A conspectus of Combretum (Combretaceae) in southern Africa, with
taxonomic and nomenclatural notes on species and sections
Two subgenera of Combretum Loefl. occur in the Flora of southern Africa (FSA) region. Previous sectional classifica-
tions were assessed in view of molecular evidence and accordingly modified. Ten sections in subgen. Combretum, 25 species
and eight subspecies are recognized. Subgen. Cacoucia (Aub!.) Exell
&
Stace comprises four sections and seven species. C.
engleri Schinz, C. paniculatum Vent. and C. tenuipes Eng!.
&
Diels are reinstated as distinct species separate from C. schu-
mannii Eng!., C. microphyllum Klotzsch and C. padoides Eng!.
&
Diels, respectively. C. schumannii occurs outside the FSA
region. Records of C. adenogonium Steud. ex A.Rich., C. platypetalum Welw. ex M.A. Lawson subsp. oatesii (Rolfe) Exell
and subsp. baumii (Eng!.
&
Gilg) Exell in Botswana are doubtfu!. C. celastroides Welw. ex M.A.Lawson subsp. orientale
Exell is elevated to species level as C. patelliforme Eng!.
&
Diels. C. grandifolium F.Hoffm. is reduced to C. psidioides Welw.
subsp. grandifolium (F.Hoffm.) Jordaan. Twenty-six names are lectotypified. The type, a full synonymy, other nomenclatural
and taxonomic information, the full distribution range and a distribution map are provided for each taxon. Selected specimens
examined are given for poorly known species. Keys to subgenera, sections and species are provided.
CONTENTS
Abstract 135
Introduction 135
Materials and method. . . . . . . . . . . . . . . . . . . . . .. 136
Taxonomy 136
Key to the southern African subgenera of Cambretum 136
A. Cambretum Laefl. subgen. Cambretum . . . . . .. 137
Key to sections of subgen. Cambretum in FSA
region 137
Group 1 (I-V) 137
I. Cambretum sect. Angustimarginata . . . . . . . .. 137
Key to species of sect. Angustimarginata .... 138
II. Cambretum sect. Spathulipetala . . . . . . . . . .. 141
Key to species of sect. Spathulipetala . . . . . .. 141
III. Cambretum sect. Macrostigmatea . . . . . . . .. 142
IV Cambretum sect. Glabripetala . . . . . . . . . . .. 143
V Cambretum sect. Ciliatipetala . . . . . . . . . . . .. 143
Key to species of sect. Ciliatipetala . . . . . . .. 144
Group 2 (VI-X) .. . . . . . . . . . . . . . . . . . . . . . . . .. 147
VI. Cambretum sect. Hypacraterapsis . . . . . . . .. 147
Key to species of sect. Hypacraterapsis . . .. 148
VII. Cambre turn sect. Metallicum 149
Key to subspp. of Cambretum collinum 149
VIII. Cambretum sect. Breviramea . . . .. 151
IX. Combretum sect. Campestria 151
X. Combretum sect. Plumbea 152
B. Combretum subgen. Cacoucia 152
Key to sections of subgen. Cacaucia 152
XI. Cambretum sect. Poivrea. . . . . . . . . . . . . 153
XII. Cambretum sect. Canniventia 153
XIII. Cambretum sect. Megalantherum . . . . . . .. 155
XIV Cambretum sect. Oxystachya 155
Specimens examined 155
*
National Herbarium, South African National Biodiversity Institute,
Private Bag
XIOI,
0001 Pretoria.
t
Student affiliation: Department of
Plant Science, University of Pretoria, 0002 Pretoria.
** H.G.W.J. Schweickerdt Herbarium, Department of Plant Science,
University of Pretoria, 0002 Pretoria.
***
Molecular Systematics Laboratory, Department of Botany and
Plant Biotechnology, APK Campus, University of Johannesburg, P.O.
Box 524, 2006 Auckland Park, Johannesburg.
MS. received: 2009-09-17.
Acknowledgements 156
References 156
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 158
Combretum Loefl. belongs to Combretaceae, one of
the 14 core families of the Myrta1es (Dahlgren & Thome
1984; Sytsma et al. 2004), one that is characterized by
mainly opposite leaves and the absence of stipules or stip-
u1es being rudimentary (Stace 2007). Combretaceae is
sister to all the other 13 families combined, and diverged
early in the evolution of the Myrtales (Angiosperm Phy-
logeny website-APweb). Cambretum (excluding Quis-
qualis L.) comprises about 250 species (Bredenkamp
2000) and occurs in tropical and subtropical regions
(America, Africa, Madagascar, India, Asia, Malesia,
Australia), but is absent from the Pacific Islands (Stace
2007). Although Bredenkamp (2000) gives the distribu-
tion as excluding Australia, C trifaliatum Vent., a vigor-
ous woody climber, was discovered in northern Australia
in the 1980s (Clarkson
&
Hyland 1986; Pedley 1990).
The greatest species diversity occurs in Africa, namely
163 in sub-Saharan Africa (Klopper et al. 2006), with 43
in Gabon (Jongkind 1999), 36 in Cameroon (Liben 1983),
and about 30 in southern Africa (Jordaan 2003). It is wide-
spread in the FSA region in all countries and provinces,
except in the Free State (rare), Lesotho and Western Cape
(Bredenkamp 2000).
Members of Cambretum are mostly deciduous or
semi-deciduous (rarely evergreen) trees, shrubs, scan-
dent shrubs (scramblers), subshrubs with woody root-
stocks (so-called geoxylic suffrutices; rare in southern
Africa) or woody climbers (lianas), sometimes with
spine-tipped lateral shoots (C imberbe Wawra). Bark
on young stems is often flaking and peeling in stringy
strips or threads in most species or in large ± cylindri-
calor hemicylindrical pieces revealing an exposed cin-
namon-red surface (C psidioides group). Leaves are
opposite, subopposite (or locally alternate), sometimes
3- or 4-whorled, exstipu1ate, simple and the margins are
always entire, rarely crenulate, or sometimes undulate
(c.
elaeagnoides Klotzsch, C. petrophilum Retief and C.
tenuipes Engl. & Diels). Indumentum on leaves, flow-
ers and fruit consists of unicellular, compartmented or
combretaceous hairs (sharp-pointed, thick-walled with
a bulbous base), multicellular stalked glands and mul-
ticellular scales. Mature scales can be classified into
three major groups and have proved to be important in
assessing taxon boundaries and phylogenetic relation-
ships (Exell
&
Stace 1972). Leaves are pinnately veined
where the lateral veins are arranged parallel to each
other, somewhat spaced and looping before they reach
the margin (brochidodromous). Hair-tuft domatia (mar-
supiiform) in the axils of the veins below are present
in a number of species (Stace 1965). Bases of leaf peti-
oles may persist as straight spines or recurved hooks as
in C. bracteosum (Hochst.) Brandis, C. mossambicense
(Klotzsch) Engl. and C. microphyllum Klotzsch. Flow-
ers are bisexual and are borne in axillary or terminal
branched or unbranched spikes, sometimes subcapitate,
and are bracteate. Flowers are 4- or 5-merous and usu-
ally sweetly scented. Petals vary from white, cream-
coloured, yellow, yellow-green in most species, but
are sometimes pale to deep pink or bright red as in C.
bracteosum, C. microphyllum, C. paniculatum Vent., C.
platypetalum Welw. ex M.A.Lawson and C. wattii Exell.
In deciduous species, the flowers appear before or with
the new leaves, e.g. C. elaeagnoides Klotzsch, C. micro-
phyllum, C. platypetalum, C. psidioides and C. zeyheri
Sond. The calyx is produced into a short, campanulate
or cup-shaped limb above the inferior ovary. Stamens
are inserted on the hypanthium, usually twice as many
as the sepals or petals, and usually exserted beyond the
petals. The stamens vary in colour from yellow, orange,
pinkish, crimson or reddish to red-brown. A glabrous or
pilose, green or red, well-developed nectariferous disc is
often present at the base of the upper hypanthium. Nec-
tar production is indicative of flowers that are pollinated
by a wide range of insects or birds (Stace 2007). The
ovary is inferior and I-locular with two pendulous ana-
tropous ovules of which only one develops into a seed.
The fruit is glabrous or covered with scales and/or hairs
and is mainly a 4-winged, or occasionally 5-winged
(c.
mossambicense, C. oxystachyum, C. wattii) indehiscent
samara, except in C. bracteosum which has wingless
fruit (nuts). In most cases winged fruit are wind-dis-
persed and the wingless fruit seem to be an adaptation to
water dispersal (Exell & Stace 1972).
Despite extensive taxonomical and anatomical Shld-
ies on Combretum in tropical and southern Africa by
Engler
&
Diels (1899), Dummer (1913), Stace (1961;
1965; 1969; 1980; 1981), Exell (1968; 1978), Verhoeven
&
Van del' Schijff (1973), Wickens (1973), Van Wyk
(1984), Ca.IT(1988), Rodman (1990) and Tilney (2002),
there are still taxonomic and nomenclatural problems
remaining, as well as new taxa to be described. Some
names are misapplied and the identity of some taxa in
southern African herbaria is uncertain. Recent molecular
work (Maurin et af. 2010) has indicated that taxa bound-
aries need revising to reflect more accurately the phy-
logeny of Combretum and its allies. Maurin et al. (2010)
deal mostly with the subgeneric, sectional and generic
delimitation of Combretum, whereas the present paper
deals with species delimitation, the status elevation of
infraspecific taxa and lectotypification of some names.
All material of Combretum in the National Herbar-
ium, Pretoria (PRE) and H.G.W.I Schweickerdt Her-
barium, University of Pretoria (PRU) was examined.
Two websites were consulted for type material: 1, www.
aluka.org and 2, the Zurich Herbarium: www.zuerich-
herbarien.unizh.ch. Types seen electronically are cited as
e! Where the holotype was destroyed in the Berlin Her-
barium (B) during World War II, lectotypification is cov-
ered by Article 9.15 of the Code (McNeill et af. 2006)
which provides for narrowing the lectotype to a single
specimen. Acronyms of herbaria where types are housed
follow Holmgren et af.
(1990).
Sectional classifications of Engler & Diels (1899),
Exell (1978) and Stace (1981) were assessed in view of
the phylogenetic studies done by Maurin et af. (2010)
and adjusted where necessary. Sections are arranged
from ancestral to derived according to the mentioned
molecular studies. C. mkuzense is treated as belonging
to sect. Spathulipetala. Species are arranged alphabeti-
cally within each section. An index is provided at the
end for easy access to species. The following species
are widespread, well defined and well known or recently
described and no voucher specimens are cited for them:
C. apiculatum Sond. subsp. apiculatum, C. elythrophyl-
lum (Burch.) Sond., C. hereroense Schinz, C. imberbe,
C. kraussii Hochst., C. molle R.Br. ex G.Don, C. vendae
A.E.van Wyk and C. zeyheri Sond. Geographical dis-
tributions mentioned in the keys refer primarily to the
FSA region, namely Botswana, Lesotho, South Africa
(RSA: Eastern Cape, Free State, Gauteng, KwaZulu-
Natal, Limpopo, Mpumalanga, Northem Cape and
North- West), Namibia and Swaziland. Species OCCUlTing
north of the FSA region are indicated by an arrow on the
distribution maps. Full range distribution, including all
countries and the relevant provinces of South Africa, is
given alphabetically under each species.
Combretum is divided into three subgenera, namely
subgen. Combretum, subgen. Cacoucia (Aubl.) Exell
&
Stace and subgen. Apetalanthum Exell
&
Stace (Stace
1981). The last-named is represented by only one spe-
cies, C. apetalum Wall. ex Kurz which is restricted to
la Scales present though sometimes inconspicuous or hidden by hairs; microscopic stalked glands absent; flowers and fruit usually 4-mer-
ous; petals cream-coloured, white, yellow or greenish, usually not red,
<
3.5 mm long. . subgen. Combretum (p. 137)
1b Scales absent; microscopic stalked glands present; flowers and fruit 5-merous or if 4-merous then petals red; petals> 4
111111
long
. . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subgen.
Cacoucia
(p.
152)
Asia. It has leaves with scales and stalked glands, pet-
als are absent and it has 10 stamens in two whorls. The
first two subgenera have scales or stalked glands, pet-
als are present, although sometimes reduced and they
have eight stamens in one or two whorls. Subgenera
are mainly based on the presence or absence of petals,
presence or absence of scales on the leaves, presence or
absence of stalked glands, the flowers being 4- or 5-mer-
ous, colour of the flowers and length of petals.
Scales present, although sometimes inconspicuous
or hidden by combretaceous hairs; microscopic stalked
glands absent; flowers usually 4-merous; petals usually
not red; stamens usually 8, in
I
or 2 whorls; fruit usu-
ally 4-winged. The structure and arrangement of scales
are of great taxonomic significance (Stace 1965). The
southern African material is divided into ten sections
(Engl. & Diels 1899; Wickens 1973; Exell 1978; Stace
1980, 1981; Rodman 1990) which fall into two groups.
The sectional grouping is based on the size of the scales
on the leaves and the resultant two groups are congru-
ent with the two major groups recovered in phyloge-
netic analyses (Maurin
et al.
2010). Each group is rep-
resented by five sections. The first five sections treated
here belong to the group with small, inconspicuous
scales, usually smaller than 100 flm,
±
circular in outline
or slightly scalloped in sect. Ciliatipetala and divided by
few radial and tangential walls. Sometimes the tangen-
tial walls are absent as in sect. Angustimarginata, sect.
Glabripetala and sometimes in sect. Ciliatipetala. (Fig-
ure 1). The last five sections have conspicuous scales,
usually larger than 100 flm, scalloped or irregularly
undulate in outline and the scales are divided by many
radial and tangential walls (Figure 2). C. imberbe is
placed in its own section, sect. Plumbea (Maurin et al.
2010). Note that although 100 flm is taken as the cut-off
measurement to separate the two groups, intermediates
do occur.
Ia Scales inconspicuous, usually
<
100 flm in diam.; scales divided by few radial and tangential walls (tangential walls sometimes absent);
stamens I-seriate (Group 1, p. 137) (Figure 1):
2a Petal apex ciliate or pilose; petals small, 0.5-1.5 mm long (if apex not ciliate then petals
<
1 mm long as in C. petrophilum); leaf api-
ces often apiculate; fruit 15-30 mmlong . V. sect. Ciliatipetala (p. 143)
2b Petal apex glabrous; petals 1.5-2.5 mm long; leaf apices not apiculate; fruit 15-50(-100) mm long:
3a Fruit up to 22 mm long; often with hair-tuft domatia in axil of veins below (except in C. erythrophyllum and C. vendae):
4a Spring leaves at least partly cream-co loured or with some leaves turning bright red in autumn; petals narrowly spathulate; disc
margin pilose; style without expanded stigma; fruit sparsely to moderately hairy, not lepidote, mature fruit tinged pink to dark
red (rare in C. erythrophyllum); stipe 4-8 mm long . . . . . . . . . . . . . I. sect. Angustimarginata (p. 137)
4b Spring leaves not cream-co loured and autumn leaves not reddish; petals broadly spathulate; disc glabrous with only a very short
free margin; style sometimes with expanded stigma; fruit green, glabrous and glutinous, yellowish green, with reddish brown
scales, giving it a satiny sheen, wings papery; stipe up to 10 mm long. . III. sect. Macrostigmatea (p. 142)
3b Fruit 35-50(-100) mm and longer; hair-tuft domatia in axil of veins below absent:
5a Leaves large, 100-200 mm long; style without swollen apex; fruit up to 35 mm long; glutinous when young, glabrous, yellowish
green tinged reddish brown; stipe up to 7 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . IV. sect. Glabripetala (p. 143)
5b Leaves medium-sized, 30-100 mm long; style with swollen apex; fruit up to 50(-100) mm long, pale green, glutinous only on
body, sparsely hairy or glabrous, drying pale brown, straw-co loured or limegreen; stipe 10-30 mm long .
. . . . . . . . . . . . . . . . . . . . . . II. sect. Spathulipetala (p. 141)
Ib Scales conspicuous and large, usually> 100 flm in diam. (84-160 flm in sect. Breviramea); scales divided by many radial and tangen-
tial walls; stamens 1- or 2-seriate (Group 2, p. 147) (Figure 2):
6a Trees; petals subcircular; fruit dark brown or reddish grey to dark purple, glabrous to densely hairy, metallic in appearance, often acute
at apex; disc with free pilose margin; cotyledons arising below soil level on a stalk formed by connate petioles .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VII. sect. Metallicum (p. 149)
6b Trees, shrubs, scrambling shrubs or woody climbers; petals linear-elliptic, obovate to spathulate; fruit densely rufous, golden or sil-
very lepidote, otherwise glabrous; disc without free margin; cotyledons arising above or below soil level:
7a Inflorescences usually terminal panicles of spikes or branched spikes; upper hypanthium little developed, flattened; stamens I-seri-
ate, inserted at margin of disc; fruit usually
<
20 mm long; stipe 1-3 mm long; cotyledons arising above or below soil level; scales
contiguous or not:
8a Scrambling shrubs or woody climbers, usually multi-stemmed, not with spine-tipped short branches; disc glabrous or pilose at
least on margin; style without stalked scales; fruit not densely silvery lepidote; stipes
<
2 mm long; cotyledons borne above soil
level; scales conspicuous but not contiguous or overlapping Vr. sect. Hypocrateropsis (p. 147)
8b Erect trees up to 15 m, occasionally up to 30 m tall, single-stemmed, with short lateral branches often spine-tipped; disc margin
densely tomentose; style with stalked scales; fruit densely silvery lepidote; stipe 2-3 mm long; cotyledons arising below soil
level; scales conspicuous, contiguous and/or overlapping . .. X. sect. Plumbea (p. 152)
7b Inflorescences usually axillary unbranched spikes; upper hypanthium well developed, campanulate or cup-shaped, not flattened; sta-
mens 2-seriate; fruit 20-35 mm long; stipe up to 11(-15) mm long; cotyledons arising above soil level; scales contiguous or overlapping:
9a Leaves with only 3 or 4(5) pairs of primary lateral veins; reticulate venation conspicuous below; hair-tuft domatia absent in axils
of veins below; leaf margin flat, usually ciliate; flowers long hairy, not conspicuously lepidote; disc with free pilose margin; fruit
densely reddish or golden lepidote . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. VIII. sect. Breviramea (p. 151)
9b Leaves with 9-13 pairs of primary lateral veins; only midrib and primary lateral veins conspicuous and prominently raised below;
hair-tuft domatia in axial of veins below; leaf margin often undulate, glabrous; flowers not bairy but densely and conspicuously
lepidote; disc without free margin; fruit silvery lepidote. . ..... IX. sect. Campestria (p. 151)
Species with small scales, usually smaller than 100
flm, divided by few radial and tangential walls or tan-
gential walls sometimes absent (Figure 1).
1. Combretum sect. Angustimarginata Engl.
&
Diels (1899)
This section is a natural taxon of six closely related
species restricted to Zimbabwe, Mozambique, Botswana,
FIGURE I.-Scales of Combretum subgen. Combretum. Group I.
Sections I-V, showing smaller size, ± circular in outline and
divided by few radial and tangential walls, and sometimes tan-
gential walls absent as in sect. Angustimarginata, sect. Glabri-
petala and sometimes in sect. Ci/iatipetala. A,
e.
elythrophyl-
lum (sect. Angustimarginala); B,
e.
zeyheri (sect. Spalhu/ipeta-
la); C,
e.
engleri (sect. Macrosligmalea); D,
e.
adenogonium
(sect. Glabripetala); E,
e.
apicl/latl/m (sect. Cilialipelala); F,
e.
albopunclalum (sect. Cilialipelala); G,
e.
molle (sect. Ci/iatip-
etala); H,
e.
mol/e. Taken from Stace (1969) and Exell (1978).
Scale bar: 80 /lm. Artist: Daleen Roodt.
South Africa and Swaziland. It is characterized by their
4-merous flowers; upper hypanthium cupulifol111 to cam-
panulate; petals subcircular, obovate, spathulate or narrowly
elliptic, glabrous, margins not ciliate; stamens 8, I-seriate,
inserted shortly above margin of disc; disc glabrous with
a pilose margin very shortly free for up to 0.5 mm; style
not expanded; fruit 4-winged; scales inconspicuous, often
obscured by hairs and/or glutinous secretions,
±
50-75
fUll
in diam., weakly scalloped, delimited by 8-16 primary
radial walls alone; cell walls very thin (Exell 1970, 1978).
Other diagnostic characters based on field obser-
vations are: the bark is ± smooth or flaking in small,
rather papery pieces in older specimens; the first spring
leaves are often partly or completely cream-coloured or
at least some leaves turning red in autumn; the flowers
have calyx lobes tinged reddish purple; the mature fruits
are usually partly or completely tinged pink to dark red
(Van Wyk 1984). The secondary xylem of older stems
has islands of interxylary phloem (Verhoeven
&
Van der
Schijff 1973).
Engler
&
Diels (1899) and Exell (1978) included
C. woodii DUmmer under C. kraussii Hochst. Rodman
(1990) considers C. woodii to be conspecific with C.
erythrophyllum. Burtt Davy (1926), Van Wyk (1984),
Carr (1988) and Jordaan (2003, 2006) consider C. wood-
ii to be a distinct species. There are ample DNA and
morphological differences to separate C. woodii from C.
kraussii and C. erythrophyllum (Maurin et al. 2010). See
Carr (1988) for differences between these three species.
Exell (1970, 1978) included C. nelsonii and C. woodii in
the synonymy of C. kraussii. Chemosystematic studies
conducted by Call' & Rogers (1987) support the recog-
nition of C. nelsonii DUmmer, C. !a-aussii and C. wood-
ii as three distinct species. C. caffrum (Eckl. & Zeyh.)
Kuntze and C. vendae also belong to this section. A fornl
of Combretum vendae with glabrous leaves occurs in the
western Soutpansberg and represents a new subspecies
to be described elsewhere.
Key to species of Combretum sect. Angustimarginata
based on Van Wyk (1984)
la Leaves glabrous below when mature, often with hair-111ft
domatia:
2a Flowers in laxly elongated spikes (25-)35-60(-85) mm
long:
3a Leaf lamina with secondary lateral veins raised but inter-
secondary veins
±
plane below; scales with 8-10 radial
cells; trees; forest; widespread 3.
e.
kraussii
3b Leaf lamina with both secondary and intersecondary
veins conspicuously raised below; scales with 10-16
radial cells; trees, shrubs or climbers; forest and
savanna; Limpopo Province, Mpumalanga, KwaZulu-
Natal and Swaziland . . . . . . . . . 6.
e.
woodii
FIGURE 2.-Scales of Combretl/m
subgen. COlllbrell/lII. Group
2. Sections VI-X, showing
larger size,
±
scalloped in
outline and divided by many
radial and tangential walls. A,
e.
pate//iforme (sect. Hypo-
crateropsis);
B,
e.
celaslroi-
des (sect. Hypocrateropsis);
C,
e.
collinum (sect. Meta//i-
cl/m); D,
e.
hereroense (sect.
Breviramea);
E,
e.
elaeag-
noides (sect. Campestria); F,
e.
imberbe (sect. Pll/mbea).
Scale bar: 80 /lm. AJiist:
Daleen Roodt.
I( '~\
)/, I
__ ~ _,-,"---i_
2b Flowers in congested subcapitate spikes (10-)15-20(-35)
mm long:
4a Usually shrubs; leaves elliptic, obovate-elliptic or obo-
vate; flowers with upper hypanthium divided into a
lower
±
campanulate part containing the disc, and an
expanded
±
cupuliform upper part; Limpopo Province
and Mpumalanga 4. C. nelsonii
4b Usually trees; leaves narrowly elliptic or lanceolate;
flowers with upper hypanthium
±
cupuliform, not
divided into a lower and upper part; Eastern Cape
. . . . . . . . . . . . . . . . . . . . . . . . . . .. I. C. cajJrum
1b Leaves distinctly hairy below when mature, at least on mid-
rib and lateral veins:
5a Reticulate venation of leaf lamina conspicuously raised
below; Venda (Limpopo Province) 5. C. vendae
5b Reticulate venation of leaf lamina plane or slightly raised
below; widespread along watercourses ..
......... 2. C.
erythrophyllum
1.
Combretum caffrum (Eck!. & Zeyh.) Kuntze,
Revisio generum p1antarum 3,2: 87 (1898); lD.Carr:
40 (1988); M.Coates Palgrave: 797 (2002). Type: South
Africa, Eastern Cape, 'ad ripas fluminum Kat et Visch-
rivier (Albany), Zondagsrivier (Uitenhage), Keys-
kamma et Keyrivier (Caffraria)', Eck/on
&
Zeyher 421
[SAM0036373-2, 1ecto. e!, designated here; FR e!, HBG
e!, K e!, Me!, We!, isolecto.].
D. conglomerata Eckl.
&
Zeyh.: 55 (1834-1835). Type: South
Africa, Eastern Cape, 'Inter frutices fluvii Katrivier prope Fort Beau-
fort Caffraria terminis', Ecklon
&
Zeyher 422 [SAM0036375-2 e!,
Iecto., designated here; K e!, Me!, isolecto.].
C. salicifolium E.Mey. ex Hook.: 1. 592 (1843); Sond.: 511 (1862);
DUmmer: 182 (1913). Type: South Africa, Eastern Cape, 'Sundays
River', Burke 592 (K, holo. e!; BM, iso. ell.
FIGURE 3.-Seedling morphology
of Combretum zeyheri (A)
and C. mkuzense (B). Solid
transverse Iines depict ground
level. In C. zeyheri cotyledons
fused to form a subcircular
peJtate structure which termi-
nates primary stem. Growth
in length is resumed by a lat-
eral bud from a point below
fused cotyledons (sympo-
dial growth). In C. mkuzense
growth in length of stem com-
mences from what seems to
be the primary growing tip
located between two oppo-
site cotyledons (monopodial
growth). Scale bar: 10 mm
Artist: Daleen Roodt.
C. dregeanum C.Presl: 73 (1844). Type: South Africa, Eastern
Cape, Klein Winterhoek between Zoutpansnek and Enon, Drege 6849a
(?PR, holo.; BM e!, HBG e!, K e!, MO e!, SAM e!, We!, iso.).
Note: Eck/on
&
Zeyher 421,422, both at SAM, are cho-
sen as lectotypes for Combretum caffrum (= Dodonaea caf
Fa), and D. cong/omerata respectively, because they have
the protologues and annotations attached to the specimens.
2. Combretum erythrophyllum (Burch.) Sond.
in Linnaea 23: 43 (1850); Sond.: 509
(J
862); Eng!.
&
Diels: 26 (1899); DUmmer: 140 (1913); Bews: 146
(1921); Exell
&
Roessler: 8 (1966); Exell: 7 (1968);
Exell: 112 (1978); lD.Carr: 69 (1988); Pooley: 356
(1993); A.E.van Wyk
&
P.van Wyk: 330 (1997); McCle-
land: 460 (2002); M.Coates Palgrave: 802 (2002). Type:
South Africa, Northern Cape, 'on the banks of the Ky-
gariep' (Griqualand West, Kalahari region, Herbert Div.,
right bank of the Vaal River at Blaauwbosch Drift),
Burchell1749 (K, holo. e!; PRE, iso.!).
C. glomeruliflorum Sond.: 47 (1850); Sond.: 509 (1862); Engl.
&
Diels: 26 (1899); Burtt Davy: 247 (1926); Codd: 130 (1951). Type:
South Africa, Natal [KwaZulu-Natal], Port Natal [Durban], Glieinzilis
62 (?S, holo.; HBG e!, K e!, P e!, We!, iso.).
C. riparillm Sond.: 47 (1850); Sond.: 511 (1862). C. glomeruliflor-
urn Sond. var. riparium (Sond.) Burtt Davy: 247 (1926); O.B.Mill.: 42
(1948). Type: South Africa, [?Gauteng], 'On Magalisriver', Zeyher
549 (S, holo. e!; BM, K e!, SAM, iso.).
C. sonderi Gerrard ~;:-Sond.: 511 (1862). Type: South Africa, Natal
[KwaZulu-Natal], 'On the Nototi River, near Port Natal', Gerrard 138
(?TCD, holo.).