Description[ edit ] Coccinia species are perennial climbing or creeping herbs. Climbing is supported by simple of unequally bifid tendrils. Most species develop a tuber from the hypocotyl , sometimes on roots. The cotyledons are simple, entire and have an blunt tip. The leaves are usually stalked , rarely sessile.
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Supplementary material 1: List of examined specimens and geo-references of the localities Data type: specimens data. Explanation note: Voucher information and predominantly inferred coordinates of geo-references in decimal degrees. Specimens included in this study, with the geographic origin of material only country and 1st administrative division given.
Herbarium acronyms follow Index Herbariorum; barcodes or other unique identifiers in brackets behind the acronym. The Open Database License ODbL is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author s are credited.
Supplementary material 2: GenBank accessions Data type: specimens data. Explanation note: Voucher information and GenBank accession numbers. Specimens included in this study, with the geographic origin of material only country and 1st administrative division given and GenBank accession numbers for all sequences. Herbarium acronyms in parentheses behind the voucher name follow Index. Abstract This monograph deals with all 95 names described in the Cucurbitaceae genus Coccinia and recognizes 25 species.
Taxonomic novelties are Coccinia adoensis var. Jeffrey Holstein, stat. Meeuse Holstein, stat. For the 25 species collections were examined, of which were georeferenced to produce distribution maps. 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. 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. 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. All species are dioecious, and one species, Coccinia grandis L.
Voigt, has heteromorphic sex chromosomes and therefore has been studied cytologically Agarwal and Roy ; ; ; Bhaduri and Bose ; Chakravorti ; Datta ; Kumar and Deodikar ; Kumar and Vishveshwaraiah ; Roy ; Roy and Roy b.
The last complete taxonomic treatment of Coccinia is by Cogniaux , more than years ago. Since then, 16 new species have been described, and the genus has only been revised regionally Hutchinson et al.
The position of Coccinia in the Benincaseae has been confirmed by molecular data Kocyan et al. The delimitation of Coccinia from other genera is difficult. The scarlet-red fruits to which the genus name — Coccinia from Latin coccineus — refers are also found in other African genera, such as Eureiandra Hook. Therefore, it is not surprising that early botanists described several species now considered to belong to Coccinia in other genera Cephalandra Schrad.
In all, names at various ranks have been proposed for what are here considered 25 species. The species concepts in the present revision are based on herbarium collections and fieldwork in Tanzania, geo-referencing of 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 effort provide a modern understanding of the evolution and species relationships in Coccinia. New collections were added if the photograph allowed identification or if misidentification appeared to be unlikely esp.
Coccinia grandis collections from the Pacific 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. Jeffrey 1 origin, 8 individuals , Coccinia microphylla Gilg 2 origins, 2 and 1 individual, respectively , Coccinia rehmannii Cogn. Jeffrey 2 origins, each 3 individuals. The present author performed crossing experiments among eight of these species.
Morphological features were documented photographically and in the form of vouchers, with 47 collections deposited in M. Field data were obtained on a trip to NE Tanzania in resulting in 28 Coccinia collections.
Phylogenies For this monograph the phylogenetic data of Holstein and Renner a ; b were augmented with 20 new sequences from 8 accessions GenBank accession numbers are given in Suppl. For Bayesian analysis, four chains were run with 2,, generations, with a sampling frequency of Distribution maps Of the examined collections, were geo-referenced and mapped in Google Earth Google Inc.
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.
Political administrative borders were taken from GADM v. The geodetic datum for the maps is WGS84; the projection in each case is equirectangular. Coordinates are given in decimal degrees in Suppl. Morphology and anatomy Habit Coccinia species are perennial climbers or creepers.
The lignification of the mature shoots differs among the species from unlignified to completely lignified. Climbing is enabled by tendrils, which are either simple or bifid.
Tendril development in young plants is delayed and emerges in Coccinia abyssinica after the 6th node Getahun a. The 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 bifid tendrils is often not fixed, but there is a strong predominance of one kind.
Bifid tendrils regularly occur or are predominant in Coccinia grandiflora Cogn. Holstein, Coccinia hirtella Cogn. Strikingly, Coccinia species with bifid tendrils occur in rather humid habitats. This 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 floristic treatments, but they may bare bifid tendrils such as Coccinia sessilifolia N. Holstein 13 and Coccinia senensis H.
Schlieben in B, K, and MO. Coccinia adoensis has bifid tendrils even in some type specimens e. Schimper in BR 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 bifid tendrils: Coccinia sessilifolia with Coccinia hirtella and Coccinia mackenii, and Coccinia adoensis with Coccinia grandiflora and Coccinia 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. Some species, such as Coccinia adoensis and Coccinia grandiflora and most likely also Coccinia senensis Klotzsch Cogn. Root tubers in Coccinia adoensis are likely to be an adaptation to fire, as this species predominantly occurs in woodlands.
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