High-throughput sequencing data have greatly improved our ability to understand the processes that contribute to current biodiversity patterns. The "vanishing refuge" diversification model is speculated for the coastal forests of eastern Africa, whereby some taxa have persisted and diversified between forest refugia, while others have switched to becoming generalists also present in non-forest habitats. Complex arrangements of geographical barriers (hydrology and topography) and ecological gradients between forest and non-forest habitats may have further influenced the region's biodiversity, but elucidation of general diversification processes has been limited by lack of suitable data. Here, we explicitly test alternative diversification modes in the coastal forests using genome-wide single nucleotide polymorphisms, mtDNA, spatial and environmental data for three forest (Arthroleptis xenodactyloides, Leptopelis flavomaculatus and Afrixalus sylvaticus) and four generalist (Afrixalus fornasini, A. delicatus, Leptopelis concolor and Leptopelis argenteus) amphibians. Multiple analyses provide insight about divergence times, spatial population structure, dispersal barriers, environmental stability and demographic history. We reveal highly congruent intra-specific diversity and population structure across taxa, with most divergences occurring during the late Pliocene and Pleistocene. Although stability models support the existence of some forest refugia, dispersal barriers and demographic models point towards idiosyncratic diversification modes across taxa. We identify a consistent role for riverine barriers in the diversification of generalist taxa, but mechanisms of diversification are more complex for forest taxa and potentially include topographical barriers, forest refugia and ecological gradients. Our work demonstrates the complexity of diversification processes in this region, which vary between forest and generalist taxa, but also for ecologically similar species with shared population boundaries.
Aims
To quantify the spatial distribution of amphibian phylogenetic endemism (PE), an indicator of potential refugia, to test PE for correlations with current and historical environmental predictors, and to evaluate the effectiveness of current protected areas at conserving evolutionary history.
Location
Coastal Forests of Eastern Africa (CFEA) and the adjacent low‐elevation Eastern Afromontane (EA).
Methods
We integrated new and existing spatial and phylogenetic data to map PE for almost the full amphibian assemblage (41 of 55 species), including 35 intraspecific lineages from several species and complexes showing high phylogeographic structure. Using spatial and non‐spatial regressive models, we tested whether PE can be predicted by measures of Quaternary climate change, forest stability, topographic heterogeneity and current climate. PE results were intersected with the protected area network to evaluate current conservation effectiveness.
Results
We detect refugia in Tanzania and coastal Kenya previously identified as CFEA centres of endemism but also new areas (lowland Tanga region and Pangani river, Zaraninge forest, Mafia island, Matumbi hills). Results show that refugia for amphibians (high PE) are located in areas with long‐term Quaternary climate stability and benign current climate (high precipitation of driest quarter, high annual precipitation), with climatically unstable areas demonstrating low PE. Conservation analyses revealed that ten PE hotspots account for over 25% of the total PE, but only small parts of these areas are under conservation protection.
Main Conclusions
Utilizing cryptic diversity from novel phylogeographic data and distribution modelling improves our understanding of endemism patterns, with climate stability being strongly correlated with the distribution of PE. Our analyses point towards high PE areas being refugia, which require an urgent need to consolidate protected areas within centres of endemism in this highly threatened biodiversity hotspot.
A new species of African reed frog (genus Hyperolius Rapp, 1842) is described from the Coastal Forests of the Eastern Africa Biodiversity Hotspot in northeastern Mozambique. It is currently only known from less than ten localities associated with the Mozambican coastal pans system, but may also occur in the southeastern corner of Tanzania. Phylogenetic reconstructions using the mitochondrial 16S marker revealed that it is the sister taxon of Hyperolius mitchelli (>5.6% 16S mtDNA sequence divergence) and forms part of a larger H. mitchelli complex with H. mitchelli and H. rubrovermiculatus. The new species is distinguished from other closely related Hyperolius species by genetic divergence, morphology, vocalisation, and dorsal colouration.
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Table S1: List of samples used in the DNA (16S) analysis, including voucher numbers, locality information and their GenBank Accession.
Clade column represents lineage membership based onFig. 2, 1 -H. cf. mitchelli, 2 -H. rubrovermiculatus, S1-H. mitchelli sensu stricto Subclade I, S2 -H. mitchelli sensu stricto Subclade II, S3 -H. mitchelli sensu stricto Subclade III, S4 -H. mitchelli sensu stricto Subclade IV, 3 -H. stictus. GenBank accession IDs for new sequences generated for this study are indicated with an asterisk.
Frogs in the genus Xenopus are ubiquitous in sub‐Saharan Africa, yet very little is recorded on their ecology. They are commonly found in anthropogenically disturbed habitats, but how do these compare to conspecifics from natural habitats? The diet of Xenopus borealis from three different sites in Taita Hills, Kenya was established based on a sample of 77 (54 females and 23 males) specimens from two disturbed and one pristine sites. Xenopus borealis from all the sites was found to be a dietary generalist, feeding predominantly on invertebrates. A total of twelve invertebrate orders both terrestrial and aquatic were recorded in addition to amphibian eggs, tadpoles and fish. Frogs from the pristine forest were smaller and had ingested more terrestrial prey items than frogs in the disturbed open habitat ponds. The stomach content (both by mass and quantity) was independent of body size. The results suggest that X. borealis is an opportunistic generalist predator which may be constrained by food availability in its natural habitat. However, disturbed habitats provide abundant food items which are enough to significantly increase the mean size of the population.
The cover image, by Christopher D. Barratt et al., is based on the Biodiversity Research Environmental correlates of phylogenetic endemism in amphibians and the conservation of refugia in the Coastal Forests of Eastern Africa, DOI:. Photo Credit: B.A. Bwong, Afrixalussylvaticus, photographed in Shimba Hills National Reserve, Kenya
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