Monograph of Coccinia (Cucurbitaceae)

Abstract This monograph deals with all 95 names described in the Cucurbitaceae genus Coccinia and recognizes 25 species. Taxonomic novelties are Coccinia adoensis var. aurantiaca (C.Jeffrey) Holstein, stat. nov., Coccinia sessilifolia var. variifolia (A.Meeuse) Holstein, stat. nov., and Coccinia adoensis var. jeffreyana 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, Coccinia grandis, extending from Senegal in West Africa east to Indonesia and being naturalized on Pacific Islands, in Australia, the Caribbean, and South America. Coccinia species are dioecious creepers or climbers with simple or bifid tendrils that occupy a range of habitats from arid scrubland, woodlands to lowland rainforest and mist forest. The 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. The globose to cylindrical fruits contain numerous grayish-beige flat 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 Coccinia grandis, of which leaves and sap are used against diabetes.


Introduction
Coccinia Wight & Arn. comprises 25 species and is the 11 th largest of the 97 genera of the Cucurbitaceae (Schaefer and Renner 2011a). Especially in the 19 th 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;Bhaduri and Bose 1947;Chakravorti 1948;Datta 1988;Kumar and Deodikar 1940;Kumar and Vishveshwaraiah 1952;Roy 1974;Roy and Roy 1971b). Th 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;Jeff rey 1967;Jeff rey and Fernandes 1986;Kéraudren-Aymonin 1975a;Kéraudren 1967;Meeuse 1962). Th e position of Coccinia in the Benincaseae has been confi rmed 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).
Th e delimitation of Coccinia from other genera is diffi cult. Th 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. Th 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 diff erent genera. In all, 113 names at various ranks have been proposed for what are here considered 25 species. Th e species concepts in the present revision are based on 3157 herbarium collections and fi 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 eff ort provide a modern understanding of the evolution and species relationships in Coccinia.

General morphology
During this study, the present author examined 3157 herbarium collections from 39 herbaria from (physical or digital) loans or in situ ( New collections were added if the photograph allowed identifi cation or if misidentifi cation appeared to be unlikely (esp. C. grandis collections from the Pacifi c area), while duplicates 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.

Phylogenies
For this monograph the phylogenetic data of Holstein and Renner (2011a; 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). Th 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. Th e fi 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/fi 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 diff erent 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 according 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 coordinates 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 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). Th 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.

Habit
Coccinia species are perennial climbers or creepers. Th e lignifi cation of the mature shoots diff ers among the species from unlignifi ed to completely lignifi ed. Climbing is enabled by tendrils, which are either simple or bifi d. Tendril development in young plants is delayed and emerges in C. abyssinica after the 6 th node (Getahun 1974a). Th e tendril arms are only rarely equally sized, as one is usually much smaller; true dichotomy of tendrils is unknown from Coccinia. Whether a species has simple or bifi d tendrils is often not fi xed, but there is a strong predominance of one kind. Bifi d tendrils regularly occur or are predominant in C. grandifl ora Cogn., C. heterophylla (Hook.f.) Holstein, C. hirtella Cogn., C. intermedia Holstein, C. mackenii Naudin ex C.Huber, C. mildbraedii Gilg, C. racemifl ora Keraudren, C. schliebenii Harms, and in some forms of C. barteri (Holstein and Renner 2011b). Strikingly, Coccinia species with bifi d tendrils occur in rather humid habitats. Th 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 fl oristic treatments, but they may bare bifi d tendrils such as C. sessilifolia (N. Holstein 13) and C. senensis (H.J.E. Schlieben 5745 in B, K, and MO). Coccinia adoensis has bifi d 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 bifi d tendrils: C. sessilifolia with C. hirtella and C. mackenii, and C. adoensis with C. grandifl ora 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 Coccinia adoensis and C. grandifl ora (and most likely also C. senensis (Klotzsch) Cogn. 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 fi re, as this species predominantly occurs in woodlands. In contrast to rather mild fi res in semiarid savannas with less infl ammable biomass, woodland fi res produce temperature rises of 60 °C in 0-3 cm depth (Bradstock and Auld 1995;Gignoux et al. 1997), so vegetative 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). Th ere is barely any research on root anatomy, 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. microphylla, C. rehmannii, C. sessilifolia, and C. trilobata, produce a lignifi ed tuber that is derived from the hypocotyl (Zimmermann 1922b;pers. observ., Fig. 1a). Th e tuber, at least of some species, contains starch as a storage nutrient (Getahun 1974b). It develops during the fi rst season, and lignifi cation may begin as soon as the appearance of the fi rst tendrils, such as in C. abyssinica (N. Holstein 132). Some species, such as C. adoensis and C. grandifl ora (and most likely also C. senensis and C. schliebenii) do not produce hypocotyl tubers but root tubers. In C. adoensis var. jeff reyana the hypocotyl is 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 fi re or grazing. Coccinia microphylla shoots can lignify completely and produce short green branches with fl 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). Th e shoots of C. grandis can become slightly succulent. Th e length of the shoots varies from 70 cm in C. microphylla to 20 m in C. grandifl ora and C. mildbraedii. Zimmermann (1922a) reports a stem of C. grandifl ora 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). Th e indumentum of Coccinia species, if present, is composed of simple, oligo-to multicellular eglandular trichomes up to 2 mm in length. Th 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 off . Th 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 petioles, 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 trichomes are rare, few-celled, not visible with the naked eye and have been found, e.g., in C. grandifl ora and C. grandis (pers. observ.;Th 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.  Holstein et al. 90); picture taken during the dry season. Th e stem is completely lignifi ed, and only green short shoots are produced b Male plant of C. sessilifolia. Th e stem is glaucous and does not lignify. Unusually, the bract is 3-lobate leaf-like.

Cotyledons
Zimmermann (1922b) reports epigeous cotyledons for C. grandifl ora and C. grandis, of which the latter is confi rmed by personal observations (Fig. 1a). Epigeous cotyledons also occur in C. abyssinica, C. adoensis var. jeff reyana, C. microphylla, C. rehmannii aff . var. littoralis, and C. sessilifolia. Th e hypocotyl and cotyledons of all observed taxa are glabrous. Th e cotyledons are elliptical to obovate and have an entire margin. Th e cotyledons are slightly fl 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 confi rmed, and if multicellular trichomes occur, then they are not visible to the naked eye. Th e fi 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
Th e leaves of Coccinia species are simple, alternate, and paired with a tendril on each node, except for the fi 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. Th 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. subsessilifl ora), or have an indumentum. Th e petiole contains several vascular bundles arranged in a U-shape (Fig. 5a). However, Hussain et al. (2011) report a ring of vascular bundles in C. grandis. Th e adaxial side of the petiole often bears two ridges above the "lateral" vascular bundles (Fig. 5a). Th ese ridges merge into the leaf margin and usually bear trichomes (Figs 3b,5b,c). Th e abaxial side of the petiole shares its indumentum with the lower leaf lamina, at least at the base of the veins (Fig. 5c).
Th 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 confi rmed for all Coccinia species except C. ogadensis, in which only the central vein in each lobe is visible.  (N. Holstein 132). Th e fi rst node is in the axilla where the glabrous cotyledons split off . Th e fi rst nodes lack probracts and tendrils b Young plant of C. sessilifolia (N. Holstein 131). Th e fi rst leaves in this species are petiolate, sessile leaves are produced later-on. Th 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. grandifl ora, stained with astra blue and safranin (3:2). Th 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. grandifl ora. Although not visible by naked-eye, the petiole is also covered with few-celled glandular trichomes c Young plant of C. adoensis var. jeff reyana. Th e trichomes are mainly occurring on the prominent veins. Th e adaxial side of the petiole bears smaller trichomes on the ridges, which fade into the leaf margin.
Young leaf buds often bear a dense indumentum, even in species that are glabrous at maturity, e.g., in C. grandis. Th e leaf lobes are linear, elliptic, (ob-)ovate to triangular. Th e incision depth of the lobes can be consistent (C. ogadensis, C. subsessilifl ora) 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 aff . var. littoralis, C. sessilifolia), a diff erentiation according to light exposure might also be possible. Th 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). Th e teeth are at the apex of lobes, lobules and smaller orders of serration or situated along the entire margin. Th e term "dentate" (toothed) is therefore ambiguous in literature describing Coccinia, as it might also refer to the margin morphology (Stearn 2004). Th 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). Th e coloration of teeth is inconspicuous in young plants and develops during maturation (as observed in C. abyssinica, C. grandis, and C. megarrhiza). Th 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 2 nd order (except those of the tip of  Torre et al. 18788 (MO). Black bars equal 1 cm. 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.
Th e upper leaf lamina is often covered with transparent to white pustules, that contain cystoliths (Avetta 1894;Solereder 1899;Zimmermann 1922a;pers. observ.). Th 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 CaCO 3 (pers. observ.). Th is can be observed in C. grandis, C. hirtella, and C. sessilifolia, hence also when the pustules are not conspicuous as in the latter two species. Th e pustules may form the base of small trichomes, such as in C. adoensis var. jeff reyana (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. Th 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.
Th e lower leaf lamina is paler than the upper side ( Fig. 3b) and can be glabrous or bear an indumentum. Th e highest density of the indumentum can be found on the prominent veins (Fig. 5c). Th e indumentum on the lower leaf surface and the abaxial surface of the petiole can consist of eglandular oligo-to multicellular trichomes. Th e trichomes are appressed or upright (Fig. 5c), usually fi liform, sometimes also narrowly conical (e.g., C. abyssinica). Filiform trichomes are straight, curved, or sinuate. Long fi liform trichomes often appear articulate when dry due to sunken lateral cell walls (Fig. 3a). Dry trichomes are hyaline, whitish, beige, or yellowish. Th e lower lamina often displays deeply colored to dark green to blackish extranuptial glands (Fig. 7b). Th e glands usually occur at the base of the leaf between the veins, sometimes also between secondary ramifi cations (C. grandis) or along the main veins (C. grandifl ora). Th 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 sterile nodes, which are called "probracts" (Zimmermann 1922b). Th e probracts can be up to 5 mm long, but also rather small (< 1 mm) or caducous. Th e fi rst nodes of the seedling lack probracts, and they are developed on later nodes. Th e shape of the probracts is ovate and entire with a round to acute apex. Th ey are often spoon-like presenting the lower surface (e.g., C. adoensis, C. barteri, C. grandis, C. megarrhiza, C. sessilifolia; Figs 3a, 8b), or they are folded in the middle with a prominent keel (C. grandifl ora; Fig. 8a). Probracts can be glabrous or bear short (< 1 mm) trichomes, and bear extranuptial glands on the lower surface ( Fig. 8a; Okoli and Onofeghara 1984).
Bracts (leaves subtending infl orescences or fl owers), if present, look like the probracts. Bracts below infl orescences are as large as probracts, while bracts below fl owers tend to be smaller. Bracts can be present or absent, the latter being an indicative character for some species.

Extranuptial glands
Th 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 fl 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.). Th is sheath is lignifi ed 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 fi nd such a statement in the citations given in that paper (eventually, this was a personal communication). Schrödter (1926), however, fi nds that young sheaths in Luff a aegyptiaca are lignifi ed but become suberinized with age, so the diff erence might be explained by diff erent stages. Chakravarty (1948) interprets the sheath as fi 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) fi 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. Th e glands secrete a clear, rarely slightly colored, sweet-tasting exudate (pers. observ. in C. grandifl ora and C. grandis). In C. grandis the exudates contain sucrose, glucose, fructose, alanine, tryptophane, threonine, and an unidentifi ed amino acid (Muhammad Ilyas 1992).

Peduncles and pedicels
Male fl owers mostly occur in racemes that are usually accompanied by 1-2 solitary fl owers on the same node (Fig. 2b). Th e fi rst fl owers in male plants of C. hirtella, C. reh mannii, and C. sessilifolia are solitary. Racemes appear later in the course of the fl owering season, although racemes are generally rare in the fi rst species (pers. observ.). If solitary fl owers and racemes are produced on the same node, then the solitary fl ower(s) precede(s) those of the racemes in time of maturity (Fig. 2b). Th e trigger to produce racemes instead of or additionally to solitary fl owers is not known. Th e racemes bear up to 35 fl owers (e.g., in C. pwaniensis, C. racemifl ora). Within the racemes, fl owering starts at the basalmost branches. If the peduncle is reduced, fl owers Figure 8. a Th e probract of C. grandifl ora is fl eshy and has a keel. Th e abaxial side bears many extranuptial nectaries. Th e structure pointing to the lower left of the picture is the tendril b Th e probract of C. trilobata (sampled as N. Holstein & P. Sebastian 9) is spoon-shaped, papery, and without a keel. appear clustered on the node. In C. grandis, which usually produces single fl owers only, fl ower clusters (short-peduncled racemes) rarely occur. Th is can be seen in plants from Ethiopia, Saudi Arabia but also from India and Sri Lanka. Th e pedicels of solitary male fl 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 fl ower upright.
Female fl owers are mostly solitary. Only in some species, female fl owers are usually in racemes, such as in C. heterophylla, C. keayana, and C. racemifl ora. Few-fl owered female racemes or clustered fl owers might also occur in C. grandifl ora, C. intermedia, and C. subsessilifl ora. In C. barteri, female fl owers can be solitary or in few-or manyfl owered racemes. Two female fl owers per node have also been observed in C. microphylla. Th e pedicels of solitary female fl owers are negatively gravitropic during fl ower development. After pollination, the pedicels of solitary female fl owers of C. grandis, C. hirtella, C. megarrhiza, C. microphylla, C. rehmannii, and C. sessilifolia exhibit positive gravitropism. Th 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 fi rm. However, only fertilized fl owers turn downwards completely, as aborted fl owers from mispollination never reach this state (pers. observ. in cultivated plants).

Perianth
Th 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 funnel. Depending on the exsertion point of the staminodes in female fl owers, parts of the tube form a hypanthium (e.g., C. grandifl ora).
Th e calyx diff erentiates as a bulge (Fig. 7a,b) with usually fi 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 diff erentiated 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-diff erentiated (congenital fusion) or postgenitally fused, is not known for Coccinia, but in the distantly related Echinopepon wrightii (A.Gray) S.Watson the perianth tube is non-diff erentiated (Leins and Galle 1971). Th e outside of the perianth tube can bear long trichomes of the type on as the lower leaf surface or the petioles (Fig. 10a). Th e calyx lobes are acute triangular to subulate or linear, rarely slightly lanceolate. Th e orientation of the calyx lobes is erect, spreading, or refl exed, although they can be curved inwards (e.g., C. rehmannii aff . var. littoralis (Fig.  10a) or outwards (e.g., C. intermedia). Th 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).
Th 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. observ. in C. megarrhiza, C. rehmannii var. rehmannii, and C. sessilifolia). Perianth tube and corolla tube are often campanulate, rarely funnel-shaped or tubular. Th e perianth tube can be urceolate in C. longicarpa, C. racemifl ora and sometimes in C. barteri. Th e tips of the (4-)5(-7) corolla lobes are rounded to acute with an apical tooth. Th e Figure 9. Leaky dioecy in a plant of C. megarrhiza. Th e plant was fl owering male (the bud on the left) through the season, but a single female fl ower developed (the second fl ower on this node was male). Th e fl ower was receptive and produced a normal-sized fruit and normal-shaped seeds. Th e probracts (left node) are spoon-shaped, the tendrils are purplish.  Holstein 126). Th e plant was raised from seeds of a female plant with ovoid fruits (B. Jarret -pers. comm.) b Male fl ower of C. adoensis var. aurantiaca (N. Holstein et al. 85). Th e halictid bee (H. Schaefer -pers. comm.) circled around the globose anther head harvesting pollen. Th e scent of the fl ower was strong and honey melon-like. 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). Th e trichomes become shorter towards the receptacle. Th e inner side of the hypanthium of female fl owers is glabrous and smooth, which suggests nectary tissue, in C. grandifl ora, C. grandis, and C. hirtella (Fig. 11a). Th e size of the corolla does not diff er conspicuously between staminate and pistillate fl owers; pistillate fl owers might be a bit smaller.

Androecium
In staminate fl owers, the three stamens originate from the base of the perianth tube, and the fi laments are fused to a central column (Fig. 10a). Th e fusion point sometimes leaves a small gap to the hollow receptacle center. Th is gap, however, can be fi lled with long multi-cellular trichomes (e.g., in C. abyssinica and C. megarrhiza). Rarely, the fi laments can also be separate. Th e fi laments are glabrous and white, greenish, yellowish, or orange. Th e number of vascular bundles in the stamens is disputed. Chakravarty (1954) reports fi 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 Deshpande et al. 1986). In C. hirtella each stamen contains a single vascular bundle (pers. observ.).
Th e anthers together form a globose head (Fig. 10b). Each anther is bithecate; sometimes one can be monothecate Chakravarty 1954). Each theca is sinuate. Deshpande et al. (1986) report a bi-layered fi brous endothecium and a secretory tapetum, which they found diff ers from distantly related Momordica charantia L. In pistillate fl owers, the three, now free, stamens are reduced to staminodes that originate from the interior perianth wall, forming a hypanthium. Introrsely, the staminodes of C. grandifl ora, C. grandis and C. hirtella bear long, multicellular trichomes, except for the apex, extrorsely the staminodes are glabrous (Fig. 11a). Th e anthers of the staminodes are strongly reduced to a slightly yellowish spot at the apex in C. grandis and C. hirtella. Th e staminodes of C. megarrhiza bear long multicellular trichomes introrsely and laterally but are glabrous extrorsely.

Pollen
Pollen in Coccinia species shows little diversity. Th e pollen is oblate-spheroidal to prolate 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. Th 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. Th e color is yellow in C. abyssinica, C. grandifl ora (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. grandifl ora 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 (I 2 KI) 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).

Gynoecium
Pistillate fl owers are epigynous and have three (rarely two or four) carpels. Th e ovary is narrowly spindle-shaped, oblong to globose. Th e surface is smooth or warty; it is glabrous or has the indumentum of the pedicel. Th 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. grandifl ora, C. grandis, C. hirtella, C. megarrhiza, C. rehmannii, and C. sessilifolia is U-shaped with the ends of the lower sides of the arms touching each other. Th e stigmatic branches can be long and free, such as in C. grandifl ora, C. grandis and C. rehmannii var. rehmannii (Fig. 11b), or short and bulbous, such as in C. hirtella, C. megarrhiza (Fig. 11c), C. microphylla, and C. sessilifolia.
Th e placentation of the ovules in Coccinia is involute, which is also discussed for other Cucurbitaceae by Leins and Galle (1971). Th 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. Th e axis-wall septum, however, might be reduced during ripening, but this needs further study.
In staminate fl owers, a pistil is not developed because the stamens fuse to a central column. Th e pedicel is narrow and reaches the perianth, and there is no indication of even a thin (sterile) inferior ovary in the fl ower.

Female gametophyte development and embryology
Th e development of the embryo has only been investigated in C. grandis. Chakravorti (1947) and Zahur (1962) report that the female gametophyte development is according to the Polygonum-type. Both observe that the synergides possess hooks instead of the fi liform apparatus. Chakravorti (1947) describes the developing endosperm as a nuclear type, which is confi rmed 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. Th en, the haustorium becomes cellular with multinuclear cells except for the apex (Chopra 1955).

Fruits
Th e fruits are many-seeded berries, which vary in size and shape between species (and within C. grandis, C. rehmannii, and C. subsessilifl ora). Th e smallest fruits occur in 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. Th e largest fruits occur in C. samburuensis and the rainforest species C. grandifl ora, 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. Th e ovules are anatropous with the micropyle facing outwards b Cross-and longitudinal section of a C. megarrhiza fruit. Th 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. Th 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. Th e left fruit is derived from pollination with C. megarrhiza pollen, whereas the smaller fruit on the right is derived from crosspollination with C. trilobata (both pollinations were conducted on the same day). curs from green with or without mottling via yellow to orange to the fi nal coloration. Th 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). Th 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 collected 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 ripening, the indumentum is usually reduced. Th e exocarp of Coccinia fruits is papery thin and has a waxy bloom when ripe. Th e endo-and mesocarp are red, fl eshy and soft (Fig. 12b,c). Th 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. Th e sieve tubes are not connected to the main vascular strands and are fi lled with a proteinaceous material. Th e authors suggest that the sieve tube network aids nutrient transport during the rapid growth of the fruit.
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 embryo 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 develops solely from the outer integument. Th 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. Th e outermost layer, the epidermis, is disintegrated, leaving the cell walls as slender rods of 500 μm length. Th is has also been noticed in other species, and the surface has been described as a fi brillose testa (Jeff rey 1967; Kéraudren 1967). De Wilde et al. (2011) also interpret the seed surface of C. grandis as having a disintegrated, pulpy, radiately striate exotesta.
Getahun contrasts his observations with those of Chakravorti (1947) in C. grandis, but the seeds are in fact similar, just incompletely described by Chakravorti. Chakravorti 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".
Th is is what Getahun calls parenchyma. Chakravorti's outermost layer, the epidermis, consists of radially elongated cells with thin walls. Th ese cells have likely just not yet disintegrated as observed by Getahun. Th e only diff erence 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. Th ese diff erent observations are explainable by two possibilities: 1) diff erent 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) diff erent staining. Chakravorti uses haematoxylin alone, which does not stain lignifi ed cell walls, whereas Getahun uses haematoxylin with safranin as counter stain, which makes lignin, and thus sclerenchyma, well visible (von Aufseß 1973).
Th 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. subsessilifl ora (pers. Figure 14. Seeds of Coccinia. Th e lack of fi 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 fi bers. Length of white bars equals 1 mm. a Seeds of C. adoensis var. jeff reyana (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 maceration) e Seeds of C. sessilifolia (harvested by mechanical extraction; taken from N. Holstein 119) f Seeds of C. grandis (harvested by mechanical extraction).
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. microphylla. 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
Th 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 germinating 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 Th ulin (3 seeds tested) did not germinate after fi ve years (pers. observ.). 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. Th 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). Th 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 declines 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 artifi cial hybrid (C. megarrhiza ♀× C. rehmannii aff . 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.

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) to 3-4 times longer (Guha et al. 2004) than the other chromosomes. Some years before Kumar and Deodikar, Sutaria (1936) reported n = 12 from pollen mother cells of C. grandis, without fi 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 mentioning whether he studied a male or a female individual. Th e author's own chromosome counts (Table 2; Fig. 15b) support McKay's report and reject the existence of heteromorphic sex chromosome in C. hirtella males. Th 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 off spring 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 fi 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. Th 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; Roy and Roy 1971b). Triploids of C. grandis with a 3n = XYY constitution are also male but bear fl 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 fl owers of the opposite sex in succeeding seasons. Two plants marked as female and one as male from observation of fl owers produced fl owers of the opposite sex in the following year (pers. observ.), making C. hirtella functionally dioecious, but genetically hermaphroditic. On the other hand, there are several observations of fl 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 fl owers are reported to develop, pollen grain development is arrested, and the male function remains suppressed. Roy and Saran (1990), however, report fully fertile hermaphroditic fl owers in an otherwise female individual. Holstein and Renner (2011a) report a collection of C. intermedia that bears male fl oral buds and female fl owers and young fruits on the same nodes. Th 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 produced a single female fl ower towards the end of the season (Fig. 9). Although two male fl owers on the same individual were open at the same time when the female fl ower was mature, the pollen sacs did not open. It is not known whether this was a coincidence or functionally signifi cant, e.g., to prevent selfi ng. Selfi ng is often discussed as being advantageous in small population sizes, e.g., when new islands are colonized. Prevention of simultaneous fl 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 fl ower was receptive and was fertilized by another male C. megarrhiza plant derived from the same fruit as the "female" plant. Th e resulting fruit and seeds developed normally.
Th e production of hermaphroditic fl 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 fl owers on two plants with otherwise female fl owers and an XX confi guration. Th ey also report the development of a normal fruit without mentioning the fertility of pollen from hermaphroditic fl owers, but interpret their fi nding as cleistogamy. However, fruit development without previous pollination (parthenogenesis) or from pseudogamy with pollen from diff erent genera as fructifi cation stimulus is described by Lal (1973). True selfi ng 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 dosages 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). Th is might indicate that the single X bears many functionally important genes in contrast to the Y, as mutations in the single X lead to an increased mortality compared to females with a balancing second X chromosome.

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. Th is means that the diff erence 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, however, 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 Cucurbitaceae (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 diff erent results as they fi 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 diff erence, which cannot be explained by diff erence 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 eff ects 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 schimperi, however, has buff petals, whereas the Asian C. grandis has snow-white petals. Both supposed species hybridized without problems. During the following two years, Naudin could not intercross within the F1 generation because plants of diff erent sexes did not fl ower at the same time, so he crossed F1 individuals with a female C. grandis from Asia, which again produced off spring. 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 buff -petaled, African C. schimperi and the white-petaled Asian C. grandis are a single species obeying the biological species 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-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). Th 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. mackenii) 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 defi ne 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, crossing experiments among species that are cultivated in Munich Botanical Garden have been performed. Positive results are given in Table 3.
Interspecifi c fertilization succeeded or failed without correlation of relatedness or co-occurrence (Table 4). Female fl 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 fl owers were discarded like non-fertilized fl 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 fl owering time and fl oral syndrome, the production of hybrids would reduce fi 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. sessilifolia, although the species neither co-occur, nor are closely related. Th e cross resulted in off spring, 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 fl owers was also not able to fertilize C. sessilifolia (1 trial). Th e occurrence of sex chromosomes in C. grandis might result in gene dosage imbalance, which interferes with the fl oral development, leading to sterile off spring. Th e inability of female C. grandis to be fertilized by C. rehmannii aff . var. littoralis and C. trilobata (Table 4) might be explained by the fact that the chromosome numbers diff er (see Table 2) and translocations lead to gene loss in hybrid genomes and thus inviability of the off spring. On the other hand, the cross between female C. hirtella and male C. trilobata (not sympatric) produced a purely intermediate F1 generation, which fl owers vigorously despite the diff erence in chromosome numbers (see Table 2 and 3). Although the anthers open like in fertile fl owers, unlike in C. grandis hybrids, the pollen of this hybrid was not able to fertilize female fl owers of C. hirtella (1 trial), C. grandis (2 trials), or C. sessilifolia (2 trials).   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 diff erent 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 aff . var. littoralis ♂ Abortion of fl ower (1 trial) C. grandis ♀ × C. trilobata ♂ Abortion of fl ower (2 trials) c C. hirtella ♀ × C. rehmannii aff . var. littoralis ♂ Onset of fruit (1 trial) c C. hirtella ♀ × C. sessilifolia ♂ Onset of fruit (1 trial) Abortion of fl 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 morphologically intermediate F1 off spring 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?] 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.

Pollination
Although bee pollination is observed for only a few species, most Coccinia species exhibit characters that support a general attraction to bees. Th 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. grandifl ora, C. grandis, C. hirtella, C. megarrhiza, C. microphylla, C. rehmannii, 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. grandifl ora at 6.30 a.m. before blooming of the fl ower, opening of the fl ower between 7 and 8 a.m. and wilting after noon. Ash (J.W. Ash 898) reports fl ower opening a.m. in C. schliebenii. Anthesis time in the other Coccinia species is not reported, but also likely to happen during the day. Th e scent is rather weak and dull sweetish, resembling that of honey melon, in C. abyssinica, C. grandifl ora, C. hirtella, C. rehmannii, 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. Th 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 fl 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. (Jeff rey 1967).
Bee pollination is confi rmed for C. rehmannii (C.J. Ward 12250), C. adoensis var. aurantiaca (Fig. 10b), C. grandifl ora, and C. grandis. Observed pollinators of C. grandis are Trigona apicalis Smith, 1857 and Trigona collina Smith, 1857 in Th ailand (Jongjitvimol and Wattanachaiyingcharoen 2006), and Megachile sp. in Cambodia (H. Schaefer, pers. comm.). Th e present author observed a halictid bee ( Fig. 10b; identifi cation 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 fl ower. Zimmermann (1922b) also observed the circling around the anther head in the largefl owered C. grandifl ora. He identifi ed the visiting small bee as Trigona sp. He also noted that a bee just having visited a Momordica fl ower walked on the inner side of the corolla loading dorsally located pollen on the anthers of a male C. grandifl ora fl ower.

Seed dispersal
Th 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 Th 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 Pacifi c Islands, dispersal of C. grandis by humans is well-known ). 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 Australia can also be explained by escape from gardens. Zimmermann reports feeding on C. grandifl ora fruits by birds, small mammals but also snails and beetles (Zimmermann 1922a), the latter two unlikely being seed dispersers. Th e forest weaver Ploceus bicolor Vieillot, 1819 was observed to feed on fruits of C. mackenii (Bleher et al. 2003). Stanford and Nkurunungi (2003) report diff ering 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.
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. jeff reyana, C. grandifl ora, C. grandis, C. hirtella, C. megarrhiza, C. microphylla, C. rehmannii, 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 fl ower) on the lower lamina of the leaves and/or on the bracts and probracts (Figs 7b,8a). Th 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. grandifl ora (Zimmermann 1922b) and in C. grandis (pers. observ.). Whereas Ilyas (1992) reports aggressive behavior of the ants on herbivores for Indian C. grandis, 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 fi nd aggressive behavior in C. grandifl ora 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 signifi cant reduction in residence time of herbivores on the cucurbit Luff a aegyptiaca when ants are patrolling on the plant. Most likely, there is no close relationship to certain ant species as guardians, and plantdefense is carried out only by a few ant species. How Coccinia species without or few probracts, bracts or sublaminal extrafl oral nectaries (e.g., C. microphylla) react when damage by herbivores occurs, is unknown. Agarwal and Rastogi (2008) found an increase of total numbers of extrafl oral 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 Pacifi c islands, in Australia, and the Neotropics, the plants can either overgrow other plants or represent a non-specifi c 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 diff erent organisms are reported to live in, on, or to feed from Coccinia species. Beetle and fl 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 fl y Bactrocera (= Dacus) cucurbitae (Coquillett, 1899), a tephritid fruit fl y (Uchida et al. 1990). Bactrocera cucurbitae larvae usually populate the fruits but are also reported to hatch from galls (Murthy 1959). Th e galls are not produced by these fl ies, however, but by the gall midge Lasioptera (= Bimba) toombii (Grover, 1962) (Bhatia and Mahto 1968). Th e gall infestation is interpreted as non-specifi c, as the female fl y might not be able to diff erentiate between the gall and an unripe fruit, which would be the usual target. In addition, also the tephritid fruit fl y Dacus ciliatus Loew, 1862 infests the galls, sometimes even together with Bactrocera cucurbitae (Bhatia and Mahto 1968). Th 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). Th e galls induced by Neolasioptera cephalandrae also appear to be gateway for a fungal infection with a mold, which is identifi ed tentatively as Cladosporium sp. (Krishnamurthy 1984).
As a result of the damage that can be done to cucurbitaceous crops and of its weedy behavior on Pacifi c islands, larvae of the clearwing moth (Sesiidae) Melittia oedipus 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). Th is moth, originating from Zanzibar (Oberthür 1878), appears to be quite specifi c 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.
Th e only known plant parasite growing on Coccinia is the hemiparasitic vine Cuscuta chinensis Lam., which is reported to grow on C. grandis in Gujarat, India (Patel and Patel 2010).  Reddy and Reddy (1988) Several fungi have been reported from Coccinia (Table 5). Th e rust fungus Puccinia windhoekensis Mennicken, Maier & Oberw. was described on Coccinia rehmannii (Mennicken et al. 2005), although Berndt (2007) noticed a great similarity of this rust to P. ctenolepidis Ramachar & Bagyanar. Berndt could not confi rm the identity of the host specimen, so it seems to be likely that it was misidentifi ed, since Ctenolepis cerasiformis looks quite similar to C. rehmannii.
Th ere are several reports of plant viruses from Coccinia species. Purcifull and colleagues (1988) tested the infectability of several Cucurbitaceae to diff erent plant viruses. Th 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 Pacifi c 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 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 diff erent 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. Th e plant is well-known in India, where its fruits had an impact even in classical Sanskrit literature. Th e red fruits are regularly used to describe lips, such as those of a beloved wife, who is described 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(Dutt -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). Th e high carotenoid value is of special importance in developing countries, as vitamin A deficiency is widespread among young children and pregnant women (WHO 2009). Social marketing has proven to be valuable in promoting the use of C. grandis to prevent vitamin A defi ciency (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 misidentifi cation or regional chemo-varieties with diff ering amounts of secondary metabolites.
Coccinia grandis has been used in Indian traditional medicine for several hundred years (Nadkarni and Nadkarni 1976;Ramachandran and Subramaniam 1983). Th ere are some studies that suggest a high potential for the use of C. grandis leaf extracts in diabetes treatment (Azad Khan et al. 1980;Kuriyan et al. 2008;Munasinghe et al. 2011). Parts of the observed eff ects are explained by inhibition of gluconeogenesis 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 eff ect on the promotor of the glucose transporter gene GLUT1 from rats is reported (Graidist and Purintrabipan 2009). Eshrat (2003) observes a positive eff ect of C. grandis in rats with hyperlipidemia, which is often connected to diabetes. However, its eff ectivity in 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 eff ects 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 (Umamaheswari and Chatterjee 2008), antipyretic, analgesic, and anti-infl ammatory (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). Antimicrobial 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 hyperprolactinemia-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 belief 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 benefi cial 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 ceremonies and celebrations and for animal fattening (Gelmesa 2010). Currently, much eff ort 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;. 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 expression diff er 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 diff erent Cucurbitaceae and found in Kenyan C. adoensis plants cucurbitacins H, I, and R in the stem but no cucurbitacins in the roots, leaves, fruits, or seeds. Coccinia trilobata was found to contain the cucurbitacins B, D, and G in the stems, cucurbitacin D, H, I, and R in the leaves, and cucurbitacin 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 cucurbitacin B (Bhakuni et al. 1962), and bitter and sweet fruited varieties are known. Guha and Sen (1973) fi nd that cucurbitacin B has an antigibberelic eff ect, and its occurrence in seeds of C. grandis might enable or increase dormancy of the seeds.

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 2 nd intron ( Fig. 18), suggest four major clades, although the backbone lacks bootstrap or posterior probability support (Holstein and Renner 2011b). Th e Coccinia rehmannii clade (IV) is well-supported in all phylogenies. Th 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). Th e plastid analysis tree separates these two clades but without support.
Th e C. rehmannii clade (IV) consists of fi ve species. Coccinia abyssinica and C. megarrhiza are sister species from Ethiopia and semi-arid parts of N Kenya and Somalia (Fig. 20). Th ey diff er ecologically with the former species occurring in the semihumid highlands and the latter one in the semi-arid lowlands. Both species diff er weakly in morphology, and hybridization cannot be ruled out. Th e plastid haplotypes of both species do not form clades in the tree, which might be explained best by incomplete lineage sorting. Th e other three species of clade IV contain several plastid haplotypes and nrDNA sequences that each also do not form clades. Th e geographical distribution of the haplotypes is not assessed. Th 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 fi 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 probability values ≥ 0.98 with values "with InDel coding" fi rst, followed by "without InDel coding." Numbers below the branches are bootstrap support values from ML analysis. Topologies from the diff erent 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.
(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 aff . var. littoralis). None of these forms cluster together. Th e other two species diff er 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 diff er morphologically from the C. rehmannii form from the dry inland. Th is scenario suggests incomplete lineage sorting and a speciation event with ecological diff erentiation in the northeastern Africa but not in southern Africa as intermediate collections between the four forms are found regularly. Th 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. Th is has been suggested several times for diff erent clades under the term "arid track" (Balinsky 1962;de Winter 1971). Th 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. Th 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 diff er morphologically but hardly produce hybrids (see chapter Hybridization and crossing experiments), a recent allopatric speciation event is probable. Th e lack of diff erentiation 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).
Th e C. adoensis clade (I) contains several morphologically and ecologically well diff erentiated species (Holstein and Renner 2011b). Th ere are three subclades in the plastid tree with accessions having the name Coccinia adoensis. Th 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 diff er 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. Th e Coccinia adoensis var. aurantiaca accessions are neither in the plastid nor in the nuclear tree monophyletic but share a dense indumentum. Th 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. jeff reyana (Fig. 17). Coccinia adoensis var. jeff reyana, however, shares the longer trichomes (Figs 3a, 5c) of some C. senensis, but it diff ers from these by lacking subulate calyx lobes and a 569 bp deletion in the trnS GCU -trnG UUC intergenic spacer region. However, one collection that does not diff er morphologically from the variety jeff reyana (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 fl ow. Additionally, gene fl 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. Th us it can be suggested that all these forms belong to one widespread species, C. adoensis, which contains diff erent plastid haplotypes. From this widespread species, several populations might have undergone ecological and morphological divergence. Coccinia grandifl ora and C. schliebenii are nested within one C. adoensis subclade, and they occupy rather humid 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 suffi ciently to be taxonomically well-separated as a species, such as C. adoensis var. aurantiaca.
Th 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 diff ers ecologically, as it occurs in mountain forest communities not in typical lowland rainforests as does the rest of the species. Th e phylogenetic position of C. intermedia is unclear as the resolution within this clade is generally poor. Coccinia intermedia shares morphological characters with C. adoensis, especially the open campanulate fl owers. Both species occur in the same habitat type with the former occurring in West Africa and the latter 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 possible that C. intermedia might have split allopatrically from a proto-C. adoensis species Figure 19. Scenario of evolution in the C. adoensis clade. Th 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. and is sister to the rest of the C. barteri clade (Fig. 19). Th 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 Pliocene 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 diff erentiation 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 genera (Table 6). Th e similarity is sometimes striking and without generative structures, one might need some experience to diff erentiate between the genera.

Characters for species discrimination
Th 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. intermedia, C. keayana), it is less useful in others (e.g., in the C. quinqueloba clade). Collections without fl owers are harder to identify. In some cases it is almost impossible to discriminate between species if fl owers are lacking. Identifi cation of only vegetative material is often possible but needs experience. Th e indumentum can be a useful character; 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 identifi cation.

Key to Coccinia species
Th e key is made from observations of herbarium material but also includes some characters from personal observations of living material and observations as given on herbarium labels. Fresh material is not needed, however, to use the key. Th e term 'articulate' refers to dried trichomes that appear wrinkled due to equatorially sunken cell walls (see Fig. 3a) but not to trichomes with ramifi cations, which have never been observed in Coccinia. In the living state, these trichomes are rather long and stiff . Th e term "dentate" refers to the sometimes colored structures (hydathodes?) at the leaf margin and leaf tip (Figs 6,7a,8a,16a,16b,21,39). L. aegyptiaca: mostly (2-)3-5-fi 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-fi 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 Habitats in this key (not the species descriptions) are given rather crudely and refl ect 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 thickets, 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. Flowers in lax many-(> 6-)fl owered racemes, western C Africa ...C. racemifl ora 9* Flowers in dense racemes, few-fl owered or on a long common peduncle that surpasses the length of the branched part; female fl owers may also be solitary . Plant rather densely covered with long (> 0.5 mm) trichomes that appear articulate 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 minute (< 0.2 mm), 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)  Plant with long (> 0.8 mm) trichomes that appear articulate when dry (Figs 3a, 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)  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) 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 conspicuous sterile apical tip ("beak"). Unripe fruits with whitish longitudinal mottling that often has a dark green halo. Seed face rather fl at, shape often asymmetrical (Fig. 14b,  Leaf reniform to lobate, rarely lobulate. Apex of leaf or central lobe retuse, obtuse, 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) (Fig. 8b). Plant from higher elevations of N Tanzania and Kenyan highlands (Fig. 33)  Lower leaf surface with long (> 0.5 mm) trichomes that appear articulate when dry or reduced to warts, rarely almost glabrous; sometimes leaves subsessile (Fig. 23)  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; Figs 17, 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 confi dently 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.
Use. Edibility of fruits is disputed and may diff er between wild and cultivated forms (E. Westphal & J.M.C. Westphal-Stevels 1951 and1953). Tuberous roots boiled for food (T. Ebba 250), young shoots and leaves are eaten when cooked (Hora 1995 Kuls 681). Th e Kefi nya name is not exclusive for C. abyssinica but also used for another crop, Plectranthus edulis (Vatke) Agnew.

Remarks.
Th e occurrence of monoecy has been reported by W.J.J. O. de Wilde et al. 7805, but the seen specimens contained male fl 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. Th 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. Th 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 Kotschy 308 specimens merely state the species name, the locality, and "frutices scandens" (= climbing on shrubs; W. Greuter -pers. comm.), which cannot be regarded as a diagnostic feature. Th e label is printed and therefore eff ectively published but not validly so. Valid publication of that name was eff ected by Achille Richard (1847), but he chose a diff erent specimen (G.H.W. Schimper 1418), which belongs to C. abyssinica. Th 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)". Th e phrase, however, is also not a validation since the species to which this variety is supposed 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.
Th e identity of Cucumis striatus A.Rich. is not obvious. Th ere are two original specimens 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. Th e P05621224 specimen consists of a ripe fruit, a drawing of the fruit, and a tiny fragment of a leaf. Cogniaux identifi ed 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. Th 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. Th e other original specimen (with a number "26" from "Sholada") contains much leaf material and fruits. Th e fruits are darker than in the fi rst type specimen. Th 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. Th 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.  (Fig. 14a, 21).  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. Remarks. Coccinia adoensis is widespread and morphologically variable. Some populations or local forms appear to be distinct, but there are intermediate individuals or similarlooking collections from diff erent parts of the overall distribution range. In East Africa (C Tanzania, Malawi), one can fi nd forms linking to C. aurantiaca, which is treated here as variety of C. adoensis, and to C. senensis. Th e latter forms have a similar plastid haplotype with C. senensis, but lack a specifi c deletion in the trnS GCU -trnG UCC intergenic spacer. Th ese forms from central and southern Tanzania are discussed here under the name C. adoensis var. jeff reyana, while another form from Kenya remains in var. adoensis. Th e non-monophyly in the plastid tree (Holstein and Renner 2011b) makes C. adoensis even more peculiar. Th 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 Coccinia adoensis var. adoensis well (sympetalous, obovate probracts). Th ey strongly resemble Eureiandra species in vegetative characters, but not in generative traits.
Taxonomic remarks. Th e lectotypifi cation of Momordica adoensis by Meeuse (1962) is not eff ected, 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.
Th e types of Bryonia jatrophaefolia are not too obvious as such. Th 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". Although the spelling has some similarities, they are quite diff erent. 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 "Djeladjeranné". 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.
Th 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-ditionally, the specimen is a fruiting female, like the specimen in K. Hence, d'Alleizette must have obtained a duplicate of the type.
Th e placement of C. hartmanniana as a synonym of C. adoensis is done with a high level of confi dence although no type specimens were found. Th e protologue contains drawings showing lenticular seeds and short calyx lobes, which match well other collections 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 duplicates 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. Th e leaves of the chosen specimen, H.J.E. Schlieben 3271, match the description well, and the specimens have been identifi ed as C. princeae when the original material was still existing. Th e specimens diff er in the generative characters (fruiting in the holotype, male fl owers in the neotype), but Gilg referred strongly to the distinctive leaves, so the neotype appears to be a good match.

Remarks.
Th 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. adoensis. Jeff rey segregated this species from the polymorphic C. adoensis because of the nonbeaked fruits and fl at seeds with a hyaline girdle. Th 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. Th e seeds are also hardly distinct from C. adoensis, perhaps somewhat larger. Seeds in Coccinia are enclosed in a hyaline aril. Jeff rey only observed the dry collapsed aril, which is not part of the seed, as a "hyaline girdle". Th 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. Th 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. Th e fl owers in the R. Polhill & S. Paulo 1274 specimen in P do not belong to Coccinia. Th e calyx appears to be Momordica foetida Schum. & Th onn. Th e HEID specimen (HEID779579) of that collection is also mistaken, eventually a mix-up while mounting the specimen. It has a completely diff erent indumentum and a narrow, almost cylindrical perianth tube.

Diagnosis.
Th is variety has affi nities with C. adoensis and C. senensis. Th 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. Th e calyx lobe length often exceeds 2 mm (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 pustulate. 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 fl owers in racemes, often accompanied by a single fl ower or one solitary fl ower. Common peduncle 5-5.5 cm, with short articulate trichomes. Pedicel of racemous fl owers 5-9 mm, with short articulate trichomes. Bracts up to 1 mm, caducous. Pedicel of solitary fl 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 fi lament column pink, anther head orange-yellow to orange, color of pollen sacs not seen. Female fl 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) 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. Th e sequenced specimens do not cluster with most other C. adoensis haplotypes from East Africa or southern Africa, and lack the typical deletion of C. senensis in the trnS GCU -trnG UCC 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. grandifl ora or C. mackenii in higher altitudes or in areas with higher precipitation. Th 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 diff er from the "typical" C. adoensis, but still belong to C. adoensis, they are treated as a new variety.
Th 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. Th is collection has a normal-sized corolla, and therefore seems to be fertile, which supports the hypothesis that the var. jeff reyana is not reproductively isolated from var. adoensis. Th is is also why the present author refrains from designating it as a paratype, namely in order to avoid confusion about the genetic defi nition of this variety.
Phylogenetically, it is uncertain whether this variety retains an ancestral morphology 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 aridifi cation 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. jeff reyana, 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.
Th e collections from Singida occur in drier habitats than those from C and S Tanzania 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 5 mm long or missing. Tendrils simple or bifi d. Male fl owers in few-to many-fl 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, refl exed, spreading or erect and adpressed to corolla, sometimes seemingly fl 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 fl owers in racemes, sometimes accompanied with a solitary fl ower or one solitary fl ower only. Peduncles and petioles in racemes like in males. Solitary female fl 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 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 fl at to fl atly lenticular.
Phenology. Flowering time: January-June, August-November. Distribution. Fig. 24. Remarks. Coccinia barteri is treated here in a wide sense as it contains several forms (see also Holstein and Renner 2011b). Th is is because data on these forms are scarce and do not unambiguously allow to refer to these as species. Th erefore, the present author refrains from creating an intraspecifi c classifi cation as a phylogeographic treatment and crossing experiments appear to be necessary to clarify this problematic taxon.
Th ere are collections in Gabon that are of intermediate morphology between C. barteri and C. racemifl ora (M.A. van Bergen 490 (WAG) = C. barteri 6 in Fig. 17). Holstein and Renner (2011b) suggested that hybridization occurs between these species. 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). Th ese were collected on that island, and they contain 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).
Th e syntypes of Coccinia macrocarpa certainly belong to diff erent taxa. Th 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 7 mm long, at the base attenuate to trun-cate and with a rounded apex. In contrast, the seeds of É. Luja 125 are subquadratic as Jeff rey already pointed out on the type specimen. A placement in Momordica by Jeff rey (on the sheet) seems to be correct, whether this is M. multifl ora Hook.f. (1871) as identifi ed by Jeff rey or M. parvifolia Cogn. (1916) as identifi ed 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 fl owers in long racemes as a new species, Coccinia subhastata Keraudren. She also described several diff erences 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 bifi d tendril. Furthermore, the C. barteri lectotype C. Barter 1525 has a subhastate leaf and simple tendrils. Th 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-fl owered racemes as C. keayana R.Fern. (Kéraudren  Description. Perennial climber. Stems up to 20 m, glabrous or (when from higher altitudes) sparsely covered with long, whitish trichomes. Petioles 2.5-13 cm, indumentum as on stem.  Lobes triangulate, ovate to oblong. Leaf margin smooth to slightly serrate, dentate. Apex obtuse to acute with fi 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 5 mm long (Fig. 8a) Remarks. Th 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. Th ese populations may represent hybrids or descendants of a non-diff erentiated common ancestor.

Coccinia grandifl ora
It is diffi cult to distinguish between C. grandifl ora and C. mildbraedii in the Central Tanzanian highlands (Eastern Arc Mts). Both species also occur in high altitude forests and are clearly delimited by fl ower size. Coccinia grandifl ora also has larger probracts than C. mildbraedii, but this is rarely well visible. Coccinia grandifl ora may also be confused vegetatively with C. barteri in Mozambique and Zimbabwe. Taxonomic remarks. Th e C. grandifl ora holotype by Holst was destroyed in the fi re of the Berlin herbarium in 1944. Th e Winkler specimen was chosen as neotype because it was already designated as type in December 2008. Th ere is no annotation on the type label, however, and it seems that this neotypifi cation was not published. However, the Winkler specimen label bears Cogniaux' handwriting. Strangely, the Winkler specimen also states "mars 1892", with the 92 crossed out. Th 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. Th e drawing is of suffi cient quality to synonymize unambiguously C. engleri with C. grandifl ora.   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 usually 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 fi 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.
Distribution  (Bharathi 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 higher than in Africa, the quick loss of surface water allows C. grandis to survive there.
Use. Fruits (raw and cooked) and shoots (cooked) edible. Th e Luo eat the leaves as spinach (Orech et al. 2005). Th 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 eff ect (see also Use, economic potential, and phytochemistry).  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 fi 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 bifi d tendrils, and both South African species lack colored leaf teeth and have erect to spreading calyx lobes instead of spreading to refl exed 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 diff er genetically (e.g., in a short sequence in the 5'-beginning of the LEAFY-like 2 nd intron) and in petal color (white in Asian, buff 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 Naudin (1862) indicate full compatibility between the African and Asian morphs though.
Taxonomic remarks. Up to the 21 st 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 collected several times during his 13-year stay in South India (Stafl eu and Cowan 1988). Th 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). Th e illegitimacy of Coccinia indica, however, does not aff ect 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 published 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-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 (1747), and only in the 2 nd and 3 rd 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. cordifolia on Rumphius' fi gure, and the epithet cordifolia is hence misapplied in Coccinia.
Th ere are four specimens in Linnaeus' herbarium, which belong to Coccinia grandis but it is unclear which are type material. Th e number 1153.2 is designated (by Linnaeus?) as Bryonia grandis on the sheet and is therefore the best choice for lectotypifi cation as is has been done by Nazimuddin and Naqvi (1984). Th e specimen 1153.13 bears the note "Bryonia foliis subrotundis angulosis, momordicae facie" on the specimen fl ip side referring to one of the citations to drawings in the protologue. Hence, this specimen is also original material. Th 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.
Th e drawing to Bryonia foliis subrotundis, angulosis, momordicae facie Burm. (Th es. zeylan.: 49, t. 19, fi g. 1. 1737), which is original material of Bryonia grandis L. appears to be a product of artistic freedom. Th e calyx lobes of the three uppermost and the lowermost fl owers are refl exed and match Coccinia/Bryonia grandis well, whereas the calyx of the other two fl 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). Th e addition "momordica facies" seems to relate to this. Strangely, the drawing of Bryonia folio anguloso acuto glabro Burm. (Th es. zeylan.: 48, t. 19, fi g. 1. 1737) matches the current defi nition 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.
Th e Forsskål 660 specimen (C10002122) has a hand-written fi eld label by Forsskål (according to notes in JStor Plant Science) on the fl ip side, stating "Cucumis incerta. Arakis, Mour." (incerta meaning "uncertain") and two hand-written identifi cations "Cucumis inedulis Fl. Yemen CXXII nr. 580" and "c. s. Arakis cent VI nr. 61 p. 169". Th e location in the text is the same as on the fi eld label: Môr/Mour [Mawr, a small town about . Th e former identifi cation is a nom. nud. with the number 580 on page CXXII of Forsskål's Flora AEgyptiaco-Arabica (1775). Th e second identifi cation links to a Cucumis sativus variety that is validly described in that book on page 169. Th e description matches well Coccinia grandis except for the tuberculate 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. Th e description of the variety also mentions that the plant is not edible, just as the supposed species epithet.
Th e genus name Turia has been created by Forsskål in his Flora AEgyptiaco-Arabica (1775). Th ere is a debate, however, whether it is validly published (Friis 1984). Th 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 fl oras (Roman page numbers), which is on page CXXI in the case of Turia. Th e fi rst species there is Turia sativa (no. 550), which is called "turia" in Arabic and is cultivated according to the epithet. Th is matches exactly the fi rst description in the descriptive part of the book (p. 165). Forsskål lists fi ve Turia species in the fl oristic part and describes fi ve species in the descriptive part. Th erefore, the genus Turia lacks a description (Jeff rey 1962), because Forsskål does not mention any character to be typical for the genus. Th e name Turia was legitimately described by Gmelin in an extension of Linnaeus' Systema naturae (1791), and so was the name Turia moghadd.
Th e typifi cation of Turia moghadd is not straight-forward, because Forsskål added little, if any marks on the sheets (Friis 1983). Th e fl ip side of the Forsskål 663 specimen has three notes: "Cucumis glandulosus", a second one with a diff erent 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 diff erent 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. Th 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). Th 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.
Th e name Momordica monadelpha Roxb. is superfl uous, because Roxburgh synonymized Bryonia moimoi Ser. in total by citing the only element of that name and Bryonia grandis. Th e other elements of M. monadelpha are also interesting, though. Roxburgh cited "Bryonia foliis subrotundis" with the citation of Burman's Th es. zeylan.: t. 19, fi g. 1. 1737 and fi g. 2, Vitis alba indica, all Coccinia grandis, and Herman's Musaeum Zeylanicum 356. Th 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 fl ower morphology (calyx lobe length and position, corolla size), rather than Cucumis maderaspatanus L., Diplocyclos palmatus L. or Cayaponia laciniosa (L.) C.Jeff rey. Th at the drawing 5: 225 (BM000595000) shows a plant with male and female fl owers on one individual might be explained best by artistic freedom.
Th e name Coccinia indica var. palmata C.B.Clarke is valid and legitimate but not obvious to typifi y. Despite C. indica being illegitimate as a nomen superfl uum, 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. Th 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 typifi cation (Art. 52.2 N1). Th e element "Bryonia palmata Linn.? herb. Madr.!" relates to a collection in Herbarium Madras that was identifi ed 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. Th 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 epithet of the variety from this misidentifi cation and not from Bryonia palmata L., which is why the name was typifi ed with this collection. Th 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.
Remarks. Th e long, subulate calyx lobes are the only good character for distinguishing 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 affi nity (introgression?) to C. racemifl ora, which also has elongated racemes, defi nes an own species, or is just a coincidental observation of intraspecifi c variation is not known.
Taxonomic remark. Th is species is the type species of Physedra Hook.f. Th 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 Jeff rey (1962) transferred P. longipes into a new genus, Ruthalicia, he indirectly lectotypifi ed Physedra.
Taxonomic remarks. Th e BR type specimen (BR0000008887184) contains two labels and is mixed. Th 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. Th e male parts on the BR specimen, however, are mixed. Th 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. Description. Perennial climber. Stem length unknown, but likely several meters, glabrous, 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 bifi d. Male fl owers in few-fl owered racemes, likely sometimes accompanied by a single fl ower. Common peduncle up to 1 cm, pedicels in racemose fl 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. Filament column and anther head not seen, pollen sacs yellowish. Female fl owers 1-3 clustered (strongly reduced raceme). Pedicels 0.6-1.2 cm, glabrous. Hypanthium tube glabrous, calyx lobes and corolla like in male fl 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 fl 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. Th is species is rather cryptic and imperfectly known. Th e leaves seem to develop conspicuous margin teeth during maturity, like e.g., C. grandis, but the darkish sublaminal glands diff er from that species. Th e erect calyx lobes with slightly recurved tips appear to be the most indicative character for C. intermedia. Th e clustered female fl owers and the fruits link to C. barteri, from which it, among other characters, diff ers in ecology. Two J.B. Hall & J.M. Lock GC 46016 specimens from K have male and female fl 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).  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 fi 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 fl owers ebracteate, in lax racemes with up to 20 fl owers, sometimes accompanied by a solitary fl ower (Fig. 32). Common peduncle up to 1.7 cm, shorter than racemous part, glabrous. Pedicels of racemous fl owers up to 1 cm, pedicels of solitary fl owers up to 1.5 cm long, each glabrous. Perianth tube glabrous. Calyx lobes linear, 2.5-3 mm, in buds spreading, later refl exed. 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 fl owers solitary or in few-fl owered lax racemes. Common peduncle 1.2-2.1 cm, glabrous. Pedicel fl owers in racemes up to 1 cm, glabrous, pedicels of solitary fl owers up to 2.7 cm, glabrous. Hypanthium glabrous, calyx lobes and corolla as in male fl owers. Ovary glabrous. Style and stigma not seen. Fruits 2-3 × 2 cm, subglobose to globose, un-   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).

Remarks.
Th e broad calyx lobes are, apart from the long cylindrical fruit, the best character for identifying this species. An urceolate corolla ) 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 fl 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.  Description. Perennial climber or creeper. Stems up to 9.5 m, glabrous. Petioles 0.7-11 cm long, glabrous or with thin trichomes.  cm, shallowly to profoundly 5-lobate, in the latter case often weakly lobulate. Lobes triangulate, lanceolate, ovate to obovate. Margin smooth, dentate, sometimes serrate to lobulate, esp. towards the apex. Apex acute with fi nal tooth. Upper leaf surface glabrous with clear to white pustules, rarely with few trichomes. Lower leaf surface glabrous or with thin, stiff or articulate trichomes, towards base usually with glands. Probracts up to 4 mm, oblong-lanceolate. Tendrils bifi d, rarely simple. Male fl owers solitary or ebracteate in few-fl owered racemes. Common peduncle 5-6.5 cm, pedicel of fl ower in raceme up to 2.5 cm, pedicel of solitary fl 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 refl exed. Corolla 1.3-2.7 cm long, cream to pale buff , corolla lobes subulate to triangulate, 0.7-1.1 cm. Filament column, anther head, and pollen sacs not seen. Female fl owers one solitary. Pedicel 0.7-5 cm long, glabrous. Hypanthium glabrous, calyx and corolla like in male fl 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 fl atly lenticular.
Remarks. Some collections with deeply lobate leaves and short petioles resemble the closely related C. quinqueloba, and some C. quinqueloba individuals have long petioles (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 interspecifi c crosses. It would be desirable to validate this observation.
Taxonomic remarks. Th e initial designation of the C. mackenii lectotype (Holstein 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 ambiguous. Th 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 aff ected 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 indica they also included a specimen tentatively identifi ed 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 diff erent 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.
Th e drawing of Coccinia dinteri in the protologue shows a bifi d 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 indicated. Th e label named it "Coccinia dinteri" after Moritz Kurt Dinter (in the protologue 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. Th 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 mentions that the seeds were sown in Kew Gardens. Th e identifi cation is given as Cephalan-dra mackenii 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 fi t the description of C. mackenii seeds, it is not possible to use them to identify the species unambiguously.  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).  cm, reniform to 3-or 5-lobate. Margin dentate (teeth at maturity brownish, when dried blackening), 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 fl 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 orangeyellow. Female fl owers 1(-2) solitary. Hypanthium with long, beigeish, upright trichomes that appear articulate when dried, calyx lobes and corolla like in male fl 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) Remarks. Coccinia megarrhiza and C. abyssinica form a species complex. Distinction between these two species can be diffi cult, especially in young plants, when the color of the marginal teeth of the leaf is not well developed. While the peduncle length diff ers, the earlier appearing solitary fl owers can have the same length in both species. Th 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 diff erentiated 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.   (Fig. 2a). Pedicels up to 0.7 cm, glabrous or with white trichomes. Ovary glabrous, with some articulate trichomes to densely wooly 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  Remarks. Th 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.

Coccinia mildbraedii
Ellis notes on the collections no. 163 and 383 a smell of rotten meat. However, it is unclear, whether this is coming from the fl 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. Simmons 64 (EA). Remarks. Morphologically this species (the only one missing DNA sequences) is close to C. senensis. Th 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. Th e racemes in C. pwaniensis have considerably more fl 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 cooccur, they might be sister species from allopatric speciation, with C. pwaniensis occurring in a refugial distribution in the northern coastal forests of East Africa. Distribution. Fig. 30. Southern and western Eastern Cape, South Africa. Elevation sea level to 1000 m. Sandy soils, also on dolomite soil. Coastal bushland, forest, dry bush, on bushes along rivers, along roadsides.

Coccinia pwaniensis
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.  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 precipitation 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 interesting to link morphological characters with haplotypes and to test the fi tness of these morphs in the diff erent habitats. In any case, the morphological diff erentiation seems to be stable in cultivated individuals, and artifi cial crosses between diff erent forms (inland vs. subglabrous from the Southeast) result in the onset of a normal fruit (resulting seeds were not used for cultivation). yellow to orange. Female fl 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 fl owers. Ovary with smutty-brownish trichomes. Style 3-6 mm, 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  Distribution. Fig. 23. Central Tanzania (Iringa, Lindi, Morogoro, Ruvuma), Malawi (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 Remarks. Th e species is recognizable by the combination of few-fl owered racemes, long subulate calyx lobes, and the often subsessile leaves. Th 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 trichome 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 (Jeff rey 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. jeff reyana in this treatment. Th 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. jeff reyana). Whether C. adoensis and C. senensis are truly separate species and the role of these intermediates needs to be tested by artifi cial hybridization, fi eld observations, and/or a phylogeographic analysis.

Coccinia racemifl ora
Taxonomic remarks. Although the holotype of C. senensis burned during the destruction 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 Jeff rey's C. fernandesiana. Th e C. senensis protologue points out "articulate" trichomes and an overall appearance like C. quinqueloba, which matches perfectly with many collections of C. fernandesiana. Interestingly, many of these collections have been identifi ed as "Coccinia quinqueloba" or "Coccinia palmata" by various collectors and scientists. Th e similarity, including the calyx lobes, is easily visible in many collections, but both species are restricted to southern Africa.
Cogniaux described var. australis of C. jatrophiifolia (synonym to C. adoensis) recognizing the similarity to the polymorphic C. adoensis. However, he diff erentiated 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 Jeff rey described 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. Th e one COI specimen is therefore paratype of C. subglabra and syntype of C. jatrophiifolia var. australis. Th e two COI specimens are also misplaced paratypes of Meeuse's C. rehmannii var. littoralis. Th e similarity of the COI specimens of Meeuse's variety to C. senensis Description. Perennial climber or creeper. Stems up to 5 m long, with slight waxy cover, glaucous (Figs 2b, 7b), glabrous (fi rst shoots may have short, white trichomes). Leaves sessile to amplexicaul (fi 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 bifi d. Male fl owers solitary or clustered in few-(rarely many-)fl owered racemes (Figs 2b,  7b). Pedicels of solitary fl owers 1-4 cm, glabrous. Peduncle 1-6 cm long, glabrous. Pedicels of fl owers in racemes 0.3-2.5 cm, glabrous. Bracts glabrous, up to 1.8 mm,   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 diff erentiation). 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 fi xation of this character. As C. sessilifolia var. sessilifolia is derived from petiolate plants, this variety might even represent a remnant population of these. Taxonomic remarks. Th ere are two Malchair 433 specimens. As they do not contain any indication of having been separated from a single specimen, they are treated as syntypes. Th e two specimens do not diff er 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. Th ese features are unusual, but the other characters match the species.  Th 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. Schweinfurth 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. As the holotype is destroyed and the description does not give enough suffi cient characters 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. 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). Th e drawing, if it represents a Coccinia (the two subsessile female fl 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.

Remarks.
Th 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. samburuensis, which diff ers in coriaceous leaves and a serrate margin with glandular teeth. Hence, this species name is not synonymous with any Coccinia species. Th e tendrils in C. longipetiolata are almost equally bifi d, which is not found in Coccinia, especially not in species not from rainforests. Th 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. Th ere are neither characters supporting a relationship with Coccinia, nor characters contradicting it, except for the tendrils. Jeff rey (1967) suggests a relationship to his Coccinia sp. E sensu F.T.E.A. (Jarman 66), but as this specimen could not be examined by the present author, it cannot be discussed. Th erefore, that species is treated as dubious. Eventually, sequencing could give disclosure about the relationships.

Coccinia glandis nom. nud.
Th is is a typographical mistake for C. grandis that has been published several times (Tewtrakul et al. 2006, Jiwajinda et al. 2002. Th is epithet should hence not be used in Coccinia.

Cucurbita laevigata Bl.?, nom. nud.?
Th is name is written on a specimen in L herbarium (L0587542). Th 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. Another label, written with a pencil, says "Cucurbita laevigata" "mihi" and "Callelet W[…]". Th 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.
Bryonopsis pedata Hassk., Cat. hort. bot. bogor.: 189. 1844, nom. nud. Hasskarl cites Noroña's Bryonia quinquefolia and a vernacular name "aroy kalanyar beurriet". Th e given description of lacinate, almost pinnatifi d leaves and male fl 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 Luff a cordifolia Bl. None of these names have been referred to Coccinia, so it seems unlikely that Bryonopsis pedata does.
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.
Th e name was used on printed labels of G.H.W. Schimper 1570 (eff ective publication), which lack a proper description (hence a nom. nud.). Under this distribution number, specimens from two diff erent shipments are included. One is taken from package "P. 16 K. no. 4", collected on 23 Apr 1841 in Djeladjeranné (label on P specimen). Th e data of this label were used for C. F. F. Hochstetter's printed labels. Th 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 notes this collecting site, hence the specimen might be from the same shipment. Specimens 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 fi 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 published it. In any case, this specimen belongs to C. grandis.
Bryonia sinuosa Wall., Numer. List 6716. 1832, nom. nud. Th 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 Coccinia grandis or Cucumis maderaspatanus. Wallich himself supposed that this collection is a mix of (Bryonia) Coccinia grandis and Melothria indica. Cogniaux cites a Schimper specimen (Iter Abyss. Sect. 3 no. 1202) that was supposed to be labeled by C. F. F. Hochstetter. Th ere are several sheets with this distribution number in Paris, but only one bears this name. Th e location is given by "In Semen" [Semien Mts]. Th e other Paris specimens with this number are from Baria Dikeno (collected on 6 Aug 1853). Th e collection is a Coccinia grandis. Remarks. Th e type does not contain much material, but the lower surface of a leaf shows a pinnatifi d venation pattern, which is unknown in Coccinia. Jeff rey (1967) transferred it correctly (Kocyan et al. 2007) to another genus, namely Cephalopentandra.

Remarks.
Th e author notes fi ve stamens, which are a good reason for not including this species in Coccinia. Th e leaves of the specimen on the picture look like these of Coccinia quercifolia, which is also excluded from Coccinia and separated by Jeff rey (1967).

Remarks.
Th e name Coccinia palmata has been applied illegitimately for C. mackenii 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 tentatively included the citation of Bryonia palmata L. More likely, however, they meant a specimen in Herbarium Madras that was identifi ed as B. palmata L. One year after Voigt's correction to Coccinia grandis (L.) Voigt, Roemer (1846)  Remarks. Jeff rey (1967) synonymized this species with Eureiandra fasciculata (Cogn.) C.Jeff rey. Remarks. Th e seeds are described as globose to subglobose, but Coccinia seeds are rather fl at. It is therefore unlikely that this species belongs to Coccinia. Jeff rey (1967) synonymized this species with Eureiandra fasciculata (Cogn.) C.Jeff rey.

Remarks.
Monoecious plant with several small subglobose fruits per node apply clearly to Diplocyclos and must therefore be synonymized as it has been done by Jeff rey (1967) to Diplocyclos decipiens.

Remarks.
Th 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 trifi d and the fruit is spherical and apple-sized, which does not fi 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 confi rmed by Kocyan et al. (2007).
most of the morphological fi ndings and the crossing experiments would not have been possible. I also thank Frank M. Mbago for his help and excellent eye on the fi eld trip in Tanzania. Th anks are given to Hanno Schäfer for communicating various observations and specimens from DNA, NY, US, Christian Bräuchler for communicating and discussion 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 exanthematica, and Brigitta and Willem de Wilde for discussion of the Momordica bicolor specimens and sharing their fi eld experience with C. grandis. I thank Eva Facher for help with staining of the petiole sections and Mila Vosyka for preparing the chromosomes. Th 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 supervision and English revision. I also thank Sandy Knapp and two anonymous reviewers for many helpful comments on the manuscript.
Th is work has been fi 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. Th is research received support (visit to L and U collections) from the SYNTHESYS Project (http://www.synthesys.info/), which is fi nanced by European 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.