creamy white or yellowish flowers, in coastal areas. D:

STP. V: Matos 7651 (LISC).

pruriens (L.) DC. var. pruriens (Macunja)

H: climber with violet flowers. E: seeds. M: 27; 98. D: ST. V:

Madureira & Martins 284 (COI); Paiva 1874 (COI).

sloanei Fawc. & Rendle

H: climber with yellow flowers. D: ST.

NEONOTONIA J.A.Lackey

wightii (Wight & Am.) J.A.Lackey

Glycine wightii (Wight & Am.) Verde.

H: perennial climber with small white or mauve flowers. D: ST.

ORMOCARPUM P.Beauv.

sennoides (Willd.) DC. subsp. hispidum (Willd.) Brenan & J. Leonard

H: shrub with red-streaked yellow flowers. M: 22; 61; 150. D: ST. V:

Madureira & Martins 332 (COI).

verrucosum P.Beauv. var. verrucosum

H: shrub with white or lilac, purple-striped flowers, in swamps and by

the sea. D: STP. V: Matos 7270 (LISC).

*?ORMOSIA Jacks.

*?dasvcarpa Jacks.

N: record (Exell 1973) based on a citation by Silva (1958) without

specimens.

*PARASERIANTHES I.C.Nielsen

*falcataria (L.) I.C.Nielsen (Acacia, Molucana)

Albizia falcata (L.) Backer

A. falcataria (L.) Fosberg

H: tree, in disturbed forest. D: ST. V: Paiva 1061 (COI).

PARKIA R.Br.

biglobosa (Jacq.) R.Br. ex G.Don (Farroba, Luba, Luba, Luva, LJluba)

oliveri J.F.Macbr.

H: tree with orange or red flowers, in savanna. E: seeds. D: ST. V:

Matos 7707 (BRLU, LISC).

*PELTOPHORUM (Vogel) Benth.

*pterocarpum (DC.) K.Heyne

H: tree up to 17 m high with yellow flowers. D: ST.

PENTACLETHRA Benth.

macrophylla Benth. (Moandim, Moandjim, Mondim, Muandi,

Muandim, Muange, Sicupira, Uba)

H: large tree with yellowish creamy flowers, in secondary forest. E:

seeds. M: 8; 95; 124; 149. D: STP. V: Paiva 11 (COI), 1475 (COI).

*PHASEOLUS L.

*lunatus L. (Feijao-cutelinho, Feijao-espadinho)

H: annual or perennial climber with white flowers. E: fruits, seeds. D:

ST.

*vulgaris L. var. vulgaris ( Feijao )

H: annual climber with white or pink flowers. E: seeds. D: ST.

PSOPHOCARPUS Neck,

scandens (Endl.) Verde.

H: perennial climbing herb with blue flowers, on roadsides. D: ST. V:

Paiva 946 (COI).

*PTEROCARPUS Jacq.

*indicus Willd.

H: tree with yellow flowers. D: ST.

*PUERARIA DC.

*phaseoloides (Roxb.) Benth. var. javanica (Benth.) Baker

H: climber. E: roots. D: ST.

RHYNCHOSIA Lour.

densiflora (Roth) DC. subsp. debilis (G.Don) Verde.

H: climber. D: ST.

minima (L.) DC. var. prostrata (Harv.) Meikle ( Sodom-campo )

H: prostrate herb with yellow flowers. M: 67; 96; 146; 167. D: ST. V:

Madureira & Martins 327 (COI).

*SAMANEA Benth.

*saman (Jacq.) Merr. (Acacia-preta, Inga-saman, Pau-chuva)

H: large tree. D: ST.

SESBANIA Scop,

sericea (Willd.) Link

H: annual or biennial herb. D: ST. V: Matos 7406 (LISC).

SOPHORA L.

tomentosa L. subsp. occidentalis (L.) Brummitt

H: shrub with yellow flowers. D: ST.

*TAMARINDUS L.

*indica L. (tree: Tamanha, Tamarindeiro; fruit: Tamarindo)

H: tree with yellow flowers with orange or red streaks. E: fruits. D:

STP. V: Paiva 817 (C 01).

TEPHROSIA Pers.

*candida (Roxb.) DC. (Banco)

H: shrub or small tree. O. M: 139. D: ST V: Madureira & Martins 569

(COI).

noctiflora Bojer ex Baker

H: annual or perennial herb. D: ST. V: Matos 7700 (LISC).

platycarpa Guill. & Perr.

flexuosa G.Don

H: annual herb woody at base, w ith pink or purple flowers. D: ST.

purpurea (L.) Pers. subsp. leptostachya (DC.) Brummitt var. pubes-

cens Baker

H: perennial herb. D: ST.

uniflora Pers. subsp. uniflora

H: perennial herb with pink flowers. D: ST. V: Matos 7328 (LISC).

vogelii Hook.f. (Bamea, Banco, Cafoto)

H: woody herb with white or violet flowers. D: STP.

TERAMNUS P.Browne

labialis (L.f.) Spreng. subsp. arabicus Verde.

H: climber with reddish flowers. D: STP. V: Paiva 979 (COI).

TETRAPLEURA Benth.

tetraptera (Schumach. & Thonn.) Taub. (Cuspila, Cuspira)

H: tree with yellowish to pinkish flowers. D: ST.

URARIA Desv.

picta (Jacq.) DC. (Fia-placela)

H: undershrub with pink, blue or reddish flowers, in savanna. D: ST.

V: Matos 7460 (LISC).

*?VICIA L.

*?faba L. (Fava, Faveira)

N: record (Exell 1973) based on a citation by Silva (1958) without

specimens.

VIGNA Savi

adenantha (G.F.Mey.) Marechal, Mascherpa & Stainier

Phaseolus adenanthus G.Mey.

H: perennial climber. D: ST.

gracilis (Guill. & Pern) Hook.f.

H: twining herb with yellow flowers. D: P. V: Stevart 14 (BRLU).

58

Bothalia 41,1 (2011)

VIGNA Savi (cont.)

luteola (Jacq.) Benth. var. luteola

oblonga sensu auct. non Benth.

H: perennial climber. D: ST. V: Matos 7517 (LISC).

racemosa (G.Don) Hutch. & Dalz.

H: perennial climbing herb with blue flowers. D: STP. V: Figueiredo

206 (LISC).

unguiculata (L.) Walp. subsp. alba (G.Don) Pasquet

alba (G.Don) Planch, ex Baker f.

unguiculata subsp. dekindtiana sensu auct. non (Harms) Verde.

H: perennial herb with white or bluish-purple flowers. E: seeds. D: ST.

V: Matos 7620 (LISC).

*ZORNIA J.F.Gmel.

Hatifolia Sm. var. latifolia

H: annual or perennial herb with yellow flowers, in savanna. D: P. V:

Matos 7764 (LISC).

FLACOURTIACEAE

CASEARIA Jacq.

barteri Mast. ( Bobo-bobo , Inhe-bobd)

mannii Mast.

H: shrub or tree, in forest and secondary forest. D: STP. V: Figueiredo

242 (LISC).

FLACOURTIA Comm, ex L’Her.

indica (Burm.f.) Merr.

flavescens Willd.

H: climbing shrub or tree, in forest. E: fruits. D: ST.

HOMALIUM Jacq.

africanum (Hook, f.) Benth. (Quebra-machado)

H: shrub, in forest. D: ST.

Ehenriquesii Gilg ex Engl. (Quebra-machado)

H: tree, in forest. M: 33; 146; 1 86. D: ST. V: Madureira & Martins 410

(COX).

ONCOBA Forssk.

spinosa Forssk. ( Dibixi , Malebouque, Malimboque)

H: shrub, in forest and savanna. M: 168. D: ST. V: Figueiredo 243

(LISC).

OPHIOBOTRYS Gilg

zenkeri Gilg (Clacia, Clacla, Cuaco-bangana. Stlala-stlala)

H: shrub or tree, in forest. M: 17. D: ST. V: Madureira & Martins 624

(COI).

*FUMARIACEAE

*FUMAR1A L

♦muralis Sond. ex W.D.J.Koch

H: herb with pink flowers. D: ST. V: Lejoly 98/214 (BRLU).

GENTIANACEAE

ANTHOCLEISTA Afzel. ex R.Br.

microphylla Wemham

H: tree or climber with white flowers, in forest and secondary forest.

D: STP. V: Figueiredo 67 (K).

nobilis G.Don

macrophylla G.Don

H: tree with white flowers, in forest and secondary forest. D: STP. V;

Matos 7449 (LISC); Oliveira 221/98 (BRLU).

scandens Hook.f.

H: tree or climber with white flowers, in forest and secondary forest.

D: ST. V: Matos 7359 (LISC).

GESNERIACEAE

EPITHEMA Blume

tenue C.B. Clarke

H: herb, in forest. D: ST.

STREPTOCARPUS Lindl.

elongatus Engl.

thomensis Exell

H: herb up to 0.3 m high with white flowers, in forest. D: ST. V: Matos

7674 (LISC).

nobilis C.B. Clarke

H: herb, in forest. D: ST.

GOODENIACEAE

SCAEVOLA L.

plumieri (L.) Vahl

H: shrub, in littoral regions. D: ST.

HERN ANDIACE AE

HERNANDIA L.

Ebeninensis Welw. ex Henriq. ( Bunga , Pau-candeia)

H: tree, in forest. M: 92; 160. D: ST. V: Madureira <£ Martins 22

(COI).

?*HYDRANGEACEAE

?*HYDRANGEA L

?sp.

N: recorded by Exell (1944) based on a citation by Chevalier; no spec-

imens known.

HYPERICACEAE

HARD NG AN A Lam.

madagascariensis Lam. ex Poir. (Pau-sangue, Po-sangue, Sangue)

H; shrub or small tree, in forest and secondary forest. M: 13; 124. D:

STP. V: Madureira & Martins 460 (COI); Paiva 1060 (COI).

*IRIDACEAE

*NEOMARICA Sprague

*caerulea (Ker Gawl.) Sprague

D: STP.

IRVINGIACEAE

IRVINGIA Hook.f.

gabonensis (Aubry-Lecomte ex O’Rorke) Baill. (Manga-macaco)

H: tree, in forest. D: P.

JUNCACEAE

LUZULA DC.

mannii (Buchenau) Kirschner & Cheek subsp. mannii

campestris (L.) DC. var. mannii Buchenau

H; perennial herb. D: ST.

LAMIACEAE

ACHYROSPERMUM Blume

oblongifolium Baker (Beganssom, Folha-belga)

H: undershrub with green-white flowers. M: 5; 51. D: ST. V:

Madureira & Martins 310 (COI).

7ELSHOLTZIA Willd.

?sp.

N: recorded for S.Tome by Exell (1944) based on a literature citation;

no specimens known.

LEONOTIS (Pers.) R.Br.

nepetifolia (L.) R.Br. (Penicano. Pinicane, Pinincano)

H: herb up to ± 1.5 m high, with pink flowers. M: 5; 22; 54; 60; 61;

77; 123; 150; 158; 174. D: STP V: Madureira & Martins 158

(COI).

♦MENTHA L.

♦rotundifolia (L.) Huds. (Letrao)

H: perennial herb. E: leaves. D: ST.

Bothalia 41,1 (201 1)

59

OCIMUM L.

*americanum L. ( Fia-mosquito )

canum L.

H: perennial herb, near cultivated areas. M: 79; 99; 134; 174. D: ST. V:

Madureira & Martins 604 (COI).

*basilicum L. (Fia-mosquito. Madlicom, Mosquito)

H: perennial herb, near cultivated areas. E: leaves. M; 79; 99; 134;

174. D: ST. V: Paiva 1035 (COI).

gratissimum L. var. gratissimum ( Fia-micoco , Micoco, Micoco-di-

-que, Pau-mosquito)

H: perennial herb with white flowers. E: leaves. M; 21; 27; 52; 79;

156; 166. D: STP. V: Paiva 1045 (COI), 1502 (COI).

^minimum L. ( Madlicom )

basilicum sensu auct. pro parte, non L.

H: perennial herb, near cultivated areas. M: 27; 52; 76; 156. D: ST. V:

Madureira & Martins 104 (COI).

PLATOSTOMA P.Beauv.

africanum P.Beauv.

H: herb up to 0.6 m high, in forest. D: ST.

*SALVIA L.

*coccinea Buc’hoz ex Etl.

H: perennial herb with red flowers. O. D: ST.

*splendens Ker Gawl.

H: perennial herb with red flowers, on roadsides. D: ST. V: Paiva 1005

(COI).

SOLENOSTEMON Thonn.

monostachyus (P.Beauv.) Briq. (Fia-manjolo. Magero, Manjo/o)

H: annual or perennial herb with bluish flowers, in forest, secondary

forest, plantations, ruderal. E: leaves. M: 79; 94; 165; 169. D: STP.

V: Paiva 355 (COI).

*STACHYS L.

*arvensis (L.) L.

H: annual herb. D: ST. V: Matos 7229 (LISC).

*LAURACEAE

*CINNAMOMUM L.

*burmanni (Nees & T.Nees) Blume (Canela-brava)

H: small tree. E: bark (spice: cassia). D: STP. V: Matos 7583 (LISC).

*camphora (L.) J.Presl (aromatic compound: Canfora\ tree: Can-

foreira)

H: tree, in secondary forest. D: ST. V: Oliveira 150 (BRLU).

*verum J.Presl ( Canela . Canela-d'obo, Caneleira. Pau-canela)

zeylanicum Blume

H: small tree with whitish flowers, in secondary forest. E: bark (spice:

canela). M: 173. D: STP. V: Figueiredo 49 (K).

*PERSEA Mill.

*americana Mill, (fruit: Abacate, Avacate. Avocado, Bacatche: tree:

Abacateiro)

H: large tree, with small yellowish flowers. E: fruits. M: 51; 57; 61;

64; 91; 137; 150; 161; 165; 168. D: STP. V: Madureira & Martins

273 (COI).

LEEACEAE

LEEA L.

guineensis G.Don

H: shrub or small tree, with red, orange or yellow flowers. E: fruits. D:

ST. V: Oliveira 1429 (BRLU).

Etinctoria Baker (Cele-ale, Cele-cele, Cele-cele)

H: shrub or small tree with red or orange flowers. D: ST. V: Figueiredo

244 (LISC); Paiva 1925 (COI).

LEMNACEAE

LEMNA L.

aequinoctialis Welw.

perpusilla sensu auct. non Torrey

H: aquatic floating herb. D: ST.

LENTIBULARIACEAE

UTRICULARIA L.

mannii Oliv.

H: small, perennial epiphyte with yellow flowers, in forest. D: STP V:

Jojfroy 209 (BRLU); Paiva 1082 (COI).

striatula Sm.

H: small epiphyte, in forest. D: P. V: Paiva 1483 (COI).

LINACEAE

HUGONIA L.

platysepala Welw. ex Oliv.

H: climbing shrub or small tree, with white or yellow flowers, in for-

est. D: P.

LOBELIACEAE

LOBELIA L.

Ebarnsii Exell

H: herb, in mountain forest. D: ST. V: Matos 7703 (LISC).

molleri Henriq.

H: herb up to 0.6 m high with white flowers, in forest. D: ST. V: Matos

7543 (LISC).

LORANTHACEAE

HELIXANTHERA Lour,

mannii (Oliv.) Danser var. mannii

H: parasitic shrub with pendent stems, in forest and plantations. D: ST.

V: Oliveira 855 (BRLU).

MALPIGH1ACEAE

ACRIDOCARPUS Guill. & Perr.

longifolius (G.Don) Hook.f. (Micando-homem. Milando-homem.

Milondd-homem)

H: small tree or climbing shrub with yellow flowers in secondary for-

est and plantations. M: 22; 167. D: STP. V: Paiva 1009 (COI).

HETEROPTERYS Kunth

leona (Cav.) Exell

H: climber with yellow flowers. D: STP.

MALVACEAE

*ABELMOSCHUS Medik.

*esculentus (L.) Moench (Iquiabo, Quiabo )

H: annual herb with yellow flowers, in plantations and on roadsides. E:

fruits. M: 95. D: STP. V: Paiva 781 (COI).

*moschatus Medik.

H: Perennial herb. E: leaves, fruits. D: ST.

ABUTILON Mill.

grandiflorum G.Don (Fid-mdliva, Maliva, Malva)

H: herb or shrub with yellow to orange flowers. M: 40; 61; 64; 89; 98;

103; 158; 174. D: ST. V; Madureira & Martins 112 (COI).

*grandifolium (Willd.) Sweet ( Mandioca-brava )

H: perennial herb with yellow to orange flowers. D: ST. V: Paiva 1062

(COI).

*striatum Dicks, ex Lindl. (Mandioca-brava. Mata-boi)

venosum Lemaire

H: shrub with orange to purple flowers in secondary forest and on

roadsides. M: 139. D: ST. V: Madureira & Martins 473 (COI).

*GOSSYPIUM L. (U-u-defu. Algodao)

*barbadense L.

H: shrub with yellow flowers. M: 69; 111; 181. D: STP.

*hirsutum L.

H: shrub with yellow flowers. M: 69; 111; 181. D: ST.

HIBISCUS L.

*acetosella Welw. ex Hiem ( Fia-mussa , Fia-mussua, Mussa, Mussa-

-blanco, Mussua)

H: annual or perennial herb or shrub, ruderal. E: leaves, flowers. M:

40; 60. D: STP. V: Madureira & Martins 649 (COI); Paiva 632

(COI).

60

Bothalia 41,1 (201 1 )

HIBISCUS L. (cont.)

donianus D.Dietr.

H: annual herb. D: ST. V: Matos 7318 (LISC).

*mutabilis L.

H: shrub with white or pink flowers. D: STP.

physaloides Guill. & Perr.

H: woody herb. D: ST.

*rosa-sinensis L.

H: shrub with red flowers. D: STP. V: Madureira & Martins 650

(COI).

sabdariffa L.

H: annual or perennial herb with white to yellow flowers. D: ST.

surattensis L.

H: annual herb with yellow flowers. M: 57. D: STP. V: Madureira &

Martins 148 (COI).

tiliaceus L. (. Fiteira )

H: shrub or tree with yellow flowers in coastal areas or near water-

courses. D: ST.

*MALVASTRUM A.Gray

*coromandelianum (L.) Garcke (Itoto-doido, Ototo, Ototo-home)

H: herb with pale yellow flowers. M: 158. D: STP. V: Madureira &

Marlins 193 (COI).

SIDA L.

acuta Burm.f. subsp. acuta (Itoto, Ototo, Ototo, Ototo-muala, Ototo-

-pequeno, Ototo-piquina)

H: herb or undershrub with yellow flowers, ruderal. M: 143; 175. D:

STP. V: Paiva 631 (COI), 827 (COI).

cordifolia L.

H: shrub with yellow or white flowers. D: ST.

javensis Cav.

pi/osa Retz.

veronicifolia Lam.

H: prostrate herb with yellow flowers. D: ST.

rhombifolia L. (Bobd-bobd, Catumba-jinji, Itoto, Ototo, Ototo-

-pequeno, Ototo-vento)

H: undershrub with yellow flowers, ruderal. D: STP. V: Figueiredo 73

(K); Paiva 637 (COI).

rigida (D.Don) D.Dietr.

H: herb or undershrub. D: ST.

urens L.

H: perennial herb or undershrub. D: ST.

*URENA L.

*lobata L. ( Ototo-fogo , Ototo-glandje, Ototo-glandji, Ototo-grande,

Ototo-ome, Ototo-vento)

H: perennial herb or undershrub with pink flowers, ruderal. M: 61; 91;

158; 175; 183. D: STP. V: Figueiredo 42 (K); Paiva 1418 (COX).

WISSADULA Medik.

rostrata (Schumach.) Hook.f. ( Ototo-vento , Ototo-vento)

amplissima (L.) R.E. Fries var. rostrata (Schumach.) R.E.Fries

H: undershrub. D: STP.

MARANTACEAE

CALATHEA G.Mey.

sp

H: herb with red-purple flowers. D: ST. V: Lejoly 93/619 (BRLU).

*MARAINTA L.

*arundinacea L.

H: herb cultivated for the fleshy rhizome. E: rhizome. D: STP.

THAUMATOCOCCUS Benth.

danicllii (Bennet) Benth. (Majunga, Mangungo)

H: rhizomatous herb. E: seed aril. M: 27; 60. D: STP. V: Madureira &

Martins 302 (COI); Paiva 626 (COI).

MELASTOMATACEAE

CALVOA Hook.f.

fconfertifnlia Exell

H: herb up to 0.4 m high, with pink flowers, in forest. D: ST.

Ecrassinoda Hook.f.

molleri Gilg

H: shrub up to 2.5 m high, with pink flowers, in forest and secondary

forest. D: ST. V: Figueiredo 199 (LISC).

grandifolia Cogn.

H: undershrub up to 1.2 m high, with pink flowers, in forest and sec-

ondary forest. D: STP. V: Figueiredo 202 (LISC); Oliveira 516

(BRLU).

hirsuta Hook.f.

henriquesii Cogn.

H: herb up to 0.3 m high, with white or pink flowers, in forest. D: STP.

V: Oliveira 569 (BRLU); Paiva 1079 (COI).

Eintegrifolia Cogn.

H: herb up to 0.4 m high, with pink flowers, in forest. D: ST.

sinuata Hook.f.

H: herb or shrub up to 1.2 m high, in forest. D: P. V: Figueiredo 160

(LISC).

DISSOTIS Benth.

barteri Hook.f. ex Triana

H: herb or shrub up to 2 m high, with pink flowers, in forest. D: P. V:

Stevart 259 (BRLU).

HETEROTIS Benth.

rotundifolia (Sm.) Jacq.-Fel.

H: decumbent herb rooting at nodes, with pink flowers, in forest. D:

STP. V: Stevart 242 (BRLU).

MELASTOMASTRUM Naudin

capitatum (Vahl) A. Fern. & R.Fern.

H: perennial herb or shrub up to 0.9 m high. D: ST. V: Oliveira 1042

(BRLU).

MEMECYLON L.

sp.

H: shrub or small tree. D: ST. V: Oliveira 1407 (BRLU).

TRISTEMMA Juss.

coronatum Benth.

H: herb or undershrub with pink or white flowers. D: P.

hirtum P.Beauv.

H: herb or undershrub with pink flowers. D: STP. V: Matos 7638

(LISC).

littorale Benth. subsp. biafranum Jacq.-Fel. var. biafranum

incompletwn sensu auct. pro parte non R.Br.

H: shrub. D: P.

Hittorale Benth. subsp. biafranum Jacq.-Fel. var. insulare Jacq.-Fel.

(Fid-vela)

incompletwn sensu auct. pro parte non R.Br.

H: shrub. M: 124; 129. D: ST. V: Madureira & Martins 503 (COI).

littorale Benth. subsp. littorale

H: shrub. D: P.

1 mauritianum J.F.Gmel. var. mildbraedii (Gilg) Jacq.-Fel.

mildbraedii Gilg ex Engl.

H: shrub with pink to mauve flowers. D: ST. V: Figueiredo 251

(LISC).

Fmauritianum J.F.Gmel. var. rozeiranum Jacq.-Fel. (Anjogo)

incompletwn sensu auct. pro parte non R.Br.

H: shrub with pink to reddish flowers. M: 173. D: ST. V: Paiva 61

(COI).

Emauritianum J.F.Gmel. var. thomense (J.H.P.B. Ferreira) Jacq.-Fel.

thomensis J.H.P.B. Ferreira

H: shrub with pink to mauve flowers. D: ST. V: Figueiredo 211

(LISC).

WARNECKEA Gilg

menibranifolia (Hook.f.) Jacq.-Fel. (Clossom-liso)

Metnecylon membranifolium Hook.f.

M. fernandianum Gilg ex Engl.

H: shrub or tree. M: 154. D: ST. V: Madureira & Martins 181 (COI).

memecylnides (Benth.) Jacq.-Fel. (Glele-nhame, Toni-fachico)

Memecylon memecyloides (Benth.) Exell

H: shrub or tree with blue flowers. M: 79. D: STP. V: Madureira &

Martins 202 (COI).

MELIACEAE

CARAPA Aubl.

procera DC. (Gogo, Gogo-vermelho, Gogo-vieme)

H: tree with white and pink flowers, in forest and plantations. M: 13;

126; 183. D: ST. V: Madureira & Martins 403 (COI).

Bothalia 41,1 (2011)

61

*CEDRELA P. Browne

*odorata L. (Cedrela, Cidlela, Cidrela, Pan-cebola, Po-cebola)

H: tree with small white flowers. M: 60; 123; 175. D: ST. V:

Madureira & Martins 441 (COI).

*MELIA L.

*azedarach L. ( Lilas-do-Cabo , Sicomoro)

H: tree with small purple or lilac flowers. D: STP.

*TOONA (Endl.) M.Roem.

*ciliata M.Roem. (Pau-alho, Po-alho)

H: tree with small white flowers. D: ST. V: Oliveira 1059 (BRLU).

TRICHILIA PBrowne

Fgrandifolia Oliv. ( Cola-de-macaco , Cola-macaco, Veludo)

H: tree, in forest. M: 22; 189. D: ST. V: Matos 7576 (LISC).

TURRAEA L.

vogelii Hook.f. ex Benth. ( Valaple , Vara-preta )

H: woody climber or scandent shrub. M: 22. D: STP. V: Matos 7630

(LISC).

MENISPERMACEAE

CHASMANTHERA Hochst.

dependens Hochst.

H: liana up to 20 m high, with yellow flowers, in cultivated ground.

D: ST.

STEPHANIA Lour.

dinklagei (Engl.) Diels

H: liana up to 20 m high, in forest. D: P.

TILIACORA Colebr.

sp.

H: liana. D: ST. V: Lejoly 98/268 (BRLU).

MORACEAE

*ARTOCARPUS J.R.Forst. & G.Forst.

*altilis (Parkinson) Fosberg (fruit: Castanha, Fluta. Fluta-pom. Fruta-

-pao\ tree: Arteira, Arvore-da-castanha, Arvore-do-pao, Pau-

castanha)

H: tree. E: fruits. M: 8; 61. D: STP. V: Paiva 257 (COI).

*heterophylla Lam. (fruit: Jaca: tree: Jaqueira, Pau-jaca )

H: tree. E: fruits. M: 27; 87. D: STP. V: Paiva 494 (COI).

*CASTILLA Cerv.

*elastica Sesse ex Cerv.

H: tree up to 18 m high. D: STP.

*CECROPIA Loefl.

*peltata L. ( Gofe , Gofe)

H: tree up to 25 m high, with yellow or green flowers. D: ST. V: Paiva

93 (COI).

FICUS L.

*carica L. (Figueira)

H: tree. E: fruits. D: ST. V: Oliveira 246 (BRLU).

chlamydocarpa Warb. ex Mildbr. & Burret subsp. chlamydocarpa

H: strangler fig, in forest. D: STP.

FchIamydocarpa Warb. ex Mildbr. & Burret subsp. fernandesiana

(Hutch.) C.C.Berg (Figo-obata)

fernandesiana Hutch.

H: strangler fig, in forest. D: ST. V: Lejoly 97/231 (BRLU, LISC).

*elastica Roxb.

H: tree. O. D: ST.

exasperata Vahl ( Meme , Po-lixa)

H: shrub or tree, in forest. D: ST. V: Figueiredo 215 (LISC).

kamerunensis Warb. ex Mildbr. & Burret (Mussanda, Mussunda)

H: tree or shrub, in forest. D: ST. V: Oliveira 36 (BRLU).

lutea Vahl

vogelii (Miq.) Miq.

H: tree or shrub, in forest. D: ST.

mucuso Welw. ex Ficalho (Figo-de-porco, Figo-ploco, Figu-plocu)

sidifolia Welw. ex Hiern

H: tree, in forest. D: ST. V: Paiva 1937 (COI).

*pumila L.

H: climber. D: ST. V: Oliveira 66/98 (BRLU).

sur Forssk. (Figo-todo, Figo-tordo, Todd)

capensis Thunb.

H: tree, in forest. E: fruits. D: STP. V: Paiva 1181 (COI), 1333 (COI).

thonningii Blume (Lemba-lemba, Mussanda)

annobonensis Mildbr. & Hutch.

H: tree or shrub, in forest. M: 57; 75; 87; 94. D: STP. V: Figueiredo

87 ( K).

IV1ESOGYNE Engl,

insignis Engl.

henriquesii Engl.

H: tree up to 16 m with greenish flowers, in forest and cultivated

ground. D: ST.

MILICIA Sim

excelsa (Welw.) C.C.Berg (Amoreira, Moreira, Mucamba-vleme,

Mulela)

Chlorophora excelsa (Welw.) Benth. & Hook.f.

H: large tree, in forest and plantations. M: 52; 87; 95; 153; 167. D:

STP. V: Oliveira 119 (BRLU).

MUSANGA C.Sm. ex R.Br.

cecropioides R.Br. (Gofe, Gofe, Gofe, Gofe-d'obo, Pau-sabrina)

H: tree up to 20 m high, with green flowers, in forest and secondary

forest. D: ST. V: Paiva 418 (COI).

TRECULIA Decne. ex Trecul

africana Decne. ex Trecul var. africana (Giquenge, Isa, Isa-quente,

Izaquente, Ogue, Opacala, Quicange, Quicuange, Zequentche)

H: tree up to 25 m high with small white flowers, in forest. E: fruits.

M: 18; 39; 40; 57; 121; 125; 158; 191. D: STP. V: Paiva 1209

(COI).

TRILEPISIUM Thouars

madagascariense DC.

H: tree, up to 30 m high, in plantations. D: ST.

MUSACEAE

*ENSETE Horan.

*ventricosum (Welw.) Cheesman

Musa ventricosa Welw.

D: ST.

*MUSA L.

* xparadisiaca L.

M: 68; 79; 103; 116; 126. D: STP.

*balbisiana Colla

rosacea Jacq.

D: ST.

MYRISTICACEAE

*MYRISTICA Gronov.

*fragrans Houtt. ( Noz-moscada )

H: small tree. E: fruits (spice). D: ST. V: Paiva 396 (COI).

PYCNANTHUS Warb

angolensis (Welw.) Warb. ( Cachao , Cashon, Cassa, Pau-caixao, Po-

-cassom, Po-casson)

H: tree, in forest, secondary forest and plantations. M: 51; 61; 79; 87;

90; 125; 167; 175. D: ST. V: Madureira & Martins 218 (COI).

STAUDTIA Warb.

Epterocarpa (Warb.) Warb. (Pau-vermelho, Vermelho, Vleme)

H: tree in forest and secondary forest. M: 13; 56; 125. D: ST. V:

Madureira & Martins 162 (COI).

MYRSINACEAE

ARDISIA Sw.

staudtii Gilg

Afrardisia cymosa (Baker) Mez

H: shrub or small tree with red flowers, in forest. D: STP. V: Matos

7440 (LISC).

62

Bothalia 41,1 (2011)

MAESA Forssk.

lanceolata Forssk. subsp. borjaeana (Henriq.) White ( Mutopa , Pau-

-cabra)

borjaeana Henriq.

H: shrub or small tree, with small white or yellowish flowers, in forest

and on roadsides. D: ST. V: Figueiredo 64 (K).

RAPANEA Aubl.

melanophloeos (L.) Mez

thomensis Exell

H: shrub or small tree, with small white or yellowish flowers, in forest.

D: ST. V: Matos 7507 (LISC).

MYRTACEAE

*EUCALYPTUS L Her.

*sp. (Cal ip to)

H: tree. D: ST. V: Oliveira 1326 (BRLU).

EUGENIA L.

*brasiliensis Lam. (Comoxama, Grumechame, Grumexava,

Grumixaba, Grumixama, Grumixameira, Grumixava)

dombeyi (Spreng.) Skeels

H: tree, in plantations. E: fruits. M: 99. D: ST. V: Madureira & Mar-

tins 669 (COI).

elliotii Engl. & Brehmer

H: shrub or small tree. D: ST.

*uniflora L. (Ameixa-do-Para, Cereja-de-Caiena, Pitanga,

Pitangueira)

H: shrub or tree. E: fruits. M: 173. D: STP. V: Matos 7690 (LISC).

*MELALEUCA L.

*leucadendron (L.) L.

H: shrub, in savanna. D: ST. V: Matos 7462 (LISC).

*MYRCIARIA O.Berg

*cauliflora (Mart.) O.Berg (Jabuticaba, Jabuticabeira)

H: small tree. E: fruits. D: ST.

*PSIDIUM L.

*cattleianum Sabine ( Gueva-quio )

littorale Raddi

H: shrub. E: fruits. M: 27. D: ST. V: Madureira & Martins 652 (COI).

*guajava L. (Gaiaba, Gaiava, Goiaba, Goiabeira, Gueva, Gaiaba)

H: shrub or tree. E: fruits. M: 19; 33; 57; 61; 67; 1 12; 135; 146; 1 75.

D: STP. V: Paiva 1520 (COI).

SYZYGIUM Gaertn.

*aromaticum (L.) Merr. & L.M. Perry (Cravinho)

H: tree with red flowers. E: flower buds (spice). D: ST.

guineense (Willd.) DC. ( Matazem , Matchanzoche, Matianzoche )

H: shrub or tree with white flowers, in forest. M: 22; 109; 125; 167;

183. D: ST. V: Madureira & Martins 328 (COI).

*jambos (L.) Alston (Jambo-jambo, Jamboeiro, Pau-jambre, Piala )

H: tree with whitish flowers, in plantations and on roadsides. E: fruits.

M: 5; 40; 79; 158; 175. D: STP. V; Paiva 1031 (COI).

staudtii (Engl.) Mildbr.

H: tree up to 30 m high, in forest. D: ST.

NYCTAGINACEAE

BOERHAVIA L

diffusa L. ( Erva-tostao , Fia-plocossom, Fia-plocssom)

coccinea Mill.

paniculata Rich.

H: perennial herb, ruderal. M: 77; 98; 1 14; 1 19; 149. D: STP. V: Paiva

384 (COI).

* BOUGAINVILLEA Comm, ex Juss.

*spectabilis Willd. (Buganvila)

H: climbing shrub with purple, red or white flowers. O. D; STP. V:

Paiva 287 (COI).

*MIRABILIS L.

*jalapa L. (Losa-bilanga, Rosa-bilanQa)

H: perennial herb up to ± 1 m high, with white or red flowers. E:

leaves. M: 4; 51; 61; 63; 98; 125; 134. O. D: ST. V: Paiva 699

(COI).

OCHNACEAE

Note: recent nomenclatural changes at generic level were not consid-

ered. The taxonomy follows AFPD (2009).

CAMPYLOSPERMUM Tiegh.

reticulatum (P.Beauv.) Farron var. reticulatum

Ouratea brunneopurpurea Gilg

H: shrub or small tree. D: STP.

vogelii (Hook.f.) Farron var. molleri (Tiegh.) Farron (Dumo, Dumo-

-vermelho, Pau-dumo)

Ouratea molleri (Tiegh.) Exell

H: shrub or tree with yellow flowers in forest. D: STP. V: Lejoly

95/020 (BRLU, LISC).

OCHNA L.

membranacea Oliv. (Pau-dumo, Po-dumo, Vilo)

H: shrub or tree in forest, secondary forest and savanna. D: ST. V:

Figueiredo 106 (K).

§multiflora DC.

H: shrub in forest. D: STP. V: Stevart 255 (BRLU).

OURATEA Aubl.

Enutans (Hiem) Exell

H: shrub with yellow flowers. D: P.

sp.

H: shrub. D: ST. V: Lejoly 98/267 (BRLU).

RHABDOPHYLLUM Tiegh.

arnoldianum (De Wild. & T.Durand) Tiegh.

Ouratea quintasii (Tiegh.) Exell

H: shrub, in forest. D: ST. V: Oliveira 1409 (BRLU).

OLACACEAE

HEISTERIA Jacq.

parvifolia Sm. (Nono, Pau-preto )

H: tree, in forest and plantations. D: STP. V: Lejoly 97/272 (BRLU).

OLAX L

gambecola Baill.

H: shrub with white or yellowish flowers. D: ST.

*XIMENIA L.

*americana L. ( Limon-ple )

H: tree, in savanna. E: fruits. M: 22. D: ST. V: Matos 7725 (LISC).

OLEACEAE

JASMINUM L.

bakeri Scott-Elliot (Codo-que-d’obd)

H: climber with white flowers, in forest. M: 79; 94. D: ST. V: Jojfroy

135 (BRLU).

fluminense Veil.

H: climber with white flowers, in forest. D: ST.

Ethomense Exell

H: climber with white flowers, in forest. D: ST. V: Oliveira 1508

(BRLU).

OLEA L.

capensis L. subsp. hochstetteri (Baker) Friis & PS. Green (Ipe, P6-

ipe)

hochstetteri Baker

H: shrub or tree with small white flowers, in forest. M: 22. D: ST. V:

Madureira & Martins 412 (COI).

capensis L. subsp. welwitschii (Knobl.) Friis & PS. Green ( Impe , Ipe)

welwitschii (Knobl.) Gilg & Schellenb.

H: shrub or tree with small white flowers, in forest. D: ST.

*europaea L. (Oliveira)

N: recorded by Exell (1973) based on Silva (1958) without specimen

citation.

ONAGRACEAE

LUDWIGIA L

erecta (L.) H.Hara (Aliba-giiia, Fia-guia)

H: annual herb with yellow flowers. M: 29; 46; 69; 88. D: ST. V:

Madureira & Martins 195 (COI).

Bothalia 41,1 (2011)

63

hyssopifolia (G.Don) Exell (Fia-placela)

H: annual herb, in secondary forest, savanna and on roadsides. D: STP.

V: Paiva 1197 (COI).

leptocarpa (Nutt.) H.Hara

H: herb with yellow flowers. D: ST.

octovalvis (Jacq.) P.H. Raven subsp. octovalvis

H: annual or perennial herb with yellow flowers, in secondary forest

and plantations. D: ST. V: Matos 7261 (LISC).

ORCHIDACEAE

(Stevart et al. 2000; Stevart & Oliveira 2001 )

AERANGIS Rchb.f.

Eflexuosa (Ridl.) Schltr.

H: epiphytic herb with white flowers, in lowland dry forest. D: ST.

ANCISTRORHYNCHUS Finet

capitatus (Lindl.) Summerh.

H: epiphytic herb with white flowers, in mid-elevation primary forest

and old secondary forest. D: P. V: Stevart 705 (BRLU).

crystalensis P.J.Cribb & Laan

H: epiphytic herb with white flowers, in mid-elevation primary rain-

forest. D: P. V: Stevart 1608 (BRLU).

metteniae (Kraenzl.) Summerh.

H: epiphytic herb with white flowers, lip green, in lowland forest and

plantations. D: STP. V; Stevart 638 (BRLU), 1212 (BRLU).

ANGRAECOPSIS Kraenzl.

Fdolabriformis (Rolfe) Schltr.

H: epiphytic herb, flower colour and habitat unknown. D: ST.

Ethomensis Stevart & P.J.Cribb

H; epiphytic herb, flowers greenish white, in old secondary forest. D:

ST. V: Oliveira 1999/138 (BRLU, STPH).

ANGRAECUM Bory

aporoides Summerh.

H: epiphytic herb with white flowers, in lowland and mid-elevation

forest. D: STP. V: Stevart 464 (BRLU), 667 (BRLU).

Eastroarche Ridl.

H: epiphytic herb, flower colour unknown, in old secondary forest. D:

ST.

distichum Lindl.

H: epiphytic herb, white flowers, habitat unknown. D: P.

doratophyllum Summerh.

H: epiphytic herb with white flowers, in mist forest. D: STP. N:

endemic to Gulf of Guinea. V; Stevart 329 (BRLU), 678 (BRLU).

infundibulare Lindl.

H: epiphytic herb with white flowers, habitat unknown. D: P.

sacciferum Lindl.

H: epiphytic herb with greenish white flowers, in mist forest. D: ST. V;

Stevart 104 (BRLU).

BOLUSIELLA Schltr.

iridifolia (Rolfe) Schltr. subsp. iridifolia

H: epiphytic herb with white flowers, in plantations. D: ST. V: Stevart

503 (BRLU).

talbotii (Rendle) Summerh.

H: epiphytic herb. D: STP. V: Primo & Stevart 77 (BRLU); Stevart

510 (BRLU).

BRACHYCORYTHIS Lindl.

Ebasifoliata Summerh.

H: terrestrial herb with purple flowers, in open vegetation. D: STP. V:

Stevart 363 (BRLU), 436 (BRLU, K).

BULBOPHYLLUM Thouars

acutibracteatum De Wild. var. acutibracteatum

H: epiphytic herb with yellow-orange flowers, in summit forest. D: P.

V: Stevart 420 (BRLU, K).

calyptratum Kraenzl.

H: epiphytic herb with greenish to yellowish flowers, habitat unknown.

D: P.

cochleatum Lindl. var. cochleatum

H: epiphytic herb with yellow to purple flowers, in montane forest. D:

ST. V; Stevart 652 (BRLU).

cochleatum Lindl. var. tenuicaule (Lindl.) JJ.Verm.

H: epiphytic herb with red to purple flowers, in montane open vegeta-

tion. D: ST. V: Stevart 417 (BRLU).

comatum Lindl. var. inflatum (Rolfe) J.J.Verm.

H: epiphytic herb with green inflorescence and purple flowers, in open

vegetation on ridges. D: STP. V: Stevart 462 (BRLU, K), 1203

(BRLU).

curvimentatum J.J.Verm.

H: pseudobulb epiphyte. D: ST.

falcatum (Lindl.) Rchb.f. var. falcatum

H: epiphytic herb with green to yellow flowers, in submontane forest.

D: STP. V: Rose 1023 (P); Stevart 71 (BRLU, K).

falcatum (Lindl.) Rchb.f. var. velutinum (Lindl.) J.J.Verm.

H: epiphytic herb with green to yellow flowers, in lowland and sub-

montane forest, also in coastal vegetation and plantations. D: STP.

V: Stevart 1211 (BRLU), 1221 (BRLU).

imbricatum Lindl.

H: epiphytic herb with violet and yellow flowers, in plantations. D: ST.

V; Stevart 489 (BRLU).

intertextum Lindl.

H: epiphytic herb with green or purple flowers, in submontane and

montane forests. D: STP. V: Stevart 681 (BRLU).

Elizae J.J.Verm.

H: epiphytic herb with green to yellow flowers, in submontane and

montane forests. D: ST. V: Stevart 1882 (BRLU).

Eluciphilum Stevart

H: epiphytic herb with green to yellow flowers, in plantations. D: ST.

V: Oliveira 1999/41 (BRLU).

maximum (Lindl.) Rchb.f.

H: epiphytic herb with yellowish to greenish flowers, in lowland and

submontane forests, also in plantations. D: STP. V: Stevart 397

(BRLU), 497 (BRLU).

mediocre Summerh.

H: epiphytic herb with orange, yellow or whitish flowers, in submon-

tane and montane forests. D: STP. N: endemic to Gulf of Guinea.

V: Stevart 622 (BRLU), 1890 (BRLU).

nigritianum Rendle

H: epiphytic herb with yellowish flowers, in plantations. D: ST. V; Ste-

vart 1158 (BRLU).

oreonastes Rchb.f.

H: epiphytic herb with orange to yellowish flowers, in ridge forests. D:

STP. V: Primo & Stevart 1 7 (BRLU); Stevart 407 (BRLU).

resupinatum Ridl.

H: epiphytic herb with yellowish to purple flowers, in lowland and

submontane forests. D: ST.

saltatorium Lindl. var. albociliatum (Finet) J.J.Verm.

H: epiphytic herb with greenish to brownish flowers with purple spots

and red pubescence, in mid-elevation ridge vegetation. D: P. V: Ste-

vart 422 (BRLU. K).

sandersonii (Hook.f.) Rchb.f. subsp. stenopetalum (Kraenzl.)

J.J.Verm.

H: epiphytic herb with yellowish to reddish flowers, in submontane

and montane forests, also in plantations. D: ST. V: Stevart 660

(BRLU).

CALANTHE R.Br.

Esylvatica (Thouars) Lindl. var. geerinckiana Stevart

H: terrestrial herb with white flowers and orange lip, in old plantations.

D: ST. V: Stevart 1194 (BRLU).

sylvatica (Thouars) Lindl. var. sylvatica

H: terrestrial herb with white flowers and violet lip, in submontane and

montane forests, and in secondary forest in old plantations. D: STP.

V: Stevart 338 (BRLU), 1195 (BRLU).

CHAMAEANGIS Schltr.

Ethomensis (Rolfe) Schltr.

H: epiphytic herb with yellowish to orange flowers, in montane for-

ests. D: ST. V: Stevart 659 (BRLU).

Evagans (Lindl.) Schltr.

H: epiphytic herb with yellowish to orange flowers, in lowland forests

and on ridges. D; P. V: Stevart 450 (BRLU, K).

CHEIROSTYLIS Blume

lepida (Rchb.f.) Rolfe

H: terrestrial herb with whitish flowers, in submontane and montane

forests. D: ST. V: Stevart 381 (BRLU).

CORYMBORKIS Thouars

corymbis Thouars

H: terrestrial herb with whitish and greenish flowers, in lowland for-

ests, and in secondary forest in old plantations. D: STP. V: Stevart

447 (BRLU), 665 (BRLU).

64

Bothalia 41,1 (2011)

CRIBBJA Senghas

confusa P.J.Cribb

H: epiphytic herb with orange flowers, in submontane and montane

forests. D: ST. V: Stevart 674 (BRLU).

1 pendula la Croix & P.J.Cribb

H: epiphytic herb with pale greenish flowers, in submontane forests,

also in secondary forest at mid-elevations. D: ST. V: Stevart 1168

(BRLU).

Ethomensis la Croix & P.J.Cribb

H: epiphytic herb with white flowers, in montane forests. D: ST. V:

Stevart 1228 (BRLU).

CYNORKIS Thouars

anacamptoides Kraenzl. var. ecalcarata P.J.Cribb

H: terrestrial herb with violet flowers, in montane forests. D: ST. V:

Stevart 385 (BRLU).

debilis (Hook.f.) Summerh.

H: terrestrial herb with purple-spotted flowers, in montane forests. D:

ST. V: Stevart 42 (BRLU, K).

gabonensis Summerh.

H: terrestrial herb with pale whitish flowers, in lowland to montane

forests, also on ridges. D: STP. V: Stevart 444 (BRLU, K), <529

(BRLU).

CYRTORCHIS Schltr.

arcuata (Lindl.) Schltr. subsp. variabilis Summerh.

H: epiphytic herb with white flowers and greenish to orange spur, in

coastal vegetation, in lowland secondary forest and plantations. D:

STP. V: Stevart 405 (BRLU, K ), 535 (BRLU).

henriquesiana (Ridl.) Schltr.

H: epiphytic herb with white to yellow flowers, in lowland secondary

forest and on ridges. D: P. V: Stevart 664 (BRLU).

monteiroae (Rchb.f.) Schltr.

H: epiphytic herb with white to yellow flowers, in lowland forest, sec-

ondary forest and plantations. D: STP. V: Stevart 205 (BRLU, K),

448 (BRLU).

ringens (Rchb.f.) Summerh.

H: epiphytic herb with white flowers and pale rose spur, in secondary

forest. D: ST.

DIAPHANANTHE Schltr.

‘acuta (Ridl.) Schltr.

H: epiphytic herb with whitish to yellowish flowers, in lowland and

mid-elevation forests. D: ST. V: Stevart 691 (BRLU).

‘papagavi (Rchb.f.) Schltr.

H: epiphytic herb, unknown flower colour, on rocky places in shade.

D: P.

pellucida (Lindl.) Schltr.

H: epiphytic herb with greenish to yellowish flowers, in lowland pri-

mary forest and old secondary forest. D: STP. V: Primo & Stevart

126 (BRLU); Stevart 1207 (BRLU).

rohrii (Rchb.f.) Summerh.

H: epiphytic herb with orange to yellowish flowers, in submontane and

montane forest and old secondary forest at high elevation. D: ST.

V: Stevart 494 (BRLU, K).

DINKLAGEELLA Mansf.

‘scandens Stevart & P.J.Cribb

H: terrestrial herb with white flowers, in swamp meadow. D: ST. V:

Stevart 1888 (BRLU).

DISPERIS Sw.

reichenbachiana Rchb.f.

H: terrestrial herb with purple to whitish flowers in submontane and

montane primary forest and mid-elevation secondary forest. D: ST.

V: Stevart 370 (BRLU).

thomensis Summerh.

H: terrestrial herb with white flowers in submontane and montane pri-

mary forest. D: ST. V: Stevart 48 (BRLU).

GENYORCHIS Schltr.

apetala (Lindl.) J.J.Verm.

H: epiphytic herb with whitish, reddish and yellowish flowers, in sec-

ondary forest and in plantations. D: P. V: Stevart 1220 (BRLU).

GRAPHORKIS Thouars

lurida (Sw.) Kuntze

H: epiphytic herb with brownish and yellowish flowers, in second-

ary forest and in plantations. D: STP. V: Stevart 302 (BRLU), 437

(BRLU).

HABENARIA Willd.

barrina Ridl.

H: terrestrial with whitish to greenish flowers, in secondary forest and

plantations. D: STP. V: Oliveira 1999/33 (BRLU); Stevart 427

(BRLU).

buettneriana Kraenzl.

H: terrestrial with greenish flowers, in secondary forest and planta-

tions. D: ST. V: Stevart 483 (BRLU).

‘letouzeyana (Szlach. & Olsz.) P.J.Cribb & Stevart

H: terrestrial with whitish flowers, in old secondary forest and lowland

primary forests. D: P. V: Stevart 631 (BRLU).

malacophylla Rchb.f. var. malacophylla

H: terrestrial with whitish to greenish flowers, in submontane and

montane primary forests. D: ST. V: Stevart 1223 (BRLU).

stenochila Lindl.

H: terrestrial with white flowers, in lowland vegetation. D: P. V: Ste-

vart 640 (BRLU).

thomana Rchb.f.

H: terrestrial with whitish to greenish flowers, in primary lowland to

montane forests. D: ST. V: Stevart 414 (BRLU); Paiva 1927 (COI).

LI PARIS Rich,

deistelii Schltr.

H: epiphytic herb with greenish to purple flowers, in secondary

and submontane forests. D: STP. V: Stevart 654 (BRLU), 1226

(BRLU).

epiphytica Schltr.

H: epiphytic herb with greenish to whitish flowers, in montane forests.

D: ST. V: Stevart 537 (BRLU).

goodyeroides Schltr.

H: terrestrial or lithophytic herb with greenish flowers, in old lowland

secondary forest. D: STP. V: Matos 7656 (BRLU); Stevart 625

(BRLU).

gracilenta Dandy

H: epiphytic herb with greenish and purple flowers, in submontane and

montane forests. D: STP. V: Stevart 672 (BRLU).

nervosa (Thunb.) Lindl.

H: terrestrial or epiphytic herb with greenish, yellowish, brownish or

purple flowers, in lowland secondary forest and in submontane for-

ests. D: STP. V: Stevart 196 (BRLU, K), 442 (BRLU).

platyglossa Schltr.

H: terrestrial herb with greenish, yellowish and purple flowers, in old

lowland secondary forest. D: P. V: Stevart 530 (BRLU).

rosseelii Stevart

H: epiphytic herb with greenish flowers, in secondary submontane for-

ests and in plantations. D: ST. V: Stevart 1188 (BRLU).

LISTROSTACHYS Rchb.f.

pertusa (Lindl.) Rchb.f.

H: epiphytic herb with white and green flowers, in secondary lowland

mid-elevation forests. D: P. V: Stevart 1216 (BRLU).

MALAXIS Sol. ex Sw.

maclaudii (Finet) Summerh.

H: terrestrial herb with reddish and purple flowers, in submontane for-

ests. D: ST.

MANN1ELLA Rchb.f.

gustavi Rchb.f.

H: terrestrial herb with brownish, pinkish or greenish flowers, in low-

land secondary forest, and in submontane and montane forests. D:

STP. V: Stevart 443 (BRLU), 642 (BRLU).

MICROCOELIA Lindl.

bulbocalcarata L.Jonss.

H: epiphytic herb with white and greenish flowers, in old mid-eleva-

tion secondary and primary forests. D: P. V: Stevart 682 (BRLU).

caespitosa (Rolfe) Summerh.

H: epiphytic herb with white and greenish to brownish flowers, in sub-

montane forests. D: ST. V: Primo & Stevart 47 (BRLU).

NERVILIA Comm, ex Gaudich.

bicarinata (Blume) Schltr.

H: terrestrial herb with yellowish to greenish flowers, in plantations.

D: STP. V: Stevart 686 (BRLU).

OBERONIA Lindl.

disticha (Lam.) Schltr.

H: epiphytic herb with whitish to yellowish flowers, in plantations. D:

ST. V: Stevart 1155 (BRLU).

Bothalia 41,1 (2011)

65

OECEOCLADES Lindl.

Hatifolia (Rolfe) Garay & P.Taylor

H: pseudobulb geophyte. D: ST.

maculata (Lindl.) Lindl.

H: terrestrial herb with whitish, yellowish to greenish flowers with a

red lip, in plantations and secondary forest. D: STP. V: Oliveira

1999/04 (BRLU, STPH); Stevart 636 (BRLU).

ugandae (Rolfe) L.A.Garray & P.Taylor

H: terrestrial herb with white and purple flowers, habitat unknown. D:

ST.

ORESTIAS Ridl.

micrantha Summerh.

H: terrestrial herb with white or pink flowers, in montane and submon-

tane forest. D: STP. V: Stevart 540 (BRLU), 642 (BRLU).

stelidostachya (Rchb.f.) Summerh.

elegans Ridl.

H: terrestrial herb with white or pink flowers, in montane and submon-

tane forest. D: STP. V: Stevart 210 (BRLU, K), 217 (BRLU).

PHAIUS Lour,

mannii Rchb.f.

H: terrestrial herb with whitish flowers with pink lip, in old planta-

tions. D: ST. V: Stevart 532 (BRLU).

PLATYLEPIS A. Rich,

glandulosa (Lindl.) Rchb.f.

H: terrestrial herb with whitish flowers, in plantations, lowland and

mid-elevation forests. D: STP. V: Stevart 501 (BRLU).

PODANGIS Schltr.

dactyloceras (Rchb.f.) Schltr.

H: epiphytic herb with white flowers, in plantations and submontane

forests. D: ST. V: Stevart 1197 (BRLU).

POLYSTACHYA Hook,

albescens Ridl. subsp. albescens

H: epiphytic herb with white and pink flowers, widespread in forest

and plantations. D: STP. V: Stevart 1178 (BRLU), 655 (BRLU).

Ealbescens Ridl. subsp. principensis Stevart

H: epiphytic or terrestrial herb with white and pink flowers, in open

vegetation and on ridges. D: P. V: Stevart 1227 (BRLU).

bifida Lindl.

farinosa Kraenzl.

H: epiphytic herb with greenish to yellowish flowers, in montane and

submontane forest. D: ST. V: Stevart 1889 (BRLU).

Ebiteaui P.J.Cribb, la Croix & Stevart

H: epiphytic herb with pink and white flowers, in plantations and sub-

montane forest. D: ST. V: Stevart 446 (BRLU).

Edisticha Rolfe

H: epiphytic herb with yellow flowers and red spots, in montane forest

and open vegetation. D: ST. V: Stevart 694 (BRLU).

Eexpansa Ridl.

H: epiphytic herb with white flowers with yellow to orange lip, in sec-

ondary forest and plantations. D: ST. V: Stevart 349 (BRLU).

fusiformis (Thouars) Lindl.

H: epiphytic herb with yellow to greenish flowers, in secondary and

submontane forest. D: STP. V: Stevart 423 (BRLU, K), 1204

(BRLU).

golungensis Rchb.f.

H: epiphytic herb with yellow flowers, in secondary forest and planta-

tions. D: ST. V: Stevart 1605 (BRLU).

moniquetiana Stevart & Geerinck

H: epiphytic herb with whitish to brownish flowers, in old mid-

elevation secondary forest and plantations. D: ST. V: Stevart 684

(BRLU).

mukandaensis De Wild.

H: epiphytic herb with yellow, brownish to greenish flowers with red

spots, in plantations. D: ST. V: Stevart 1191 (BRLU).

paniculata (Sw.) RolH: epiphytic herb with yellow to orange flowers,

in plantations. D: ST. V: Stevart 499 (BRLU).

Eparviflora Summerh.

H: epiphytic herb with yellow to brownish flowers, in submontane and

montane forest. D: STP. V: Stevart 484 (BRLU).

pobeguinii (Finet) Rolfe

H: epiphytic herb with pink flowers with lilac lip, in plantations or

near the sea. D: P. V: Stevart 1222 (BRLU).

polychaete Kraenzl.

H: epiphytic herb with yellow to greenish flowers with red spots, in

plantations. D: ST. V: Stevart 1156 (BRLU).

principia Stevart & P.J.Cribb

H: epiphytic herb with yellow orange flowers in old secondary forest.

D: P. V: Stevart 648 (BRLU).

pyramidalis Lindl.

H: epiphytic herb with yellow flowers with red spots, in plantations, on

ridges and in submontane forest. D: STP. V: Stevart 451 (BRLU,

K), 7225 (BRLU).

ridleyi Rolfe

H: epiphytic herb with brown, reddish or greenish flowers, in planta-

tions, and in submontane and montane forest. D: ST. N: endemic to

Gulf of Guinea. V: Stevart 1175 (BRLU).

Esetifera Lindl.

H: epiphytic herb with yellow flowers with red spots, in ridge vegeta-

tion. D: P. V: Stevart 1215 (BRLU).

tessellata Lindl.

H: epiphytic herb with greenish, yellowish, violet or purple flowers,

widespread in lowland and mid-elevation forest. D: STP. V: Stevart

454 (BRLU), 639 (BRLU).

Ethomensis Summerh.

H: epiphytic herb with yellow to purple flowers and white lip, in open

vegetation near summit. D: ST. V: Primo & Stevart 37 (BRLU).

RANGAERIS (Schltr.) Summerh.

trilobata Summerh.

H: epiphytic herb with brown to white flowers, in lowland and mid-

elevation forests. D: ST. V: Stevart 703 (BRLU).

RHIPIDOGLOSSUM Schltr.

Ebrevifolium Summerh.,

Diaphananthe brevifolia (Summerh.) Summerh.

H: facultative epiphytic herb with translucent whitish flowers, in

open vegetation on ridges in montane forest and in crater of Lagoa

Amelia. D: ST. V: Stevart 295 (BRLU).

curvatum (Rolfe) Garay

Diaphananthe curvata (Rolfe) Summerh.

H: epiphytic herb with greenish to yellowish flowers, in old secondary

lowland forest. D: P. V: Primo & Stevart 49 (BRLU).

rutilum (Rchb.f.) Schltr.

Diaphananthe rutila (Rchb.f.) Summerh.

H: epiphytic herb with whitish, greenish, brownish or violet flowers,

in submontane primary forest and in plantations. D: ST. V: Stevart

1606 (BRLU).

SOLENANGIS Schltr.

clavata (Rolfe) Schltr.

H: epiphytic herb with white flowers, in meadow of Lagoa Amelia. D:

ST. V: Stevart 1231 (BRLU).

scandens (Schltr.) Schltr.

H: epiphytic herb with whitish, yellowish to greenish flowers, in plan-

tations. D: ST.

STOLZIA Schltr.

elaidum (Lindl.) Summerh.

H: epiphytic herb with whitish to greenish flowers, in plantations and

on ridges. D: STP. V: Stevart 458, 521 (BRLU).

Epeperomioides (Kraenzl.) Summerh. subsp. thomensis (Stevart &

P.J.Cribb) Stevart, Droissart & Simo

thomensis Stevart & P.J.Cribb

H: epiphytic herb with green translucent flowers in evergreen mist for-

est. D: ST. V: Stevart 1891 (BRLU).

TRIDACTYLE Schltr.

armeniaca (Lindl.) Schltr.

H: epiphytic herb with white to orange flowers; habitat unknown. D:

ST.

Eaurantiopunctata P.J.Cribb & Stevart

H: epiphytic herb with white and orange flowers, in ridge forests. D: P.

V: Stevart 1224 (BRLU).

Eexellii P.J.Cribb & Stevart

H: epiphytic herb with whitish and greenish flowers, in submontane

and montane forests. D: ST. V: Stevart 708 (BRLU).

latifolia Summerh.

H: epiphytic herb with yellow flowers, in Pandanus forest on ridges.

D: P. V: Stevart 478 (BRLU).

Ethomensis P.J.Cribb & Stevart

H: epiphytic herb with white flowers, in plantations and submontane

forests. D: ST. V: Stevart 488 (BRLU, K).

tridactylites (Rolfe) Schltr.

H: epiphytic herb with whitish to orange flowers, widespread in forest

and plantations. D: STP. V: Stevart 1229 (BRLU), 1230 (BRLU).

66

Bothalia 41,1 (201 1 )

VANILLA Plum, ex Mill,

crenulata Rolfe

H: terrestrial herb with white, yellowish to orange flowers with pink

lip, in lowland forest and ridges. D: P. V: Stevart 647 (BRLU).

grandifolia Lindl.

H: terrestrial herb with yellowish flowers, on ridges. D: P. V: Stevart

328 (BRLU).

*planifoIia Andr.

H: terrestrial herb with yellowish flowers, cultivated or in old planta-

tions D: STP. V: Stevart 270 (BRLU).

ZEUXINE Lindl.

elongata Rolfe

H: terrestrial herb with white to greenish flowers, in secondary forest.

D: STP. V: Stevart 135 (BRLU, K), 334 (BRLU).

heterosepala (Rchb.f.) Geerinck

Hetaeria heterosepala (Rchb.f.) Summerh.

H: terrestrial herb with white and pink flowers, in submontane and

mid-elevation forests. D: STP. V: Stevart 428 (BRLU, K).

stammleri Schltr.

H: terrestrial herb with pink, white to greenish flowers, in lowland for-

ests. D: P. V: Stevart 633 (BRLU).

OXALIDACEAE

OXALIS L.

corniculata L.

H: annual or perennial herb with yellow flowers. D: STP. V: Paiva 567

(COI), 974 (COI).

*corymbosa DC. ( Fid-viola, Lamera, Madringueiro)

H: perennial herb with pink or purple flowers, in secondary forest,

plantations, and on roadsides. M: 33; 79. D: STP. V: Paiva 1386

(COI). 1945 (COI).

PANDANACEAE

PANDANUS Parkinson

candelabrum P.Beauv.

H: tree. D: P.

Ethomensis Henriq. (Paia-sela, Pau-esteira, Po-paia-cela)

H: tree ± 15 m high. D: ST. V: Paiva 1162 (COI).

PAPAVERACEAE

*ARGEMONE L.

*mexicana L. ( Cundu-de-muala-ve )

H: annual herb with yellow flowers. N: common name indicated by

Santo (1969) is same as for Acanthus montanus. D: ST.

PASSIFLORACEAE

ADENIA Forssk.

cissampeloides (Planch, ex Hook.) Harms ( Fissandja )

H: climber. M: 79; 95; 140; 183. D: ST. V: Madureira & Martins 486

(COI).

lobata (Jacq.) Engl.

H: climber with greenish yellow flowers. D: STP. V: Lejoly 97/230B

(BRLU, LISC).

*PASSIFLORA L.

*edulis Sims ( Macuja , Malacuja, Maracuja, Maracuja-pequeno,

Maracuja-roxo)

H: climber. E: fruits. M: 136. D: ST. V: Paiva 188 (COI).

*foetida L. ( Macuja-de-mato , Malacuja-blavo, Maracuja-do-mato,

Martirios-fetidos )

H: climber. E: fruits. M: 79; 98; 114; 136; 179. D: STP. V: Paiva 816

(COI).

*quadrangularis L. ( Grenadilha , Macuja, Malacuja, Maracuja,

Maracuja-grande)

H: climber. E: fruits. M: 42; 98. D: STP. V: Paiva 968 (COI).

PHYTOLACCACEAE

*PETIVERIA L.

*alliacea L.

H: perennial herb. D: ST. V: Oliveira 883 (BRLU).

PHYTOLACCA L.

dodecandra L'Her. ( Fia-mati , Matri)

dodecandra L’Her. var. apiculata (Engl.) Baker & C.H. Wright

H: climbing shrub, in secondary forest and on roadsides. M: 139. D:

ST. V: Paiva 1066 (COI).

PIPERACEAE

PEPEROMIA Ruiz & Pav.

fernandopoiana C.DC.

H: herb, creeping or trailing, mostly epiphytic, in forest. D: P. V: Jof-

froy 197 (BRLU).

molleri C.DC.

H: epiphytic herb, in forest. D: STP. V: Paiva 1399 (COI).

pellucida (L.) Kunth ( Alfabaca , Alfavaca, Fia-alfobaca)

H: erect or procumbent, annual herb, in forest. M: 57; 61; 79; 98; 114;

123; 149. D: STP. V: Paiva 1406 (COI), 1890 (COI).

retusa (L.f.) A.Dietr. var. retusa

retusa (L.f.) A.Dietr. var. mannii (Hook.f. ) Dull

H: perennial, creeping or epiphytic herb, in forest. D: ST. V: Matos

7565 (LISC).

tetraphylla (G.Forst.) Hook. & Am.

reflexa (L.f.) A.Dietr.

H: perennial, usually epiphytic herb, in forest. D: ST. V: Matos 7555

(LISC).

thomeana C.DC.

vaccinifolia C.DC.

H: creeping herb. D: ST.

vulcanica Baker & C.H. Wright

hygrophila Engl.

H: creeping herb. D: ST. V: Lejoly 94/528 (BRLU, LISC).

PIPER L.

capense L.f. ( Fia-boba-piquina )

H: shrub or undershrub, sometimes liana, in forest, secondary forest

and plantations. M: 43; 53; 80; 104; 154; 166. D: STP. V: Figuei-

redo 59 (K); Paiva 523 (COI).

guineense Schumach. & Thonn. ( Pau-pimenta , Pimenta-de-S.Tome,

Po-pimenta)

H: liana, in forest. E: fruits (spice). M: 51; 63; 108; 115. D: STP. V:

Madureira & Martins 463 (COI).

*nigrum L. (Pimento, Pimenteira )

H: climber. E: fruits (spice). M: 63; 152; 164. D: ST. V: Paiva 654

(COI).

umbellatum L. ( Fia-boba , Fia-bdba-d'obo, Fia-bobo, Folha-boba,

Folha-boba-branca)

Pothomorphe umbellata (L.) Miq.

H: shrub or woody herb, in forest, secondary forest and on roadsides.

M: 16; 51; 150; 159; 166; 167. D: STP. V: Figueiredo 40 (K);

Paiva 1421 (COI).

*PLANTAGINACEAE

*PLANTAGO L.

*major L.

H: herb. D: ST.

PLUMBAGINACEAE

PLUMBAGO L.

*auriculata Lam.

H: shrub with blue flowers. O. D: ST.

zeylanica L.

H: herb with white flowers, in secondary forest. M: 145; 183. D: ST.

V: Madureira & Marlins 498 (COI).

POACEAE

ACROCERAS Stapf

zizanoides (Kunth) Dandy

H: perennial grass, over 1 m high. D: ST.

7AGROSTIS L.

?tropica P.Beauv.

N: recorded by Exell (1944, 1973) based on literature citations; no

specimens known.

Bothalia 41,1 (2011)

67

ANTHEPHORA Schreb.

cristata (Doll) Hack, ex De Wild. & T.Durand

H: annual grass, up to 1 m high. D: ST.

AXONOPUS P.Beauv.

*compressus (Sw.) P.Beauv.

H: perennial grass up to 0.6 m high. D: ST.

flexuosus (Peter) C.E.Hubb. ex Troupin

H: perennial grass, up to I m high. D: P.

*BAMBUSA Schreb.

*vulgaris Schrad. ex J.C.Wendl. ( Bambu )

H: bamboo up to 25 m high. M: 22. D: STP V: Paiva 65 (COl).

*BRIZA L.

*maxima L.

H: annual grass, up to 1 m high. D: ST.

CENTOTHECA Desv.

lappacea (L.) Desv.

H: annual grass up to 0.6 m high, in secondary forest, plantations and

on roadsides. D: STP. V: Matos 7594 (LISC).

CHLORIS Sw.

pilosa Schumach.

H: annual grass up to 1 m high. D: STP.

pycnothrix Trin.

H: annual grass up to 0.5 m high. D: STP V: Lejoly 95/064 (LISC).

*COIX L.

*lacryma-j obi L. (Capim-de-Nossa-Senhora)

H: annual grass up to 1.3 m high. D: ST.

*CYMBOPOGON Spreng.

*citratus (DC.) Stapf (Belgata, Capim-cheiroso, Capim-do-Gabao,

Cha-do-Gabao, Cha-do-Principe, Fia-chalela, Folha-Gabao )

H: perennial grass to 2 m high. M: 22; 52; 76; 107; 150. D: STP V:

Madureira & Martins 335 (COI).

CYNODON Rich,

dactylon (L.) Pers.

H: perennial grass up to 0.4 m high. D: ST. V: Paiva 278 (COI).

DACTYLOCTENIUM Willd.

aegyptium (L.) Willd.

H: annual grass up to 0.6 m high. D: ST. V: Paiva 395 (COI).

DIGITARIA Haller

horizontalis Willd.

H: annual grass up to 0.6 m high. D: STP.

longiflora (Retz.) Pers.

H: annual grass up to 0.3 m high. D: ST. V: Paiva 402 (COI).

ECHINOCHLOA P.Beauv.

colona (L.) Link

H: annual grass up to 1.2 m high. D: ST.

ELEUSINE Gaertn.

indica (L.) Gaertn. ( Aliba-cagu , Queblancana)

H: annual grass up to 0.6 m high. M: 61; 79; 104; 135; 175. D: STP. V:

Madureira & Martins 303 (COI).

ERAGROSTIS Wolf

amabilis (L.) Wight & Am. var. amabilis

tenella (L.) P.Beauv. ex Roem. & Schult.

H: annual grass up to 0.5 m high. D: STP. V: Lejoly 97/289 (LISC).

domingensis (Pers.) Steud. ( Fia-vador, Folha-de-voador )

H: annual grass up to 1.2 m high, in coastal areas. D: ST. V: Paiva 810

(COI).

superba Peyr.

H: perennial grass up to 1.2 m high, ruderal. D: ST. V: Paiva 646

(COI).

ERIOCHLOA Kunth

fatmensis (Hochst. & Steud.) Clayton

nubica (Steud.) Hack. & Stapf ex Thell.

H: annual grass up to 0.7 m high. D: ST.

HETEROPOGON Pers.

contortus (L.) P.Beauv. ex Roem. & Schult.

H: perennial grass up to 1 m high. D: STP. V: Lejoly 93/656 (BRLU).

?*HORDEUM L

?*vulgare L.

N: recorded by Silva (1958) but no specimens are known.

HYPARRHENIA Andersson ex E.Loum.

diplandra (Hack.) Stapf

H: perennial grass up to 3 m high. D: ST.

rufa (Nees) Stapf

rufa var. major (Rendle) Stapf

H: perennial grass up to 2.5 m high. D: ST. V: Matos 7315 (LISC).

ISACHNE R.Br.

buettneri Hack.

H: perennial grass up to 0.5 m high. D: STP. V: Paiva 1182 (COI).

mauritiana Kunth

H: perennial grass up to 0.6 m high. D: ST.

LEPTASPIS R.Br.

zeylanica Nees ex Steud.

cochleata Thwaites

H: rhizomatous grass up to 1 m high. D: STP. V: Matos 7364 (LISC).

MEGASTACHYA P.Beauv.

mucronata (Poir.) P.Beauv.

H: annual or perennial herb up to 0.9 m high. V: Matos 7767 (LISC).

MELINIS P.Beauv.

minutiflora P.Beauv.

H: perennial grass up to 1.5 m high. D: ST.

repens (Willd.) Zizka

Rhynchelytrum repens (Willd.) C.E.Hubb.

H: annual or perennial grass up to 1.5 m high. D: P.

*OLYRA L

*latifolia L. (Fid-piala. Impiala)

H: perennial grass up to 4 m high. M: 180; 183. D: STP. V: Madureira

& Martins 276 (COI).

OPLISMENUS P.Beauv.

burmannii (Retz.) P.Beauv.

H: annual grass up to 0.6 m high, in forest and on roadsides. D: ST. V:

Paiva 1309 (COI).

hirtellus (L.) P.Beauv. ( Erva-colchao )

H: perennial grass up to 1 m high, in forest. D: STP V: Paiva 585

(COI), 1137 (COI).

?*ORYZA L.

?*sativa L. ( Arroz )

N: recorded by Silva (1958), but no specimens are known.

PANICUM L.

brevifolium L.

H: annual grass up to 0.6 m high. D: STP. V: Paiva 1184 (COI).

hoehstetteri Steud. ( Er\<a-de-senhora )

H: perennial grass up to 1 m high, in forest. D: ST. V: Paiva 1095

(COI).

maximum Jacq. ( Capim-gigante )

H: perennial grass up to 3 m high, on roadsides. D: STP. V: Paiva 964

(COI).

PASPALUM L.

conjugatum P.J.Bergius (Gutchim-duatche, Muala-sende-mom-sende-

-ope)

H: perennial grass up to 0.6 m high, in forest. M: 27; 52; 150. D: STP.

V: Paiva 1383 (COI).

paniculatum L. (Albargao, Aliba-cagu, Aliba-queblancana, Comida-

-de-cachequeque, Erva-cacho, Mansuensuen, Muambe)

H: perennial grass up to 1.2 m high. D: STP. V: Paiva 966 (COI).

scrobiculatum L.

orbiculare G.Lorst.

H: perennial grass up to 0.6 m high, in forest. D: ST. V: Lejoly 93/642

(BRLU).

vaginatum Sw.

H: perennial grass up to 0.6 m high. D: ST.

PENNISETUM Rich

laxior (Clayton) Clayton

Beckeropsis laxior Clayton

H: annual grass up to 0.8 m high. D: ST.

68

Bothalia 41,1 (2011)

PENNISETUM Rich, (cont.)

polystachion (L.) Schult.

H: annual or perennial grass up to 2 m high. D: STP. V: Paiva 514

(COI).

purpureum Schumach.

H: perennial grass up to 8 m high. D: ST. V: Paiva 737 (COI).

POA L.

annua L.

H: annual grass up to 0.3 m high. D: STP.

PSEUDECHINOLAENA Stapf

polystachya (Kunth) Stapf

H: annual grass up to 0.3 m high. D: ST.

ROTTBOELLIA L.f.

cochinchinensis (Lour.) Clayton

exaltata (L.) L.f.

H: annual grass up to 2.5 m high. D: ST.

*SACCHARUM L.

*officinarum L. ( Cana-de-agucar, Cana-such, Canam )

H: perennial grass up to 4 m high. D: STP. V: Paiva 1038 (COI).

SACCIOLEPIS Nash

africana C.E.Hubb. & Snowden

H: rhizomatous perennial grass up to 1.8 m high. D: ST.

?*SECALE L.

?*cereale L.

N: recorded by Silva (1958), but no specimens are known.

SETARIA P.Beauv.

barbata (Lam.) Kunth

H: annual grass up to 1 .5 m high. D: STP.

longiseta P.Beauv.

H: perennial grass up to 0.9 m high. D: STP.

megaphylla (Steud.) T.Durand & Schinz ( Erva-boi , Oga-oga, Uaga-

-uaga, Uga-uga )

chevalieri Stapf

H: perennial grass up to 3 m high. M: 29; 31; 62; 64; 129. D: STP. V:

Paiva 544 (COI), 1318 (COI).

SORGHUM Moench

arundinaceum (Desv.) Stapf (Mansuensue, Sorgo)

H: annual or perennial up to 4 m high. D: ST. V: Paiva 121 (COI).

* xdrummondii (Steud.) Millsp. & Chase

H: annual grass. D: P.

SPOROBOLUS R.Br.

molleri Hack.

H: annual grass up to 0.3 m high. D: STP.

pyramidalis P.Beauv.

H: perennial grass up to 1.6 m high. D: STP. V: Paiva 170 (COI).

tenuissimus (Mart, ex Schrank) Kuntze

H: annual grass up to 0.9 m high. D: ST.

virginicus (L.) Kunth

H: perennial grass up to 0.6 m high. D: STP.

STENOTAPHRUM Trin.

secundatum (Walter) Kuntze

H: perennial grass, in coastal areas. D: STP. V: Oliveira 66/99

(BRLU).

?*TRITICUM L.

?*aestivum L.

N: recorded by Silva ( 1958); no specimens known.

*ZEA L.

*mays L. (Mi, Milho )

H: annual grass up to 4 m high. E: fruit. M: 10; 64; 183. D: STP.

*ZOYSIA Willd.

*matrella (L.) Merr.

H: perennial grass. D: ST. V: Lejoly 98/270 (BRLU).

POLYGONACEAE

*ANTIGONON Endl.

Heptopus Hook. & Am. ( Brinco-de-casamento , Rosa-de-casamento)

H: climber with pink llowers. D: ST. V: Paiva 1521 (COI).

PERSICARIA (L.) Mill,

decipiens (R.Br.) K.L. Wilson

Polygonum salicifolium Brouss. ex Willd.

H: annual herb. D: ST. V: Matos 7500 (LISC).

senegalensis (Meisn.) Sojak

Polygonum senegalense Meisn.

H: perennial succulent herb, near water. D: ST.

RUMEX L.

abyssinicus Jacq.

H: perennial herb. D: ST. V: Matos 7366 (LISC).

crispus L.

H: herb, ruderal. D: ST. V: Lejoly 97/236 (BRLU, LISC).

PORTULACACEAE

PORTULACA L.

oleracea L. (Beldroega, Bodlega, Bodlega-piquina, Fia-bodlega, Fia-

-bondlega)

H: annual succulent herb, ruderal. E: leaves. M: 51; 135; 179. D: ST.

V: Paiva 563 (COI), 1283 (COI).

quadrifida L.

N: recorded by Exell (1973) based on Silva (1958) without specimen

citation.

*TALINUM Adans.

*triangulare (Jacq.) Willd. ( Bodlega-grande , Budelega)

H: perennial succulent herb with purple flowers. E: leaves. M: 185. D:

ST. V: Paiva 652 (COI).

*PROTEACEAE

*GREVILLEA R.Br. ex Knight

*robusta A.Cunn. ex R.Br. (Grevilia)

H: tree with orange flowers. O. D: ST. V: Oliveira 85 (BRLU).

*PUNICACEAE

*PUNICA Smook