FLORA OF SOUTHERN AFRICA
VOLUME 9
Editor G. Germishuizen
m
Part: Urticaceae
by I. Friis and K.L. Immelman
r
?8o
T A N I C A L
Digitized by the Internet Archive
in 2016
https://archive.org/details/floraofsoutherna09unse_0
FLORA OF SOUTHERN AFRICA
which deals with the territories of
SOUTH AFRICA, LESOTHO, SWAZILAND, NAMIBIA AND BOTSWANA
VOLUME 9
PART: URTICACEAE
by
I. Friis and K.L. Immelman
Scientific editor: G. Germishuizen
Technical editor: E. du Plessis
NATIONAL
Botanical
INSTITUTE
Pretoria
2001
Editorial Board
B.J. Huntley National Botanical Institute, Cape Town, RSA
R.B. Nordenstam Swedish Museum of Natural History, Stockholm, Sweden
W. Greuter Botanischer Garten und Botanisches Museum Berlin-
Dahlem, Berlin, Germany
Typesetting and page layout by S.S. Brink, NBI, Pretoria
Reproduction by 4 Images, P.O. Box 34059, Glenstantia, 0010 Pretoria
Printed by United Litho, P.O. Box 40900, 0007 Arcadia
© published by and obtainable from the
National Botanical Institute, Private Bag X101, Pretoria, 0001 South Africa
Tel. (012) 804-3200 Fax (012) 804-3211
ISBN 1-919795-55-3
CONTENTS
New taxa, new combinations and new statuses published in Volume 9, Part: Urticaceae
Introduction
Urticaceae
Urtica
Urera
Obetia
Laportea
Girardinia
Pilea
Pouzolzia
Parietaria
Forsskaolea
Droguetia
Didymodoxa
References
Index
Appendix:
Plan of Flora of southern Africa
FSA contributions in Bothalia
Flora of southern Africa : alphabetical list of published taxa
iv
v
1
2
7
8
10
14
15
17
21
23
26
29
32
35
A-l
A-3
A-4
NEW TAXA, NEW COMBINATIONS AND NEW STATUSES
PUBLISHED IN VOLUME 9, PART: URTICACEAE
None.
Date of publication: January 2001.
INTRODUCTION
This part was compiled in accordance with the Guide for contributors to the Flora of southern
Africa (compiled by Leistner, Ross & De Winter and available from the Editor, National Botanical
Institute, Private Bag X101, Pretoria, 0001 South Africa).
The maps show the distribution of the various taxa in the FSA region only. In the text, exotic
taxa are marked with an asterisk.
The numbering of the genus is according to De Dalla Torre & Harms in their Genera siphonoga-
marum (1900-1907), as adapted by Arnold & De Wet (1993, Plants of southern Africa: names and
distribution).
v
Urticaceae
URTICACEAE
by I. FRIIS* and K.L. IMMELMAN**
Herbs, shrubs, lianas or small trees, monoecious, dioecious or rarely polygamous, sometimes aniso-
phyllous, some genera with stinging hairs, sometimes with stiff non-stinging hairs which may be
curled or sharply hooked; epidermal cells usually with cystoliths, these punctiform, elongated or
linear. Leaves alternate or opposite, petiolate or sessile, simple or 3-5(-7)-lobed, usually tripli-
nerved from base, sometimes penninerved, margin entire or variously dentate; stipules usually pre-
sent, lateral or often intrapetiolar, often fused; cystoliths usually present. Inflorescences very varied,
mostly lax or condensed racemes, often with flowers in small cymose glomerules, or condensed
cymes in leaf axils, sometimes subtended by involucral bracts; inflorescence axis sometimes flat-
tened into a disc-shaped, fleshy receptacle. Flowers minute, unisexual or rarely bisexual, actino-
morphic or (especially female flowers) zygomorphic, with single whorl of tepals or rarely female
flowers naked; pedicel often articulated below perianth. Female flowers', tepals 3-5, free or united,
often very unequal, often accrescent after pollination, rarely absent; staminodes, if present, rudi-
mentary and scale-like; ovary superior, usually somewhat laterally compressed, sometimes asym-
metrical, unilocular, unicarpellate, placentation basal; stigma capitate, brush-like or linear. Male
flowers: tepals (1)2-5; stamens equalling tepals or solitary, indexed in bud; rudimentary ovary
often present. Fruit an achene, sometimes enclosed by persisting, accrescent perianth which some-
times becomes fleshy.
About 50 genera and 1 000 species; almost cosmopolitan but most numerous in the tropics. Most
occur in humid habitats, for example on forest floors or at forest margins, often along streams, but
some genera are adapted to arid areas ( Obetia and Forsskaolea).
There are 1 1 genera and 21 species represented in the FSA area. Boehmeria nivea (L.) Gaudich.
has been collected twice in KwaZulu-Natal (both specimens in NU) but, although not stated on the
specimens, the plants are almost certainly cultivated. This species is widely cultivated for fibre. It
has large (up to 140 x 80 mm) leaves with dense white felt beneath, similar to but usually not as
dense as in Pouzolzia mixta , the apical tooth is long and the leaf base decurrent, and the young
stems and leaves are covered with dense golden hairs.
A few species have been documented only by old collections, the southern African origin of
which is not beyond doubt. Such cases are discussed under the generic description of Laportea (no.
4) and the excluded names under Parietaria (no. 8).
la Plants with stinging hairs, at least on inflorescences and petioles; inflorescence a panicle
with flowers in minute cymose clusters along inflorescence axes:
2a Stipules free, lateral:
3a Trees; leaves alternate 3. Obetia (p. 8)
3b Herbs; leaves opposite 1. Urtica (p. 2)
2b Stipules fused, intrapetiolar or lacking:
* Botanical Museum and Library, University of Copenhagen, Gothersgade 130, DK-1123 Copenhagen K, Denmark.
** Formerly of the National Botanical Institute, Pretoria. Present address: Range and Forage Institute, Agricultural
Research Council, Private Bag X05, East Lynne, 0039 South Africa.
2
Urticaceae
4a Climbers; stems woody throughout; fruit surrounded by persisting fleshy perianth . .
2. Urera (p. 7)
4b Herbs; stems lignified at base only; fruit dry:
5a Female flowers with 3 almost completely united tepals; stinging hairs mostly longer
than 5 mm and very dense 5. Girardinia (p. 14)
5b Female flowers with 4 free, unequal tepals — 2 larger lateral ones and 2 smaller dor-
sal and ventral ones; stinging hairs mostly shorter than 5 mm, moderately dense to
sparse 4. Laportea (p. 10)
lb Plants completely without stinging hairs; inflorescence a ± sessile axillary cluster:
6a Leaves opposite, slightly to definitely anisophyllous; stipules completely fused, intra-
petiolar; female and male flowers with 3 tepals 6. Pilea (p. 15)
6b Leaves alternate or rarely a few on each plant opposite or leaves opposite throughout,
but then male flowers with 1 tepal and 1 stamen only, and female flowers naked;
stipules free, lateral or lacking (in Parietaria ):
7a Bracts below inflorescence prominent, covering at least three quarters of inflores-
cence and forming a ± fused, campanulate involucre:
8a Involucral bracts free for most of their length; leaves alternate, usually with numer-
ous hooked hairs above; shrubs or herbs of arid areas of Namibia and the Northern
Cape, Western Cape and western part of the Eastern Cape 9. Forsskaolea (p. 23)
8b Involucral bracts almost completely fused; leaves opposite or alternate, without or
with very few hooked hairs; herbs of forest margins and clearings in KwaZulu-
Natal and the Western and Eastern Cape 10. Droguetia (p. 26)
7b Bracts below inflorescence small and insignificant:
9a Shrubs; flowers unisexual; male flowers globular, 4(5)-merous; female flowers with
tubular perianth constricted below stigma 7. Pouzolzia (p. 17)
9b Annual or short-lived perennial herbs; flowers polygamous or unisexual; male flow-
ers tubular and with 4 stamens, or club-shaped and with 1 stamen; female flowers
with tubular perianth or naked:
10a Flowers polygamous, male and bisexual flowers tubular or campanulate, with 4
stamens; leaves entire; in summer-rainfall areas 8. Parietaria (p. 21)
10b Flowers unisexual; male flowers club-shaped, with 1 stamen only; female flowers
naked; leaves crenate or dentate or, if entire, then in winter-rainfall area
11. Didymodoxa (p. 29)
1973000 1. URTICA
Urtica L„ Species plantarum 2: 983 (1753); Benth.: 381 (1880); Engl.: 104 (1888); Hutch.: 181
(1967); Friis: 5 (1989a); Friis: 302 (1990); Friis: 620 (1993). Type species: U. dioica L.
Annual or perennial herbs, monoecious or dioecious. Stems with stinging and sometimes stiff
hairs, with punctiform cystoliths. Leaves opposite, petiolate, simple; stipules lateral, free, or (in
Africa outside FSA region) interpetiolar, fused. Inflorescences axillary, often paired in each leaf
axil, bisexual or unisexual, lax cymose panicles, flowers usually clustered in small cymose
glomerules. Flowers unisexual, 4-merous; tepals free. Female flowers in 2 slightly unequal pairs;
staminodes absent; ovary ovoid, laterally compressed, symmetrical; stigma sessile, penicillate.
Male flowers in 2 subequal to unequal pairs; rudimentary ovary present. Achenes enclosed in or
released from persisting perianth, lenticular, with a raised central area on each face.
Urticaceae
3
About 80 species, mostly in the northern temperate regions, with a few in the tropics and south-
ern temperate regions; three species are found in southern Africa, of which one is indigenous and
the other two are introduced weeds.
la Leaves usually as broad as or broader than long, slightly to deeply cordate; petiole
(20— )35— 1 1 2 mm long; achenes 1. 5-2.0 mm long 3. U. lobulata
lb Leaves usually longer than broad, cuneate, truncate or rarely cordate at base; petiole up
to 30(-50) mm long; achenes 1-2 mm long:
2a Achenes 1.0- 1.5 mm long; tepals of female flowers pubescent over whole exterior sur-
face; leaves (20— )50— 1 50 mm long, base truncate to subcordate, rarely cordate, teeth
never lobed; dioecious perennial 1. U. dioica*
2b Achenes 1 .5-2.0 mm long; tepals of female flowers glabrous except for marginal fringe
and single central stinging hair; leaves 22^15 mm long, base cuneate or rarely truncate,
teeth often lobed; monoecious annual 2. U. urens*
1 . *Urtica dioica L., Species plantarum 2:
984 (1753); N.E.Br.: 543 (1925). Type: from
Sweden, collector unknown (LINN; microfiche
in PRE, Nos. 1111.6!, 7!, 8!). Several possible
lectotypes exist, none of which have been so
designated.
U. dioica var. capensis Wedd.: 78 (1856). Syntypes:
Eastern Cape, Queenstown Div., Swart Kei River, Drege
2844 ( K!, P!, both as 8244, there seems to be misprint in
Wedd. l.c.); South Africa, further locality unknown, Zeyher
1547 (BM!, P!); Somerset Div., Bowker s.n. (K!).
U. eckloniana Blume: 142 (1856). U. dioica var. eckloni-
ana (Blume) Wedd.: 51 (1869). Type: Cap bon. sp., further
locality unknown, Ecklon 12.116.12 (L!, P! ).
U. eckloniana var. flavovirens Blume: 143 (1856). Type:
South Africa, not further indicated; no type material traced.
Erect perennial herb up to 1 m high, growing
from horizontal, ± woody rhizome, dioecious.
Stems almost always with dense stinging hairs;
stiff hairs present or absent. Leaves lanceolate,
(20— )50— 1 50 x (12-)25-90 mm, apical tooth
shorter to longer than broad, margin with 1 1-25
pairs of teeth, with stinging hairs scattered on
upper surface or absent, and on veins below,
with stiff hairs densely scattered below; petiole
(5—) 1 0—40 mm long, with dense stinging hairs,
with or without stiff hairs; stipules oblong to
broadly lanceolate, 2-6 x 1-3 mm, acute to
nearly obtuse, membranous, with up to 5 paral-
lel veins. Inflorescence lax, drooping, with
cymose clusters of subsessile flowers, up to 90
mm long; peduncle up to 15 mm long. Female
flowers with hairs as in key, 1.0- 1.5 mm long;
pedicel with stinging and stiff hairs. Male flow-
ers with stiff hairs, 1.25-1.5 mm in diameter.
Achenes ovoid, laterally compressed to lenticu-
lar, with a raised central area on each face,
smooth or minutely punctate, pale ochre.
An introduced weed in southern Africa,
recorded mainly from the Great Karoo of the
Western and Eastern Cape, with a single record
from the Northern Cape; originally from
Europe, where the species is much more vari-
able in general morphology and indumentum.
Southern African specimens almost uniformly
have a dense indumentum of stinging and stiff
hairs. Grows near rivers and along paths in for-
est, and on rocky mountain slopes, presumably
with some shade. Map 1 .
4
Urticaceae
Figure 1. — Urtica lobulata: 1, branch with pairs of inflorescences at each node, leaves with teeth unlobed, x 1 (Hoener
1715 , Archibald 4484 ); 2, leaf with lobed teeth, x 1 ( Dieterlen 59a)\ 3, female flower, x 10; 4, male flower, x 15; 5, fruit
with dark speckling, x 10 ( Archibald 4484). Artist: Gillian Condy.
Urticaceae
5
Vouchers: Acocks 14129 (PRE); Dieterlen
59b (PRE); Pappe PRE52599 (PRE); Theron 54
(PRE).
2. *Urtica urens L., Species plantarum 2:
984 (1753); Rendle: 242 (1917a); N.E.Br.: 543
(1925); Roessler: 6 (1967); Friis: 5 (1989a).
Type: from Sweden, collector unknown (LINN;
microfiche in PRE, No. 1111.5!).
Erect, ascending or procumbent annual herb
up to 0.5 m high, sometimes profusely branched
from base, monoecious. Stems with stiff hairs
when young, later glabrescent; stinging hairs
dense to sparse. Leaves elliptic, 22^-5 x 1 8-28
mm, apical tooth as long as or longer than
broad, margin with 9-13 pairs of long narrow
teeth which are often trilobed, with stinging
hairs scattered on upper surface and on veins of
lower surface, with stiff hairs absent or some-
times a few present on young leaves; petiole
(7-)30-50 mm long, with stinging and usually
with stiff hairs; stipules narrowly lanceolate, ±
1.5 x 0.5 mm, acute, pubescent. Inflorescence at
first dense, later becoming lax, cylindrical,
5-20 x 3-5 mm, bisexual, with male and female
flowers mixed, frequently with stinging and
stiff hairs; peduncle short. Female flowers with
hairs as in key, 1.75-2.0 mm long; pedicel with
stinging and stiff hairs. Male flowers 0.8-1. 0 mm
in diameter; pedicel ± 0.5 mm long. Achenes
smooth or minutely punctate, pale ochre.
An introduced weed in southern Africa, with
scattered records from Gauteng, Mpumalanga,
the Free State, Lesotho, KwaZulu-Natal,
Eastern, Western and Northern Cape and south-
ern Namibia; originally from Europe. A weed of
open disturbed places, in shade or sun. Map 2.
Vouchers: Acocks 3571 (PRE); Flora 1862
(PRE); Pegler 18 (PRE); Pole Evans & Smith
s.n. (PRE); Potts 3571 (PRE).
3. Urtica lobulata Blume in Museum
botanicum Lugduno-Batavum 2,10 & 11: 143
(1856); Wedd.: 84 (1856); N.E.Br.: 544 (1925).
Type: ‘Liliefontijn’, ‘Camisberge’, Drege 2987
(L, holo.!). Blume stated that his type material
came from ‘Leliefontein’, which would indicate
that specimens marked ‘ Urtica lobulata E.M.a.’
(BM!, K!, P! ), also stated to come from that
locality, would represent Drege collections
which are isotypes. It is uncertain where the
‘2987’ cited in the protologue and on the speci-
men comes from.
U. lobata Blume: 144 (1856). Type: Cap. bon. sp., further
locality unknown, Ecklon 11.10.9 (L).
U. meyeri Wedd.: 64 (1856); Wedd.: 44 (1869). Type:
locality unknown. Drege s.n. (Herb. Delessert, as stated by
Weddell).
U. burchellii N.E.Br.: 544 (1925). Syntypes: Cape,
Uitenhage, Zeyher s.n. (K !); Cape, Bathurst Div., near
Barville Park, Burchell 4092 (K!).
Erect, probably annual herb up to 0.8 m high.
Stems unbranched or branched from base only,
with dense stinging hairs; stiff hairs sometimes
present. Leaves broadly ovate to broader than
long, ( 1 0— )20— 1 70 x ( 1 0— )20— 1 80 mm, apical
tooth usually longer than broad, margin with
11-18 pairs of teeth which may be trilobed, or
with irregular lobes, upper surface with scat-
tered stinging hairs only, lower surface with
scattered stiff hairs, with dense stinging hairs
along veins; petiole (20— )35— 1 1 2 mm long, with
stinging hairs, with or without stiff hairs; stip-
ules pubescent, narrowly lanceolate, ±3x1
mm, acute. Inflorescence cylindrical, dense,
later becoming lax, erect to drooping, 20-100
Urticaceae
Figure 2. Urera trinervis: 1 , branch with inflorescences and branching aerial roots, x 1 (composite of Staner 1568 ,
Pegler 1533 and Wells & Edwards 41)\ 2, female inflorescence, x 2 (Leeuwenburg 71 87)\ 3, female flower, x 10
(Leeuwenburg 7187 ); 4, male flower, x 10 ( Breteler 1491)\ 5, fruit, x 10 ( Leeuwenburg 7187). Artist: Gillian Condy.
Urticaceae
7
mm long, apparently unisexual, with cymose
clusters of sessile or shortly pedicellate flowers;
peduncle 4-15 mm long, with stinging hairs.
Female flowers with scattered bristly hairs
which are denser along margin, with single dor-
sal stinging hair, ± 2 mm long; pedicel with
stinging hairs. Male flowers with or without
bristly hairs, up to 2 mm in diameter; pedicels
up to 1 mm long. Achenes smooth to rugose,
pale ochre to brownish, may have pale to dark
brown speckling. Figure 1 .
Indigenous species occurring in the eastern
Free State, KwaZulu-Natal Drakensberg and
Lesotho and the Eastem,Westem and Northern
Cape. Very variable; probably most closely
related to U. urens (no. 2), with which it is some-
times confused. Found among rocks, probably in
seepage areas or along streams, and occasional-
ly on the floor and margins of forests. Map 3.
Vouchers: Galpin 2496 (PRE); Henderson
617 (PRE); Hilliard & Bum 8998 (PRE);
Hoener 1715 (PRE); Shearing 367 (PRE).
1978000 2. URERA
Urera Gaudich. in Freycinet, Voyage autor du monde ... execute sur les corvettes de S.M. 1 ’Uranie
et la Physicienne Pillet-aine: 496 (1830); Benth.: 383 (1880); Engl.: 105 (1888); Hutch.: 181
(1967); Friis: 6 (1989a); Friis: 304 (1990); Friis: 80 (1991 ); Friis: 621 (1993). Type species: U. bac-
cifera (L.) Gaudich.
Woody climbers, often fixed to substratum by adventitious, axillary roots from stems, dioecious,
with stinging hairs on petioles and inflorescences, stiff hairs absent, with punctiform cystoliths.
Leaves alternate, petiolate, simple; stipules intrapetiolar, fused but with tips free. Inflorescence a
lax axillary raceme with flowers in small cymose glomerules. Female flowers with 4 somewhat
unequal tepals fused for three quarters of their length or more; ovary ovoid; stigma almost sessile,
penicillate. Male flowers 4- or 5-merous; rudimentary ovary present. Fruit an achene enclosed by
persisting 4-lobed to 4-lobulate perianth which enlarges and becomes fleshy and red in fruit.
Some 35 species, widespread in tropical Africa, on Madagascar, in tropical America and on the
Pacific Islands. One species occurs in southern Africa.
Urera trinervis (Hochst.) Friis & Immel-
man in Friis, Immelman & Wilmot-Dear in
Nordic Journal of Botany 7,2: 126 (1987); Friis:
6 (1989a); Friis: 81 (1991). Type: KwaZulu-
Natal, Umlaas River, July 1839, Krauss 1267
(TUB, holo.!; C, K, PRE, photo.!).
Elatostemma trinerve Hochst.: 88 (1845).
U. cameroonensis Wedd.: 97 (1869); Rendle: 261 (1917a);
Hauman: 185 (1948); Keay; 618 (1958); Letouzey: 76 (1968);
Agnew: 321 (1974); Friis: 549 (1985a). U. acuminata var.
cameroonensis (Wedd.) Leandri: 24 (1965) excl. spec. cit.
Type: Cameroun, Mt Cameroon, 1 500 m, Mann 2173 (K!).
U. woodii N.E.Br.: 96 (1911); N.E.Br.: 549 (1925).
Syntypes: Eastern Cape, in a cutting to the lighthouse near
Port St Johns. Pegler 1533 (BMl.^BOL!, K!); KwaZulu-
Urticaceae
Natal, near Umzinyati Falls, Wood 1803 (K!); without pre-
cise locality, Sanderson 594 (K!).
Robust climber, sometimes creeping among
rocks. Stems softly woody, with wide spongy
pith or hollow in centre, with large leaf scars on
young branches, up to 100 mm in diameter at
base. Bark longitudinally striate, brown to pur-
plish. Leaves elliptic to very broadly elliptic,
63-120 x 40-65 mm, apex acute to long- or
short-attenuate, base cuneate, truncate or slight-
ly cordate, margin entire, main veins prominent,
side veins scalariform, glabrous, often discolor-
ous; petiole 20-50 mm long, glabrous or with a
few large stinging hairs, often mounted on pro-
tuberances; stipules early caducous, ± 7 mm
long. Female inflorescence lax, paniculate, ± 20
mm long, with small cymose clusters of flow-
ers; peduncle with a few stinging hairs. Male
inflorescence lax, paniculate, ± 65 mm long,
with flowers densely clustered at intervals along
axes and branches of inflorescence; peduncle
mostly with a few stinging hairs. Female j low-
ers sessile; perianth cylindrical, with constric-
tion at apex, with 4 blunt but clearly marked
teeth, glabrous, ± 1 mm long. Male flowers on
pedicels up to 1 mm long, glabrous, ± 1.75 mm
in diameter. Achenes glabrous, 1. 5-2.0 mm
long, without markings, enclosed in persisting,
accrescent, fleshy, orange perianth. Figure 2.
In South Africa it occurs in KwaZulu-Natal
and as far south as Port St Johns in the Eastern
Cape, restricted to areas near the coast; also
from Ghana through central Africa to Ethiopia,
through East Africa to South Africa. A climber
in coastal or riverine forest, especially near
margins and in clearings, or sprawling in forest
margins and among boulders. Map 4.
Vouchers: Ross 1872 (PRE); Vahrmeijer 497
(PRE); Vahrmeijer & Tolken 312 (PRE); Wells
& Edwards 41 (PRE); Wood 1872 (PRE).
1979000 3. OBETIA
Obetia Gaudich., Voyage autour du monde execute pendant les annes 1836 & 1837 sur la corvette
La Bonite - Botanique, Atlas: t. 82 (1844); Benth.: 382 (1880); Engl.: 106 (1888); Hutch.: 182
(1967); Friis: 221 (1983); Friis: 10 (1989a); Friis: 306 (1990); Friis: 85 (1991); Friis: 620 (1993).
Type species: O. ficifolia (Savigny) Gaudich.
Small to medium-sized shrubs or small trees, completely deciduous in dry season, dioecious.
Stems herbaceous when young, wood soft and juicy, pith often wide; cystoliths dot-like or slightly
elongated. Leaves alternate, petiolate, simple or lobed; stipules free, lateral. Inflorescence a
bracteate panicle, in axils of fallen leaves or sometimes of young leaves, developing before leaves
appear; male inflorescence much smaller than female. Female flowers 4-merous, outer pair of
tepals smaller than inner pair, enlarging and becoming thinly membranous in fruit; ovary ovoid,
with sessile penicillate stigma. Male flowers 5-merous, regular; rudimentary ovary present.
Achenes compressed, enclosed in persisting membranous perianth.
Eight species in eastern and southern Africa, Madagascar, the Mascarenes and Aldabra. In
southern Africa there are two species, which are closely related.
Urticaceae
9
Stipules 2. 5-4.0 mm broad, with scattered stiff hairs and often with stinging hairs along
midrib; lamina often bullate; Namibia 1.0. carruthersiana
Stipules 1.0-2. 5 mm broad, entirely glabrous; lamina not bullate; North-West, Gauteng,
Northern Province, Mpumalanga, Swaziland, KwaZulu-Natal and the Eastern Cape
2. O. tenax
1. Obetia carruthersiana (Hieni) Rendle,
Flora of tropical Africa 6,2: 245 (1917a); Roess-
ler: 5 (1967); Coates Palgrave: 121 (1977). Type:
Angola, Golungo Alto, Delamboa, Welwitsch
6267 ( BM!, K!, P!, SAM!).
Urticastrum carruthersianum Hiern: 987 (1900). Lapor-
tea carruthersiana (Hiern) K.Schum.: 463 (1902).
Urera engleriana Dinter: 54 (1909), name invalidly pub-
lished. Based on the following original material: Nambia,
Waterberg, near the spring; Teufelsbacher Schlucht, Farm
Dobra; Spitskoppies; Otavi; etc., no collector given, ?
Dinter. No original material has been traced, but from cir-
cumstantial evidence, this is certainly O. carruthersiana.
O. australis Engl.: 424, t. 2 (1914); Engl.: 54, t. 35 (1915).
Syntypes: Angola, Huila, Antunes 241 ( B ! ; COI!); Namibia,
Hereroland, Gaub, Dinter & Engler 1913\ Namibia, Otavi,
Kalkbergen, Dinter 629\ Nambia, Waterberg Distr.,
Okosongo-Muingo. flat-topped sandstone mountain, no col-
lector given (possibly also Dinter ?); Namibia, Otjim-
bingue. Use Fischer s.n.\ Nambia, near Okahandja, mica-
ceous schist, Dinter s.n. (Friis has verified that only the
Antunes collection survives).
Shrub or small tree up to 3 m high. Branches
densely covered with long stinging hairs, longi-
tudinally striate, brown, with large leaf scars,
wood soft, fibrous, with wide, spongy pith or
hollow centre. Leaves with lamina broadly
ovate in outline, up to 140 x 190 mm, some-
times shallowly lobed, base cordate, margin
with 16-29 pairs of broad irregular teeth, usual-
ly bullate, upper surface with scattered stinging
and stiff' hairs, the latter sometimes very dense,
lower surface with scattered bristly hairs, sting-
ing hairs concentrated along veins; petiole up to
100 mm long, with short stiff hairs and stinging
hairs; stipules ovate, 6-11 (-18) x 2. 5-4.0 mm,
apex acuminate, with scattered stiff hairs and
stinging hairs along midrib. Inflorescence a
laxly and profusely branching panicle, forming
a tangled mass ± 150 mm long, on short pedun-
cle. Female flowers single or in small clusters,
shortly pedicellate, glabrous, ± 1 mm long.
Map 5. — • Obetia carruthersiana
▲ O. tenax
Male flowers not seen. Achenes laterally com-
pressed, rugose over whole surface, 1.0- 1.5
mm long, brown, papery perianth persisting and
enlarging to length of ± 2 mm.
In southern Africa it is restricted to a few
localities in northern and central Namibia; also
in southwestern Angola. Occurs on rocky slopes,
probably on calcrete. Map 5.
Vouchers: Esterhuyse 436 (PRE); Gibson
30 (PRE, WIND); Strey 2681 (PRE); Viljoen
WIND4361 (PRE, WIND).
2. Obetia tenax (N.E.Br.) Friis in Kew
Bulletin 38,2: 226 (1983); Friis: 85 (1991).
Type: KwaZulu-Natal, Inanda, Wood 3837 (K!.
holo., vide Friis 1983: 226; BOL!, without no.).
Urera tenax N.E.Br.: t. 1748 (1888); N.E.Br.: 548 (1925);
Coates Palgrave: 121 (1977).
Shrub or small tree up to 5 m high. Branches
densely covered with long stinging hairs or
10
Urticaceae
glabrous, purplish brown to grey, often longitu-
dinally striate, gnarled with large leaf scars,
wood soft and fibrous, with wide spongy pith or
hollow in centre. Leaves with lamina broadly
ovate in outline, 35— 1 30(— 170) x 35—1 15(— 190)
mm, sometimes shallowly lobed, base usually
truncate, rarely deeply cordate, margin with
1 2-22 pairs of broad irregular teeth, upper sur-
face with scattered stinging and stiff hairs,
lower surface with scattered stiff hairs and
stinging hairs along veins; petiole up to 120 mm
long; stipules lanceolate, 4—8 x 1-2.5 mm, with
long-acuminate apex, glabrous. Inflorescence of
profusely branched panicles, up to 100 mm
long; peduncle with stinging hairs. Female
flowers single or in small clusters along axes
and branches of inflorescence, shortly pedicel-
late, glabrous, ± 1.25 mm long. Male flowers in
denser inflorescences, ± 2 mm in diameter, usu-
ally with stinging hairs on tepals. Achenes later-
ally compressed, rugose over whole surface, ±
1.5 mm long, ochre to brown, papery perianth
persisting and enlarging to length of ± 2 mm.
In southern Africa it is found in the Northern
Province, Mpumalanga, North-West, Gauteng,
Swaziland, KwaZulu-Natal and the Eastern Cape;
also in Zimbabwe and Mozambique. Grows on
rocky hillsides and in forest margins. Map 5.
Vouchers: Coetzee 238 (PRE); Leistner,
Thom & Gillham 3312 (PRE); Mogg 12380
(PRE); Van der Schijff 4985 (PRE); Ward 5830
(PRE).
1980000 4. LAPORTEA
Laportea Gaudich. in Freycinet, Voyage autor du monde ... execute sur les corvettes de S.M.
l’Uranie et la Physicienne Pillet-aine: 496 (1830), nom. cons.; Benth.: 383 (1880); Engl.: 106
(1888); Chew: 200 (1965); Hutch.: 183 (1967); Chew: 111 (1969); Friis: 156 (1981); Friis: 203
(1985b); Friis: 15 (1989a); Friis: 308 (1990); Friis: 88 (1991); Friis: 620 (1993). Type species: L.
canadensis (L.) Wedd. (lecto., vide Chew 1969: 115).
Fleurya Gaudich.: 496 (1830); Benth.: 382 (1880); Engl.: 106 (1888); Hutch.: 183 (1967). Type species: F. spicata
Gaudich. (lecto. of sect. Fleurya , vide Chew 1969: 115).
Annual or perennial herbs or shrubs, monoecious or dioecious. Stems with numerous stinging
hairs. Leaves alternate, petiolate, simple, serrate; cystoliths linear or punctiform; stipules intrapeti-
olar, fused almost to apex. Inflorescence unisexual, paniculate or appearing spike-like because of
very reduced branches, may have interrupted ‘spikes’. Female flowers pedicellate; pedicel some-
times winged, either laterally or on dorsal and ventral side (as defined by symmetry of flower);
tepals 4, unequal, lateral pair usually much larger than median pair, median pair of unequal size;
staminodes absent; ovary asymmetrical, ovoid, laterally compressed; stigma sessile, filiform or
deeply trifid with filiform branches. Male flowers usually pedicellate, 4- or 5-merous; rudimenta-
ry ovary present. Achenes compressed, often with characteristic sculpturing on sides, consisting of
a ring-like ridge enclosing a rugose surface, stipitate and shed without the persisting perianth (pre-
viously placed in sect. Laportea) or sessile and shed with the persisting perianth (previously placed
in sect. Fleurya (Gand.) Chew).
About 22 species, pantropical, extending into the temperate regions of North America and East
Asia; three species with two subspecies in southern Africa. L. interrupta (L.) Chew has not been
found in the FSA region for more than 150 years, and has therefore been excluded from this treat-
ment. Chew (1969: 148) records one specimen from southern Africa but this is based on a misiden-
tification of the specimen cited [Schlechter 2791 , collected at Durban, June 1893 (BM!)], which is
L. peduncularis subsp. latidens, as verified by us. However, at Kew there is another specimen of
this species from the FSA region, marked: ‘Cap. bon. spei, Roxburgh' , collected about 1794. L.
interrupta is authentically recorded from Mozambique, and could occur in southern Africa, but no
Urticaceae
other specimen was seen by either author. The distinguishing characters are the trifid stigma (one
branch longer than the other two) and the inflorescence which resembles an interrupted spike
owing to reduction of the lateral branches.
la Leaf margins each with 4 — 7( — 9) equal-sided triangular teeth; lamina usually with white
blotches with a stinging hair in the centre of each blotch; stinging hairs mounted on
long, slender protuberances with long-decurrent bases 3. L. grossa
lb Leaf margins each with (5-) 15-40 unequal-sided teeth; lamina without white blotches;
petiole and young stems with stinging hairs hardly ever mounted on protuberances and
(if so) these never with decurrent bases;
2a Leaf margins with 25-40 pairs of teeth; plant erect, up to 2 m high; male flowers with
4 tepals; female flowers in fruit with pedicels greatly expanded at right angles to
plane of flattened achene (laterally winged) 1 . L. alatipes
2b Leaf margins with 5-25 pairs of teeth; plant ascending or scrambling or sometimes
erect when young; male flowers with 5 tepals; female flowers in fruit with pedicels
slightly expanded parallel to the plane of the flattened achene (dorsiventrally winged)
2. L. peduncularis
1. Laportea alatipes Hook.f. in Journal of
the Linnean Society, Botany 7: 215 (1864);
Wedd.: 79 (1869); Rendle: 252 (1917a); Hau-
man: 194 (1948); Robyns: 71, t. 6 (1948); Keay:
620 (1958); Letouzey: 117, t. 18 (1968); Chew:
124 ( 1969); Agnew: 321 (1974); Friis: 16 (1989a);
Friis: 93 (1991). Type: Cameroun, Cameroon
Mountain, 7 000 ft, Mann 1973 (K!, holo.).
Urticastrum alatipes (Hook.f.) Kuntze: 635 (1891).
Fleurya alatipes (Hook.f.) N.E.Br.: 547 ( 1925).
Robust annual or short-lived perennial herb,
monoecious. Stems soft, erect or decumbent, up
to 2 m long, with sparse to dense stinging hairs
when young, glabrescent, stiff hairs sometimes
present. Leaves broadly lanceolate to ovate,
100-160 x 58-104 mm, apical tooth longer than
broad, base cordate, margin with 25-40 pairs of
small close-set teeth, surface often bullate, with
dense or sparse stinging hairs scattered on upper
surface and concentrated along veins on lower
surface, stiff hairs sparse, usually on lower sur-
face; petiole 40-125 mm long, with both stinging
and stiff hairs; stipules deciduous, linear-lanceo-
late to lanceolate, 3-8 x ± 2.5 mm, with long-
acuminate apex, with bristly hairs. Inflorescences
loosely branching panicles, with clusters of flow-
ers; female in upper axils, up to 180 mm long;
male in axils below female ones, up to 100 mm
long. Female flowers with pedicels broadly, lat-
erally winged, stinging hairs dense to sparse.
bristly hairs usually in two lines along peduncle
or sometimes over whole peduncle. Male flowers
4-merous, usually with stinging hairs, stiff hairs
present, ± 0.5-1. 0 mm in diameter. Achenes
shortly stipitate, laterally compressed, ± 2 mm in
diameter, centre of flattened sides irregularly
rugose and pale brown, margin smooth and
black-brown, dispersed without perianth.
In southern Africa it is found in two discrete
areas: from the Northern Province, and from
southwestern KwaZulu-Natal and the Eastern
Cape; also widespread in East and West tropical
Africa. Found in undergrowth of forest, usually
near water. Map 6.
12
Urticaceae
Vouchers: Codd 7859 (PRE); Kerfoot 8375
(PRE); Moll 3002 (PRE); Van Warmelo 223
(PRE); Vorster 279 (PRE).
2. Laportea peduncularis (Wedd.) Chew
in Gardens' Bulletin, Singapore 21: 201 (1965);
Chew: 152 (1969); Friis: 205 (1985b); Friis: 19
(1989a); Friis: 93 (1991). Type: KwaZulu-
Natal, Port Natal (Durban), at Yellowwood
River, Kachu, Drege s.n., marked ‘ Urtica pe-
duncularis E.M.’ (G!, lecto., vide Chew 1965:
201, K!).
Annual herb, monoecious, or dioecious by
abortion. Stems soft, decumbent or scrambling,
sometimes rooting at lower nodes, up to 1.5 m
long, with few to fairly many ± 1 mm long
stinging hairs, stiff hairs sometimes present
when young. Leaves lanceolate to ovate,
17-125 x 11-77 mm, apical tooth longer than
broad, base rounded, truncate or rarely cordate,
with long stinging hairs scattered above and on
veins of lower surface, with stiff hairs; petiole
6-115 mm long, with stiff hairs and sometimes
with stinging hairs, these occasionally mounted
on short, slender protuberances without decur-
rent base; stipules as in L. alatipes but 4-10 x
1-2 mm. Inflorescence paniculate, mostly with
flowers in small cymose clusters; female in
upper axils, usually repeatedly dichotomously
branched, or with one branch taking the lead
over the other, up to 110 mm long; peduncles
glabrous or with stinging hairs; male in lower
axils, up to 35 mm long, often with two or more
glomerules of equal size terminating the
branches; peduncles as in female. Female f low-
ers pedicellate, 1-2 mm long. Male flowers 5-
merous, shortly pedicellate; pedicel with a nar-
row dorsiventral wing, 1.75-3.0 mm in diame-
ter. Achenes almost sessile, laterally com-
pressed, centre of flattened sides warted, with
smooth circular marginal ridge, ±1.5 mm long,
dispersed with perianth attached.
Occurs in the Northern Province, Mpu-
malanga, Swaziland, KwaZulu-Natal and the
Eastern and Western Cape; also in Mozambique
to tropical East Africa. Grows in forest, often in
clearings or along the margins.
Two subspecies are recognised:
Leaf margin with (5-) 12-25 pairs of small
teeth; lamina ovate, base rounded,
truncate or rarely cordate, never cu-
neate, indumentum usually of stiff
hairs and stinging hairs mounted on
small protuberances
2a. subsp. peduncularis
Leaf margin with 5— 8(— 12) pairs of broad
triangular teeth; lamina ovate to
rhomboid, base cuneate or some-
times truncate, usually glabrous and
without mounted stinging hairs . . .
2b. subsp. latidens
2a. subsp. peduncularis.
Fleurya peduncularis Wedd.: 75 (1869).
Urtica mitis Hochst.: 88 (1845), nom. nud. F. mitis Wedd.:
183 (1854), nom. nud. F. capensis var. mitis Wedd.: 118
(1856). F. peduncularis var. mitis (Wedd.) Wedd.: 76 (1869).
F. mitis (Wedd.) N.E.Br.: 546 (1925). Syntypes: Cape
Province, Galgebosch, forest, Drege s.n., marked ‘ Urtica
mitis E.M.a.' (BM!, K!, P!); Strandfontein & Matjiesfontein,
Drege s.n., marked ‘Urtica mitis variet. E.M.b.’ (K!, P!);
Doukamma. forest, Drege s.n., marked ‘ Urtica mitis variet.
E.M.a.’ (SAM!); Natal, Yellowwood River, Rachu, Drege
s.n., marked ‘Urtica mitis E.M.b.’ (not the same as the type
of L. peduncularis) (K!, P!); Van Staadens Rivier, Drege s.n.,
marked ‘ Urtica mitis variet. E.M.c.’ (not traced).
F. capensis auct.: Wedd.: 117, t. la, figs 7, 8 (1856);
Rendle: 249 (1917a); excl. spec, typ., not of Wedd. (1854).
Urticaceae
13
In South Africa it is found from KwaZulu-
Natal to the Eastern Cape, more or less in
coastal areas; also in Mozambique. Found in
dune forest or coastal scrub. Map 8.
Vouchers: Flanagan 1225 (PRE); Lambinon
82/213 (PRE); McClean 122 (PRE); Strey 5108
(PRE); Watmough 410 (PRE).
3. Laportea grossa (Wedd.) Chew in Gar-
dens’ Bulletin, Singapore 21: 200 (1965). Type:
Eastern Cape, Uitenhage Div., Zeyher 3865 (P!,
lecto., vide Chew 1965: 200; FI, SAM!).
Fleurva grossa Wedd.: 119, t. 1A, figs 1-3 (1856);
N.E.Br.: 545 (1925).
L. caffra Chew: 155 (1969). Type: Western Cape,
Outeniquaberg Distr., near George, forest margin, Schlechter
2331 (A, BOL!, BM!, BR!, GRA!, K!, P!, PRC, PRE!, S!,
SAM!, STE!, UPS!,Z!).
Leaves ovate, indumentum usually of stiff
hairs and stinging hairs mounted on small pro-
tuberences, base rounded, truncate or rarely cor-
date, never cuneate, margin with (5-) 12-25
pairs of small teeth.
Occurs in southern Africa in the Northern
Province, Mpumalanga, Swaziland, KwaZulu-
Natal and the Eastern and Western Cape; also in
tropical East Africa as far north as northern
Tanzania. In forest from the Drakensberg down
to sea level. Map 7.
Vouchers: Meeuse 9765 (PRE); Rogers 1966
(PRE); Scheepers 588 (PRE); Strey 8531 (PRE);
Theron 1145 (PRE); Van Wannelo 5159/7 (PRE).
2b. subsp. latidens Friis: 206 (1985b); Friis:
94 (1991). Type: Mozambique, Inhaca Island,
near sea level, Mogg 30920 (K, holo.!; LMU!).
L. interrupta sensu Chew (1969), p.p. quoad spec.
Schlechter 2791 (BM!), non Urtica interrupta L.
Leaves ovate to rhomboid, usually glabrous
and without mounted stinging hairs, base
cuneate or sometimes truncate, margin with
5— 8(— 12) pairs of broad triangular teeth.
Annual herb, monoecious. Stems soft, juicy,
sometimes decumbent and rooting at nodes, up
to 1 m high, glabrous or young parts with few
stinging hairs mounted on long protuberances
and with decurrent bases, deeply striate. Leaves
broadly ovate, sometimes broader than long,
(35— )65— 130 x (35— )45— 1 20 mm, each side with
4— 7 (-9) large, equal-sided, triangular teeth, api-
cal tooth longer than broad, base cuneate to
truncate, with whitish blotches, upper surface
glabrous except for a single stinging hair in cen-
tre of each white blotch, lower surface with a
few stinging hairs along 3 main veins especial-
ly near petiole; petiole 30-125 mm long, some-
times pink, with stinging hairs up to 1.5 mm
14
Urticaceae
long, on protuberances up to 2.5 mm long with
long, usually decurrent bases on petiole, stiff
hairs absent; stipules minute, early caducous.
Female inflorescence paniculate, regularly
branching, up to 80 x 30 mm; peduncle and
axes with sparse long, stinging hairs. Male
inflorescence paniculate, branching less, up to
50 mm long; peduncle glabrous or with a few
stinging hairs. Female flowers 1.5 mm long.
Male flowers 4- or 5-merous, 2 mm in diameter.
Achenes with circular ridge near edge, smooth,
± 1.75 mm long, dispersed with perianth.
Occurs in Mpumalanga, KwaZulu-Natal and
the Eastern Cape, with one record from the
Western Cape. Erroneously indicated for
Mozambique by Chew (1969: 159), the speci-
mens cited being L. mooreana (Hiem) Chew.
Found in undergrowth of forest and near
streams, also in waste places where it is moist
and shady. Map 9.
Vouchers: Acocks 13572 (PRE); Galpin
5681 (PRE); Heinecken K92 (PRE); Marais
783 (PRE); Nicholson 1513 (PRE).
1983000 5. GIRARDINIA
Girardinia Gaudich. in Freycinet, Voyage autor du monde ... execute sur les corvettes de S.M.
l’Uranie et la Physicienne ... Pillet-aine: 498 (1830); Endl.: 283 (1837); Benth.: 384 (1880); Engl.:
107 (1888); Hutch.: 183 (1967); Friis: 143 (1981); Friis: 12 (1989a); Friis: 306 (1990); Friis: 85
(1991); Friis: 621 (1993). Type species: G. leschenaultiana Decne. (= G. diversifolia).
Erect annual (or very short-lived perennial) herbs, monoecious, or dioecious by abortion, with
long stinging hairs on all aerial parts; cystoliths punctiform. Leaves alternate, petiolate, simple or
mostly variously divided, margin coarsely serrate; stipules intrapetiolar, fused almost to apex.
Inflorescence unisexual, of dense, elongated cymes in axils of upper leaves. Female flowers ses-
sile, with 3 almost completely fused tepals ± enclosing ovary, sometimes with 1 minute mem-
branaceous free tepal, without staminodes; ovary ± asymmetrical, reflexed, laterally compressed,
with sessile, filiform stigma. Male flowers pedicellate, 4- or 5-merous; rudimentary ovary present.
Achenes laterally compressed, rugose.
Two species distributed in the mountains of the Old World tropics from West Africa to South
China and Taiwan; one species found in the Northern Province, Mpumalanga and Swaziland. This
genus has the longest stinging hairs in the family, but apparently the effect is less severe than that
of some other genera, for example Laportea.
Girardinia diversifolia (Link) Friis in Kew
Bulletin 36,1: 145 (1981); Friis: 13 (1989a); Friis:
86 (1991). Type: plant from India, cultivated in
Berlin Botanical Garden, not preserved. Neotype
(see Friis, ibid.): India, Sikkim, 5-7 000 ft.
Hooker s.n. (K!, neo., vide Friis 1981: 145; iso-
neo. B!,C!,CAL!,L!).
Urtica diversifolia Link: 385 (1822) non Blume (1825).
U. heterophylla D.Don: 59 (1825), nom. illegit. non Vahl
(1790). Type: Nepal, Wallich 4603 (K!).
G. heterophylla Decne.: 151, t. 153 (1844); Letouzey:
1 10, t. 17 (1968). Based on U. heterophylla Vahl: 76 (1790),
nom. illegit. superfl., including the type of U. palmata
Forssk. (1775). Syntypes: Yemen, Forsskal s.n. (C!);
Rheede-tot-Drakenstein, Hort. Ind. Malab. 2: t. 41 (1679).
G. condensata (Steud.) Wedd.: 181 (1854); Agnew: 323
(1974). U. condensata Steud.: 260 (1850). Type: Ethiopia,
Mt Sellueda near Adua, Schimper Series III, 1888 (BM!,
BR!, FI!, G!, K!, L!, LE!, P [in Herb. Steudel], P!, S!).
G. palmata Blume: 158 (1856), nom. illegit. non U. pal-
mata Forssk. (1775). Type: India, Nilgiri Hills, Leschenault
54 (P!, also holotype of G. leschenaultiana Decne. 1844).
G. vahlii Blume: 158 (1856), nom. illegit. superfl. Based
on U. palmata Forssk. (1775).
Urticaceae
15
Erect, probably annual herb up to 1 m high.
Stems with stinging and stiff hairs. Leaves broad-
ly ovate in outline, deeply 3-5-lobed, or unlobed
near base of plant or on side shoots, 111-170 x
90-170 mm, apical tooth longer than broad, base
cuneate to truncate, margin with 19-24 pairs of
broad teeth, each tooth sometimes with a single
adaxial lobe, densely covered with long stinging
hairs 3-7 (-9) mm long, these scattered on upper
surface, concentrated along veins on lower sur-
face, stiff hairs also present; petiole 55-90 mm
long, with stinging and adpressed stiff hairs; stip-
ules lanceolate, ±8x3 mm, fused, except for the
free, attenuate apices. Female inflorescence dense-
ly cylindrical, 1 or 2 per node, 35-75 mm long;
peduncles densely covered with long, hard, pun-
gent stinging hairs up to 4 mm long. Male inflo-
rescence not seen in material from FSA region,
thin and spike-like, 1 or 2 per node, 50-80 mm
long; peduncle covered with stiff hairs. Female
flowers sessile, without free fourth tepal, ± 2 mm
long, with thick unilateral dorsal ridge. Male
flowers on pedicels ± 1 mm long; tepals without
subapical horn-like appendage. Achenes ovoid,
laterally compressed, strongly rugose over whole
surface, 3-4 mm in diameter, dark brown when
mature.
Map 10. — Girardinia diversifolia
From a few scattered localities in the North-
ern Province, Mpumalanga and Swaziland;
widespread from East and West tropical Africa,
Madagascar, Yemen, India to East China, Tai-
wan and Indonesia. Grows in shady places, often
in disturbed habitats. Map 10.
Vouchers: Strey & Schlieben 8612 (PRE);
Theron 2105 (PRE); Thorncroft 2050 (PRE).
1984000 6. PILEA
Pilea Lindl., Collectanea botanica: t. 4 (1821), nom. cons.; Benth.: 384 (1880); Engl.: 108 (1888);
Hutch.: 185 (1967); Friis: 648 (1988); Friis: 25 (1989a); Friis: 557 (1989b); Friis: 310 (1990); Friis: 96
(1991); Friis: 622 (1993). Type species: P. muscosa Lindl., nom. illegit. (- P. microphylla (L.) Liebm.).
Annual or perennial herbs, monoecious. Stems ± juicy, without stinging hairs. Leaves opposite,
petiolate or subsessile, the two of a pair often unequal, margin usually serrate; cystoliths linear; stip-
ules intrapetiolar, completely fused thus appearing as one. Inflorescences unisexual or bisexual,
dense cymes or ± lax panicles. Female flowers with 3 free tepals, one usually considerably larger,
usually cucullate or with dorsal horn-like appendage, the 2 lateral ones usually much smaller, flat,
rarely cucullate or with hom-like appendage; staminodes 3, scale-like, inflexed, later reflexing to
eject the achene; ovary symmetrical, erect, with sessile penicillate stigma. Male flowers 3-5-mer-
ous; tepals with a dorsal hom-like appendage; rudimentary ovary present. Achene ovate, ± smooth.
About 250 species, almost pantropical in distribution. One species is found in southern Africa.
There is also a single specimen in PRE of P. microphylla (L.) Liebm. (= P. muscosa Lindl.) from
Durban, growing between bricks in a garden. This species is widely distributed in tropical Africa
(Friis 1989b: 596) and is probably an adventive there, but there is no evidence that the species has
16
Urticaceae
Figure 3. — Pilea rivularis: 1 , branch with sessile inflorescences and unequal leaves, x 1 ; 2, two pairs of fused intrapeti-
olar stipules, x 2; 3, female flower, x 18; 4, 6 scale-like staminodes in female flower, x 18; 5, male flower, x 18; 6, fruit, x
18 (Scheepers 590). Artist: Gillian Condy.
Urticaceae
17
become naturalised in the FSA area. It is a small, prostrate or creeping, mat-forming, anisophyllous
plant with leaves ±3x2 and 1.75 x 1.25 mm respectively. It is originally from North and Central
America, but possibly came to South Africa via East Africa.
Pilea rivularis Wedd. in Archives du
Museum National d’Histoire Naturelle, Paris
9,1: 266 (1856); Letouzey: 163, t. 27 (1968);
Friis: 29 (1989a); Friis: 565 (1989b); Friis: 98
(1991). Type: Comoro Ils, Boivin s.n. (P, holo.!).
P. ceratomera Wedd.: 132 (1869); Rendle: 269 (1917a);
Hauman: 203 (1948); Agnew: 323 (1974). Type: Fernando
Po, Clarence Peak, Mann 626 (K!).
P. worsdellii N.E.Br.: 550 (1925). Syntypes: Northern
Province, Houtbosch, Schlechter 4740 (K!); Northern
Province, Soutpansberg, Worsdell s.n. (K!).
Perennial herb, up to 0.6 m high. Stems erect,
usually unbranched, slightly anisophyllous,
arising from creeping stolons which may root at
nodes, glabrous, fleshy. Leaves broadly ovate,
55-75 x 40-60 mm, terminal tooth longer than
broad, base truncate to slightly cordate, margin
with 10 or 11 pairs of teeth, lower surface
glabrous, upper surface glabrous or with scat-
tered broad translucent hairs; petioles distinctly
unequal, 22-38 (short) or 35-55 mm (long),
glabrous; stipules broadly ovate to broader than
long, 4—6 x 4-7 mm, apex obtuse, base cordate,
membranous. Inflorescence a dense, sessile,
cymose cluster of flowers in each axil, 7-10
mm long. Female flowers pedicellate, with 3
somewhat unequal boat-shaped tepals each with
a prominent dorsal horn; horn on largest tepal ±
as long as tepal, glabrous except for sessile
glands, ± 1.5 mm long; staminodes present.
Male flowers pedicellate, 3-merous; tepals
equal, glabrous except for sessile glands, ± 2
mm long; rudimentary ovary absent. Achenes
shortly stipitate, ± 2 mm long, smooth, pale
ochre to brown with pale narrow rim round
apex and both margins. Figure 3.
Occurs in the Northern Province and Mpuma-
langa, with one record from the Eastern Cape;
widespread in the mountains of West and East
tropical Africa, the Comoro Islands and Mada-
gascar. Found in forests and in forest margins, in
damp places and on streambanks. Map 11.
Vouchers: Kluge 1604 (PRE); Kluge 2501
(PRE); Mohle 266 (PRE); Scheepers 590
(PRE); Schlechter s.n. (PRE, 21063 in Tvl Mus.
Herb).
1992000 7. POUZOLZIA
Pouzolzia Gaudich. in Freycinet, Voyage autor du monde ... execute sur les corvettes de S.M.
l’Uranie et la Physicienne Pillet-aine: 503 (1830); Benth.: 387 (1880); Engl.: 112 (1888); Hutch.:
188 (1967); Friis & Jellis: 587 (1984); Friis: 46 (1989a); Friis: 317 (1990); Friis: 108 (1991); Friis:
623 (1993). Type species: P. laevigata (Poir.) Decne. (= Parietaria laevigata Poir., as lP. levigata')-
Erect perennial herbs or shrubs, sometimes annual, monoecious. Leaves alternate, petiolate,
entire or dentate; cystoliths punctate; stipules free, lateral. Inflorescence of compact axillary bisex-
18
Urticaceae
Figure 4.— Pouzolzia mixta: 1, branch apex showing sessile inflorescences, x 1; 2, female inflorescence, with pair of
free stipules, x 10; 3, male flower, x 10 ( Immelman 635). Artist: Gillian Condy.
Urticaceae
19
ual glomerules. Female flowers sessile, with an indefinite number of completely fused tepals
enclosing the ovary; staminodes absent; stigma filiform with lateral papillae, deciduous. Male flow-
ers pedicellate, 4- or 5-merous; ovary rudimentary or absent. Achenes ovoid, erect, completely
enclosed in persistent membranous perianth.
About 70 species, mainly in the tropics of the Old World; in southern Africa there are two wide-
spread species.
Leaves entire, usually densely grey-white-felted below; shrub or tree up to 5 m high 1 . P. mixta
Leaves with dentate margins, not felted below; herb up to 1 m high or scrambling ... 2. P. parasitica
1. Pouzolzia mixta Solms in Sitzungs-
Bericht der Gesellschaft Naturforschender
Freunde zu Berlin 1864: 21 (1864); Solms: 188
(1867); Friis & Jellis: 590 (1984); Friis: 48
(1989a); Friis: 109 (1991). Type: border between
Sudan and Ethiopia, Fazugly, Cienkowsky s.n.
(B, holo., destroyed; LE!, lecto., vide Friis &
Jellis 1984: 590).
P. hypoleuca Wedd.: 227 (1869); Rendle: 291 (1917a);
N.E.Br.: 551 (1925); Roessler: 6 (1967). Type: Mozam-
bique, Miramballa, Kirk s.n. (Kl).
Shrub or small tree up to 3(-5) m high. Stems
longitudinally striate, greyish or reddish brown,
with wide spongy pith or hollow centre, wood
soft, inflorescence and leaf scars prominent
especially on young stems, with dense curved or
arachnoid hairs when young, glabrescent.
Leaves deciduous, ovate to lanceolate, 48-115 x
25-75 mm, apex attenuate-acute, base cuneate
to truncate, upper surface roughly velvety or
scabrid, with dense hairs, lower surface densely
to slightly white-felted, felt sparse on veins;
petiole 15-22 mm long, pubescent to white-felt-
ed; stipules ovate to lanceolate, membranous,
red-brown, apex acute to acuminate, midrib pro-
nounced, hairs scattered or restricted to midrib
and margin. Inflorescence appearing with young
leaves, sessile in leaf axils or scattered along
twigs. Female flowers sessile, pubescent, 2-3
mm long. Male flowers pedicellate, pubescent,
1.5-2. 5 mm long. Achenes ± compressed fusi-
form, smooth, shiny, dark brown, ± 2.5 mm
long, sometimes dispersed with persisting mem-
branaceous perianth. Figure 4.
Occurs in Namibia, Botswana, North-West,
the Northern Province, Gauteng, Mpumalanga,
Swaziland and KwaZulu-Natal; also in the
southern Arabian Peninsula, Ethiopia, East trop-
ical Africa and Angola. Found on rocky hillsides
in open bushveld, often in sandy soil. Map 12.
Vouchers: Fourie M010 (PRE); Oliver,
Schweickerdt & Verdoorn 337 (PRE); Rogers
30020 (PRE); Van der Schijff 3605 (PRE);
Verdoorn 2258 (PRE).
2. Pouzolzia parasitica (Forssk.) Schweinf.
in Bulletin de l’Herbier Boissier 4, App. 2: 145
(1896); Rendle: 293 (1917a); Hauman: 215
(1948); Robyns: 84 (1948); Keay: 763 (1958);
Letouzey: 194 (1968); Agnew: 325 (1974); Friis
& Jellis: 593 (1984); Friis: 51 (1989a); Friis:
111 (1991). Type: Yemen, Hadie, on walls of
coffee plantations, Forsskal s.n. (C, holo.!).
Urtica parasitica Forssk.: 160 (1775).
20
Urticaceae
Figure 5 — Parietaria debilis: 1, branch, x 1; 2, male flower, x 20; 3, female flower with developing fruit, x 20; 4,
bisexual flower, x 20 ( Merxmiiller & Giess 28148). Artist: Gillian Condy.
Urticaceae
21
Margarocarpus procridioides Wedd.: 204 (1854).
Boehmeria procridioides (Wedd.) Blume: 204 (1856).
P. procridioides (Wedd.) Wedd.: 412 (1857); N.E.Br.: 551
(1925). Syntypes: Eastern Cape, Pondoland, between Mtata
(Umtata) and Umsimubu (Port St John’s) Rivers,
1 000-2 000 ft, Drege s.n., marked ‘ Urtica ?procridioides
E.M.’ (BM!, K!, P!); Umsimubu (Port St John’s) River,
Drege s.n., marked 1 Urtica ?procridioides E.M.b.’ (P!).
Erect or ascending perennial herb up to 1 m
high, sometimes scrambling, often with slen-
der shoots from a woody base, up to 50 mm in
diameter. Stems often rooting at lower nodes,
with long, dense, patent, often pale golden yel-
low hairs, glabrescent. Leaves lanceolate to
ovate, 60-125 x 40-70 mm, apical tooth
longer than broad, base broadly cuneate, mar-
gin with 10-23 pairs of teeth, both surfaces
with fairly dense scattered hairs, these denser
on veins of lower surface; petiole slender,
(20-)40-80 mm long, densely patent-hairy;
stipules prominent, lanceolate, 5-10 x 1. 5-3.0
mm, membranous, persistent, apex attenuate
and recurved, base sessile and cordate, with
long hairs on margin and midrib. Female flow-
ers hairy, ± 2 mm long. Male flowers with
perianth deeply 4-lobed, acuminate, hairy, ±
2.5 mm long. Achenes fusiform, 1.5-2. 5 mm
long, enclosed in hairy accrescent perianth;
seed pale cream, smooth, glistening.
Occurs in the Northern Province, Mpu-
malanga and Swaziland, with scattered records
from KwaZulu-Natal and the Eastern Cape;
widespread in mountains of tropical Africa and
Yemen. Found in undergrowth of forests, usual-
ly near water, recorded once as a garden weed
(Mbabane). Map 13.
Vouchers: Acocks 13949 (PRE); Gerstner 5719
(PRE); Kluge 812 (PRE); Pegler 733 (PRE);
Pole Evans 3942 (PRE).
2007000 8. PARIETARIA
Parietaria L., Species plantarum 2: 1052 (1753); Benth.: 392 (1880); Engl.: 115 (1888); Hutch.:
193 (1967); Friis: 52 (1989a); Friis: 320 (1990); Friis: 627 (1993). Type species: P. officinalis L.
Annual or perennial herbs, polygamous. Leaves alternate, petiolate, entire; cystoliths punctate; stip-
ules absent. Inflorescences of paired sessile or shortly pedunculate clusters in leaf axils; bracts some-
times very prominent; flowers bisexual or unisexual. Female flowers with ± tubular, 3- or 4-lobed peri-
anth; ovary symmetrical; stigma sessile, penicillate, deciduous. Male and hermaphrodite flowers 3- or
4-merous; tepals almost free; rudimentary ovary present. Achenes enclosed in persistent perianth.
Some 10 species widely distributed in the tropics, subtropics and warm temperate regions; one
polymorphic species occurs in mountains of tropical and southern Africa.
Parietaria debilis G.Forst., Florulae insu- Letouzey: 208 (1968); Agnew: 325 (1974);
larum praustralium prodromus: 73 (1786); Friis: 52 (1989a). Type: from islands in South
Rendle: 298 (1917a); Roessler: 5 (1967); Pacific, not yet designated.
22
Urticaceae
Annual herb. Stems prostrate, delicate, some-
times rooting at nodes, glabrous or with short stiff
hairs, sometimes with stalked glands when
young, stinging hairs absent. Leaves broadly
ovate to broader than long, 5-30 x 3-23 mm,
apex obtuse or slightly obtuse-acuminate, base
truncate to slightly cordate, margin entire, gla-
brous or with short stiff hairs, hairs denser along
veins and margin, sometimes with long curved
hairs intermixed; petiole glabrous or with curved
stiff hairs. Inflorescence of 1-few sessile or short-
ly pedicellate flowers in leaf axils, with male,
female and bisexual intermixed. Flowers with
bracts having glandular hairs and long curved
hairs. Female flowers 4-merous, shortly pedicel-
late; perianth ± 0.5 x 1.5 mm long, with minute
hairs on outer surface. Bisexual flowers with peri-
anth with short hairs on outer surface, ± 1 mm
long. Male flowers subsessile, 4-merous, up to 1.5
mm in diameter. Achenes ± 1.5 mm long, smooth,
shiny, dark brown. Figure 5.
Occurs in the Free State, Lesotho, KwaZulu-
Natal and the Eastern Cape, with two records
from Namibia; widespread in warm temperate
regions and mountains of tropical regions.
Found in wet shady areas such as caves and
dense forest, also on edges of vleis. Map 14.
It has been suggested (B.L. Burtt pers.
comm.) that the specimens of Parietaria from
the mountains of tropical and South Africa are
conspecific with those from the Himalayas, and
that all this material should be determined as P.
micrantha Ledeb., as Hara (1975: 23) has done
Names
Parietaria capensis Thunb., Prodromus plantarum capen-
sium: 31 (1794); Thunb.: 155 (1823). Boehmeria capensis
(Thunb.) Spreng.: 844 (1826). Droguetia thunbergii N.E.Br.: 80
(1913); N.E.Br.: 558 (1925), nom. illegit. superfl. Type: Cape
Province, Grootvadersbosch, specimen marked ‘ Urtica cajfra
alpha’, Thunberg s.n. (UPS-THUNB No. 22129!, lecto., vide
Friis 1986: 703). This is Droguetia iners (Forssk.) Schweinf.
Parietaria lanceolata Thunb., Prodromus plantarum
capensium: 31 (1794); Thunb.: 155 (1823). Didymodoxa
debilis var. lanceolata (Thunb.) Wedd.: 235.61 (1869).
Australina lanceolata (Thunb.) N.E.Br.: 555 (1925). Type:
specimen in UPS-THUNB (No. 24019!) marked ‘ Parie-
taria lanceolata ’, and marked ‘Cap. bon. spei’ on reverse.
for the eastern Himalayan specimens. We quite
agree with the view that the South African spec-
imens belong to the same species as the western
Himalayan material, but they are also very simi-
lar to at least part of the material from New
Zealand and Polynesia, the region from which
the type of P. debilis originated. In absence of a
more complete study of the tropical and southern
hemisphere species of the genus, we are very
reluctant to change the name of the African
species, and have therefore retained the epithet
debilis.
Vouchers: Hilliard & Burtt 6730 (PRE);
Hilliard & Burtt 14998 (PRE); Hilliard & Burtt
15072 (PRE); Merxmiiller & Giess 28148
(PRE); Schlechter 4889 (PRE).
This is Pouzolzia zeylanica (L.) Benn., which does not
occur in the FSA region. The species is widespread in trop-
ical and warm temperate Asia, and may have been collect-
ed by Thunberg on his travels in Indonesia and Japan. See
Friis: 703 (1986).
Parietaria pilosa Willd., Species plantarum 4: 954
(1806). Type: ‘Caput b. spei’, stated to have been collected
by Thunberg (B-WILLD No. 18900!, holo.). See Friis: 704
(1986). This is Parietaria judaica L., which at present does
not seem to occur in the FSA region. Another specimen that
appears to be the same species (or P. officinalis L.),
Oldenburg 1405 at BM!, is also stated to have been collect-
ed in South Africa (dated 1772).
Urticaceae
23
2012000
9. FORSSKAOLEA
Forsskaolea L., Opobalsamum declaratum: 17 (1764); Benth.: 393 (1880); Engl.: 117 (1888);
Hutch.: 194 (1967); N.E.Br.: 556 (1925); Roessler: 17 (1967); Merxm. & Roessler: 263 (1980);
Lobin & Roessler: 373 (1985); Friis & Wilmot-Dear: 25 (1988); Friis: 54 (1989a); Friis: 321
(1990); Friis: 628 (1993). Type species: F. tenacissima L.
Annual or perennial herbs or shrublets, monoecious, often covered with hispid hairs. Leaves
alternate, petiolate, variously serrate; cystoliths punctate, prominent; stipules lateral, free.
Inflorescence bisexual, rarely unisexual (female), sessile in leaf axils, enclosed in a campanulate
involucrum of bracts; bracts lanceolate, ovate or obovate, free or fused at base, almost covered by
long dense woolly indumentum, but hairs at base long and stiff. Female flowers few at centre of
inflorescence, without perianth; ovary erect; stigma sessile and filiform. Male flowers pedicellate,
mostly near edge of inflorescence; perianth irregularly 3-lobed; stamen 1, inflexed, later reflexing;
rudimentary ovary absent. Achenes ovate, covered by woolly indumentum.
Three species are found in southern Africa, two of them endemic to the FSA region; there are
five or six species from the drier parts of Africa, the Macaronesian Islands, Spain and extending
to Pakistan and northwestern India. All occur in dry areas and are taxonomically difficult.
Merxmiiller & Roessler (1980) have shown that, in the FSA region, there are frequently inter-
mediate specimens between the three taxa, but that these intermediates are localised to compar-
atively restricted transition zones, a phenomenon which they explain by assuming introgressive
hybridisation. We have consequently chosen to maintain the three taxa at specific rank.
la Involucre of inflorescence usually 8-13 mm long; lobes 3.3-10 times as long as broad;
leaf margin broadly and shallowly dentate with up to 4 pairs of teeth; mostly restrict-
ed to the Namib Desert of southern Namibia and the Northern Cape 1. F. hereroensis
lb Involucre of inflorescence usually 4—9 mm long; lobes 1 .3-3.3 times as long as broad; leaf
margin either broadly and shallowly dentate or crenate, with 3-8 pairs of teeth; from
northern Namibia to the Northern, Western and Eastern Cape:
2a Feaf margins crenate; plant usually herbaceous; Namibia north of Windhoek ... .2 . F. viridis
2b Feaf margins with broad shallow irregular teeth; plant usually woody at base; Namibia
mainly south of Windhoek, the Northern Cape, Western Cape and western part of the
Eastern Cape 3. F. Candida
1. Forsskaolea hereroensis Schinz in
Bulletin de l’Herbier Boissier 4, App. 3: 51
(1896); Rendle: 300 (1917a); N.E.Br.: 557
(1925); Roessler: 4 (1967); Friis & Wilmot-
Dear: 35 (1988). Type: Damaraland, Lindner
s.n. (BR, holo.!; K!).
Woody annual herb up to 1 m high, much
branched. Stems with curved broad-based, stiff
hairs interspersed with shorter puberulous hairs.
Leaves lanceolate to ovate, 1 7 — 40 x 10-26 mm,
apex broadly acute, base decurrent, margin with
teeth obsolete or in up to 4 pairs, with broad-
based curved hairs, upper surface with scattered
slender hooked hairs, lower surface usually
with dense white wool, wool thinner on veins
which often have curved broad-based, stiff
hairs; petiole ± 10-30 mm long; stipules broad-
ly ovate, apex abruptly acuminate, base cordate,
membranous with thicker midrib, fringed with
long stiff hairs. Inflorescence with (6 — )8— 1 2-
lobed involucre, dimensions of lobes as in key,
otherwise as for genus. Flowers as for genus.
Achenes shortly stipitate, ovoid, 2. 5-3.0 x
1.50-2.25 mm, dark brown. Figure 6: 7, 8.
Endemic to Namibia and the Northern Cape.
On rocky hillsides and in riverbeds, usually in
24
Urticaceae
Figure 6. — Forsskaolea Candida: 1, branch apex showing sessile inflorescences, x 1; 2, inflorescence x 4; 3, female
flower, x 15; 4, male flower, x 15 (Oliver, Muller & Schweickerdt 6328). F. viridis: 5, leaf with inflorescence, x 1 ; 6, slen-
der curved hair on leaf margin, x 40 ( Giess 13510). F. hereroensis: 7, inflorescence, x 4; 8, broad-based sharply curved
hair on upper surface of leaf, x 40 (Jensen 341). Artist: Gillian Condy.
Urticaceae
25
shade. Transitional forms between F. hereroen-
sis and F. viridis (no. 2) are known from north-
ern Namibia, and between F. hereroensis and F.
Candida (no. 3) from central and southern
Namibia (see Merxmiiller & Roessler 1980).
Map 15.
Vouchers: Giess & Midler 14440 (PRE);
Hardy 672 (PRE); Jensen 341 (PRE); Strey
2291 (PRE); Visser 00230 (PRE).
2. Forsskaolea viridis Webb in Hook.,
Niger Flora: 179 (1849); Wedd.: 235.56 (1869);
Rendle: 302 (1917a); Roessler: 4 (1967); Friis:
727 (1982); Friis & Wilmot-Dear: 35 (1988).
Type: plant grown at the Paris Botanical Garden
from seed collected at the Red Sea coast of
Ethiopia by Ehrenberg s.n. (FI-WEBB!, lecto.,
vide Friis 1982: 727).
F. eenii Rendle: 203 (1917b); Rendle: 301 (1917a). Type:
Damaraland, Een s.n. (BM!).
Annual herb up to 1 .2 m high, base usually
herbaceous. Stems with sparse broad-based
curved hairs and denser shorter often hooked
stiff hairs. Leaves lanceolate to ovate, 18^10
(-95) x 12— 20(— 55) mm, apex broadly acute,
base decurrent, margin crenate with 4-8 pairs of
teeth, with curved hairs, upper surface with
scattered curved hairs, lower surface usually
with dense white wool, wool thinner on veins
which often have curved hairs; petiole 13-30
(—45) mm long; stipules broadly ovate, margin
ciliate. Inflorescence with (3— )4 — 7(-8)-lobed
involucre, dimensions of lobes as in key, other-
wise as for genus. Flowers as for genus.
Achenes shortly stipitate, ovoid, 2-3 x 1.25-
2.00 mm, dark brown. Figure 6: 5, 6.
Occurs mainly in Namibia in the area north
of Windhoek; also widely distributed in south-
ern Angola, tropical East Africa, Yemen, Saudi
Arabia, southern Egypt and the Cape Verde
Islands. Occurs on rocky slopes and in dry
watercourses in shade. Transitional forms to F.
hereroensis (no. 1) exist (see under that spe-
cies). Map 16.
The involucral bracts of the type of F. eenii
approach those of F. hereroensis, as the
length/width ratio is ± 3.5 and the length of the
involucrum 8-9 mm, but the general morpholo-
gy is most similar to that of F. viridis. For this
reason we have placed it under F. viridis, while
Roessler (1967: 3) has placed it under F. Candi-
da (no. 3). As the type belongs to the group of
intermediate specimens, the choice of position
will remain somewhat arbitrary.
Vouchers: De Winter & Leistner 5222 (PRE);
Dinter 492 (PRE); Giess 13308 (PRE); Goyns
64 (PRE); Schoenfelder S623 (PRE); Volk 2331
(PRE).
26
Urticaceae
3. Forsskaolea Candida L.f., Supple-
mentum plantarum: 245 (1782); Rendle: 301
(1917a); N.E.Br.: 556 (1925); Roessler: 3
(1967); Friis & Wilmot-Dear: 35 (1988). Type:
Cap. bon. spei, Thunberg s.n. A specimen in
LINN (No. 605.3) might be a suitable lectotype;
there is also a specimen, perhaps a duplicate, at
S-LINN (No. 195.13).
F. Candida var. virescens Wedd.: 235.56 (1869); N.E.Br.:
557 (1925). Type: South Africa, Drege s.n., marked
' Forsskaolea viridis Ehrenb. ?’ (P, ex Herb. Lenormand!).
F. scabra Retz.: 31 (1783). Type: Cape, Thunberg s.n. (A
specimen at LD, in the Herb. Retzius, would be a suitable
lectotype.)
Description as for F. viridis except the fol-
lowing: Base of plant often woody. Leaves
10-75 x 6^40 mm, margin with 3-7 pairs of
broad irregular teeth; petiole 6-30 mm long.
Inflorescence with (4 — )5— 9(— 1 1 )-lobed involu-
cre, dimensions of lobes as in key, otherwise as
for genus. Figure 6: 1-4.
Confined mainly to south-central Namibia
and the dry areas of the Northern and Western
Cape, with a few records from northern Namibia
Map 17. — Forsskaolea Candida
and the Eastern Cape. On rocky hillsides and in
riverbeds, usually in shade. Transitional speci-
mens between this species and F. hereroensis
exist (see under that species). Map 17.
Vouchers: Giess 14591 (PRE); Giess & Muller
12297 (PRE); Oliver, Muller & Steenkamp 6328
(PRE); Oliver & Steenkamp 6237 (PRE); Van der
Westhuizen 226 (PRE).
2013000 10. DROGUETIA
Droguetia Gaudich. in Freycinet, Voyage autor du monde ... execute sur les corvettes de S.M.
PUranie et la Physicienne Pillet-aine - Botanique: 505 (1830); Benth.: 394 (1880); Engl.: 117
(1888); Hutch.: 195 (1967); Friis, Immelman & Wilmot-Dear: 125 (1987); Friis & Wilmot-Dear:
36 (1988); Friis: 56 (1989a); Friis: 322 (1990); Friis: 112 (1991); Friis: 628 (1993). Type species:
D. leptostachys (Pers.) Wedd. (lecto., vide Friis & Wilmot-Dear 1988: 36).
Slender annual or perennial herbs, monoecious, or apparently dioecious by abortion. Leaves
alternate or opposite, petiolate, serrate; cystoliths dot-like; stipules lateral, free. Inflorescences ses-
sile axillary clusters, either bisexual and surrounded by a common campanulate involucre of fused
bracts, or a cluster of very small involucres each with 1(2) female flowers (involucres then appear-
ing like perianths); bisexual involucres usually with many male and 1-few female flowers (rarely
all male); female involucres much smaller than bisexual ones; bracts and flowers with long wool-
ly hairs. Female flowers pedicellate, naked; ovary covered with a woolly tomentum; stigma sessile,
filiform; staminodes absent. Male flowers pedicellate, clavate; perianth 3-lobed, 1 lobe apiculate
and with prominent setae; stamen single, inflexed in bud. Achenes enclosed in involucrum, brown,
shiny, often crowned by persistent stigma.
Seven species in tropical and southern Africa, Yemen, South India and on Java; in southern Africa
the two species and two subspecies are found in KwaZulu-Natal and the Eastern and Western Cape.
Urticaceae
27
Leaves always opposite 1 . D. iners
At least uppermost leaves on stem or some of the leaves on side branches alternate ... 2. D. ambiguci
1. Droguetia iners (Forssk.) Schweinf. in
Bulletin de l’Herbier Boissier 4, App. 2: 146
(1896); Rendle: 303 (1917a); Letouzey: 213
( 1968); Agnew: 325 (1974); Friis, Immelman &
Wilmot-Dear: 126 (1987); Friis & Wilmot-
Dear: 38 (1988); Friis: 57 (1989a); Friis: 112
(1991). Type: Yemen, Forsskal s.n. (C, holo.!;
microfiche in PRE, No. 774!).
Perennial herb or subshrub. Stems prostrate
and erect, up to ± 1 m high, indumentum very
varied (see key to subspecies). Leaves ovate, up
to 45(-80) x 25(-45) mm, apex acute or acumi-
nate, base cuneate, margin serrate with 3-12
pairs of teeth, upper surface with scattered stiff
hairs above, lower surface with stiff hairs on
veins or sometimes also scattered; petiole up to
20(^15) mm long, with indumentum as in
branches; stipules lanceolate, apex acuminate,
up to 3 mm long, midrib brown and prominent,
membranous. Inflorescences in upper axils con-
sisting of clusters of bisexual, female or mixed
inflorescences, in lower axils often only female
inflorescences, bisexual inflorescences up to 4
mm in diameter. Flowers as for genus. Achenes
enclosed in persistent, glabrous or lanate in-
volucre.
Found in KwaZulu-Natal and the Eastern
and Western Cape; also through East tropical
Africa to the Sudan, northern Ethiopia, Macias
Nguema (Fernando Po), Cameroon, and in
Angola; also in the Yemen and in the mountains
of southern India and Indonesia. Occurs in
montane forests and evergreen scrub.
There are two subspecies in the FSA area:
Young stems and petioles subglabrous or
with an indumentum of stiff sparse
hairs or stiff appressed hairs; termi-
nal tooth usually more than twice as
long as broad; stipules lanceolate,
not prominent la. subsp. iners
Young stems and petioles with a dense
indumentum of erect or slightly curly
fine hairs; terminal tooth rarely
more than 1.5 times as long as
broad; stipules prominent and broad
lb. subsp. burchellii
la. subsp. iners.
Urtica iners Forssk.: 160 (1775). U. urens var. iners
(Forssk.) Wedd.: 40 (1869).
D. thunbergii N.E.Br.: 80 (1913); N.E.Br.: 558 (1925).
Type: Cape Province, probably Grootvadersbosch,
Thunberg specimen marked 'Urtica capensis alpha’ ( UPS-
THUNB No. 22129!, lecto., vide Friis 1986: 703).
D. woodii N.E.Br.: 561 (1925). Type: KwaZulu-Natal,
Inanda, Wood 1243 (K!).
Slender or robust herb or subshrub 0.1-1 m
high. Stems with hairs as in key. Leaves
18-45(-80) x 10-25(-45) mm, margin with
4-8(-12) pairs of teeth, upper surface glabrous
to sparsely hairy, lower with a few stiff hairs on
veins, sometimes also with scattered stiff hairs;
petiole 5-20(-40) mm long; stipules lanceolate,
1-2 x up to 2 mm. Inflorescence with involu-
crum always lanate on the inside, otherwise as
for species. Flowers and fruit as for species.
28
Urticaceae
Distribution and ecology as for the species,
with exception of the Western Cape and the
Indian and Indonesian part of the range, where
it is replaced by subsp. burchellii and subsp.
urticoides (Wight) Friis & Wilmot-Dear respec-
tively. Map 18.
Vouchers: Duthie s.n. ( 13059 in STE, PRE);
Rattray 334 (PRE).
lb. subsp. burchellii (N.E.Br.) Friis &
Wilmot-Dear in Friis, Immelman & Wilmot-
Dear in Nordic Journal of Botany 7,2: 126
(1987); Friis & Wilmot-Dear: 41 (1988). Type:
Eastern Cape, Bathurst Division, Barville Park,
Burchell 4084 (K, holo.!).
D. burchellii N.E.Br.: 561 (1925).
Differs from subsp. iners as follows: Plants
up to 0.5 m high; pubescence differing as given
in key. Leaves 15-60 x 6-25 mm, margin with
3-10 pairs of teeth, apex and pubescence differ-
ing as given in key; petiole 5-25 mm long; stip-
ules broadly lanceolate to ovate, especially in
upper part of stems, sometimes broader than
long, up to 3 x 4 mm.
Endemic to South Africa, from southern
KwaZulu-Natal and the Eastern and Western
Cape. Found in coastal forest and scrub, some-
times in dune forest or evergreen scrub. Map 19.
Vouchers: Flanagan 1224 (PRE); Rousseau
67 (PRE); Taylor 10401 (PRE); Tyson PRE 12643
(PRE); Wager s.n. (PRE, 24109 in Tvl Mus.).
Note on variation in D. iners : There is a
degree of overlap in all characters used to dis-
tinguish taxa in this widespread species. Friis &
Wilmot-Dear have therefore recognised only
three taxa (subspecies iners, burchellii and
urticoides), and only at the subspecific level.
Subsp. burchellii was recognised on its different
indumentum and larger stipules; it also occupies
a limited distribution along the southern coast
of South Africa. The type of D. thunbergii is
interpreted as a small-leaved form of D. iners.
The placement of the type of D. woodii has
proved somewhat difficult, as it has exception-
ally large leaves and does not fit well into any
of the subspecies, but it was decided that it was
closest to subsp iners.
2. Droguetia ambigua Wedd. in Annales
des Sciences Naturelles, Paris. Botanique, Ser.
4,1: 211 (1854); N.E.Br.: 560 (1925) excl. spec-
im. cit. ; Friis & Wilmot-Dear: 41 (1988). Type:
South Africa, without further locality, Drege
s.n., specimen marked ‘ Parietaria urticaefolia
L. a’ (P!, lecto., vide Friis & Wilmot Dear 1988:
42; BM!, E!, G!, K!).
D. urticaefolia Wedd.: 235.58 (1869); N.E.Br.: 559
(1925), nom. illegit. superfl. based on D. ambigua.
Herb, apparently annual but possibly some-
times perennial. Stems prostrate and ascend-
ing, up to 0.4 m high; young branches with
sparse to dense erect stiff or sometimes curly
hairs. Leaves broadly ovate to rhomboid, up to
40 x 22 mm but usually much smaller, almost
glabrous above, with a few stiff hairs along
veins below, apex bluntly acute, terminal
tooth ± as long as broad, base broadly cuneate,
margin crenate to bluntly serrate with 2-7
pairs of teeth; petiole up to 14 mm long but
usually shorter; stipules broadly lanceolate,
up to 2 x 1.5 mm, green, ciliate. Inflores-
cences bisexual and almost globular in upper
Urticaceae
29
outside, densely lanate on inside, up to 2.5
mm in diameter. Achenes enclosed in marked-
ly lanate involucre.
Endemic to South Africa (KwaZulu-Natal
and Eastern Cape). Found in coastal bush, often
on dunes, or in forest. Map 20.
Often confused in the literature with D. iners
(no. 1), but can easily be distinguished by the
deep green leaves, the characteristic leaf shape,
the crenate leaf margin and the usually almost
globular bisexual involucres, which have a
characteristic pale pubescent or lanate indu-
mentum on the outer surface.
Vouchers: Ross & Moll 2197 (PRE); Ross &
Moll 2315 (PRE); Ward 4901 (PRE); Watmough
414 (PRE).
2014010 11. DIDYMODOXA
Didymodoxa Wedd. in Archives du Museum National d’Histoire Naturelle, Paris 9: 547 (1857);
Wedd.: 235.61.8 (1869); Friis: 209 (1985b); Friis, Immelman & Wilmot-Dear: 126 (1987); Friis &
Wilmot-Dear: 45 (1988); Friis: 60 (1989a); Friis: 114 (1991); Friis: 628 (1993). Type species: D.
integrifolia Wedd. (lecto., vide Friis & Wilmot-Dear 1988: 45).
Australina auct., non Gaudich.; Benth.: 394 (1880), p.p.; Engl.: 117 (1888), p.p.; Rendle: 305 (1917a) quoad A. acumi-
nata; N.E. Br.: 552 (1925); Hutch.: 194 (1967), p.p. and other authors on the flora of southern and East Africa.
The name Didymotoca E.Mey. is recorded in the Index kewensis and elsewhere, but has not been validly published.
Annual herbs, monoecious, or dioecious by abortion. Leaves alternate, petiolate, serrate, crenate
or entire; cystoliths punctiform; stipules free, lateral. Inflorescence usually bisexual, axillary, ses-
sile, bracteate; bracts free, not prominent, usually shorter than flowers. Female flowers without
perianth; ovary erect, with a subcapitate or shortly linear style. Male flowers with a boat-shaped,
sometimes almost bract-like perianth with erect tip and slightly fused cylindrical base; stamen 1,
indexed (later reflexed); rudimentary ovary absent. Achenes slightly winged along one side, other
side rounded, 2 ovaries or achenes frequently joining along rounded side thus forming a double
fruit much larger than 2 single fruits, both seeds developing in this case.
Two species in southern Africa and East tropical Africa, as far north as northern Ethiopia.
Widespread in southern Africa but not often collected, from northern and southern Namibia, the
Northern Province, Gauteng, Mpumalanga, the Free State, KwaZulu-Natal, Fesotho and the
Eastern, Western and Northern Cape.
What was formerly known as the genus Australina, distributed in Australia and Africa, has been
divided into two genera by Friis & Wilmot-Dear (1988: 30), who resuscitated Weddell's genus
Didymodoxa , applying the name to two of the African species, while restricting Australina sensu
axils, unisexual (female) only in lower axils;
bisexual inflorescences pubescent or lanate on
30
Urticaceae
stricto to two species, one from Australia and New Zealand and the other from Ethiopia and north-
western Kenya.
Stipules broadly lanceolate to ovate, ( 1 . 5— )2 — 4(— 5 ) mm wide and up to 5.5 mm long; leaves
ovate with entire to crenate margin or with up to 5(— 7) blunt to pointed teeth per side;
apical tooth often indiscernible, rarely longer than broad; inflorescence often with
11-20 flowers; Western Cape and southern Namibia (winter rainfall) 1. D. capensis
Stipules relatively narrow, lanceolate, up to 0.7(-l) mm wide and up to 2.5 mm long; leaves
lanceolate to ovate, always with 6-12 pointed teeth per side; apical tooth clearly dis-
cernible, usually 2-3 times as long as broad; inflorescences with few-10 flowers;
northern Namibia, Northern Province, Gauteng, Mpumalanga, Free State, KwaZulu-
Natal, Lesotho and Eastern Cape (summer rainfall) 2. D. cajfra
1. Didymodoxa capensis (L.f.) Friis &
Wilmot-Dear in Friis, Immelman & Wilmot-
Dear in Nordic Journal of Botany 7,2: 126
(1987); Friis & Wilmot-Dear: 46 (1988). Type:
South Africa, without precise locality, Thunberg
s.n., specimen marked ‘ Urtica capensis beta’
(UPS-THUNB No. 22,131 !, lecto., vide Friis &
Wilmot-Dear 1987).
Urtica capensis L.f.: 417 (1782). Mercurialis capensis
(L.f.) Sond.: 112 (1850), excl. descr. et syn. heterotyp.
Fleurya capensis (L.f.) Wedd.: 183 (1854). Leidesia capen-
sis (L.f.) Miill.Arg.: 793 (1866), excl. descr., specim. cit. et
syn. heterotyp. Acalypha capensis (L.f.) Prain in Prain &
Hutch.: 15(1913) excl. descr., specim. cit. et syn. heterotyp.
Australina thunbergii N.E.Br.: 554 (1925), nom. illegit.
superfl. Type: South Africa, without further locality,
Thunberg s.n., specimen marked ‘ Urtica capensis beta’
(UPS-THUNB No. 22 131, lecto.).
Note: the lectotypification of Urtica capensis L.f. made
by Prain (1913: 386, 387) and accepted by Prain &
Hutchinson (1913: 15, 16) with a specimen of Acalypha
decumbens Thunb. in LINN (No. 1111.26) is in conflict
with the protologue and has therefore been superseded, see
Friis & Wilmot-Dear: 47 (1988).
Prostrate to erect annual herb up to 0.35 m
high. Stems glabrous when young, or with
sparse to dense stiff hairs. Leaves ovate to ellip-
tic, 8M-5(-55) x 6-38 mm, apex bluntly acute,
acute or sometimes acuminate, base broadly
cuneate, margin entire or with 5-15 pairs of cre-
nations or teeth, upper surface with sparse stiff
hairs, lower with long stiff hairs on veins and
short stiff scattered hairs; petiole 3-38 mm
long; stipules triangular, ovate or broadly lan-
ceolate, up to 5.5 mm long, membranous with
green midrib, reticulation or margin, ciliate.
Inflorescence usually bisexual, occasionally of
a single female flower only, up to 4 mm in
diameter; bracts with cilia up to 2 mm long,
lanceolate, often translucent. Female flowers
with stiff hairs, up to 2 mm long, may fuse in
pairs. Male flowers acute to shortly acuminate,
ciliate, up to 2 x 1 mm long; perianth wider than
in D. caffra. Achenes up to 2.5 mm long, double
fruits up to 3.5 mm long.
Endemic to the winter-rainfall area of the
Western and Northern Cape, from the Cape
Peninsula to Namaqualand. Growing among
rocks and on sandy flats.
Two varieties are recognised:
Leaves with well-defined teeth
la. var. capensis
Leaves entire or very shallowly toothed . .
lb. var. integrifolia
la. var. capensis.
Australina procumbens N.E.Br.: 553 (1925). Type:
Northern Cape, Het Kruis, Stephens & Glover 8776 ( K!,
BOL).
Erect or procumbent herb, often fairly robust.
Leaves broadly lanceolate to ovate, up to 45 x
37 mm, margin serrate or crenate; stipules up to
2.25 mm wide, at least on upper nodes. Inflores-
cence many-flowered, not on morphologically
distinguished part of plant.
Urticaceae
31
Map 21. — Didymodoxa capensis var. capensis
Both this and the following variety are dis-
tributed in the winter-rainfall area of the Cape,
from the Cape Peninsula in the Western Cape to
Namaqualand in the Northern Cape; also found
in southern Namibia. Growing among rocks and
on sandy flats. Map 21.
Vouchers: Acocks 14922 (PRE); Acocks
23318 (PRE); Bolus 9455 (PRE); Bolus 12827
(PRE); Mittendorf 104 (PRE).
1 b. var. integrifolia Friis & Wilmot-Dear in
Friis, Immelman & Wilmot-Dear in Nordic
Journal of Botany 7,2; 126 (1987); Friis &
Wilmot-Dear: 49 (1988). Type: South Africa,
without further indication; Cape Province, Drege
s.n ., marked ‘ Parietaria lanceolata Thunb. ?c’ (P!,
lecto., vide Friis & Wilmot-Dear 1987; P!, K!).
Australina integrifolia Wedd.: 212 (1854). D. integrifolia
(Wedd.) Wedd.: 549(1857).
A. capensis Wedd.: 212 (1854); N.E.Br.: 554 (1925). D.
debilis Wedd.: 548, t. 20B (1857); Wedd.: 235 (1869), nom.
illegit. superfl. Type: Cape Province, Drege s.n., specimen
marked 'Didymodoxa debilis E.M. a' (BM!, E!, G!, K!,
MEL!, P!, S!).
A. paarlensis N.E.Br.: 553 (1925). Type: Western Cape,
Paarl Div., Paarl Mountain, Drege s.n., marked 'Didy-
modoxa debilis E.M.c.’ (BM!, E!, G!, K!, SAM!).
Erect or procumbent herb, often delicate.
Leaves ovate to broadly ovate, usually less than 20
x 15 mm, margin entire to slightly crenate, rather
membranous; stipules lanceolate, up to 1.25 mm
wide. Inflorescences comparatively few-flowered,
often situated on part of plant with longer inter-
nodes and shorter leaves than the infertile part.
Distribution (except for southern Namibia)
and habitat as for typical variety. Map 22.
Vouchers: Galpin 11506 (PRE); Hafstrom &
Acocks 379 (PRE); Marloth 146 (PRE); Oliver,
Tolken & Venter 512 (PRE); Taylor 7126 (PRE).
2. Didymodoxa eaffra (Thunb.) Friis &
Wilmot-Dear in Friis in Boletim Sociedade
Broteriana 58: 210 (1985b); Friis & Wilmot-Dear:
49 (1988); Friis: 60 (1989a); Friis: 114 (1991).
Type: South Africa, without further locality,
Thunberg s.n., specimen marked ‘ Urtica eaffra beta'
(UPS-THUNB!, lecto., vide N.E.Br. 1913: 80).
Urtica eaffra Thunb.: 31 (1794); Thunb.: 155 (1823).
Australina eaffra (Thunb.) Prain: 388 (1913); Fourc.: 80
(1941).
A. acuminata Wedd.: 212 (1854); Rendle: 306 (1917a);
N.E.Br.: 555 (1925); Roessler: 2 (1967); Agnew: 325 (1974).
D. acuminata (Wedd.) Wedd.: 549 (1857). D. cuneata Wedd.:
235.62.8 (1869), nom. illegit. superfl.. based on A. acumina-
ta. Type: KwaZulu-Natal. Yellowwood River, Drege s.n.,
specimen marked 'Parietaria cuneata E.M. a' (G ! , K!, P!).
Slender annual herb. Stems up to 0.5 m high,
much branched, sometimes rooting at nodes,
32
Urticaceae
moderately to sparsely pubescent with patent
stiff or slender curly hairs, sometimes also with a
few hooked hairs. Leaves lanceolate, 23-75 x
14-40 mm, base cuneate, margin clearly crenate
with 5-11 pairs of teeth, upper surface with
sparse appressed stiff hairs, lower with numerous
hairs on veins, few scattered hairs between veins;
petiole 25^17 mm long; stipules lanceolate, 2-5
x 0.3-0.6 mm, attenuate, glabrous or ciliate.
Inflorescence up to 5 mm in diameter; bracts lin-
ear to narrowly lanceolate, ciliate. Achenes cov-
ered with broad hooked hairs and glands, ± 1.75
x 1 mm, double fruits up to ± 3 x 2.5 mm present
in Namibian material.
In northern Namibia, the Northern Province,
Gauteng, Mpumalanga, the Free State, Kwa-
Zulu-Natal, Lesotho and the Eastern Cape; trop-
ical East Africa as far north as Ethiopia. Among
rocks or in forest in moist places. Map 23.
12 14 16 18 20 22 24 26 28 30 32 34
Map 23. — Didymodoxa caffra
Vouchers: Dinter 5638 (PRE); Giffen 868
(PRE); Kluge 1747 (PRE); Repton 252 (PRE);
Repton 703 (PRE).
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Urticaceae
35
INDEX*
Acalypha capensis (L.f.) Prain, 30
decumbens Thunb., 30
Australina auct., 29
Australina acuminata Wedd., 31
caffra (Thunb.) Prain, 31
capensis Wedd., 31
integrifolia Wedd., 31
lanceolata (Thunb.) N.E.Br., 22
paarlensis N.E.Br., 31
procumbens N.E.Br., 30
thunbergii N.E.Br., 30
Boehmeria capensis (Thunb.) Spreng., 22
nivea (L.) Gaudich., 1
procridioides (Wedd.) Blume, 21
Didymodoxa Wedd., 29
acuminata (Wedd.) Wedd., 31
caffra (Thunb.) Friis & Wilmot-Dear, 31
capensis (L.f. ) Friis & Wilmot-Dear , 30
var. capensis, 30
var. integrifolia Friis & Wilmot-Dear, 3 1
cuneata Wedd., 3 1
debilis Wedd., 31
var. lanceolata (Thunb.) Wedd., 22
integrifolia Wedd., 29
integrifolia (Wedd.) Wedd., 31
Didymotoca E.Mey., 29
Droguetia Gaudich., 26
ambigua Wedd., 28
burchellii N.E.Br., 28
iners (Forssk.) Schweinf, 22, 27
subsp. burchellii (N.E.Br.) Friis & Wilmot-Dear, 28
subsp. iners, 27
leptostachys (Pers.) Wedd., 26
thunbergii N.E.Br., 22, 27
urticaefolia Wedd., 28
woodii N.E.Br., 27
Elatostemma trinerve Hochst., 7
Fleurya Gaudich., 10
aiatipes (Hook.f.) N.E.Br., 11
capensis auct., 12
capensis (L.f.) Wedd., 30
var. mitis Wedd., 12
grossa Wedd., 13
mitis Wedd., 12
mitis (Wedd.) N.E.Br., 12
peduncularis Wedd., 12
var. mitis (Wedd.) Wedd., 12
spicata Gaudich., 10
Forsskaolea L., 23
Candida L.f, 26
var. virescens Wedd., 26
eenii Rendle, 25
hereroensis Schinz, 23
scabra Retz., 26
* Synonyms are in italics; naturalised taxa are marked with
an asterisk.
tenacissima L., 23
viridis Webb, 25
Girardinia Gaudich., 14
condensata (Steud.) Wedd., 14
diversifolia (Link) Friis, 14
heterophylla Decne., 14
leschenaultiana Decne., 14
palmata Blume, 14
vahlii Blume, 14
Laportea Gaudich., 10
aiatipes Hook.f, 1 1
caffra Chew, 13
canadensis (L.) Wedd., 10
carruthersiana (Hiem) K.Schum., 9
grossa (Wedd.) Chew, 13
interrupta (L.) Chew, 10, 13
mooreana (Hiem) Chew, 14
peduncularis (Wedd.) Chew, 12
subsp. latidens Friis, 10. 13
subsp. peduncularis, 12
Leidesia capensis (L.f.) Miill.Arg., 30
Margarocarpus procridioides Wedd., 21
Mercurialis capensis (L.f.) Sond., 30
Obetia Gaudich., 8
australis Engl., 9
carruthersiana (Hiem) Rendle, 9
ficifolia (Savigny) Gaudich., 8
tenax (N.E.Br.) Friis, 9
Parietaria L., 21
capensis Thunb., 22
debilis G.Forst., 21
judaica L., 22
laevigata Poir., 17
lanceolata Thunb., 22
micrantha Ledeb. 22
officinalis L., 21, 22
pilosa Willd., 22
Pile a Lindl., 15
ceratomera Wedd., 17
microphylla (L.) Liebm., 15
muscosa Lindl. 15
rivularis Wedd., 17
worsdellii N.E.Br., 17
Pouzolzia Gaudich., 17
hypoleuca Wedd., 19
laevigata (Poir.) Decne., 17
mixta Solms, 1, 19
parasitica (Forssk.) Schweinf. , 19
procridioides (Wedd.) Wedd., 21
zeylanica (L.) Benn., 22
Urera Gaudich., 7
acuminata var. cameroonensis (Wedd.) Leandri. 7
baccifera (L.) Gaudich., 7
cameroonensis Wedd., 7
engleriana Dinter, 9
tenax N.E.Br., 9
trinervis (Hochst.) Friis & lmmelman, 7
woodii N.E.Br., 7
36
URTICACEAE
Urtica L„ 2
burchellii N.E.Br., 5
cajfra Thunb., 31
capensis L.f., 30
condensata Steud., 14
dioica L .*, 2, 3
var. capensis Wedd., 3
var. eckloniana (Blume) Wedd., 3
diversifolia Link, 14
eckloniana Blume, 3
var. flavovirens Blume, 3
heterophylla D.Don, 14
heterophylla Vahl, 14
iners Forssk., 27
interrupta L., 13
lobata Blume, 5
lobulata Blume , 5
meyeri Wedd., 5
mitis Hochst., 12
palmata Forssk. 14
parasitica Forssk., 19
urens L.*, 5
var. iners (Forssk.) Wedd., 27
URTICACEAE, 1
Urticastrum alatipes (Hook.f.) Kuntze, 1 1
carruthersianum Hiem, 9
A-l
APPENDIX
PLAN OF FLORA OF SOUTHERN AFRICA
Cryptogam volumes will in future not be numbered, but will be known by the name of the group they cover. The number
assigned to the volume on Charophyta therefore becomes redundant. Occasional contributions to the Flora are published in
Bothalia under the title FSA contributions.
Exotic families are marked with an asterisk.
Published volumes and parts are shown in bold.
INTRODUCTORY VOLUMES
The genera of southern African flowering plants, Vols 1 (1975) and 2 (1976). Replaced by Seed plants of southern
Africa: families and genera, published as Strelitzia 10 (2000).
Botanical exploration of southern Africa (1981)
CRYPTOGAM VOLUMES
Charophyta (as Vol. 9 in 1978)
Bryophyta: Part 1: Musci: Fascicle 1: Sphagnaceae, Andreaeaceae, Fissidentaceae, Nanobryaceae, Archidiaceae,
Ditrichaceae, Seligeriaceae, Dicranaceae, Calymperaceae, Encalyptaceae,
Pottiaceae, Bryobartramiaceae, Grimmiaceae (1981)
Fascicle 2: Gigaspermaceae, Ephemeraceae, Funariaceae, Splachnaceae, Bryaceae, Mnia-
ceae, Eustichiaceae, Rhizogoniaceae, Aulacomniaceae, Bartramiaceae (1987)
Fascicle 3: Erpodiaceae, Rhachitheciaceae, Ptychomitriaceae, Orthotrichaeeae, Rhabdowei-
siaceae, Racopilaceae, Fontinalaceae, Wardiaceae, Hedwigiaceae, Cryphaeaceae,
Leucodontaceae, Prionodontaceae, Trachypodaceae, Pterobryaceae, Meteoria-
ceae, Leptodontaceae, Neckeraceae, Thamnobryaceae, Hookeriaceae (1998)
Fascicle 4: Fabroniaceae, Leskeaceae, Thuidiaceae, Rigodiaceae, Amblystegiaceae, Brachy-
theciaceae, Entodontaceae, Plagiotheciaceae, Catagoniaceae, Sematophyllaceae,
Hypnaceae, Hylocomiaceae, Polytrichaceae
Hepatophyta: Part 1: Marchantiopsida: Fascicle 1: Targioniaceae, Lunulariaceae, Aytoniaceae, Cleveaceae, Exormo-
thecaceae, Marchantiaceae, Oxymitraceae, Ricciaceae (1999)
Anthocerotophyta
Pteridophyta (1986)
FLOWERING PLANTS VOLUMES
Vol. 1: Stangeriaceae, Zamiaceae, Podocarpaceae, Pinaceae*, Cupressaceae, Welwitschiaceae, Typhaceae, Zoster-
aceae, Potamogetonaceae, Ruppiaceae, Zannichelliaceae, Najadaceae, Aponogetonaceae, Juncaginaceae,
Alismataceae, Hydrocharitaceae (1966)
Vol. 2: Poaceae
Vol. 3: Cyperaceae, Arecaceae, Araceae, Lemnaceae, Flagellariaceae
Vol. 4: Part 1: Restionaceae
Part 2: Xyridaceae, Eriocaulaceae, Commelinaceae, Pontederiaceae, Juncaceae (1985)
Vol. 5: Part 1: Fascicle 1: Aloaceae (First part): Aloe (2000)
Colchicaceae, Eriospermaceae, Asphodelaceae (Chortolirion, 1995 in Bothalia 25: 31-33; Poellnitzia, 1995
in Bothalia 25: 35, 36)
Part 2: Alliaceae, Liliaceae*, Hyacinthaceae, Agavaceae (1996 in Bothalia 26: 31-35)
Part 3: Dracaenaceae, Asparagaceae, Luzuriagaceae, Smilacaceae (1992)
A-2
Vol. 6:
Vol. 7:
Vol. 8:
Vol. 9:
Vol. 10:
Vol. 11:
Vol. 12:
Vol. 13:
Vol. 14:
Vol. 15:
Vol. 16:
Vol. 17:
Vol. 18:
Vol. 19:
Vol. 20:
Vol. 21:
Vol. 22:
Vol. 23:
Vol. 24:
Haemodoraceae, Amaryllidaceae, Hypoxidaceae, Tecophilaeaceae, Velloziaceae, Dioscoreaceae
Iridaceae: Part 1: Nivenioideae, Iridoideae
Part 2: Ixioideae: Fascicle 1: Ixieae (First part): Ixiinae, Tritoniinae (1999)
Fascicle 2: Syringodea, Romulea (1983)
Musaceae, Strelitziaceae, Zingiberaceae (1998 in Bothalia 28: 35-39), Cannaceae*, Burmanniaceae, Orchidaceae
(Holothrix, 1996 in Bothalia 26: 125-140)
Part: Urticaceae (2001)
Casuarinaceae* (2000 in Bothalia 30: 143-146), Piperaceae (2000 in Bothalia 30: 25-30), Salicaceae, Myrica-
ceae, Fagaceae*, Ulmaceae (1999 in Bothalia 29: 239-247), Moraceae, Cannabaceae* (1999 in Bothalia 29:
249-252), Proteaceae
Part 1 : Loranthaceae, Viscaceae ( 1979),
Santalaceae, Grubbiaceae, Opiliaceae, Olacaceae, Balanophoraceae, Aristolochiaceae, Rafflesiaceae, Hydno-
raceae, Polygonaceae, Chenopodiaceae, Amaranthaceae, Nyctaginaceae
Phytolaccaceae, Aizoaceae, Mesembryanthemaceae
Portulacaceae, Basellaceae, Caryophyllaceae, Illecebraceae, Cabombaceae, Nymphaeaceae, Ceratophyllaceae
(1997 in Bothalia 27: 125-128), Ranunculaceae, Menispermaceae, Annonaceae, Trimeniaceae, Lauraceae,
Hemandiaceae, Papaveraceae, Fumariaceae
Brassicaceae, Capparaceae, Resedaceae, Moringaceae, Droseraceae, Roridulaceae, Podostemaceae, Hydro-
stachyaceae (1970)
Crassulaceae (1985)
Vahliaceae, Montiniaceae, Escalloniaceae, Pittosporaceae, Cunoniaceae, Myrothamnaceae, Bruniaceae, Hama-
melidaceae, Rosaceae, Connaraceae
Fabaceae: Part 1: Mimosoideae (1975)
Part 2: Caesalpinioideae ( 1977)
Part 3: Papilionoideae: Fascicle 1:
Fascicle 2:
Fascicle 3:
Fascicle 4:
Fascicle 5:
Fascicle 6:
Fascicle 7:
Fascicle 8:
Fascicle 9:
Swartzieae-Robinieae
Indigofereae
Desmodieae, Phaseoleae
Psoraleeae-Galegeae
Loteae-Liparieae
Crotalarieae ( Aspalathus ) (1988)
Crotalarieae (Bolusia-Lebeckia)
Crotalarieae (Lotononis-Wiborgia)
Crotalarieae ( Pearsonia-Argyrolobium ), Genisteae ( Cytisus-Ulex )
Geraniaceae, Oxalidaceae
Part 1 : Linaceae, Erythroxylaceae, Zygophyllaceae, Balanitaceae
Part 2: Rutaceae
Part 3: Simaroubaceae, Burseraceae, Ptaeroxylaceae, Meliaceae (Aitoniaceae), Malpighiaceae (1986)
Part 1 : Polygalaceae, Dichapetalaceae
Part 2: Euphorbiaceae, Callitrichaceae, Buxaceae (1996 in Bothalia 26: 37-40)
Part 3: Anacardiaceae: Fascicle 1: Rhus (1993)
Fascicle 2: remaining genera
Aquifoliaceae (1994 in Bothalia 24: 163-166)
Celastraceae, Icacinaceae, Sapindaceae, Melianthaceae, Greyiaceae, Balsaminaceae, Rhamnaceae, Vitaceae
Part 1: Tiliaceae (1984)
Malvaceae, Bombacaceae, Sterculiaceae
Ochnaceae, Clusiaceae, Elatinaceae, Frankeniaceae, Tamaricaceae, Canellaceae, Violaceae, Flacourtiaceae,
Turneraceae, Passifloraceae, Achariaceae, Loasaceae, Begoniaceae, Cactaceae (1976)
Geissolomataceae, Penaeaceae, Oliniaceae, Thymelaeaceae, Lythraceae, Lecythidaceae
Rhizophoraceae, Combretaceae, Myrtaceae, Melastomataceae, Onagraceae (1997 in Bothalia 27: 149-165),
Trapaceae (1998 in Bothalia 28: 11-14), Haloragaceae, Gunneraceae, Araliaceae, Apiaceae, Comaceae
A-3
Vol. 25: Ericaceae
Vol. 26: Myrsinaceae, Primulaceae, Plumbaginaceae, Sapotaceae, Ebenaceae, Oleaceae, Salvadoraceae, Loganiaceae,
Gentianaceae, Apocynaceae (1963)
Vol. 27: Part 1: Periplocaceae, Asclepiadaceae (Microloma-Xysmalobium)
Part 2: Asclepiadaceae (Schizoglossum-Woodia)
Part 3: Asclepiadaceae ( Asclepias-Anisotoma )
Part 4: Asclepiadaceae ( Brachystelma , Ceropegia, Riocreuxia) (1980)
Asclepiadaceae (remaining genera)
Vol. 28: Parti: Con vol vulaceae (2000)
Part 2: Hydrophyllaceae, Boraginaceae
Part 3: Stilbaceae, Verbenaceae ( Vitex , 1996 in Bothalia 26: 141-151)
Part 4: Lamiaceae (1985)
Part 5: Solanaceae, Retziaceae
Vol. 29: Scrophulariaceae
Vol. 30: Parti: Bignoniaceae, Pedaliaceae, Martyniaceae, Orobanchaceae
Part 2: Gesneriaceae, Lentibulariaceae
Part 3: Acanthaceae: Fascicle 1: Justiciinae (1995)
Acanthaceae (remaining genera), Myoporaceae
Vol. 31: Part 1: Fascicle 1: Plantaginaceae (1998 in Bothalia 28: 151-157), Rubiaceae (Rubioideae — First part)
Fascicle 2: Rubiaceae (Rubioideae — Second part): Paederieae, Anthospermeae, Rubieae (1986)
Fascicle 3: Ixoroideae, Chinchonoideae
Part 2: Valerianaceae, Dipsacaceae, Cucurbitaceae
Vol. 32: Campanulaceae, Sphenocleaceae (2000 in Bothalia 30: 31-33), Lobeliaceae, Goodeniaceae
Vol. 33:
Asteraceae: Part 1:
Part 2:
Part 3:
Part 4:
Part 5:
Part 6:
Part 7:
Part 8:
Part 9:
Lactuceae, Mutisieae, ‘Tarchonantheae’
Vemonieae, Cardueae
Arctotideae
Anthemideae
Astereae
Calenduleae
Inuleae: Fascicle 1: Inulinae
Fascicle 2: Gnaphaliinae (First part) (1983)
Heliantheae, Eupatorieae
Senecioneae
FSA CONTRIBUTIONS IN BOTHALIA
FSA contributions 1: Aquifoliaceae. S. ANDREWS. 1994. Bothalia 24: 163-166.
FSA contributions 2: Asphodelaceae/Aloaceae, 1029010 Chortolirion. G.F. SMITH. 1995. Bothalia 25: 31-33.
FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia. G.F. SMITH. 1995. Bothalia 25: 35, 36.
FSA contributions 4: Agavaceae. G.F. SMITH & M. MOSSMER. 1996. Bothalia 26: 31-35.
FSA contributions 5: Buxaceae. H.F. GLEN. 1996. Bothalia 26: 37^40.
FSA contributions 6: Orchidaceae: Holothrix. K.L. IMMELMAN. 1996. Bothalia 26: 125-140.
FSA contributions 7: Verbenaceae: Vitex. C.L. BREDENKAMP & D.J. BOTHA. 1996. Bothalia 26: 141-151.
FSA contributions 8: Ceratophyllaceae. C.M. WILMOT-DEAR. 1997. Bothalia 27: 125-128.
FSA contributions 9: Onagraceae. P. GOLDBLATT & P.H. RAVEN. 1997. Bothalia 27: 149-165.
FSA contributions 10: Trapaceae. B. VERDCOURT. 1998. Bothalia 28: 11-14.
FSA contributions 11: Zingiberaceae. R.M. SMITH. 1998. Bothalia 28: 35-39.
FSA contributions 12: Plantaginaceae. H.F. GLEN. 1998. Bothalia 28: 151-157.
FSA contributions 13: Ulmaceae. C.M. WILMOT-DEAR. 1999. Bothalia 29: 239-247.
FSA contributions 14: Cannabaceae. C.M. WILMOT-DEAR. 1999. Bothalia 29: 249-252.
FSA contributions 15: Piperaceae. K.L. IMMELMAN. 2000. Bothalia 30: 25-30.
FSA contributions 16: Sphenocleaceae. W.G. WELMAN. 2000. Bothalia 30: 31-33.
FSA contributions 17: Casuarinaceae. C.M. WILMOT-DEAR. 2000. Bothalia 30: 143-146.
A -4
FLORA OF SOUTHERN AFRICA
ALPHABETICAL LIST OF PUBLISHED TAXA
* exotic families
Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc.l (1995)
Achariaceae, Vol. 22 (1976)
Agavaceae (Bothalia 26, 1 996)
Alismataceae, Vol. 1 (1966)
Aloaceae (first part): Aloe , Vol. 5, Part 1, Fasc. 1 (2000)
Aloe, Aloaceae (first part), Vol. 5, Part 1, Fasc. 1 (2000)
Anacardiaceae: Rhus, Vol. 19, Part 3, Fasc. 1 (1993)
Andreaeaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Anthospermeae, Rubiaceae: Rubioideae (second part), Vol.
31, Part 1, Fasc. 2 (1986)
Apocynaceae, Vol. 26 (1963)
Aponogetonaceae, Vol. 1 (1966)
Aquifoliaceae (Bothalia 24, 1994)
Archidiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Asclepiadaceae: Brachystelma-Riocreuxia, Vol. 27, Part 4
(1980)
Aspalathus, Fabaceae: Papilionoideae, Vol. 16, Part 3, Fasc.
6 (1988)
Asparagaceae, Vol. 5 (1992)
Asphodelaceae: Chortolirion, Poellnitzia (Bothalia 25,
1995)
Asteraceae: Inuleae: Gnaphaliinae (first part), Vol. 33, Part
7, Fasc. 2 (1983)
Aulacomniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Aytoniaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Bartramiaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Begoniaceae, Vol. 22 (1976)
Brachystelma, Asclepiadaceae, Vol. 27, Part 4 (1980)
Brassicaceae, Vol. 13 (1970)
Bryaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Bryobartramiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Bryophyta (three fascicles published 1981, 1987, 1998: see
plan of FSA )
Burseraceae, Vol. 18 (1986)
Buxaceae (Bothalia 26, 1996)
Cactaceae, Vol. 22 (1976)
Caesalpinioideae, Fabaceae, Vol. 16, Part 2 (1977)
Calymperaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Canellaceae, Vol. 22 (1976)
Cannabaceae (Bothalia 29, 1999)
Capparaceae, Vol. 13 (1970)
Casuarinaceae (Bothalia 30, 2000)
Ceratophyllaceae (Bothalia 27, 1997)
Ceropegia, Asclepiadaceae, Vol. 27, Part 4 (1980)
Charophyta, Cryptogams ‘Vol. 9’ (1978)
Chortolirion, Asphodelaceae ( Bothalia 25, 1995)
Cleveaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Clusiaceae, Vol. 22 (1976)
Commelinaceae, Vol. 4 (1985)
Convolvulaceae, Vol. 28, Part 1 (2000)
Crassulaceae, Vol. 14 (1985)
Crotalarieae, Aspalathus, Fabaceae: Papilionoideae, Vol.
16, Part 3, Fasc.6 (1988)
Cryphaeaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Cupressaceae, Vol. 1 (1966)
Dicranaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Ditrichaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Dracaenaceae, Vol. 5 (1992)
Droseraceae, Vol. 13 (1970)
Ebenaceae, Vol. 26 (1963)
Elatinaceae, Vol. 22 (1976)
Encalyptaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Ephemeraceae, Bryophyta, Part 1, Fasc. 2 (1987)
Eriocaulaceae, Vol. 4 (1985)
Erpodiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Eustichiaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Exormothecaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Fabaceae: Caesalpinioideae, Vol. 16, Part 2 (1977)
Fabaceae: Mimosoideae, Vol. 16, Part 1 (1975)
Fabaceae: Papilionoideae, Crotalarieae, Aspalathus, Vol.
16, Part 3, Fasc. 6 (1988)
Fissidentaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Flacourtiaceae, Vol. 22 (1976)
Fontinalaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Frankeniaceae, Vol. 22 (1976)
Funariaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Gentianaceae, Vol. 26 (1963)
Gigaspermaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Gnaphaliinae (first part), Asteraceae: Inuleae, Vol. 33, Part
7, Fasc. 2 (1983)
Grimmiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Hedwigiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Hepatophyta, Part 1, Fasc. 1 (1999)
HolothrLx, Orchidaceae (Bothalia 26, 1996)
Hookeriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Hydrocharitaceae, Vol. 1 (1966)
Hydrostachyaceae, Vol. 13 (1970)
Inuleae, Asteraceae: Gnaphaliinae (first part), Vol. 33, Part
7, Fasc. 2 (1983)
Iridaceae: Ixieae (first part): Ixiinae, Tritoniinae, Vol. 7, Part
2, Fasc. 1 (1999)
Iridaceae: Syringodea, Romulea, Vol. 7, Part 2, Fasc. 2 (1983)
Ixieae (first part), Iridaceae: Ixiinae, Trinoniinae, Vol. 7,
Part 2, Fasc. 1 (1999)
Ixiinae, Iridaceae: Ixieae (first part), Vol. 7, Part 2, Fasc. 1
(1999)
A-5
Juncaceae, Vol. 4 (1985)
Juncaginaceae, Vol. 1 (1966)
Justiciinae, Acanthaceae, Vol. 30, Part 3, Fasc. 1 (1995)
Lamiaceae, Vol. 28 (1985)
Leptodontaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Leucodontaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Loasaceae, Vol. 22 (1976)
Loganiaceae, Vol. 26 (1963)
Loranthaceae, Vol. 10 (1979)
Lunulariaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Luzuriagaceae, Vol. 5 (1992)
Malpighiaceae, Vol. 18 (1986)
Marchantiaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiales, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiidae, Hepatophyta, Part 1, Fasc. 1 (1999)
Marchantiopsida, Hepatophyta, Part 1 (1999)
Meliaceae, Vol. 18 (1986)
Meteoriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Mimosoideae, Fabaceae, Vol. 16, Part 1 (1975)
Mniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Moringaceae, Vol. 13 (1970)
Myrsinaceae, Vol. 26 (1963)
Nanobryaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Najadaceae, Vol. 1 (1966)
Neckeraceae, Bryophyta, Part 1, Fasc. 3 (1998)
Ochnaceae, Vol. 22 (1976)
Oleaceae, Vol. 26 (1963)
Onagraceae (Bothalia 27, 1997)
Orchidaceae: Holothrix ( Bothalia 26, 1996)
Orthotrichaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Oxymitraceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Paederieae, Rubiaceae: Rubioideae (second part), Vol. 31,
Part 1, Fasc. 2 (1986)
Passifloraceae, Vol. 22 (1976)
Pinaceae*, Vol. 1 (1966)
Piperaceae ( Bothalia 30, 2000)
Plantaginaceae ( Bothalia 28, 1998)
Plumbaginaceae, Vol. 26 (1963)
Podocarpaceae, Vol. 1 (1966)
Podostemaceae, Vol. 13 (1970)
Poellnitzia, Asphodelaceae (Bothalia 25, 1995)
Pontederiaceae, Vol. 4 (1985)
Potamogetonaceae, Vol. 1 (1966)
Pottiaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Primulaceae, Vol. 26 (1963)
Prionodontaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Ptaeroxylaceae, Vol. 18 (1986)
Pteridophyta (1986) (for list of families, see p. v of Pteri-
dophyta volume)
Pterobryaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Ptychomitriaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Racopilaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Resedaceae, Vol. 13 (1970)
Rhabdoweisiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Rhachitheciaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Rhizogoniaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Rhus, Anacardiaceae, Vol. 19, Part 3, Fasc. 1 (1993)
Ricciaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Riocreuxia, Asclepiadaceae, Vol. 27, Part 4 (1980)
Romulea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983)
Roridulaceae, Vol. 13 (1970)
Rubiaceae: Rubioideae (second part): Paederieae, Antho-
spermeae, Rubieae, Vol. 31, Part 1, Fasc. 2 (1986)
Rubieae, Rubiaceae: Rubioideae (second part), Vol. 31, Part
1, Fasc. 2 (1986)
Rubioideae (second part), Rubiaceae, Vol. 31, Part 1, Fasc.
2 (1986)
Ruppiaceae, Vol. 1 (1966)
Salvadoraceae, Vol. 26 (1963)
Sapotaceae, Vol. 26 (1963)
Seligeriaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Simaroubaceae, Vol. 18 (1986)
Smilacaceae, Vol. 5 (1992)
Sphagnaceae, Bryophyta, Part 1, Fasc. 1 (1981)
Sphenocleaceae ( Bothalia 30, 2000)
Splachnaceae, Bryophyta, Part 1, Fasc. 2 (1987)
Stangeriaceae, Vol. 1 (1966)
Syringodea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983)
Tamaricaceae, Vol. 22 (1976)
Targioniaceae, Hepatophyta, Part 1, Fasc. 1 (1999)
Thamnobryaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Tiliaceae, Vol. 21 (1984)
Trachypodaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Trapaceae (Bothalia 28, 1998)
Tritoniinae, Iridaceae: Ixieae (first part), Vol. 7, Part 2, Fasc.
1 (1999)
Tumeraceae, Vol. 22 (1976)
Typhaceae, Vol. 1 (1966)
Ulmaceae (Bothalia 29, 1999)
Urticaceae, Vol. 9, Part: Urticaceae (2001)
Verbenaceae: Vitex (Bothalia 26, 1996)
Violaceae, Vol. 22 (1976)
Viscaceae, Vol. 10 (1979)
Vitex, Verbenaceae (Bothalia 26, 1 996)
Wardiaceae, Bryophyta, Part 1, Fasc. 3 (1998)
Welwitschiaceae, Vol. 1 (1966)
Xyridaceae, Vol. 4 (1985)
Zamiaceae, Vol. 1 (1966)
Zannichelliaceae, Vol. 1 (1966)
Zosteraceae, Vol. 1 (1966)