Phylogeny of the genus raphionacme (Apocynaceae)

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Date
2013-07
Authors
Pienaar, Magdil
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Publisher
University of the Free State
Abstract
English: The phylogeny and biogeography of Raphionacme were investigated in this first comprehensive study which included virtually all the species. Raphionacme is the largest genus in the subfamily Periplocoideae (Apocynaceae sensu lato) and consists of 36 species and two subspecies. Sequence data from the nuclear ITS gene region and 45 morphological characters were used to determine relationships between 30 Raphionacme species. Raphionacme is an African genus and widely distributed throughout this continent with the highest concentration of species in southern Africa, the Cape Floristic Region excluded. Only one species, R. arabica, is found outside Africa on the Arabian Peninsula. The most common habitat in which members of this genus occur is grassland or savanna. The majority of species are herbaceous geophytes, with four species that have a climbing habit. Four areas of species richness were identified. These are the Tongaland-Pondoland Mosaic in northeastern South Africa, which includes the Maputaland Centre of Endemism, the Zambezian Region, the Karoo-Namib Region and Kalahari-Highveld Transitional Zone both in the west of southern Africa. Nine endemic species were identified of which seven occur south of the Equator. These are R. chimanimaniana, R. elsana, R. haeneliae, R. namibiana, R. sylvicola, R. villicorona and R. zeyheri, with only R. moyalica occurring in the north of Kenya and R. arabica in Oman. The cladistic results of this investigation indicated that the first and therefore oldest clade to diverge contain species from southern Africa, leading to the assumption that Raphionacme originated in this region. The geophytic habit of the species would probably be an indication that this coincided with the aridification of this region. Thereafter diversification and radiation may have occurred northwards following the savanna that developed east and north of the remaining Equatorial rain forest. Some taxonomic implications, based on the phylogenetic results, have been included. The outgroup Schlechterella abyssinica was found to be nested in the Raphionacme clade, in both the molecular and combined cladograms. The removal of Raphionacme abyssinica to the genus Schlechterella would seem to have been incorrect and the species S. abyssinica may have to be returned to Raphionacme. Three Raphionacme species, R. bingeri, R. excisa and R. splendens were combined under the name R. splendens. Subsequently R. splendens was subdivided into two subspecies. The cladistical results of this study support the creation of Raphionacme splendens subsp. splendens and R. splendens subsp. bingeri.
Afrikaans: Die filogenie en biogeografie van Raphionacme was ondersoek in hierdie eerste omvattende studie wat naastenby al die spesies ingesluit het. Raphionacme is die grootste genus in die subfamilie Periplocoideae (Apocynaceae sensu lato) en bestaan uit 36 spesies en twee subspesies. Data van basispaaropeenvolgingsbepaling van die kern ITS geenstreek en 45 morfologiese kenmerke is gebruik om die verwantskappe tussen 30 Raphionacme spesies vas te stel. Raphionacme is ‘n Afrika genus en kom wydversprei oor hierdie kontinent voor met die hoogste spesiekonsentrasie in suidelike Afrika, maar is afwesig in die Kaap Floristiese Streek. Slegs een spesie, R. arabica, word buite Afrika op die Arabiese Skiereiland aangetref. Die meeste lede van die genus word in savanna of grasveld aangetref. Die meerderheid spesies is kruidagtige geofiete, maar vier spesies is klimplantagtig. Vier gebiede met ‘n hoë spesiekonsentrasie is geidentifiseer, naamlik die Tongaland-Pondoland Mosaiek in die noordooste van Suid-Afrika, wat die Maputaland Sentrum van Endemisme insluit, die Zambeziestreek, die Karoo- Namibstreek en Kalahari-Hoëveldoorgangsone, beide in die weste van suidelike Afrika. Nege endemiese spesies is geïdentifiseer waarvan sewe suid van die ewenaar voorkom. Hierdie spesies is R. chimanimaniana, R. elsana, R. haeneliae, R. namibiana, R. sylvicola, R. villicorona en R. zeyheri, met slegs R. moyalica wat in die noorde van Kenia voorkom en R. arabica in Oman. Die kladistiese resultate van hierdie ondersoek toon aan dat die eerste en dus oudste klade wat gevorm het, spesies van suidelike Afrika bevat, wat daarop dui dat Raphionacme in hierdie gebied ontstaan het. Die geofitiese groeivorm is waarskynlik ‘n aanduiding dat dit saamgeval het met die uitdroging van die streek. Daarna het diversifisering en noordwaartse verspreiding van Raphionacme spesies moontlik die ontwikkeling van savanna oos en noord van die oorblywende ekwatoriale reënwoud gevolg. Etlike taksonomiese toepassings, gegrond op die filogenetiese resultate, is aangedui. Die buitegroep Schlechterella abyssinica het in beide die molekulêre en gekombineerde kladogramme deel van die Raphionacme klade gevorm. Dit wil dus voorkom asof die oorplasing van Raphionacme abyssinica na die genus Schlechterella foutief was en dat die spesie S. abyssinica weer na Raphionacme oorgedra moet word. Drie Raphionacme spesies, R. bingeri, R. excisa en R. splendens is onder die naam R. splendens saamgevoeg. Daarna is R. splendens weer in twee subspesies verdeel. Die kladistiese resultate van hierdie studie ondersteun die skepping van Raphionacme splendens subsp. splendens en R. splendens subsp. bingeri.
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Dissertation (M.Sc. (Plant Sciences (Botany))--University of the Free State, 2013, Apocynaceae -- Phylogeny, Plants -- Evolution, Schlechterella affinity, Raphionacme, Molecular taxonomy, Geophyte, Biogeography, African flora, Phylogeny
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