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Monograph of Coccinia (Cucurbitaceae) 1
Monograph of Coccinia (Cucurbitaceae)
Norbert Holstein1
1 Nees-Institute for Biodiversity of Plants, Meckenheimer Allee 170, 53115 Bonn, Germany
Corresponding author: Norbert Holstein (holstein@lrz.uni-muenchen.de)
Academic editor: Sandra Knapp|Received 25 April 2012|Accepted 23 September 2014|Published 3 August2015
Citation: Holstein N (2015) Monograph of Coccinia (Cucurbitaceae). PhytoKeys 54: 1–166. doi: 10.3897/
phytokeys.54.3285
Abstract
is monograph deals with all 95 names described in the Cucurbitaceae genus Coccinia and recognizes
25 species. Taxonomic novelties are Coccinia adoensis var. aurantiaca (C.Jerey) Holstein, stat. nov.,
C. sessilifolia var. variifolia (A.Meeuse) Holstein, stat. nov., and C. adoensis var. jereyana Holstein,
var.nov. For the 25 species 3157 collections were examined, of which 2024 were georeferenced to produce
distribution maps. All species are distributed in sub-Saharan Africa with one species, C. grandis, extending
from Senegal in West Africa east to Indonesia and being naturalized on Pacic Islands, in Australia, the
Caribbean, and South America. Coccinia species are dioecious creepers or climbers with simple or bid
tendrils that occupy a range of habitats from arid scrubland, woodlands to lowland rainforest and mist
forest. e corolla of Coccinia species is sympetalous, usually pale yellow to orange, and 1 to 4.5 cm
long. Pollination is by bees foraging for pollen or nectar. After pollination, the developing ovary often
exhibits longitudinal mottling, which usually disappears during maturation. All species produce berries
with a pericarp in reddish colors (orange-red through to scarlet red), hence the generic name. e globose
to cylindrical fruits contain numerous grayish-beige at to lenticular seeds. Chromosome numbers are
2n=20, 24, and 22 + XX/XY. Many Coccinia species are used for food, either as roasted tubers, greens as
spinach, or the fruits as vegetables. Medicinal value is established in C. grandis, of which leaves and sap
are used against diabetes.
Keywords
Cucurbitaceae, Coccinia, molecular phylogeny, biogeography, taxonomy, morphology, sex expression,
useful plants
PhytoKeys 54: 1–166 (2015)
doi: 10.3897/phytokeys.54.3285
www.phytokeys.com
Copyright Norbert Holstein. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Norbert Holstein / PhytoKeys 54: 1–166 (2015)
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Table of contents
Introduction ............................................................................................................. 4
Materials and methods ............................................................................................. 4
General morphology .......................................................................................... 4
Phylogenies ........................................................................................................ 5
Distribution maps .............................................................................................. 5
Morphology and anatomy ........................................................................................ 6
Habit ................................................................................................................. 6
Roots ................................................................................................................. 6
Hypocotyl and shoots ........................................................................................ 7
Cotyledons ......................................................................................................... 9
Leaves ................................................................................................................ 9
Probracts and bracts ......................................................................................... 13
Extranuptial glands .......................................................................................... 13
Peduncles and pedicels ..................................................................................... 14
Perianth ........................................................................................................... 15
Androecium ..................................................................................................... 17
Pollen ............................................................................................................... 18
Gynoecium ...................................................................................................... 19
Female gametophyte development and embryology.......................................... 19
Fruits ............................................................................................................... 19
Seeds ................................................................................................................ 21
Germination .................................................................................................... 23
Genome, chromosomes, and hybridization ............................................................. 23
Chromosomes and sex determination ............................................................... 23
Genome of C. grandis ....................................................................................... 26
Hybridization and crossing experiments ........................................................... 26
Plant – biotic environment interactions .................................................................. 29
Pollination ....................................................................................................... 29
Seed dispersal ................................................................................................... 30
Interaction with ants ........................................................................................ 31
Diseases and parasites ....................................................................................... 31
Use, economic potential, and phytochemistry .................................................. 34
Evolution and phylogeny ........................................................................................ 36
Identication of Coccinia species ............................................................................ 41
Possible confusion with other genera ................................................................ 41
Characters for species discrimination ................................................................ 41
Key to Coccinia species ..................................................................................... 41
Taxonomic treatment ............................................................................................. 54
1. Coccinia abyssinica (Lam.) Cogn. ........................................................... 56
2a. Coccinia adoensis (Hochst. ex A.Rich.) Cogn. var. adoensis ................... 59
2b. Coccinia adoensis var. aurantiaca (C.Jerey) Holstein ........................... 67
Monograph of Coccinia (Cucurbitaceae) 3
2c. Coccinia adoensis var. jereyana Holstein .............................................. 70
3. Coccinia barteri (Hook.f.) Keay .............................................................. 72
4. Coccinia grandiora Cogn. ..................................................................... 75
5. Coccinia grandis (L.) Voigt ..................................................................... 78
6. Coccinia heterophylla (Hook.f.) Holstein ................................................ 89
7. Coccinia hirtella Cogn. ........................................................................... 92
8. Coccinia intermedia Holstein .................................................................. 94
9. Coccinia keayana R.Fern. ....................................................................... 95
10. Coccinia longicarpa Jongkind ............................................................... 97
11. Coccinia mackenii Naudin ex C.Huber ................................................ 99
12. Coccinia megarrhiza C.Jerey ............................................................. 102
13. Coccinia microphylla Gilg ................................................................... 104
14. Coccinia mildbraedii Gilg ................................................................... 107
15. Coccinia ogadensis ulin ................................................................... 109
16. Coccinia pwaniensis Holstein .............................................................. 111
17. Coccinia quinqueloba (unb.) Cogn. ................................................ 113
18. Coccinia racemiora Keraudren .......................................................... 114
19. Coccinia rehmannii Cogn. .................................................................. 115
20. Coccinia samburuensis Holstein .......................................................... 122
21. Coccinia schliebenii Harms ................................................................. 123
22. Coccinia senensis (Klotzsch) Cogn. ...................................................... 124
23a. Coccinia sessilifolia (Sond.) Cogn. var. sessilifolia ............................... 129
23b. Coccinia sessilifolia var. variifolia (A.Meeuse) Holstein...................... 132
24. Coccinia subsessiliora Cogn. .............................................................. 133
25. Coccinia trilobata (Cogn.) C.Jerey .................................................... 135
Insuciently known taxa ............................................................................... 137
Dubious names .............................................................................................. 138
Invalid names ................................................................................................. 140
Exluded taxa .................................................................................................. 145
Acknowledgements ............................................................................................... 150
References ............................................................................................................ 151
Supplementary material 1 ..................................................................................... 165
Supplementary material 2 ..................................................................................... 165
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
4
Introduction
Coccinia Wight & Arn. comprises 25 species and is the 11th largest of the 97 genera
of the Cucurbitaceae (Schaefer and Renner 2011a). Especially in the 19th century, it
drew gardeners’ attention probably because of its striking fruits (André 1900; Edwards
1815; Huber 1864; 1865; Koch 1865; Sims 1816). All species are dioecious, and one
species, C. grandis (L.) Voigt, has heteromorphic sex chromosomes and therefore has
been studied cytologically (Agarwal and Roy 1975; 1983; 1984; Bhaduri and Bose
1947; Chakravorti 1948; Datta 1988; Kumar and Deodikar 1940; Kumar and Vish-
veshwaraiah 1952; Roy 1974; Roy and Roy 1971b). e last complete taxonomic
treatment of Coccinia is by Cogniaux (1881), more than 130 years ago. Since then,
16 new species have been described, and the genus has only been revised regionally
(Hutchinson et al. 1954; Jerey 1967; 1978; 1995; Jerey and Fernandes 1986; Ké-
raudren-Aymonin 1975a; Kéraudren 1967; Meeuse 1962). e position of Coccinia in
the Benincaseae has been conrmed by molecular data (Kocyan et al. 2007; Schaefer
and Renner 2011b), and the monophyly has been tested with almost complete species
sampling by Holstein and Renner (2011b).
e delimitation of Coccinia from other genera is dicult. e scarlet-red fruits to
which the genus name – Coccinia from Latin coccineus – refers are also found in other
African genera, such as Eureiandra Hook.f. erefore, it is not surprising that early
botanists described several species now considered to belong to Coccinia in other genera
(Cephalandra Schrad. ex Eckl. & Zeyh., Physedra Hook.f., and Staphylosyce Hook.f.),
or species described in Coccinia now belong to dierent genera. In all, 113 names at
various ranks have been proposed for what are here considered 25 species. e species
concepts in the present revision are based on 3157 herbarium collections and eldwork
in Tanzania, geo-referencing of 2024 collections and cultivation of 10 species in the
greenhouse. In combination, plastid and nuclear data obtained for multiple accessions
representing most species and ecological information coming from the mapping eort
provide a modern understanding of the evolution and species relationships in Coccinia.
Materials and methods
General morphology
During this study, the present author examined 3157 herbarium collections from 39
herbaria from (physical or digital) loans or in situ (B, BM, BR, BRI, C, CANB, CBG,
COI, DSM, EA, FR, FT, G, GAT, GOET, H, HBG, HEID, JE, K, L, LISC, LISU, M,
MO, MSB, NHT, P, PERTH, PR, PRC, S, U, UBT, UPS, W, WAG, Z, ZT). Additional
collections were obtained via personal communication (BO, DNA, TUB, US), from
web pages such as like JStor Plant Science (JPS, http://plants.jstor.org/), Chinese Virtual
Herbarium (CVH, http://www.cvh.org.cn/), and homepages of the following herbaria:
A, AAU, BAR, CAY, FLAS, FSU, FTG, GH, HAST, NTUF, NY, PDA, TAIF, and USF.
Monograph of Coccinia (Cucurbitaceae) 5
New collections were added if the photograph allowed identication or if misidentica-
tion appeared to be unlikely (esp. C. grandis collections from the Pacic area), while du-
plicates were added without visual inspection of the specimen photo. Online availability
of specimen images is mentioned in the list of exsiccatae (Suppl. material 1). Ten species
were cultivated in greenhouses of Munich Botanical Garden: C. abyssinica (Lam.) Cogn.
(1 origin, 4 individuals), C. adoensis var. jereyana (1 origin, 4 individuals), C. grandiora
Cogn. (1 origin, 1 individual), C. grandis (3 origins with 2, 61, 3 individuals, respec-
tively), C. hirtella Cogn. (1 origin, 12 individuals), C. megarrhiza C.Jerey (1 origin, 8
individuals), C. microphylla Gilg (2 origins, 2 and 1 individual, respectively), C. rehman-
nii Cogn. with two varieties (2 origins, each 3 individuals), C. sessilifolia (Sond.) Cogn.
var. sessilifolia (1 origin, 8 individuals), and C. trilobata (Cogn.) C.Jerey (2 origins, each
3 individuals). e present author performed crossing experiments among eight of these
species. Morphological features were documented photographically and in the form of
vouchers, with 47 collections deposited in M. Field data were obtained on a trip to NE
Tanzania in 2009 resulting in 28 Coccinia collections.
Phylogenies
For this monograph the phylogenetic data of Holstein and Renner (2011a; 2011b) were
augmented with 20 new sequences from 8 accessions (GenBank accession numbers are
given in Suppl. material 2), and new phylogenies were calculated using RAxML v. 7.2.6
(Stamatakis 2006) and MrBayes v. 3.2 (Ronquist et al. 2012). e substitution model was
GTR+Γ as used before, and 1000 ML replicates were used to infer statistical support for
the nodes via bootstrapping. For Bayesian analysis, four chains were run with 2,000,000
generations, with a sampling frequency of 1000. e rst 25% of the trees were discarded
as burn-in, and the rest were plotted as a 50% majority rule consensus tree using FigTree
1.3.1 (http://tree.bio.ed.ac.uk/software/gtree/). Gaps in the plastid matrix occurring in
more than one accession were coded as “0”, “1”, or “?”, with “?” when data were missing
or when shorter gaps were coded in the same place, but in dierent accessions.
Distribution maps
Of the examined collections, 2024 were geo-referenced and mapped in Google Earth
(Google Inc., Mountain View, CA, USA). Cultivated plants were geo-referenced ac-
cording to the original collection site. If collecting sites were given as distances from
locations, a path along major roads was used, beginning from the center of the starting
location. Collecting sites were geo-referenced according to the description even if co-
ordinates were given on the label, except for cases in which the coordinates were clearly
derived from GPS or if the description did not allow further improvement. Location
names were cross-validated from printed maps and then imported into DIVA-GIS
7.1.6.2 (http://www.diva-gis.org). If collecting sites of specimens appeared to be too
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
6
useful to ignore in the distribution maps despite the large uncertainty of the position
(radius > 5 km) or if the collecting site was only given as “nearby” a distinct locality,
then geo-referenced coordinates are given in brackets. Political administrative borders
were taken from GADM v.1 (Jan 2009) or v.2 (Jan 2012) (http://www.gadm.org/) and
elevation data (1 km resolution) from CGIAR Consortium for Spatial Information
(Jarvis et al. 2008). e geodetic datum for the maps is WGS84; the projection in each
case is equirectangular. Coordinates are given in decimal degrees in Suppl. material 1
with WGS84 as geodetic datum.
Morphology and anatomy
Habit
Coccinia species are perennial climbers or creepers. e lignication of the mature
shoots diers among the species from unlignied to completely lignied. Climbing is
enabled by tendrils, which are either simple or bid. Tendril development in young
plants is delayed and emerges in C. abyssinica after the 6th node (Getahun 1974a). e
tendril arms are only rarely equally sized, as one is usually much smaller; true dichot-
omy of tendrils is unknown from Coccinia. Whether a species has simple or bid ten-
drils is often not xed, but there is a strong predominance of one kind. Bid tendrils
regularly occur or are predominant in C.grandiora Cogn., C. heterophylla (Hook.f.)
Holstein, C.hirtella Cogn., C. intermedia Holstein, C.mackenii Naudin ex C.Huber,
C. mildbraedii Gilg, C. racemiora Keraudren, C.schliebenii Harms, and in some forms
of C. barteri (Holstein and Renner 2011b). Strikingly, Coccinia species with bid ten-
drils occur in rather humid habitats. is suggests an adaptive advantage, because more
tendril arms increase stability, as the leaves of rainforest species are larger, coriaceous,
and thus heavier than leaves of species from drier habitats. Some species are regularly
described as having simple tendrils in oristic treatments, but they may bare bid
tendrils such as C.sessilifolia (N. Holstein 13) and C.senensis (H.J.E. Schlieben 5745
in B, K, and MO). Coccinia adoensis has bid tendrils even in some type specimens
(e.g., G.H.W. Schimper 166 in BR8886781 and on the sheet with a drawing in K) and
is still listed as simple-tendrilled. All three species with this polymorphism, however,
have predominantly simple tendrils. Interestingly, these species are also closely related
to species with predominantly bid tendrils: C. sessilifolia with C. hirtella and C. mac-
kenii, and C. adoensis with C. grandiora and C. schliebenii.
Roots
Coccinia species have perennial roots. Most (if not all) species are woody at the base,
and most of them produce hypocotyl tubers (Fig. 1a). Some species, such as Coc-
cinia adoensis and C. grandiora (and most likely also C. senensis (Klotzsch) Cogn.
Monograph of Coccinia (Cucurbitaceae) 7
and C. schliebenii), however, produce globular subterranean root tubers, much like
potatoes, but smaller in size (Holstein and Renner 2011b; Zimmermann 1922b; pers.
observ.). Root tubers in Coccinia adoensis are likely to be an adaptation to re, as this
species predominantly occurs in woodlands. In contrast to rather mild res in semi-
arid savannas with less inammable biomass, woodland res produce temperature rises
of 60 °C in 0–3 cm depth (Bradstock and Auld 1995; Gignoux et al. 1997), so vegeta-
tive buds near the ground (hemicryptophytes) might be damaged, whereas root tubers
(geophytes) have a higher chance of survival.
Coccinia grandis and C. barteri produce adventitious roots if stems touch the soil
(Fig. 1b). Coccinia hirtella, C. sessilifolia and their F1 hybrids with C. grandis appear
to lack this ability (pers. observ.). Adventitious roots also occur along the hypocotyl
of C.abyssinica seedlings (Getahun 1974a). ere is barely any research on root anat-
omy, solely Getahun (1974a) reports tetrarch vascular bundles in the primary roots of
C.abyssinica seedlings and di- to triarch bundles in secondary roots.
Hypocotyl and shoots
Many species, such as C. abyssinica, C. grandis, C.hirtella, C. megarrhiza, C. micro-
phylla, C. rehmannii, C.sessilifolia, and C. trilobata, produce a lignied tuber that is
derived from the hypocotyl (Zimmermann 1922b; pers. observ., Fig. 1a). e tuber,
at least of some species, contains starch as a storage nutrient (Getahun 1974b). It de-
velops during the rst season, and lignication may begin as soon as the appearance
of the rst tendrils, such as in C. abyssinica (N. Holstein 132). Some species, such as
C.adoensis and C. grandiora (and most likely also C. senensis and C. schliebenii) do not
produce hypocotyl tubers but root tubers. In C. adoensis var. jereyana the hypocotyl is
Figure 1. a Young plant of C. grandis. e hypocotyl is already thickened and lignies later-on to a
tuber. e cotyledons are glabrous, have an entire margin and an obtuse apex b Adventitious root on
C.grandis; the beige structure to the left is a dry tendril.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
8
minute (N. Holstein 130), which prevents the development of a tuber. Whether West
and Central African forest species produce tubers is unknown.
Each plant produces one to several shoots, which can persist or die back completely
during the dry season or due to re or grazing. Coccinia microphylla shoots can lignify
completely and produce short green branches with owers and small leaves during the dry
season (Fig. 2a), whereas shoots of C. sessilifolia do not lignify at all (pers. observ. from
greenhouse cultivation over 4 years). e shoots of C. grandis can become slightly succu-
lent. e length of the shoots varies from 70 cm in C. microphylla to 20 m in C. grandiora
and C. mildbraedii. Zimmermann (1922a) reports a stem of C. grandiora being 6 cm in
diameter. Usually, the bark of the hypocotyl tuber and the shoots is grayish in color. Fresh
shoots and twigs are usually deep green to brownish green, sometimes speckled with pale
to whitish pustules. In C. abyssinica and C.megarrhiza the shoots and tendrils can turn
purple during maturity. Coccinia sessilifolia produces glaucous shoots that bear a waxy
bloom (Fig. 2b). e indumentum of Coccinia species, if present, is composed of simple,
oligo- to multicellular eglandular trichomes up to 2 mm in length. e long trichomes
consist of oblong cells that may appear articulate when dried (Fig. 3a). Shorter trichomes
can be lineal to conical (Fig. 3b). Sometimes, trichomes have a thickened base that appear
warty when the trichomes break o. e density of the trichomes is often increased on the
nodes. Trichome type and length on shoots are like those of the abaxial surface of the peti-
oles, but usually less dense. Young shoots often exhibit short (< 0.5 mm), weak trichomes,
even in species that are later glabrous, e.g., in C. grandis or C. sessilifolia. Glandular tri-
chomes are rare, few-celled, not visible with the naked eye and have been found, e.g., in C.
grandiora and C. grandis (pers. observ.; anki 1989; Fig. 3c). Glandular trichomes are
also observed in young stems of C. abyssinica (Getahun 1974a), which are usually covered
with long multicellular eglandular trichomes.
Figure 2. a Female ower bud of C. microphylla (N. Holstein et al. 90); picture taken during the dry season.
e stem is completely lignied, and only green short shoots are produced b Male plant of C.sessilifolia. e
stem is glaucous and does not lignify. Unusually, the bract is 3-lobate leaf-like.
Monograph of Coccinia (Cucurbitaceae) 9
Cotyledons
Zimmermann (1922b) reports epigeous cotyledons for C. grandiora and C. grandis, of
which the latter is conrmed by personal observations (Fig. 1a). Epigeous cotyledons
also occur in C. abyssinica, C. adoensis var. jereyana, C. microphylla, C. rehmannii
a. var. littoralis, and C. sessilifolia. e hypocotyl and cotyledons of all observed taxa
are glabrous. e cotyledons are elliptical to obovate and have an entire margin. e
cotyledons are slightly eshy and green, which is also observed in those of C. abyssinica
(Getahun 1974a), and the cotyledonous apex has a pale marking and is obtuse to retuse.
Getahun reports that the prominent veins and the margins on the lower cotyledon
surface in C. abyssinica are covered with multicellular trichomes. However, prominent
veins in C. abyssinica cotyledons cannot be conrmed, and if multicellular trichomes
occur, then they are not visible to the naked eye. e rst normal leaf in this species,
however, emerges in the axilla of the cotyledons (N. Holstein 132, Fig. 4a), and thus
might have been confused.
Leaves
e leaves of Coccinia species are simple, alternate, and paired with a tendril on each
node, except for the rst nodes (Figs 1a, 4a, b). Leaves of all species are petiolate, except
for C. sessilifolia var. sessilifolia, which only develops petioles when young (N. Holstein
131, Fig. 4b) or rarely subsessile leaves when older; full petioles in this species are
only realized in C. sessilifolia var. variifolia (A.Meeuse) Holstein. Subsessile leaves are
common in C. quinqueloba and C. senensis, while the other species’ leaves are usually
distinctly petiolate. e petioles’ surface can be glabrous, at maturity speckled with
hyaline to white cell clusters (C. grandis, C. heterophylla, C. intermedia, C. quinqueloba,
C. rehmannii, C. samburuensis, C. senensis, C. subsessiliora), or have an indumentum.
e petiole contains several vascular bundles arranged in a U-shape (Fig. 5a). However,
Figure 3. Trichomes on Coccinia species. a Node of C. adoensis var. jereyana (J.C. Lovett 1197). e black
bar equals 1 mm. b Part of a leaf C. adoensis var. jereyana (A.F. Stolz 504). e white bar equals 1 mm.
cGlandular trichome from C. grandis.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
10
Hussain et al. (2011) report a ring of vascular bundles in C. grandis. e adaxial side of
the petiole often bears two ridges above the “lateral” vascular bundles (Fig. 5a). ese
ridges merge into the leaf margin and usually bear trichomes (Figs 3b, 5b, c). e
abaxial side of the petiole shares its indumentum with the lower leaf lamina, at least at
the base of the veins (Fig. 5c).
e venation in C. grandis is reticulate, and the mid rib is reported to contain three
bicollateral vascular bundles with xylem and phloem arranged in a ring (Hussain et al.
2011). Reticulate venation can be conrmed for all Coccinia species except C. ogaden-
sis, in which only the central vein in each lobe is visible.
Figure 4. a Young plant of C. abyssinica (N. Holstein 132). e rst node is in the axilla where the
glabrous cotyledons split o. e rst nodes lack probracts and tendrils b Young plant of C. sessilifolia
(N.Holstein 131). e rst leaves in this species are petiolate, sessile leaves are produced later-on. e
glabrous cotyledons are already dried (plant had the same age as the one in Fig. 4a).
Figure 5. a Cross-section of a petiole of C. grandiora, stained with astra blue and safranin (3:2). e
bicollateral vascular bundles are arranged in a U-shape b Trichome on the adaxial ridge of a cross-section
of a petiole of C. grandiora. Although not visible by naked-eye, the petiole is also covered with few-celled
glandular trichomes c Young plant of C. adoensis var. jereyana. e trichomes are mainly occurring on
the prominent veins. e adaxial side of the petiole bears smaller trichomes on the ridges, which fade into
the leaf margin.
Monograph of Coccinia (Cucurbitaceae) 11
Young leaf buds often bear a dense indumentum, even in species that are glabrous
at maturity, e.g., in C. grandis. e leaf lobes are linear, elliptic, (ob-)ovate to triangular.
e incision depth of the lobes can be consistent (C. ogadensis, C. subsessiliora) or
highly variable (C. adoensis, C. grandis, C. senensis (Fig. 6)), even within a single
individual. In taxa with a variable degree of lobation, young leaves tend to be not
or shallowly lobed (e.g., C. grandis, C. megarrhiza, C. rehmannii a. var. littoralis,
C.sessilifolia), a dierentiation according to light exposure might also be possible. e
leaf margin usually is beset conspicuously with small teeth and often bears trichomes
(Figs 3b, 5b, c), even in otherwise glabrous species (e.g., C. grandis, C. sessilifolia). e
teeth are at the apex of lobes, lobules and smaller orders of serration or situated along
the entire margin. e term “dentate” (toothed) is therefore ambiguous in literature
describing Coccinia, as it might also refer to the margin morphology (Stearn 2004).
e teeth are often pale, but can also be colored, esp. when dry, such as in C.abyssinica,
C. grandis, C. intermedia, C. longicarpa, C. megarrhiza, and C. samburuensis (Fig.
7a). e coloration of teeth is inconspicuous in young plants and develops during
maturation (as observed in C.abyssinica, C. grandis, and C. megarrhiza). e teeth are
interpreted as hydathodes by Zimmermann (1922a), because he observed water drops
in C. grandis and C. trilobata on the teeth of the 2nd order (except those of the tip of
Figure 6. Leaf shape variability in Coccinia senensis. a H.J. Schlieben 5259 (HBG) b E.M.C. Groenendijk
et al. 1031 (WAG) c A.R. Torre et al. 18788 (MO). Black bars equal 1 cm.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
12
the lobes) in the morning. A white deposit at the teeth on the upper side of the leaf of
a C. adoensis plant (P. Quarré 75; PR) seems to support Zimmermann’s interpretation.
e upper leaf lamina is often covered with transparent to white pustules, that con-
tain cystoliths (Avetta 1894; Solereder 1899; Zimmermann 1922a; pers. observ.). e
pustules consist of up to 25 cells in C. mackenii (Avetta 1894) but are larger and denser
in glabrous species from dry habitats (esp. C. ogadensis). As they develop over time (they
are smaller and less well visible in forest species), it can be assumed that the pustules
are an adaptation towards protection against high solar radiation. When acetic acid is
applied to microscopic sections of the leaves, heavy gas development suggests that the
cystoliths consist of CaCO3 (pers. observ.). is can be observed in C. grandis, C. hir-
tella, and C.sessilifolia, hence also when the pustules are not conspicuous as in the latter
two species. e pustules may form the base of small trichomes, such as in C.adoen-
sisvar.jereyana (Fig. 3b) or C. microphylla. In some species, the upper surface is usually
covered with an indumentum (C. hirtella, C. schliebenii, C. senensis, and C. trilobata),
but it may also be reduced, and other species rarely exhibit a trichome-bearing upper
surface, e.g., C. adoensis. In each case, the trichomes are simple, < 1 mm, and whitish.
e veins on the upper surface are either glabrous to the naked eye or are covered with
small <0.5 mm long simple trichomes. Zimmermann (1922a) observed in C.grandis
that the glabrous surface of the lamina is only slightly wettable, whereas a drop of eosine
disperses along the veins rapidly. Zimmermann argues that these “capillary drainage
lines” might serve to transport water to the hydathodes during the dry season.
e lower leaf lamina is paler than the upper side (Fig. 3b) and can be glabrous or
bear an indumentum. e highest density of the indumentum can be found on the
Figure 7. a Male ower of C. grandis (N. Holstein 37). Apices of the petals and calyx lobes, as well
as major teeth on the leaf margin are colored in red. Minor margin teeth are inconspicuously colored.
e calyx lobes in C. grandis are spreading in ower buds and reexed in mature owers b Male plant
of C.sessilifolia. Darkish glands (extranuptial nectaries) are commonly found at the base of a lower leaf
lamina. e calyx lobes are unusually large in this specimen (cp. Fig. 2b).
Monograph of Coccinia (Cucurbitaceae) 13
prominent veins (Fig. 5c). e indumentum on the lower leaf surface and the abaxial
surface of the petiole can consist of eglandular oligo- to multicellular trichomes. e
trichomes are appressed or upright (Fig. 5c), usually liform, sometimes also narrowly
conical (e.g., C. abyssinica). Filiform trichomes are straight, curved, or sinuate. Long
liform trichomes often appear articulate when dry due to sunken lateral cell walls
(Fig. 3a). Dry trichomes are hyaline, whitish, beige, or yellowish. e lower lamina
often displays deeply colored to dark green to blackish extranuptial glands (Fig. 7b).
e glands usually occur at the base of the leaf between the veins, sometimes also
between secondary ramications (C. grandis) or along the main veins (C. grandiora).
e epidermis of the lower leaf lamina in C. grandis consists of cells with undulating
anticlinal cell walls and anomocytic stomata (pers. observ.).
Probracts and bracts
In addition to the foliose leaves, most Coccinia species have bracteose prophylls on ster-
ile nodes, which are called “probracts” (Zimmermann 1922b). e probracts can be up
to 5 mm long, but also rather small (< 1 mm) or caducous. e rst nodes of the seed-
ling lack probracts, and they are developed on later nodes. e shape of the probracts
is ovate and entire with a round to acute apex. ey are often spoon-like presenting
the lower surface (e.g., C. adoensis, C. barteri, C. grandis, C. megarrhiza, C. sessilifolia;
Figs3a, 8b), or they are folded in the middle with a prominent keel (C.grandiora;
Fig.8a). Probracts can be glabrous or bear short (< 1 mm) trichomes, and bear ex-
tranuptial glands on the lower surface (Fig. 8a; Okoli and Onofeghara 1984).
Bracts (leaves subtending inorescences or owers), if present, look like the pro-
bracts. Bracts below inorescences are as large as probracts, while bracts below owers
tend to be smaller. Bracts can be present or absent, the latter being an indicative char-
acter for some species.
In rare cases, probracts and bracts can be leaf-like (e.g., N. Holstein 126, Fig. 2b;
P.C.M. Jansen 2065; H. Wanntorp & H.E. Wanntorp 1159) indicating that the (pro-)
bracts are likely derived from usual leaves.
Extranuptial glands
e conspicuous glands on the lower leaf surface, probracts, and bracts (Figs 7b, 8a)
are of the Benincasa-type (sensu Zimmermann 1932), meaning that they are at and
consist of several layers of secretory cells, which are surrounded by a single-layered
sheath (Muhammad Ilyas 1992; Okoli and Onofeghara 1984; pers. observ.). is
sheath is lignied in C.microphylla and C.trilobata (Zimmermann 1922b). However,
Zimmermann (1932) cites Nieuwenhuis von Üxküll-Güldenbandt as saying that the
sheath in C.grandis is suberinized, but the present author did not nd such a statement
in the citations given in that paper (eventually, this was a personal communication).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
14
Schrödter (1926), however, nds that young sheaths in Lua aegyptiaca are lignied
but become suberinized with age, so the dierence might be explained by dierent
stages. Chakravarty (1948) interprets the sheath as lter tissue that is surrounded by
an “external osmotic tissue”. Also Muhammad Ilyas (1992) interprets these radially
elongated cells as secretory and notes that they have a connection to the vascular
strand. However, Okoli and Onofeghara (1984) nd that the glands in C. barteri
are too distant to be interpreted as vascularized. Zimmermann (1922b) observes
intermediate forms between few-celled, stalked glandular hairs and the Benincasa-type
glands in C.microphylla and C. trilobata, including the sheath that forms the base
of the protruding glandular tissue. e glands secrete a clear, rarely slightly colored,
sweet-tasting exudate (pers. observ. in C.grandiora and C.grandis). In C.grandis the
exudates contain sucrose, glucose, fructose, alanine, tryptophane, threonine, and an
unidentied amino acid (Muhammad Ilyas 1992).
Peduncles and pedicels
Male owers mostly occur in racemes that are usually accompanied by 1–2 solitary
owers on the same node (Fig. 2b). e rst owers in male plants of C. hirtella,
C. reh mannii, and C. sessilifolia are solitary. Racemes appear later in the course of
the owering season, although racemes are generally rare in the rst species (pers.
observ.). If solitary owers and racemes are produced on the same node, then the
solitary ower(s) precede(s) those of the racemes in time of maturity (Fig. 2b). e
trigger to produce racemes instead of or additionally to solitary owers is not known.
e racemes bear up to 35 owers (e.g., in C.pwaniensis, C.racemiora). Within the
racemes, owering starts at the basalmost branches. If the peduncle is reduced, owers
Figure 8. a e probract of C. grandiora is eshy and has a keel. e abaxial side bears many extranuptial
nectaries. e structure pointing to the lower left of the picture is the tendril b e probract of C.trilobata
(sampled as N. Holstein & P. Sebastian 9) is spoon-shaped, papery, and without a keel.
Monograph of Coccinia (Cucurbitaceae) 15
appear clustered on the node. In C.grandis, which usually produces single owers only,
ower clusters (short-peduncled racemes) rarely occur. is can be seen in plants from
Ethiopia, Saudi Arabia but also from India and Sri Lanka. e pedicels of solitary male
owers of C.hirtella, C.megarrhiza, and C.rehmannii exhibit a negative gravitropism.
In creeping plants, pedicels that grow downwards in the beginning make a sharp bent
upwards to present the ower upright.
Female owers are mostly solitary. Only in some species, female owers are usually
in racemes, such as in C.heterophylla, C.keayana, and C.racemiora. Few-owered
female racemes or clustered owers might also occur in C.grandiora, C.intermedia,
and C.subsessiliora. In C.barteri, female owers can be solitary or in few- or many-
owered racemes. Two female owers per node have also been observed in C.micro-
phylla. e pedicels of solitary female owers are negatively gravitropic during ower
development. After pollination, the pedicels of solitary female owers of C.grandis,
C.hirtella, C.megarrhiza, C.microphylla, C.rehmannii, and C.sessilifolia exhibit posi-
tive gravitropism. e downturn is not due to slackness caused by the weight of the
developing fruit but an active process, as the pedicels thicken and remain rm. How-
ever, only fertilized owers turn downwards completely, as aborted owers from mis-
pollination never reach this state (pers. observ. in cultivated plants).
Perianth
e perianth of all Coccinia species is synsepalous and sympetalous. At the base, calyx
tube and corolla tube are connected with each other and form a perianth tube or fun-
nel. Depending on the exsertion point of the staminodes in female owers, parts of the
tube form a hypanthium (e.g., C.grandiora).
e calyx dierentiates as a bulge (Fig. 7a, b) with usually ve lobes, or only the
lobes emerge from the perianth tube (Fig. 9). If the calyx emerges as a bulge, then it and
the perianth tube are rather conspicuously dierentiated from the corolla in terms of
color. If only the calyx lobes emerge, then the color of perianth tube fades to green color
towards the receptacle, with the veins of the corolla remaining more intensely colored.
Whether calyx and corolla are non-dierentiated (congenital fusion) or postgenital-
ly fused, is not known for Coccinia, but in the distantly related Echinopepon wrightii
(A.Gray) S.Watson the perianth tube is non-dierentiated (Leins and Galle 1971). e
outside of the perianth tube can bear long trichomes of the type on as the lower leaf
surface or the petioles (Fig. 10a). e calyx lobes are acute triangular to subulate or
linear, rarely slightly lanceolate. e orientation of the calyx lobes is erect, spreading, or
reexed, although they can be curved inwards (e.g., C.rehmannii a. var. littoralis (Fig.
10a) or outwards (e.g., C.intermedia). e color of the calyx lobes can be more intense
(green) than the perianth tube or the pedicel (Fig. 7b). In C.grandis, the tip of the calyx
lobes is brownish to reddish just as the teeth on the leaves and the corolla (Fig. 7a).
e petals of Coccinia species are fused at the base, usually for at least one third of
the total length. Rarely, the petals are free down to the height of the calyx lobes (pers.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
16
observ. in C.megarrhiza, C.rehmannii var. rehmannii, and C.sessilifolia). Perianth
tube and corolla tube are often campanulate, rarely funnel-shaped or tubular. e peri-
anth tube can be urceolate in C.longicarpa, C.racemiora and sometimes in C.barteri.
e tips of the (4–)5(–7) corolla lobes are rounded to acute with an apical tooth. e
Figure 9. Leaky dioecy in a plant of C. megarrhiza. e plant was owering male (the bud on the left)
through the season, but a single female ower developed (the second ower on this node was male). e
ower was receptive and produced a normal-sized fruit and normal-shaped seeds. e probracts (left
node) are spoon-shaped, the tendrils are purplish.
Figure 10. a Male plant of C. rehmannii [a. var. littoralis] (N. Holstein 126). e plant was raised
from seeds of a female plant with ovoid fruits (B. Jarret – pers. comm.) b Male ower of C. adoensis var.
aurantiaca (N. Holstein et al. 85). e halictid bee (H. Schaefer – pers. comm.) circled around the globose
anther head harvesting pollen. e scent of the ower was strong and honey melon-like.
Monograph of Coccinia (Cucurbitaceae) 17
apical tooth can be inconspicuous or colored claret red or brown, such as in C.adoensis
var. aurantica or C.grandis. Outside, the perianth tube and the corolla is glabrous or
covered with short (< 10 globose cells in C.grandis) trichomes. Inside, the corolla is
covered with long trichomes (up to 20 cells in C.grandis), sometimes with a glandular
apical cell (Fig. 10b). e trichomes become shorter towards the receptacle. e in-
ner side of the hypanthium of female owers is glabrous and smooth, which suggests
nectary tissue, in C.grandiora, C.grandis, and C.hirtella (Fig. 11a). e size of the
corolla does not dier conspicuously between staminate and pistillate owers; pistillate
owers might be a bit smaller.
Androecium
In staminate owers, the three stamens originate from the base of the perianth tube,
and the laments are fused to a central column (Fig. 10a). e fusion point sometimes
leaves a small gap to the hollow receptacle center. is gap, however, can be lled with
long multi-cellular trichomes (e.g., in C.abyssinica and C.megarrhiza). Rarely, the la-
ments can also be separate. e laments are glabrous and white, greenish, yellowish,
or orange. e number of vascular bundles in the stamens is disputed. Chakravarty
(1954) reports ve vascular bundles in C.grandis: two stamens have two bundles each,
and the third stamen has a single bundle. Later research shows three vascular bundles
for the same species with one per stamen (Bhuskute et al. 1986; Deshpande et al.
1986). In C.hirtella each stamen contains a single vascular bundle (pers. observ.).
e anthers together form a globose head (Fig. 10b). Each anther is bithecate;
sometimes one can be monothecate (Bhuskute et al. 1986; Chakravarty 1954). Each
theca is sinuate. Deshpande et al. (1986) report a bi-layered brous endothecium
and a secretory tapetum, which they found diers from distantly related Momordica
charantia L.
Figure 11. a Longitudinal section of a female C. hirtella ower (size of the small squares is 1 mm²). Peri-
anth and style were detached. e pollen sacs on the staminodes are highly reduced. e introrse side of the
staminodes bears long trichomes that in intact owers touched the style b Female ower of C. rehmannii var.
rehmannii with bilobate stigma arms c Female ower of C. megarrhiza with bulging stigma arms.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
18
In pistillate owers, the three, now free, stamens are reduced to staminodes that
originate from the interior perianth wall, forming a hypanthium. Introrsely, the stami-
nodes of C.grandiora, C.grandis and C.hirtella bear long, multicellular trichomes,
except for the apex, extrorsely the staminodes are glabrous (Fig. 11a). e anthers of
the staminodes are strongly reduced to a slightly yellowish spot at the apex in C.gran-
dis and C.hirtella. e staminodes of C.megarrhiza bear long multicellular trichomes
introrsely and laterally but are glabrous extrorsely.
Pollen
Pollen in Coccinia species shows little diversity. e pollen is oblate-spheroidal to pro-
late with a reticulate exine (Table 1). Additionally, the pollen of C.pwaniensis (Holstein
and Renner 2010), C.hirtella, and C.trilobata is prolate, the exine texture is unknown.
e sampling of the examined species covers all clades and suggests uniformity in shape
and exine texture, which negates systematic value of pollen in Coccinia. e color is yel-
low in C.abyssinica, C.grandiora (Zimmermann 1922b; and pers. observ.), C.grandis,
C.hirtella, C.megarrhiza, C.microphylla, C.rehmannii, C.sessilifolia, and C.trilobata,
and orange in C.adoensis var. aurantiaca. Zimmermann (1922b) reports that pollenkitt
of C.grandiora contains a yellow colorant that is soluble in peanut oil, but not in water
and only slightly in heated chloral hydrate solution. It changes its color in concentrated
sulfuric acid to blue, in Lugol’s iodine (I2KI) to green, and in osmic acid to brown. As
in several other cucurbit species, in vitro germination of C.grandis pollen increases from
pH = 7 towards alkalinity and is maximal at pH = 8.5 (Zaman 2009).
Table 1. Pollen characters in Coccinia species.
shape Size (P × E) [µm] Exine texture source
C. abyssinica Prolate-spheroidal 60–70 × 56–65 Reticulate Marticorena 1963
Oblate-spheroidal to
spheroidal 76 × 81 Reticulate Khunwasi 1998
C. adoensis Prolate 66–73 × 45–50 Reticulate Marticorena 1963
Prolate-spheroidal 72 × 61 Reticulate Khunwasi 1998
C. barteri Prolate 70–80 × 50–60 Reticulate Marticorena 1963
Prolate-spheroidal 71 × 58 Reticulate Khunwasi 1998
C. grandiora Prolate Zimmermann 1922b
C. grandis Prolate 60–63 × 34–40 Reticulate Marticorena 1963
Prolate-spheroidal 58 × 52 Reticulate Khunwasi 1998
Prolate 47.61–64.62 × 35.91–44.80 Coarsely reticulate Perveen and
Qaiser2008
Subprolate to prolate 34–52 × 28–35 Reticulate Awasthi 1962
Prolate 41.20 ± 0.61 × 34.00 ± 0.45 Reticulate Datta 1988
C. megarrhiza Oblate-spheroidal to
spheroidal 92 × 92 Reticulate Khunwasi 1998
C. mildbraedii Prolate 55–60 × 35–41 Reticulate Marticorena 1963
C. sessilifolia
var.sessilifolia
Prolate-spheroidal to
prolate 70 × 58 Reticulate Khunwasi 1998
Monograph of Coccinia (Cucurbitaceae) 19
Gynoecium
Pistillate owers are epigynous and have three (rarely two or four) carpels. e ovary
is narrowly spindle-shaped, oblong to globose. e surface is smooth or warty; it is
glabrous or has the indumentum of the pedicel. e style is often greenish-white or
pale-yellowish; the stigmas are frequently in yellowish colors and covered with long
trichomes (Fig.11c). Each stigma in C.grandiora, C.grandis, C.hirtella, C.megar-
rhiza, C.rehmannii, and C.sessilifolia is U-shaped with the ends of the lower sides of
the arms touching each other. e stigmatic branches can be long and free, such as in
C.grandiora, C.grandis and C.rehmannii var. rehmannii (Fig. 11b), or short and bul-
bous, such as in C.hirtella, C.megarrhiza (Fig. 11c), C.microphylla, and C.sessilifolia.
e placentation of the ovules in Coccinia is involute, which is also discussed for
other Cucurbitaceae by Leins and Galle (1971). e funicle appears to be attached to
the outer wall, but actually attaches to a septum coming from the axis (Fig. 11a, b, c),
which itself is connected to the outer wall. e axis-wall septum, however, might be
reduced during ripening, but this needs further study.
e anatomy of the ovules is barely surveyed in Coccinia. In C. abyssinica and
C.grandis, the ovules are reported to be anatropous, bitegmic, and crassinucellate (Ge-
tahun 1973; Zahur 1962). Anatropus ovules also occur in C.hirtella (Fig. 12a). e
position of the ovules is horizontal in C.grandis (Kirkwood 1904), C.hirtella (Fig. 12a),
and C.megarrhiza (Fig. 12b) with the micropyle facing outwards. Horizontal ovules are
regularly reported in the Benincaseae and the Cucurbiteae (Schaefer and Renner 2011a).
In staminate owers, a pistil is not developed because the stamens fuse to a central
column. e pedicel is narrow and reaches the perianth, and there is no indication of
even a thin (sterile) inferior ovary in the ower.
Female gametophyte development and embryology
e development of the embryo has only been investigated in C.grandis. Chakravorti
(1947) and Zahur (1962) report that the female gametophyte development is accord-
ing to the Polygonum-type. Both observe that the synergides possess hooks instead of
the liform apparatus. Chakravorti (1947) describes the developing endosperm as a
nuclear type, which is conrmed by Chopra (1955). By formation of a large central
vacuole, the nuclei become displaced to the periphery. After the endosperm becomes
cellular, the often lateral chalazal haustorium remains coenocytic.en, the hausto-
rium becomes cellular with multinuclear cells except for the apex (Chopra 1955).
Fruits
e fruits are many-seeded berries, which vary in size and shape between species (and
within C. grandis, C. rehmannii, and C. subsessiliora). e smallest fruits occur in
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
20
C.rehmannii var. rehmannii and C.microphylla with globose berries as small as 1 cm in
diameter at maturity. However, in both species larger globose fruits (up to 2.5 cm in diam.)
and in C.rehmannii ovoid fruits may occur additionally, in the latter case especially in
more humid habitats. e largest fruits occur in C.samburuensis and the rainforest species
C.grandiora, C.longicarpa, C.mildbraedii, and C.schliebenii, which have long elliptical
to cylindrical (sausage-shaped) fruits up to 20 cm long and up to 5 cm in diameter.
Immature fruits often have a white or pale-green (C.hirtella, C.sessilifolia) or dark
green (C.adoensis) longitudinal mottling or lines, even when the ovary and the ripe
fruit is single-colored (Figs 2a, 13a, b, c). In the C.rehmannii clade, the white spots
or lines become surrounded by a dark green halo during ripening (Fig. 13c). Rarely,
if no white mottling develops, e.g., in some C.microphylla, dark green spots develop
nevertheless. In any case, the mature fruit in species of the C.rehmannii clade is usually
uniformly colored red (Zimmermann 1922b; and pers. observ.). Ripening usually oc-
Figure 12. a Cross-section through an ovary of C. hirtella. e ovules are anatropous with the micropyle
facing outwards b Cross- and longitudinal section of a C. megarrhiza fruit. e seeds are enclosed in a
hyaline hull (aril) and seemingly attached to the periphery c Cross-section through a fruit of C. sessilifolia.
Note that the vascular bundles in the lower left of the picture bend in the periphery, so the placentation
is not parietal but involute.
Figure 13. a Ripening fruit of C. hirtella. Note the typical lobulate leaves of this species in the lower
right b Ripening fruit of C. sessilifolia. e fruit, like the plant, bears a waxy bloom c Ripening fruits of
C. megarrhiza have a dark green halo around the white longitudinal mottling. e left fruit is derived
from pollination with C. megarrhiza pollen, whereas the smaller fruit on the right is derived from cross-
pollination with C. trilobata (both pollinations were conducted on the same day).
Monograph of Coccinia (Cucurbitaceae) 21
curs from green with or without mottling via yellow to orange to the nal coloration.
e color changes from the apex of the fruit downwards (Fig. 13a, b, c), independent
of the position (hanging vs. horizontal) in C.sessilifolia. In C.megarrhiza, pendulous
fruits ripen from the apex to the base, which sometimes remains green even when the
apex already turns soft. In lying fruits from creeping C.megarrhiza plants, ripening
does not proceed from the apex, but starts from point that is closest to the source of
either warmth or light (pers. observ. from greenhouse cultivation). e degree of the
yellow to orange ripening zone varies. In C.sessilifolia, fruits directly turn red, whereas
in C.grandis the color change includes a well visible yellow zone. Unripe fruits col-
lected of C.grandis tend to turn yellow outside and pink to red inside (Imbumi 2004).
Mature fruits are in deep red colors (hence the genus name) or orange-red. Rarely, a
white longitudinal mottling is described in ripe fruits (e.g., C.mackenii).
Immature fruits are glabrous or have the same indumentum as the ovary. By ripen-
ing, the indumentum is usually reduced. e exocarp of Coccinia fruits is papery thin
and has a waxy bloom when ripe. e endo- and mesocarp are red, eshy and soft
(Fig.12b, c). e pulp is nerved with a dense network of tubular tissue. Shah et al.
(1983) report that such a network consists of sieve tubes in C.grandis. e sieve tubes
are not connected to the main vascular strands and are lled with a proteinaceous ma-
terial. e authors suggest that the sieve tube network aids nutrient transport during
the rapid growth of the fruit.
Seeds
e seeds (Fig. 14) in Coccinia species are beige to grayish with a small margin, which
often has a darker coloration. e shape is more (esp. in the C.rehmannii clade) or less
asymmetrically ovate (especially C.adoensis, C.pwaniensis, and C.senensis). e sur-
face is at, esp. in the C.rehmannii clade, to lenticular (esp. C.adoensis, C.pwaniensis,
and C.senensis). e size varies from 4.5–7 × 3–5 × 1–2 mm (L/W/H). Seed numbers
per fruit vary drastically from about 10 (C.microphylla) to c.100 (+ c.20 infertile) in
C.sessilifolia (N. Holstein 119). Species with larger fruits might contain more seeds.
Detailed observations of the seed anatomy have been made by Getahun (1973) for
C.abyssinica and by Chakravorti (1947) for C.grandis. Getahun describes the mature seed
as consisting of the embryo, a membrane-like structure (pellicle) closely adhering the em-
bryo but separated from the hard testa. However, Chakravorti does not recognize a pellicle
in C.grandis. Both authors agree that the inner integument disappears and the testa devel-
ops solely from the outer integument. e testa of C.abyssinica is described as comprising
four layers (from center outwards): (1) a thin-walled parenchyma, (2) a sclerenchyma of
macrosclereids, (3) a thick-walled parenchyma, and (4) an epidermal layer. e outer-
most layer, the epidermis, is disintegrated, leaving the cell walls as slender rods of 500 µm
length. is has also been noticed in other species, and the surface has been described as
a brillose testa (Jerey 1967; Kéraudren 1967). De Wilde et al. (2011) also interpret the
seed surface of C.grandis as having a disintegrated, pulpy, radiately striate exotesta.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
22
Getahun contrasts his observations with those of Chakravorti (1947) in C.grandis,
but the seeds are in fact similar, just incompletely described by Chakravorti. Chakra-
vorti draws a four-layered testa, but does not name the innermost layer that has the
same hatching as the third layer, which he describes as “cells with thickened walls”.
is is what Getahun calls parenchyma. Chakravorti’s outermost layer, the epidermis,
consists of radially elongated cells with thin walls. ese cells have likely just not yet
disintegrated as observed by Getahun. e only dierence between both observations
is the second layer, which consists of macrosclereids in C.abyssinica and of radially
slightly elongated cells with thin walls in C.grandis. ese dierent observations are
explainable by two possibilities: 1) dierent developmental stages of the seeds, since
Chakravorti surveys the seed development, so the layers are immature, while Getahun
surveys mature seeds and germination, or 2) dierent staining. Chakravorti uses hae-
matoxylin alone, which does not stain lignied cell walls, whereas Getahun uses hae-
matoxylin with safranin as counter stain, which makes lignin, and thus sclerenchyma,
well visible (von Aufseß 1973).
e seeds in Coccinia, at least in C.abyssinica (Hora 1995), C.grandis, C.hirtella,
C.megarrhiza, C.microphylla, C. mildbraedii, C.sessilifolia, and C. subsessiliora (pers.
Figure 14. Seeds of Coccinia. e lack of bers in e and f are preparation artifacts due to mechanical
removal of the hyaline aril. Maceration (coarse crushing of the fruit and soaking of the mass in water for
2–3 weeks; R. Brüggemann – pers. comm.) retains the surface bers. Length of white bars equals 1 mm.
aSeeds of C. adoensis var. jereyana (plant derived from seed of the same fruit: N. Holstein 130). Note the
lenticular face and symmetrical shape of the seed b seeds of C. abyssinica (plant derived from seed of the
same fruit: N.Holstein 120 and 132) c Seeds of C. trilobata d Seeds of C. sessilifolia (harvested by macera-
tion) e Seeds of C. sessilifolia (harvested by mechanical extraction; taken from N. Holstein 119) f Seeds of
C. grandis (harvested by mechanical extraction).
Monograph of Coccinia (Cucurbitaceae) 23
observ.) are surrounded by a hyaline red juicy envelope (Fig. 12b). As the ovule is
bitegmic (see above), one might assume the hyaline envelope is the testa. Chakravorti
(1947), however, observes that the juicy envelope is derived from carpellary tissue.
However, Getahun (1973) does not recognize the hyaline hull in C.abyssinica, which
is surprising as it also occurs in the closely related species C.megarrhiza and C.micro-
phylla. Similar structures to the hyaline hull are also found in other Cucurbitaceae, esp.
in Momordica. Van der Pijl (1982) interprets these as “endocarp-pulpa” taking over
the function of an aril for seed dispersal as the fruits of Momordica species dehisce at
maturity. However, Coccinia fruits disintegrate and do not dehisce, e.g., into valves.
Germination
e seeds of C.abyssinica maintain a high germination rate (100%) after four years of
storage at room temperature (Getahun 1973). However, time from watering until ger-
minating increases from 4 days (after one year of storage) to 16 days (after four years of
storage). Seeds of C.grandis are also able to germinate after four years of storage, while
seeds of C.ogadensis ulin (3 seeds tested) did not germinate after ve years (pers. ob-
serv.). Getahun (1973) reports that C.abyssinica seeds do not germinate below 10 °C and
above 35 °C.In the latter case, he observes thermal damage to hypocotyls and primary
roots. e optimum for germination in C.abyssinica is between 20 and 30 °C and that of
C.grandis is 35 °C, whereas temperatures <23.5 °C and >40 °C inhibit germination (Li
et al. 2001). e germination rate of C.abyssinica seeds in the light is decreased by 35%
compared to germination in darkness (Getahun 1973). In C.sessilifolia, seed viability de-
clines after 9 months, and germination is at a maximum after 10–20 min smoke exposure
or red:far red light treatment, followed by burying and a long-day cycle (Weiersbye and
Witkowski 2003). Rotting of a crushed ripe fruit in water (for seed extraction) resulted in
germination of two seeds in an articial hybrid (C.megarrhiza ♀× C.rehmannii a. var.
littoralis ♂) after 3 weeks of soaking (pers. observ.). Coccinia grandis seeds do not exhibit
dormancy (Motooka et al. 2003); for the other species there is no information available.
Genome, chromosomes, and hybridization
Chromosomes and sex determination
Coccinia is one of the few examples in the plant kingdom, in which at least one species
has heteromorphic sex chromosomes (Ming et al. 2011). Coccinia grandis contains 22
autosomes plus 2 gonosomes. Female individuals have homomorphic XX, whereas
male individuals have heteromorphic XY chromosomes (Fig. 15a). Although Kumar
and Deodikar (1940) report males to have two large “X” and females a large X and a
smaller Y chromosome, later studies (Chakravorti 1948; Kumar and Vishveshwaraiah
1952) reveal that males are heteromorphic and the Y is 2.5 (Bhaduri and Bose 1947)
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
24
to 3–4 times longer (Guha et al. 2004) than the other chromosomes. Some years be-
fore Kumar and Deodikar, Sutaria (1936) reported n = 12 from pollen mother cells of
C.grandis, without nding the large Y chromosome. Although scientists from India
conducted some research for C.grandis, chromosome work in other Coccinia species is
almost none-existent. McKay (1930) reports n = 12 for C.hirtella, without mention-
ing whether he studied a male or a female individual. e author’s own chromosome
counts (Table 2; Fig. 15b) support McKay’s report and reject the existence of hetero-
morphic sex chromosome in C.hirtella males. is is also the case for C.sessilifolia
(Fig.15c). Two counts in the C.rehmannii clade reveal a reduction of chromosome
number and the non-existence of a heteromorphic Y chromosome there.
Due to the sex chromosomes, sex expression in C.grandis plants is pre-determined
and sex ratios in the ospring basically follow Mendelian inheritance of a single allele.
However, Agarwal and Roy (1983) report that of 500 planted seeds only 181 (36.2%)
were male, and their interpretation is that there might be a genetic mechanism to
reduce the number of male plants. As they do not report XY females, their nding
might be explained rather by an increased lethality of XY embryos due to deleterious
mutations on the single X or on the Y in Y-containing pollen. e Y chromosome
in C.grandis is dominant, as the presence of a single Y results in male phenotypes,
regardless of the number of X chromosomes (Agarwal and Roy 1975; Kumar and
Figure 15. Chromosome preparations xed in 3:1 EtOH – acetic acid and stained with orcein, objective
lens: 100×. a Mitotic plate of a male C. grandis (2n = 22 + XY) b Mitotic plate of a male plant of C. hirtella
(2n = 24) c Phase contrast image of a mitotic plate of a male plant of C. sessilifolia (2n = 24).
Table 2. Chromosome numbers of Coccinia species and sexes of surveyed individuals.
Species Sex of individual Chromosomes (2n)Voucher
C. grandiora female 24 N. Holstein 114 (EA, M)
C. grandis male 22 + XY (Fig. 15a) N. Holstein 32 (M)
C. grandis female 22 + XX N. Holstein 25 (M)
C. hirtella male 24 (Fig. 15b) N. Holstein 29 (M)
C. rehmannii a.
var.littoralis male 20 N. Holstein 126 (M)
C. sessilifolia male 24 (Fig. 15c) N. Holstein 13 (M), N. Holstein 109 (M (3))
C. sessilifolia female 24 N. Holstein 119 (B, M)
C. trilobata male 20 (A. Sousa, pers. comm.) N. Holstein & P. Sebastian 9 (M)
Monograph of Coccinia (Cucurbitaceae) 25
Vishveshwaraiah 1952; Roy and Roy 1971b). Triploids of C.grandis with a 3n = XYY
constitution are also male but bear owers in clusters instead of the usually solitary
ones and exhibit leaf deformations (Roy and Roy 1971b).
Evidence for Mendelian inheritance of sex in C.hirtella is not so clear, as the same
plant can produce owers of the opposite sex in succeeding seasons. Two plants marked as
female and one as male from observation of owers produced owers of the opposite sex
in the following year (pers. observ.), making C.hirtella functionally dioecious, but geneti-
cally hermaphroditic. On the other hand, there are several observations of owers of the
opposite sex in otherwise unisexual plants. Kumar and Vishveshwaraiah (1952) report a
gynodioecious form of C.grandis that has homomorphic chromosomes (XX). Although
bisexual owers are reported to develop, pollen grain development is arrested, and the male
function remains suppressed. Roy and Saran (1990), however, report fully fertile hermaph-
roditic owers in an otherwise female individual. Holstein and Renner (2011a) report a
collection of C.intermedia that bears male oral buds and female owers and young fruits
on the same nodes. is observation can be interpreted as ‘leaky dioecy’ (Baker and Cox
1984). Among the author’s own cultivated plants, a male individual of C.megarrhiza pro-
duced a single female ower towards the end of the season (Fig. 9). Although two male
owers on the same individual were open at the same time when the female ower was ma-
ture, the pollen sacs did not open. It is not known whether this was a coincidence or func-
tionally signicant, e.g., to prevent selng. Selng is often discussed as being advantageous
in small population sizes, e.g., when new islands are colonized. Prevention of simultaneous
owering of both sexes on the same plant implies that leaky dioecy would not immediately
aid the establishment of new distant populations per se. It might require the establishment
of several plants or clonal separation. In any case, the single female ower was receptive
and was fertilized by another male C.megarrhiza plant derived from the same fruit as the
“female” plant. e resulting fruit and seeds developed normally.
e production of hermaphroditic owers in X-radiation studies (Agarwal and Roy
1983; Roy 1974) shows that dicliny in Coccinia is kept up actively. Agarwal and Roy
found hermaphroditic owers on two plants with otherwise female owers and an XX
conguration. ey also report the development of a normal fruit without mentioning
the fertility of pollen from hermaphroditic owers, but interpret their nding as cleis-
togamy. However, fruit development without previous pollination (parthenogenesis)
or from pseudogamy with pollen from dierent genera as fructication stimulus is
described by Lal (1973). True selng from own pollen (in irradiated XX individuals)
would mean that the Y chromosome is not important for fertile pollen development.
Furthermore, this means that it only carries at least one gene for suppression of the
development of the female organs, and the occurrence of a second X suppresses pollen
development in “normal” plants. Agarwal and Roy (1983) also report that X-ray dos-
ages of 5 to 50 R [1.29 × 10-3 to 12.9 × 10-3 C/kg] result in a drastic reduction of the
sex ratio (11 males, 2 hermaphrodites, and 127 females out of 500 irradiated seeds).
is might indicate that the single X bears many functionally important genes in con-
trast to the Y, as mutations in the single X lead to an increased mortality compared to
females with a balancing second X chromosome.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
26
Genome of C. grandis
Aside from research on the sex chromosomes, a few studies on the genome of C.grandis
have been undertaken. Guha et al. (2004) report the 4C nuclear DNA content of female
C.grandis as 8.37 ± 0.14 pg, whereas that of male C.grandis is 10.17 ± 0.24 pg. is means
that the dierence between X and Y chromosome adds about 20% to the complete DNA
content. Patankar et al. (1985) report a DNA content of 1C = 2.75 pg for C.grandis, how-
ever, they do not report the sex of the analyzed individual. Interphase nuclear structure in
C.grandis is chromocentric with 14 ± 0.25 chromocenters (Patankar et al. 1985).
Surveys on the reassociation kinetics in Cucurbitaceae suggest that C.grandis has
the lowest amount of repetitive DNA among the six species studied in the Cucurbita-
ceae (Bhave et al. 1984). Fragments of 550 bp length have 25% of repetitive elements
(Bhave et al. 1984), whereas 7400 bp long fragments consist of 49% repetitive DNA
(Bhave et al. 1986). However, Pasha and Sen (1995) report dierent results as they
nd 400–600 bp long fragments to comprise 38% highly repetitive DNA (52% total
repetitive DNA), which appears to be average in the Cucurbitaceae. Pasha and Sen do
not discuss this dierence, which cannot be explained by dierence in sex, as the large
amounts of plant material used (1 kg seeds or 100 seeds respectively) suggest that it
must have comprised elements of both sexes. A sex discriminating study of C.grandis
reassociation kinetics has not been undertaken yet.
Hybridization and crossing experiments
Charles Naudin’s famous work on the eects of hybridization included crosses between
Coccinia plants. He reports successful crosses between Asian C.indica (nom. illeg. for
C.grandis) and the NE African C.schimperi (P06809214, P06809215, P06809216;
Naudin 1862), which is now seen as a synonym of C.grandis. Naudin’s Coccinia schim-
peri, however, has bu petals, whereas the Asian C.grandis has snow-white petals. Both
supposed species hybridized without problems. During the following two years, Nau-
din could not intercross within the F1 generation because plants of dierent sexes did
not ower at the same time, so he crossed F1 individuals with a female C.grandis from
Asia, which again produced ospring. As Naudin erroneously supposed that he dealt
with two species, he deduced that hybrids between species could be fully fertile, have
a reduced fertility or be sterile, and that there was no clear boundary between species
and varieties. However, he proved rather that the bu-petaled, African C.schimperi
and the white-petaled Asian C.grandis are a single species obeying the biological spe-
cies concept.
Naudin also crossed other Coccinia species that he had in cultivation. Coccinia
quinqueloba and C.mackenii, although sometimes not easily distinguishable, were not
amenable to crossing (Naudin 1866). Only 1 out of 20 crossing trials resulted in a fruit
that developed poorly. Naudin did not publish whether the hybrid seeds were fertile
or even viable, but his observations are valuable as each one accession of C.quinque-
Monograph of Coccinia (Cucurbitaceae) 27
loba and C.mackenii were not distinguishable using more than 3500 bp of plastid
sequences, and hence might share the same plastid haplotype (Holstein and Renner
2011b). ere are collections that share characters of C.mackenii and C.quinqueloba,
but these are not intermediates. In these collections, long petioles (typical for C.mack-
enii) are coupled with simple tendrils (typical for C.quinqueloba), and thus cannot be
unambiguously allocated to either species. However, if both typical forms are indeed
reproductively isolated, then they are species sensu Mayr (1942), and the crossing
behavior of these species needs to be tested reciprocally to dene the morphological
scope of the two species.
Naudin also crossed male C.diversifolia (C.abyssinica) with a female C.mackenii,
which are rather distantly related and do not co-occur in nature. However, the cross
resulted in onset of mediocre fruits with only few, but well-developed and viable seeds
(Naudin 1866). Naudin did not report further results for this cross either.
As reproductive isolation between species is often assumed but rarely tested, cross-
ing experiments among species that are cultivated in Munich Botanical Garden have
been performed. Positive results are given in Table 3.
Interspecic fertilization succeeded or failed without correlation of relatedness or
co-occurrence (Table 4). Female owers of C.sessilifolia could not be fertilized with
pollen of C.megarrhiza (4 trials), C.trilobata (4 trials), C.rehmannii (2 trials), C.hirtella
(3 trials) or Diplocyclos palmatus (2 trials). Pollinated female owers were discarded like
non-fertilized owers. Hence, hybridization seems to be prevented prezygotically in
female C.sessilifolia with members of the C.rehmannii clade and C.hirtella as pollen
donor. As C.sessilifolia and C. rehmannii co-occur widely in their range and share
owering time and oral syndrome, the production of hybrids would reduce tness
drastically. Coccinia sessilifolia and C.hirtella do not co-occur, but belong to the same
clade (see chapter Evolution and phylogeny). Although a female C.sessilifolia could
not be fertilized by pollen from C.hirtella, pollination of a female C. hirtella with
pollen from C.sessilifolia resulted in fruit onset.
In contrast to C.sessilifolia, C.grandis is fertilized easily by C.hirtella and C.sessili-
folia, although the species neither co-occur, nor are closely related. e cross resulted
in ospring, which was growing vigorously but sterile, as pollen sacs did not open
(Table 3). Hybrid pollen that was extracted from the pollen sacs of fully open owers
was also not able to fertilize C.sessilifolia (1 trial). e occurrence of sex chromosomes
in C.grandis might result in gene dosage imbalance, which interferes with the oral
development, leading to sterile ospring. e inability of female C.grandis to be ferti-
lized by C.rehmannii a. var. littoralis and C.trilobata (Table 4) might be explained by
the fact that the chromosome numbers dier (see Table 2) and translocations lead to
gene loss in hybrid genomes and thus inviability of the ospring. On the other hand,
the cross between female C.hirtella and male C.trilobata (not sympatric) produced a
purely intermediate F1 generation, which owers vigorously despite the dierence in
chromosome numbers (see Table 2 and 3). Although the anthers open like in fertile
owers, unlike in C.grandis hybrids, the pollen of this hybrid was not able to fertilize
female owers of C.hirtella (1 trial), C.grandis (2 trials), or C.sessilifolia (2 trials).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
28
Table 3. Description of the F1 from crosses and a natural hybrid between Coccinia species. Species used
for the articial crosses are not sympatric.
Parent species Ospring Occurrence
C. grandis ♀ ×
C.hirtella ♂
F1 vegetatively morphologically intermediate; owers
are either aborted or sterile (pollen sacs remain closed);
pollen globose; corolla is smaller than in each parent
species (Fig. 16a)
Articially in Munich Botanical Garden;
voucher: N. Holstein 108 (M)
C. hirtella ♀ ×
C.grandis ♂
F1 vegetatively morphologically intermediate; owers
smaller (Fig. 16b), sterile
Articially in Munich Botanical Garden;
voucher: N. Holstein 116 (M)
C. grandis ♀ ×
C.pwaniensis ♂F1 morphologically intermediate; owers sterile
(pollen sacs remain closed) (Fig. 16c)
Naturally in Pugu Hills, Dar es Salaam,
Tanzania; vouchers: N. Holstein et al.
102 (DSM, M), 103 (M), 104 (M), 105
(DSM, M)
C. grandis ♀ ×
C.sessilifolia ♂
F1 vegetatively morphologically intermediate; owers
are either aborted or sterile (pollen sacs remain closed);
corolla is smaller than in each parent species (Fig. 16d)
Articially in Munich Botanical Garden;
voucher: N. Holstein 113 (B, M);
N.Holstein 115 (M)
C. hirtella ♀ ×
C.trilobata ♂
F1 morphologically intermediate; males owering
vigorously with intermediate owers, pollen sacs open,
but pollen is sterile
Articially in Munich Botanical Garden;
N. Holstein 121 (M)
Table 4. Observations on fruit development of crosses between Coccinia species (except for crosses men-
tioned in Table 3). Viability and morphology of the F1 is not known so far. a = sympatrically distributed
species, b = close relatives/sister species, c = species occur in the same area but in dierent habitats.
Parent species crossability
C. grandis ♀ × C. abyssinica ♂Onset of fruit (1 trial) c
C. grandis ♀ × C. megarrhiza ♂Onset of fruit (1 trial) a
C. grandis ♀ × C. rehmannii a. var. littoralis ♂Abortion of ower (1 trial)
C. grandis ♀ × C. trilobata ♂Abortion of ower (2 trials) c
C. hirtella ♀ × C. rehmannii a. var. littoralis ♂Onset of fruit (1 trial) c
C. hirtella ♀ × C. sessilifolia ♂Onset of fruit (1 trial)
C. megarrhiza ♀ × C. abyssinica ♂Onset of fruit (1 trial) b & c
C. megarrhiza ♀ × C. hirtella ♂Abortion of ower (1 trial)
C. megarrhiza ♀ × C. rehmannii a. var. littoralis ♂Onset of fruit (1 trial)
C. megarrhiza ♀ × C. sessilifolia ♂Abortion of ower (2 trials)
C. megarrhiza ♀ × C. trilobata ♂Onset of fruit (Fig. 13c; 1 trial)
C. microphylla ♀ × C. megarrhiza ♂Abortion of ower (1 trial) a
C. microphylla ♀ × C. trilobata ♂Onset of fruit (1 trial) b & c
C. rehmannii var. rehmannii ♀ × C. rehmannii a. var. littoralis ♂Onset of fruit (1 trial) b
C. rehmannii var. rehmannii ♀ × C. trilobata ♂Abortion of ower (2 trials) b
Roy and Roy (1971a) report an intergeneric cross between a female C.grandis
and a male individual of the monoecious Diplocyclos palmatus, resulting in a morpho-
logically intermediate F1 ospring in 5% of the trials. All F1 individuals are female,
indicating that the X chromosome bears at least one gene for maleness suppression,
which is dominant over the maleness genes of D. palmatus. Whether the F1 is fertile, is
not clear, as the authors report successful back-crossing only with a female [sic, male?]
Monograph of Coccinia (Cucurbitaceae) 29
C.grandis but not with D. palmatus. To the author’s knowledge, there are no reports
of the F2 generation. However, if the parental sexes were the other way around (male
C.grandis × female D. palmatus), fertilization was not possible.
Plant – biotic environment interactions
Pollination
Although bee pollination is observed for only a few species, most Coccinia species ex-
hibit characters that support a general attraction to bees. e petal color is commonly
pale yellow but can also range to white, pale pink or bright orange, with green, yellow,
orange to purple venation. Anthesis is during the day in C.abyssinica, C.adoensis var.
aurantiaca, C.grandiora, C.grandis, C.hirtella, C.megarrhiza, C.microphylla, C.re-
hmannii, C.sessilifolia, and C.trilobata (pers. observ.) but often only for a few hours
(in, e.g., C.megarrhiza, C.rehmannii). Zimmermann (1922b) reports pollen release
in C.grandiora at 6.30 a.m. before blooming of the ower, opening of the ower
Figure 16. Hybrids of Coccinia species. a Coccinia grandis ♀ × C. hirtella ♂ b Coccinia hirtella ♀ ×
C.grandis ♂ c Coccinia grandis ♀ × C. pwaniensis ♂; note that the pollen sacs are not open although the
ower is in full bloom indicating sterility d Left ower: male C. sessilifolia, right ower: male ower of
C.grandis ♀ × C. sessilifolia ♂ (C. grandis owers are about equally sized as C. sessilifolia).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
30
between 7 and 8 a.m. and wilting after noon. Ash (J.W. Ash 898) reports ower open-
ing a.m. in C.schliebenii. Anthesis time in the other Coccinia species is not reported,
but also likely to happen during the day. e scent is rather weak and dull sweetish,
resembling that of honey melon, in C.abyssinica, C.grandiora, C.hirtella, C.rehman-
nii, C.sessilifolia, and C.trilobata, weak but fresh in C.grandis, and intense, sweet, and
fruit-like (like honey melon) in C.adoensis var. aurantiaca and C.megarrhiza. e only
evidence of an alternative to bee pollination is C.ogadensis, which is reported to smell
of rotten meat (P. Ellis 163 and 383). However, it is unclear whether the owers emit
a fetid scent or whether the smell comes from crushed vegetative parts, as it is known
from Momordica foetida or Kedrostis foetidissima (Jacq.) Cogn. (Jerey 1967).
Bee pollination is conrmed for C.rehmannii (C.J. Ward 12250), C.adoensis var.
aurantiaca (Fig. 10b), C.grandiora, and C.grandis. Observed pollinators of C.grandis
are Trigona apicalis Smith, 1857 and Trigona collina Smith, 1857 in ailand (Jongjitvi-
mol and Wattanachaiyingcharoen 2006), and Megachile sp. in Cambodia (H. Schaefer,
pers. comm.). e present author observed a halictid bee (Fig. 10b; identication by
H. Schaefer, pers. comm.) in a male C.adoensis var. aurantiaca walking on the globose
anther head and collecting pollen in the corbicula. Stigmas in Coccinia are lobed (Fig.
11b) or bulging (Fig. 11c), and nectaries are located presumably in the hypanthium, so
one can assume stripping of the pollen from the venter when crawling into the ower.
Zimmermann (1922b) also observed the circling around the anther head in the large-
owered C.grandiora. He identied the visiting small bee as Trigona sp. He also noted
that a bee just having visited a Momordica ower walked on the inner side of the corolla
loading dorsally located pollen on the anthers of a male C.grandiora ower.
Seed dispersal
ere are no observations of actual seed dispersal but mammals and birds appear to
be attracted by the fruits and likely act as seed dispersers. Fruit bats such as Cynopterus
sphinx (Vahl, 1797) feed on C.grandis fruits in ailand (Elangovan et al. 2001; Ruby
et al. 2000). Fruits of C.grandis are also taken up by birds (Bhatt and Kumar 2001) and
eaten by humans (Voigt 1845). Elephants also feed on C.grandis (Mubalama 2000)
and are possibly also seed dispersers. From its introduction to Pacic Islands, dispersal
of C.grandis by humans is well-known (Muniappan et al. 2009). Human dispersal, in
many cases likely to be intentional (Starr et al. 2010), also explains the occurrence of
this species in the Neotropics and even in Missouri, USA. Some occurrences in Aus-
tralia can also be explained by escape from gardens. Zimmermann reports feeding on
C.grandiora fruits by birds, small mammals but also snails and beetles (Zimmermann
1922a), the latter two unlikely being seed dispersers. e forest weaver Ploceus bicolor
Vieillot, 1819 was observed to feed on fruits of C.mackenii (Bleher et al. 2003). Stan-
ford and Nkurunungi (2003) report diering preference of Coccinia plant parts by
gorillas. Whereas the gorillas feed on the leaves and fruit pulp of C.mildbraedii but not
the seeds, they take only the leaves of C.barteri.
Monograph of Coccinia (Cucurbitaceae) 31
Successful seed germination in Munich Botanical Garden indicates that passage
through a digestive tract is not necessary at least for C.abyssinica, C.adoensis var.
jereyana, C.grandiora, C.grandis, C.hirtella, C.megarrhiza, C.microphylla, C.reh-
mannii, C.sessilifolia, and C.trilobata. However, whether seeds would survive intestine
passage and the role of endozoochoric dispersal is also not known.
Interaction with ants
Many species of Coccinia bear extranuptial glands (nectar producing glands outside of
the ower) on the lower lamina of the leaves and/or on the bracts and probracts (Figs
7b, 8a). e glands are sunken into the surface and are surrounded by cells with a
thicker cell wall (Muhammad Ilyas 1992; pers. observ.). Ants take up the sweet-tasting
sap in C.grandiora (Zimmermann 1922b) and in C.grandis (pers. observ.). Whereas
Ilyas (1992) reports aggressive behavior of the ants on herbivores for Indian C.gran-
dis, the present author could not observe this in Tanzanian C.grandis. Nieuwenhuis
von Üxküll-Güldenbandt (1907) found a weak attraction of ants and heavy damage
by herbivores in C.grandis in Bogor Botanical Garden (Java, Indonesia). In addition,
Zimmermann (1922b) does not nd aggressive behavior in C.grandiora either but
reports that the ants attacked a caterpillar he had placed onto the plant. Agarwal and
Rastogi (2008), on the other hand, report a signicant reduction in residence time
of herbivores on the cucurbit Lua aegyptiaca when ants are patrolling on the plant.
Most likely, there is no close relationship to certain ant species as guardians, and plant-
defense is carried out only by a few ant species. How Coccinia species without or few
probracts, bracts or sublaminal extraoral nectaries (e.g., C.microphylla) react when
damage by herbivores occurs, is unknown. Agarwal and Rastogi (2008) found an in-
crease of total numbers of extraoral nectaries over time but did not discuss changes of
nectary density as reaction to grazing.
Diseases and parasites
Some research has been undertaken on parasites and diseases for C.grandis for its status as
crop but also as weed. As C.grandis is naturalized on several Pacic islands, in Australia, and
the Neotropics, the plants can either overgrow other plants or represent a non-specic host
for diseases of cucurbitaceous crops (Bamba et al. 2009; Muniappan et al. 2009). Its rapid
growth can become problematic, as Pangelinan (2002) reports that C.grandis covered 35%
of the vegetation of the island of Saipan only eleven years after its introduction.
Many dierent organisms are reported to live in, on, or to feed from Coccinia spe-
cies. Beetle and y larvae are either a disease for Coccinia, or in some cases, they are used
to eradicate C.grandis. Fruits of C.grandis are a host for the larvae of the melon y Bac-
trocera (= Dacus) cucurbitae (Coquillett, 1899), a tephritid fruit y (Uchida et al. 1990).
Bactrocera cucurbitae larvae usually populate the fruits but are also reported to hatch from
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
32
galls (Murthy 1959). e galls are not produced by these ies, however, but by the gall
midge Lasioptera (= Bimba) toombii (Grover, 1962) (Bhatia and Mahto 1968). e gall
infestation is interpreted as non-specic, as the female y might not be able to dierenti-
ate between the gall and an unripe fruit, which would be the usual target. In addition,
also the tephritid fruit y Dacus ciliatus Loew, 1862 infests the galls, sometimes even
together with Bactrocera cucurbitae (Bhatia and Mahto 1968). e galls in C.grandis
do not only result from Lasioptera toombii but can also be produced by the Itonidid gall
midge Neolasioptera cephalandrae Mani, 1934 (Dharmamaraju 1968), which is reported
to be the major disease in C.grandis in India (Unni et al. 1976). e galls induced by
Neolasioptera cephalandrae also appear to be gateway for a fungal infection with a mold,
which is identied tentatively as Cladosporium sp. (Krishnamurthy 1984).
Other major cucurbit pests can also use C.grandis as a host such as Diaphania
(= Palpita) indica (Saunders, 1851) (Lepidoptera: Pyralidae), Aulacophora foveicollis
(Lucas, 1849) (Coleoptera: Chrysomelidae), Leptoglossus australis (Fabricius, 1775)
(Hemiptera: Coreidae), Aphis gossypii Glover, 1877 (Hemiptera: Aphididae), Liri-
omyza spp. leafminers (Diptera: Agromyzidae), Bemesia spp. white ies (Hemiptera:
Aleyrodidae) (Bamba et al. 2009), and Epilachna vigintioctopunctata (Fabricius, 1775)
(Coleoptera: Coccinellidae) (Maurice and Ramteke 2012).
As a result of the damage that can be done to cucurbitaceous crops and of its
weedy behavior on Pacic islands, larvae of the clearwing moth (Sesiidae) Melittia oed-
ipus Oberthür, 1878 and the weevil (Curculionidae) species Acythopeus burkhartorum
O’Brian, 1998 and Acythopeus cocciniae O’Brian, 1998 were introduced to Hawaii for
biological pest control against C.grandis (Muniappan et al. 2002). Immediately after
hatching from the eggs, Melittia oedipus larvae bore into the stems, where they live and
pupate after two to four months (Chun 2002). is moth, originating from Zanzibar
(Oberthür 1878), appears to be quite specic as larvae only rarely develop on Cucumis
sativus L. (Chun 2001). Also Zehneria guamensis (Merrill) Fosberg, a Guam endemic,
is not attacked by M. oedipus (Bamba et al. 2009; Reddy et al. 2009). As C.grandis is
a noxious weed in Hawaii (Hawaiian Department of Agriculture 1992) active search
for pests for biological control was undertaken, which led to the discovery of two
new beetle species from Kenya: Acythopeus burkhartorum whose larvae produce galls
in young shoots, and A. cocciniae whose larvae mine the leaves (Chun 2002; O’Brian
and Pakaluk 1998). O’Brian and Pakaluk report a close morphological similarity of
both Acythopeus species to A. cucurbitae (Marshall), which is a major pest on various
cucurbitaceous crops in Africa, the Middle East, and South India.
Many crop plants are attacked by root parasites or diseases, but there is little
known from Coccinia. Only root lesion nematodes Pratylenchus dasi Fortuner, 1985
(nom. nov. for P. capitatus Das & Sultana, 1979) and P. crassi Das & Sultana, 1979
were described from the soil around the roots of C.grandis (Das and Sultana 1979;
Siddiqi 2000), but it is not known if they harm the plants.
e only known plant parasite growing on Coccinia is the hemiparasitic vine Cus-
cuta chinensis Lam., which is reported to grow on C.grandis in Gujarat, India (Patel
and Patel 2010).
Monograph of Coccinia (Cucurbitaceae) 33
Table 5. List of fungi reported from Coccinia species (sorted by phylum of the fungus).
Fungus Host Symptom Citation
Plasmopara cubensis (Berk. & M.A.Curtis)
C.J.Humphrey (Peronosporales, Oomycota) C. grandis Downy
mildew Selby (1899)
Alternaria pluriseptata (P.Karst. & Har. ex Peck) Jørst.
(Saccharomycetales, Ascomycota) C. grandis Fruit rot Chagale and Bhale (2010)
Cercospora cocciniae Munjal, Lall & Chona
(Dothideales, Ascomycota) C. grandis Leaf spot
disease
Rangaswami and
Chandrasekaran (1961)
Cercospora elaterii Pass. C. grandis Leaf spot
disease
Rangaswami and
Mahadevan (2004)
Colletotrichum gloeosporioides (Penz.)
Sacc.(Glomerellales, Ascomycota) C. grandis Fruit rot Bhagavan Reddy and
Reddy (1987)
Colletotrichum orbiculare (Berk. & Mont.) Arx C. grandis Anthracnose
fruitrot Imbumi (2004)
Corynespora cassiicola (Berk. & M.A.Curtis) C.T.Wei
(Pleosporales, Ascomycota) C. grandis Leaf blight Philip et al. (1972)
Curvularia pallescens Boedijn (Pleosporales) C. grandis Black rot Imbumi (2004)
Erysiphe cichoracearum DC. ex Merat
(Erysiphales,Ascomycota) C. grandis Powdery
mildew Imbumi (2004)
Fusarium moniliforme J.Sheld. (Hypocreales,
Ascomycota) C. grandis Fruit rot Kapoor et al. (1981)
Geotrichum candidum Link (Pleosporales) C. grandis Fruit rot Chagale and Bhale (2010)
Sphaerotheca fuliginea (Schltdl.) Pollacci (Erysiphales) C. grandis Powdery
mildew Imbumi (2004)
Puccinia cephalandrae ümen
(Uredinales,Basidiomycota) C. quinqueloba Rust Berndt (2007)
Puccinia cephalandrae-indicae Syd. & P.Syd. C. grandis Rust Berndt (2007)
Puccinia physedrae Syd. C. barteri Rust Berndt (2007)
Puccinia windhoekensis Mennicken, Maier & Oberw. C. rehmannii? Rust Mennicken et al. (2005)
Rhizoctonia solani Khun (Cantharellales,
Basidiomycota) C. grandis Fruit rot Bhagavan Reddy and
Reddy (1988)
Several fungi have been reported from Coccinia (Table 5). e rust fungus Puc-
cinia windhoekensis Mennicken, Maier & Oberw. was described on Coccinia rehman-
nii (Mennicken et al. 2005), although Berndt (2007) noticed a great similarity of this
rust to P.ctenolepidis Ramachar & Bagyanar. Berndt could not conrm the identity
of the host specimen, so it seems to be likely that it was misidentied, since Ctenolepis
cerasiformis looks quite similar to C.rehmannii.
ere are several reports of plant viruses from Coccinia species. Purcifull and col-
leagues (1988) tested the infectability of several Cucurbitaceae to dierent plant vi-
ruses. ey found that C.grandis can be infected by the papaya ringspot virus type
W (PRSV-W) and the Trichosanthes virus but not by the cucumber mosaic virus,
squash mosaic virus, watermelon mosaic virus-2, and the zucchini yellow mosaic virus.
PRCV-W infections of C.grandis are also reported from several Pacic islands (Davis
and Ruabete 2010).
Verma et al. (1983) suggest a yet undescribed mosaic virus, which is expressed in
the occurrence of deformed leaves and a mosaic pattern in C.grandis leaves. A strain
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
34
of the Moroccan watermelon mosaic virus, a Potyvirus, can infest Coccinia barteri
(Owolabi et al. 2012), whereas the infection of C.sessilifolia with this virus, maybe a
dierent strain, failed (van der Meer and Garnett 1987).
Use, economic potential, and phytochemistry
Several Coccinia species are used by tribal communities, mainly as a food source but
also for cultural applications (for details see species descriptions). Coccinia grandis is
notable for its economic value (although often cited erroneously as Coccinia cordifolia
or C.indica), whereas the importance of C.abyssinica is only regional. Other species
are used by local tribes only.
Coccinia grandis is used in a wide variety of applications. e plant is well-known
in India, where its fruits had an impact even in classical Sanskrit literature. e red
fruits are regularly used to describe lips, such as those of a beloved wife, who is de-
scribed by her husband in Kālidāsa’s poem Meghadūta (Wilson 1867) or those of the
goddess Sita and the god Rama in the epic Ramayana (Dutt 1891–1894). However,
the fruits are also edible (raw when ripe and cooked when unripe) and are valued for
their high content of carotenoids, esp. lycopene (Barua and Goswami 1979). Also
young shoots and leaves are eaten as spinach and contain high amounts of lutein and
other carotenoids (Addis et al. 2009; Wasantwisut and Viriyapanich 2003). e high
carotenoid value is of special importance in developing countries, as vitamin A de-
ciency is widespread among young children and pregnant women (WHO 2009). So-
cial marketing has proven to be valuable in promoting the use of C.grandis to prevent
vitamin A deciency (Chittchang et al. 1999). Domestication of C.grandis is in an
early stage but promising cultivars are developed in South and SE Asia (Bharathi 2007;
Engle et al. 1998; Ramachandran and Subramaniam 1983). Additionally, the leaves
seem to be a good source of selenium and potassium, as well as vegetable protein (Xu
et al. 2003; Xu et al. 2004). In Africa, C.grandis is mostly used from wild collections
(Addis et al. 2009; Imbumi 2004). Contraindications to the use of C.grandis are also
reported (Adanson 1757; Orech et al. 2005), but these might also be the result from
either misidentication or regional chemo-varieties with diering amounts of second-
ary metabolites.
Coccinia grandis has been used in Indian traditional medicine for several hun-
dred years (Nadkarni and Nadkarni 1976; Ramachandran and Subramaniam 1983).
ere are some studies that suggest a high potential for the use of C.grandis leaf ex-
tracts in diabetes treatment (Azad Khan et al. 1980; Kuriyan et al. 2008; Munasinghe
et al. 2011). Parts of the observed eects are explained by inhibition of gluconeo-
genesis in the liver due to repression of glucose-6-phosphatase (Hossain et al. 1992)
and fructose-1,6-bisphosphatase (Shibib et al. 1993). Also an activating eect on the
promotor of the glucose transporter gene GLUT1 from rats is reported (Graidist and
Purintrabipan 2009). Eshrat (2003) observes a positive eect of C. grandis in rats
with hyperlipidemia, which is often connected to diabetes. However, its eectivity in
Monograph of Coccinia (Cucurbitaceae) 35
diabetes treatment and the overall experimental design is in dispute (Ramachandran
and Subramaniam 1983; Sadikot 2009), and more research to test the medical value
is necessary. Since 2005, more than 15 studies researched chemical compounds in
C.grandis and tested their validity in folk medicine. Some applications by tribal people
could be reproduced ex situ but research is still in its infancy. Suggested eects are, e.g.,
anti-anthelmintic (Dewanjee et al. 2007b), anti-tussive (Pattanayak and Sunita 2009),
hepatoprotective (Moideen et al. 2011; Vadivu et al. 2008), antioxidative (Umama-
heswari and Chatterjee 2008), antipyretic, analgesic, and anti-inammatory (Niazi
et al. 2009), anti-ulcerogenic (Mazumder et al. 2008), and antimicrobial (Bulbul et
al. 2011; Dewanjee et al. 2007a; Farrukh et al. 2008; Shaheen et al. 2009). Antimi-
crobial activity is explained by the occurrence of a protease inhibitor (Satheesh and
Murugan 2011). Observed xanthine oxidase inhibition and antiuricaemical activity
(Umamaheswari et al. 2007) suggests a use for gout treatment. Female rats with hyper-
prolactinemia-caused infertility regain fertility when treated with an aqueous extract of
C.grandis stems and leaves (Jha et al. 2010).
Coccinia abyssinica is mainly an Ethiopian tuber crop. Under the name anchote, its
starch containing (c.20%) tubers are an important staple food in the SW semi-humid
highland regions (Aga and Badada 1997; Asfaw 1997; Hora 1995). Additionally, the
tubers contain a relatively high amount of calcium, which might explain the local be-
lief that the plant helps with repairing bone fractures and displaced joints (Hora 1995).
Locally (around Dembi Dolo, Oromia), young shoots and leaves are also eaten (Hora
1995). Although the fruits of the cultivated landraces are not eaten (Getahun 1973),
the use might be benecial due to the carotenoid content of the fruits, which are likely
to be comparable to those of C.grandis. However, fruits of wild races of C.abyssinica
are already used (Asfaw and Tadesse 2001). In Wollega (W Ethiopia), C.abyssinica is
also used to treat gonorrhea, tuberculosis, and cancer, as well as in traditional ceremo-
nies and celebrations and for animal fattening (Gelmesa 2010). Currently, much eort
is put into the development of anchote to increase the yield by selection of cultivars
with larger tubers and by improving crop growing with better suited fertilizers (Abera
and Guteta 2007; Bekele et al. 2014; Mengesha et al. 2012).
Also other species of Coccinia are used as food sources but if so, then only locally.
In these species, such as C.sessilifolia, some wild landraces lack bitter substances (Bosch
2004; Dinter 1912). Bitterness in Cucurbitaceae is mainly caused by triterpenoids
called cucurbitacins, although not all cucurbitacins are bitter. Cucurbitacins are often
cytotoxic and often exist as β-glucosides (Miró 1995). All Coccinia species screened so
far contain cucurbitacins, although the cucurbitacin type, organ, and time of expres-
sion dier greatly. Whereas fruits of C.hirtella and C.quinqueloba contain glycosidic
cucurbitacin B, C.adoensis from South Africa contains aglycosidic cucurbitacin B only
in unripe fruits and traces of cucurbitacin D but not in ripe fruits (Rehm et al. 1957).
Unripe fruits of C.rehmannii and C.sessilifolia are not bitter, and therefore lack bitter
cucurbitacins (Enslin et al. 1956). Njoroge and Newton (1994) tested the type and
distribution of cucurbitacins within the plant in dierent Cucurbitaceae and found in
Kenyan C.adoensis plants cucurbitacins H, I, and R in the stem but no cucurbitacins
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
36
in the roots, leaves, fruits, or seeds. Coccinia trilobata was found to contain the cucur-
bitacins B, D, and G in the stems, cucurbitacin D, H, I, and R in the leaves, and cu-
curbitacin G in the fruits, with no cucurbitacins in the roots and seeds. However, there
seems to be much variability, as there are reports of edible (non-bitter) C.trilobata
leaves (Coilly? 24, F. Msajiri 19). Coccinia grandis is also reported to contain cucurbi-
tacin B (Bhakuni et al. 1962), and bitter and sweet fruited varieties are known. Guha
and Sen (1973) nd that cucurbitacin B has an antigibberelic eect, and its occurrence
in seeds of C.grandis might enable or increase dormancy of the seeds.
Cucurbitaceae are also known for the occurrence of non-coded amino acids, such
as citrulline in Citrullus lanatus (unb.) Matsum. & Nakai (Wada 1930). In a survey
of such amino acids in Cucurbitaceae C.grandis and C.hirtella seeds were found to
contain citrulline in low amounts, β-(pyrazol-1-yl)-L-alanine in very high amounts and
the peptide Γ-L-glutamyl-β-(pyrazol-1-yl)-L-alanine in intermediate amounts (Dunnill
and Fowden 1965). is pattern is similar to those that were found in Diplocyclos tenuis
(Klotzsch) C.Jerey, Acanthosicyos horridus Welw. ex Hook.f., Peponium hirtellum Ke-
raudren, Ruthalicia eglandulosa (Hook.f.) C.Jerey, Dactyliandra welwitschii Hook.f.,
and Ctenolepis cerasiformis (Stocks) C.B.Clarke (all in the same tribe as Coccinia).
Evolution and phylogeny
Recent phylogenetic analyses (Kocyan et al. 2007; Schaefer and Renner 2011b) show
that Coccinia belongs to the tribe Benincaseae with a moderately supported sister
group relationship to the genus Diplocyclos. However, the backbone of the tribe is not
resolved and the relationship of the Coccinia-Diplocyclos clade to the other genera is
unknown. Citrullus, Cucumis, or Scopellaria cluster with this clade but each without
support, and morphological characters also do not seem to suggest any closer relatives.
Both phylogenies, plastid (Fig. 17) and the nuclear LEAFY-like 2nd intron
(Fig. 18), suggest four major clades, although the backbone lacks bootstrap or
posterior probability support (Holstein and Renner 2011b). e Coccinia rehmannii
clade (IV) is well-supported in all phylogenies. e Coccinia quinqueloba group (II) is
well-supported in the plastid DNA analysis, and consists of C.hirtella, C.mackenii,
and C.quinqueloba. Additionally, C.sessilifolia belongs to this group, but it is only
supported here by the nrDNA data. According to the nuclear data, the C.barteri clade
(III) is nested within the C.adoensis clade (I). e plastid analysis tree separates these
two clades but without support.
e C. rehmannii clade (IV) consists of ve species. Coccinia abyssinica and
C. megarrhiza are sister species from Ethiopia and semi-arid parts of N Kenya and
Somalia (Fig.20). ey dier ecologically with the former species occurring in the semi-
humid highlands and the latter one in the semi-arid lowlands. Both species dier weakly
in morphology, and hybridization cannot be ruled out. e plastid haplotypes of both
species do not form clades in the tree, which might be explained best by incomplete
lineage sorting. e other three species of clade IV contain several plastid haplotypes and
Monograph of Coccinia (Cucurbitaceae) 37
nrDNA sequences that each also do not form clades. e geographical distribution of the
haplotypes is not assessed. e three species, however, are distinct. Coccinia rehmannii
occurs in southern Africa while the other two species occur in NE Africa. In Coccinia
rehmannii four forms can be recognized, which are included in the plastid phylogeny: (1)
an inland form from dry habitats with small globose fruits (type form / var. rehmannii),
Figure 17. Phylogenetic relationships in Coccinia based on ve plastid DNA loci (matK, ndhF–rpl32
intergenic spacer (IS), rpl20–rps12 IS, trnL intron, trnL–trnF IS, trnS–trnG IS) obtained for 75 accessions
from 24 species. Shown is the topology of the 50% majority rule consensus tree obtained from Bayesian
analysis including simple gap coding for ingroup InDels. Numbers above the branches are posterior prob-
ability values ≥ 0.98 with values “with InDel coding” rst, followed by “without InDel coding.” Numbers
below the branches are bootstrap support values from ML analysis. Topologies from the dierent analyses
were not contradictive, although some clades were not resolved without gap coding. Roman numbers
indicate clades as discussed in the text: I = C. adoensis clade, II = C. quinqueloba group, III = C. barteri
clade, and IV = C. rehmannii clade.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
38
(2) a form similar to var. rehmannii, but with larger globose fruits (described by Dinter
and Gilg as C.ovifera), (3) a long-petiolate and long-peduncled coastal form from the
(semi-)humid Southeast (described by Meeuse as var. littoralis), and (4) plants with
oblong fruits occurring in all semi-humid areas from the Southeast to the northern parts
in the periphery of the C.rehmannii distribution (C.rehmannii 5; here referred to Coccinia
rehmannii a. var. littoralis). None of these forms cluster together. e other two species
dier morphologically and ecologically from each other: C.trilobata has, e.g., oblong
fruits and occurs in the semi-humid uplands, and C.microphylla has globose fruits and
occurs in the semi-arid lowlands (Fig. 33). Interestingly, C.microphylla does not dier
morphologically from the C.rehmannii form from the dry inland. is scenario suggests
incomplete lineage sorting and a speciation event with ecological dierentiation in the
northeastern Africa but not in southern Africa as intermediate collections between the
four forms are found regularly. e distribution of these three species and the estimated
age of this clade of 3.2 Ma suggest either a long distance dispersal or vicariance. As each
of the three species contains several plastid haplotypes, vicariance is more likely, which
indicates that semi-arid conditions might have prevailed between today’s Tanzania and
Zimbabwe. is has been suggested several times for dierent clades under the term
“arid track” (Balinsky 1962; de Winter 1971).
Figure 18. Phylogenetic relationships in Coccinia based on 505 nucleotides of the nuclear LEAFY-like
2nd intron, obtained for 37 accessions from 23 species analyzed under maximum likelihood (ML) and the
GTR + Γ model. Numbers below branches refer to ML bootstrap support > 80% from 1000 replicates.
e dots at nodes and behind the two accessions refer to uniquely shared indels. Roman numbers indicate
clades as discussed in the text: I = C. adoensis clade, II = C. quinqueloba group, III = C. barteri clade, and
IV = C. rehmannii clade.
Monograph of Coccinia (Cucurbitaceae) 39
e C.quinqueloba clade (II) is only supported in the nrDNA phylogeny, as plastid
sequences of C.sessilifolia and its distinctly petiolate variety variifolia lack synapomorphies
that support a closer relationship to any clade in Coccinia. e two varieties of C.sessilifolia
occur in the semi-arid and sub-semi-humid inland (Fig. 40; see species description),
whereas the other three species prefer more humid habitats in the Southeast (Fig. 30; see
species description; Holstein and Renner 2011b). One species, C.hirtella, occurs in the
rather open habitats, especially in the Drakensberg Mountains, which receive high amounts
of rainfall. Coccinia mackenii occurs in remnant forest sites in the humid Southeast of
southern Africa, whereas C.quinqueloba occurs only in coastal bushlands of the Eastern
Cape, where it receives less precipitation than the other two species but has a more evenly
distributed water availability all over the year (Holstein and Renner 2011b). As C.mackenii
and C.quinqueloba do not co-occur but have similar ecologies, and as they only slightly
dier morphologically but hardly produce hybrids (see chapter Hybridization and crossing
experiments), a recent allopatric speciation event is probable. e lack of dierentiation in
the plastid sequences over 3500 bp between two accessions might support this hypothesis.
In contrast to the C.rehmannii clade, all species of this clade occur exclusively in southern
Africa, although the clade is older (c. 5.0 Ma vs. 3.2 Ma).
e C. adoensis clade (I) contains several morphologically and ecologically well
dierentiated species (Holstein and Renner 2011b). ere are three subclades in the
plastid tree with accessions having the name Coccinia adoensis. e type form (Fig. 21;
see species description) from Ethiopia (no DNA sequences available) is morphologically
inseparable from South African forms (C.adoensis 1 and 6). Geographically between
those two populations, however, there are many populations that mostly dier gradually
in length and density of trichomes. Two forms (with especially dense and long trichomes,
respectively) could be assessed geographically and are accepted by the present author as
varieties. e Coccinia adoensis var. aurantiaca accessions are neither in the plastid nor
in the nuclear tree monophyletic but share a dense indumentum. ese forms cluster in
the nuclear tree with collections that have a less dense indumentum and thus are rather
referred to as C.adoensis var. adoensis (Fig. 18). In the plastid tree, these collections
cluster together with a Kenyan specimen of C.adoensis var. adoensis and var. jereyana
(Fig. 17). Coccinia adoensis var. jereyana, however, shares the longer trichomes (Figs
3a, 5c) of some C.senensis, but it diers from these by lacking subulate calyx lobes
and a 569 bp deletion in the trnSGCU–trnGUUC intergenic spacer region. However, one
collection that does not dier morphologically from the variety jereyana (R.E. Gereau
and C.J. Kayombo 3582) clusters within the East African forms of C.adoensis, which
indicates either homoplasy of the trichome length or gene ow. Additionally, gene ow
among the C.adoensis clades might also occur. Holstein and Renner (2011b) found a
collection from Namibia (C.adoensis 5) that contained ITS sequences that are otherwise
found exclusively in the South African and East African plastid haplotypes. us it can
be suggested that all these forms belong to one widespread species, C.adoensis, which
contains dierent plastid haplotypes. From this widespread species, several populations
might have undergone ecological and morphological divergence. Coccinia grandiora and
C.schliebenii are nested within one C.adoensis subclade, and they occupy rather humid
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
40
habitats while C.grandis and C.ogadensis occupy more arid habitats. Some populations
probably evolved parapatrically in former times with morphological shifts (C.senensis,
C.pwaniensis, C.samburuensis) or evolved in allopatry (C.intermedia) (Holstein and
Renner 2011b; Fig. 19). Some populations, however, did not diverge suciently to be
taxonomically well-separated as a species, such as C.adoensis var. aurantiaca.
e C. barteri clade (III) mostly consists of rainforest species from West and
Central Africa, except for the recently described C.intermedia (Holstein and Renner
2011a). Coccinia mildbraedii (including C. ulugurensis) also diers ecologically, as
it occurs in mountain forest communities not in typical lowland rainforests as does
the rest of the species. e phylogenetic position of C.intermedia is unclear as the
resolution within this clade is generally poor. Coccinia intermedia shares morphologi-
cal characters with C.adoensis, especially the open campanulate owers. Both species
occur in the same habitat type with the former occurring in West Africa and the lat-
ter north and east of the Central African rainforests. If the C.barteri clade is indeed
nested in the C.adoensis clade, as suggested by the nuclear phylogeny, then it is pos-
sible that C.intermedia might have split allopatrically from a proto-C.adoensis species
Figure 19. Scenario of evolution in the C. adoensis clade. e green line surrounds today’s distribution of
C. adoensis. Blue lines surround today’s distributions of C. senensis and C. pwaniensis. Blue arrows indicate
peripatric speciation without shift in precipitation preference. Yellow arrows indicate speciation with shifts
towards more arid habitats. Black arrows indicate speciation with shift towards more humid habitats.
Monograph of Coccinia (Cucurbitaceae) 41
and is sister to the rest of the C.barteri clade (Fig. 19). en, the common ancestor of
the other species of the barteri clade might have shifted the habitat preference towards
perhumidity once and evolved allopatrically in refugia during arid periods of the Plio-
cene and Pleistocene. Alternatively, the habitat of C.intermedia would be explained
as a reversal from a rainforest distributed common ancestor of the C.barteri clade. As
the frequency of the Pleistocene climatic oscillations increased, reproductive isolation
did not always occur, leading to weak morphological dierentiation of interbreeding
populations, such as in the polymorphic C.barteri (Holstein and Renner 2011b).
Identification of Coccinia species
Possible confusion with other genera
Some Coccinia species are easily confused with collections of other Cucurbitaceae gen-
era (Table 6). e similarity is sometimes striking and without generative structures,
one might need some experience to dierentiate between the genera.
Characters for species discrimination
ere is no character that is useful for all species. For example, whereas the direction
of the calyx lobes can be a useful character for some species (e.g., C.grandis, C.inter-
media, C.keayana), it is less useful in others (e.g., in the C.quinqueloba clade). Col-
lections without owers are harder to identify. In some cases it is almost impossible
to discriminate between species if owers are lacking. Identication of only vegetative
material is often possible but needs experience. e indumentum can be a useful char-
acter; especially the trichomes (length, somewhat also the shape) on the abaxial side of
the petiole and the lower leaf lamina can be helpful. However, the trichomes on the
adaxial side of the petiole and the leaf margin do not seem to have any purpose for
species identication.
Key to Coccinia species
e key is made from observations of herbarium material but also includes some char-
acters from personal observations of living material and observations as given on her-
barium labels. Fresh material is not needed, however, to use the key. e term ‘articu-
late’ refers to dried trichomes that appear wrinkled due to equatorially sunken cell walls
(see Fig. 3a) but not to trichomes with ramications, which have never been observed
in Coccinia. In the living state, these trichomes are rather long and sti. e term “den-
tate” refers to the sometimes colored structures (hydathodes?) at the leaf margin and
leaf tip (Figs 6, 7a, 8a, 16a, 16b, 21, 39).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
42
Table 6. African taxa that Coccinia species might be misidentied with.
Coccinia species Similar taxon Dierences
C. barteri, C. heterophylla,
C.racemiora
Bambekea
racemosa Cogn.
B. racemosa: petals free, has veins running along the leaf margin
Coccinia: petals connate, veins not running directly along the margin
C. barteri, C. heterophylla,
C.racemiora Cogniauxia spp.
Cogniauxia: petals free and up to 8 cm long, has veins running along
the leaf margin, prophylls lanceolate, fruits up to 15 × 8 cm (L × D)
large, seeds up to 2 cm long (Schaefer and Renner 2011a)
Coccinia: petals connate and < 3.5(–4.5 cm), leaf veins not directly
running along the leaf margin, probracts ovate or missing, fruit
diameter < 4(–5) cm, if length > 6 cm, then not ovate, seeds
<0.8cm long
W African rainforest
species Ruthalicia spp.
Ruthalicia: bracts lanceolate, petals free, seeds black or dark brown;
R.eglandulosa with trichomes with a claret-red color (coloration is
often at the ends of the long, centrically sunken-in cells)
Coccinia: bracts ovate or missing, petals connate, seeds gray to beige;
trichomes whitish, beige, yellowish or rarely light brownish
rainforest species Peponium spp.
Peponium: petals free, male owers with long-stretched hypanthium
and three free stamens, which connect only with the long-stretched
anthers, seeds dark colored; P. vogelii: sessile probracts and bracts are
round and up to 3 cm long
Coccinia: petals connate, male owers with perianth tube of which
the length does not exceed two times the diameter, three already
connected laments and a globose anther head, seeds gray to
beige; short petiolate probracts and bracts ≤0.5 cm in C & W
African species
C. schliebenii Lua aegyptiaca
Mill.
L. aegyptiaca: mostly (2–)3–5-d tendrils (check as many as possible),
petals free, petals bright yellow (in L. acutangula (L.) Roxb. also dull
yellowish), stamens 5
C. schliebenii: (1–)2-d tendrils, petals connate, petals dull yellowish
or yellow-orange, stamens 3
C. schliebenii Lagenaria spp.
Lagenaria: often tooth-like glands at the base of the lamina or
along the petiole, trichomes > 1 mm, petals free and white, anthers
serpentine
C. schliebenii: never glands as above, trichomes < 1 mm, petals
connate and dull yellowish or yellow-orange, anthers S-shaped
C. adoensis Eureiandra spp.
Eureiandra: petals free, calyx lobes triangulate to lanceolate, stamens
5, seeds almost globose and whitish (Jerey 1967)
Coccinia: petals connate, calyx lobes lineal, subulate to narrowly
triangulate, stamens 3 in a central column, seeds grayish to beige,
attened
C. microphylla,
C.rehmannii
Ctenolepis
cerasiformis
C.B.Clarke
C. cerasiformis: large roundish, sinuate-ciliate probract, petals <5mm
(Jerey 1967; Meeuse 1962)
Coccinia: probract < 3 mm, ovate or missing, petals > 1 cm
C. microphylla,
C.rehmannii
Dactyliandra
spp.
Dactyliandra: large roundish, sinuate-ciliate probract, petals < 5 mm;
D. stefaninii (Chiov.) C.Jerey from N Africa lacks the probracts
but the seed shape is conspicuously rounded (Jerey 1967; Meeuse
1962)
Coccinia: probract < 3 mm, ovate or missing, petals > 1 cm; seeds
asymmetrical (almost falcate)
Monograph of Coccinia (Cucurbitaceae) 43
Habitats in this key (not the species descriptions) are given rather crudely and
reect the vegetation that would be found naturally. Savannas and woodlands (tree
stands with not largely overlapping canopies) can also include mopane, but also dry
forests (larger amounts of deciduous trees and overlapping canopies), deciduous thick-
ets, tall grasslands, and secondary vegetation derived from these. “Rainforests” include
gallery forests, semi-deciduous forests derived from rainforests, e.g., in relict areas,
perhumid savanna types, and open areas, in which rainforest would be predominant if
it was not for human impact, or swamps.
A local key for Coccinia from West Africa is provided separately by Holstein and
Renner (2011a). If the plant is collected from outside of Africa, then it is C.grandis.
1 Mature leaves sessile (rst leaves may be petiolate), rarely subsessile; alive usu-
ally bluish-green; glabrous; male owers solitary or in few-owered racemes,
female owers solitary; fruit long ovoid, elliptical to spindle-shaped; preferring
dry habitats; from S Africa (Figs 2b, 7b, 13b) ..... C.sessilifolia var. sessilifolia
1* All leaves petiolate; plant not like in 1 .........................................................2
2 Tendrils mostly bid; usually forest species or from Drakensberg Mts or hu-
mid coastal bushland in SE Africa (in E and W Africa also in woodlands or
savannas) ....................................................................................................3
3 Plant with owers ....................................................................................... 4
4 Corolla ≥ 4 cm long, calyx lobes > 3 mm long; ovaries and fruits long ovoid
to cylindrical; E Africa or Ethiopia .............................................................. 5
5 Leaf surface usually glabrous, rarely with sparse weak thin trichomes on the
abaxial side; leaves profoundly lobed (Fig. 8a); E African (rain) forests of C
Zimbabwe and Mozambique to S Kenya ...............................C.grandiora
5* Leaf surface, at least below (secondary and tertiary veins) densely covered
with small trichomes; leaf shallowly or rarely profoundly lobed; margin of
humid forests and in forests; from N Mozambique to C–S Tanzania or Ethi-
opia to South Sudan ............................................................... C.schliebenii
4* Corolla < 4 cm long ....................................................................................6
6 Calyx lobes (> 2.5 mm) subulate (Fig. 28), western C Africa ... C.heterophylla
6* Calyx lobes < 2.5 mm, if longer then from S Africa ....................................7
7 Calyx lobes > 3 mm, plant from S Africa ....................................................8
8 Leaf lamina and stem usually densely covered with long (> 0.5 mm) tri-
chomes; lamina profoundly lobate and lobulate; lobe tips usually rounded;
pedicels covered with long (> 0.5 mm) trichomes (Fig. 13a) ........ C.hirtella
8* Leaf lamina and stem glabrous or rarely sparsely covered with long trichomes,
with lobes often tapering into an acute tip, only side lobes with a slight lobule
on outer side; pedicels glabrous ................................................. C.mackenii
7* Calyx lobes < 3 mm, plant not from S Africa ..............................................9
9 Flowers in lax many-(> 6-)owered racemes, western C Africa ... C.racemiora
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
44
9* Flowers in dense racemes, few-owered or on a long common peduncle
that surpasses the length of the branched part; female owers may also be
solitary .................................................................................................. 10
10 Male owers on a long common peduncle that surpasses the length of the
branched part; female owers solitary with cylindrical ovary; mountain for-
ests of Kivu Mts, Livingstone Mts, and Eastern Arc Mts, introduced into
Kenyan high mts ...................................................................C.mildbraedii
10* Male owers in a raceme, in which the common peduncle is shorter than the
branched part; female owers in racemes, clustered or if solitary, then with
subglobose to elliptical ovary ....................................................................11
11 Corolla campanulate, calyx lobes erect with recurved tips. Lower leaf surface
at maturity often with white speckles and leaf margin with colored teeth. W
African semi-humid savannas and woodlands ........................ C.intermedia
11* Corolla urn-, cup-, funnel-shaped or narrow campanulate. Calyx lobes vari-
able but not as above. Lower leaf surface rarely with white speckles, teeth on
leaf margin not conspicuously colored. Rainforests of W Africa, C Africa, and
in relict forests to Angola, Zambia?, W Tanzania, Uganda, and the Chiman-
imani Mts (Mozambique, Zimbabwe) .......................................... C.barteri
3* Plant with fruits or vegetative parts only ...................................................12
12 Plant with fruits ........................................................................................ 13
13 Fruit oblong to cylindrical (mature > 5 cm long), plant from E or NE Africa ... 14
14 Lower leaf surface, often also upper surface densely covered with short tri-
chomes; N Mozambique, C and S Tanzania or W Ethiopian to SE South
Sudanian mts .......................................................................... C.schliebenii
14* Upper leaf surface glabrous, lower leaf surface glabrous or rarely with some
trichomes; plant from E Africa, incl. Kivu Mts and Chimanimani Mts (Mo-
zambique, Zimbabwe); hard to dierentiate in shared mountain ranges ...15
15 Probracts > 3.5 mm (Fig. 8a); corolla > 3 cm, calyx lobes > 3 mm; forests and forest
relicts in S Kenya, Mt Meru to Usambara Mts, Eastern Arc Mts, SE Tanzania, N
Mozambique, Chimanimani Mts (Mozambique, Zimbabwe) ...... C.grandiora
15* Probracts < 3.5 mm; corolla < 3 cm, calyx lobes < 3 mm; forests of Kivu Mts,
Livingstone Mts, and Eastern Arc Mts, introduced into Kenyan high mts ....
............................................................................................. C.mildbraedii
13* Fruits ovoid, if long elliptical, then from S Africa .....................................16
16 Fruits in lax racemes, plant from western C Africa .................C.racemiora
16* Fruits in dense racemes or solitary .............................................................17
17 Plant from S Africa; fruits solitary ............................................................. 18
18 Leaf surface and stem usually densely covered with long (> 0.5 mm) trichomes;
lamina profoundly lobate and lobulate; lobe tips usually rounded (Fig. 13a) ..
....................................................................................................... C.hirtella
18* Leaf surface and stem glabrous or sparsely covered with long (> 0.5 mm)
trichomes, with lobes often tapering into an acute tip, only side lobes with a
slight lobule on outer side ......................................................... C.mackenii
Monograph of Coccinia (Cucurbitaceae) 45
17* Plant from W to C Africa to Chimanimani Mts (Mozambique, Zimbabwe) ... 19
19 Plant from western C Africa, not distinguishable with condence without
owers ............................................................C.heterophylla or C.barteri
19* Plant not from western C Africa ...............................................................20
20 Lower leaf surface at maturity often with white speckles and leaf margin with
colored teeth when dry. W African semi-humid savannas and woodlands .....
.............................................................................................. C.intermedia
20* Lower leaf surface rarely covered with white speckles, teeth on leaf margin not
conspicuously colored. Rainforests of W to C Africa to Chimanimani Mts
(Mozambique, Zimbabwe) ...........................................................C.barteri
11* Plant vegetative only ................................................................................. 21
21 Lower leaf surface, often also upper surface densely conspicuously covered
with short trichomes; N Mozambique, C and S Tanzania or W Ethiopian to
SE South Sudanian mts .......................................................... C.schliebenii
21* Leaves glabrous, or if covered with trichomes, then they are long (> 0.7 mm)
or inconspicuous ....................................................................................... 22
22 Plant from S Africa ................................................................................... 23
23 Leaf surface and stem usually densely covered with long (> 0.5 mm) tri-
chomes; leaves profoundly lobate and lobulate; lobe tips usually rounded
(Fig.13a) ...................................................................................... C.hirtella
23* Leaf surface and stem glabrous or sparsely covered with long (> 0.5 mm)
trichomes, with lobes often tapering into an acute tip, only side lobes with a
slight lobule on outer side ......................................................... C.mackenii
22* Plant from W to E Africa .......................................................................... 24
24 Plant from E African (incl. S Kenyan) rainforests or forest relicts, Mt Meru to
Usambara Mts, Eastern Arc Mts to Chimanimani Mts (Mozambique/Zim-
babwe); probracts > 3 mm (Fig. 8a) ....................................... C.grandiora
24* Plant from W, C, or E Africa, if from E Africa, then probracts < 3 mm .... 25
25 From mountain forests of E Africa, incl. Kivu Mts ...............C.mildbraedii
25* Rather from lowland rainforests from W Africa, C Africa, or from rainforests
surrounding the Western Rift ...................................................................26
26 Plant from western C Africa not condently distinguishable without owers
............................................ C.heterophylla, C.racemiora, or C.barteri
26* Plant not from western C Africa ...............................................................27
27 Lower leaf surface at maturity often with white speckles and leaf margin with
colored teeth. W African semi-humid savannas and woodlands ...C.intermedia
27* Lower leaf surface rarely with white speckles, teeth on leaf margin not con-
spicuously colored. Rainforests of W to C Africa to Chimanimani Mts (Mo-
zambique, Zimbabwe) .................................................................. C.barteri
2* Tendrils usually simple, if not, then from semi-arid habitats or E and NE-
African woodlands ....................................................................................... 28
28 Leaves deeply palmately lobed with lineal lobes. If lobes lobulate, then leaf
lamina at lobe base as broad as vein. E Ethiopia and C Somalia ... C.ogadensis
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
46
28* Leaves profoundly, but not deeply lobed, or if deeply lobed, then lobes lan-
ceolate or lobe base broader than vein ....................................................... 29
29 Leaves deeply lobed with lanceolate lobes. Male owers in racemes with short
peduncle and pedicels. Rainforests of C Africa and around the Western Rift
..........................................................................................C.subsessiliora
29* Plant not as above ..................................................................................... 30
30 Leaves 7-lobate, rarely 5-lobate. Outer side of lobes serrate to lobulate with
conspicuously colored tips. Calyx lobes > 4 mm, corolla > 2.5 cm, fruit ripe
>10 cm long, from Samburu area (C Kenya, E Africa) ........C.samburuensis
30* Leaves cordate or 3-lobate. If 5-lobate, then plant not as above. Calyx lobes
shorter than above or if longer then corolla shorter. Fruit shorter than above
or if longer then from S Africa .................................................................. 31
31 Plant glabrous; leaves usually subsessile, 5-lobate. Plant from coastal bush-
lands of Eastern Cape (South Africa) ...................................C.quinqueloba
31* Plant with trichomes or if glabrous, then from dierent region ................. 32
32 Plant glabrous, glaucous. Fruit long (> 6 cm) elliptical to spindle-shaped.
Limpopo Province (South Africa) .................... C.sessilifolia var. variifolia
32* Plant with trichomes, or if glabrous then not glaucous and from dierent region .. 33
33 Lower leaf surface, often also upper surface densely conspicuously covered
with short trichomes; bracts > 3 mm; calyx > 1 cm, corolla > 4 cm; fruit ob-
long to short cylindrical, > 5 cm long; N Mozambique, C and S Tanzania or
W Ethiopian to SE South Sudanian mts..................................C. schliebenii
33* Lower leaf surface glabrous or sparsely covered with trichomes. If densely
covered with trichomes, then bracts < 3 mm and owers smaller.................34
34 Plant with male owers .............................................................................35
35 Plant glabrous, rarely some trichomes on adaxial petiole and leaf margin. Lower
leaf lamina with pale (rarely also black when oxidized) glands towards the base,
sometimes also between secondary veins. Margin of mature leaves with claret-
red or brownish (black when dry) teeth, when young pale. Flowers solitary,
rarely clustered, calyx lobes spreading to reexed, corolla white or bu (Fig. 7a).
NW (Senegal) to NE and E Africa, southern Arabia, S and SE Asia, naturalized
or likely to become so in (sub)tropical regions worldwide ............... C.grandis
35* Plant with trichomes, or if glabrous then dierent from above .................. 36
36 Plant glabrous or rarely with soft multicellular trichomes. Flowers in lax ebracteate
racemes, calyx lobes lineal, > 2 mm long, in buds spreading, when mature reexed
(Fig. 32). Rainforests of W Africa (W of the Dahomey Gap) .............. C.keayana
36* Plant not as above ..................................................................................... 37
37 Plant glabrous. Leaves cordate to subhastate, rarely 3-lobate. Flowers in
ebracteate racemes. Calyx lobes erect, at base broader than 0.75 mm, corolla
urceolate. Rainforests of W Africa ........................................... C.longicarpa
37* Plant dierent and not from W Africa, or if from W Africa then calyx lobes
narrower or spreading to reexed .............................................................. 38
38 Plant glabrous, at maturity often with white speckles on stem, petiole, and
lower leaf lamina. Flowers in racemes or 1 solitary. Calyx lobes erect with
Monograph of Coccinia (Cucurbitaceae) 47
recurved tips. Corolla campanulate. Semi-humid savannas and woodlands of
W Africa ................................................................................ C.intermedia
38* Plant not from W Africa or if so, then rainforest species (sometimes hard to
distinguish from C.intermedia), or lower leaf surface conspicuously covered
with trichomes .......................................................................................... 39
39 Plant glabrous (or puberulous), leaves usually coriaceous. W or C (or western
E) African rainforests ................................................................................40
40 Male owers in racemes with common peduncle shorter than racemose part.
Lowland rainforests or rainforest relicts in higher altitudes or along rivers.....
..................................................................................................... C.barteri
40* Male owers in racemes with common peduncle longer than racemose part.
Mountain forests from Kivu Mts, Eastern Arc Mts, Livingstone Mts, also
introduced in Kenyan high mts ............................................C.mildbraedii
39* Plant conspicuously covered with trichomes or if glabrous, then leaves papery
or from NE, E, or S Africa ........................................................................ 41
41 Plant from S Africa (except C and N Mozambique) ..................................42
42 Plant (esp. stem oder petioles) with or without white speckles, perianth tube/
hypanthium with long (> 0.7 mm) trichomes or if glabrous then calyx lobes
> 2 mm (Fig. 10a) ...................................................................C.rehmannii
42* Plant without white speckles, perianth tube/hypanthium with short (< 0.7
mm) trichomes or if glabrous then calyx lobes < 2 mm (Fig. 21)...................
............................................................................. C.adoensis var. adoensis
41* Plant from E (incl. C and N Mozambique), NE, or NC Africa.................43
43 Flowers clustered, common peduncle < 1 cm, if ower solitary then pedicel
usually < 1 cm. NE Africa (incl. N Tanzania) ...........................................44
44 Upper and lower leaf surface rather densely covered with multicellular trichomes.
Plant usually from higher elevations of N Tanzania and Kenya ........C.trilobata
44* Upper leaf surface pustulate but without trichomes, or with minute trichomes
from pustules. Plant rather from dry habitats and lower elevations. May be
hard to distinguish .................................................................................... 45
45 Leaf margin in mature leaves with conspicuously colored teeth. Plant densely cov-
ered with long trichomes that appear articulate when dry (Fig. 9) .........................
..........................................................................................................C.megarrhiza
45* Leaf margin in mature leaves without conspicuously colored teeth. Plant less
densely covered with trichomes or if densely, then trichomes minute (< 0.2 mm)
or if longer then not appearing articulate when dry (Fig. 2a) ..... C.microphylla
43* Flowers in racemes with peduncle > 1 cm (if smaller then from C Tanzania),
or if solitary then either pedicel >1cm or plant from C Tanzania ............ 46
46 Calyx lobes subulate to narrowly triangulate with pointed tip, > 2.5 mm.
Petiole and lower leaf surface not puberulous. Plant from E Africa (Tanza-
nia, Mozambique, Malawi) ..................................................................... 47
47 Leaves 3-lobate, distinctly petiolate, often with few short trichomes on the
main nerves of the lower leaf surface. Racemes with > 8 owers. Coastal for-
ests of Kenya or NE Tanzania (Fig. 23) ..................................C.pwaniensis
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
48
47* Leaves subcordate to 3- or 5-lobate, subsessile or distinctly petiolate (Fig. 6).
Lower leaf surface glabrous or nerves with short (wart-like) to long trichomes.
Racemes < 10 owers (Fig. 39). SE Tanzania, C and N Mozambique, or Ma-
lawi (Fig. 23) ...............................................................................C.senensis
46* Calyx lobes < 2.5 mm or if longer, then not pointed (may be lineal though)
or petiole and lower leaf lamina puberulous, or plant from NE Africa (Kenya,
Ethiopia, Somalia) ................................................................................... 48
48 Plant with long (> 0.5 mm) trichomes or with short, narrowly conical tri-
chomes, calyx lobes > 2 mm, lineal. NE Africa ......................................... 49
49 Apex of the cordate leaf or central lobe tapering into a long, acute tip. Male
owers solitary or in racemes with a long common peduncle. Plant from high
elevations (Fig. 20).................................................................. C.abyssinica
49* Apex of leaf or central lobe retuse, obtuse, or rather abruptly tapering into a
short acute tip (Fig. 9). Male owers solitary, clustered or if in racemes, then
peduncle short. Plant rather from lower elevations (Fig. 20) ...C.megarrhiza
48* Plant glabrous or with short trichomes (< 0.8 mm), if with longer trichomes
then not from NE Africa. Calyx lobes < 2.5 (–3.5) mm long. Taxa in E Africa
not easily distinguishable (complex around C.adoensis) ........................... 50
50 Plant with long (> 0.8 mm) trichomes (Figs 3a, 5c), calyx lobes 1.5–3.5 mm
long but not with pointed tip. Malawi, C, and S Tanzania, maybe also N
Mozambique (Fig. 23) ....................................... C.adoensis var. jereyana
50* Plant glabrous or with short (< 0.8 mm) trichomes .................................. 51
51 Lower leaf surface and usually also herbaceous stems, petioles, and upper leaf
surface densely covered with short (< 0.5 mm) trichomes. Peduncle often
shorter than pedicelled part. Calyx lobes < 2 mm. Corolla orange, rarely yel-
low? E Africa (C Tanzania; Fig. 23) ................. C.adoensis var. aurantiaca
51* Lower leaf surface glabrous to densely covered with trichomes, but if so then
peduncle longer than pedicelled part. Calyx lobes usually < 2 mm. E, NE, or
NC Africa (Fig. 21) ..............................................C.adoensis var. adoensis
34* Plant with female owers, fruits or vegetative ........................................... 52
52 Plant with female owers ......................................................................... 53
53 Flowers solitary or in ebracteate racemes. Calyx lobes lineal, spreading in
buds, reexed in mature owers, > 2 mm long. W African rainforests (W of
Dahomey Gap) (Fig. 32) ........................................................... C.keayana
53* Plant not as above. If with spreading to reexed calyx lobes, then not from
rainforest regions or < 2 mm long ............................................................ 54
54 Ovary cylindrical. Calyx lobes broader than 0.75 mm at base, corolla urceo-
late. W African rainforests .......................................................C.longicarpa
54* Ovary shortly elliptical, (ob-)ovoid or globose, if cylindrical then not from W
African rainforests. Calyx lobes narrower at base ...................................... 55
55 Flowers in bracteate or ebracteate racemes or solitary. Ovary globose or (ob-)
ovoid, if longer then from E Africa. Calyx lobes < 2 mm long. W and C
Africa but also in rainforest relicts or mountain forests in E Africa ........... 56
Monograph of Coccinia (Cucurbitaceae) 49
56 Female owers solitary or in racemes. Corolla cup-, urn- or funnel-shaped,
not open campanulate. Ovary globose to (ob-)ovoid. W or C (or western E)
Africa. Lowland or in relict rainforests in highlands ...................... C.barteri
56* Female owers solitary. Corolla cup-shaped to campanulate. Ovary long
spindle-shaped to oblong. Mountain forests of Kivu Mts, Eastern Arc Mts,
Livingstone Mts, also introduced in Kenyan high mts ..........C.mildbraedii
55* Flowers solitary. Calyx lobes > 2 mm or if shorter then plant not from rain-
forests from regions as above ..................................................................... 57
57 Calyx lobes spreading to reexed, lower leaf surface glabrous with pale glands be-
tween main veins. Leaf margin with colored teeth (Fig. 7a). Calyx lobes lineal, in
buds spreading, later reexed. Corolla white or bu. Plant from NW (Senegal) to
NE and E Africa, southern Arabia, S and SE Asia, naturalized or likely to become so
in (sub)tropical regions worldwide but natively not from C or S Africa.... C.grandis
57* Plant with trichomes or if glabrous, then with darkish glands or without
glands on lower leaf surface. Calyx lobes not reexed. Corolla in various
colors but not snow-white .........................................................................58
58 Calyx lobes erect with recurved tips, lower leaf lamina with dark glands between
veins, sometimes with white pustules on veins and petiole. Margin of mature leaves
with colored teeth. Plant from woodlands or savannas of W Africa .... C.intermedia
58* Plant not as above and not from W Africa or if so, then not with white pus-
tules and dark teeth...................................................................................59
59 Calyx lobes subulate to narrowly triangulate with pointed tip, > 2.5 mm
(Fig. 39); petiole and lower leaf surface not puberulous. E Africa (Tanzania,
Mozambique, Malawi; Fig. 23) ................................................................. 60
60 Leaves 3-lobate, distinctly petiolate, often with few short trichomes on the
main nerves of the lower lamina. Coastal forests of Kenya or NE Tanzania ..
...............................................................................................C.pwaniensis
60* Leaves cordate to 3- or 5-lobate, subsessile or distinctly petiolate. Lower leaf
lamina glabrous or nerves with short (wart-like) to long articulate trichomes.
SE Tanzania, C and N Mozambique, or Malawi .........................C.senensis
59* Calyx lobes < 2.5 mm or if longer, then not with pointed tip or then petiole
and lower leaf lamina puberulous, or plant from NE Africa (Kenya, Ethiopia,
Somalia) ........................................................................................................61
61 Plant from S Africa (S Angola, Zimbabwe, C Mozambique and further S) ... 62
62 Plant (esp. stem and petioles) with or without white speckles, perianth tube/
hypanthium with long (> 0.7 mm) trichomes or if glabrous then calyx lobes
> 2 mm (Fig. 10a) ...................................................................C.rehmannii
62* Plant without white speckles, perianth tube/hypanthium with short (< 0.7
mm) trichomes or if glabrous then calyx lobes < 2 mm (Fig. 21)...................
............................................................................. C.adoensis var. adoensis
61* Plant from NC, NE or E Africa ................................................................ 63
63 Plant with long (> 0.5 mm) trichomes or straight, narrowly conical trichomes
or if trichomes short (< 0.2 mm) then ovary globose. Plant from NE Africa
(Ethiopia, Kenya, Somalia, N Tanzania) ................................................... 64
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
50
64 Apex of leaf or central lobe tapering into a long, acute tip. Plant from high
(>800 m) elevations of Ethiopia (Fig. 20) ...............................C.abyssinica
64* Apex of leaf or central lobe retuse, obtuse, or rather abruptly tapering into a
short, acute tip. Plant rather of dry habitats in lower elevation .................. 65
65 Plant rather densely covered with long (> 0.5 mm) trichomes that appear articu-
late when dry. Leaf apex retuse, obtuse, or rather abruptly tapering into a short
acute tip (Fig. 9). Leaf margin of mature leaves with dark glands. Ovary ellipsoid,
never globose. N Kenya, Ethiopia and likely also Somalia ...........C.megarrhiza
65* Plant rather laxly covered with trichomes, if denser then trichomes usually min-
ute (< 0.2mm), if longer then not appearing articulate when dry. Leaf apex rarely
obtuse (e.g., around the Usambaras), often abruptly tapering into a short acute
tip. Leaf margin never with dark glands. Ovary globose, rarely (ob-)ovoid. N
Tanzania, Kenya, Ethiopia and likely also Somalia (Fig. 2a) .......C.microphylla
63* Plant glabrous or with short (< 0.5 mm) trichomes, if with longer trichomes
then not from NE Africa, ovary not globose .............................................66
66 Leaves on upper lamina with short trichomes (Fig. 8b). Calyx lobes 2–5 mm
long. N Tanzania or Kenya ........................................................ C.trilobata
66* Leaves on upper lamina glabrous (but with white pustules) or if with short
trichomes, then from C Tanzania. Calyx lobes < 2 mm or if longer, then
lower leaf lamina with long (> 0.8 mm) trichomes or puberulous. Taxa in E
Africa not easily distinguishable (complex around C.adoensis) .................. 67
67 Plant with long (> 0.8 mm) trichomes that appear articulate when dry
(Figs3a, 5c), calyx lobes 1.5–3 mm long but not pointed (as in Fig. 39). C
and S Tanzania, Malawi, maybe also N Mozambique (Fig. 23) ....................
.......................................................................... C.adoensis var. jereyana
67* Plant glabrous or with short (< 0.8 mm) trichomes only ........................... 68
68 Lower sometimes also upper leaf lamina densely covered with short tri-
chomes. Ovary densely covered with short (< 0.5 mm) trichomes. Calyx lobes
< 2 mm. Corolla orange, rarely yellow? E Africa (C Tanzania; Fig. 23) .........
.........................................................................C.adoensis var. aurantiaca
68* Lower leaf lamina glabrous or covered with short trichomes (Fig. 21). Ovary gla-
brous or only laxly (inconspicuously) covered with short trichomes. Corolla yellow-
ish, pinkish or orange. E, NE, or NC Africa (Fig. 22) .... C.adoensis var. adoensis
52* Plant with fruits only or vegetative ............................................................ 69
69 Plant with fruits ........................................................................................ 70
70 Fruit long elliptical to cylindrical (> 8 cm). Forest species ......................... 71
71 Plant from West African rainforests ........................................ C.longicarpa
71* Plant from mountain forests of Kivu Mts, Eastern Arc Mts, Livingstone Mts,
also introduced in Kenyan high mts ......................................C.mildbraedii
70* Fruit globose to oblong (< 8 cm) ..............................................................72
72 Fruit globose. Plant from dry habitats ....................................................... 73
73 Plant from N Tanzania, Kenya, S and SE Ethiopia or Somalia ... C.microphylla
73* Plant from S Africa ................................................................. C.rehmannii
Monograph of Coccinia (Cucurbitaceae) 51
72* Fruit obovoid to oblong, if (sub-)globose, then from humid habitats ........ 74
74 Leaf margin at maturity with colored teeth (blackening when dry), lower leaf
surface glabrous and with pale glands between main veins, petioles and veins
at maturity often with white pustules. Fruit (ob-)ovoid to elliptical. Plant
natively not from C or S Africa .................................................... C.grandis
74* Lower leaf surface without glands or with darkish glands or if with pale
glands, then mature leaves without colored teeth on leaf margin ............... 75
75 Fruit subglobose to obovoid-elliptical, in raceme or if solitary, then rainforest
species ....................................................................................................... 76
76 Fruit in ebracteate raceme or solitary. Plant from W Africa (W of Dahomey Gap)
(hardly distinguishable in shared distribution range) ..... C.keayana or C.barteri
76* Fruit in bracteate or ebracteate raceme. Plant from W and C Africa and in
relict rainforest patches along the Western Rift ............................. C.barteri
75* Fruit solitary or 1–3 clustered but not in raceme. Plant not from rainforests . 77
77 Plant glabrous, at maturity often with white speckles on stem, petiole, and
lower leaf lamina. Fruit subglobose to obovoid-elliptical, solitary or 1–3 clus-
tered. Semi-humid savannas and woodlands of W Africa ........ C.intermedia
77* Plant dierent and not from W Africa. If from (eastern) W Africa then fruit
often with sterile apex (“beak”) .................................................................78
78 Fruit elliptical to oblong, often with sterile apical tip (“beak”). Unripe with
dark green/light green longitudinal stripes or mottling. Seeds rather lenticular
and with symmetrical shape (Fig. 14a). Lower leaf lamina glabrous or with
trichomes, if trichomes appearing articulate and > 0.5 mm, then from C and
S Tanzania, maybe also Malawi or N Mozambique ..................................79
79 Leaves 3-lobate. Leaf surface glabrous but veins and petiole often with
few short trichomes. Plant from coastal forests of SE Kenya to E Tanzania
(Fig.23) .................................................................................C.pwaniensis
79* Plant dierent or from dierent region (hardly distinguishable) ...............80
80 Plant with long (> 0.8 mm) trichomes (Figs 3a, 5c) or if reduced, then sti but
not bent (as in Fig. 21), warty or subglabrous. C and S Tanzania, Malawi or C
to N Mozambique (Fig. 23) ...........C.senensis or C.adoensis var. jereyana
80* Plant glabrous or with short (< 0.8 mm), but not warty or articulate appear-
ing trichomes ............................................................................................ 81
81 Stem, petiole, lower leaf lamina, and ovary/young fruit densely covered with
short trichomes. C Tanzania (Fig. 23) ..............C.adoensis var. aurantiaca
81* Stem, petiole, lower leaf lamina glabrous or with short trichomes, but young fruit only
with lax indumentum. S, E, NE, or NC Africa (Fig. 22) ... C.adoensis var. adoensis
78* Fruit obovoid, shortly to long elliptical, but not oblong and not with conspicu-
ous sterile apical tip (“beak”). Unripe fruits with whitish longitudinal mottling
that often has a dark green halo. Seed face rather at, shape often asymmetrical
(Fig. 14b, c). S Africa or NE Africa, incl. Kenya and N Tanzania ................ 82
82 Plant from S Africa ................................................................. C.rehmannii
82* Plant from NE Africa, incl. Kenya and N Tanzania .................................. 83
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
52
83 Upper leaf lamina with ne, short trichomes. Lower leaf lamina with rather
shortly (< 0.8 mm) articulate (Fig. 8b) or with narrowly conical trichomes.
Plant from higher elevations of N Tanzania and S to C Kenya ... C.trilobata
83* Upper leaf lamina glabrous (but with pustules), rarely with narrowly conical
trichomes. Lower leaf lamina with often long (> 0.8 mm), articulate or with
narrowly conical trichomes .......................................................................84
84 Apex of leaf or central lobe tapering into a long acute tip. Plant from high
elevations (> 900 m) (Fig. 20) ..................................................C.abyssinica
84* Apex of leaf or central lobe retuse, obtuse, or rather abruptly tapering into a
shortly acute tip (Fig. 9). Plant rather from low elevations ........................85
85 Plant rather densely covered with trichomes that appear articulate when dry.
Leaf apex retuse, obtuse, or rather abruptly tapering into a shortly acute tip
(Fig. 9). Leaf margin of mature leaves with dark teeth. N Kenya, Ethiopia and
likely also Somalia (Fig. 20) ...................................................C.megarrhiza
85* Plant rather laxly covered with trichomes or if densely, then trichomes minute (<
0.5 mm). Leaf apex, rarely obtuse (e.g., around the Usambaras), often abruptly
tapering into a short acute tip. Leaf margin never with dark teeth. Fruit globose
to elliptical. N Tanzania, Kenya, Ethiopia and likely also Somalia (Figs 2a, 33) ...
.......................................................................................................C.microphylla
69* Plants with vegetative characters only .......................................................86
86 Plant glabrous, lower leaf surface with pale glands (if strongly oxidized then
also dark, but then leaf margin also with black markings/teeth) between veins,
veins at maturity often with white speckles. Leaf margin at maturity with color-
ed teeth (Fig. 7a). Plant from W to NE Africa, incl. Kenya and N Tanzania or
from outside of Africa ...................................................................... C.grandis
86* Plant with trichomes or if glabrous, then with darkish glands between veins
or without glands on lower leaf surface .....................................................87
87 Plant from W Africa ................................................................................. 88
88 Leaves cordate to 5-lobate, rarely broader than 10 cm, rather papery, lobes
triangulate to narrowly lanceolate or oblong. Lower leaf surface without white
speckles, glabrous or with often short, bent trichomes (Fig. 21). Semi-humid
savannas and woodlands of N Cameroon, N Nigeria, distribution in the area
imperfectly known ................................................ C.adoensis var. adoensis
88* Leaves cordate to subhastate to 3- or 5-lobate, mature often > 10 cm wide.
Leaf lobes triangulate to broad lanceolate, but not narrowly lanceolate or ob-
long. Rainforest species or if from semi-humid savannas or woodlands (rarely
dry forests), then margin of mature leaves with conspicuously colored teeth
and lower leaf lamina often with white speckles ........................................ 89
89 Plant glabrous, at maturity often with white speckles on stem, petiole, and
lower leaf surface. Margin of mature leaves with conspicuously colored teeth.
Semi-humid savannas and woodlands of W Africa ................. C.intermedia
89* Plant not as above, from rainforests or gallery forests. Species not condently
distinguishable .............................. C.barteri, C.keayana or C.longicarpa
87* Plant not from W Africa ........................................................................... 90
Monograph of Coccinia (Cucurbitaceae) 53
90 Leaves coriaceous. Plant glabrous or puberulous on abaxial side of petiole.
Rainforest or mountain forest plant from C Africa or western E Africa (along
the Western Rift, Livingstone Mts, Eastern Arc Mts), plants vegetatively hardly
distinguishable ..............................................................................................91
91 Lowland rainforest (in relict sites of western E Africa also in mountains) plant
from C Africa, incl. areas around Kivu Mts, Chimanimani Mts, and forests
(mountain ranges) along the Western Rift, incl. Uganda ..............C.barteri
91* Plant from mountain forests of Kivu Mts, Eastern Arc Mts, Livingstone Mts,
also introduced in Kenyan high mts ......................................C.mildbraedii
90* Plant not from C African rainforests ......................................................... 92
92 Plant from S Africa ................................................................................... 93
93 Lower leaf surface with usually bent trichomes (Fig. 21), never white speckled,
rarely subglabrous ................................................... C.adoensis var. adoensis
93* Lower leaf surface with straight, in herbarium collections often articulate appear-
ing trichomes, often with white speckles towards maturity or glabrous ...............
........................................................................................................C.rehmannii
92* Plant from E, NE, or NC Africa ............................................................... 94
94 Plant from NE or NC Africa ....................................................................95
95 Teeth on leaf margin conspicuously colored. NE Africa ............................96
96 Apex of leaf or central lobe tapering into a long acute tip. Plant from high
elevations (> 900 m; Fig. 20) ...................................................C.abyssinica
96* Apex of leaf or central lobe retuse, obtuse, or rather abruptly tapering into a shortly
acute tip (Fig. 9). Plant rather from lower elevation (< 1200 m) (Fig. 20) .............
........................................................................................................ C.megarrhiza
95* Teeth on leaf margin not conspicuously colored. Plant from NC or NE Africa .. 97
97 Lower leaf surface glabrous or with short (often bent) trichomes, cordate to deep-
ly lobate (Figs 2a, 21). Plant from NC and NE Africa (incl. N Tanzania) ........ 98
97* Lower leaf surface with long (> 0.5 mm) trichomes that appear articulate
when dry (such as in Figs 8b, 9), or with narrowly conical trichomes. Plant
from NE Africa ........................................................................................ 99
98 Stem sometimes pustulate. Leaf shape variable, if lobate then lobes extending and
not pointing forward, lobes not oblong to linear. Leaves usually with trichomes, of-
ten also on upper lamina and then minute (< 0.2 mm). Trichomes on lower lamina
often not only restricted to the veins. (Fig. 2a) Plant rather from low elevation dry-
lands (rarely in higher elevations) of NE Africa (incl. N Tanzania) ...C. microphylla
98* Stem glabrous or with short (< 0.5 mm) trichomes but not pustulate. Leaf
shape variable (cordate to deeply lobate) but also with oblong to linear lobes
that point forward. Leaves glabrous or with (often bent) trichomes, on upper
leaf lamina rarely beset with trichomes in this distribution area. Trichomes on
lower leaf lamina usually restricted to the veins (Fig. 21). Plant from wood-
lands and semi-humid habitats ..............................C. adoensis var. adoensis
99 Leaves cordate to lobate but not lobulate. If profoundly lobate, then central
lobe lanceolate or ovate tapering into an acute tip. Lower leaf surface with
long (> 0.5 mm) trichomes that appear articulate when dry, or with narrow
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
54
conical trichomes. Plant from higher elevations (> 900 m) or cultivated, from
Ethiopia (Fig. 20) ....................................................................C. abyssinica
99* Leaf reniform to lobate, rarely lobulate. Apex of leaf or central lobe retuse, ob-
tuse, rather abruptly tapering into a shortly acute tip (Fig. 9), if longer tapering
into an acute tip then lobes lobulate. Lower leaf surface with long (> 0.5 mm)
trichomes that appear articulate when dry, only rarely with conical trichomes.
Plant rather from lower elevations (< 1200 m) (Fig. 33) ........... C.microphylla
94* Plant from E Africa .................................................................................100
100 Leaves 3- or 5-lobate. Lobes extending, not pointing towards apex, broadly
triangulate, elliptical, ovate or somewhat angulate but not narrow, oblong, or
lineal. Upper and lower leaf surface with short, white trichomes that appear
articulate when dry (Fig. 8b). Plant from higher elevations of N Tanzania and
Kenyan highlands (Fig. 33) ........................................................ C.trilobata
100* Upper leaf surface glabrous (but pustulate) or if with trichomes then leaf
shape dierent or from dierent region...................................................101
101 Stem sometimes pustulate. Leaf shape variable, if lobate then lobes extending and
not pointing forward, lobes not oblong to linear. Leaves usually with trichomes, of-
ten also on upper lamina and then minute (< 0.2 mm). Trichomes on lower lamina
often not only restricted to the veins. (Fig. 2a) Plant rather from low elevation dry-
lands (rarely in higher elevations) of E Africa (Kenya, N Tanzania) ...C. microphylla
101* Stem not pustulate, leaves with or without oblong to elliptical lobes. If upper
lamina densely covered with minute trichomes then from C Tanzanian wood-
lands. If plant with long articulate appearing trichomes then from highlands of
C to S Tanzania, Malawi or N Mozambique................................................ 102
102 Leaves 3-lobate (rather small auriculate), upper surface glabrous (but pustulate),
lower lamina glabrous, but often with short trichomes on main veins. Coastal
forests of SE Kenya and NE to E Tanzania (Fig. 23) ...................C.pwaniensis
102* Plant not as above or from dierent area (in some cases hard to distinguish) ... 103
103 Lower leaf surface with long (> 0.5 mm) trichomes that appear articulate
when dry or reduced to warts, rarely almost glabrous; sometimes leaves sub-
sessile (Fig. 23) ............................C.senensis or C.adoensis var. jereyana
103* Lower leaf surface glabrous or with short, thin, straight or bent trichomes .... 104
104 Lower leaf surface (also often upper lamina), petiole and stem rather densely
covered with short trichomes. C Tanzania ........C.adoensis var. aurantiaca
104* Lower leaf lamina glabrous or covered with trichomes, if densely then up-
per lamina glabrous (but pustulate) or with few straight trichomes but not
tomentose. Widespread in E Africa ....................... C.adoensis var. adoensis
Taxonomic treatment
Herbarium abbreviations follow Index Herbariorum (http://sciweb.nybg.org/science2/
IndexHerbariorum.asp). Digital collections were accessed from the homepages of the
corresponding herbaria, except for “JPS” (= JStor Plant Science; http://plants.jstor.org/)
and “CVH” (= Chinese Virtual Herbarium; http://www.cvh.org.cn/).
Monograph of Coccinia (Cucurbitaceae) 55
Species concepts in this treatment mainly follow the morphospecies concept but also
include ecological aspects (habitats) and biogeography. Apart from easily recognizable
distinct forms, it was tried to include molecular data (plastid and nuclear; Figs17,18)
from as many forms as possible to check whether they cluster together or not. Accessions in
polytomies are treated as one species as long as they are not morphologically or ecologically
(habitat) distinct or are distantly distributed, if not contra-indicated otherwise (e.g., full
crossing compatibility in Asian and African C.grandis). Names have been synonymized
if no character was found to separate condently the collections from the type material.
Names have been changed in status (in this treatment to varieties), when characters to
separate the collections change in degree, rather than absence/presence.
e minimum leaf size and petiole length were taken from leaves on the same node
as open owers or fruits. Leaf length is measured from the attachment point of the
petiole on the lamina to the apex.
Coccinia Wight & Arn., Prodr. . Ind. orient.: 347. 1834.
Cephalandra Schrad. ex Eckl. & Zeyh., Enum. pl. afric.austral. 2: 280. 1836.
Type species: Bryonia quinqueloba unb.
Physedra Benth. & Hook.f., Gen pl. 1(3): 827. 1867. Indirectly lectotypied by Jerey
(1962: 361)
Type species: Physedra heterophylla Hook.f.
Staphylosyce Benth. & Hook.f., Gen pl. 1(3): 828. 1867.
Type species: Staphylosyce barteri Hook.f.
Type species. see Bryonia grandis L.
Description. Dioecious. Perennial climbers or creepers. Stems up to 20 m, glabrous or
covered with simple smutty-white to yellowish trichomes. Leaves alternate, simple, paired
with a tendril. Leaves sessile (C.sessilifolia var. sessilifolia), subsessile to distinctly petiolate.
Petioles up to 16.5 cm. Petioles glabrous or covered with simple trichomes. Leaves 0.7–20
× 1.1–23 cm, reniform, cordate to deeply palmately 3- to 7-lobate, sometimes lobulate.
Lobes triangulate, ovate, elliptical to linear. Margin entire to more or less densely serrate,
dentate. Teeth inconspicuous or colored. Leaf apex obtuse, acute to acuminate. Upper leaf
surface with clear or whitish pustules, sometimes with trichomes emerging from the lamina
or from pustules. Nerves glabrous or with simple trichomes. Lower leaf surface paler than
upper surface, glabrous or with simple trichomes. Probracts caducous or persistent, ovate,
up to 4.5 mm long. Lower surface keeled or bulging outwards, often with extranuptial
glands. Tendrils simple or unequally bid. Flowers and inorescences emerging from leaf
axils. Male owers solitary, fascicled or in up to 20-owered racemes. If solitary owers
and racemes are developed, then solitary owers occurring before the racemes (within
the plant and per node). Common peduncle of raceme 0.5–10 cm, pedicel of owers
in racemes 0.3–1.8 cm, glabrous or with indumentum as on stem but often less dense.
Bracts ovate, up to 4 mm long or missing. Pedicel of solitary owers 0.2–8.5 cm, glabrous
or with simple trichomes. Perianth tube glabrous or more or less densely covered with
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
56
trichomes. Calyx connate, campanulate, rarely cupulate or urceolate, glabrous, puberulous
or with long, simple trichomes. Calyx lobes 0.5–15 mm, triangulate, lineal or subulate;
reexed, spreading to erect. Corolla connate, campanulate, urn-shaped or tubular, 0.7–6.2
cm long; white, dull yellow to orange, salmon; lobes 0.3–4.7 cm, inside densely covered
with multicellular trichomes, of which some end with a glandular endcell. Filament
column (greenish-)white or orange, anther head pale yellowish green to orange, pollen sacs
S-shaped. Female owers solitary, in pairs or in racemes. Common peduncle 0.3–2.1 cm,
glabrous or puberulous. Pedicel of owers in racemes 0.3–1 cm, glabrous or with simple
trichomes, pedicels of solitary owers 0.7–5 cm, glabrous or puberulous. Calyx and corolla
as in males but with hypogynous ovary. Calyx in few cases urn-shaped. Style columnar,
greenish yellow, yellow, or orange. Stigmas bulging or 2-lobed, greenish yellow, yellow, or
orange. Staminodes 3, attached to the perianth, white (also yellowish or orange?), anthers
reduced. Ovary glabrous or with simple, short to long trichomes that then appear articulate
when dry. Fruits 1.8–30 × 1.4–5 cm, globose, ovoid, elliptical, or cylindrical; glabrous
or with sparse trichomes. Unripe fruits glaucous green to green, sometimes with white,
white-and-green or rarely green longitudinal mottling. Ripe fruits orange-red to scarlet
red; unicolored or rarely with white to yellowish longitudinal mottling. Seeds enclosed
in a hyaline hull, 4.5–7 × 2–3.5 × 1–1.5 mm (L/W/H), symmetrically or asymmetrically
obovate, apex round, base narrowed, obtuse, round or square-edged. Face at to lenticular.
Seed surface, depending on the extraction mode, rugulose or lamentose.
1. Coccinia abyssinica (Lam.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 536. 1881.
Bryonia abyssinica Lam., Encycl. 1(2): 497. 1785.
Type: Cultivated. Unknown, from seeds sent by Bruce (Jerey 1962) from Ethiopia,
cultivated in Paris Royal Botanical Garden, male, , Anon. in herb. J.-B. Lamarck
s.n. (Holotype: P-LAM! [P00307815, digital image: P-LAM]).
Bryonia macrophylla Ser. in DC.,Prodr. 3: 308. 1828.
Type: without location [probably Ethiopia]. Male and female, , 1815, Anon. in coll.
[E.]ibaud s.n. (Holotype: G-DC!).
Cucumis striatus A.Rich., Tent. Fl. Abyss. 1: 295. 1847.
Type: Ethiopia. [Tigray]: Mt Sholada near Adwa, fr, [Aug], R. Quartin-Dillon s.n.
(Lectotype, designated here: P! [from “Sholada”]).
Type: Ethiopia. [Tigray]: [Mt] Selleuda [Mt Sholada near Adwa], fr, R. Quartin-Dillon
s.n. (Syntype: P! [P05621224, digital image: P]).
Cucurbita exanthematica Fenzl ex A.Rich., Tent. Fl. Abyss. 1: 296. 1847.
Type: Ethiopia. Without detailed location, female, , G.H.W. Schimper 1418 (Lec-
totype, designated here: W! [digital image: WU]; isolectotypes: BM!, G!, P!
[P05621261, digital image: P], TUB! [TUB004727], non TUB-004726!).
Coccinia diversifolia Naudin ex C.Huber, Cat. Print. 1864: 6. 1864. Cephalandra
diversifolia (Naudin ex C.Huber) Naudin, Ann. Sci. Nat. Bot. ser. 5, 5: 19. 1866.
Type: Cultivated. From seeds sent by Schimper from Ethiopia, cultivated in Paris Bo-
tanical Garden and Huber’s Garden in Olbia [Hyères, France], C.V. Naudin s.n.
Monograph of Coccinia (Cucurbitaceae) 57
(Lectotype, designated by Jerey (1962: 347): P! [P06745719, digital image: P];
isolectotypes: P [P06745720, digital image: P], G-DC (2)!, K).
Coccinia diversifolia var. glabrescens Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 537. 1881.
Type: Ethiopia. Chaqou-Choada, 2000 m, in thicket, male, , 21 Jul 1852, G.H.W. Schimper
250 (Lectotype, designated here: P! [sheet with descriptive text]; isolectotypes: P (2)!).
Description. Perennial climber. Stems up to 5 m, covered with more or less dense, ar-
ticulate, dirty-white to yellowish trichomes, rarely glabrous. Petioles 1.5–14 cm, at least
on nerves more or less densely covered with articulate trichomes, rarely glabrous. Leaves
7.5–12 × 6.5–12 cm, often cordate to profoundly 3- or 5-lobate. If lobate then central
lobe dominating, over-all shape rather (long) cordate (Fig. 4a). Lobes triangulate, ovate to
elliptical. Margin more or less densely serrate, dentate. Teeth rarely (if so then small) pale
brownish colored in living state or blackish when dried. Leaf apex acute, or if leaf lobate
then central lobe acute to long acuminate. Upper leaf surface with clear or whitish pus-
tules, sometimes with some trichomes. Lower leaf surface with soft trichomes articulate
appearing when dry or sparsely with sti narrowly conical trichomes, which can appear
warty when short or broken o. Probracts up to 3 mm long. Tendrils simple. Male ow-
ers solitary or in long-pedicelled few-owered racemes. Pedicel with indumentum as on
stem. Common peduncle of raceme 2.5–10 cm, pedicel of owers in racemes up to 1.5
cm, indumentum as on stem or less dense. Bracts up to 1.7 mm long or missing. Solitary
owers with up to 5 cm long pedicel with trichomes as on stem. Perianth tube more
or less densely covered with articulate trichomes. Calyx lobes 2–4 mm, lineal-subulate,
upright. Corolla c.1.4 cm long, yellow to slightly orange, darker on the lobes, lobes up
to 5 mm. Filament column white, anther head pale yellowish green, pollen sacs yellow.
Female owers solitary. Pedicel up to 3.5 cm long, indumentum as on stem to glabrous.
Style not seen. Stigma shape not seen, yellow. Staminodes not seen. Ovary with long tri-
chomes, often appearing articulate when dry. Hypanthium more or less densely covered
with articulate trichomes, calyx lobes and corolla as in males. Fruits 5.5–6 × 3.5–4 cm,
short elliptical, glabrous, orange-red sometimes with yellow longitudinal mottling. Seeds
5–6 × 3 × 1.5 mm (L/W/H), slightly asymmetrically obovate, face at (Fig. 14b).
Phenology. Flowering time: June–October.
Distribution. Fig. 20. Ethiopia (Amhara, Oromia, Southern Nations, Nationali-
ties and People’s Region, Tigray). Elevation 1300–2800 m. On limestone, sandstone,
black soil, chromic nitisol (Mengesha et al. 2012), loam, on deep to shallow soil. Along
lake shores among Typha sp., in Podocarpus-Celtis forest (clearings) and degraded forms
of these, evergreen shrubs (e.g., Euclea sp.).
Use. Edibility of fruits is disputed and may dier between wild and cultivated
forms (E. Westphal & J.M.C.Westphal-Stevels 1951 and 1953). Tuberous roots boiled
for food (T. Ebba 250), young shoots and leaves are eaten when cooked (Hora 1995).
For details see chapter Use, economic potential, and phytochemistry.
Vernacular names. Dawuro: shushe, ushushe (Hora 1995); Galinya [Oromo]: an-
chote (Getahun 1974b); Kenya [Kaa]: ajjo (Hora 1995); Tigrinya: wouchich (G.H.W.
Schimper 1048); Wollamo [Wolleyta]: ušuše (W. Kuls 681). e Kenya name is not ex-
clusive for C.abyssinica but also used for another crop, Plectranthus edulis (Vatke) Agnew.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
58
Remarks. e occurrence of monoecy has been reported by W.J.J.O. de Wilde et
al. 7805, but the seen specimens contained male owers only. If both sexes are found
on the same individual, this is likely to be a case of leaky dioecy (see also section on
Chromosomes and sex determination).
Taxonomic remarks. e C.abyssinica specimen in the Lamarck herbarium must
be the holotype, since there is only one specimen of Coccinia abyssinica in the herbarium
of Lamarck in Paris and none in the herbarium of Sonnerat, which he has seen, too. e
specimen in the Linnaean herbarium was not annotated with a corresponding name.
Cucurbita exanthematica Fenzl ex A.Rich. is commonly recognized as a synonym
of C. grandis with a K.G.T. Kotschy collection as type. However, the label on the
Figure 20. Distribution map of C. abyssinica (pale yellow dots; based on 23 collections) and C. megarrhiza
(blue dots; based on 28 collections). For Ethiopia the borders of the regions are given.
Monograph of Coccinia (Cucurbitaceae) 59
Kotschy 308 specimens merely state the species name, the locality, and “frutices scan-
dens” (= climbing on shrubs; W. Greuter – pers. comm.), which cannot be regarded
as a diagnostic feature. e label is printed and therefore eectively published but not
validly so. Valid publication of that name was eected by Achille Richard (1847), but
he chose a dierent specimen (G.H.W. Schimper 1418), which belongs to C.abyssinica.
e Schimper 1418 specimens bear printed labels on which Fenzl designated a variety
of his invalid name with the phrase “var. foliis superioribus integris (non lobatis)”. e
phrase, however, is also not a validation since the species to which this variety is sup-
posed to belong, is not validly published either (Art. 41.3a and b ICN). Naudin (1859)
suggested that Eduard Fenzl mixed-up some specimens. He accepted the Kotschy 308
specimen as a synonym of his C.schimperi and recognized the similarity of the Schimper
specimen to Lamarck’s Bryonia (Coccinia) abyssinica and Cucumis striatus.
e identity of Cucumis striatus A.Rich. is not obvious. ere are two original speci-
mens with this name in P herbarium: one from Selleuda (P05621224) and the other
one from Sholada, both names for the same mountain near the city of Adwa. e
P05621224 specimen consists of a ripe fruit, a drawing of the fruit, and a tiny frag-
ment of a leaf. Cogniaux identied this specimen as C.adoensis. However, the fruit is
ovoid, which would be unusual for that species in which fruits are long ovoid to short
cylindrical and often have a sterile apex (“beak”). Since there are no seeds, which would
help to clear this problem up easily, the fruit shape is the only usable character. e leaf
fragment might be C.adoensis but it is too small to be certain, and it is loose so it might
also be debris from another specimen. e other original specimen (with a number
“26” from “Sholada”) contains much leaf material and fruits. e fruits are darker than
in the rst type specimen. e indumentum of the lower leaf lamina matches certain
C.abyssinica collections, as does the leaf shape (cf. G. Negri 703, G.H.W. Schimper 250)
although they are not very typical. is specimen is not close to C.adoensis, therefore
the present author chose it to be the lectotype and to synonymize the name Cucumis
striatus with C.abyssinica.
Specimens examined. (Selection, in total: 57) Ethiopia. Amhara: Sanka-
Berr [vicinity of Reb river] and Begemder [highland], G.H.W. Schimper 1446 (E
[E00303229], S [S08-12052], S [S08-12057], W, Z (3)). Oromia: 32 km from Ad-
dis Abeba on road to Debre Zeit [Debre Zeyit], E. Westphal & J.M.C.Westphal-Ste-
vels 1951 (BR [BR0000008914613], EA, MO, PRE, WAG [WAG0225550], WAG
[WAG0225551], WAG [WAG0225552]) & 1953 (MO, WAG [WAG0225546], WAG
[WAG0225547]). SNNPR: Bonga, near Roman Catholic Mission, W.J.J.O. de Wilde &
B.E.E. de Wilde-Duyes 7805 (MO, WAG [WAG0225537], WAG [WAG0225538],
WAG [WAG0225539]). Tigray: 18 km along road from Adu Abun to Axum, 14°09'N,
38°49'E, J.J.F.E. de Wilde 7059 (M, WAG [WAG0225544], WAG [WAG0225545]).
2a. Coccinia adoensis (Hochst. ex A.Rich.) Cogn. var. adoensis
Momordica adoensis Hochst. ex A.Rich., Tent. Fl. Abyss. 1: 293. 1847. Coccinia adoensis
(Hochst. ex A.Rich.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 538. 1881.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
60
Type: Ethiopia. [Tigray]: Adwa, near church “Eta Mariam”, mixed specimens, male
and female, , fr, 5 Jun 1837, G.H.W. Schimper Iter Abyss. sect. 1 no. 166 (Lec-
totype, designated here: P! [P00346261, digital image: JPS, P]; isolectotypes:
BM! [BM001010004], BR [BR0000008351036], BR! [BR0000008351039,
digital image: BR, JPS], BR! [BR0000008886781, digital image: BR, JPS], G!
[G00301597], G! [G00301598], G-DC!, HBG! [HBG506429, digital image:
JPS], K! [K000542642, digital image: JPS, K], K! [K000542643, digital image:
JPS, K], K! [K000542644, digital image: JPS, K], L! [L0057303, digital image: JPS,
L], LG! [LG0000090028250; digital image: JPS], M! [M0105779, digital image:
JPS], M! [M0105780, digital image: JPS], P! [P00346261, digital image: JPS,
P], P! [P00346262, digital image: JPS, P], S! [S10-19076, digital image: S], TUB
[TUB004719, digital image: JPS], TUB [TUB004720, digital image: JPS], W! [W
0011091, digital image: WU], W! [digital image: WU]).
Type: Ethiopia. [Tigray]: near Adwa, in thicket, R. Quartin-Dillon s.n. (Syntype: P!).
Type: Ethiopia. [Amhara?]: Ouodgerate Province, A. Petit no.? (Syntype: P?).
Bryonia convolvuloides A.Rich., Tent. Fl. Abyss. 1: 289. 1847.
Type: Eritrea/Ethiopia. [Gash-barka/Tigray]: Chiré [borderland between both regions,
between Tekezé river and Mareb/Gash river], male, , R. Quartin-Dillon & A. Petit
s.n. (Lectotype, designated here: P! [P05621227, digital image: P]; isolectotype: P!
[P05621226, digital image: P]).
Type: Ethiopia. No location, male, 1844, R. Quartin-Dillon & A. Petit s.n. (Syntype:
P! [P05621255, digital image: P]).
Bryonia jatrophiifolia A.Rich. [sphalm. Bryonia jatrotrophæfolia A.Rich.], Tent. Fl.
Abyss. 1: 289. 1847. Coccinia jatrophiifolia (A.Rich.) Cogn. [sphalm: Coccinia
jatrophæfolia (A.Rich.) Cogn.] in A.DC. & C.DC., Monogr. Phan. 3: 535. 1881.
Type: Ethiopia. [Tigray]: in valley near Adwa, male, , Aug 1839, R. Quartin-Dillon &
A. Petit s.n. (Lectotype, designated here: P! [P00346263, digital image: JPS, P; K
neg. 2990], syntype: P! [P05621222, digital image: P]).
Type: Ethiopia. [Tigray]: near Tchélatchekanné [Djeladjeranné, in Tekezé river valley
(Gillett 1972), on label as “Tchessu Hetchequenné”], male, , R. Quartin-Dillon
s.n. (Syntype: P! [P00346260, digital image: JPS, P; K neg. 4467], P! [P05621223,
digital image: P]).
Cephalandra pubescens Sond. in Harv. & Sond., Fl. Cap. 2: 493. 1862. Coccinia
pubescens (Sond.) Eyles, Trans. Roy. Soc.South Africa 5(4): 498. 1916. Coccinia
pubescens (Sond.) Cogn. ex Harms in Fries, Notizbl. Bot. Gart. Berlin-Dahlem 8:
491. 1923. nom. super.
Type: South Africa. [North West]: Magaliesberg, male and female, , Dec, J. Burke 408
(Lectotype, designated here: K! [K000313229, digital image: JPS, K]; isolectotype:
BM! [BM000815207], K! [K000313227, digital image: JPS, K], NBG, Z!).
Type: South Africa. North West/Gauteng: at Magalies river, male and female, , C.L.P.
Zeyher 588 (Syntype: K! [K000313228, digital image: JPS, K], S! [S08-12187,
digital image: S]).
Coccinia hartmanniana Schweinf., Reliq. Kotschy.: 42, t. 27, t. 28. 1868.
Type: drawing in protologue, t. 27 (Lectotype, designated here).
Monograph of Coccinia (Cucurbitaceae) 61
Type: Sudan. Sinnar Province: no detailed location given, 1860, R. Hartmann s.n.
(Syntype: ?, if B, then destroyed).
Type: South Sudan. At White Nile in the region of the Tschier people [a nilotic tribe
also called: Chir, Kir, Mandari, Mondari, Mundari, Shir], [possibly the area be-
tween Tombe and Mongalla in Central Equatoria state], 1861, W. von Harnier s.n.
(Syntype: B destroyed, ?).
Type: South Sudan. [al-Qadarif]: Gallabat at Gendua [river], Jun 1861, H. Steudner
843 (Syntype: ?, if B, then destroyed).
Type: Ethiopia. [Amhara]: near Matamma at border to Sudan, after beginning of Jun
1865, G.A. Schweinfurth Flora of Gallabat 62 (Syntype:?, if B, then destroyed).
Coccinia rigida Cogn., Bot. Jahrb. Syst. 21: 210. 1895.
Type: Tanzania. [Tabora]: Ugunda, near Gonda [Igonda], on ground in wet corn
elds, R. Böhm 176 (Holotype: B destroyed, lectotype, designated here: BR!
[BR0000008886804, digital image: BR, JPS]).
Coccinia djurensis Schweinf. et Gilg, Bot. Jahrb. Syst. 34: 357. 1904.
Type: South Sudan. [West Bahr al-Ghazal or Warab]: Seriba Ghattas, male, , 24 May
1869 or Jun 1869, G.A. Schweinfurth 1878 (Lectotype, designated here: P!; isolec-
totypes: B [destroyed], E! [E00303230], G!, K! [K000542640, digital image: JPS,
K], PRE [PRE0592944-0, digital image: JPS], S! [S08-12060, digital image: S], Z!
[Z-000073403, digital image: Z]).
Type: South Sudan. [West Bahr al-Ghazal or Warab]: Seriba Ghattas, male, , 24 May
1869, G.A. Schweinfurth 1867 (Syntype: if B, then destroyed).
Type: South Sudan. [West Bahr al-Ghazal]: Djur realm, Seriba Agad Wau [Waw], fr,
May, G.A. Schweinfurth 1688 (Syntype: B [destroyed], K! [K000542641, digital
image: JPS, K], L!).
Coccinia princeae Gilg, Bot. Jahrb. Syst. 34: 358. 1904.
Type: Tanzania. Iringa: Uhehe highland, no detailed location given, fr, M. von Prince
s.n. (Holotype: B, destroyed).
Type: Tanzania. Morogoro: Uluguru Mts, NW slopes, male, , 19 Jan 1933, H.J.E.
Schlieben 3271 (Neotype, designated here: B!; isoneotypes: G!, HBG!, M! [M-
0210964], P! [P05621242, digital image: P], S! [S08-12183]).
Coccinia parvifolia Cogn. in Schinz, Vierteljahresschr. Naturforsch. Ges. Zürich 52:
433. 1907.
Type: South Africa. Limpopo: [Mopani District], [Leydsdorp area], Mt Marovounge [Mt
Marovougne], male, , May 1904, H.A. Junod 2491 (Holotype: Z! [Z-000004444,
digital image: Z], isotype: BR! [BR0000008886811, digital image: BR, JPS]).
Coccinia homblei Cogn., Bull. Jard. Bot. État Bruxelles 5: 114. 1916.
Type: D. R. Congo. Katanga: Lualaba region, Kapanda conuence plains, male, ,
Dec 1912, H. Homblé 992 (Lectotype, designated here: BR! [BR0000008887146,
digital image: BR, JPS]; isolectotype: BR! [BR0000008887474, digital image:
BR, JPS]).
Type: D. R. Congo. Katanga: Kapiri valley, fr, Feb 1919, H. Homblé 1198 (Syntype:
BR! [BR0000008886927, digital image: BR], BR! [BR0000008886859, digital
image: BR, JPS]).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
62
Type: D. R. Congo. Katanga: Kapiri valley, male, , Feb 1919, H. Homblé 1199 (Syntype:
BR! [BR0000008887177, digital image: BR, JPS], BR! [BR0000008887504, digital
image: BR, JPS], BR! [BR0000008886842, digital image: BR, JPS, K neg. 4875]).
Coccinia subspicata Cogn., Bull. Jard. Bot. État Bruxelles 5: 115. 1916.
Type: D. R. Congo. Katanga: Lualaba region, Kapanda conuence plains, male, , Dec
1912, H. Homblé 992a (Holotype: BR! [BR0000008887528, digital image: BR,
JPS, K neg. 4876]).
Coccinia roseiora Suess., Proc.Trans. Rhodes. Sci. Ass. 43: 134. 1951.
Type: Zimbabwe. [Mashonaland East]: Marandellas [Marondera], Cave Tatooma, 31
Nov 1941, G. Dehn 188A (Holotype: M! [M0105778, digital image: JPS]).
Description. Perennial climber or creeper. Stems up to 6 m, glabrous to densely covered
with trichomes. Indumentum whitish to beige. Trichomes < 0.5 mm, sometimes curved
(Fig. 21). Petioles 0.6–3.5 cm, glabrous, pubescent to tomentose with indumentum
like on stem. Leaves 4.2–13.5 × 4.7–16 cm, cordate, shallowly to deeply 3- or
5-lobate, rarely elliptical with hastate base. If profoundly to deeply lobate, then lobes
often (slightly) pointing towards apex (as in C.keayana; Fig. 32). Lobes triangulate,
lanceolate, lineal, elliptical to obovate. Leaf margin entire to serrate (especially on lateral
sides) to lobulate. Tips of lobes retuse, obtuse to acute, often with apical tooth. Upper
leaf surface clear to white pustulate, rarely with whitish trichomes. Lower leaf surface
soft pubescent with short, whitish trichomes, rarely glabrous, often with black to dark
brown glands between nerves towards the base, rarely also along main nerves. Probracts
missing or obovoid up to 2.5 mm long. Tendrils simple, rarely bid. Male owers in
few to many-owered racemes, often accompanied by one solitary ower. Common
peduncle 1–11 cm, glabrous or with short trichomes. Pedicel of racemous ower 0.4–
1.3 cm, indumentum like peduncle, pedicel of solitary ower 1.5–4 cm glabrous or
with short, white trichomes. Bracts missing (caducous?) or up to 2 mm. Perianth tube
glabrous or with short (< 0.5 mm), white trichomes. Calyx lobes 1.2–2.5(–3) mm,
linear to broadly triangular, adpressed to spreading, apex obtuse to acute. Corolla 0.9–
1.6 cm, yellow, salmon-pink, orange, maroon, veins sometimes purplish-brown, lobes
2–5 mm. Filament column whitish to orange, anthers whitish?, yellowish to orange,
pollen sacs yellow to orange. Female owers solitary. Pedicel 0.8–3.5 cm, often glabrous
or with short, white trichomes. Hypanthium glabrous or with short (< 0.5 mm), white
trichomes. Calyx lobes and corolla like in male owers. Ovary glabrous. Style and
stigmas not seen. Fruits 3–7 × 1–1.5 cm, ovoid, oblong to shortly cylindrical, while
ripening (usually?) with dark green longitudinal mottling), ripe orange-red to red, often
with sterile tip (“beaked”). Seeds 4–6 × 3–4 × 1–1.7 mm (L/W/H), symmetrically
obovate, face lenticular (Fig. 14a, 21).
Phenology. Flowering time: January–May, August–December.
Distribution. Fig. 22. Angola? (likely in the South and East since the species
occurs close-by), Botswana (North-West District), Burundi?, Cameroon (Extreme
North), N Central African Republic, S Chad, Democratic Republic of Congo (Ka-
tanga, along the Western Rift), Eritrea (Gash-Barka, likely wider distributed as relicts
Monograph of Coccinia (Cucurbitaceae) 63
in the highlands), Ethiopia (except the dry southeast), Kenya (in the west and central
highlands), Malawi, Mozambique, N Namibia, Nigeria (only known from Adamawa
State, but likely more widely distributed), Rwanda (Eastern Province, maybe wid-
er distributed), South Africa (Gauteng, KwaZulu-Natal, Limpopo, Mpumalanga, E
North West, E Free State), South Sudan, Sudan (West Darfur, maybe also in wood-
land relict sites in other provinces), Swaziland, Tanzania, Uganda (Northern Region),
Zambia, Zimbabwe. Elevation 130–3450 m. On sandy and silty soils, clay loam, loam,
laterite, syenite soils, dolerite soil, dolomite soil, limestone. Hyparrhenia cymbaria sa-
vanna; Crossopteryx tree savanna; Acacia-Combretum-Stereospermum-Cussonia wood-
land; Pseuodoprosopis scheri woodland, Anogeissus leiocarpus woodland, Brachystegia
woodland, sourveld grassland, Melhania rehmannii-Enneapogon scoparius mixed bush-
veld (Siebert et al. 2010).
Use. Roots are boiled and drunk for fever (J.C.Lovett & C.J. Kayombo 3434). e
potato-like tubers are eaten (F.W. Andrews 1310), also raw (T. Scudder 56). e greens
are used as spinach, among others by the Venda (N.J. van Warmelo s.n. Mar 1960,
J.Gerstner 5838) and also eaten by the Luo (Johns and Kokwaro 1991). Ripe fruits are
edible (J.C.Lovett et al. 3842, J. Pawek 11008, T. Scudder 56). According to de Boer et
Figure 21. Male inorescence and leaf of C. adoensis; picture taken from lectotype (G.H.W. Schimper
166 (P00346261)). Note the short bent trichomes, which are a good indicator for this species (but gla-
brous collections or other kinds of trichomes may occur in this species, too).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
64
al. (2005), the Pare people in Tanzania use an infusion of leaves and stems for abortions,
uterus cleansing, and against chickenpox. Coccinia adoensis is quite variable and so there
are likely many varieties, of which some might not be edible. Rehm et al. (1957) reports
the cytotoxic cucurbitacin B and traces of cucurbitacin D in immature fruits, but edible
ripe fruits are known from South Africa. Gradé et al. (2009) report that the usability or
toxicity of the tuberous roots is disputed, suggesting chemical variability.
Figure 22. Distribution map of C. adoensis var. adoensis (based on 311 collections).
Monograph of Coccinia (Cucurbitaceae) 65
Vernacular names. Bokora tribe [Karamojong?]: edaldalakisin (Gradé et al.
2009); Kiluo: mutkuru (Johns and Kokwaro 1991), nyatunduguwoge (J.G.B. New-
bould 5745); Kipare: mlushi (de Boer et al. 2005); Kisafwa: tandandala (J.C.Lovett &
C.J. Kayombo 3434), ndandala (J.C.Lovett & C.J. Kayombo 3773); Kisagara [Kisagala]:
lutetere (F. Haerdi 563/0); Nhungoé [Cinyungwe]: mugwingwi (L. Macuácua 1441);
Sotho [most likely Northern Sotho]: sephu (J. Gerstner 5838); Tigrinya: entatakh
(de G.H.W. Schimper (Schweinfurth 1893)), entota (de A. Richard (Schweinfurth
1893)); Tshivenda: tshiphu (N.J. van Warmelo s.n. Mar 1960); Zande: bawiriokoro
(F.W. Andrews 1601).
Remarks. Coccinia adoensis is widespread and morphologically variable. Some popula-
tions or local forms appear to be distinct, but there are intermediate individuals or similar-
looking collections from dierent parts of the overall distribution range. In East Africa (C
Tanzania, Malawi), one can nd forms linking to C.aurantiaca, which is treated here as
variety of C.adoensis, and to C.senensis. e latter forms have a similar plastid haplotype
with C.senensis, but lack a specic deletion in the trnSGCU–trnGUCC intergenic spacer. ese
forms from central and southern Tanzania are discussed here under the name C.adoensis
var. jereyana, while another form from Kenya remains in var. adoensis. e non-monophy-
ly in the plastid tree (Holstein and Renner 2011b) makes C.adoensis even more peculiar.
e scenario given in the chapter Evolution and phylogeny might explain this pattern, but
without phylogeographic analysis and crossing experiments, this will remain speculative.
Some specimens from the Kingupira area (Lindi, Tanzania; K. Vollesen 3182,
3212, 3384, 4320) have an unusual morphology by having veins that run along the
leaf margin, which is unique in Coccinia. Except for this character, they match Coccin-
ia adoensis var. adoensis well (sympetalous, obovate probracts). ey strongly resemble
Eureiandra species in vegetative characters, but not in generative traits.
Taxonomic remarks. e lectotypication of Momordica adoensis by Meeuse
(1962) is not eected, as he did not specify which specimen was supposed to be the
lectotype. However, the present author follows his suggestion and chose among the
two Schimper 166 specimens from P.
e types of Bryonia jatrophaefolia are not too obvious as such. e protologue states
“Tchélatchekanné”, but Paris Herbarium holds two Quartin-Dillon and Petit specimens
(P00346260 and a non-barcoded one) with a location “Tchessu Heckequenné”. Al-
though the spelling has some similarities, they are quite dierent. On the other hand, the
Ge’ez letter sat [S] and läwe [L] look similar and might have been mistranscribed if the
name was written down in Ge’ez script. Additionally, the two specimens bear the species
name in Richard’s handwriting (C.Bräuchler, pers. comm.). Gillett (1972) notes that
Achille Richard, the author of the botanical treatment of Quartin-Dillon's and Petit's
journey, wrote consistently “Tchelatchekanné”, although Schimper calls it “Djeladjera-
nné”. However, there is a locality with that name [Tchelatchekenneh at c.13°43'N,
38°22'E] in Johnston’s atlas (Johnston 1861), to which Gillett (1972) refers.
e G.A. Schweinfurth 1668 specimen in L is not obviously a type specimen as
it lacks the original label. However, the Herb. D’Alleizette label mentions “Coccinia
djurensis”, and the location is the same as the K duplicate with the original label. Ad-
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
66
ditionally, the specimen is a fruiting female, like the specimen in K. Hence, d’Alleizette
must have obtained a duplicate of the type.
e placement of C.hartmanniana as a synonym of C.adoensis is done with a high
level of condence although no type specimens were found. e protologue contains
drawings showing lenticular seeds and short calyx lobes, which match well other col-
lections of the C.adoensis complex. According to Ascherson (in Schweinfurth 1867),
von Harnier’s collections consisted of two duplicates, one of them in B. Other du-
plicates of von Harnier are in BM and K but not this collection apparently. Since no
type specimen was seen, despite extensive search, a drawing from the protologue was
chosen as lectotype.
For C.princeae, a neotype was selected because the holotype was destroyed. e
leaves of the chosen specimen, H.J.E. Schlieben 3271, match the description well, and
the specimens have been identied as C.princeae when the original material was still
existing. e specimens dier in the generative characters (fruiting in the holotype,
male owers in the neotype), but Gilg referred strongly to the distinctive leaves, so the
neotype appears to be a good match.
e holotype of C.roseiora is the drawing in M, which has the number 188a. e
protologue states: “descriptio sec. tabulam cl. Dehniae” (described following/based
on the illustration of Ms. Dehn). ere are (at least) two specimens with the number
188 (K, SRGH), which partially might have served as basis for the drawing, but they
are not types. e drawing contains all necessary characters to synonymize it with
certainty with C.adoensis: seed shape, fruit maculation and calyx lobe morphology.
Specimens examined. (Selection, in total: 483) Botswana. North-West Dis-
trict: Ngamiland, Motantanyane, H.H. Curson 784 (M). Central African Repub-
lic.Nana-Grébizi: Gribingui (Ft-Crampel) [=Kaga-Bandoro], A.J.B. Chevalier 6325
(P [P05621208]). D. R. Congo. Orientale: Faradje (Kibali-Ituri), J. Lebrun 3406 (BR,
WAG [WAG0225534]). Eritrea. Gash-Barka: Seraé [Seraè, a former province], in Tu-
cul region, A. de Benedictis 519 (FT). Ethiopia. Tigray: Edaga Sciaba, E. Chiovenda 581
(FT). Malawi. Northern Region: Karonga district, 17 mls [27.2 km] N of Chilumba,
J. Pawek 11008 (DSM, MO, PRE, WAG [WAG0234129]). Mozambique. Zambézia:
Massingire [Morrumbala district], M’bôbo [river at Morrumbala], on road to Mopeia,
A.R. Torre 5337 (M). Nigeria. Adamawa: 10 miles [16 km] from Mubi to Toyola,
P. Wit et al. 1797 (BR, MO (2), P [P05621253], WAG [WAG0041392], WAG
[WAG0041393]). Rwanda. Eastern Province: Kibungo prefecture, Rusumo [Ruzu-
mo Falls], savanna park, on opposite slope of A.I.D.R. [Association internationale de
développement rural] camp, J. Lambinon 74/1568 (M, MO, WAG [WAG0225516]).
South Africa. Gauteng: [City of Johannesburg], N of Eikenhof, Johannesburg, Walk-
erville rd., L.E. Davidson 3781 (B, M). Sudan. West Darfur: Zalingei, G.E. Wick-
ens 1800 (K). Tanzania. Lindi: Selous Game Reserve, Kingupira, 8°28'S, 38°33'E,
[38°34'E], K. Vollesen MRC 3384 (DSM, EA, WAG [WAG0234135]). Ruvuma:
Songea, E. Milne-Redhead & P. Taylor 7876 (EA); ibid., E. Milne-Redhead & P. Tay-
lor 7950 (B, EA, K, P [P05621243], S [S08-11819]). Zambia. Lusaka: c.10 km S of
Monograph of Coccinia (Cucurbitaceae) 67
Chilanga and 23 km S of Lusaka, c.1 km N of Kafue Road near Chilanga Cement
housing; Shimabala Cave, 15°39'01"S, 28°14'15"E, D.K. Harder & M.G. Bingham
2584 (K, MO). Zimbabwe. Matabeleland North: Matetsi Safari Area Headquarters
House no. 2, P. Gonde 256 (E, G, MO, P [P05621269]).
2b. Coccinia adoensis var. aurantiaca (C.Jerey) Holstein, stat. nov.
urn:lsid:ipni.org:names:77148917-1
Coccinia aurantiaca C.Jerey, Kew Bull. 17: 169. 1963.
Type: Tanzania. Dodoma: Kondoa District, Great North road, 15 miles S of Kondoa,
1310 m, , fr, 19 Jan 1962, R. Polhill & S. Paulo 1221 (Holotype: K! [K000313236,
digital image: JPS, K], isotypes: B! [B 10 0154922, digital image: B, JPS], B! [B
10 0154923, digital image: B, JPS], BR! [BR0000008887252, digital image: BR,
JPS], EA (2)!, PRE! [PRE0592951-1, digital image: JPS], PRE! [PRE0592951-2,
digital image: JPS]).
Type: Tanzania. Dodoma: 1 ml. [1.6 km] S of Dodoma, Imagi hill, R. Polhill & S. Paulo
1274 (Paratypes: B! [B 10 0154921, digital image: B, JPS], BR!, EA!, K (3)!, P! p.p.,
PRE!, S! [S08-11841, digital image: S]).
Type: Tanzania. Mwanza: Mwanza, Ilemera, Butimba, R.E.S. Tanner 1902 (Paratypes:
BR!, EA!, K (2)!).
Type: Tanzania. Mwanza: Mbarika [chiefdom], Buzomo, R.E.S. Tanner 1068 (Paraty-
pes: BR!, COI (2)!, EA!, K!, NY!).
Type: Tanzania. Mwanza: Mwanza, R.E.S. Tanner 646 (Paratype: K!).
Type: Tanzania. Shinyanga: near Shinyanga, R.D. Bax 57 (Paratypes: K (2)!).
Type: Tanzania. Shinyanga: hills near Shinyanga, B.D. Burtt 2517 (Paratype: K!).
Type: Tanzania. Shinyanga: Shinyanga, H. Koritschoner 1823 (Paratypes: EA (2)!, K!).
Description. Perennial climber. Stems up to 10 m, almost tomentose with short
(< 0.5 mm), sti, whitish trichomes. Petiole 1.5–3.5 cm, indumentum similar to
stems. Leaves 5.2–12.5 × 6.4–14.5 cm, cordate, shallowly to profoundly 3- or 5-lo-
bate. Lobes triangulate, ovate, elliptical to obovate. Margin serrate to lobulate. Apex
obtuse, rarely acute, with nal tooth. Upper leaf surface white-pustulate sometimes
more or less pubescent with short, whitish trichomes. Lower leaf surface usually
densely covered with curved or short, straight trichomes on nerves. Probracts up
to 1.5 mm, often caducous. Tendrils simple. Male owers solitary or in racemes.
Common peduncle 0.4–3 cm, pedicels in racemes up to 0.5 cm, pedicel of solitary
owers 1–2 cm, indumentum as on stem. Bracts up to 1.5 mm, persisting. Perianth
tube densely covered with short (< 0.5 mm) trichomes. Calyx lobes 1–3 mm, nar-
row triangular to dentate, spreading. Corolla 1.6–2.4 cm long, pale yellow-brown
to orange, rarely? yellow, with green to orange venation, lobes 0.6–1 cm. Filament
column, anther head, and pollen sacs more or less pale orange, rarely yellowish? (Fig.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
68
10b). Female owers solitary, pedicel 1.2–4 cm, indumentum as on stems. Hypan-
thium densely covered with short (< 0.5 mm) trichomes, calyx lobes, and corolla as
in male owers. Style shape not seen, green. Stigma shape not seen, yellow to orange.
Ovary with short trichomes. Unripe fruits pale green with irregular lighter spots and
Figure 23. Distribution map of C. adoensis var. aurantiaca (pale yellow triangles; based on 18 collec-
tions), C. adoensis var. jereyana (pale yellow dots; based on 18 collections), C. pwaniensis (blue triangles;
based on 11 collections, including a natural hybrid), and C. senensis (blue dots; based on 30 collections).
For Tanzania the borders of the regions are given.
Monograph of Coccinia (Cucurbitaceae) 69
dark green longitudinal lines. Fruits 5–9 × 1.5–3.5 cm, long ovoid, apex sometimes
beaked, when ripe orange-red. Seeds 6–6.5 × 3.5–4 × 1.5–1.7 mm [L/W/H], slightly
asymmetrically obovate, face atly lenticular.
Phenology. Flowering time: January, March, July, October, December.
Distribution. Fig. 23. Tanzania (Dodoma, Iringa, Manyara, Morogoro, Mwanza,
Shinyanga). Elevation 600–1200 m. Red sandy soil, red clay, granite. White clay. Gray
sand. Brown sandy loam. Dry Miombo woodland, Acacia tanganyikensis-A. tortilis
subsp. spirocarpa-Adansonia digitata-Maerua crassifolia-Balanites aegyptiaca woodland,
long grass savannas, dry (Commiphora-Acacia) bushland, thickets (e.g., Combretum
thickets), among rocks on hills.
Use. Leaves are boiled and eaten (J.L. Newman 62). Fruits edible when ripe and
dry (E.S. Macha 600).
Vernacular names. Sandawe language: koba (J.L. Newman 62).
Remarks. e status of this taxon as species is unclear, therefore it is treated as a
variety of the polymorphic C.adoensis. Coccinia adoensis var. aurantiaca specimens as
listed here are usually more densely covered with trichomes than C.adoensis var. adoen-
sis. Jerey segregated this species from the polymorphic C.adoensis because of the non-
beaked fruits and at seeds with a hyaline girdle. e beak is a sterile part of the ovary
with variable length, but it does not occur in all populations. Two of the paratypes
(R. Polhill & S. Paulo 1274 (BR, P)), which match other C.adoensis var. aurantiaca
collections vegetatively, have a slightly beaked fruit, although most other collections
do not. e seeds are also hardly distinct from C.adoensis, perhaps somewhat larger.
Seeds in Coccinia are enclosed in a hyaline aril. Jerey only observed the dry collapsed
aril, which is not part of the seed, as a “hyaline girdle”. e orange color of the petals,
even with purple venation also occurs in individuals of C.adoensis var. adoensis that
have a less dense indumentum. e corolla is thus not a good distinguishing character
either. However, this variety seems to occur in a drier part of the range of the overall
C.adoensis distribution (Holstein and Renner 2011b).
Taxonomic remarks. e owers in the R. Polhill & S. Paulo 1274 specimen
in P do not belong to Coccinia. e calyx appears to be Momordica foetida Schum.
& onn. e HEID specimen (HEID779579) of that collection is also mistaken,
eventually a mix-up while mounting the specimen. It has a completely dierent indu-
mentum and a narrow, almost cylindrical perianth tube.
Specimens examined. (Selection, in total: 29) Tanzania. Dodoma: Dodoma–
Kondoa road, c.20 km S of Kondoa, 05°16'31.5"S, 35°53'01.1"E, N. Holstein et al.
85 (DSM, M), and 86 (M). Iringa: Iringa Rural District, along road Iringa–Moro-
goro road and Lukosi River, at bottom of Kitonga Gorge, c.6 km W of Mahenge
village at milepost 253 km from Morogoro, 7°38'S, 36°14'E, [7°34'S, 36°19'E], C.M.
Taylor et al. 8485 (K, MO); [Ruaha National Park], Msembi [near aireld], P.J.
Greenway & K. Kanuri 14811 (EA (2), K, M). Manyara: Tarangire National Park,
road Tarangire camp–Babati, 1 ml. from camp, H.M. Richards 24817 (EA, K). Mo-
rogoro: Kilosa district, Elphon’s Pass, 7°22'S, 36°42'E, J.C.Lovett & T.C.E. Congdon
2931 (K, MO).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
70
2c.Coccinia adoensis var. jereyana Holstein, var. nov.
urn:lsid:ipni.org:names:77148914-1
Type: Tanzania. Iringa, Mundi District, Ngwazi, 8°30'S, 35°15'E, 1830 m, female,
, fr, 25 Feb 1987, J.C.Lovett 1597 (Holotype: MO!, isotype: EA!).
Diagnosis. is variety has anities with C.adoensis and C. senensis. e abaxial
side of the petiole and the lower leaf surface bears simple trichomes with long cells,
which appear crumpled or articulate when dry. Most of the trichomes, especially on
the nodes, exceed 0.8 mm (–1.2 mm), whereas trichomes of C.adoensis var. adoensis
and var. aurantiaca are shorter < 0.5(–0.8) mm. e calyx lobe length often exceeds
2mm (in contrast to other C.adoensis varieties), but the lobes are not subulate or
narrowly acute as in C.senensis but rather linear or if narrowly triangulate, then not
with a pointed tip.
Description. Perennial creeper or climber. Stems up to 3 m, more or less densely
covered with long (at least on the nodes > 0.8 mm, Figs 3a, 5c) trichomes that appear
articulate when dry. Petiole 0.25–3.5 cm, subsessile to distinctly petiolate, with long
patent trichomes. Leaves 3.2–10.5 × 2.6–12 cm, shallowly to profoundly 3- or 5-lobate,
lobes triangular, ovate to elliptical, margin dentate, slightly serrate, apex acute to obtuse
with apical tip. Upper leaf surface glabrous or with few trichomes, hyaline to white pus-
tulate. Lower leaf surface more or less densely covered with articulate trichomes, rarely
almost glabrous with white pustules on veins. Probracts up to 3 mm. Tendrils simple.
Male owers in racemes, often accompanied by a single ower or one solitary ower.
Common peduncle 5–5.5 cm, with short articulate trichomes. Pedicel of racemous ow-
ers 5–9 mm, with short articulate trichomes. Bracts up to 1 mm, caducous. Pedicel
of solitary ower 2.2–7.8 cm, with short articulate trichomes. Hypanthium with short
trichomes. Calyx lobes 1–3.5 mm, narrowly triangular but not subulate, erect. Corolla
1.1–1.65 cm, yellow, orange, to dark crimson with darker veins outside, lobes 4–7 mm.
Color of lament column pink, anther head orange-yellow to orange, color of pollen sacs
not seen. Female owers solitary. Pedicel 0.6–1.7 cm long, puberulous. Ovary with short
to long, articulate trichomes. Fruit size c.2–6×c.1 cm long, elliptical, often with sterile
apical tip (“beaked”), glabrous, green with white spots when unripe, red when ripe. Seeds
4–5.5 × 3–3.5 × 1.5 mm (L/W/H), symmetrically obovate, face lenticular (Fig. 14a).
Phenology. Flowering time: January–March, November, December.
Distribution. Fig. 23. Malawi (Northern Region, Southern Region), Tanzania
(Dodoma?, Iringa, Mbeya, Morogoro?, Singida), Kenya (southern Rift Valley Province).
1300–2600 m. Soil preferences unknown. With Dodonea viscosa; under pines; in Euca-
lyptus plantation, highland grassland, in open woodland with Combretum sp., Grewia
sp., Strophanthus emenii, Acacia tortilis, Tapiphyllum obtusifolium, Burttia sp., Cassia sp.
Etymology. e epithet was chosen to honor Charles Jerey, who worked exten-
sively on the Cucurbitaceae and the ora of East Africa.
Use. Unripe and ripe fruits are reported to be edible (C.J. Kayombo 296, P. Kuchar
22631), roots taken to make stomach medicine (P. Kuchar 22631).
Monograph of Coccinia (Cucurbitaceae) 71
Vernacular names. Kihehe: mtumbulansoka (W. Carmichael 171); Kinyaturu:
mukunguhi (P. Kuchar 22631).
Remarks. Morphologically, this variety closely matches C.senensis (with rather short
petiolate to subsessile leaves, and a C.senensis-like indumentum), but it has the calyx
lobes rather of C.adoensis var. adoensis, with the lobe length being intermediate between
C.senensis and C.adoensis var. adoensis. e sequenced specimens do not cluster with
most other C.adoensis haplotypes from East Africa or southern Africa, and lack the typi-
cal deletion of C.senensis in the trnSGCU–trnGUCC intergenic spacer (Holstein and Renner
2011b). A C.adoensis var. adoensis-like collection (S.A. Robertson 1925) also clusters with
this variety, but it lacks the long trichomes. Long trichomes also appear in populations of
C.grandiora or C.mackenii in higher altitudes or in areas with higher precipitation. e
collections of this variety are distributed above 1300 m and thus receive higher amounts
of rainfall, so the long trichomes could be an adaptation. On the other hand, very similar
trichomes regularly occur in C.senensis, sometimes short though, but that species does
not occur in such high altitudes. As the collections of this variety dier from the “typical”
C.adoensis, but still belong to C.adoensis, they are treated as a new variety.
e collection R.E. Gereau & C.J. Kayombo 3582 (K, MO; C.adoensis 4 in Fig.17)
is morphologically inseparable from this variety, and the plastid haplotype clusters within
East African C.adoensis. is collection has a normal-sized corolla, and therefore seems to
be fertile, which supports the hypothesis that the var. jereyana is not reproductively iso-
lated from var. adoensis. is is also why the present author refrains from designating it as
a paratype, namely in order to avoid confusion about the genetic denition of this variety.
Phylogenetically, it is uncertain whether this variety retains an ancestral morphol-
ogy of the common ancestor of C.adoensis var. adoensis and C. senensis or whether
the longer trichomes are homoplastic due to an adaptive nature or this is a case of
incomplete lineage sorting. Given the strong impact of aridication caused by the ice
ages, the ancestor of C.adoensis and C.senensis presumably survived during an arid
era in more humid coastal “forests” and woodlands of East Africa, where it evolved to
C.senensis and C.pwaniensis. Other morphs evolved in woodlands rather in the inland,
and are now pooled as C.adoensis. Interestingly, the distribution of C.adoensis var.
jereyana, C.senensis, and the allied C.pwaniensis (shares the subulate calyx lobes with
C.senensis) is very similar to that of the Apocynaceae species Carvalhoa campanulata
K.Schum. (Leeuwenberg 1985), which suggests shared ecological preferences.
e collections from Singida occur in drier habitats than those from C and S
Tanzania. Collections with an indumentum like C.adoensis var. jereyana also occur
in NE D. R. Congo (A. Taton 128, G. Troupin 570), but it is uncertain whether these
are also genetically linked to C.adoensis var. jereyana, so they are listed here under
C.adoensis var. adoensis.
Specimens examined. (selection, in total: 26) Kenya. Rift Valley Province: Na-
manga, cultivated in Munich Botanical Garden, N. Holstein 125 (M) and 130 (M).
Malawi. Northern Region: Mzimba Dist., 3 mi. W of Mzuzu at Katoto, J. Pawek
10404 (MO, WAG [WAG0234128]). Tanzania. Iringa: Ludewa district. Livingstone
Mountains, E slope of Msalaba Mountain, above stand of Acacia abyssinica on foot trail
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
72
from mission at Luana, 09°59'S 34°36'E, R.E. Gereau & C.J. Kayombo 3535 (DSM,
EA, MO, NHT, PRE); Great North Road, Sao Hill, 61ml. S of Iringa, R. Polhill
& S. Paulo 1722 (B, EA, P [P05621244], PRE). Mbeya: Nyassa Hochland, Station
Kyimbila, A.F. Stolz 504 (JE, M, U, W). Singida: 8½ km along road from Singida to
Sepuka, 04°46'35"S 034°40'00"E, P. Kuchar 23919 (MO, S [S08-12129]).
3. Coccinia barteri (Hook.f.) Keay, Kew Bull. 8: 82. 1953.
Staphylosyce barteri Hook.f. in Oliv., Fl. trop. Afr. 2: 554. 1871. Physedra barteri
(Hook.f.) Cogn. in A.DC.& C.DC.,Monogr. Phan. 3: 525. 1881.
Type: Nigeria. Nupe [Niger State]: exact locality not specied, male, , C.Barter 1525
(Lectotype, designated by Jerey (1967: 60): K!, digital image! [K]).
Type: Equatorial Guinea. [Fernando Po] Bioko Island, female, C.Barter no. ? (Syntype:
K?), see Taxonomic remarks.
Coccinia macrocarpa Cogn., Bull. Jard. Bot. État Brux. 5: 113. 1915–1919. Pro parte
[except E. Luja 125].
Type: D. R. Congo. Sankuru river [tributary of Kasai river], no detailed location given,
on farmland and in bushland, male, , Jul 1904, E. Luja 205 (Lectotype, designated
by Jerey (1967: 61): BR! [BR0000008888570, digital image: BR, JPS, photo: K!).
Type: Sankuru river [tributary of Kasai river], no detailed location given, female, fr, Nov
1903, E. Luja 125 (Syntype: BR! [BR0000008888228, digital image: BR, JPS]).
Coccinia subhastata Keraudren, Fl. Cam. 6: 131. 1967.
Type: Cameroon. South Region: Bitye, male, , 1917, G.L. Bates 1469 (Holotype: BM!).
Description. Perennial climber. Stems up to 10 m, glabrous or puberulous. Petioles
1–3.5(–8.5) cm, glabrous to puberulous, adaxial side rarely with trichomes. Leaves
3.5–20 × 4–23 cm, cordate, subhastate, shallowly to deeply 3- or 5-lobate. Lobes
triangulate, ovate to oblong. Margin entire with few to many teeth to serrate. Apex
obtuse to acute, with apical tooth. Upper leaf surface glabrous with clear or white
pustules, lower leaf surface glabrous to puberulous on main nerves, esp. towards base,
with or without small dark glands. Probracts ovate to elliptical, up to 5mm long or
missing. Tendrils simple or bid. Male owers in few- to many-owered racemes.
Common peduncle up to 3–8 mm long, glabrous to puberulous. Pedicel < 8 mm,
indumentum like peduncle. Flowers without or with up to 1.5 mm long bracts.
Perianth tube glabrous to puberulous. Calyx lobes 1–2.5 mm, subulate, lineal, rarely
somewhat lanceolate, reexed, spreading or erect and adpressed to corolla, sometimes
seemingly eshy. Corolla 1.1–2.4 cm, salmon, yellow to orange-yellow, lobes up to
3–10 mm. Filament column, anther head, and pollen sac color not seen. Female
owers in racemes, sometimes accompanied with a solitary ower or one solitary
ower only. Peduncles and petioles in racemes like in males. Solitary female owers
with up to 1.5 cm long glabrous to puberulous pedicel. Ovary glabrous. Hypanthium
glabrous to puberulous, calyx lobes and corolla as in males. Style not seen. Stigma
Monograph of Coccinia (Cucurbitaceae) 73
shape not seen, more or less dark yellow. Fruit 1.5–2.5 × 1.5 cm, shortly elliptical to
subglobose, unripe green with pale spots, ripe red. Seeds 5.5 × 2.5–3 × 1–1.5 mm
(L/W/H), more or less symmetrically obovate, face at to atly lenticular.
Phenology. Flowering time: January–June, August–November.
Distribution. Fig. 24. Humid tropical West Africa, Angola (Cabinda, Cuanza
Norte), Burundi?, C and S Cameroon, C and S Central African Republic, D. R. Congo,
R. Congo, Equatorial Guinea, Gabon, Mozambique (Manica), S South Sudan?, Ugan-
da (Western, Central), Rwanda?, W Tanzania, Zambia (Northern Province), Zimba-
bwe (Manicaland). Elevation from sea level to 1650 m. Soil preference not well known,
on loam soil, on granite (J.B. Gillett 15298). (Newtonia) rainforest; forest margins; near
open water with Pandanus candelabrum, Oxystigma mannii, and Raphia vinifera; near
river with Saba comorensis; on border of gallery forest and Terminalia glaucescens wood-
land; in riverine bushes, on river islands with Alchornea cordifolia; in fallows.
Use. e Turumbu people mash young leaves, mixed with white argil, and put the
paste onto the heads of ill children (W. Kesler 1034).
Vernacular names. Lissongo [Mbati]: makpo (C.Tisserant (Équipe) 2250); Twi:
isamaŋ kyẽkyẽa (F.R. Irvine 2604); Turumbu: eliki e litoko (J. Louis 2253), ndombo
di ilo (W. Kesler 1034)
Remarks. Coccinia barteri is treated here in a wide sense as it contains several forms
(see also Holstein and Renner 2011b). is is because data on these forms are scarce
and do not unambiguously allow to refer to these as species. erefore, the present
author refrains from creating an intraspecic classication as a phylogeographic treat-
ment and crossing experiments appear to be necessary to clarify this problematic taxon.
ere are collections in Gabon that are of intermediate morphology between
C.barteri and C.racemiora (M.A. van Bergen 490 (WAG) = C.barteri 6 in Fig. 17).
Holstein and Renner (2011b) suggested that hybridization occurs between these spe-
cies. Whether the hybrids are fertile or sterile is not known.
Taxonomic remarks. Coccinia barteri (Hook.f.) Keay is type species of the genus
Staphylosyce Hook.f.
Joseph Dalton Hooker mentions collections from Fernando Po [Bioko Island] and
Nupe in the protologue of C. barteri. He only gives the name of Barter, whose Nupe
specimen is in K, but there are no Coccinia specimens by Barter from Fernando Po.
However, there are two specimens collected by G. Mann (Mann N199! and N1166!)
in Hooker’s herbarium (now in K). ese were collected on that island, and they con-
tain drawings that were most likely the basis for Hooker’s description of Staphylosyce
barteri. Possibly, Hooker mistakenly left out Mann’s name in the protologue, whose
collections contain many type specimens (Hooker 1871).
Keay published in error Coccinea barteri [sic] in his new combination, but accepted the
species as belonging to Coccinia in Hutchinson and Dalziel’s Flora of Tropical Africa (1954).
e syntypes of Coccinia macrocarpa certainly belong to dierent taxa. e present
author concurs with Kéraudren, who placed the male specimen É. Luja 205 into the
polymorphic Coccinia barteri (1967). However, the female plant É. Luja 125 is clearly
not part of Coccinia. Coccinia seeds are up to 7mm long, at the base attenuate to trun-
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
74
cate and with a rounded apex. In contrast, the seeds of É. Luja 125 are subquadratic
as Jerey already pointed out on the type specimen. A placement in Momordica by
Jerey (on the sheet) seems to be correct, whether this is M. multiora Hook.f. (1871)
as identied by Jerey or M. parvifolia Cogn. (1916) as identied by Kéraudren is
beyond the present author's knowledge.
Coccinia subhastata was described under the assumption that C.barteri has long calyx
lobes, as it can be seen in Flore du Cameroun (Kéraudren 1967). Monique Kéraudren
in her research on western Central African Cucurbitaceae (Kéraudren-Aymonin 1975a;
Kéraudren 1967) treated Coccinia/Physedra barteri and P. heterophylla as synonymous.
However, she confused the long subulate calyx lobes of P. heterophylla as a character for
C.barteri, describing a specimen with short calyx lobes and owers in long racemes as
a new species, Coccinia subhastata Keraudren. She also described several dierences of
C.subhastata to C.barteri, which are not supported when carefully examined. Coccinia
subhastata should only have simple tendrils, but the holotype of C.subhastata also has a
bid tendril. Furthermore, the C.barteri lectotype C.Barter 1525 has a subhastate leaf
and simple tendrils. e description of C.subhastata thus is thus wrong and the species
is a synonym of C.barteri, as it has been pointed out by Holstein and Renner (2010). In
addition to the confusion of C.heterophylla and C.barteri, Kéraudren separated the western
Central African specimens with few-owered racemes as C.keayana R.Fern. (Kéraudren
Figure 24. Distribution map of C. barteri (based on 99 collections).
Monograph of Coccinia (Cucurbitaceae) 75
1967). Coccinia keayana, however, does in fact not occur in this region but only in West
Africa, and her C.keayana specimens from Cameroon belong to the polymorphic C.barteri.
Specimens examined. (Selection, in total: 153) Angola. Cuanza Norte: Cazengo
municipality, near Agricultural Station Cazengo, J. Gossweiler 5492 (COI, LISU), and
5507 (LISU). Benin. Atlantique: Allada commune, Dahounkpa (Niaouli), 6°44'N,
2°07'E, A. Akoègninou & F. Bada 2992 (WAG [WAG0314946]). Cameroon. South
Region: Bipinde [Bipindi], G.A. Zenker 1657 (E, G (4), HBG, P [P05621189], P
[P05621192], W, Z). Central African Republic.Lobaye: Boukoko, C.Tisserant (Équipe)
2176 (BM, P [P05620598], P [P05621175]). D. R. Congo. Katanga: [Haut Katanga
district], 40 km on road from Lubumbashi to Sakanja, A. Schmitz 4465 (EA (2), WAG
[WAG0225507], WAG [WAG0225508]). Equatorial Guinea. Bioko Norte: Malabo–
Punta Hermosa km 9, 32NMK8114, F.J. Fernández Casas 12077 (BM, MA n.v., MO,
WAG [WAG0069018]). Gabon. Haut-Ogooué: 21 km on road from Okonja to Akié-
ni, 0°45.84'S, 13°47.01'E, J.J. Wieringa et al. 6387 (M, WAG [WAG0250956], WAG
[WAG0250957], WAG [WAG0250958]). Ghana. Volta: Agumatsa Wildlife Sanctuary,
at town Wli-Agorviefe, W of Park Guard HQ, 7°06'46"N, 0°35'25"E, H.H. Schmidt
et al. 2192 (K, MO). Guinea. Nzérékoré: [Beyla préfecture], Bola [Boola], Famondou
[Famodougou], 8.48068°N 8.70129°W, E. Achigan-Dako 07 NIA 899 (GAT (3)). Ivo-
ry Coast. Lagunes: Abidjan, Banco Forest Reserve, in marshy valley, near the entrance,
J. de Koning 6144B (WAG [WAG0099439]). Nigeria. Rivers: Old GRA [Government
reservation area], Port Harcourt, B.E. Okoli 150 (IFE (cited and picture of plant in
Okoli 1984)). R. Congo. Sangha: 36.89 km E-SE Bomassa, 2°11.85'N, 16°31.10'E,
S.T. Ndolo Ebika 401 (E [E00486113]). Sierra Leone. Kono: Tingi Mts, top of E ridge,
J.K. Morton & D. Gledhill SL1886 (WAG [WAG0225480], WAG [WAG0225481]).
Tanzania. Kigoma: Kigoma Rural District, Gombe Stream National Park, Mitumba
Valley, research sta houses, 4°39'11"S, 29°38'09"E, G. Gobbo et al. 471 (MO). Zim-
babwe. Manicaland: Chirinda Forest, B. Goldsmith 39/62 (COI, K, NY).
4. Coccinia grandiora Cogn., Bot. Jahrb. Syst. 21: 211. 1895.
Coccinia grandiora Cogn. ex Engl., Abh. K. Preuss. Akad. Wiss.: 34. 1894. nom. nud.
Type: Tanzania. [Tanga]: Mlalo, dry hill range. C.H.E.W. Holst 506a (Holotype: B,
destroyed).
Type: Tanzania. Tanga: Usambara, near Amani, male, , [H.J.P.?] Winkler 3611 (Neotype,
designated here: BR!).
Coccinia engleri Gilg, Bot. Jahrb. Syst. 34: 354. 1904.
Type: Tanzania. [Tanga]: West Usambara, Sakare [Sakarre], at waterfall in primeval forest,
1100m, , fr, Sep, A. Engler, Reise nach Ostafrika 948 (Holotype: B, destroyed).
Type: drawing in protologue (Lectotype, designated here).
Description. Perennial climber. Stems up to 20 m, glabrous or (when from higher alti-
tudes) sparsely covered with long, whitish trichomes. Petioles 2.5–13 cm, indumentum
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
76
as on stem. Leaves 12–20 × 11–20 cm, profoundly 5-lobate. Lobes triangulate, ovate
to oblong. Leaf margin smooth to slightly serrate, dentate. Apex obtuse to acute with
nal tooth. Upper leaf surface glabrous with small hyaline pustules. Lower leaf surface
glabrous, rarely with few trichomes on the main nerves esp. at base, with blackish glands
scattered esp. along main nerves. Probracts up to 5mm long (Fig. 8a). Tendrils bid.
Male owers solitary or in (usually few-(–10-)owered) racemes. Common peduncle
4–12 cm, glabrous. Pedicels of owers in racemes 0.2–1.3cm, glabrous. Bracts up to 3
mm or missing. Pedicel of solitary owers 4–15 cm, glabrous. Perianth tube glabrous.
Calyx lobes (2–)4–13 mm, lineal, narrowly lanceolate to triangulate, tip subulate to sub-
acute. Corolla 4–6.5 cm long, apricot, salmon, yellowish-bu to yellow, lobes 2–4.7 cm.
Filament column and anther head not seen. Pollen sac yellow-orange. Female owers
solitary, rarely in racemes, glabrous. Common peduncle < 1 cm, pedicel in racemes, pedi-
cel of solitary owers 3–4 cm. Ovary long cylindrical, glabrous or with hyaline–whitish
pustules. Hypanthium glabrous, calyx lobes and corolla like in male owers. Style colum-
nar, yellowish to bu. Stigmas 2-lobed, yellow. Fruits < 30 × 2–4 cm, long cylindrical,
glabrous, when unripe green, ripe (orange-)red. Seeds 4.5 × 2.5 × 1–1.2 mm (L/W/H),
symmetrically obovate, face atly lenticular.
Phenology. Flowering time: January–December.
Distribution. Fig. 25. Southeastern Kenya (Coast Province: Taita Hills, coastal
forests), Tanzania (SE Dodoma; Iringa: Uzungwa Mts; Kilimanjaro; Lindi; Moro-
goro: Nguru Mts, Ukaguru Mts, Uzungwa Mts; Mtwara; Tanga: Usambara Mts), Ma-
lawi (Northern Region: Misuku Hills; Central Region: Nchisi Mts; Southern Region:
Lisao Hill), Mozambique (Manica: East African Highlands with foothills), Zimbabwe
(Manicaland: East African Highlands with foothills). Elevation 30–1900 m. On rich
red-brown clay, over limestone, on diabase outcrops. Coastal and lowland forests and
forest margins (Parinari sp.-Newtonia buchanii forests and others), riverine forests
(with Cola clavata, Synsepalum msolo, Sorindeia madagascariensis), rarely in montane
forests (Cassipourea malosana-Teclea simplicifolia-Teclea nobilis-Olea mildbraedii-Tab-
ernaemontana forest), or Miombo from degraded forests.
Use. Fruits are reported to be either poisonous (A. Peter 56598) or edible (W.J.
Kindeketa 630). Leaves cooked in water used against fever (K. Braun 714).
Vernacular names. Kihehe: mudesselema (F. Haerdi 617/0), Kipare: hotwe (W.J.
Kindeketa 630), Kishamba: matombo shanga (G.R. Williams G43), Kisamba: matom-
bo ya nyoka “snake breasts” (M.A. Mwangoka & A. Kalage 1578).
Remarks. e southern distributed individuals in Zimbabwe and C Mozambique often
bear short trichomes, and the leaves are rather shallowly lobate, just as in C.schliebenii. ese
populations may represent hybrids or descendants of a non-dierentiated common ancestor.
It is dicult to distinguish between C.grandiora and C.mildbraedii in the Cen-
tral Tanzanian highlands (Eastern Arc Mts). Both species also occur in high altitude
forests and are clearly delimited by ower size. Coccinia grandiora also has larger pro-
bracts than C.mildbraedii, but this is rarely well visible. Coccinia grandiora may also
be confused vegetatively with C.barteri in Mozambique and Zimbabwe.
Monograph of Coccinia (Cucurbitaceae) 77
Figure 25. Distribution map of C. grandiora (based on 62 collections). For Tanzania the borders of
the regions are given.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
78
Taxonomic remarks. e C.grandiora holotype by Holst was destroyed in the
re of the Berlin herbarium in 1944. e Winkler specimen was chosen as neotype
because it was already designated as type in December 2008. ere is no annotation on
the type label, however, and it seems that this neotypication was not published. How-
ever, the Winkler specimen label bears Cogniaux’ handwriting. Strangely, the Winkler
specimen also states “mars 1892”, with the 92 crossed out. is is the date when Holst
collected his specimen; but H. J. P. Winkler collected in Tanzania in 1910.
As the holotype of C.engleri also was destroyed, the original material left is a
drawing in the publication of the protologue. e drawing is of sucient quality to
synonymize unambiguously C.engleri with C.grandiora.
Specimens examined. (Selection, in total: 105) Kenya. Coast Province: Taita-De-
veta District, Taita Hills, Mbololo Forest, 3°19'S, 38°27'E, Mwambirwa Forest Station,
R.B. Faden et al. 799 (EA, WAG [WAG0234141]). Malawi. Southern Region: Chirad-
zulu district, Lisau Hill above Njuli P.O., R.K. Brummitt & I.H. Patel 18534 (K, WAG
[WAG0361192]). Mozambique. Manica: Chimoio, Garuzo mountain ridge, J.G. Garçia
522A (LISC, MO). Tanzania. Arusha: above Saje, N side of Ngurdoto Crater rim, P.J.
Greenway & K. Kanuri 13444 (EA, K, M, PRE). Kilimanjaro: Chome [ward], Njokava
forest, R. Abdallah 814 (EA, NHT). Lindi: 40 km W of Lindi, Lake Lutamba, Mirola
Valley, H.J.E. Schlieben 5905 (B, HBG, G, M, S [S08-12068], Z (2)). Morogoro: Kilosa
district, Ukaguru Mts, along track between Mandege and Ihanga rock, c.6°24'S, 36°56'E,
M. ulin & B.E. Mhoro 2877 (DSM, EA, K, MO). Tanga: western Usambara Mts, Sha-
gai forest, forestry house near Sunga, R.B. Drummond & J.H. Hemsley 2783 (B, EA, K,
S [S08-12075]); [eastern Usambara Mts], Amani, c.50 m before gate of headquarters,
05°05'58.3"S, 38°39'11.2"E, N. Holstein et al. 96 (B, DSM, M), 97 (BR, DSM, M), 98
(DSM, M, WAG), 99 (DSM, M), and 100 (M). Zimbabwe. Manicaland: Chipinge dis-
trict, outskirts of Chirinda [forest] above Msilizwe river, B. Goldsmith 2/63 (B, BR, S [S08-
12072]); [near Mutare], S slope of Murahwa’s Hill, N.C.Chase 8008 (COI, K, LISC, MO).
5. Coccinia grandis (L.) Voigt, Hort. suburb. Calcutt.: 59. 1845.
Bryonia grandis L., Mant. Pl.: 126. 1767. Coccinia indica Wight & Arn., Prodr. . Ind.
orient.: 347. 1834. nom. illeg. [nom. super. as epithet has not been adopted].
Coccinia grandis M.Roem., Syn. Pepon.: 93. 1846. nom. illeg. [nom. super.]
Cephalandra indica Naudin, Ann. Sci. Nat. Bot. ser. 5, 5: 16. 1866. nom. illeg.
Cephalandra grandis (L.) Kurz, J. As. Soc.Beng. 46(2): 103. 1877.
Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., es.
zeylan.: 49, t. 19, g. 2. 1737 (Type: drawing in l.c.).
Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448,
t. 166, g. 1. 1747 (Type: drawing in l.c.).
Type: No detailed information, female, , Anon. in Herb. Linn. 1153.2 (Typolectotype,
designated by Nazimuddin and Naqvi (1984): LINN! [digital image: LINN]).
Monograph of Coccinia (Cucurbitaceae) 79
Type: India. Gottori, male and female, , Anon. in Herb. Linn. 1153.3 (Type?: LINN!
[digital image: LINN]).
Type: India. [Gujarat]: Suratt [Surat], female, , Anon. in Herb. Linn. 1153.12 (Type?:
LINN! [digital image: LINN]).
Type: India. No detailed information, female, , Anon. in Herb. Linn. 1153.13 (Ty-
potype: LINN! [digital image: LINN]).
Cucumis sativus var. arakis Forssk., Fl. aegypt.-arab.: 169. 1755.
Type: Yemen. [Al-Hudaydah Governorate]: Lohaja [Al-Luhayyah] [region?], Môr
[Mawr], male, , P. Forsskål 660 (Holotype: C [C10002122, digital image: JPS,
microche IDC: 35 II, 7–8]).
Turia moghadd Forssk. ex J.F.Gmel., Syst. nat. 2(1): 403. 1791. Turia moghadd
Forssk., Fl. aegypt.-arab.: 166. 1755. nom. inval. Coccinia moghadd (Forssk. ex
J.F.Gmel.) Asch. in Schweinf., Beitr. Fl. Aethiop.: 251. 1867. Cephalandra mog-
hadd (Forssk. ex J.F.Gmel.) Broun et Massey, Fl. Sudan: 105. 1929.
Type: Yemen. [Al-Hudaydah Governorate]: Lohaja [Al-Luhayyah], female, , P. For-
sskål 663 (Lectotype, designated here: C! [microche: IDC 110 I, 1–2]).
Type: ibid., male, , P. Forsskål 662 (Syntype: C! [microche: IDC 109 III, 7–8]).
Type: ibid., P. Forsskål 666 (Syntype: C! [microche: IDC 110 I, 3–4]).
Bryonia alceifolia [sphalm. alceaefolia] Willd. in Rottler, Neue Schriften d. Ges. Naturf.
Freunde Berlin 4: 223. 1803. Coccinia cordifolia (L.) Cogn. var. alceifolia [sphalm.
alceaefolia] (Willd.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 531. 1881.
Type: India. [Tamil Nadu]: Tiruchinapally [Tiruchirappalli], male and female, , Nov
1793, Anon. [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. J.G. Klein 177 (Lecto-
type, designated here: B-W! [B-W 18065], isolectotype: K!).
Momordica covel Dennst., Schlüssel Hortus malab.: 23. 1818. Cucumis pavel Kostel.,
Allg. med.-pharm. Fl. 2: 738. 1833. nom. illeg. [nom. super.]
Type: Covel Rheede, Hort. malab. 8: 27, t. 14. 1688 (Holotype: drawing in l.c.).
Momordica bicolor Blume, Bijdr. . Ned. Ind.: 928. 1825–26.
Type: Indonesia. [Java], Kuripan, in calcareis [on calcareous ground?], K.L. Blume
1012 (Holotype: L! [L 0587745]).
Momordica bicolor var. a Blume, Bijdr. . Ned. Ind.: 928. 1825–26. nom. inval. Indo-
nesia. Maluku Province, Timor, A. Zippelius s.n. (L! [L 0587743]).
Momordica bicolor var. b Blume, Bijdr. . Ned. Ind.: 928. 1825–26. nom. inval.
[Indonesia]. [Java], Parang Mts, calcareous mountains. K.L. Blume 1016 (L! [L
0587744]).
Bryonia moimoi Ser. in DC.Prodr. 3: 305. 1828. Moï-moï Adans., Hist. nat. Sénégal:
159, Paris. 1757. nom. inval. Coccinia moimoi (Ser.) M.Roem., Syn. Pepon.: 93.
1846.
Type: Sri Lanka. Bryonia folio anguloso acuto glabro Burm., es. zeylan.: 48, t. 19, g.
1. 1737 (Holotype: drawing in l.c.).
Momordica monadelpha Roxb., Fl. ind. [2nd ed.], 3: 708. 1832. nom. illeg. [nom. su-
per.] pro parte. Bryonia foliis cordatis oblongis angulatis dentatis glabris. Fl. Zeyl.
356. 1747. nom. inval.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
80
Type: Sri Lanka. P. Hermann Mus. Zeyl. 2: 37 (Type: BM [BM000621642], BM
[BM000621643]) [these two specimens are Cucumis maderaspatanus L.].
Type: Sri Lanka. P. Hermann Mus. Zeyl. 5: 225 (Lectotype, designated here:: BM
[BM000595000]).
Type: Sri Lanka. P. Hermann Mus. Zeyl. 5: 321 (Type: BM [BM000621089]).
Type: Sri Lanka. Bryonia folio anguloso acuto glabro Burm., es. zeylan.: 48, t. 19, g. 1.
1737. (Type: drawing in l.c.).
Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., es. zey-
lan.: 49, t. 19, g. 2. 1737 (Type: drawing in l.c.).
Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448, t.
166, g. 1. 1747 (Type: drawing in l.c.).
Coccinia loureiriana M.Roem., Syn. Pepon.: 93. 1846. Bryonia grandis Lour., Fl. cochinch.
1(2): 595. 1790. nom. illeg. and Fl. cochinch. 2: 731. 1793. nom. illeg.
Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., es. zey-
lan.: 49, t. 19, g. 2. 1737. (Type: drawing in l.c.).
Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448, t.
166, g. 1. 1747 (Lectotype, designated here: drawing in l.c.).
Coccinia wightiana M.Roem., Syn. Pepon.: 93. 1846. Coccinia cordifolia (L.) Cogn.
var. wightiana (M.Roem.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 531.
1881. Coccinia grandis var. wightiana (M.Roem.) Greb. in R. Mansfeld & J.
Schultze-Motel, Verz. Landwirtsch. u. Gaertn. Kulturp. 2: 929. 1986. Coccinia
indica Wight et Arn. β, Prodr. . Ind. orient.: 347. 1834.
Type: India. [Tamil Nadu]: [= Wall. Cat 6711a], [J.G. Klein, B. Heyne or J.P. Rottler] in
Herb. Madras s.n. (Syntype: E! [E00174668, digital image: E, JPS]).
Type: India. [Tamil Nadu]: [Chennai, Saidapet], Nopalry [= in Wall. Cat. 6711b ore],
female, , R. Wight 1124 (Lectotype, designated here: E! [E00174667, digital
image: E, JPS]).
Type: India. [Tamil Nadu]: Negapatam [Nagapattinam], female, , fr, R. Wight 1124
(Syntype: E! [E00174666, digital image: E, JPS]).
Type: unknown. R.Wight 1124 (Syntypes: E! [E00174664], NY! [00172358, digital
image: NY]).
Coccinia grandis var. quinqueangularis Miq., Fl. Ned. Ind. 1(4): 673. 1856.
Type: [Indonesia]. [Central Java]: near Soerakarta [Surakarta], T. Horseld s.n. (Holo-
type: U!).
Coccinia schimperi Naudin, Ann. Sci. Nat. bot., ser. 4, 8: 366. 1857. Cephalandra
schimperi (Naudin) Naudin, Ann. Sci. Nat. Bot. ser. 5, 5: 16. 1866.
Type: Ethiopia. In Semen [Semien Mts], female, , 1854, G.H.W. Schimper Herb.
Abyss. 1215 (Lectotype, designated here: P! [the specimen with thick branch and
fruit], isolectotype: P!).
Type: Ethiopia. Biria Dekeno et Dschadscha, 5000’, female, , 1853, G.H.W. Schimper
Herb. Abyss. 1215 (Syntype: P!).
Type: Ethiopia. Dschadscha, 5000’–5500’, male and female, , fr, 13 and 21 Jul 1853,
G.H.W. Schimper Herb. Abyss. 1215 (Syntype: P!).
Monograph of Coccinia (Cucurbitaceae) 81
Type: Ethiopia. Dschadscha, 5000’, male and female, , fr, 13 and 21 Jul 1853, G.H.W.
Schimper Herb. Abyss. 1215 (Syntype: P!).
Type: Ethiopia. without details, G.H.W. Schimper Herb. Abyss. 1215 (Syntypes:
BR! [BR0000005113958, digital image: BR, JPS], BR! [BR0000005114641,
digital image: BR], BR! [BR0000008251053, digital image: BR, JPS], BR
[BR0000008351050], G-DC!).
Type: Ethiopia. Dschadscha, female, 1853, G.H.W. Schimper Herb. Abyss. 1215 (Syn-
type: BR! [BR0000005111923, digital image: BR]).
Coccinia palmatisecta Kotschy, Sitzungsber. K. Akad. Wiss. Math.-Naturwiss. Cl. Abt.
1, 51: 360 –361. 1865.
Type: [South Sudan], Kyk [Ciec (a Dinka subtribe) realm, S of conuence of Bahr
al-Ghazal and White Nile], male, , no date given, M.L. Hansal s.n. (Lectotype,
designated here: W! [K neg. 4837]).
Type: [South Sudan]. marshes in Noer [Nuer] realm [S to E of Malakal], no detailed
location given, F. Binder s.n. (Syntype: W! [K neg. 4838]).
Cephalandra indica var. palmata C.B.Clarke in Hook.f., Fl. Brit. Ind. 2(6): 621. 1879.
Bryonia alceifolia [sphalm. alceaefolia] Willd. in Rottler, Neue Schriften d. Ges. Naturf.
Freunde Berlin 4: 223. 1803. Coccinia wightiana M.Roem., Syn. Pepon.: 93. 1846.
Type: India, [Tamil Nadu]: Tiruchinapally [Tiruchirappalli], male and female, , Nov
1793, Anon. [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. J.G. Klein 177 (Syntypes:
B-W! [B-W 18065], K!)
Type: India. [Tamil Nadu]: [= Wall. Cat 6711a], [J.G. Klein, B. Heyne or J.P. Rottler] in
Herb. Madras s.n. (Lectotype, designated here: E! [E00174668, digital image: E, JPS]).
Type: India. [Tamil Nadu], [Chennai, Saidapet], Nopalry [= in Wall. Cat. 6711b
ande], female, , R. Wight 1124 (Syntype: E! [E00174667, digital image: E, JPS]).
Type: India. [Tamil Nadu]: Negapatam [Nagapattinam], female, , fr, R. Wight 1124
(Syntype: E! [E00174666, digital image: E, JPS]).
Type: unknown. R.Wight 1124 (Syntypes: E! [E00174664], NY! [00172358, digital
image: NY]).
Coccinia cordifolia (L.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 529. 1881. pro
parte majore, non Bryonia cordifolia L.
Coccinia cordifolia (L.) Cogn. var. genuina Cogn. in A.DC. & C.DC., Monogr. Phan.
3: 531. 1881. nom. invalid.
Description. Perennial climber or creeper. Stems up to 5 m, glabrous, when older
often white pustulate. Petioles 0.5–5.5 cm, glabrous, rarely some trichomes on adaxial
side. Leaves 3–11 × 3–13 cm, cordate to 3-lobate or 5-edged to 5-lobate, sometimes
lobulate. Lobes triangulate, ovoid, oblong, to obovoid. Leaf margin dentate, teeth usu-
ally with yellowish-reddish to brownish gland (Fig. 7a), becoming black when dried.
Margin rarely with short (< 1 mm), whitish trichomes. Apex obtuse to acute with nal
tooth. Upper leaf surface glabrous, more or less dense hyaline to white pustulate. Lower
leaf surface glabrous, with glands that are usually framed with lighter color between
major nerves, nerves sometimes with white pustules. Probracts < 1.5 mm or missing.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
82
Tendrils simple. Male owers 1(–3) solitary, rarely in short racemes. Peduncle 0.3–1.5
cm, glabrous, pedicels of owers in racemes up to 3.2 cm, glabrous, pedicels of solitary
owers up to 4.5 cm, glabrous. Bracts inconspicuous (< 1 mm), often absent. Perianth
tube glabrous. Calyx lobes 1.2–3.5 mm long, lineal, spreading to reexed, often with
reddish to brownish gland on lower side at the acute tip (Fig. 7a). Corolla 1.7–4 cm
long, yellowish bu (mostly African collections) to snow-white (esp. outside Africa).
Corolla lobes 0.7–1.7 cm. Filament column and anther head pale greenish, pollen sacs
yellow. Female owers 1(–2) solitary. Pedicel up to 1 cm, glabrous. Hypanthium gla-
brous, calyx and corolla like in males. Ovary glabrous. Style columnar, yellowish-green.
Stigmas 2-lobed, greenish. Fruit globose to ellipsoid, cultivated also shortly cylindrical,
3–4.5 × 1.5 cm, glabrous, unripe greenish with few pale spots and/or lines, becoming
scarlet red when ripe. Seeds 5–7 × 2.5–3.5 × 1.2mm (L/W/H), asymmetrically obo-
vate, face at (Fig. 14f).
Phenology. Flowering time: All over the year, but not at the end of the dry season
or in cold seasons. Seems to need 1 –2 weeks of at least 10 hours daylight with sunny
weather for ower induction (pers. observ. from greenhouse cultivation).
Distribution. Figs 26, 27. Benin, N Cameroon, S Chad, D. R. Congo (in Great
African Rift valley), Djibouti, Egypt (along Nile, Elba Mts), Eritrea, Ethiopia, Kenya,
Mali (except N), S Mauritania, Niger, Senegal, Somalia, South Sudan, S and E Sudan,
N Tanzania (Arusha, Dar es Salaam, Manyara, Morogoro, Mwanza, Pwani, Tanga,
Zanzibar), Uganda, mountains and oases of the southern and western Arabian Penin-
sula, tropical and subtropical India, subtropical Nepal, Pakistan, Sri Lanka, South East
Asia, S China incl. Hainan. Occurrence likely due to recent introduction in: Australia
(Northern Territory, Queensland, Western Australia), R. China, Maldives, Mauritius,
Mozambique, USA (Florida, Guam, Hawaii, Wake Island), many tropical Pacic is-
lands, Caribbean area, Central and tropical South America. 0 to 1600 m. Black clay soil,
black cotton soil, sand, on limestone. ere seems to be a certain salt tolerance (Bhar-
athi 2007). Grasslands, bushlands, (semi-arid) savannas, thickets, along rivers, ruderal
sites, seemingly avoiding closed canopies (woodlands and forests). Coccinia grandis is,
especially in tropical Asia, often reported from sand or from calcareous grounds (karst
areas), which are well-drained. Although precipitation in South East Asia is much high-
er than in Africa, the quick loss of surface water allows C.grandis to survive there.
Use. Fruits (raw and cooked) and shoots (cooked) edible. e Luo eat the leaves
as spinach (Orech et al. 2005). e sap is used against diabetes (Ramachandran and
Subramaniam 1983) due to its hypoglycemic activity (Chopra and Bose 1925; Shibib
et al. 1993). It is used in traditional Indian medicine in India for various diseases,
and seems to have a general antibacterial eect (see also Use, economic potential, and
phytochemistry).
Vernacular names. Agau or Chomir [most likely: Khamir/Xamtanga language of
Agaw language family]: amballa gosa (G.H.W. Schimper 365, Schweinfurth 1893); Ara-
bic: mogad (Getahun 1974b); Bagirmi: na odio (A.J.B. Chevalier 9527); Bengali: tala-
kucha (Nadkarni and Nadkarni 1976), tela kucha [tēlākucā] (J. Sinclair 4414); Canare-
se [Kannada]: tonde-konde, tonde-kayee [toṇḍe kāyi] (Nadkarni and Nadkarni 1976);
Chinese: hong gua [Hóng guā] (Wu et al. 2011); Dassanetch [Daasanach]: dali (C.J.
Monograph of Coccinia (Cucurbitaceae) 83
Carr & C.Metolo 867); Galinya [Oromo]: raho (T. Ebba 624; Getahun 1974b); Gujara-
ti: gholi (Nadkarni and Nadkarni 1976); Hindi: kan-duriki-bel (Nadkarni and Nadkarni
1976), kundru [kundrū] (Bharathi 2007); Karen [in Mae Hong Son Province, ai-
land]: khae-do (de Wilde and Duyes 2008); Khmer: (sloek’) bah (Kéraudren-Aymonin
1975b); Kizigua: lukewja (R.E.S. Tanner 2691), pondwa (R.E.S. Tanner 2030); Lao:
(khùa ‘phăk) tăm ling, tăm nhing (Kéraudren-Aymonin 1975b), phak tam nin (J.-M.
Dubost 200); Luo: nyathud gush (S.H. Padwa 71); Mahrati [Marathi]: ran-tondula, ton-
dli (Nadkarni and Nadkarni 1976); Malayalam: kova (Nadkarni and Nadkarni 1976),
kovakka (Bharathi 2007); Masai: olaposhi (C.H. Pecler 3); Nepali: gol kankri (Aryal
2007); Ngadha (Baba-Ngadha): `uta kala (J.A.J. Verheijen 5415); Ngadha (Mataloko-
Ngadha): uta kobho (J.A.J. Verheijen 5415); Persian [Farsi]: kabare-hindi (Nadkarni and
Nadkarni 1976); Punjabi: kanduri (Nadkarni and Nadkarni 1976); Sanskrit: vimboshta,
Figure 27. Partial distribution map (Pacic Ocean and Americas) of C. grandis (based on 629 collections).
State borders are given for the USA.
Figure 26. Partial distribution map (Africa to Australia) of C. grandis (based on 629 collections).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
84
vimbaja, bimba, tundika [tuṇḍikā] (Nadkarni and Nadkarni 1976); Serer: ayandyaloup
(J.-F. Ezanno Cucurbitaceae 9); Sinhala: kowakka (D. Philcox et al. 10454, D. Philcox et
al. 10458); Somali: masskar (T. Ebba 804); Songhai: lombaria (A.J.B. Chevalier 2996);
Sunda: aroi papassang (Miquel 1855); Tamasheq [Touareg]: seel (H. Lhote 2). Tamil:
kovai [kowai] (Nadkarni and Nadkarni 1976), kovakkai (Bharathi 2007); Telugu: don-
dakaya [doṇḍa kāya] (Bharathi 2007), dondatiga, kakidonda (Nadkarni and Nadkarni
1976); ai: in Central Region: phak tamlueng, in Northern Region: phak khaep (de
Wilde and Duyes 2008); Tigrinya: asumbeh (G.A. Schweinfurth & D. Riva 1007);
Vietnamese: northern: hoa bát, southern: (dây) bai bác, bình bát, ma'nh bát, lá bát (Ké-
raudren-Aymonin 1975b); Wolof: barbouf (J. Trochain 663).
Remarks. With the exception of South and South East Asia, C. grandis is easily
recognizable, especially by the lack of an obvious indumentum and the pale framed
(in living state) glands in the axils of the nerves at the base of the lower leaf lamina. In
NE Africa, collections with nely dissected leaves can be similar to C.ogadensis. When
compared to collections from South Africa, C.sessilifolia var. variifolia and some forms
of C.mackenii are also similar, but C.sessilifolia var. variifolia is glaucous, C.mackenii
plants have bid tendrils, and both South African species lack colored leaf teeth and
have erect to spreading calyx lobes instead of spreading to reexed calyx lobes. In South
and South East Asia, some vegetatively similar Gymnopetalum species (e.g., G. chinense)
can be mistaken for C. grandis, as in both taxa the leaf shape is 5-edged to cordate,
and glands on the lower leaf lamina can be found. However, Gymnopetalum species are
rather densely beset with trichomes, have ribbed fruits and are monoecious, whereas
C. grandis is glabrous, has smooth fruits, and is dioecious, at least in wild populations.
Asian and (at least most of the) African C.grandis dier genetically (e.g., in a short
sequence in the 5’-beginning of the LEAFY-like 2nd intron) and in petal color (white in
Asian, bu in African individuals). Hence, the distribution in Asia (at least India) is not
due to human impact. Whether C.grandis is introduced or native to Malesia, northern
Australia and southern China and Taiwan is not known. Crossing experiments by Nau-
din (1862) indicate full compatibility between the African and Asian morphs though.
Taxonomic remarks. Up to the 21st century (e.g., Bulbul et al. 2011; Hussain et
al. 2011), there has been quite a lot of confusion about the valid name of the species
that is now called Coccinia grandis. Wight and Arnott (1834) established the genus
Coccinia, with the name based on the scarlet-red fruits of a species that Wight had col-
lected several times during his 13-year stay in South India (Staeu and Cowan 1988).
e name of the only species they described in their new genus, Coccinia indica Wight
& Arn., is illegitimate since they included Bryonia grandis L., of which they ought to
have adopted the epithet (Art. 52.1 ICN). e illegitimacy of Coccinia indica, how-
ever, does not aect the legitimacy of the genus name Coccinia (Art. 42.2), although
there was also some confusion about which genus name to use. In 1845, Voigt pub-
lished the correct combination Coccinia grandis (L.) Voigt with a description exactly
matching Bryonia grandis.
Although C.indica is not valid, and the problem seemed to have been solved, a
third species name was brought into discussion by Cogniaux. He thought that Bryo-
Monograph of Coccinia (Cucurbitaceae) 85
nia cordifolia L. and Bryonia grandis L. referred to the same species as Linnaeus cited
Rumphius’ Vitis alba indica (C. grandis) under Bryonia cordifolia (Linnaeus 1763).
Hence, Cogniaux created a more broadly circumscribed Coccinia cordifolia (L.) Cogn.
(1878). However, Linnaeus described Bryonia cordifolia beforehand (1753) citing the
description from his Flora Zeylanica number 354 (1747), and only in the 2nd and 3rd
edition of his Species Plantarum did Linnaeus synonymize Rumphius’ Vitis alba indica
(1747). Cogniaux therefore erred, when he stated that Linnaeus had based his B. cor-
difolia on Rumphius’ gure, and the epithet cordifolia is hence misapplied in Coccinia.
ere are four specimens in Linnaeus’ herbarium, which belong to Coccinia gran-
dis but it is unclear which are type material. e number 1153.2 is designated (by
Linnaeus?) as Bryonia grandis on the sheet and is therefore the best choice for lectotypi-
cation as is has been done by Nazimuddin and Naqvi (1984). e specimen 1153.13
bears the note “Bryonia foliis subrotundis angulosis, momordicae facie” on the speci-
men ip side referring to one of the citations to drawings in the protologue. Hence,
this specimen is also original material. e other two specimens (1153.3 and 1153.12)
do not bear indications that Linnaeus referred to them as Bryonia grandis, but there is
no contraindication either, so they might be original material, too.
e drawing to Bryonia foliis subrotundis, angulosis, momordicae facie Burm. (es.
zeylan.: 49, t. 19, g. 1. 1737), which is original material of Bryonia grandis L. appears
to be a product of artistic freedom. e calyx lobes of the three uppermost and the
lowermost owers are reexed and match Coccinia/Bryonia grandis well, whereas the
calyx of the other two owers appears to consist of almost free elliptic petals, quite like
in Momordica foetida (except for the soft spines that are missing in the drawing). e
addition “momordica facies” seems to relate to this. Strangely, the drawing of Bryonia
folio anguloso acuto glabro Burm. (es. zeylan.: 48, t. 19, g. 1. 1737) matches the
current denition of Bryonia grandis also well, but has not been cited by Linnaeus.
Eventually, the synonyms that Burman used, which also include Bryonia cordifolia L.
(Cucumis maderaspatanus L.), made it hard to interpret the species and also lead to the
confusion of Cogniaux.
e Forsskål 660 specimen (C10002122) has a hand-written eld label by Forsskål
(according to notes in JStor Plant Science) on the ip side, stating “Cucumis incerta.
Arakis, Mour.” (incerta meaning “uncertain”) and two hand-written identications
“Cucumis inedulis Fl. Yemen CXXII nr. 580” and “c.s. Arakis cent VI nr. 61 p. 169”.
e location in the text is the same as on the eld label: Môr/Mour [Mawr, a small
town about 30 km E of al-Luhayyah]. e former identication is a nom. nud. with
the number 580 on page CXXII of Forsskål’s Flora Ægyptiaco-Arabica (1775). e
second identication links to a Cucumis sativus variety that is validly described in that
book on page 169. e description matches well Coccinia grandis except for the tuber-
culate ovary. Additionally the collection is supposed to be from Loheja [al-Luhayyah],
but might only indicate the region, in which Mawr is localized. However, the Arabic
name of Cucumis inedulis and Cucumis sativus var. arakis are both: Arakis [3raqīs],
so they can be cross-referenced. e description of the variety also mentions that the
plant is not edible, just as the supposed species epithet.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
86
e genus name Turia has been created by Forsskål in his Flora Ægyptiaco-Ara-
bica (1775). ere is a debate, however, whether it is validly published (Friis 1984).
e descriptions of this part of Forsskål’s book (page number in Latin) must be used
in consideration with the corresponding parts in the lists of local oras (Roman page
numbers), which is on page CXXI in the case of Turia. e rst species there is Turia
sativa (no. 550), which is called “turia” in Arabic and is cultivated according to the
epithet. is matches exactly the rst description in the descriptive part of the book
(p. 165). Forsskål lists ve Turia species in the oristic part and describes ve species
in the descriptive part. erefore, the genus Turia lacks a description (Jerey 1962),
because Forsskål does not mention any character to be typical for the genus. e name
Turia was legitimately described by Gmelin in an extension of Linnaeus’ Systema
naturae (1791), and so was the name Turia moghadd.
e typication of Turia moghadd is not straight-forward, because Forsskål added
little, if any marks on the sheets (Friis 1983). e ip side of the Forsskål 663 specimen
has three notes: “Cucumis glandulosus”, a second one with a dierent handwriting:
“Bryonia Turia 35 Forsk” and a third one with another handwriting “Turia gijef Forsk
Cent. 6 no. 38” with all words except for “Turia” crossed out. It can be hypothesized
that these are dierent trials to identify the specimen, but not by Forsskål himself.
Aside from these, the specimen lacks any written marks, but it matches exactly the
description of Turia moghadd, as do the specimens Forsskål 662 and 666, which are
syntypes. e Forsskål 663 specimen is chosen to be the lectotype, because the original
description mentions the occurrence of fruits and this specimen is the only female of
the three.
Willdenow describes Bryonia alceifolia in a travel report by J. P. Rottler from 1799,
but he only mentions that he separates Rottler’s Bryonia epigaea from another new
species, viz. B. alceifolia, so the type was not necessarily collected in 1799. Willdenow
knew B. alceifolia from Klein’s specimens in his herbarium. Rottler was missionary in
Tranquebar, the same place in which J. G. Klein was surgeon (Jensen 2005). Both are
known to have collected together (with B. Heyne). e lectotype collection is in the
describer’s herbarium under the Klein number 177, but it is not certain, whether Klein
collected the specimen himself, of if it was by Rottler or even Heyne. A duplicate with
the same label data as in B-W is deposited in K. For Willdenow described the species,
the specimen in his herbarium was chosen to be lectotype.
e name Momordica monadelpha Roxb. is superuous, because Roxburgh syn-
onymized Bryonia moimoi Ser. in total by citing the only element of that name and
Bryonia grandis. e other elements of M. monadelpha are also interesting, though.
Roxburgh cited “Bryonia foliis subrotundis” with the citation of Burman’s es. zey-
lan.: t. 19, g. 1. 1737 and g. 2, Vitis alba indica, all Coccinia grandis, and Herman’s
Musæum Zeylanicum 356. e latter one consists of two specimens (2: 37), which are
Cucumis maderaspatanus L., however, there are also two drawings (5: 225 and 5: 321)
with the number 356. Both drawings represent Coccinia grandis because of the fruit
size, fruit shape, and the ower morphology (calyx lobe length and position, corolla
size), rather than Cucumis maderaspatanus L., Diplocyclos palmatus L. or Cayaponia
Monograph of Coccinia (Cucurbitaceae) 87
laciniosa (L.) C.Jerey. at the drawing 5: 225 (BM000595000) shows a plant with
male and female owers on one individual might be explained best by artistic freedom.
e name Coccinia indica var. palmata C.B.Clarke is valid and legitimate but not
obvious to typiy. Despite C.indica being illegitimate as a nomen superuum, the
variety is legitimate and validly described. Clarke cites Bryonia alceifolia Willd. and
the C.indica protologue with page 348 although C.indica was described on page 347
(Wight and Arnott 1834). Page 348, however, only comprises the β variety, which
itself was the basis for C.wightiana M. Roem. is unnamed variety consists of several
elements, of which the literature citations of Bryonia palmata are mentioned with a
question mark and are thus not eligible for typication (Art. 52.2 N1). e element
“Bryonia palmata Linn.? herb. Madr.!” relates to a collection in Herbarium Madras
that was identied a Bryonia palmata, but Wight and Arnott appear to have been
in doubt whether the name was used sensu Linnaeus, hence the question mark after
the name. is specimen (or a duplicate) is found in E and represents a deeply lobed
C.grandis just as the protologue of the β variety says. Clarke obviously takes the epi-
thet of the variety from this misidentication and not from Bryonia palmata L., which
is why the name was typied with this collection. at Clarke meant a deeply lobed
C.grandis is evident since he also cites B. alceifolia, which is also deeply lobed. It is thus
not a new combination and status change but a new variety.
e name Coccinia cordifolia var. genuina Cogn. cannot be regarded as intended to
represent a new variety, because Cogniaux divides all specimens he had seen into three
varieties. Hence, the epithet “genuina” and his “genuina” variety is just synonymous
with the autonym.
Specimens examined. (Selection, in total: 1066). Australia. Queensland: Weipa,
Awonga Court, 12°37'28"S, 141°52'39"E, B.M. Waterhouse 7428 (BRI, CANB
[CANB 682857]). Bangladesh. Sylhet: Sillet, F. de Silva in Wall. list 6700g (M [“439”],
M [“440”]). Barbados. St. Michael: Bridgetown, old railway yard, Sep 1940, E.G.B.
Gooding s.n. (BAR00002591). Brazil. Minas Gerais: No detailed location, <1840,
P.C.D. Clausen s.n. (G-DC). Cameroon. North Region: Pitowa [Pitoa], c.17 km
NE of Garoua, W.J.J.O. de Wilde et al. 4932 (MO, WAG [WAG0225471]). Chad.
Chari-Baguirmi: Baguirmi et région du Lac Fittri, Tjecna, A.J.B. Chevalier 9527 (G,
P [P05620497]). P. R. China. Hainan: Ch’ang-kiang District [Changjiang country],
[Bawangling Nature Reserve], Ka Chik Shan [Qicha hill], S.K. Lau 3008 (GH n.v., P
[P06394498], S). Hong Kong: near Ouang-nei-Tchong [Wang nai chung] – Happy
Valley, E.M. Bodinier 1367 (E). Yunnan: Nan-chiao [Meng che], C.W. Wang 76988
(A n.v., IBSC [0207527], KUN n.v., PE [01178321], PE [01178322]). R. China.
Pingtung County: Hsinpi [Sinpi], Y.-P. Cheng 4603 (BR [BR0000005164189],
TAIF). Cuba. Havana: Santiago de las Vegas, A. O’Donovan 5106 (G, S [S08-
12050]). D. R. Congo. Orientale: Mahagi territory, Mahagi-Port, J. Lebrun 3800
(BR (2), EA, WAG [WAG0225470]). East Timor. Dili: just W of Dili, Tasutolu
area, 8°33'57"S, 125°30'05"E, L.D. Cowie 10658 (L). Egypt. Red Sea Governorate:
[Hala’ib triangle], Gebel Shendib, G.W. Murray 3857 (K [K000037307]). Eritrea.
Gash-Barka: Mai Mentai [at Sciagolgol River, SE of Agordat], N. Beccari 118 (FT).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
88
Northern Red Sea Region: [NE of Ghimda/Gimda], Pianura Sabarguma, A. Pappi
3970 (G, MO, P [P05620558], S [S08-12133], W). Ethiopia. Oromia: 85 km
NE of Nazareth, along road to Awash, c.5 km W of Metahara, 8°55'N, 39°55'E,
J.J.F.E. de Wilde 6870 (BR, MO, WAG [WAG0225448], WAG [WAG0225449],
WAG [WAG0225450]). Tigray: near Djeladjeranné, 23 Apr 1841, G.H.W.
Schimper 1570 (P [P05620544], specimens from BM, G, MO, S [S08-12147], and
W [W 0011063] might also be from this location, for details see under Cucurbita
schimperiana). Guyana. Demerara-Mahaica: Georgetown, W. Hahn 4810 (MO, US).
India. West Bengal: Sibpur [Shibpur in Howrah city] near Calcutta [Kolkata], J.G.
Hallier s.n. (M [“999”]). Indonesia. East Java: Pasoeroean [Pasuruan], 17 Jul 1928,
J.J. Ochse s.n. (B, BR [BR0000009938076], L [L 0587578], P [P06394462], U).
Mali. Kayes: Bafoulabé Cercle, Bamafélé arondissement, 3.2 km ESE of Manantali,
13°10.847'N, 10°26.260'W, C.S. Duvall 130 (MO (2)). Mozambique. Maputo: Vila
Luísa [Marracuene], along Incomáti river, A. Balsinhas 1273 (COI). Nepal. Madhya
Pashchim: [Rapti], Tulsipur, O. Polunin et al. 5906 (E). Nicaragua. Chontales: St
Tomas [Santo Tomàs], 1841, E. von Friedrichsthal s.n. (W). Niger. Agadez: [Aïr
Mountains], Mt Bagwezan [Mt Bagzan, Idoukal-n-Taghès], 31 May 1920, A.
Buchanan s.n. (MO [acc.no. 1667128]). Pakistan. Sindh: 7 mls [c.11.2 km] from
Hyderabad to Tando M.[Mohammad] Khan, S. ul-Abedin 3839 (B, KUH n.v.,
MO). Senegal. Dakar: Jof [Yo], J.-G. Adam 1806 (MO (2), [non P [P00694028/
P05590411], which is a Citrullus sp.]). Singapore. South East District: Potong Pasir
o Serangoon Road, J. Sinclair 5434 (E, L [L 0587600], P [P0639450]). Somalia.
Togdheer: [c.54 km SSW of Berbera], foot of Sheikh Pass, P.R.O. Bally 11827 (G
(3), K, PRE). Sri Lanka. North Central Province: Anuradhapura, F.R. Fosberg & N.
Balakrishnan 53431 (MO, PDA n.v.); ibid., W. Forstner s.n. W13705 (W). Sudan.
North Kurdufan: Chursi [Khursi], K.G.T. Kotschy Iter nub. 308 (BM, BR (2), E, G
(2), G-DC, GOET, K, L, M [“400”], M [“401”], M [“782”], MO, P [P05620478],
P [P05620484], P [P05620578], PRC, S [S08-12152], TUB [TUB004721], TUB
[TUB004722], W [W 0011066], W (2), WAG [WAG0234161]). Tanzania.
Tanga: Tanga town, near public beach, 200 m E of 05°03'19.2"S, 39°07'31.3"E,
N. Holstein et al. 94 (B, DSM, M, WAG), and 95 (DSM, M). Zanzibar Urban/
West: Chukwani, H.G. Faulkner 2782 (BR, COI, K n.v., P [P05620529]). ailand.
Bangkok: Bangkok, R. Zimmermann 66 (B (2), BR [BR0000009938878], G (2),
H, L [L 0587657], M, MO, P [P06394565], PR [PR 801377], PRC, S, U, W).
USA. Florida: along Indian River [lagoon], Coco [Cocoa], 9 May 1918, J.K. Small
s.n. (FLAS [FLAS 29152], G, MO [acc.no. 1161454], US). Hawaii: O’ahu Island,
Dole St, behind U Hawaii Manoa dorms, B. Kennedy 47 (MO, US [00674433]).
Midway Atoll: Sand Island, 4208 Commodore Ave., F. Starr & K. Martz 080601-01
(BISH, PTBG [PTBG1000013099]). Vietnam. Da Nang: Tourane [Da Nang] and
vicinity, no detailed location, J. Clemens & M.S. Clemens 3257 (A n.v., G, MO, P
[P06394668], U, W, Z). Yemen. Al-Hudaydah: Mor [Mawr], 15°40'N, 28–30 Mar
1825, F.W. Hemprich s.n. (P [P06394550], S [S08-12138]).
Monograph of Coccinia (Cucurbitaceae) 89
Figure 28. Reconstruction of the habit of a female raceme of Coccinia heterophylla based on T.-J. Klaine
414 (P). Black bar equals 0.5 cm.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
90
6. Coccinia heterophylla (Hook.f.) Holstein, Kew Bull. 65(3): 435–441. 2010
[published in 2011].
Physedra heterophylla Hook.f. in D.Oliv., F. T. A. 2: 553. 1871.
Type: Angola. [Cuanza Norte]: Golungo Alto, along the banks of the stream Casaballa,
at the base of the mountains in Sobato de Bumba, male, , Oct 1855, F.M.J. Wel-
witsch 791 (Lectotype: BM! [BM000948006, digital image: BM, JPS], selected in
Holstein and Renner (2010: 440); isolectotype p.p.: LISU! [LISU214547, digital
image: JPS]).
Type: Angola. No detailed location, female, fr, Jan 1856, F.M.J. Welwitsch 791 (Syn-
type: BM! [BM000948008, digital image: BM, JPS]).
Type: Angola. In rugged places at Delamboa river, with Coea melanocarpa, no clear
date given, F.M.J. Welwitsch 791 (Syntype: BM! [BM000948007, digital image:
BM, JPS]).
Type: Angola. No detailed location and date, F.M.J. Welwitsch 791 (Syntype: COI!
[COI00005515, digital image: JPS]).
Type: Angola. No detailed location, male, , no date, F.M.J. Welwitsch 791 (Syntype: K!
[K000313234, digital image: JPS]).
Type: Angola. No detailed location, female, , no date, F.M.J. Welwitsch 791 (Syntype:
LISU! [LISU214548, digital image: JPS]).
Type: Angola. No detailed location, female, fr, Jan 1856, F.M.J. Welwitsch 791 (Syn-
type: LISU! [LISU214549, digital image: JPS]).
Type: Angola. No detailed location, male, , no date, F.M.J. Welwitsch 791 (Syntype:
LISU! [LISU214550, digital image: JPS]).
Type: Angola. In rugged places at Delamboa river, with Coea melanocarpa, Sep 1855,
F.M.J. Welwitsch 791 (Syntype: LISU! [LISU214551, digital image: JPS]).
Type: Angola. At Delamboa river, no date, F.M.J. Welwitsch 791 (Syntype: LISU!
[LISU214552, digital image: JPS]).
Type: Angola. No detailed location and date, F.M.J. Welwitsch 791 (Syntype: LISU!
[LISU214553, digital image: JPS]).
Type: Angola. No detailed location and date, F.M.J. Welwitsch 791 (Syntype: P! [digital
image: JPS]).
Type: Angola. No detailed location and date, F.M.J. Welwitsch 791 (Syntype: G-DC!).
Physedra heterophylla var. hookeri Hiern in Cat. Welw. Afr. Pl. 1(2): 400. 1898.
Type: Angola. [Cuanza Norte]: Golungo Alto, near Ponte de Felix Simões, female,
, Dec 1855, F.M.J. Welwitsch 792 (Holotype: BM! [BM000948009, digital
image: BM]).
Description. Perennial climber. Stem up to 5–6 m, glabrous, sometimes whitish-speckled.
Petiole 1–2 cm long, puberulous on abaxial side with few tiny trichomes, rarely with up to
0.8 mm long, yellowish trichomes, sometimes white-speckled, on adaxial side with small,
yellowish-dirty trichomes. Leaves 7.5–12.5 × 10–14 cm, cordate, deltoid-subhastate to
Monograph of Coccinia (Cucurbitaceae) 91
5-lobate, auriculate, or 7-lobate. Lobes triangulate to ovoid. Leaf margin dentate, often
serrate. Apex acute with nal tooth to acuminate. Upper leaf surface glabrous, small hya-
Figure 29. Distribution map of C. heterophylla (based on 19 collections).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
92
line pustulate. Lower leaf surface glabrous often with blackish glands, nerves glabrous, ex-
cept for basis with up to 0.8 mm long, yellowish trichomes and sometimes white-speckled.
Probracts up to 5 mm. Tendrils bid. Male owers in short racemes. Common peduncle
2–13 mm long, not exceeding the pedicel bearing part in length, puberulous with tiny
yellowish-dirty trichomes (magnifying glass!). Pedicels < 4 mm, glabrous to puberulous.
Bracts up to 3.5 mm. Perianth tube glabrous. Calyx lobes 5–7 mm, subulate, erect. Co-
rolla up to 1.6 cm, dirty yellowish, dirty orange to reddish-orange. Corolla lobes 4–7 mm.
Color of lament column, anther head, or pollen sacs not seen. Female owers solitary or
in short or long (up to 15 cm) racemes (Fig. 28). Raceme and bracts as in males, except
raceme can be elongated (see remarks). Hypanthium glabrous, calyx lobes and corolla
like male owers. Ovary glabrous. Style columnar to 3-parted to the middle, color not
seen. Stigma bulging, pale yellow. Fruits up to 4–6 × 2–2.5 cm, ovoid-elliptical to shortly
cylindrical, glabrous. Fruits ripen from green? via orange-colored with longitudinal green
bands, ripe fruits unknown. Seed size not known, rather symmetrically obovate, face at.
Phenology. Flowering time: February–April, June, September–December.
Distribution. Fig. 29. Western Gabon (Estuaire), southern R. Congo (Kouilou),
western D. R. Congo (Bas-Congo), western Angola (Cuanza Norte, Namibe). Eleva-
tion 10 to 900 m. Soil preference unknown. Transition between tropical lowland
rainforest and woodlands, and its relict sites along the Angola highland escarpment, in
secondary regrowths, on shrubs, along rivers.
Remarks. e long, subulate calyx lobes are the only good character for distin-
guishing this species, which otherwise can be easily confused with C.barteri. Female
collections from Libreville (Gabon) and R. Congo have elongated racemes while the
racemes are more condensed in the south. Whether this character shows anity (intro-
gression?) to C.racemiora, which also has elongated racemes, denes an own species,
or is just a coincidental observation of intraspecic variation is not known.
Taxonomic remark. is species is the type species of Physedra Hook.f. e
genus was described by J. D. Hooker (Bentham and Hooker 1867) with three
species belonging to it. However, in Oliver’s Flora of Tropical Africa (Hooker
1871), Hooker only describes two species, P. heterophylla and P. longipes. When
Jerey (1962) transferred P.longipes into a new genus, Ruthalicia, he indirectly
lectotypied Physedra.
Monique Kéraudren regarded P. heterophylla as synonymous to C.barteri (Kéra-
udren-Aymonin 1975a; Kéraudren 1967) from which it, in fact, diers by the long
calyx lobes. is, however, led to the creation of C.subhastata with short calyx lobes,
which is a synonym to C.barteri (Holstein and Renner 2010).
Specimens examined. (Selection; in total: 37) Angola. Cabinda: Maiombe [Forest],
Belize, J. Gossweiler 7653 (COI, K, LISU). Cuanza Norte: Cazengo municipality, near
Agricultural Station Cazengo, J. Gossweiler 5655 (COI, LISU [LISC 031407], LISC
[LISC 031408], LISC [LISC 031409], LISC [LISC031410], LISC [LISC 031411],
LISU]), and 5707 (COI, LISC [LISC 031412], LISU). Huíla: Châo da Chela, between
[Lago] Tchivinguiro and Bruco, on middle slope of Serra da Chela escarpment, E.J.
Mendes 925 (BM, COI, LISC); ibid. L.A. Grandvaux Barbosa 9448 (COI). D. R.
Monograph of Coccinia (Cucurbitaceae) 93
Congo. Bas-Congo: Lukula territoire, Temvo, F.M.C.Vermoesen 1824 (BR, EA, WAG
[WAG0225504]). R. Congo. Kouilou: on left bank of Kouilou river, 4 km upstream
of Kakamoeka, Sounda on path to level meter, C. Farron 4980 (P [P05621154],
P [P05621155], P [P05621156]). Gabon. Estuaire: Libreville, T.-J. Klaine 414 (P
[P05620605], P [P05620610], P [P05621146], P [P05621147], P [P05621149], P
[P05621150], P [P05621152], P [P05621153]); Libreville, I.R.A.F. building, 0°25'N,
9°26'E, J.M. Reitsma & B. Reitsma 2120 (MO, NY, WAG [WAG0225493]).
7. Coccinia hirtella Cogn. in Schinz, Bull. Herb. Boiss. 4: 821. 1896.
Type: [South Africa]. KwaZulu-Natal: Howick, 3400’, male, , 18 Feb 1895, R. Schlech-
ter 6775 (Lectotype, designated by Meeuse (1962: 101): Z! [Z-000004443, digital
image: Z]; isolectotypes: BR! p.p. [BR0000008887184, digital image: BR, see tax-
onomic remarks], GRA [GRA0002851-0, digital image: JPS], K! [K000542639,
digital image: K]).
Description. Perennial creeper or climber. Stems up to 3 m, densely covered with long
(> 0.5 mm), upright, whitish trichomes. Petioles 1.5–4 cm long, indumentum like
on stem. Leaves 3–10× 2.5–11 cm, 5-lobate. Lobes obovate or elliptical in outline,
rarely ovate. Margin lobulate or coarsely serrate, (at maturity pale brownish) dentate
(Fig.13a). If margin serrate then lobe tips acute, else often rounded. Upper and lower
leaf surface, esp. on the nerves, densely covered with long, upright, whitish trichomes.
Probracts up to 3 mm. Tendrils simple or bid. Male owers solitary, rarely in few-
owered raceme. Common peduncle 1.8–4.5 cm, with white trichomes, pedicel of
racemous owers 0.2–3cm, with white trichomes. Bracts up to 2 mm, caducous.
Pedicel of solitary owers 3–8.5 cm, with white trichomes. Perianth tube with white
trichomes. Calyx lobes narrow triangulate to lineal, 3.5–6.5 mm long, spreading.
Corolla 2–3.7cm long, bu, lobes 1–2 cm. Filament column white, anther head pale
yellowish, pollen sacs yellow to orange. Female owers one solitary. Pedicel 0.5–5.5cm,
with long, white trichomes. Hypanthium with white trichomes, calyx lobes and corolla
like in male owers. Style columnar, whitish. Stigma bulging, yellow. Ovary with
some long trichomes. Fruit oblong ovoid, 5–6 cm × 2.5–3 cm, sparsely covered with
long trichomes to glabrous, ripening from green via green with longitudinal whitish
mottling, via yellow, orange to red when ripe (Fig.13a). Seeds 5.5–6.5 × 3–3.5 mm
(L/W), seed height not seen, rather symmetrically obovate, face atly lenticular.
Phenology. Flowering time: January–April, September, November, December.
Distribution. Fig. 30. Lesotho, South Africa (S KwaZulu-Natal, SE Free State).
Afromontane scrubland and sourveld grassland. Elevation 110–1900 m. On sand,
sandstone, loam, well-drained soils, full sun, grazing maybe tolerated.
Use. It is said that if a Masuto dreams unpleasantly about an ancestor (“balimo”),
then relief is given after a bath with sun-dried C.hirtella roots and ironstone in a hole
on the threshold of the “lelopa” (the circular fence around the hut) (Phillips 1933).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
94
Vernacular names. Sotho: leraka-la-balimo (Jacot Guillarmod 1971; Phillips
1933), Sotho: monyaku (Jacot Guillarmod 1971).
Taxonomic remarks. e BR type specimen (BR0000008887184) contains two la-
bels and is mixed. e female parts on the sheets have most likely been detached from the
lectotype in Z, because it is a female branch with shallowly lobate leaves, just as part of
the lectotype. e male parts on the BR specimen, however, are mixed. e leaves with
the obtuse lobules are also likely type material, whereas the leaf with the acute lobules is
very similar to the leaves of the R. Schlechter 6708 collection, which is not a type.
Specimens examined. (Selection; in total: 36). Lesotho. Leribe: Léribé [Hlot-
se], H. Dieterlen & A. Dieterlen 145 (BR [BR0000008887467], P [P05620661], P
Figure 30. Distribution map of C. hirtella (pale yellow dots; based on 22 collections), C. mackenii (blue
triangles; based on 50 collections), and C. quinqueloba (blue dots; based on 38 collections). For South
Africa the borders of the provinces are given.
Monograph of Coccinia (Cucurbitaceae) 95
[P05620662], Z). Maseru: Roma, Map[h]otong, M. Schmitz 8039 (PRE). South Af-
rica. Free State: Rooiberge area, Ross Kloof, M. Jacobs 8565 (L, LISU, PRE). Kwa-
Zulu-Natal: 17 km from Nottingham Road on road to Loteni, E. Retief 1638 (MO,
PRE); Albert Falls, A.K. Meebold 13160 (M); Giants Castle Game Reserve (Dinosaur
Footprint area), W.R. Trauseld 930 (PRE).
8. Coccinia intermedia Holstein, PhytoKeys 7: 28. 2011.
Type: Benin. Atakora: Natitingou, Kouaténa (Perma), 10°12.00'N; 1°30.18'E, riv-
er bed, female, , fr, 3 Oct 2000, A. Akoègninou et al. 3625 (Holotype: WAG!
[WAG0278370]; isotype: WAG [WAG0278369]).
Type: Ghana. Greater Accra: Shai Hills Game Reserve, monoecious, , fr, 25 May
1976, J.B. Hall & J.M. Lock GC 46016 (Paratypes: K! (4), MO!).
Type: Ivory Coast. Zanzan: Bouna, male, , 10 Aug 1967, C.Geerling & J.Bokdam
662 (Paratypes: MO!, WAG! [WAG0225492]).
Type: Togo. Maritime: between Lomé and Aného, female, fr, 25 Jun 1994, L. Aké Assi
18983 [typographical error in orig. publication stated 18982] (Paratype: MO!).
Description. Perennial climber. Stem length unknown, but likely several meters, gla-
brous, at maturity with clear to white pustules. Petioles 2.8–10.8 cm, glabrous, when
older with clear to white pustules. Leaves 6–15 × 7–18 cm, shallowly to profoundly
5-lobate, more or less auriculate. Margin conspicuously dentate, blackening when
dried. Apex acute. Upper leaf surface glabrous with clear to whitish pustules. Lower
leaf surface glabrous, often with small dark glands near the leaf base. Probracts up to
2.5 mm long. Tendrils simple or bid. Male owers in few-owered racemes, likely
sometimes accompanied by a single ower. Common peduncle up to 1 cm, pedicels
in racemose owers 2–4 mm, each glabrous. Bracts up to 1.5 mm. Perianth tube
glabrous, calyx lobes c.1.5 mm, lineal to narrowly triangulate, erect with slightly
recurved tips. Corolla 1.6 cm long, pale reddish-yellow to yellow, lobes 0.7 cm. Fila-
ment column and anther head not seen, pollen sacs yellowish. Female owers 1–3
clustered (strongly reduced raceme). Pedicels 0.6–1.2 cm, glabrous. Hypanthium
tube glabrous, calyx lobes and corolla like in male owers. Ovary glabrous. Style and
stigmas not seen. Fruit 4.5 × 2.5 cm, elliptical to oblong. Unripe fruit green with
pale green longitudinal mottling, ripe orange?, more likely becoming red via orange
ripening stage. Size of mature seeds unknown (≥ 5.5 × 3.5 × 1.3 mm (L/W/H)),
symmetrically (to slightly asymmetrically) obovate, face at.
Phenology. Flowering time: May, August, October.
Distribution. Fig. 31. NE Ivory Coast, SE Ghana (likely also in the north), S
Togo (likely also in the north), NW Benin. Elevation sea level to 415 m. Wooded
grasslands (semi-humid savanna), woodlands, dry forests, in riverbeds.
Remarks. is species is rather cryptic and imperfectly known. e leaves seem
to develop conspicuous margin teeth during maturity, like e.g., C.grandis, but the
darkish sublaminal glands dier from that species. e erect calyx lobes with slightly
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
96
recurved tips appear to be the most indicative character for C.intermedia. e clus-
tered female owers and the fruits link to C.barteri, from which it, among other
characters, diers in ecology. Two J.B. Hall & J.M. Lock GC 46016 specimens from
K have male and female owers/fruits on one twig and are thus monoecious. As all
other Coccinia intermedia collections are dioecious, this could be a case of “leaky
dioecy” (Baker and Cox 1984), which also has been observed in other Cucurbitaceae
(Schaefer and Renner 2010).
9. Coccinia keayana R.Fern., Bol. Soc.Brot. ser. 2, 33: 191. 1959.
Type: Guinea-Bissau. Tombali: region Cacine, rainforest, , Aug 1933, J.V.G. do Es-
pírito Santo 603 (Holotype: COI).
Type: ibid., J.V.G. do Espírito Santo 631 (Paratypes: LISC [LISC 011640, digital im-
age: IICT, JPS], LISC! [LISC 011641, digital image: IICT, JPS], LISC [LISC
011642, digital image: IICT, JPS], LISJC).
Type: Guinea-Bissau. Tombali: between Cacine and Guileje, plantation, 1 Aug 1945,
J.V.G. do Espírito Santo 2151 (Paratypes: COI, LISC [LISC 002508, digital image:
IICT, JPS], LISC! [LISC 002509, digital image: IICT, JPS]).
Type: Liberia. [Margibi County]: Firestone plantation, at Du River, 29 Jul 1926, D.H.
Linder 121 (Paratype: K!).
Type: Sierra Leone. Jigaya, c.350m, 28Sep 1914, W. omas 2844 (Paratype: K!).
Type: Sierra Leone. [Northern Province]: Bumban National Park, 30 Aug 1928,
F.C.Deighton 1221 (Paratype: K!).
Type: Sierra Leone. [Southern Province]: Moyamba District, Moyamba, 25 Aug 1931,
F.C.Deighton 2217 (Paratype:K!).
Coccinia sp. A Keay, Fl. W. Trop. Afr. 1: 216. 1954.
Figure 31. Distribution map of C. intermedia (pale yellow dots; based on 4 collections), C. keayana (blue
dots; based on 23 collections), and C. longicarpa (blue triangles; based on 22 collections).
Monograph of Coccinia (Cucurbitaceae) 97
Description. Perennial? climber. Stem up to 5 m, glabrous. Petiole 1.5–5 cm, with short,
few-cellular trichomes on adaxial side, glabrous on abaxial side. Leaves 5–11 × 3.5–11
cm, (shallowly to) profoundly 3-(or 5-)lobate, auriculate, rarely long cordate. Margin
rather remotely dentate to slightly serrate. Lobe apex acute or subacute with nal tooth.
Upper leaf surface tiny hyaline pustulate. Lower leaf surface with blackish glands, dried
often with bluish-green tinge, glabrous or rarely with soft multicellular trichomes on
nerves. Probracts up to 3 mm. Tendrils simple. Male owers ebracteate, in lax racemes
with up to 20 owers, sometimes accompanied by a solitary ower (Fig. 32). Common
peduncle up to 1.7 cm, shorter than racemous part, glabrous. Pedicels of racemous ow-
ers up to 1 cm, pedicels of solitary owers up to 1.5 cm long, each glabrous. Perianth
tube glabrous. Calyx lobes linear, 2.5–3 mm, in buds spreading, later reexed. Corolla
1.7–2 cm long, white, yellow, dirty orange, salmon to dull pinkish. Corolla lobes 3–5
mm long. Filament column and anther head not seen, pollen sacs pink–orange. Female
owers solitary or in few-owered lax racemes. Common peduncle 1.2–2.1 cm, gla-
brous. Pedicel owers in racemes up to 1 cm, glabrous, pedicels of solitary owers up
to 2.7cm, glabrous. Hypanthium glabrous, calyx lobes and corolla as in male owers.
Ovary glabrous. Style and stigma not seen. Fruits 2–3 × 2 cm, subglobose to globose, un-
Figure 32. Reconstruction of the habit of Coccinia keayana based on C.C.H. Jongkind et al. 6542
(WAG). Black bar equals 1 cm.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
98
ripe glaucous green, ripening via yellow to pinkish [rather glaucous?] red. Seeds 4.5×2.5
× ? mm (L/W/H), rather symmetrically obovate, face atly lenticular.
Phenology. Flowering time: March to November.
Distribution. Fig. 31. Tropical West Africa: Guinea-Bissau (Tombali), Sierra
Leone, Liberia, Guinea (Nzérékoré), S Ivory Coast, S Ghana, S Togo? Elevation sea
level to 1250 m. On sandy soil, lateritic soils. Tropical rainforest, in high trees, high
bushes, on roadsides.
Vernacular names. Cf. Koranko: nala (W. omas 2844), Limba: ngolibwe (W. omas
2844), Mende: ndogbo-gojai (F.C.Deighton 2217), Temne: efosa (W. omas 2844).
Remarks. e long racemes with ebracteate owers and the linear, reexed calyx
lobes are good characters for this species. It is barely distinguishable from C.barteri
without owers. Coccinia keayana collections often have a bluish green tinge and the
lobes conspicuously point forwards (see Fig. 32), which might only rarely occur in
C.barteri. e corolla is rather tubular, sometimes somewhat inated.
Taxonomic remarks. e name C.keayana R.Fern. is misapplied for the Flora of
Cameroon or western Central Africa in general (for details, see taxonomical remarks
of Coccinia barteri).
Specimens examined. (Selection; in total: 33) Ghana. Eastern Region: Asiakwa
district, Atewa Range Forest Reserve, Accra-Kumasi highway 5–6 km along forest road
that intersects the highway at Sagyimase village, 6°13'48"N, 0°32'42"W, M. Merello
et al. 1179 (MO). Volta: Amedzofe, J.B. Hall GC40053 (P [P05620653]). Western
Region: Bia National Park, J.B. Hall & J.M. Lock GC46493 (WAG [WAG0046501]).
Guinea. Nzérékoré: Nzérékoré Préfecture, [WSW of] Nzérékoré, 7°43'35.54"N,
8°51'21.28"W, E. Achigan-Dako 07 NIA 917 (GAT). Guinea-Bissau. Tombali:
Bedanda sector, Cantanhez, J. Alves Pereira 3172 (H, LISC). Ivory Coast. Lacs:
Oroumba Boca [Orombo Bocca Mt.], H.C.D. de Wit 5772 (WAG [WAG0234139],
WAG [WAG0234140]). Lagunes: Abidjan, Banco Forest Reserve, W.J.J.O. de Wilde
893 (BR, EA, WAG [WAG0044624], WAG [WAG0044625], Z). Liberia. Bong: 3
miles NE of Suacoco, Z.D. Traub 256 (BR, G, MO). Nimba: Yéképa, Mt Nimba, J.-
G. Adam 21232 (MO, P [P00694038/P05590407], P [P05620652], PRE).
10. Coccinia longicarpa Jongkind, Blumea 49: 83. 2004.
Type: Ivory Coast. Lagunes: Forêt du Banco, S of Arboretum, near river, male and fe-
male, fr, 20 Jul 1973, J. de Koning 1965 (Holotype: WAG! [WAG0099441, digital
image: JPS], isotype: K! [K000035540, digital image: K]).
Type: Ivory Coast. Lagunes: Forêt du Banco, Route Martineau, secondary forest, 10
Oct 1974, J. de Koning 4077 (Paratype: WAG! [WAG0099430]).
Type: Ivory Coast. Lagunes: Forêt du Banco, near swampy secondary forest, 8 Aug 1975,
J. de Koning 5904 (Paratypes: K [K000035541], MO!, WAG! [WAG0099438,
digital image: JPS]).
Monograph of Coccinia (Cucurbitaceae) 99
Type: Ivory Coast. Lagunes: Forêt du Banco, N of center, near Banco river, in for-
est clearing on clear spot, 16 Jun 1975, W.J. van der Burg 551 (Paratype: WAG!
[WAG0062627]).
Type: Ivory Coast. Lagunes: near Abidjan, 6 Sep 1967, C.Geerling & J. Bokdam 829
(Paratypes: WAG! [WAG0011282], WAG! [WAG0011283]).
Type: Ivory Coast. Lagunes: [W of Abidjan], Adiopodomé [Adiopo-Doumé],
margin of bush pathway, 3 Aug 1956, J.J.F.E. de Wilde 183 (Paratypes: WAG!
[WAG0044616], WAG! [WAG0044617], WAG! [WAG0044618], WAG!
[WAG0044619]).
Type: Ghana. Eastern Region: near Kibi, Atewa Range Forest Reserve, Jun 1976, J.M.
Lock GC 43991 (Paratype: K).
Type: Liberia. Gbapolu/Lofa: Gbanga, Sep 1926, D.H. Linder 576 (Paratype: K).
Type: Ghana. Nsuta, no detailed location, 1500 ft, May 1929, C.Vigne 1735 (Para-
types: K, P!).
Coccinia sp. A C.Jerey, Key to the Cucurbitaceae of West Tropical Africa. J. W. Afri-
can Sci. Assoc.9: 87 p.p., 1964.
Coccinia sp. B Keay, Fl. W. Trop. Afr. 1(1): 216 p.p., 1954. Nigeria. [Ogun]: Ijebu-Ode
District, toward the head of extraction Rd. from Grace Camp [in Omo Forest Re-
serve] westwards toward the R. Omo [Shasha], about 1.5 mls from the Grace Camp, 2
May 1946, S. Tamajong FHI 16938 (K!); D.H. Linder 576 (K); C.Vigne 1735 (K, P!).
Coccinia sp. D Keay, Fl. W. Trop. Afr. 1(1): 216 p.p., 1954. Nigeria. Oyo: Lagos
[Colony], Ibadan forest, 1 Dec 1900, C.Punch 46 (K!).
Description. Perennial climber. Stems up to 5 m, glabrous. Petiole 0.9–4.5 cm,
glabrous. Leaves 7–12 cm × 2.5–11 cm, 5-angularly subcordate, subhastate, rarely
3-lobate, auriculate. Lobes elliptical. Margin suspiciously dentate, whitish in living
state, blackening when dried. Apex acute with nal tooth. Upper leaf surface pale to
white pustulate. Lower leaf surface glabrous, nerves often white-speckled. Probracts <
1 mm, often missing. Tendrils simple, rarely bid. Male owers ebracteate, in glabrous
racemes, occasionally accompanied a solitary ower. Common peduncle 0.7–1.5 cm,
pedicel of owers in raceme 2.5–8 mm, pedicel of solitary ower 5–10 mm. Perianth
tube glabrous, calyx lobes 2–3 mm long, (0.75–)1.2–1.8mm broad at base, erect.
Corolla 0.8–1.5 cm long, yellow to (pale) orange, sometimes with greenish nerves.
Corolla lobes up to 2 mm long. Filament column and anther head not seen, pollen sacs
yellow to orange. Female ower solitary. Petiole 1.5–2.2cm, glabrous. Hypanthium
glabrous, calyx lobes and corolla not seen, but likely as in male plants. Ovary glabrous.
Style and stigmas not seen. Fruit up to 20 cm long, c.1 cm in diam., long cylindrical,
unripe waxy green, ripe unknown. Seed size and shape unknown, face at.
Phenology. Flowering time: May–December.
Distribution. Fig. 31. Tropical West Africa: SW Ivory Coast, S Ghana, SE Benin,
SW Nigeria. Elevation sea level to 460 m. Soil preference not known. In primary or
secondary forests, in disturbed places (roadsides, near rivers).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
100
Remarks. e broad calyx lobes are, apart from the long cylindrical fruit, the best
character for identifying this species. An urceolate corolla (Jongkind 2004) also occurs
in C.barteri.
A single collection (H.J. Beentje 602 from M) mentions a lilac corolla color, which
would be unique in Coccinia. Although this might be possible, since there are also
pinkish owers reported in C.adoensis, the fact that the WAG duplicate with the same
collection number is a Ruthalicia makes it more likely that the observation is due to a
mixed collection, eventually from a Convolvulaceae.
Specimens examined. (Selection; in total: 24) Benin. Plateau: Pobe,
6°57'52.56"N, 2°40'19.70"E, E. Achigan-Dako 07 NIA 731 (GAT). Ghana.
Ashanti: Bobiri Forest Reserve, 6°41'N, 1°21'W, C.C.H. Jongkind 3970 (WAG
[WAG0023747], WAG [WAG0023748]). Central Region: Twifo/Hemang/Low-
er Denkyira District, Kakum, 5°20'54.31"N, 1°23'1.39"W, E. Achigan-Dako 07
NIA 734 (GAT). Eastern Region: Atewa Range Forest Reserve, 06°15'N, 00°33'W,
C.C.H. Jongkind et al. 1538 (MO, WAG [WAG0020265]). Western Region: Jo-
moro District, Fawoman, 5°19'32.63"N, 2°43'28.13"W, E. Achigan-Dako 06 NIA
050 (GAT); ibid., E. Achigan-Dako 06 NIA 051 (GAT). Ivory Coast. Lagunes: Abid-
jan. Banco Forest Reserve, J.J. Wieringa 5386 (WAG [WAG015997]). Sud-Comoé:
Aboisso, 5 km NNW of Nganda-Nganda [Forest Reserve], near lagoon Aghien [la-
gune Aby?, because lagune Aguien is NE of Abidjan], 5°14'N, 3°20'W, H.J. Beentje
602 (M, but not WAG). Nigeria. Ogun: Ijebu-Ode District, Omo Forest Reserve,
Compartment 8, J.R. Charter FHI 38635 (K).
11. Coccinia mackenii Naudin ex C.Huber [sphalm. Mac-Kennii, after John
M’Ken, ICN 60C.5], Cat. Print.: 5. 1865.
Cephalandra mackenii (Naudin ex C.Huber) Naudin [sphalm. mac kennii], Ann. Sci.
Nat. Bot. 5: 17, ser. 5. 1866.
Type: Cultivated. Cultivated in Paris Botanical Garden from seeds from Huber’s Gar-
den in Olbia [Hyères, France] who obtained the seeds from M’Ken from near Port
Natal [Durban, KwaZulu-Natal, South Africa], female, , 1864, C.V. Naudin s.n.
(Syntypes: P!, G-DC! [G00211343, digital image: G]), P [P06745731, digital im-
age: P], P [P06745732, digital image: P].
Type: Cultivated. Cultivated in Paris Botanical Garden from seeds from Huber’s Gar-
den in Olbia [Hyères, France] who obtained the seeds from M’Ken from near
Port Natal [Durban, KwaZulu-Natal, South Africa], 1863, C.V. Naudin s.n. (Syn-
types: P! [P06745737, digital image: P], P! [P06745739, digital image: P], P
[P06745740, digital image: P]).
Type: Cultivated. Cultivated in Huber’s Garden in Olbia [Hyères, France] who ob-
tained the seeds from M’Ken from near Port Natal [Durban, KwaZulu-Natal,
South Africa], male and female, , 1864, C.V. Naudin s.n. (Lectotype, designated
here: P! [P06745735, digital image: P; K neg. 2993]; isolectotypes: G-DC! [3
Monograph of Coccinia (Cucurbitaceae) 101
sheets, all G00211344, digital images: G], K! [K000542637, digital image: K],
K! [K000542638, digital image: K], K (2)!, P! [P06745730, digital image: P], P!
[P06745733, digital image: P]).
Coccinia palmata (Sond.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 540. 1881.
Nom. illeg. Momordica palmata E.Mey. ex Drège, Zwei panzengeogr. Dokum.
[addition to Flora 26(2)]: 156, 159, 202. 1843. Nom. nud. Cephalandra palmata
Sond. in Harv. & Sond., Fl. Cap. 2: 493. 1862.
Type: [South Africa]. [KwaZulu-Natal]: near Port Natal [Durban; cited in l.c.p. 159], male
and female, , fr, 7 Apr 1832, J.F. Drège s.n. (Lectotype, designated by Meeuse (1962:
96): S! [S08-12155, digital image: S], isolectotype: P! [P00748835, digital image: P]).
Type: [South Africa]. Without location and date, male, , J.F. Drège s.n. (Syntypes: G!
[G00226835, digital image: G], HBG! [HBG506427], K! [K000313198, digital
image: K], K! [K000313199, digital image: K], L!, P! [P00748833, digital image:
P], P!, PRC!, W! [W 0026939, digital image: WU]).
Type: [South Africa]. [KwaZulu-Natal]: Omsamculo [Umzimkulu], between shrubs
and thickets, near river mouth, female, fr, 5 Mar 1832, J.F. Drège 4637 (Syntype:
P! [P00748834, digital image: P])
Coccinia dinteri André, Rev. hort. [Paris] 72: 276. 1900.
Type: Unnumbered plate in l.c.
Description. Perennial climber or creeper. Stems up to 9.5 m, glabrous. Petioles
0.7–11 cm long, glabrous or with thin trichomes. Leaves 3–13.5 × 3–15.5 cm,
shallowly to profoundly 5-lobate, in the latter case often weakly lobulate. Lobes
triangulate, lanceolate, ovate to obovate. Margin smooth, dentate, sometimes ser-
rate to lobulate, esp. towards the apex. Apex acute with nal tooth. Upper leaf
surface glabrous with clear to white pustules, rarely with few trichomes. Lower leaf
surface glabrous or with thin, sti or articulate trichomes, towards base usually with
glands. Probracts up to 4 mm, oblong-lanceolate. Tendrils bid, rarely simple. Male
owers solitary or ebracteate in few-owered racemes. Common peduncle 5–6.5
cm, pedicel of ower in raceme up to 2.5 cm, pedicel of solitary owers 6–9 cm, all
glabrous, rarely with long trichomes. Perianth tube glabrous. Calyx lobes 1.5–6.5
mm, lineal, subulate to narrowly triangulate, when young erect, later spreading to
reexed. Corolla 1.3–2.7 cm long, cream to pale bu, corolla lobes subulate to tri-
angulate, 0.7–1.1 cm. Filament column, anther head, and pollen sacs not seen. Fe-
male owers one solitary. Pedicel 0.7–5 cm long, glabrous. Hypanthium glabrous,
calyx and corolla like in male owers. Ovary glabrous. Style columnar, color not
seen. Stigma bulging, color not seen. Fruits elliptical to oblong, c.10 × 2–2.5 cm.
Unripe green with white mottling, ripe red-orange to red, sometimes? with white
mottling. Seeds 6–7 × 4–4.5 × 1.5 mm (L/W/H), slightly asymmetrically obovate,
face atly lenticular.
Phenology. Flowering time: January–April, July, November, December.
Distribution. Fig. 30. South Africa (E Eastern Cape, KwaZulu-Natal, Mpuma-
langa, Limpopo), Swaziland. Elevation sea level to 1750 m. Clay, Berea red sand,
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
102
sandstone, quartzite, poorly drained soils. Afromontane forests, coastal forests, littoral
forests, forest margins, sometimes grassland. In frost-free areas (Meeuse 1962).
Use. Leaves and fruits are eaten by Tsonga people (Shackleton et al. 1998).
Vernacular names. Xitsonga: Gomo, XipapaXipapana (Shackleton et al. 1998).
Remarks. Some collections with deeply lobate leaves and short petioles resemble
the closely related C.quinqueloba, and some C.quinqueloba individuals have long peti-
oles (C.V. Naudin s.n. 1863, C.V. Naudin s.n. 1863–1865, E. Retief 1215). However,
Naudin (1866) reports considerable problems with seed production in interspecic
crosses. It would be desirable to validate this observation.
Taxonomic remarks. e initial designation of the C.mackenii lectotype (Hol-
stein and Renner 2010: 440) is not valid, because it erroneously designated a female
specimen from Paris Botanical Garden. However, Naudin stated that all plants from
Paris Botanical Garden were male (P06745733), so the former designation was am-
biguous. e new lectotype was chosen from Olbia [Hyères] material. In contrast
to Naudin’s statement, that Olbia material was female, there is a male K specimen
(K000542638), from Huber’s Garden in Olbia. Eventually, this specimen is from
Paris Botanical Garden but incorrectly labeled, because the lack of the opposite sex
aected Naudin’s crossing experiments.
Due to an overlooked published combination C.mackenii bore the illegitimate name
C.palmata for more than 120 years. When Wight and Arnott published Coccinia in-
dica they also included a specimen tentatively identied as Bryonia palmata L. Although
without relevance for the genus Coccinia itself, it lead to further complications. One year
after Voigt’s publication of the correct combination Coccinia grandis, Roemer (1846) also
recognized the apparently missing combination and that Linnaeus’ B. palmata and B.
grandis indeed referred to dierent species. Roemer treated them, amongst other species,
as C.grandis (L.) M.Roem. (nom. illeg.) and C.palmata (L.) M.Roem. In addition to the
name Coccinia palmata (L.) M.Roem. another species from South Africa was described with
the name Cephalandra palmata E.Mey. ex Sond. (Harvey and Sonder 1862). Cogniaux
(1881) accepted this species in Coccinia, overlooking Coccinia palmata (L.) M.Roem. He
thus created an illegitimate Coccinia palmata (E.Mey. ex Sond.) Cogn., which has since
been used for this species. Holstein and Renner (2010) called attention to this erroneous
usage by resurrecting the correct name, Coccinia mackenii Naudin ex C.Huber.
e drawing of Coccinia dinteri in the protologue shows a bid tendril. Since all
other characters match C.mackenii and the resemblance was already discussed in the
protologue, it is feasible to synonymize it with that species. M. Proschowsky grew this
plant in the Fabron quarter of Nice, France, but the origin of the seeds was not indi-
cated. e label named it “Coccinia dinteri” after Moritz Kurt Dinter (in the proto-
logue erroneously spelled as “Hurt Dinter”), who was curator in La Mortola (Giardini
Botanici Hanbury, Liguria, Italy) where many South African plants were cultivated.
Hence, it is reasonable to assume this origin as done by André there, which again would
match C.mackenii. ere is a specimen in K herbarium containing only seeds and a
label indicating that they were sent from Hanbury, La Mortola in 1897. A note men-
tions that the seeds were sown in Kew Gardens. e identication is given as Cephalan-
Monograph of Coccinia (Cucurbitaceae) 103
dramackenii with a question mark and a later note with the Coccinia dinteri citation.
It is plausible to assume that these seeds come from the same plant stock that was used
to grow and to describe C. dinteri. Although the seeds t the description of C. mackenii
seeds, it is not possible to use them to identify the species unambiguously.
Specimens examined. (Selection, in total: 71) South Africa. Eastern Cape: Port St.
Johns, Jan 1933, A.O.D. Mogg s.n. (L, PRE [PRE 42990], Z). Kwazulu-Natal: Durban
district, Isipingo North, C.J. Ward 3747 (COI, PRE); Umzinto district, Vernon Crook-
es Nature Reserve, far end of Golden Valley, K. Balkwill et al. 10930 (E [E00264193],
MO); Pietermaritzburg, Fernclie Forest, J. Bodenstein 92 (PRE); Nkandlha [Nkandla],
Qudeni Forest, 5 mls [8 km] S of Qudeni P.O., L.E.W. Codd 6991 (PRE). Limpopo:
near Lydenburg, near Echo Cave, R.G. Strey 3762 (M, PRE, WAG [WAG0234163]);
Woodbush [Forest Reserve], 6 Aug 1925, A.O.D. Mogg s.n. (COI, L, PRE [PRE 43066],
Z). Mpumalanga: Letaba district, E side of shoulder extending northwards from ridge
above Weltevreden, J.C.Scheepers 1110 (M, PRE). Swaziland. Hhohho: about 20 km
N of Mbabane, Ngwenya Hills, Castle peak, north slopes, B. Maguire 7553 (B [B 10
0019799], E). Manzini: Usutu Forests, R.H. Compton 32287 (PRE).
12. Coccinia megarrhiza C.Jerey, Kew Bull. 15(3): 347. 1962.
Type: Kenya. Northern Province [Eastern Province]: Moyale, 3800 ft, male, , 28 Apr
1952, J.B. Gillett 12967 (Holotype: K! [K000313235, digital image: JPS, K], iso-
types: B! [B 10 0154926, digital image: B, JPS], S! [S08-12479]).
Type: Kenya. Northern Province [North Eastern Province]: [western Mandera District],
Dandu, 2600 ft, 10 Apr 1952, J.B. Gillett 12759 (Paratype: K! [K000354130]).
Type: Kenya. Northern Province [North Eastern Province]: [western Mandera District],
Dandu, 2700 ft, 9 May 1952, J.B. Gillett 13122 (Paratypes: EA!, K! [K000354129]).
Type: Kenya. Northern Province [North Eastern Province]: [western Mandera Dis-
trict], Dandu, 3000 ft, 14 May 1952, J.B. Gillett 13191 (Paratypes: EA!, K!).
Type: Kenya. Northern Province [Eastern Province]: Moyale, 3200 ft, 3 Oct 1952, J.B.
Gillett 13986 (Paratype: K! [K000354128]).
Type: Kenya. Northern Province [Eastern Province]: Moyale, 3600 ft, 14 Oct 1952,
J.B. Gillett 14036 (Paratypes: B! [B 10 0154927, digital image: B, JPS], BR!, EA!,
K! [K000354131], S! [S08-12482]).
Type: Kenya. ibid., J.B. Gillett 14037 (Paratypes: BR! [BR0000008914033], K!
[K000354133], K! [K000354134], S! [S08-12481]).
Type: Kenya. ibid., J.B. Gillett 14038 (Paratypes: K! [K000354125], K! [K000354132]).
Type: Kenya. ibid., J.B. Gillett 14039 (Paratypes: K! [K000354126], S! [S08-12480]).
Description. Perennial climber or creeper. Stem up to 6 m, with long, whitish to
beigeish patent trichomes, which appear articulate when dried. Petioles 1.5–5.6
cm, indumentum as on stem (Fig. 9). Leaves 3–11 × 4.2–17 cm, reniform to 3- or
5-lobate. Margin dentate (teeth at maturity brownish, when dried blackening),
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
104
serrate to lobulate. Upper leaf surface glabrous with pale pustules or with short,
whitish to beigeish trichomes, lower leaf surface with indumentum as on stem,
rarely glabrous. Probracts up to 3 mm long. Tendrils simple. Male owers
clustered. Pedicel < 1.5 cm, indumentum as on stem. Perianth tube with long,
beigeish, upright trichomes that appear articulate when dried. Calyx lobes 2.5–4
mm, subulate to lineal, spreading. Corolla 1.2–1.3 cm, yellow to pale orange, lobes
4–6 mm. Filament column greenish, anther head pale greenish, pollen sacs orange-
yellow. Female owers 1(–2) solitary. Hypanthium with long, beigeish, upright
trichomes that appear articulate when dried, calyx lobes and corolla like in male
owers. Ovary green with whitish spots. Style columnar, green. Stigma bulging,
yellow (Fig. 11c). Fruit ovoid–ellipsoid, up to 6.5 cm long, unripe green with
longitudinal white mottling. During ripening mottling partly developing a dark
green corona (Fig. 13c). Ripe red (Fig. 12b). Seeds 7 × 4 × 1.5 mm (L/W/H),
asymmetrically obovate, face at.
Phenology. Flowering time: March–May, August–October.
Distribution. Fig. 20. Ethiopia (Oromia, Somali Region), Kenya (Eastern
Province, North Eastern Province). Elevation 800–1600[–2000] m. On granite
outcrops and red lateritic soils, Acacia-Commiphora bushland, Dracaena vegetation,
Balanites vegetation, Euphorbia candelabrum woodlands, dry Juniperus forest, graz-
ing is tolerated.
Remarks. Coccinia megarrhiza and C.abyssinica form a species complex. Distinc-
tion between these two species can be dicult, especially in young plants, when the
color of the marginal teeth of the leaf is not well developed. While the peduncle length
diers, the earlier appearing solitary owers can have the same length in both species.
e broad leaves with an emarginate, obtuse to cuspidate tip (C.megarrhiza) versus
rather long leaves with an acute tip (C.abyssinica) seems to be the best character. At
maturity, the teeth coloration in C.megarrhiza is also much more conspicuous than in
C.abyssinica. A phylogeographic analysis and crossing experiments would shed light
on the question, whether these are ecologically dierentiated forms or true species.
Plants from the mountains near Yebelo with very large leaves are almost glabrous and
occur, untypically, in dry Juniper “forests”. However, they have the typical cuspidate
to obtuse central lobes and bear the colored leaf margin teeth. As larger leaves are also
observed in high altitude individuals of C.microphylla, these forms might be regarded
as mast specimens.
Specimens examined. (Selection; in total: 29) Ethiopia. Oromia: 38 km S
of Neghelli [Negele Boran] on Wachelli road, J.W. Ash 814 (EA (2), K); Arero
(Meta-Gafersa), G. Cufodontis 273 (FT, W); Bombal ca. 40 km on the way to
Jijiga from Harar, T. Ebba 622 (K, WAG [WAG0285707]); c. 20 km NW of
Moyale on the road to Mega, just after the turn o to Tuqa (and Sololo in Kenya)
(3°39'N, 38°56'E), I. Friis et al. 8736 (K); c. 36 km from Harar to Jijiga and
then c.20 km to S, J.J.F.E. de Wilde 4793 (B, K, MO, WAG [WAG0285708],
WAG [WAG0285709]). Somali Region: 95 km from Negele of Filtu road, 5°00'N,
40°12'E, M.G. Gilbert & B.M.G. Jones 110 (K).
Monograph of Coccinia (Cucurbitaceae) 105
13. Coccinia microphylla Gilg, Bot. Jahrb. Syst. 34: 357. 1904.
Type: Tanzania. [Kilimanjaro]: at base of Pare Mountains, between Kiswani [Kisiwani]
and Maji ya Juu [Madji-ja-juu], mix of thornbush and wooded grassland [“gemi-
schte Dornbusch- und Obstgartensteppe”], 700 m, , Oct, A. Engler, Reise nach
Ostafrika 1587 (Syntype: B destroyed).
Type: Tanzania. [Kilimanjaro]: between Kihuiro [Kihurio] and Gonja, thornbush
steppe, , Oct, A. Engler, Reise nach Ostafrika 948 (Syntype: B destroyed).
Type: Kenya. Coast Province: near Mariakani, NW of Mombasa, male, , 15 Oct
1955, E. Milne-Redhead & P. Taylor 7104 (Neotype, designated here: LISU!, iso-
neotypes: B!, EA!)
Coccinia buikoensis Zimm., Die Cucurbitaceen 2: 177, 24, 51, 84, 96, 114, g. 17 I–III,
g. 63 II, g. 74 VII–XII, g. 81 XVI, XVII. 1922.
Type: Tanzania. [Tanga]: Lushoto District, [S of Pare Mts], [between Hedaru and
Mkomazi], near Buiko, steppe, male and female, , fr, Dec 1919, P.W.A. Zimmer-
mann G6595 (Holotype: B destroyed, lectotype designated by Jerey (1967: 68):
EA [EA000002132, digital image: JPS]).
Coccinia sp. C in C.Jerey, F. T. E. A.: 69. 1967. Kenya, Northern Province: Furroli, lava
plateau, semi-desert, Acacia-Commiphora shrub, on sand, female, , fr, 12 Sep 1952,
J.B. Gillett 13820 (B!, EA!, K!, P!, S! [S08-12180]) and J.B. Gillett 13826 (K!).
Description. Perennial creeper or climber. Stems up to 4 m, glabrous or more or less
densely covered with short, white trichomes, when older often densely white pustu-
late. Petiole 0.45–4 cm, with erect, often thick, when longer sometimes bent trichomes
that are sometimes soft spiny (<1 mm) or only wart-like. Leaves 0.7–7.5 × 1.1–12
cm wide, usually rather small, shallowly to deeply 3- or 5-lobate, sometimes lobulate,
rarely reniform. Lobes narrow to broadly triangulate to lanceolate. Upper leaf surface
more or less densely white pustulate, pustules sometimes with a short, thick trichome
(Fig. 2a). Lower leaf surface glabrous or more or less dense with often sti, pale yel-
lowish trichomes, sometimes with darker glands between nerves, nerves usually with
thick, erect trichomes, sometimes reduced to wart-like appearance. Leaf margin rather
remotely denticulate, usually with minute, bent trichomes. Apex acute to obtuse, with
nal tooth. Probract usually absent, if present then < 1.5 mm. Tendrils simple. Male
owers 1–2 solitary, if fasciculate or in few-owered racemes, then accompanied by 1
solitary ower. Common peduncle < 5 mm, glabrous or with minute to long, articulate
trichomes. Pedicel of solitary owers 2–7(–25 mm), pedicel of owers in inorescences
<5 mm, glabrous or especially at apex with long, yellowish, articulate trichomes. Peri-
anth tube usually with long, yellowish, articulate trichomes, rarely almost glabrous. Ca-
lyx lobes 1.5–4 mm, narrowly lanceolate to lineal, when young erect, later spreading to
reexed. Corolla 0.7–1.2 cm long, greenish white to yellowish-orange with darker green
veins, lobes 4–7 mm. Filament column and anther head pale greenish yellow. Pollen
sacs yellow. Female owers 1(–2) solitary (Fig. 2a). Pedicels up to 0.7 cm, glabrous or
with white trichomes. Ovary glabrous, with some articulate trichomes to densely wooly
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
106
Figure 33. Distribution map of C. microphylla (blue dots; based on 49 collections) and C. trilobata (pale
yellow dots; based on 51 collections).
with long (when dry articulate) trichomes. Style columnar, pale green. Stigmas bulging,
greenish yellow. Ripe fruit globose to shortly obovate, 1.8–2.5× 1.4–2.5 cm, glabrous
or with few articulate trichomes, unripe green sometimes with longitudinal, whitish
Monograph of Coccinia (Cucurbitaceae) 107
mottling (Fig. 2a), which obtains a dark green corona during ripening, when ripe bright
orange-red to red. Seeds 4.5–6 × 2–3 × 1–1.5 mm (L/W/H), asymmetrically obovate to
somewhat falcate, face attened.
Phenology. Flowering time: January, April, May, July, October–December.
Distribution. Fig. 33. NE Tanzania (Arusha, Kilimanjaro, Manyara, Tanga),
Kenya (Coast, Eastern, North Eastern, Rift Valley), S Ethiopia (S and C Oromia),
Somalia. 70–1300(–1600) m. Savanna, open Acacia-Commiphora bushland, degrad-
ed Combretum bushland, open grassland, cultivated land, roadsides. Red sand, dark
brownish-black soil. Limestone.
Vernacular names. [Akiek; Ogiek]: notoku (A.S. Vincent 29, A.S. Vincent 221),
Maa [Maasai language]: ndegegeya (A.S. Vincent 29), sikuni (Kiamba et al. KEFRI 112).
Remarks. Some collections have a mixed (not intermediate) phenotype with
C.trilobata: the calyx lobes are unusually long (up to 7 mm), which speaks for C.tri-
lobata, but the indumentum matches C.microphylla. However, these do not occur
in a single location, but are found in the Ndoto Mts (O. Kerfoot 2644), in Kiboko
(P. Kirika et al. 002/020/2011), and around Voi (M. Hucks 579, B. Verdcourt 3888).
Whether these are hybrids (F2 or later) or just a variety is not known. ese collections
also resemble C.megarrhiza, which occurs in northern Kenya and Ethiopia, however,
the indumentum does not match either.
Despite the epithet, the leaves can become quite large, especially at higher alti-
tudes. en, collections may resemble C.trilobata, which has a denser indumentum.
In dry low altitude areas, leaves and owers emerge quickly, e.g., soon after a rain
shower. e leaves are thus not well developed and small. Collections from more arid
locations tend to be smaller in many characters, but whether these represent a new spe-
cies is doubtful. ere are only few collections of the proposed species (C.sp. Burger
2947A, C.sp. Gilbert & Jones 129 (Jerey 1995)), but the natural scope of C.micro-
phylla is hardly assessed. is will not be resolved without a phylogeographic analysis
and more intensive collecting from Ethiopia.
Specimens examined. (Selection, in total: 72) Ethiopia. Oromia: 105 km on
road from Negelli [Negele] to Filtu, J.J.F.E. de Wilde & M.G. Gilbert 346 (K, WAG
[WAG0285710], WAG [WAG0285711], UPS). Kenya. Coast Province: near Mari-
akani, NW of Mombasa, E. Milne-Redhead & P. Taylor 7105 (B, EA, LISU). Eastern
Province: E side of Lake Rudolf, between Koobi Fora and Shin (hill), 3°57’–58'N,
36°12'–20'E, R.B. Faden & A. Evans 71/301 (EA, K). North Eastern Province: Wajir
District, Catholic Girl’s Town 2 km E of Wajir, J.B. Gillett 21273 (EA, K, WAG
[WAG0234120]). Rift Valley Province: Turkana District, I. Ohta 24 (EA). Somalia.
Togdheer: Malol [Mt Malool] near Sheikh, J.R. Ironside 5/73/31 (K). Tanzania. Arusha:
Monduli district, Longido division, SE of Longido, c.100–300 m from Arusha Mu-
nicipality, 2°43'14"S, 36°42'02"E, C.J. Kayombo & K. Kitaba 4242 (MO). Dodoma:
Tarangire National Park, Kalima Hill, S. Chuwa et al. 5329 (NHT). Kilimanjaro: Same
district, Mkomazi Game reserve, Ibaya Hill, 3°58'S, 37°48'E, R. Abdallah & K. Vollesen
95/198 (BR, K, P [P05620649]). Tanga: Korogwe District, 2 km W of Mkomazi, un-
der power line, 4°38'53.7"S, 38°03'26.7"E, N. Holstein et al. 90 (B, DSM, M).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
108
Figure 34. Distribution map of C. mildbraedii (based on 51 collections). e individual in Kenya is
introduced. For Tanzania the borders of the regions are given.
14. Coccinia mildbraedii Gilg, Wissenschaft. Ergebn. Deutsch. Zentral-Afrika-
Exped. 1907–1908 Herzog Adolf Friedrich zu Mecklenburg, Bot. 2(4): 343. 1914.
Type: Rwanda. [Western Province]: Kissenye [Gisenyi]. Bugoy [Bugoyi] forest, mixed bam-
boo forest, c.2500 m, , fr, 30 Oct 1907, J. Mildbraed 1425 (Holotype: B, destroyed).
Type: Burundi. Muramvya: [Mt] Teza, 3°13'S, 29°33'E, female, , fr, M. Reek-
mans 7399 (Neotype, designated here: K!; isoneotypes: BR!, EA!, MO!, WAG!
[WAG0225430], WAG! [WAG0225433]).
Coccinia ulugurensis Harms in Mildbraed, Notizbl. Bot. Gart. Berlin-Dahlem 11:
1091. 1934.
Type: Tanzania. [Morogoro]: Uluguru Mts, northwestern side, c.1350 m, over shrubs
at forest margin, male, , 14 Mar 1933, H.J.E. Schlieben 3643 (Holotype: B! [B
Monograph of Coccinia (Cucurbitaceae) 109
10 0154929, digital image: B, JPS], isotypes: B! [B 10 0154930, digital image:
B, JPS], BM! [BM000815208], BR! [BR0000008886163, K neg. 5264, digital
image: JPS], BR! [BR0000008887498, digital image: BR, JPS], G! [G00301594],
HBG! [HBG506425, digital image: JPS], LISC! [LISC 002496, digital image:
IICT, JPS], M! [M0105771, digital image: JPS], MA! [MA386121, digital image:
JPS], MO!, P! [P00346275, digital image: JPS, P], S [S-G-1519, digital image:
JPS], Z! [Z-000004448, digital image: Z], photo of isotype from BR! [EA, K]).
Description. Perennial climber. Stems up to 20 m, when young sometimes villose
with whitish, articulate trichomes, later often subglabrous to glabrous. Petioles 4–8
cm long, glabrous or with pale, articulate trichomes. Leaves 9.5–16.5 × 10–16.5 cm,
shallowly to profoundly 3- or 5-lobate. Lobes triangulate, ovate to elliptical. Leaf mar-
gin entire and denticulate to serrate. Upper leaf surface glabrous or with hyaline to
white pustules. Lower leaf surface glabrous or sometimes villose with whitish, articu-
late trichomes, sometimes with white pustules on the main veins. Probracts up to 3.5
mm. Tendrils simple or bid. Male owers in racemes, rarely accompanied by one
solitary ower, or one single ower only. Common peduncle 3–4.5 cm, pedicels up
to 7 mm, bracts up to 1mm, caducous. Pedicels of solitary owers up to 2.5 cm, each
glabrous. Perianth tube glabrous, calyx lobes up to 2.5 mm, triangulate to lineal, in
buds adpressed to corolla, later spreading. Corolla 1.2–2.9 cm long, orange bu, lobes
0.3–1cm. Filament column and anther head not seen. Pollen sacs cream yellow. Fe-
male owers solitary, pedicel 3–8 cm, glabrous. Hypanthium glabrous, calyx lobes and
corolla like in male owers. Ovary glabrous, ribbed. Style not seen, stigma bulging,
yellow. Fruit up to 20 cm long and 5 cm in diameter, unripe green with white mottling
and longitudinal green lines, ripening via yellow, orange into deep red. Seeds 6–7 × 5
× 1.5 mm (L/W/H), symmetrically obovate, face atly lenticular.
Phenology. Flowering time: January–April, June, August, September, November,
December, likely throughout the year.
Distribution. Fig. 34. NW Burundi, D. R. Congo (North Kivu, South Kivu),
Rwanda (Western Province), Tanzania (Iringa: Kipengere Range, Uzungwa Mts; Kigo-
ma: Mahali Mts; Mbeya: Kipengere Range; Morogoro: Uluguru Mts; Ukaguru Mts,
Uzungwa Mts), Uganda (Western Region). Elevation 1200–2600 m. Afromontane
cloud forests and mist forests, upland rainforests, bamboo forest, in Macaranga kili-
mandsharica shrubs, rarely in Pennisetum savannas. On basalt and graphitic schist soils,
lateritic clay. Introduced in Kenya.
Vernacular names. Kihunde: mutanga (Deru 485), Kindanda: mwore (Deru 485),
Kinande: mombowa (P. Gille 218), Kinyarwanda: umuvunguvungu (G. Bouxin 820),
umufungofungo (G. Troupin 11163), umwonkalere (Deru 485), Kisafwa: itangalulu
(C.J. Kayombo 1003).
Remarks. Coccinia ulugurensis cannot be denitely distinguished from
C.mildbraedii. e leaves are 3-lobate with rather triangulate lobes towards central
Tanzania (C. ulugurensis), whereas in the western areas the leaves may be deeper
lobate with lanceolate lobes (C.mildbraedii). Collections of the C.ulugurensis form
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
110
also occur in the Ukaguru Mts (M. ulin & B. Mhoro 2933) from which a close-
by located population has been recollected for sequencing (N. Holstein et al. 76)
because collections from the Uluguru Mts were not available. However, forms similar
to C.ulugurensis also occur in the Western Rift area. Vice-versa, 5-lobate leaves also
occur in central Tanzania. Jerey (1967) used to distinguish the species also by the
occurrence of “crisped hairs”, but these do not seem to be specic.Such trichomes also
occur in other species, such as C.adoensis, C.grandiora and C.mackenii, and the ne
pubescence as described for C.ulugurensis regularly occurs in young shoots and often
disappears later on. Collections that represent the two species cluster together (Fig. 17)
and either both species share major parts of their distribution ranges and only dier
morphologically in nuances of lobation depth, or they belong to a single species, of
which here the latter case is assumed here.
Specimens examined. (Selection, in total: 76) D.R. Congo. North Kivu: Lu-
bero territory, Bingi, A. Léonard 5415 (BR, EA, WAG [WAG0225422], WAG
[WAG0225423], WAG [WAG0225424]). South Kivu: Kabare territory, Marais Mu-
sisi, 28°42'E, 2°16'S, P. Bamps 2844 (BR, EA, WAG [WAG0225420]). Kenya. Central
Province: Limuru, tea estate, introduced from Tanzania, J.B. Gillett 20185 (EA, MO).
Rwanda. Western Province: Shangugu territoire, Mont Bigugu, A.R. Christiaensen
1616 (EA, WAG [WAG0225419]). Southern Province: Rutovu, km 64 on Astrida
[Butare]–Shangugu [Cyangugu] route, M. Reynders 394 (BR). Tanzania. Iringa: Da-
baga Highlands, Ihangana Forest Reserve, near Kibangu, 18 mls [29 km] S of Dabaga,
R. Polhill & S. Paulo 1476 (B, BR, EA, K, P [P05620648], PRE). Kigoma: Mpanda
district, Mahali Mts, Sisaga, c.6°S 30°E, T.G. Jeord & J.G.B. Newbould 1924 (COI,
EA). Mbeya: Mbeya rural district, Umalila Forest Reserve, c.7 km W of Ruanda II on
road to Izumbwe (2 km SSE of main peak of Mbogo Mt.), 9°11'S, 33°18'E, R.E. Ger-
eau et al. 5060 (K, MO). Morogoro: Kilosa district, Ukaguru Mts, between Mandege
and Masenge, 6°22'S, 36°58'E, M. ulin & B.E. Mhoro 2792 (DSM, EA, K, MO).
Uganda. Western Region: Kigezi district, Virunga chain, northern foot of Mzhavura
Mt., Nkanda, H.U. Stauer 931 (BR, M, Z).
15. Coccinia ogadensis ulin, Kew Bull. 64: 485. 2009.
Type: Ethiopia. Somali Region: Harerge, 5 km S of Qarsonney, female, fr, 15 May
2006, M. ulin et al. 11183 (Holotype: ETH; isotypes: K! [K000543219, digital
image: K], UPS!).
Type: [Ethiopia]. [Somali Region]: Somaliland, Harradigi [Harradigit], Mar 1885,
F.L. James & J.G. rupp s.n. (Paratype: K!).
Type: [Ethiopia]. [Somali Region]: Somaliland, Harradiqi [Harradigit] or Boobi, Mar or
Apr 1885, F.L. James & J.G. rupp s.n. (Paratype: K!).
Type: [Ethiopia]. [Somali Region]: Agar Ven [Agar Uen], 6°30'N, 45°20'E, 2500
ft [c.760 m], red sandy soil, bushland, 25 Oct 1953, P. Ellis 163 (Paratypes:
FT!, K (2)!).
Monograph of Coccinia (Cucurbitaceae) 111
Type: [Ethiopia]. [Somali Region]: W of Shillavo (Scillave) [Shilabo], 6°25'N,
44°42'E, 1300 ft [c.400 m], sandy soil, bushland, Nov 1955, P. Ellis 383 (Para-
type: K!).
Type: [Ethiopia]. [Somali Region]: E of Gorrahei [Korahe], 700 m, 1 Nov 1967,
P.R.O. Bally 12989 (Paratypes: G!, K!).
Type: [Ethiopia]. [Somali Region]: Scillave [Shilabo]–Wardere road, 6°13'N, 44°45'E,
1130 ft [c.344 m], red sandy soil, open bushland, male, , 2 Apr 1956, J. Simmons
S63 (Paratypes: EA!, K!).
Type: [Ethiopia]. [Somali Region]: 11 km NE Scillave [Shilabo], 6°10'N, 44°52'E,
1300 ft [c.400 m], red sandy soil, open bushland, 13 Apr 1956, J. Simmons S179
(Paratypes: EA!, K!).
Type: Somalia. [Mudug]: 47 miles [75 km] from Galkayo [Gaalkacyo] on Garoe [Ga-
roowe] road, c.1000 ft [c.300 m], red sandy loam and limestone ridges, 15 Oct
1959, C.F. Hemming 1713 (Paratypes EA!, K!).
Description. Perennial? climber or trailer. Stems up to 2 m or longer. Stems gla-
brous, except for nodes with short trichomes, sometimes white pustulate. Petioles
4–15 mm long, glabrous or nearly so. Leaves deeply (3- or) 5-pedately lobate. Cen-
tral lobe 2–8.5 cm long, 1–8 mm wide, lateral lobes shorter. Lobes entire or dentate
to lobulate, linear to oblanceolate. Leaf margins often revolute, apex obtuse with
Figure 35. Distribution map of C. ogadensis (blue dots; based on 8 collections) and C. samburuensis (pale
yellow dots; based on 4 collections). For Ethiopia the borders of the regions and for Kenya the borders
of the provinces are displayed.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
112
nal (brownish? colored) tooth to acute. Upper leaf surface glabrous, pale to white
pustulate; pustules up to 5 mm in diam. Lower leaf surface glabrous, at base with pale
aureolate glands between nerves. Probracts < 1 mm with short, whitish trichomes.
Tendrils simple. Male owers solitary, clustered, or in few-owered racemes. Com-
mon peduncle up to 2 cm, glabrous. Pedicels up to 4–20 mm long, subglabrous
to glabrous. Perianth tube glabrous, calyx lobes 1–6 mm long, in buds erect, later
reexed, glabrous or nearly so, lineal to narrowly triangulate. Corolla 1.7–2.5 cm,
white with green veins or yellow, lobes 0.7–1.3 cm. Petals inside with multicellular
trichomes, outside with short, oligocellular trichomes. Color of lament column,
anther head, and pollen sacs not seen. Female owers not seen, but very likely soli-
tary, pedicels, hypanthium/perianth tube, calyx lobes and petals not largely diering
from male owers. Fruits spindle-shaped to shortly cylindrical, 4.5–5.5 cm long,
c.1.5 cm in diameter, sometimes with short apical tip (“beaked”). Unripe green
with elongate with spots, turning red with whitish, elongate spots. Seeds 4–5.5 ×
2–2.5 × 1–1.5 mm (L/W/H), slightly asymmetrically obovate, face atly lenticular.
Phenology. Flowering time: Imperfectly known, owering in April and in Octo-
ber and November during rainy seasons.
Distribution. Fig. 35. Eastern Ethiopia (C and E Somali Region), Central So-
malia (Mugud). Elevation 300 to 800 m. Red sand and sandy loam, limestone soils.
Open Acacia-Commiphora bushland, Cordeauxia edulis bushland, semi-desert.
Use. Fruits are reported to be edible, juicy, and thirst-quenching (P.R.O. Bally 12989).
Vernacular names. Somali: dudu (P.R.O. Bally 12989), ilgeel (ulin 2009), So-
mali?: lehailu (J. Simmons S179), Somali?: salo dudub (J. Simmons S63).
Remarks. is species is similar to collections of C.grandis with deeply lobate
leaves (described as Coccinia palmatisecta). However, the lobules in C. grandis are
much more distinct when the lobulation is that deep. Apart from this, fruit and seed
shape of C.ogadensis resemble that of C.adoensis.
Ellis notes on the collections no. 163 and 383 a smell of rotten meat. However,
it is unclear, whether this is coming from the owers or from crushed leaves. Several
cucurbit species have a putrid smell when crushed, such as Kedrostis foetidissima or
Momordica foetida, but this has never been reported for a Coccinia species.
Specimens examined. (in total: 10). Ethiopia. Somali Region: Ogaden, J.Sim-
mons 64 (EA).
16. Coccinia pwaniensis Holstein, Kew Bull. 65(3): 435. 2010 [publ. 1 Jan 2011].
Type: Kenya. [Coast Province]: Kwale District, Buda Masini forest, 8 miles [12.9
km] WSW of Gazi, 80 m, male, , 22 Aug 1953, R.B. Drummond & J.H. Hemsley
3953 (Holotype: K! [3 sheets, K000309479, the other two sheets without bar-
code], isotype: EA!).
Type: Kenya. Coast Province: Kili District, Mangea Hill, 39°42'E, 03°16'S, 450
m, dry bushland with Cynometra sp., Brachylaena sp., Manilkara sp., Brachystegia
Monograph of Coccinia (Cucurbitaceae) 113
sp., Julbernardia sp., Diospyros sp., Xylopia sp., Inhambanella sp., 28 Dec 1988,
W.R.Q. Luke 1601 (Paratype: EA!).
Type: Kenya. [Coast Province]: Kwale District, Cha Simba forest, 300 m, fr, 1 Feb
1953, R.B. Drummond & J.H. Hemsley 1078 (Paratype: K!).
Type: Kenya. [Coast Province]: Kwale District, Shimba Hills, Giriama Point area,
1250 ft [c.381 m], forest edge, 17 Mar 1968, F.C.Magogo & P. Glover 315 (Para-
types: EA!, K!).
Type: Kenya. [Coast Province]: Kwale District, Shimba Hills, Pengo Hill area, 1500 ft
[c.457 m], forest, 27 Mar 1968, F.C.Magogo & P. Glover 493 (Paratypes: EA!, K!).
Type: Kenya. Shimba Hills, SE-part of Longomagandi Forest, 350 m, lowland rainfor-
est, 13 Nov 1988, R. Schmidt 1203 (Paratype: EA!).
Type: Kenya. Kwale District, no detailed location given, 15 Jun 1957, Saunders 11241
(Paratype: EA!).
Type: Tanzania. Pwani: Bagamoyo District, Zaraninge Forest in Kiono Plateau, 38°36'E,
6°09'S, 1000 ft [c.305 m], dry evergreen coastal forest, on sand, 14 Mar 1990, Frontier-
Tanzania Coastal Forest Research Programme 1041 (Paratype: K!).
Type: Tanzania. [Pwani]: Kirasawe District: Pugu Hills Forest Reserve on Dar es Sa-
laam–Kisarawe road. Roadside in forest, 100–270 m, 12 May 1970, K.H. Macau-
ley CVL 102 (Paratypes: DSM!, EA!).
Type: Tanzania. [Pwani]: Pugu Hills, 19 Mar 1939, J.H. Vaughan 2774 (Paratype: EA!).
Type: Tanzania. [Pwani]: Pugu Hills Forest Reserve, road W from road-tunnel,
100m, in bushes by car-track through forest, 23 Jul 1972, R.C.Wingeld 2056
(Paratypes: DSM!, EA!).
Coccinia sp. B in C.Jerey, F. T. E. A.: 69. 1967. R.B. Drummond & J.H. Hemsley
1078 (K!); R.B. Drummond & J.H. Hemsley 3953 (K!, EA!); Saunders 11241 (EA!);
J.H. Vaughan 2774 (EA!).
Description. Perennial climber or creeper. Stems up to 3 m long, glabrous. Petiole
0.6–4.1 cm, adaxial side glabrous or with short, sti trichomes, abaxial side with sti,
patent trichomes that can be quite reduced, then appearing wart-like or subglabrous.
Leaves 2–10.4 × 2.7–11.4 cm, shallowly to profoundly 3-(or 5-)lobate, lobes broadly
triangulate to elliptic, margin minutely dentate, tips acute. Upper leaf surface minutely
hyaline pustulate, nerves sometimes with tiny trichomes, lower leaf surface glabrous,
rarely with blackish glands at base, nerves towards the base with sti, patent trichomes
that can be quite reduced, then appearing wart-like or subglabrous. Probracts 2–3mm
long. Tendrils simple. Male owers in racemes, sometimes accompanied by 1–2 soli-
tary owers. Peduncle 3.2–7.7 cm, glabrous, pedicels of owers in racemes 0.2–1cm,
bracts 1–1.5 mm, pedicels of solitary owers up to 3.8 cm. Perianth tube glabrous,
calyx lobes 2.5–3.5 mm long, subulate and spreading, corolla pale yellow to pale or-
ange-yellow, 1.7–2.6 cm, lobes 1–2 cm. Color of lament stalk, anther head, and
pollen sacs not seen. Female ower not seen, perianth likely like in male owers. Style
and stigmas not seen. Fruit solitary, petioles at maturity 20–33 mm long, fruit shape
oblong-fusiform, 6.2–8 × 1.8–2.3 cm, rarely (?) with an up to 5.5 cm long sterile apical
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
114
tip, immature green with pale longitudinal mottling, at maturity becoming orange-red
to scarlet-red with pale mottling. Seeds 6.5–7 × 4–4.5 × c.1.5 mm (L/W/H), more or
less symmetrically obovate, face lenticular.
Phenology. Flowering time: January–March, June–August.
Distribution. Fig. 23. Kenya (Coast Province), Tanzania (Pwani, but likely also
in Dar es Salaam region and Tanga). Elevation 80–460 m. On sandy soil. Open and
disturbed places of East African coastal forests and woodlands (Brachystegia sp., Julber-
nardia sp., Diospyros sp.).
Vernacular names. Kidigo: mnokonyoka (F.C.Magogo & P. Glover 493), mtam-
baa (F.C.Magogo & P. Glover 315), Kijibana: muri ya nyoka (L.J. Lap 258).
Remarks. Morphologically this species (the only one missing DNA sequences)
is close to C.senensis. e indumentum is reduced in prominence and in extent to
the petiole and leaves in C.pwaniensis, and the leaves are rather 3-lobate and long
petiolate, in contrast to often 5-lobate and shortly petiolate leaves in C.senensis. e
racemes in C.pwaniensis have considerably more owers than in C.senensis. However,
both species share the subulate calyx lobes, and fruit and seed shape suggest that both
species are closely related with C.adoensis. As C.pwaniensis and C.senensis do not co-
occur, they might be sister species from allopatric speciation, with C.pwaniensis occur-
ring in a refugial distribution in the northern coastal forests of East Africa.
Specimens examined. (in total: 13) Kenya. Coast Province: Kili district,
Kaya Jibana, entering southern forest patch of Kaya Jibana following the path from
shop/hoteli at Mwarakaya–Ribe road, 3°50'0"S, 39°40'30"E, L.J. Lap 258 (WAG
[WAG0195516], WAG [WAG0195517]); Kwale district, Shimba Hills, Lon-
gomagandi Forest, R. Schmidt 527 (EA, UBT).
17. Coccinia quinqueloba (unb.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3:
533. 1881.
Bryonia quinqueloba unb., Prodr. Pl. Cap. 1: 13. 1794. Cephalandra quinqueloba
(unb.) Schrad. ex Eckl. & Zeyh., Enum. pl. afric.austral. 2: 280. 1836. Momor-
dica quinqueloba (unb.) E.Mey. ex Drège, Zwei panzengeogr. Dokum.: 126,
132, 133, 137, 202. 1843.
Type: [South Africa]. [Eastern Cape]: sylva Krakakamma, male, , Dec, C.P. unberg
22836 (Lectotype, designated by Meeuse (1962: 99) and here: UPS-THUNB! [K
neg. 2978]).
Type: ibid., male, , Dec, C.P. unberg 22837 (Syntype: UPS-THUNB! [K neg. 2977]).
Type: Cap. b. spei [Cape of Good Hope colony], C.P. unberg s.n. (Syntype: S! [S08-
12379]).
Description. Perennial creeper or climber. Stems up to 9 m, glabrous (rarely with re-
mote trichomes). Leaves usually subsessile, petiole 3–8(–17) mm, glabrous (rarely with
remote trichomes). Leaves, 3–9.5 × 4–10 cm, 3- or 5-lobate, auriculate. Lobes oblong,
Monograph of Coccinia (Cucurbitaceae) 115
elliptical to obovate. Leaf margin remotely dentate, apices towards lobe often serrate.
Lobe apices obtuse with a point. Upper leaf surface pale pustulate, lower leaf surface
glabrous, rarely a few blackish glands near base, nerves rarely white-speckled. Probracts
< 1 mm or missing. Tendrils simple. Male owers solitary or in racemes. Common pe-
duncle 0.5–2 cm, petiole in racemous owers up to 1.8 cm, bracts > 1 mm or missing,
solitary owers with petiole 1.8–4 cm, all glabrous. Perianth tube glabrous, calyx lobes
1.5–3 mm, narrow triangulate, erect to spreading. Corolla 1.2–2.2 cm long, pale yel-
low, corolla lobes 0.8–1.2 cm. Color of lament stalk, anther head, and pollen sacs not
seen. Female owers one solitary. Petiole 1–2.5 cm, glabrous. Hypanthium glabrous,
calyx lobes and corolla like in male owers. Style and stigmas not seen. Fruits 3.5–9 ×
3–4 cm, elliptical to oblong, sometimes short elongated tip, unripe green with longitu-
dinal, white mottling, ripe (orange-)red. Seeds 6–7.5 × 3–3.5 × 1–1.2 mm (L/W/H),
slightly asymmetrically obovate, face (atly) lenticular.
Phenology. Flowering time: January, February, April, July, September, November,
December.
Distribution. Fig. 30. Southern and western Eastern Cape, South Africa. Eleva-
tion sea level to 1000 m. Sandy soils, also on dolomite soil. Coastal bushland, forest,
dry bush, on bushes along rivers, along roadsides.
Remarks. See also under C.mackenii.
Taxonomic remarks. Cephalandra quinqueloba is the type species of the genus
Cephalandra. Meeuse (1962) designated the lectotype of Bryonia quinqueloba to UPS
but did not choose a specimen, which is done here.
Specimens examined. (Selection, in total: 77) South Africa. Eastern Cape: East
London, Dec 1916, H.G. Breyer s.n. TRV23225 (PRE); Amatle Mts, Hogsback Pass,
32°36'50"S, 26°55'25"E, P.B. Phillipson 1079 (MO, PRE); Glen Avon, Feb 1923,
Mrs. J.E. Brown s.n. (PRE [“PRE43005”], Z); Grahamstown, Old Quarry, R.D.A.
Bayliss 8470 (G (2), M, MO, Z); 28 mls [45 km] from Grahamstown on Port Eliza-
beth road, R. Story 2346 (B [B 10 0019800], L, M (2) , MO, PRE, S [S08-12378]);
near Port Alfred, J.L. Sidey 1095 (PRE, S [S08-12464]).
18. Coccinia racemiora Keraudren, Adansonia, ser. 2, 8(1): 41. 1968.
Type: Gabon. [Moyen-Ogooué]: Abanga, C. E. F. A. [Compagnie d’Exploitations
Forestière Africaine] lot, male, , Jun 1963, N. Hallé 2425 (Holotype: P!
[P00346267, digital image: JPS], P! [P00748828, digital image: JPS, P]).
Type: Gabon. Ibid., female, , N. Hallé 2305 (Paratype: P! [P00748829, digital image:
JPS, P], P! [P00748830, digital image: JPS]).
Description. Perennial climber or prostate creeper. Stems up to 5 m, glabrous. Petioles
0.5–2.5 cm, on adaxial side often with line of thin smutty-beige trichomes or glabrous,
abaxial side glabrous. Leaves 6.5–11 × 5–9.5 cm, hastate to 3-lobate with central lobe
dominating, auriculate (auricles may reach the stem). Lobes triangulate. Leaf margin
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
116
entire to somewhat angulate, remotely dentate. Teeth darkening when dried. Upper
leaf surface with waxy cover glabrous with few-celled clear pustules. Lower leaf surface
glabrous with dispersed blackish glands. Probracts up to 2 mm. Tendrils bid. Male
owers ebracteate in lax, glabrous racemes. Common peduncle up to 2.5 cm, pedicels
0.3–1 cm. Perianth tube glabrous, calyx lobes 0.5 mm, shortly lineal, spreading.
Corolla c.1.2 cm, yellowish to orange, lobes 2–4 mm. Color of lament column,
anther head, and pollen sacs not seen. Female owers ebracteate in lax, glabrous
racemes, like in males. Hypanthium glabrous, calyx lobes, and corolla like in males.
Ovary glabrous. Style and stigmas not seen. Unripe fruits glabrous, glaucous, globose.
Figure 36. Distribution map of C. racemiora (blue dots; based on 8 collections, including 1 supposed hybrid).
Monograph of Coccinia (Cucurbitaceae) 117
Ripe fruits unknown, size c.1.5 cm in diam.? Seeds 5 × 3 × 1.5 mm (L/W/H), rather
symmetrically obovate, atly lenticular.
Phenology. Flowering time: January–March, imperfectly known.
Distribution. Fig. 36. Gabon, S Cameroon. Tropical lowland rainforest.
Taxonomic remarks. One Hallé 2425 specimen bears a type label but it is not clear
whether it was attached by Kéraudren herself. Since both specimens appear to be part of
the same individual, both are best to be treated to be holotypes mounted on two sheets.
Remarks. A collection from the Gamba area in S Gabon (M.A. van Bergen 490
(WAG [WAG0151338])) is morphologically close to C.racemiora but shares the
calyx lobes with C.barteri and thus may represent a hybrid. Using plastid markers,
this collection (C.barteri 6) clusters within C.barteri, while in the nuclear LEAFY-like
tree, it clusters with one representative of C.racemiora, but not with both (Figs 17,
18; Holstein and Renner 2011b). Another collection (F.J. Breteler et al. 8835 (MO,
WAG) from Saint Germain area, C Gabon, has calyx lobes typical for C.racemiora,
but a more condensed raceme as in the M.A. van Bergen 490 collection. e hypanthia
of the female owers are urceolate, whereas in male owers they are cup-shaped. Ur-
ceolate hypanthia are also found in some C.barteri forms, and a phylogenetic signi-
cance, e.g., by introgressive hybridization, cannot be ruled out.
Specimens examined. (Selection; in total: 10) Cameroon. South Region: 3 km
N of km 20 Kribi-Lolodorf, high forest exploitation, 3°01'N, 10°03'E, J.J. Bos 6590
(WAG [WAG0225514], WAG [WAG0225515]). Gabon. Estuaire: 12 km SW of
Kinguélé Falls, N. Hallé & J.F. Villiers 5357 (K, P [P05620813]). Ngounié: 35 km
on road from Lebamba to Yéno, 1°58'S, 11°25'E, J.J.F.E. de Wilde & M. Sosef 10456
(WAG [WAG0044628]). Ogooué-Maritime: Rabi North, 1°51.6'S, 9°51'E, I. van
Nek 536 (WAG [WAG0044627]).
19. Coccinia rehmannii Cogn. in Schinz, Bull. Herb. Boiss. 3: 418. 1895.
Type: South Africa. Transvaal: boshveld ad Klippan [according to Meuse (1962)
in Limpopo: Greater Sekhukhune District Municipality, Doornpoort; 24°37'S,
29°26'E], 1875–1880. A. Rehmann 5156 (Syntype: Z! [Z-000004445, digital
image: Z]; syntype: BR! [BR0000005111602, digital image: BR, JPS]).
Type: South Africa. Ibid., A. Rehmann 5157 (Syntype: Z! [Z-000060807, digital
image: Z]).
Type: South Africa. Ibid., A. Rehmann 5168 p.p. (Lectotype, designated by
Meeuse (1962: 102): Z! [Z-000060808, digital image: Z]; isolectotype: BR!
[BR0000005111930, digital image: BR, JPS], K! [K000313196, digital image:
JPS, K]).
Type: South Africa. At Eland river, A. Rehmann 4944 [sic, must be A. Rehmann 4954,
see Taxonomic remarks] (Syntype: Z! [Z-000060806, digital image: Z]).
Coccinia rehmannii Cogn. var. littoralis A.Meeuse, Bothalia 8: 104. 1962. pro parte ex
R. de Carvalho s.n. (Paratypes: COI (2)!).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
118
Type: South Africa. [Eastern Cape]: [Amatole District Municipality], Komgha, Kei
Mouth, H.G. Flanagan 457 (Holotype: PRE [PRE0190559-0, digital image: JPS],
isotypes: BOL?, NBG?).
Type: South Africa. [Eastern Cape]: Cape Morgan, H.G. Flanagan 457 (Paratype:
GRA [GRA0002852-0, digital image: JPS], BOL?, NBG?).
Type: South Africa. [Eastern Cape]: East London, Nahoon, M.W. Nanni 151 (Paratype:
PRE!).
Type: South Africa. [Eastern Cape]: Coee Bay, W. Tyson 24 (Paratypes: B!, COI!,
GRA, MO!, NY!, PRE!, S! [S08-12380]).
Type: South Africa. [KwaZulu-Natal]: 10 mls NW of Mtubatuba, L.E.W. Codd 9620
(Paratypes: COI!, M! [M-0198513], PRE).
Type: South Africa. [Kwa-Zulu Natal]: Umhlanga Rocks, A. Dohse & B. de Winter
223 (Paratypes: NH, PRE!).
Type: South Africa. [Kwa-Zulu Natal]: Manaba Store, J. Gerstner 3407 (Paratype: NH).
Type: South Africa. [Kwa-Zulu Natal]: Dhlebe, J. Gerstner 4261 (Paratypes: NH, PRE).
Type: South Africa. [Kwa-Zulu Natal]: near Durban, T.J. Jenkins TRV7092 (Para-
type: PRE).
Type: South Africa. [Kwa-Zulu Natal]: Mtunzini, S.M. Johnson 612 (Paratype: NBG).
Type: South Africa. [Kwa-Zulu Natal]: Stella Bush, W.E. Marriott herb. no. 24341
(Paratype: ?).
Type: South Africa. [Kwa-Zulu Natal]: ibid., W.E. Marriott herb. no. 27143 (Paratype:
NH).
Type: South Africa. [Kwa-Zulu Natal]: Shelly Beach, A.O.D. Mogg 11941 (Paratype: ?).
Type: South Africa. [Kwa-Zulu Natal]: ibid., A.O.D. Mogg 12070 (Paratypes: M!
[M-0198511], M! [M-0198512], PRE!).
Type: South Africa. [Kwa-Zulu Natal]: without detailed location, T.B. Oatley C 15
(Paratype: PRE).
Type: South Africa. [Kwa-Zulu Natal]: Berea, Small herb. no. 34714 (Paratype: NH).
Type: South Africa. [Kwa-Zulu Natal]: Ubombo coastal veld, P.A. Tosh 28 (Paratype: NU).
Type: South Africa. [Kwa-Zulu Natal]: Ndumu Game Reserve, Ward 3169 (Paratype: ?).
Type: South Africa. [Kwa-Zulu Natal]: ibid., Ward 3170 (Paratype: ?).
Type: South Africa. [Kwa-Zulu Natal]: Umvoti, orns near Greytown, J.M. Wood
5318 (Paratype: NH).
Type: South Africa. [Kwa-Zulu Natal]: Durban, Berea, J.M. Wood 6350 (Paratypes:
BOL, L!, NBG, NH, PRE).
Type: South Africa. [Kwa-Zulu Natal]: Doonside, J. Wylie herb. no. 23299 (Paratype: NH).
Type: Mozambique. Maputo: Lourenço Marques [Maputo], J.M. Borle 253 (Paratypes:
M! [M-0198510], MO!, P! [P05620807, digital image: P]).
Type: Mozambique. Maputo: ibid., J.M. Borle 427 (Paratype: ?).
Type: Mozambique. Maputo: ibid., J.M. Borle 442 (Paratype: PRE!).
Type: Mozambique. Maputo: Inhaca Island, H.G. Breyer TRV20506 (Paratype: PRE).
Type: Mozambique. Maputo: Inhachingo, A.W. Exell et al. 630 (Paratype: SRGH).
Type: Mozambique. Maputo: Massinga, A.W. Exell et al. 645 (Paratype: SRGH).
Monograph of Coccinia (Cucurbitaceae) 119
Type: Mozambique. Maputo: Lourenço Marques [Maputo], A.J.W. Hornby 4599
(Paratype: PRE!).
Type: Mozambique. Maputo: Delagoa Bay [Maputo Bay], H.A. Junod 20 (Paratypes:
BR!, G (2)!, Z! [Z-000073406, digital image: Z]).
Type: Mozambique. Maputo: Inhaca Island, 6 Jul 1958, A.O.D. Mogg s.n. (Paratype: PRE!).
Type: Mozambique. Maputo: ibid., 14 Dec 1955, A.R.A. Noel s.n. (Paratype: PRE!).
Type: Mozambique. Maputo: Lourenço Marques [Maputo], R. Schlechter 11555
(Paratypes: BOL, COI!, G (3)!, GRA, HBG! [HBG518897], PR! [PR 801378],
WAG! [WAG0234182], Z! [Z-000073405, digital image: Z]).
Type: Mozambique. Maputo: Katembe [Catembe], R. Schlechter 11614 (Paratypes: G
(2)!, GRA, PRE!, Z! [Z-000073407, digital image: Z]).
Coccinia ovifera Dinter & Gilg in Dinter, Veg. Veldkost Südw.-Afrik.: 16. 1912.
Type: [Namibia]. Karas: Sandverhaar, M.K. Dinter 1214 (Syntype: ?).
Type: [Namibia]. Otjozondjupa: Otjiwarongo, female, , fr, Jan 1912, M.K. Dinter
s.n. (Syntype: SAM [SAM0072115-0, digital image: JPS]).
Type: [Namibia]. Waldau, female, fr, 3 Feb 1917, M.K. Dinter 432 (Lectotype, desig-
nated here: SAM [SAM0066515-0, digital image: JPS]).
Further possible syntypes (cited in Dinter 1919/20) if collected before end of 1912:
Oshikoto: Gaub, M.K. Dinter 2412 (?); Tsumeb, M.K. Dinter s.n. (?). Unknown:
Hereroland, Palmenwald, M.K. Dinter s.n. (?); Hereroland, Wilhelmsberg, M.K.
Dinter s.n. (?).
Description. Perennial climber or creeper. Stems up to 4 m, glabrous or with broad-
based trichomes, when old often densely white pustulate (esp. in drier areas). Petiole
0.2–4.2 cm, glabrous or with erect, broad-based or often up to 1.5 mm long, articulate
trichomes or only wart-like, when old sometimes dense white pustulate (esp. in drier
areas). Leaves 0.9–9.7 × 1.4–16.6 cm, shallowly to deeply 3- or 5-lobate, auriculate,
sometimes lobulate, rarely cordate. Lobes and lobules usually extending, rarely pointing
towards tip, narrowly to broadly triangulate to lanceolate. Leaf margin rather remotely
denticulate. Apex acute to obtuse, apiculate. Upper leaf surface more or less densely white
pustulate, pustules sometimes with a thick, small trichome, on nerves often with thick,
small trichomes. Lower leaf surface glabrous, sometimes with small, blackish glands be-
tween nerves, nerves usually with erect trichomes, sometimes wart-like. Probract usu-
ally absent, if present then up to 3.5 mm. Tendrils simple. Male owers 1–3 solitary,
if fasciculate or in few-owered racemes then accompanied by 1–2 owers. Common
peduncle 0.7–4.5(–8.5) cm, glabrous or with long, articulate trichomes. Pedicel of ow-
ers in inorescences 0.6–2.8 cm, bracts up to 2.5 mm or missing. Pedicel of solitary ow-
ers (0.2–)0.5–5(–9) cm, glabrous or especially at apex with long, articulate trichomes.
Perianth tube usually with long (> 0.5 mm) trichomes, rarely almost glabrous. Calyx
lobes 0.2–7 mm, narrowly lanceolate or lineal, when young erect, later also spreading to
reexed. Corolla 0.8–2.5 cm long, bu to more or less pale yellow, sometimes with green
venation. Lobes 0.3–1.1 cm. Filament column pale bu, anthers bu, pollen sacs yellow
(Fig. 10a). Female ower solitary. Pedicel 0.4–1.5 cm, glabrous or with long, articulate
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
120
trichomes. Hypanthium usually with long (> 0.5 mm) trichomes, rarely almost glabrous,
calyx lobes and corolla like in male owers. Ovary rarely glabrous, often more or less
densely covered with articulate trichomes. Style columnar, green. Stigmas 2-lobed, yellow
(Fig. 11b). Immature fruit, rarely also at maturity, with whitish, longitudinal mottling
that develops a dark green corona during ripening. Ripe fruit globose to elliptical 1–6.2
× 1–2.8 cm, glabrous or with few articulate trichomes, bright orange to red. Seeds 4.5–7
× 2–3.5 × 1–1.2 mm (L/W/H), asymmetrically obovate to somewhat falcate, face at.
Phenology. Flowering time: January–April, June, October–December.
Distribution. Fig. 37. South Africa (except Western Cape and SW Eastern
Cape), Namibia (except hyperarid regions), Swaziland, southern Mozambique (Gaza,
Inhambane, Maputo), Botswana, Zimbabwe (Manicaland, Masvingo, Matabeleland
South), southern Angola (Namibe, Huila, Cunene, Cuandocubango). Elevation sea
level to 1850 m. Limestone, dolomitic, quartzitic, granitic, and ultrabasic soils. Possibly
some tolerance to Ni and Cu. On loam, clay, white and red sand, sandstone, and gravel,
but prefers sandy (well drained) soils (Meeuse 1962). Full sun to shade. Coastal dunes,
riverbanks, Acacia sclerocarya–A. cara woodland, Combretum apiculatum bushland,
Grewia ava bushland, mopane, Kalahari thornveld, grassland, semi-desert, dunes.
Grazing is tolerated. Light frost seems to be tolerated (Meeuse 1962).
Use. Tuber edible after baking (Dinter 1912, M.E. Keith 50, B. de Winter & O.A.
Leistner 5598). Leaves used as spinach by Shangane people in Gaza province of Mo-
zambique (K.L. Tinley 3000). Fruits edible (R.H.W. Seydel 938, R. Story 5029, B. de
Winter & O.A. Leistner 5598).
Vernacular names. Muchope [Xichope?]: fuculumué (L.A. Grandvaux Barbosa &
F. de Lemos 8502), Otjiherero: otjimaga (M.K. Dinter s.n. Jan 1912), Ronga dialect [of
Xitsonga language]: inyamgwazi (A.O.D. Mogg 31308), shiracarana (L. Macuácua 73
and 75), Tsonga [Xitsonga]: inyamwazi (A.O.D. Mogg 31538), nyampape (C.Liengme
491), Zulu [isiZulu]: uselwa-iwenyoka (Sewram et al. 2006).
Remarks. e (sub-)glabrous “littoralis” form can be easily confused with the pol-
ymorphic C.adoensis (Hochst. ex A.Rich.) Cogn., which diers in shorter calyx lobes
and lenticular seeds, and with C.senensis, which also has lenticular seeds and usually
long-peduncled male racemes.
Meeuse’s variety littoralis is hard to dene as the paratypes are variable, and char-
acters for delimitation are unclear. For example, although the variety should lack white
speckles on the stems, there are some individuals with white speckles along with long
pedicels or conspicuous racemes in KwaZulu-Natal and southern Mozambique as in
the variety littoralis. e holotype of var. littoralis is, in the present author’s opin-
ion, rather intermediate between the holotype of var. rehmannii and the subglabrous
forms, e.g., from Inhaca Island. However, the tendency that Meeuse describes is ap-
parent. Other characters in the collections of his variety, viz. relatively long calyx lobes
and petals, also occur in the high mountains of Namibia but also in the whole periph-
ery of the C.rehmannii distribution range. Strangely, Meeuse does not mention the
most striking dierence between C.rehmannii collections from the inland/dry areas
and coastal/peripheral collections being the globose fruit in inland/dry area individuals
Monograph of Coccinia (Cucurbitaceae) 121
or long elliptical fruit in coastal/peripheral individuals (shown and mentioned in this
treatment as "C. rehmannii a. var. littoralis"; Fig. 10a) respectively. Collections with
this fruit morph also occur in Angola and Zimbabwe but do not follow Meeuse’s other
characters of the var. littoralis. On the other hand, collections with long calyx lobes
and long petiolate owers can have globose fruits. Like the area of the southeastern
coast of South Africa, areas in the north of southern Africa receive more and longer
rainfall per year than the inland, so there is a possible correlation between precipita-
tion and fruit morphology. Elliptical fruits also occur in the closely related C.trilobata
from mountainous and thus more humid habitats but not in C.microphylla from the
dry bushlands of NE Africa whose fruit is globose. However, the characterization by
Meeuse that C.rehmannii is an aggregate species seems to be true. It might be interest-
ing to link morphological characters with haplotypes and to test the tness of these
morphs in the dierent habitats. In any case, the morphological dierentiation seems
to be stable in cultivated individuals, and articial crosses between dierent forms (in-
land vs. subglabrous from the Southeast) result in the onset of a normal fruit (resulting
seeds were not used for cultivation).
Figure 37. Distribution map of C. rehmannii (blue dots; based on 251 collections). For South Africa the
borders of the provinces are given.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
122
e placement of C.rehmannii var. littoralis forms with other C.rehmannii forms
in plastid and nuclear phylogenies (Figs 17, 18) also rejects the hypothesis of a hybrid
origin (e.g., with C.adoensis or C.senensis), but rather suggests parallel evolution due
to ecological factors.
Taxonomic remarks. e protologue contains a literal mistake for the syntype
from Eland River and must be corrected from 4944 to 4954. On the one hand, A.
Rehmann 4944 (GRA, K) is a Malpighiaceae. On the other hand, there is A. Rehmann
4954 (a C.rehmannii) in Z from Eland river and with a remark by Cogniaux ‘sp. nov.’
Hence, A. Rehmann 4954 is the syntype of Coccinia rehmannii, not A. Rehmann 4944.
e GRA specimen of H.G. Flanagan 457, which is supposed to be the isotype
(cited by Meeuse) of C.rehmannii var. littoralis, is in fact merely a paratype. Meeuse
stated clearly the location as “Komgha: Kei Mouth” and chose the PRE specimen from
there as the holotype, but the GRA specimen is from the nearby located Cape Morgan.
Apparently, Flanagan used the same collection number for dierent gatherings. e
GRA specimen thus cannot be regarded as a duplicate despite the same number. As
the GRA specimen does not have a label by Meeuse, he just cited the specimen without
seeing it.
e similarity of C.rehmannii var. littoralis to C.senensis led to a misplaced para-
type. One of the two specimens by R. de Carvalho is a syntype of C.jatrophiifolia
var. australis Cogn. and the two R. de Carvalho specimens from COI are paratypes of
C.subglabra, which are both synonyms of C.senensis.
Coccinia ovifera is a validly published name, although the description is a little cryptic,
hence the species name is not a nomen nudum. Dinter writes that he has found, viz. col-
lected, the species around Grootfontein, in Hereroland (not in the narrow sense of the
1968 homeland) and in Sandverhaar (Namaland). erefore, the requirements for valid-
ity are met (37.3 Note 2). e latter site is cited by him explicitly in a later publication
(Dinter 1919/20). e collections designated as syntypes above match the description as
given in the protologue. As the present author did not see the specimens designated as
“possible syntypes”, they are tentative and might have been destroyed in Berlin.
Specimens examined. (Selection, in total: 321) Angola. Huíla: Cambos, near Chi-
ange, A. Menezes 3629 (LISC [LISC 031347], P). Namibe: andados ca. 55 km de Moça-
medes [Namibe] para Dois Irmãos [Caraculo], E.J. Mendes 3969 (COI, LISC). Botswana.
Ghanzi: 200 mls [320 km] NW of Molepolole, R. Story 5029 (COI (2), EA (2), PRE (2),
Z). Kgalagadi: c.50 mls [80 km] NNW of Tsabong, O.A. Leistner 3120 (LISU, M, PRE).
Ngamiland: 80 km W of Tsau on Cae Cae road, D.G. Long & D.A.H. Rae 416 (E). Mo-
zambique. Gaza: Vila de João Belo [Xai-Xai], entre Chicumbane e a Barra do Limpopo:
próximo da povoação commercial da Barra, F. de Lemos & A. Balsinhas 131 (BM, COI) and
133 (BM, COI (2)). Inhambane: Pomene, in hotel area, P.C.M. Jansen et al. 7533 (G n.v.
[G00305107], MO, WAG [WAG0234198], WAG [WAG0234199]). Maputo: between
Costa do sol and Marracuene, Mutanhane, A. Balsinhas 230 (BM, COI, PRE). Namibia.
Erongo: Farm Anschluss, 150 km E of Swakopmund on Khomas road to Windhoek, B.
de Winter & D. Hardy 8001 (M, PRE, WAG [WAG0234173]). Hardap: c.20 mls [32
km] from Kalkrand on road to Rehoboth, B. de Winter 3538 (COI (2), L, M, PRE).
Monograph of Coccinia (Cucurbitaceae) 123
Karas: Dassiefontein Farm, 2–3 km E of highway, in foothills of Groot Karasberge, c.64
km NNE of Grünau, G. Davidse & A. Loxton 6240 (M, MO, S). Omaheke: [Farm] Brei-
tenberg, border of Kalahari, R.H.W. Seydel 2513 (COI, M, WAG [WAG0234172]). Ot-
jozondjupa: 160 mls [257.5 km] E of Grootfontein, Gautscha Pan, R. Story 6219 (PRE);
ibid., R. Story 6238 (M (2), PRE (2), S [S08-12417]). South Africa. Eastern Cape: Port St.
Johns distr., First Beach, M.J. Wells 3434 (MO). Gauteng: Pretoria, Brummeria: BRI, A.
Balsinhas 3474 (MO, PRE, WAG [WAG0234189]). KwaZulu-Natal: Lower Tugela Val-
ley, below Maqumbi, D. Edwards 3053 (B, M, HEID, PRE); 5mls [8 km] on Nkonko-
ni–Pongola road, M.J. Wells 2162 (B, EA, M, Z [Z-000060820]). Limpopo: Immerpan,
near post oce on roadside, A.D.J. Meeuse 9452 (B, COI, L, PRE, Z [Z-000060813]).
Mpumalanga: Impala, siding, E. Retief 1260 (PRE); ibid., E. Retief 1261 (MO, PRE);
Steelport, Burgersfort, 2 km E of town, 24°40'S, 30°22'E, H.J. Venter & A. Venter 10260
(S [S08-12382], WAG [WAG0234178]). Northern Cape: 25 ml. [40.2 km] W of Kim-
berley, H.J.E. Schlieben & H.R. Tölken 11017 (G (2), HEID, M, PRE, S). Northwest:
Rooisloot, 6 Apr 1935, A.O.D. Mogg s.n. (B, COI, L, PRE); Farm Welgevonden, 6 Apr
1935, A.O.D. Mogg s.n. (B, L, PRE [PRE43077], Z [Z-000060812]). Swaziland. Lubom-
bo: saneni [Tjaneni], I.F. La Croix 4909 (MO, WAG [WAG0234170]). Zimbabwe.
Manicaland: Chipinga district, Sabi Valley Experimental Station, C.Soane 162 (COI
(3)). Matabeleland South: Beit Bridge [Beitbridge], A.W. Exell et al. 425 (LISC); ibid.,
L.C.Leach 10700 (COI, MO).
20. Coccinia samburuensis Holstein, Kew Bull. 65(3): 438. 2010 [published on 1
Jan 2011].
Type: Kenya. [Rift Valley Province]: Samburu East District, on Wamba-Isiolo road,
0.7 km S of turno to Maralal, c.1300 m, female, , fr, 4 Jul 1974, R.B. Faden &
A.J. Faden 74/948 (Holotype: MO!, isotype: WAG! [WAG0234153]).
Type: Kenya. Rift Valley Province: Samburu District, Mt Nyiru, southern slopes, near
a river, 2°03'N, 36°51'E, 1600 m, 1 Apr 1995, B. Bytebier et al. 355 (Paratypes:
EA (2)!).
Type: Kenya. Operoi, 1°12'N, 36°49'E, 1350 m, rocky outcrop in Acacia woodland,
23 Dec 2004, W.R.Q. Luke & P.A. Luke 10787 (Paratypes: EA!, K!).
Type: Kenya. Near Maralal, Lowaweregoi [Lowua Werekoi Mt] 4000 ft [c.1220 m],
rocks in bushland, 5 Dec 1958, J.G.B. Newbould 3233 (Paratype: K!).
Description. Perennial climber. Stems up to 5 m, glabrous, except for minute few-cellular
trichomes visible under 5–10× magnication. Petioles glabrous, at base white speckles
may occur. Leaves 6–14 cm × 10–17 cm wide, (5- or)7-lobate. Leaf lobes elliptical, mar-
gin serrate (to lobulate), teeth (lobule tips) with yellowish glands. Lobe apex subacute, api-
culate. Upper leaf surface glabrous, more or less clear to white pustulate. Lower leaf sur-
face glabrous, nerves white-speckled. Probracts up to 4 mm. Tendrils simple. Male owers
1–2 solitary. Pedicel up to 5 cm long, glabrous. Perianth tube glabrous, calyx lobes 6.5
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
124
mm long, linear, erect. Corolla 3.7–4 cm long, brownish yellow, lobes 2.2–2.5 cm. Female
owers solitary. Pedicel 4–5 mm, glabrous. Hypanthium tube glabrous, calyx lobes and
corolla like in males. Ovary narrowly cylindrical, glabrous. Fruits c.14 × 1.5–2 cm, long
cylindrical, unripe green with lighter spots, color of ripe fruit unknown but likely red.
Seeds 6.5–7 × 3.5–4.5 × (≥ 1) mm (L/W/H), symmetrically obovate, face atly lenticular.
Phenology. Flowering time: Imperfectly known. Flowering in April, July, and
December, but likely to ower as long water is available (rainy seasons).
Distribution. Fig. 35. Only known from Samburu area in Kenya (hence the epi-
thet). Only known from seepage line in rocky (granite) outcrops in Acacia-Commiphora
deciduous bushland.
21. Coccinia schliebenii Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 685. 1932.
Type: Tanzania. Morogoro: Ulanga district, Mahenge ward, Mtimaliassi near Mahenge
station. 900–1000 m, male, , 14 Jan 1932, H.J.E. Schlieben 1620 (Holotype:
B! [B 10 0154928, digital image: B, JPS], isotypes: BM! [BM001010003], BR!
[BR0000008886828, digital image: BR, JPS], BR! [BR0000008887580, digi-
tal image: BR, JPS], G! [G00301595], M! [M0105774, digital image: JPS], MA
[MA386129, digital image: JPS], P! [P00346274, digital image: JPS, P], S [S-G-
1518, digital image: S], Z! [Z-000004447]).
Coccinia calophylla Harms in Mildbraed, Notizbl. Bot. Gart. Berlin-Dahlem 12: 522. 1935.
Type: Tanzania. Lindi: Muera plateau, Bakari, , 26 Oct 1934, H.J.E. Schlie-
ben 5551 (Holotype: B! [B 10 0154924, digital image: B, JPS], isotypes: BM!
[BM001010001], BM! [BM001010002], BR! [BR0000008887153, digital image:
BR, JPS], BR! [BR0000008887511, digital image: BR, JPS], G! [G00301767], G!
[G00301768], HBG! [HBG506428, digital image: JPS], M! [M0105775, digital
image: JPS], MA [MA386127, digital image: JPS], P! [P00346272, digital image:
JPS, P], P! [P00346273, digital image: JPS, P], S [S08-11865, digital image: JPS,
S], Z! [Z-000073408, digital image: Z], Z! [Z-000073409, digital image: Z]).
Description. Perennial climber. Stems up to 12 m, densely covered with short, sti,
smutty-brownish trichomes. Petioles 1.5–11 cm, indumentum as on stem. Leaves 5–18
× 4.5–18 cm, slightly to deeply palmately 5-lobate. Lobes broadly triangulate to long
elliptical, margin dentate, tips acute or obtuse. Upper leaf surface usually densely cov-
ered with short, thin trichomes. Lower leaf surface densely (esp. on nerves) covered with
short, sti, dirty-brownish-beige trichomes. Probracts up to 4.5 mm. Tendrils simple or
bid. Common peduncle 1.1–6.5 cm, with indumentum like on stem to puberulous,
pedicels of owers in racemes with up to 4 mm, indumentum as on peduncle, bracts 3–4
mm. Pedicels of solitary owers 1.2–5 cm, indumentum as on peduncle. Perianth tube
with indumentum like on stem to puberulous. Calyx lobes lineal to narrowly lanceolate
10–15 mm. Corolla 4–6.2 cm long, yellow, apricot, pale orange, sometimes marked with
purple, lobes 2–3.2 cm. Filament color not seen, anther head not seen, pollen sacs dark
Monograph of Coccinia (Cucurbitaceae) 125
yellow to orange. Female owers solitary, pedicels 2.5–4.5 cm long, densely covered with
short trichomes. Hypanthium with indumentum like on stem to puberulous, calyx lobes,
and corolla like in male owers. Ovary with smutty-brownish trichomes. Style 3–6mm,
color not seen. Stigmas 2-lobed, orange-yellow. Fruit 7–9 × c.2.5 cm long, oblong to
shortly cylindrical, ripening from green with 10 more deeply colored ribs via yellow to
red. Seeds 5.5–6 × 2.5–3 × 1 mm (L/W/H), symmetrically obovate, face lenticular.
Phenology. Flowering time: January–March, May–July, December.
Distribution. Fig. 38. Ethiopia (Benishangul-Gomaz?, Gambela, Oromia,
SNNPR), Mozambique (Cabo Delgado), South Sudan (Eastern Equatoria), Tanzania
Figure 38. Distribution map of C. schliebenii (pale yellow dots; based on 27 collections). For Ethiopia
and Tanzania the borders of the regions are given.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
126
(Iringa, Lindi, Morogoro, Mtwara?, Ruvuma). Elevation 300–1900 m. Black cotton
soil, reddish soil, volcanic underground. Markhamia-Dombeya woodland, Chlorophora-
Albizia woodland, Dalbergia-Pterocarpus-Combretum-Acacia woodland, woodland
with Acanthus sennii, Baphia abyssinica forests, lower afromontane forests, termite hills,
gallery forests.
Use. Fruits edible (W.J. Kindeketa et al. 2793).
Vernacular names. Didinga: moroich (J.G. Myers 10918), Kipogoro: mdalla (W.J.
Kindeketa 2747), Mokonde: ncauedi (M.F. Correia 92).
Specimens examined. (Selection, in total: 31) Ethiopia. Gambela: 20 km E of Pu-
nido, along the new road to Gog, 7°34'N, 34°24'E, I. Friis et al. 7317 (C, K). Oromia:
c.25 km E of Lekemti [Nekemte], W.J.J.O. de Wilde & B.E.E. de Wilde-Duyes 7184
(K, MO, WAG [WAG0225409], WAG [WAG0225410], WAG [WAG0225411]).
SNNPR: Ghibie [Gibe] or upper Omo gorge, 182 km SW of Addis Abeba on road
to Jimma, north bank, J.W. Ash 898 (EA (2), K). Mozambique. Cabo Delgado: Ma-
condes, 2 km from Mueda to Negomano, near Santo António mission, M.F. Correia
92 (LISC). South Sudan. Eastern Equatoria: Didinga Mts, Mt Lotuke, Char, J.G.
Myers 10918 (K). Tanzania. Iringa: Mundi district, Lulanda village, N and NW of
Ihili forest patch, 8°35'59"S, 35°37'12"E, M.A. Mwangoka & C.J. Kayombo 63 (MA
n.v., MO, P [P05620800]). Lindi: Rondo Plateau, E. Milne-Redhead & P. Taylor 7630
(EA, K). Morogoro: Ulanga district, Kitonga subvillage, 8°46'46"S, 36°42'34"E, G.S.
Laizer et al. 1449 (BM, MO). Ruvuma: [near Gumbiro], by R. Mtandazi [river], E.
Milne-Redhead & P. Taylor 8538 (B [B 10 0312902], EA, K, LISC, P [P05620801])
and 8539 (EA, P [P05620802]).
22. Coccinia senensis (Klotzsch) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 535. 1881.
Cephalandra senensis Klotzsch in W.C.H. Peters, Naturw. Reise Mossambique: 151. 1862.
Type: Mozambique. [Zambézia Province]: Rios de Sena [province], without detailed
locality, in grassland, W.C.H. Peters s.n. (Holotype: B, destroyed).
Type: Tanzania, Lindi Region: 40 km W of Lindi, Lake Lutamba, hill, woodland,
climbing over bushes, c.240 m, male, , 6 Sep 1934, H.J.E. Schlieben 5259 (Neo-
type, designated in Holstein and Renner (2010: 441): M! [M-0165202]; iso-
neotypes: B! [B 10 0379053, digital image: B], BM!, G! [G00301596], HBG!
[HBG518898], MO!, P!, S! [S08-12156, digital image: S], Z!).
Coccinia jatrophiifolia (A.Rich.) Cogn. var. australis Cogn. [sphalm.: Coccinia jatrophæ-
folia var. australis Cogn.] Bol. Soc.Brot. ser. 1, 7: 228. 1889.
Type: Mozambique. [Nampula]: Mossuril et Cabaceira (Zambézia), male, , 1884,
R.de Carvalho 15 (Lectotype, designated here: BR!).
Type: Mozambique. Ibid., male, , 1884–1885, R. de Carvalho s.n. (isolectotype: COI!).
Coccinia fernandesiana C.Jerey, Kew Bull. 30(3): 478. 1975.
Type: Mozambique. Niassa: Erati, between Namapa and Ocúa, near river Lúrio
bridge, female, , fr, 9 Mar 1960, F. de Lemos & L. Macuácua 29 (Holotype: COI,
Monograph of Coccinia (Cucurbitaceae) 127
isotypes: BM! [BM001010006], BM! [BM001010007], K!, LISC! [LISC 002485,
digital image: IICT, JPS], LMA, PRE! [PRE0592949-0, digital image: JPS], PRE!
[PRE0592950-0, digital image: JPS], SRGH! [SRGH0106711-1, digital image:
JPS], SRGH! [SRGH0106711-2, digital image: JPS], SRGH! [SRGH0106711-3,
digital image: JPS]).
Type: Tanzania. Mtwara: Masai Distr. [sic, must be Masasi Distr.], W of R. Bangala,
390 m, in woodland on gravelly soil, 17 Dec 1955, E. Milne-Redhead & P. Taylor
7703 (Paratypes: EA!, K (2)!, LISC!, P!).
Type: Tanzania. Lindi: Mlinguru, 275 m, shrub woodland, 18 Dec 1934, H.J.E.
Schlieben 5745 (Paratypes: B! [B 10 0379052, digital image: B], B!, BR!, EA!,
HBG! [HBG518899], K (2)!, P!, LISC!, MO!, PRE!, SRGH).
Type: L. cl., F. de Lemos & L. Macuácua 30 (Paratypes: BM! [BM001010005], COI!
[2 sheets], K!, LISC! [LISC 002482, digital image: IICT, JPS], LMA, P!, SRGH).
Type: Mozambique. Niassa: Erati, between Namapa and Nacarea, F.A. Mendonça 1128
(Paratypes: LISC! [LISC 002483, digital image: IICT, JPS]).
Type: Ibid., F.A. Mendonça 1129 (Paratypes: LISC! [LISC 002484, digital image:
IICT, JPS]).
Type: Mozambique. Zambézia: Milange, 95 km towards Quelimane, A.R. Torre &
M.F. Correia 14060 (Paratypes: K! [K000313233, digital image: JPS, K], LISC!
[LISC 002481, digital image: IICT, JPS]).
Type: Tanzania. Lindi: Nachingwea, Pterocarpus-Combretum woodland, B. Anderson
815 (Paratypes: EA!, K!, NHT!).
Type: Tanzania. Lindi: Mbemkuru [also called Mbwenburu, Mto Bwamkuro], in de-
ciduous thicket by roadside, 135 m, E. Milne-Redhead & P. Taylor 7473 (Paratypes:
Figure 39. Inorescence of a male Coccinia senensis, note the long triangulate (may be narrower in other
collections, then subulate) calyx lobes in contrast to the calyx lobes of C. adoensis var. adoensis in Fig. 21;
picture taken from the neotype (H.J.E. Schlieben 5259 (M)). Black bar equals 1 cm.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
128
BR!, EA!, K (2)!, P!), E. Milne-Redhead & P. Taylor 7473A (Paratypes: K (2)!),
E.Milne-Redhead & P. Taylor 7473B (Paratype: K), E. Milne-Redhead & P. Taylor
7473C (Paratype: K), E. Milne-Redhead & P. Taylor 7473D (Paratype: K).
Coccinia subglabra C.Jerey, Kew Bull. 30(3): 479. 1975.
Type: Mozambique. Nampula: Nacala, 11km from Itoculo towards Nacala, 130m,
male, , 4 Dec 1963, A.R. Torre & J. Paiva 9417 (Holotype: LISC [LISC 002486,
digital image: IICT, JPS]; isotypes: COI, K! [K000313458, digital image: JPS, K],
LMA).
Type: Mozambique. Ibid., A.R. Torre & J. Paiva 9417A (Paratypes: COI, K!
[K000313457, digital image: JPS, K], LISC, LMA).
Type: Mozambique. Niassa: Ruvuma River, J. Kirk s.n. (Paratype: K!).
Type: Mozambique. Nampula: Mossuril e Cabaceira, R. de Carvalho s.n. (Paratypes:
COI (2)!).
Type: Mozambique. Zambézia: 23 km on road [from vila] Maganja da Costa [=
Olinga] towards Namacurra, A.R. Torre & M.F. Correia 14176 (Paratypes: EA!,
LISC [LISC 002487, digital image: IICT, JPS], MO!).
Description. Perennial climber or creeper. Stems up to 3 m, glabrous or with erect,
sti, articulate, pale trichomes, glabrescent, when older sometimes with white pus-
tules. Leaves subsessile or distinctly but not long petiolate. Petiole 0.4–4 cm long,
abaxial side more or less densely covered with erect, sti, articulate, pale trichomes,
sometimes glabrous. Leaves 4–14 × 5–16 cm, cordate or shallowly to deeply 3- or
5-lobate, sometimes auriculate (Fig. 6). Lobes triangulate, lanceolate to lineal, some-
times with broader end. Finely to coarsely serrate-dentate, especially towards the lobe
tips, sometimes lobulate. Lobe apices acute to subacute, with an apical tooth. Upper
leaf surface with erect, sti, articulate, pale trichomes or with whitish pustules. Lower
leaf surface glabrous or covered with erect, sti, articulate, pale trichomes, nerves with
same indumentum or with white pustules. Between the main nerves at lamina base
often with blackish glands. Probracts up to 4 mm. Tendrils simple, rarely bid. Male
owers (Fig. 39) in few-owered racemes, sometimes accompanied with 1–2 solitary
owers. Common peduncle 0.3–6.2 cm long, often more or less densely covered with
erect, sti, articulate, pale trichomes. Petiole of owers in racemes 0.2–1.2 cm. Bracts
up to 2.5 mm, often missing. Petioles of solitary owers 3.2–6.2 cm long. Petioles
in each case with indumentum of peduncle. Perianth tube glabrous or with erect,
sti, articulate, pale trichomes. Calyx lobes 2.5–6.5 mm long, subulate to narrowly
triangulate-lanceolate. Corolla 1.1–3 cm, yellow, orange to salmon, lobes 0.9–1.1 cm.
Color of lament column, anther head, and pollen sacs not seen. Pedicel of female
ower 1.2–2.3 cm. Hypanthium, calyx lobes, and corolla like in male owers. Ovary
glabrous or more or less covered with erect, articulate trichomes. Style and stigma not
seen. Fruit 3.5–4.4 × 1–1.5 cm, long ovoid to shortly cylindrical, often with apical
sterile tip. Unripe pale green with dark green longitudinal lines, ripe red. Seeds 5.5–7
× 3–5 × 1.2–1.3 mm (L/W/H), symmetrically obovate, face lenticular.
Phenology. Flowering time: January–April, September, December.
Monograph of Coccinia (Cucurbitaceae) 129
Distribution. Fig. 23. Central Tanzania (Iringa, Lindi, Morogoro, Ruvuma), Ma-
lawi (Southern Region), Mozambique (Cabo Delgado, Nampula, Tete, Zambezia).
Elevation 0–700 m. Sandy soil. Coccinia senensis seems to be a typical element of the
Zambezian center of endemism (White 1983b). Deciduous woodlands, termitaria,
riverine thickets, coastal forests.
Vernacular names. Kihehe: mtumbulansoka (W. Carmichael 171), Macua
[Makhuwa]: muuco-uco (F. de Lemos & L. Macuácua 29)
Remarks. e species is recognizable by the combination of few-owered racemes,
long subulate calyx lobes, and the often subsessile leaves. e trichome type (often
appearing articulate when dried) is the same as in C.rehmannii, where (sub-)glabrous
collections also occur (see also the Taxonomic remarks). Except for the degree of tri-
chome density, a subglabrous collection (E.M.C.Groenendijk et al. 1031) from 11 km
from the collecting site of the C.subglabra holotype was neither morphologically nor
genetically (Holstein and Renner 2011b) distinguishable from C.fernandesiana, and
C.senensis (sensu Flora Zambesiaca (Jerey 1978) and sensu Holstein and Renner
(2010)). Fruit shape and length as well as the length of the female pedicel are variable,
so C.subglabra is synonymized. Without calyx lobes, C.senensis is only hardly, if at
all, distinguishable from glabrous C.adoensis collections or those with long articulate
trichomes, which are described as C.adoensis var. jereyana in this treatment. e fruit
and seed shape also match the variable C.adoensis. Usually, C.adoensis var. adoensis
has short trichomes and calyx lobes are ≤ 2 mm, but where both species meet (Malawi,
NW Mozambique, S Tanzania), exceptions can be found (listed and further discussed
as C.adoensis var. jereyana). Whether C.adoensis and C.senensis are truly separate
species and the role of these intermediates needs to be tested by articial hybridization,
eld observations, and/or a phylogeographic analysis.
Taxonomic remarks. Although the holotype of C.senensis burned during the de-
struction of the Berlin herbarium in 1943, and the name appears to have been lost,
the protologue mentions several characters that allow C. senensis to be synonymized
with Jerey’s C.fernandesiana. e C.senensis protologue points out “articulate” tri-
chomes and an overall appearance like C.quinqueloba, which matches perfectly with
many collections of C. fernandesiana. Interestingly, many of these collections have
been identied as “Coccinia quinqueloba” or “Coccinia palmata” by various collectors
and scientists. e similarity, including the calyx lobes, is easily visible in many collec-
tions, but both species are restricted to southern Africa.
Cogniaux described var. australis of C.jatrophiifolia (synonym to C.adoensis) rec-
ognizing the similarity to the polymorphic C. adoensis. However, he dierentiated
between the R. de Carvalho specimens with long lineal lobes (BR, COI) and specimens
with lanceolate lobes (BR, COI), which he determined as C.senensis. When Jerey de-
scribed C.subglabra, he cited the two COI specimens (as deduced from his ID labels),
but he did not refer to Cogniaux’ variety, which must have been overlooked. e one
COI specimen is therefore paratype of C.subglabra and syntype of C.jatrophiifolia var.
australis. e two COI specimens are also misplaced paratypes of Meeuse’s C.rehman-
nii var. littoralis. e similarity of the COI specimens of Meeuse’s variety to C.senensis
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
130
is striking, but the long peduncles and the conspicuous black sublaminal glands refer
rather to C.senensis than to C.rehmannii.
Specimens examined. (Selection, in total: 49) Malawi. Northern Region: Rumphi
district, Nyika Plateau, 20 mls N of M1, J. Pawek 13339B (MO). Southern Region:
Bvumbwe, I.F. La Croix 2653 (MO); Lengwe National Park, near Mukanyu ravine,
A. Hall Marker 1051 (K). Mozambique. Cabo Delgado: Mueda Plateau, 11°22'S,
39°20'E, W.R.Q. Luke et al. 10084 (EA, K). Nampula: Monapo district, Monapo,
forest reserve of Mr. Wolf, E.M.C.Groenendijk et al. 1031 (WAG [WAG0104327]).
Tete: Cabora bassa [Cahora bassa], police post no. 3, 5 km from barrage, A.R. Torre
et al. 18788 (MO). Tanzania. Lindi: Selous Game Reserve, Kingupira, 8°28'S,
38°33'E, K. Vollesen 1908 (EA); ibid., K. Vollesen MRC 4316 (DSM, EA, K, WAG
[WAG0234144]); ibid., R.C.Wingeld et al. 3466 (DSM). Morogoro: Kilosa district,
Ilonga Research Institute, 9.5 km NNE of Kilosa on road to Dumila, 6°46'31.3"S
37°2'23"E, N. Holstein et al. 66 (DSM, M).
23a. Coccinia sessilifolia (Sond.) Cogn. var. sessilifolia
Cephalandra sessilifolia Sond. in Harv. & Sond., Fl. Cap. 2: 493. 1862. Coccinia sessilifo-
lia (Sond.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 534. 1881.
Type: South Africa. Vaal river, J. Burke 289 (Syntypes: K! [K000313207, digital
image: K], PRE, SAM).
Type: South Africa. Slengerfontein in Nieuwe Hantom [area where the provinces
Western Cape, Eastern Cape and Free State meet], on rocks, 4500–5000 ft, fe-
male, fr, 1839, J.F. Drège 3375 (Lectotype, designated here: P! [P00346268, digital
image: P]).
Type: South Africa. Nieuwe Hantom, on rocks, 4500–5000 ft, 1839, J.F. Drège s.n.
(Syntypes: BR! [BR0000005111596, digital image: BR], G! [G00301769], K!
[K000313206, digital image: K], L!, P! [P00346270, digital image: P], S! [S08-
12468, digital image: JPS, S], W! [W 0026938: digital image: WU]).
Type: South Africa. Transvaal, C.L.P. Zeyher 580 (Syntypes: BM! p.p., E! [E00303259],
K! p.p. [K000313205, digital image: K], P! p.p. [P00346271, digital image: P]).
Coccinia sessilifolia var. major Cogn. in Schinz, Verh. Bot. Ver. Provinz Brandenburg
30: 152. 1889.
Type: Namibia. Hereroland, male, , 1885, A. Lüderitz 133 (Lectotype, designated
here: Z!).
Type: Namibia. Walvis bay to Odyitambi, Dec 1885–Feb 1886, A. Lüderitz 1a
(Syntype: Z!).
Coccinia schinzii Cogn., Bull. Herb. Boiss. 3: 419. 1895.
Type: South Africa. Transvaal: Klippan [according to Meuse (1962) in Limpopo:
Greater Sikhukhune District Municipality, Doornpoort; 24°37'S, 29°26'E], bush-
veld, 1875–1880, A. Rehmann 5162 (Lectotype, designated by Meeuse (1962:
Monograph of Coccinia (Cucurbitaceae) 131
98): Z! [Z-000004446, digital image: Z], isolectotype: BR! [BR0000005112265,
digital image: BR, JPS]).
Description. Perennial climber or creeper. Stems up to 5 m long, with slight
waxy cover, glaucous (Figs 2b, 7b), glabrous (rst shoots may have short, white
trichomes). Leaves sessile to amplexicaul (rst leaves after appearance of stem can
be distinctly petiolate, rarely also when mature (up to 1.5 cm; Figs 2b, 4b, 7b)),
glaucous, 1.5–12.5 × 2.2–13.5 cm, (cordate to) deeply palmately 5-lobate. Lobes
linear, lanceolate to elliptic. Leaf margin remotely denticulate, with or without
lobules. Lobe apex obtuse to acute, apiculate. Upper leaf surface glabrous, clear to
white pustulate. Lower leaf surface glabrous, sometimes with dark glands near base
of lamina. Probracts up to 1.7 mm or missing. Tendrils simple, very rarely bid.
Male owers solitary or clustered in few-(rarely many-)owered racemes (Figs 2b,
7b). Pedicels of solitary owers 1–4 cm, glabrous. Peduncle 1–6 cm long, glabrous.
Pedicels of owers in racemes 0.3–2.5 cm, glabrous. Bracts glabrous, up to 1.8 mm,
Figure 40. Distribution map of C. sessilifolia var. sessilifolia (pale yellow dots; based on 133 collections)
and C. sessilifolia var. variifolia (blue dots; based on 7 collections). For South Africa the borders of the
provinces are given.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
132
or missing. Perianth tube glabrous, calyx lobes 1.5–3.5 mm long, lanceolate to
(narrow) triangulate, erect to reexed. Corolla 1.5–3 cm long, whitish cream to pale
yellow, rarely dull orange-brown with conspicuous green venation, lobes 0.9–2 cm.
Filament column not seen, anther head not seen, pollen sacs yellow. Female owers
one solitary. Pedicel 1–3 cm, glabrous. Hypanthium glabrous, calyx and corolla like
in males. Ovary glabrous. Style columnar, greenish yellow. Stigmas bulging, greenish
yellow. Fruit 8–12 × 3–4cm, ellipsoid to oblong, when immature green with white
longitudinal spots to stripes with waxy bloom, ripe red (Fig. 13b). Seeds 6–8 ×
3–3.5 × 1–1.5 mm (L/W/H), symmetrically to slightly asymmetrically obovate, face
at (Fig. 14d, e).
Phenology. Flowering time: January–May, October–December.
Distribution. Fig. 40. South Africa, Botswana, Namibia, except high mountains,
hyper-arid regions, and Cape oristic region. Elevation 300–1500 m. Stony soil, sand,
sandy loam, clay loam. Granite or calcareous substrate. Semi-desert, grassland (e.g.,
Rhynchelytrum sp.), bushland, open woodland. Full sun to shade. Moderate distur-
bance tolerated.
Use. Unripe fruits are baked in ashes and eaten (Dinter 1912), ripe fruits edible
(N. Holstein 119, R.H.W. Seydel 3829, R.H.W. Seydel 4100).
Vernacular names. Afrikaans: bobbejaan komkommer (C.A. Smith 3981), Otji-
herero: ekungu (singular), omakungu (plural) (Dinter 1912; 1919/20), Tswana:
mogábalá (J. Snyman & C.Noailles 229), !Kun [Kung]: kitwa (R. Story 5167).
Taxonomic remarks. e C.L.P. Zeyher 580 specimens (syntype) in BM, K, and
P are mixed with a Trochomeria sp.
Specimens examined. (Selection, in total: 184) Botswana. Central District: Ma-
halapye, 2 mls [3.8 km] SW of Kalamare, H.J. van Rensburg B4019 (PRE). Kgatleng: 15
km SE of Artesia (Mosomane), D.T. Cole 1542 (PRE). North-West District: Aha Hills,
H. Wild & R.B. Drummond 6953 (COI). South-East District: Lobatsi [Lobatse], F.A.
Rogers 6281 (G, Z). Namibia. Erongo: Karibib, Okongawa, R.H.W. Seydel 3026 (B (2),
COI, FR (2), G (3), H, HEID, M, WAG [WAG0234195], WAG [WAG0234197]).
Khomas: [Farm] Aris, mountain in the west, R.H.W. Seydel 4100 (B (2), M, MO)
and 4100a (B (2)). Oshana: Amboland, Uukuanjama [Oukwanyama], Omupanda, A.
Wulfhorst 18 (Z). Otjozondjupa: 32 mls [51.2 km] N of Nurugas on road to Karaku-
wisa, B. de Winter 3710 (M, PRE). South Africa. Eastern Cape: in valley near Graa-
Reinet, H. Bolus 364 (S [S08-12381]). Free State: Kroonstad townland, NE of conu-
ence of Blomspruit and Vals river, J.C.Scheepers 1720 (EA, LISU, PRE, S [S08-12461).
Gauteng: Pretoria, Brummeria, Botanical Garden, A. Balsinhas 3406 (MO, PRE, WAG
[WAG0234191]); ibid., A. Balsinhas 3476 (MO, PRE, WAG [WAG0234190]). Lim-
popo: Penge mine, E. Retief 1354 (MO, PRE, WAG [WAG0234188]); c. 30 mls
[48 km] W of Louis Trichardt, western part of Zoutpansberg, near Mara, Buysdorp,
H.J.E. Schlieben 7453 (B, G, HBG, M). Mpumalanga: Blyderivierspoort Nature Re-
serve, Sybrand van Niekerk resort at camp area, E. Retief 1340 (PRE, MO, WAG
[WAG0234187]), ibid., E. Retief 1341 (PRE). Northern Cape: Colesburg, Achtertang,
16 Apr 1934, J.O. Swinford s.n. (PRE [PRE42988]). North West: near Klerksdorp,
Monograph of Coccinia (Cucurbitaceae) 133
H.J.E. Schlieben 10695 (HEID, PRE, S [S08-12463]); 60 mls [96 km] NW of Vry-
burg, Farm Palmyra, R.J. Rodin 3605 (MO, P [P05620787], PRE).
23b. Coccinia sessilifolia var. variifolia (A.Meeuse) Holstein, stat. nov.
urn:lsid:ipni.org:names:77148916-1
Coccinia variifolia A.Meeuse, Bothalia 8: 100. 1962.
Type: South Africa. [Limpopo]: Waterberg, Vaalwater, about 2.25 km from Vaalwater
on road to Hermanusdoorns, male, , 6 Jan 1959, A.D.J. Meeuse & R.G. Strey
10413 (Holotype: PRE [2 sheets: PRE0188239-1 and PRE0188239-2, digital im-
age: JPS], isotypes: BOL, L, SRGH).
Type: South Africa. Ibid., A.D.J. Meeuse & R.G. Strey 10413bis (Paratype: PRE!).
Type: South Africa. Limpopo: Palala river, M.G. Breyer[-Brandwijk] TRV25226 (Para-
type: ?).
Type: South Africa. Limpopo: Rietspruit near Nylstroom [Modimolle], G.P.F. van
Dam TRV23372 (Paratype: PRE!).
Type: South Africa. Limpopo: Nabomspruit, Mosdene, E.E. Galpin s.n. (Paratype: ?).
Type: South Africa. Limpopo: 11 km from Warmbaths [Bela Bela] on Nylstroom road, R.
Story 1525 (Paratype: ?).
Type: South Africa. Limpopo: Warmbaths, c.3600 ft [1100 m], grassland/bush veld,
H. Bolus 11893 (Paratype: BR!).
Type: South Africa. Ibid.?, R. Leendertz TRV7579 (Paratype: ?).
Type: South Africa. [Limpopo]: Waterberg, 5.5 mls [8.85 km] NNE of Warmbaths,
c.1220 m, sour bushveld, J.P.H. Acocks 13903 (Paratype: S! [S08-12475]).
Description. Perennial climber. Stems up to 1.2 m, likely also longer, glabrous. Petiole
0.7–1.6 cm, glabrous. Leaves 5.2–6 × 6–7.5 cm, deeply to shallowly 5-lobate, lobes out-
wards lobulate. Leaf margin remotely dentate, apex obtuse with nal tooth. Upper leaf
surface glabrous, with clear to whitish pustules. Lower leaf surface glabrous, with glands
at base between nerves. Probracts up to 2 mm. Tendrils simple. Male owers in racemes,
accompanied by a solitary ower. Common peduncle 1–1.4 cm, pedicel in racemes 3–6
mm, each glabrous. Bracts up to 1.5 mm, narrowly ovate. Pedicel of solitary owers
0.9–2 cm, glabrous. Perianth tube glabrous. Calyx lobes 2.5–4 mm, subulate to narrowly
triangulate, erect. Corolla c.2 cm, pale bu, lobes not measured. Filament column,
anther head, and pollen sacs not seen. Female owers solitary or clustered in reduced
2-owered racemes. Common peduncle 1 mm, pedicel in racemes 0.9–1 mm, pedicel of
solitary ower not seen, each glabrous. Hypanthium most likely glabrous and perianth
as in male owers. Ovary glabrous. Style and stigma not seen. Fruit and seeds not seen.
Phenology. Flowering time: January–March, November, December. Likely as in
C.sessilifolia var. sessilifolia.
Distribution. Fig. 40. Only known from Limpopo Province in South Africa.
Eleva tion 800–1200 m. On sandstone, well-drained stony sand. Low closed woodland.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
134
Remarks. e new status of C.variifolia was chosen due to the minor dierences
to C.sessilifolia s.str. Subsessile leaves spontaneously occur in C.sessilifolia (H. Bolus
364, F.A. Rogers 19262) and young individuals usually (always?) have petiolate leaves
(N. Holstein 131, Fig. 4b). However, these collections of mature plants with distinctly
petiolate leaves have only been observed in Limpopo Province in South Africa. Sessile
C.sessilifolia leaves can be quite variable, profoundly to deeply lobate, sometimes also
lobulate. Compared to the rather uniform C.quinqueloba (Meeuse 1962), the leaves
thus appear to be extraordinarily variable. Meeuse’s C.variifolia shares the sublaminal
glands (cp. Fig. 7b) and the calyx lobes of C.sessilifolia, and it is geographically nested
within this species (hence no climatic dierentiation). Acocks (J.P.H. Acocks 13903)
also reports a “stark glaucous” appearance, just as in C.sessilifolia. As petiolate leaves
also occur in young C.sessilifolia plants, and subsessile leaves also occur in mature
plants, it is more likely that the distinctly petiolate C.sessilifolia individuals represent a
local xation of this character. As C.sessilifolia var. sessilifolia is derived from petiolate
plants, this variety might even represent a remnant population of these.
Specimens examined. (in total: 8) South Africa. Limpopo: Waterberg Distr., F.A.
Rogers 24932 (Z [Z-000073427]), N. Rooyen 1667 (PRE), R.H. Westfall 2136 (PRE).
24. Coccinia subsessiliora Cogn., Bull. Jard. Bot. État Brux. 4(1): 225. 1914.
Type: D. R. Congo. [Équateur Province]: around Likimi, male, , 15 Oct 1910,
L.C.E. Malchair 433 (Lectotype, designated here: BR! [BR0000008887481, digi-
tal image: BR, JPS], syntype: BR! [BR0000008886835, digital image: BR, JPS]).
Coccinia sp. D in C.Jerey, F. T. E. A.: 70. 1967. Uganda. [Western Region]: Kigezi
District [(Kanungu District/Kisoro District)], Kayonza Forest Reserve [Bwindi
Forest Reserve / Impenetrable Central Forest Reserve], S. Paulo 644 (EA!, K!,
MO!); [Central Region]: Mengo district, Mabira forest, M.V. Loveridge 87 (?);
[Central Region]: Mabira Forest, near Kiwala, R.A. Dummer 3195 (?).
Description. Perennial creeper or climber. Stems up to 4 m, glabrous. Petiole
2–12cm, glabrous, sometimes with white pustules. Leaves 5.5–15 × 6.5–17.5 cm,
almost to the base palmately 5-lobate. Lobes lanceolate, sometimes lobulate; tip acute,
acuminate. Margin serrate-lobulate, denticulate. Upper leaf surface glabrous with
clear to white pustules, rarely with few ne (up to 1.5 mm long) trichomes. Lower
leaf surface glabrous, rarely with dispersed small blackish glands, rarely with tiny tri-
chomes; sometimes nerves with white pustules. Probracts up to 1.5 mm or missing.
Tendrils simple. Male owers in glabrous, dense, compact racemes. Peduncles up to 6
mm. Pedicels up to 4 mm. Bracts 2–2.5 mm. Perianth tube glabrous. Calyx lobes 1–2
mm, subulate, triangulate to lineal, erect to reexed. Corolla 1.2–1.3 cm, orange, pale
yellow-orange, yellow, lobes c.3 mm. Filament column, anther head, and pollen sacs
not seen. Female owers solitary or in few owered racemes. Common peduncle up 1
cm, glabrous. Pedicel of owers in racemes up to 4 mm. Bracts up to 2 mm or missing.
Monograph of Coccinia (Cucurbitaceae) 135
Pedicel of solitary owers up to 1.1 cm, glabrous. Hypanthium glabrous, calyx lobes
and corolla like in males. Ovary glabrous. Style columnar, pale yellow. Stigma 2-lobed,
yellow. Fruit 2–2.4(–7) × 1.7 cm, globose to long ovoid, unripe green with glaucous
waxy cover, ripe color not known, most likely red. Seeds ≥ 4.5 × 2–2.5 × 1–1.5 mm
(L/W/H), asymmetrically obovate, face at.
Phenology. Flowering time: January, April, July, August, October, December.
Distribution. Fig. 41. Congo basin (Central African Republic, D. R. Congo). Forested
mountains of NW Burundi, D. R. Congo (North Kivu, South Kivu), W Rwanda, Uganda
(Western Province), South Sudan (Eastern Equatoria/Sharq al-’Istiwa’iyah: Lotti Forest).
Elevation 300–1950 m. Soil preferences unknown. Tropical rainforests. Macrolobium [=
Gilbertiodendron?] forest, swamp forest, disturbed ground in open forest.
Vernacular names. Kihunde: mutangatanga (R. Gutzwiller 965), Lissongo [Map-
ti]: kanganga (C.Tisserant (Équipe) 1103), Turumbu: ndombo (J. Louis 2709).
Taxonomic remarks. ere are two Malchair 433 specimens. As they do not con-
tain any indication of having been separated from a single specimen, they are treated as
syntypes. e two specimens do not dier in quality of the material, so the specimen
with the original label was chosen to be the lectotype.
Remarks. Collections from the eastern parts of the distribution (esp. E of the
Western Rift) have longer fruits but it appears to be a variable character.
Rarely (J. Louis 5672, J. Louis 13030), the lower leaf lamina and the adaxial petiole
side have short trichomes and the upper lamina has some long trichomes. ese fea-
tures are unusual, but the other characters match the species.
Figure 41. Distribution map of C. subsessiliora (blue dots; based on 18 collections). For D.R. Congo
the borders of the provinces (until 1988) are given.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
136
Although C.subsessiliora is nested within C.barteri in the molecular tree from
plastid markers (Fig. 17), it can be regarded as a proper morphospecies. e deeply
lobate leaves are a distinct character, and the species is distributed only in the Congo
Basin and the eastern rainforests. e present author supposes that C.subsessiliora
might have evolved peripatrically in an arid period of the Pliocene/early Pleistocene
period, eventually near the Kivu Mts, and C.barteri populated these areas later on.
Specimens examined. (Selection; in total: 21) Burundi. Bubanza: Bubanza, J. Lewalle
6504 (BR, EA); ibid., M. Reekmans 1477 (BR (2)). Central African Republic.Lobaye:
Boukoko, C.Tisserant (Équipe) 1103 (BM, G, P [P05620797], P [P05620798]); ibid.,
2276 (BM, G, P [P05620796], P [P05620799], P [P05621163]). D.R. Congo. Équa-
teur: [Nord-Ubangi district] Businga territoire, between Karawa and Businga, J. Lebrun
1928 (BR, P [P05620794], WAG [WAG0225403]). Maniema: between Kindu and
Katakokombe, J. Lebrun 6011 (P [P05620791], WAG [WAG0225402]). North Kivu:
Beni territory, Kiandolili river, Gongobotsi Camp of Albert National Park guards, H.
Fredericq in Herb. G.F. de Witte 8288 (BR, M, PRE, WAG [WAG0225407]). Ori-
entale: Haut-Uélé district, Faradje territoire, Kurukwata (Aba), P. Gerard 3564 (BR,
EA, WAG [WAG0225405]); Ituri district, Mambasa territoire, Réserve de Faune à
Okapi, Epulu, 1°25'N, 28°35'E, C.E.N. Ewango 2290 (M, MO); Yangambi, J. Louis
13030 (BR [BR0000008912916], BR [BR0000008913272], P [P05620795], WAG
[WAG0225404]). South Sudan. Eastern Equatoria: Torit district, Lotti Forest, J.K. Jack-
son 3026 (K). Uganda. Western Region: [Masindi district], Bunyoro, Bujenje county,
Budongo Forest, A.B. Katende K2801 (MO); ibid. T.J. Synott 1322 (EA).
25. Coccinia trilobata (Cogn.) C.Jerey, Kew Bull. 15: 349. 1962.
Peponia parviora var. trilobata Cogn., Bot. Jahrb. Syst. 21: 210. 1895. Peponia trilo-
bata (Cogn.) Engl., Panzenw. Ost-Afr. C: 399. 1895, nom. illeg. [Peponia is a
diatom genus]. Peponium trilobatum (Cogn.) Engl., Engl. & Prantl, Panzenfam.,
Nachtr.: 318. 1897.
Type: Tanzania. Kilimanjaro: Mkuu [c.3°10'S, 37°36'E], 1500 m, in hedges, , fr,
Mar 1894, G. Volkens 1956 (Holotype: B, destroyed; lectotype acc.to sheet, but
not published, so designated here: BR! [BR0000008887160, digital image: BR,
JPS]; isolectotype: BR!).
Coccinia kilimandjarica Cogn. ex Harms in Fries, Notizbl. Bot. Gart. Berlin-Dahlem
8: 489. 1923.
Type: Tanzania. Kilimanjaro: Kibohöhe [farm at c.3°15'50"S, 37°12'0"E], 1100–
1200m, , R. Endlich52a (Holotype; B, destroyed; lectotype, designated here: M!
[M0105772, “1122”, digital image: JPS], isolectotype: H!).
Coccinia kilimandjarica var. subintegrifolia Cogn. ex Harms in Fries, Notizbl. Bot.
Gart. Berlin-Dahlem 8: 490. 1923.
Type: Tanzania. Kilimanjaro: Kibohöhe, 1100–1200 m, , R. Endlich 52 (Holotype:
B, destroyed; lectotype, designated here: M! [M0105773, “1121”, digital image:
JPS], isolectotype: H!).
Monograph of Coccinia (Cucurbitaceae) 137
Description. Perennial climber. Stems up to 3 m, with soft, whitish trichomes, at least
along nerves. Petiole 1.5–16.5 cm, with whitish trichomes (Fig. 8b). Leaves 2.6–14.5
× 3.2–18 cm, cordate, 5-angulate to 5-lobate, sometimes lobulate. Margin denticulate.
Apex at least of central lobe acute, often acuminate. Upper leaf surface with hyaline to
white pustules and usually with white trichomes. Lower leaf surface more or less cov-
ered with soft trichomes, denser on nerves. Probracts up to 1.5 mm or missing. Tendrils
simple. Male owers solitary or in short few-owered racemes. Common peduncle up
to 1.7 cm long, pedicel in raceme up to 3.2 cm, each glabrous or with short trichomes.
Bracts up to 1 mm. Pedicel in solitary owers 0.7–4.7 cm, indumentum as in racemes.
Perianth tube with articulate trichomes. Calyx lobes 2–5 mm long, lineal, erect to re-
exed. Corolla 0.7–2.2 cm, orange-yellow to reddish-orange, deeper colored on the in-
ner side of the lobes, outside with green venation, lobes 0.7–1.3 cm. Filament column
pale yellowish green, anther head a bit darker than lament column. Pollen sacs yellow.
Female owers 1(–2) solitary, pedicel 0.5–4 cm, glabrous or sparsely covered with short
trichomes. Hypanthium with articulate trichomes, calyx lobes and corolla like in males.
Ovary with trichomes, becoming glabrous towards fruit ripening. Style and stigmas not
seen. Fruits ovoid to oblong, 4–4.7(–9) × 2 cm, unripe green with white longitudinal
stripes that develop a dark green corona during ripening, ripe fruits orange-red. Seeds
6.5–7 × 2.5–3.5 × 1.5 mm (L/W/H), more or less asymmetrically obovate, face at
(Fig. 14c).
Phenology. Flowering time: January, May–July, October–December.
Distribution. Fig. 33. Tanzania (Arusha, Kilimanjaro, Tanga), Kenya (Central,
Coast, Eastern, Nairobi, southern Rift Valley Province). Elevation 1100–2100 m. Red
soil, black soil. Open forest, savannas, evergreen bushland, shrubland, grassland.
Use. Leaves eaten as vegetable (Coilly? 24, F. Msajiri 19).
Vernacular names. Dholuo: angwe (G.R. Williams 307), Kinandi: notondwe (G.R.
Williams 307), Meru: katakeru (Coilly? 24), Kikuyu: kigerema (P. Njogu EA13835),
Kipare: itotwe (W.J. Kindeketa 648).
Remarks. e fruits are reported to be poisonous (G.R. Williams 307).
ere are some collections that have a mixed (not intermediate) phenotype with
C.microphylla: the calyx lobes are unusually long (up to 7 mm), which is a strong argu-
ment for C.trilobata, but the indumentum matches C.microphylla. ese morphs do not
occur in single location but are found in the Ndoto Mts (O. Kerfoot 2644), in Koboko
(P. Kirika et al. 002/020/2011), and around Voi (M. Hucks 579, B. Verdcourt 3888, R.
Polhill & S. Paulo 962). Whether these are hybrids (F2 or later) or just a variation is not
known. ese collections look also quite like C.megarrhiza, which occurs in northern
Kenya and Ethiopia, however, the indumentum does not match either. A clarication
where these collections belong to would require sequence data and a better understand-
ing of the plastid and nuclear haplotypes in the three species, which is not available so far.
e collections in the Usambara Mts are often quite glabrous or the trichomes are
minute and thus easy to mix up with C.microphylla.
Specimens examined. (Selection, in total: 63) Kenya. Central Province: South
Nyeri district, S of road (D450), c.4 km E of Nairobi–Nanyuki road, 3 km N of
Kiganjo, S.S. Hooper & C.C.Townsend 1697 (K [K000353353]). Eastern Province:
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
138
Nkunga Crater Lake, P.A. Luke et al. 7256 (EA). Nairobi: Nairobi river Valley, Chi-
romo, 1°16'30"S, 36°48'E, R.B. Faden & A.J. Faden 74/822 (BR, DSM, EA, MO,
WAG [WAG0234201]). Rift Valley Province: Naivasha District [Nakuru district],
Ol Longonot Estate, O. Kerfoot 3543 (EA, S [S08-12472]). Tanzania. Arusha: Small
Momela Lake, H.M. Richards 20036 (EA (2), K [K000353413]). Kilimanjaro: valley
slopes near Alt Moschi [Old Moshi], A. Peter 56453 (B). Tanga: western Usambara
Mts, Mombo–Soni road, R.B. Drummond & J.H. Hemsley 3007 (B, EA, K, LISC, S
[S08-12484]).
Insufficiently known taxa
Coccinia sp. A C.Jerey, Fl. Zambes.: 450. 1978.
Zambia. Northern Province: Chilongowelo, Tasker’s Deviation waterfall, 4900 ft, fe-
male, , 27 Feb 1952, H.M. Richards 883 (K!).
Distribution. Only known from single collection.
Remarks. Jerey (1978) suggested that this collection is allied with C.barteri.
e inorescence matches that of C.racemiora, which diers in an urceolate perianth
tube/hypanthium, more coriaceous leaves, and a glabrous surface. Simple tendrils also
occur in C.barteri, so this is not a good distinctive character. As the plastid haplotypes
of C.racemiora are nested in C.barteri, and the distribution of C.sp. A is within the
C.barteri range, Jerey’s hypothesis is sound. However, the corolla lobes are quite
long, and it seems that the corolla is open campanulate. erefore, there is some simi-
larity with C.mildbraedii and even with C.grandiora.
Coccinia sp. B C.Jerey, Fl. Zambes.: 450. 1978.
Zambia. Southern Province: Mazabuka, on Nanga Estate near Kafue pilot polder
[c.15°45'S, 27°54'E], female, , 7 Mar 1963, H.J. van Rensburg 1620 (K!).
Distribution. Only known from single collection.
Remarks. Like C.sp. A, Jerey (1978) suggested a closer relationship to C.barteri.
e tubular corolla and the coriaceous leaves support that. e collection was found
in riverine bush in Acacia woodland and Hyparrhenia/Setaria grassland in a ood plain
area, which is unusual for C.barteri as it rather occurs in (rain) forests. White (1983a)
calls the phytochorion of that region “edaphic grassland with semi-aquatic vegetation”.
On the one hand, the local soil conditions are not known, and water might be avail-
able throughout the year, on the other hand this individual might also represent a local
adaptation towards increased drought tolerance.
Monograph of Coccinia (Cucurbitaceae) 139
Dubious names
Coccinia aostae Buscal. & Muschl., Bot. Jahrb. Syst. 59: 499. 1913.
Type: [Eastern Africa]. At Mbusi river [authors state that this river ows into the Indian
Ocean in Mozambique, the collector went upstream towards Zambezi river and
Victoria falls; most likely the Buzi River is meant], tree steppe, , 14 Dec 1909, H.
von Aosta [H.L.F.H. d’Orléans] 105 (Syntype: B destroyed; duplicate ?).
is species is supposed to be from Mozambique. However, the describing author,
Muschler, provoked a scandal with this work as Georg Schweinfurth (1915) and his
former supervisor Adolf Engler (Engler et al. 1915; Ryding 2001) accused him of fraud.
Gilg, who contributed corrections in the Cucurbitaceae, suggested that Coccinia aostae
had been described using the G.A. Schweinfurth 578 specimen from Eritrea, which bore
the ms. name Coccinia lalambae Schweinf. A drawing of this species exists in BR! (K
neg. 4887), which likely represents a C.adoensis. However, the name for G.A. Schwein-
furth 578 remains unpublished and the von Aosta 105 specimen is destroyed, and the
name remains dubious. According to the describing authors, duplicates of the von Aosta
specimens have been distributed, and White (1962) found some in Florence (FI or FT).
However, a loan from FT did not contain any Coccinia collections by von Aosta.
Coccinia buettneriana Cogn., Bull. Acad. Roy. Sci. Belgique ser. 3, 14: 351. 1887.
Type: Gabon. No detailed information given, Sep 1884, R. Büttner 18 (Holotype: B
destroyed).
Cogniaux and Harms synonymize (1924) C. buettneriana under Momordica gabonii
Cogn. (1881) as it was collected in close vicinity of Büttner 17 (Momordica gabonii), which
is, according to Cogniaux himself almost not distinguishable from C.buettneriana.
Coccinia calantha Gilg, Bot. Jahrb. Syst. 34: 358. 1904.
Type: Tanzania. [Tanga]: Usambara Mts, Duga, near Nikunde village, 100m, in bush
and on fencing, , Jul, C.H.E.W. Holst 3190 (Holotype: B destroyed; duplicates ?).
As the holotype is destroyed and the description does not give enough sucient char-
acters to relate C.calantha to other species, the name remains dubious. Zimmermann
(1922b) presents a drawing of an anther, but the thecae are too narrow for a Coccinia
but would match Eureiandra species. On the other hand, Eureiandra has free petals,
whereas C.calantha ought to be sympetalous.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
140
Coccinia helenae Buscal. & Muschl., Bot. Jahrb. Syst. 59: 498. 1913.
Type: [Eastern Africa]. At Mbusi river [authors state that this river ows in the Indian
Ocean in Mozambique, the collector went upstream towards Zambezi river and
Victoria falls; likely the Buzi river is meant], steppe, , fr, 3 Dec 1910, H. von Aosta
[H.L.F.H. d’Orléans] 87 (Holotype: B, destroyed; duplicates ?).
As for C.aostae, C.helenae seems to be mistaken. Gilg (Engler et al. 1915) suggested
that Coccinia helenae had been described using the G.A. Schweinfurth 932 collection
from Blue Nile. A drawing of this species exists in BR (K neg. 4846). e drawing, if it
represents a Coccinia (the two subsessile female owers on one node are suspicious), does
not match any species of the present author’s knowledge from Blue Nile area entirely. It
might be C.abyssinica if it was collected in the Ethiopian highlands or C.adoensis but
the fruit would be unusually ovoid. However, if it is from the area as given by Muschler,
it might be C.rehmannii. As the von Aosta 87 specimen is destroyed the name remains
dubious. According to the describing authors duplicates of the von Aosta specimens have
been distributed, and White (1962) found some von Aosta specimens in Florence (FI or
FT). However, a loan from FT did not contain any Coccinia specimens by von Aosta.
Coccinia longipetiolata Chiov., Fl. somala 2: 223. 1932.
Type: Somalia. [Jubbada Dexhe/Jubbada Hoose border]: between Afmadù [Afmadow]
and Saamoggia, 1926, P. Gorini 149 (Syntype: FT [K neg. 4851, digital image: JPS]).
Type: Somalia. [Jubbada Dexhe/Jubbada Hoose border]: between Afmadù [Afmadow]
and Saamoggia, 1926, P. Gorini 150 (Syntype: FT [K neg. 4850, digital image: JPS]).
Remarks. e specimens are quite poor. No leaf is spread out, and generative characters are
missing. However, 7-lobate leaves, according to description, do only occur in C.samburu-
ensis, which diers in coriaceous leaves and a serrate margin with glandular teeth. Hence,
this species name is not synonymous with any Coccinia species. e tendrils in C.longipeti-
olata are almost equally bid, which is not found in Coccinia, especially not in species not
from rainforests. e drawing accompanying the protologue shows stipules, but this can
only be seen in a single node of P. Gorini 149, while the other nodes are more typical of
Cucurbitaceae. It shows, however, more likely a bud and a probract of similar sizes that give
the impression of stipules. In all, the specimens are likely to belong to the Cucurbitaceae.
ere are neither characters supporting a relationship with Coccinia, nor charac-
ters contradicting it, except for the tendrils. Jerey (1967) suggests a relationship to
his Coccinia sp. E sensu F.T.E.A. (Jarman 66), but as this specimen could not be ex-
amined by the present author, it cannot be discussed. erefore, that species is treated
as dubious. Eventually, sequencing could give disclosure about the relationships.
Monograph of Coccinia (Cucurbitaceae) 141
Invalid names
Coccinia abdallai Zimm., nom. nud.
e name is mentioned on P.W.A. Zimmermann G6594 (EA!) and in Die Cucurbita-
ceen 2: 8 (1922b) but not described. is is a Coccinia trilobata.
Bryonia acerifolia D.Dietr., Syn. pl. 5: 367. 1852, nom. nud.
Dietrich mentions this species in his synopsis as a name by Willdenow. However, no
such name by Willdenow is known. As Cogniaux (1881) synonymizes B. acerifolia
with B. alceaefolia Willd., and Dietrich uses the exact same words for B. acerifolia as
for B. alceaefolia, this name is likely just mistaken.
Bryonia barbata Buch.-Ham. ex Cogn. in A.DC.& C.DC., Monogr. Phan. 3:
530. 1881, nom. nud.
If the plate 625 in the East India Company’s Museum is additioned by a printed label
with description, then this name might be valid, but Cogniaux (1881) mentioned the
name as unpublished. He synonymized it with Coccinia cordifolia (see also there), so
it is not clear whether B. barbata is a C.grandis or a Cucumis maderaspatanus. How-
ever, the epithet suggests the existence of rigid trichomes, which does not match with
C.grandis.
Coccinia cordifolia var. triangularis A.Chev., nom. nud.
is name appeared on labels in P specimens of A.J.B. Chevalier 8886, 9527, and
10934, and it is apparently unpublished.
Coccinia crassifolia H.K.Walter, Naturwissenschaft und Landwirtschaft 9: 33.
1926, nom. nud.
is is a typographical error of Caccinia crassifolia Kuntze, a Boraginaceae.
Coccinia dubia Palacký, Lotos 10(4): 70. 1860, nom. nud.
Palacký cites a Coccinia dubia, which was supposed to be described by von Bunge in
his “Reliquiae lehmanniae” (von Bunge 1854). However, Palacký mistyped the genus,
which is in fact called Caccinia, a Boraginaceae.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
142
Coccinia glandis nom. nud.
is is a typographical mistake for C.grandis that has been published several times
(Tewtrakul et al. 2006, Jiwajinda et al. 2002). is epithet should hence not be used
in Coccinia.
Physedra gracilis A.Chev., Explor. Bot. Afrique occ.franç. 1: 292. 1920, nom. nud.
Ivory Coast: Bassin de la Moyenne-Sassandra, at Guidéko, A.J.B. Chevalier 16416 (P!
[P05591866, digital image: P]). Ivory Coast: Bassin de la Moyenne-Sassandra, at
Guidéko, A.J.B. Chevalier 19013 (P! [P05591864], P! [P05591865]).
is is a nom. nud. (ICN 32.3) because the note that the plants have yellow owers
cannot be regarded as intended to describe a new species. e specimens are Coc-
cinia keayana.
Cephalandra indica var. triangularis A.Chev., nom. nud.
is name appeared on a label by Chevalier from October 1908 on A.J.B. Chevalier
9527, but apparently was not published.
Cucumis inedulis Forssk., Fl. aegypt.-arab.: CXXII. 1755, nom. nud.
For details, see the Taxonomic remarks of Coccinia grandis, to which the name would belong.
Cephalandra ivorensis A.Chev., Explor. Bot. Afrique occ. franç. 1: 295. 1920,
nom. nud.
Collections connected to this nomen nudum have been synonymized with Physedra eg-
landulosa (Hook.f.) Hutch. & Dalziel (now Ruthalicia eglandulosa (Hook.f.) C.Jerey).
Bryonia lagenaria E.Mey. ex Drège, Zwei panzengeogr. Dokum. 54, 169. 1843,
nom. nud.
is name appears on some Drège collections and is only listed in the work of Meyer.
However, the collections are type specimens Coccinia sessilifolia.
Coccinia medica M.T.H.Khan in: Gottschalk-Batschkus and Green, Handbuch
der Ethnotherapien: 377, 384, 517, 537. 2002, nom. nud.
Khan used this name but without taxonomic context. Most likely, he meant Coccinia
indica, an illeg. name for C.grandis.
Monograph of Coccinia (Cucurbitaceae) 143
Coccinia monteroi Hort., Catalogue des graines du Jardin botanique de
Bordeaux. 1866.
e protologue was not available to the present author, but Cogniaux (1881) listed
this name as nomen tantum. It is, however, not mentioned in the notes sections of the
Bordeaux Garden catalogues from 1866 or 1867 where Naudin described some new
species, so it might well be just a nomen nudum.
Coccinia moshiensis Zimm., nom. nud.
Mentioned on P.W.A. Zimmermann G6599 (EA!). is is a Coccinia trilobata.
Coccinia natalensis Burtt-Davy, A manual of the owering plants and ferns of the
Transvaal with Swaziland, South Africa 1: 237. 1926, nom. nud.
Cephalandra natalensis Oliv., unknown.
Coccinia natalensis (Oliv.) Cogn., unknown.
e names of Cogniaux und Oliver are mentioned in Burtt-Davy and Pott-Leendertz
in Ann. Transvaal Mus. 3(3): 121. 1912. However, no citation is given. Neither Cepha-
landra natalensis in mentioned in Daniel Oliver’s Flora of Tropical Africa, nor any
Cogniaux publication with this name is known. Burtt-Davy writes in 1926 that the
name ‘appears to have been an unpublished MS. name’.
Coccinia lalambensis Schweinf. ex Penzig, Atti Congr. Bot. Int. Genova (1892):
342. 1893, nom. inval.
Eritrea. [Anseba Province]: Monte Lalambensis near Keren, c.2000m, 20 Mar, G.A.
Schweinfurth 568 (B, destroyed). [Northern Red Sea Province]: Habab, J.M.
Hildebrandt plant. Habab 1802 (LE?).
No description given, therefore this name is not validly published.
Cucurbita laevigata Bl.?, nom. nud.?
is name is written on a specimen in L herbarium (L0587542). e specimen was
part of the collection of C.G. C.Reinwardt but lacks collector, collecting site, and
date. One ink-written label solely states “1766.E.5138.” and the species name. Anoth-
er label, written with a pencil, says “Cucurbita laevigata” “mihi” and “Callelet W[…]”.
e last word is unreadable to the present author. Another specimen (L0587515) bears
a similar label with “1766.E.5138.”, however without a pencil-written label. Since
both specimens are Coccinia grandis, Cucurbita laevigata would be a synonym, if it had
been validly published. A Waitz collection (L0587563) bears the names “Cucurbita
laevigata Bl.” and “Callelet Bl.”, so the former name is maybe a Blume manuscript
name and the latter one is indigenous.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
144
Bryonia quinquefolia Noronha, Verh. Bat. Genootsch. 5: 155(8). 1790, nom. nud.
Miquel (1855) synonymizes this nomen nudum under C.wightiana M.Roem., which
is C.grandis (see Taxonomic remarks there). However, he also synonymizes Bryonopsis
pedata Hassk., which cites Bryonia quinquefolia. e description of Bryonopsis pedata
mentions lacinate, almost pinnatid leaves and male owers in oblong clustered ra-
cemes. is does not match C.grandis at all but eventually Diplocyclos palmatus, which
Roemer excluded from C.indica as Coccinia palmata M.Roem. e identity of Bryonia
quinquefolia might be solved, if one nds a Noroña specimen stating “Bryonia quin-
queloba” or “Oyot-kekèp”, the latter name being the Javanese term for this species.
However, the epithet rather links to the deeply lobate leaves of D. palmatus.
Bryonopsis pedata Hassk., Cat. hort. bot. bogor.: 189. 1844, nom. nud.
Hasskarl cites Noroña’s Bryonia quinquefolia and a vernacular name “aroy kalanyar
beurriet”. e given description of lacinate, almost pinnatid leaves and male owers
in oblong clustered racemes does not match C.grandis. According to Filet (1859), the
vernacular name is used in Sundanese and refers to Bryonia [sic] pedata Hassk., two
Trichosanthes species and Lua cordifolia Bl. None of these names have been referred to
Coccinia, so it seems unlikely that Bryonopsis pedata does.
Coccinia peterii Zimm., nom. nud.
is is an unpublished ms. name on R. Soleman 6046 (EA!). e specimen, however,
is a C.grandis.
Bryonia ruderalis Zipp. ex Span., Linnaea 15: 206. 1841, nom. illeg. & nom. nud.
In L herbarium, there is a specimen determined as “Bryonia ruderalis Zp.” from Timor
(L0587573), which is a Zippelius collection of Coccinia grandis. However, the name is
a later homonym of Bryonia ruderalis Salisb. Additionally, it lacks a description in the
publication, so it is a nomen nudum, too.
Cucurbita schimperiana Hochst., nom. nud.
e name was used on printed labels of G.H.W. Schimper 1570 (eective publication),
which lack a proper description (hence a nom. nud.). Under this distribution number,
specimens from two dierent shipments are included. One is taken from package “P.
16 K. no. 4”, collected on 23 Apr 1841 in Djeladjeranné (label on P specimen). e
data of this label were used for C.F. F. Hochstetter’s printed labels. e TUB-004724
and TUB-004725 specimens bear a Schimper label from package “P. 10 D. no. 23”
from “Landschaft Modat” collected in April 1839. An unnumbered W specimen also
Monograph of Coccinia (Cucurbitaceae) 145
notes this collecting site, hence the specimen might be from the same shipment. Speci-
mens of both collections are Coccinia grandis.
Coccinia schultzei Gilg, Namaland & Kalahari: 697. 1907, nom. nud.
Apparently a collection by L. Schultze (Schultze 320a) in B herbarium, but not validly
published by Gilg afterwards. However, if so, then the holotype was burned in the
Berlin herbarium re in 1943.
Coccinia sericea Zimm., nom. nud.
Zimmermann marked the specimen P.W.A. Zimmermann G6600 (EA!) to be a new
species, but B. Verdcourt pointed out on the specimen that Zimmermann never pub-
lished it. In any case, this specimen belongs to C.grandis.
Bryonia sinuosa Wall., Numer. List 6716. 1832, nom. nud.
e present author did not see a specimen with this number, so it cannot be decided
whether Cogniaux’ (1881) partial synonymization of Coccinia cordifolia refers to Coc-
cinia grandis or Cucumis maderaspatanus. Wallich himself supposed that this collection
is a mix of (Bryonia) Coccinia grandis and Melothria indica.
Cephalandra sylvatica A.Chev., Explor. Bot. Afrique occ. franç. 1: 295. 1920,
nom. nud.
Collections connected to this nomen nudum (ICN 32.3) have been synonymized with
Physedra eglandulosa (Hook.f.) Hutch. & Dalziel (now Ruthalicia eglandulosa (Hook.f.)
C.Jerey).
Cucurbita triangulata Hochst. ex Cogn. in A.DC.& C.DC.Monogr. Phan. 3:
532. 1881, nom. nud.
Cogniaux cites a Schimper specimen (Iter Abyss. Sect. 3 no. 1202) that was supposed to
be labeled by C.F. F. Hochstetter. ere are several sheets with this distribution num-
ber in Paris, but only one bears this name. e location is given by “In Semen” [Semien
Mts]. e other Paris specimens with this number are from Baria Dikeno (collected on
6 Aug 1853). e collection is a Coccinia grandis.
Coccinia wightii Miq., Fl. Ned. Ind. 1(1): 1112. 1855, nom. nud.
Name variation of Coccinia wightiana M.Roem. in the index of the book.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
146
Exluded taxa
Physedra bequaertii De Wild., Pl. Bequart. 1: 569. 1922.
Type: D.R. Congo. Along the Semliki [river], female, , 16 Jun 1914, J. Beq-
uaert 4791 (Syntypes: BR [BR0000008886330, digital image: BR, JPS], BR
[BR0000008887122, digital image: BR, JPS], BR [BR0000008887245, digital
image: BR, JPS]).
Type: D.R. Congo. Along Ruthuru river, female, , 17 Nov 1914, J. Bequaert 6315
[sic, should be 6215] (Syntypes: BR [BR0000008886477, digital image: BR, JPS],
BR [BR0000008886521, digital image: BR, JPS]).
Remarks. As Physedra is a synonym of Coccinia but the specimens are belonging to
the genus Bambekea, the name P. bequaertii has to be excluded. Jerey (1962: 364)
published the new combination Bambekea bequaertii (De Wild.) C.Jerey, however, it
is not clear whether this is a species separate from Bambekea racemosa Cogn.
Coccinia cordifolia (L.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 529. 1881. pro parte.
Bryonia foliis cordatis oblongis quinquangularibus dentatis scabris L., Fl. zeyl.: 168. 1747.
Bryonia cordifolia L., Sp. pl. 2: 1012. 1763.
Type: Sri Lanka. No detailed location given, P. Hermann 354 (Typotype: Herm. Flora
zeylanica 2:22, BM! [BM000521582, digital image: BM]).
Remarks. Cogniaux cites Bryonia cordifolia sensu Linnaeus’ Species plantarum 2nd
edition, where Linnaeus synonymizes Rumphius’ Vitis alba indica, which is Coccinia
grandis. If B. cordifolia would have been originally described in 1763, Cogniaux’ choice
would have been valid, but B. cordifolia was described in 1753 (Species plantarum 1st
edition), where Vitis alba indica is not mentioned, but only a collection from Hermann
herbarium, which is Cucumis maderaspatanus L.
Coccinia decipiens (Hook.f. in Oliv.) Cogn. in A.DC. & C.DC., Monogr. Phan.
3: 539. 1881.
Cephalandra decipiens Hook.f. in Oliv., F. T. A. 2: 552. 1871.
Type: Angola. [Cuanza Norte]: Pungo Adongo, grassland, F.M.J. Welwitsch 816 (Holo-
type: BM! [BM000799218], isotypes: COI [COI00005507, digital image: JPS], K
[K000313451, digital image: JPS, K], LISU [LISU00214555, digital image: JPS]).
Remarks. e sessile beaked fruits match to the genus Diplocyclos, which has been cor-
rectly observed by Jerey (1962) as Diplocyclos decipiens (Hook.f.) C.Jerey.
Monograph of Coccinia (Cucurbitaceae) 147
Coccinia ecirrhosa Cogn., Bull. Herb. Boissier 4(12): 822. 1896.
Type: Somalia. Abdallah, 1891, C. Keller 106 [sphalm. 116 in l.c.] (Type: BR!
[BR0000008885999, digital image: BR, JPS], Z! [Z-000004442, digital image: Z,
K neg. 4832]).
Remarks. e type does not contain much material, but the lower surface of a leaf
shows a pinnatid venation pattern, which is unknown in Coccinia. Jerey (1967)
transferred it correctly (Kocyan et al. 2007) to another genus, namely Cephalopentandra.
Coccinia gabonensis Keraudren, Adansonia 8: 40. 1968.
Type: Gabon. Ogooué-Ivindo: Bélinga, 950–1000 m, male and female, , Nov 1964,
N. Hallé 3018 (Holotype: P [P00348266, digital image: JPS, P], isotypes: P!
[P00348264, digital image: JPS, P], P! [P00348265, digital image: JPS, P], K!
[K000313237, digital image: JPS, K]).
Remarks. is species does certainly not belong to Coccinia. It is monoecious, has free
petals, and rather large bracts, in contrast to the dioecious, sympetalous Coccinia species
with much smaller bracts. It rather belongs to Momordica.
Coccinia macrantha nom. nud.
Physedra macrantha Gilg, Bot. Jahrb. Syst. 34: 356. 1904.
Type: Liberia. Gran Bassa: Fishtown, in bush, sand, , 10 m, 27 Aug 1898, M. Dinklage
2019 [Cucurbitaceae no. 1846] (Holotype: B! [B 10 0154925, digital image: B, JPS]).
Remarks. e combination Coccinia macrantha was only used in B herbarium for stor-
age but apparently never published. Physedra macrantha Gilg has been synonymized,
correctly, with Physedra eglandulosa Hutch. & Dalziel (1928), which is now in the
genus Ruthalicia.
Coccinia obbadiensis (Chiov.) Cufod., Supplem.: Enum. Pl. Aethiop. Spermato-
phyta 35(2): 1050. 1965.
Cephalopentandra obbadiensis Chiov., Fl. somala: 187, tab. 20, g. 1. 1929.
Type: Somalia. [Mudug]: Obbia [Hobyo] Sultanate, between Dolobscìo and Mag-
ghiòle, 27 Apr, G. Stefanini & N. Puccioni 458 (Holotype: FT [FT003569, K neg.
4845, digital image: JPS], isotype: K! [K000313183, digital image: JPS, K).
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
148
Remarks. e author notes ve stamens, which are a good reason for not including this
species in Coccinia. e leaves of the specimen on the picture look like these of Coccinia
quercifolia, which is also excluded from Coccinia and separated by Jerey (1967).
Coccinia quercifolia Hutch. et E.A.Bruce, Bull. Misc.Inform. Kew 2: 99. 1941.
Type: Somalia. [Nugaal]: Boundary Pillar 93, 45°9'E, 8°37'N, 990 m, , 6 Oct, J.B.
Gillett 4194 (Holotype: K! [K000313174, digital image: JPS, K], K! [K000313174,
digital image: JPS, K]).
Type: Ethiopia. [Somali Region]: Harradigit [c.7°45'N, 45°30'E], Apr, F.L. James &
J.G. rupp no.? (Paratype: K?).
Remarks. e leaves have a pinnatid venation, just like Cephalopentandra (Coccinia)
ecirrhosa, but are deeply lobed. Since the leaf form may vary in Cucurbitaceae, it is
likely that these specimens belong together, so that the odd leaf venation excludes this
type from Coccinia. Jerey already synonymized this name with C. ecirrhosa.
Coccinia palmata (L.) M.Roem., Syn. pepon.: 93. 1846.
Bryonia palmata L., Sp. pl. 2: 1012. 1753. Bryonia zeylanica, folio quinquepartito
Burm., es. zeylan.: 49. 1737. Bryonia foliis palmatis lævibus quinquepartitis:
laciniis lanceolatis repando-serratis L., Fl. zeyl.: 146. 1747.
Type: [Sri Lanka]. No location given. P. Hermann 25 (Type lost?).
Type: [Sri Lanka]. No location given. P. Hermann 41 (Type lost?).
Type: [Sri Lanka]. No location given. P. Hermann 62 (Type lost?).
Type: [Sri Lanka]. No location given. P. Hermann 353 [Musæum zeylonicum 2:58]
(Lectotype, designated by Jerey (1962: 352): BM [BM-000621700]).
Remarks. e name Coccinia palmata has been applied illegitimately for C.mack-
enii for a long time due to an overlooked combination. When Wight and Arnott
published the name Coccinia indica, they cited Bryonia grandis L. and also tenta-
tively included the citation of Bryonia palmata L. More likely, however, they meant
a specimen in Herbarium Madras that was identied as B. palmata L. One year
after Voigt’s correction to Coccinia grandis (L.) Voigt, Roemer (1846) also recog-
nized the seemingly missing combination and that Linnaeus’ B. palmata and B.
grandis indeed referred to dierent species. Roemer treated them, amongst other
species, as C.grandis (L.) M.Roem. (nom. illeg.) and C.palmata (L.) M.Roem.
Roemer cited the second edition of Linnaeus’ Species plantarum (1763), which
has the identical description for this species as the rst edition, so he explicitely
meant Bryonia palmata to be a part of Coccinia. e Bryonia palmata typolectotype
belongs to what is now widely called Diplocyclos, due to the globose striped fruits
Monograph of Coccinia (Cucurbitaceae) 149
easily recognizable as being not part of Coccinia. Apart from that, another spe-
cies from South Africa was described with the name Cephalandra palmata E.Mey.
ex Sond. (Harvey and Sonder 1862). Cogniaux (1881) accepted this species in
Coccinia but overlooked Coccinia palmata (L.) M.Roem. He thus created an il-
legitimate Coccinia palmata (E.Mey. ex Sond.) Cogn., which has since been used
for this species. Holstein and Renner (2010) called attention to this erroneous
usage by resurrecting the correct name, Coccinia mackenii Naudin ex C.Huber,
while C.palmata is a synonym of Diplocyclos palmatus in spite of its wide usage for
C.mackenii since then.
Coccinia petersii Gilg, Bot. Jahrb. Syst. 34: 356. 1904.
Type: Mozambique. [Zambézia]: Rios de Sena and Boror, without detailed locality,
male and female, on dry ground, 1846, W.H.C.Peters s.n. (Holotype: B destroyed,
isotype: K).
Remarks. Jerey (1967) synonymized this species with Eureiandra fasciculata (Cogn.)
C.Jerey.
Coccinia polyantha Gilg, Bot. Jahrb. Syst. 34: 356. 1904.
Type: Tanzania. Lindi: Kilwa district, near Mariwe, upon low shrub, in light, slightly
wet ground pori, , Dec, W. Busse 512 (Syntype: B destroyed, isosyntype: EA
[EA000002139, digital image: JPS]).
Type: Tanzania. [South central Tanzania], Kissaki steppe at Ruji river, 250 m, on
sandy laterite soil, fr, Nov, Götze 80 (Syntype: B destroyed).
Remarks. e seeds are described as globose to subglobose, but Coccinia seeds are
rather at. It is therefore unlikely that this species belongs to Coccinia. Jerey (1967)
synonymized this species with Eureiandra fasciculata (Cogn.) C.Jerey.
Cucumis rheedii Kostel., Allg. med.-pharm. Fl. 2: 738. 1833.
Schunambuvalli Rheede, Hort. malab. 8: 21, t. 11. 1688.
Type: drawing in l.c.
Remarks. Cogniaux and Harms (1924) synonymized this species with Coccinia cor-
difolia (L.) Cogn., a name that Cogniaux used for C.grandis (L.) Voigt. However, the
drawing shows 3–5-d tendrils and small? subglobose fruits on an ebracteate panicle.
is does not correspond to Coccinia.
Norbert Holstein / PhytoKeys 54: 1–166 (2015)
150
Coccinia stefaninii Chiov., Result. Sc.Miss. Stefan.-Paoli Somal. Ital. 1: 84. 1916.
Type: Somalia. Somali Republic, Jubaland, Dintorni di El Uré, G. Paoli 1069 (Syntype:
FT! [2 sheets, FT003512, digital image: JPS]).
Type: Somalia. Fra Jagdoudou e Duddumai, G. Paoli 1179 (Syntype: FT! [FT003513,
digital image: JPS]).
Remarks. is name has been transferred to the genus Dactyliandra by Jerey (1985).
Coccinia stolzii Harms, Bot. Jahrb. Syst. 57: 241. 1923.
Type: Nyasaland [Tanzania]. Kyimbila district, Mbaku, 600m, , fr, Jun 1913, A.F.
Stolz 2028 (Holotype: ?, isotype: BM!, G! [G00301602], K [K000313456, digital
image: JPS, K], L!, P! [P05590096, digital image: P], PR!, PRE [PRE0592846-0,
digital image: JPS], S [S08-12469, digital image: JPS], U! [U0074536], W!, Z!
[Z-000004440, digital image: Z]).
Remarks. Monoecious plant with several small subglobose fruits per node apply clear-
ly to Diplocyclos and must therefore be synonymized as it has been done by Jerey
(1967) to Diplocyclos decipiens.
Coccinia suburceolata Cogn., Symb. Antill. (Urban) 1: 450. 1900.
Type: Haiti. Near Port-au-Prince, in Tête bois de pin, 1800 m, male, , Nov, L. Picarda
1498 (Holotype: BR! [BR0000009939141, digital image: JPS], isotypes: B, L, NY, S?).
Remarks. is plant was found on Haiti and is therefore geographically far away from
the natural distribution range of the genus Coccinia. According to the protologue, the
tendrils are often trid and the fruit is spherical and apple-sized, which does not t
to the morphospace of any Coccinia species. Urban (1921) put the specimen and thus
species, amongst others, into a new genus: Penelopeia, which has been conrmed by
Kocyan et al. (2007).
Acknowledgements
I thank the curators of B, BAR, BM, BR, BRI, C, CANB, CBG, COI, DSM, EA, FR,
FT, G, GAT, GOET, H, HBG, HEID, JE, K, L, LISC, LISU, M, MO, MSB, NHT, P,
PERTH, PR, PRC, S, U, UBT, UPS, W, WAG, Z, ZT, and Charlie Jarvis for making
the collections available, sending loans or scans, or helpful comments about the col-
lections. I am grateful to Jean-Luc Gatard for sending seeds and plants, without which
Monograph of Coccinia (Cucurbitaceae) 151
most of the morphological ndings and the crossing experiments would not have been
possible. I also thank Frank M. Mbago for his help and excellent eye on the eld trip
in Tanzania.
anks are given to Hanno Schäfer for communicating various observations and
specimens from DNA, NY, US, Christian Bräuchler for communicating and discus-
sion of Schimper specimens from TUB, and Eberhard Fischer for communicating
material from Rwanda. Shixiao Luo is thanked for help with translations from Chinese
language, Werner Greuter for clarifying doubts with the validity of Cucurbita exan-
thematica, and Brigitta and Willem de Wilde for discussion of the Momordica bicolor
specimens and sharing their eld experience with C.grandis. I thank Eva Facher for
help with staining of the petiole sections and Mila Vosyka for preparing the chromo-
somes. anks are given particularly to Marc Gottschling for many fruitful discussions
and advice on the ms., esp. in taxonomic questions and Susanne S. Renner for supervi-
sion and English revision. I also thank Sandy Knapp and two anonymous reviewers for
many helpful comments on the manuscript.
is work has been nanced by Deutsche Forschungsgemeinschaft (RE603/6-1
and 6-2 given to S.S. Renner). Travel to the herbaria, BR, P, and W have been funded
by travel grants by Munich Graduate School for Evolution, Ecology and Systematics
of the LMU Munich. is research received support (visit to L and U collections) from
the SYNTHESYS Project (http://www.synthesys.info/), which is nanced by Europe-
an Community Research Infrastructure Action under the FP7 “Capacities” Program.
I also like to acknowledge the Tanzania Commission for Science and Technology
(COSTECH) for providing research clearance to undertake fieldwork in Tanzania.
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Supplementary material 1
List of examined specimens and geo-references of the localities
Authors: Norbert Holstein
Data type: specimens data.
Explanation note: Voucher information and (predominantly inferred) coordinates of
geo-references (in decimal degrees). Specimens included in this study, with the geo-
graphic origin of material (only country and 1st administrative division given).
Herbarium acronyms follow Index Herbariorum; barcodes or other unique identi-
ers in brackets behind the acronym. Indication of digitized images refer to the
homepage of the corresponding institution indicated by the herbarium acronym,
except for JPS (JSTOR plant science) and CVH (= Chinese Virtual Herbarium).
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Supplementary material 2
GenBank accessions
Authors: Norbert Holstein
Data type: specimens data.
Explanation note: Voucher information and GenBank accession numbers. Specimens
included in this study, with the geographic origin of material (only country and 1st
administrative division given) and GenBank accession numbers for all sequences.
Herbarium acronyms (in parentheses behind the voucher name) follow Index.
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Available via license: CC BY 4.0
Content may be subject to copyright.
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