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Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot

Authors

  • Krystal A. Tolley,

    Corresponding author
    1. Applied Biodiversity Research Division, South African National Biodiversity Institute, Claremont 7735, Cape Town, South Africa
    2. Department of Botany and Zoology, University of Stellenbosch, Matieland 7602, South Africa
      Correspondence: Krystal A. Tolley, Applied Biodiversity Research Division, South African National Biodiversity Institute, Claremont 7735, Cape Town, South Africa.
      E-mail: k.tolley@sanbi.org.za
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  • Colin R. Tilbury,

    1. Department of Botany and Zoology, University of Stellenbosch, Matieland 7602, South Africa
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  • G. John Measey,

    1. Applied Biodiversity Research Division, South African National Biodiversity Institute, Claremont 7735, Cape Town, South Africa
    2. Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa
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  • Michele Menegon,

    1. Tropical Biodiversity Section, Museo Tridentino di Scienze Naturali, Via Calepina 14, I-38100 Trento, Italy
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  • William R. Branch,

    1. Bayworld, P.O. Box 13147, Humewood 6013, South Africa
    2. Department of Zoology, P O Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth 6031, South Africa
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  • Conrad A. Matthee

    1. Department of Botany and Zoology, University of Stellenbosch, Matieland 7602, South Africa
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Correspondence: Krystal A. Tolley, Applied Biodiversity Research Division, South African National Biodiversity Institute, Claremont 7735, Cape Town, South Africa.
E-mail: k.tolley@sanbi.org.za

Abstract

Aim  East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia.

Location  East Africa.

Methods  We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal.

Results  Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre-dates the Pliocene (> 6 Ma). These dates are much older than the Plio-Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages.

Main conclusions  The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa.

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