[Show abstract][Hide abstract] ABSTRACT: The Eastern Afromontane Region (EAR) contains numerous endemic species, yet its reptile diversity remains relatively poorly understood. We used molecular data to examine species diversity of the Sub-Saharan chameleon genus Trioceros. In particular, we focus on establishing species boundaries for taxa with disjunct distributions across the fragmented mountains of the EAR, including T. affinis, T. balebicornutus, T. deremensis, T. harennae, T. tempeli and T. werneri. We applied three species-delimiting approaches, General Mixed Yule-Coalescent (GMYC), a Bayesian implementation of the GMYC, and Bayes Factor Delimitation to estimate species diversity. Using a dated phylogeny, we also examined spatial and temporal diversification patterns in Trioceros. We found strong congruence between different species delimitation approaches, with all methods suggesting that species diversity is currently underestimated. In particular, T. werneri consists of at least four candidate species (i.e. species awaiting description) with some mountain ranges (Uluguru and Udzungwa) having potentially more than one species. Most interspecific divergences between extant Trioceros lineages are estimated to be >5 Mya, consistent with a Pliocene origin of the endemic montane fauna, as exhibited in other taxonomic groups. Multiple, overlapping geographic events (climate and/or geomorphological changes) might account for speciation patterns in Trioceros given the dating results.
Molecular Phylogenetics and Evolution 08/2014; · 4.07 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Euparkeria capensis is resolved as the sister taxon to Archosauria in many cladistic phylogenies and provides a key outgroup which may approximate the ancestral archosaur morphology. Several other taxa have been referred to the family Euparkeriidae, but the monophyly of this taxon remains doubtful and largely untested. To test this monophyly, the archosauriform and putative euparkeriid Dorosuchus neoetus from the Mid-Triassic of Russia is re-examined in the light of recent work on the evolution of stem archosaurs. Dorosuchus neoetus is found to possess a number of morphological features that place it close to Archosauria, including a sigmoidal femur with a clear attachment region for the m. caudifemoralis musculature, but no unambiguous archosaurian apomorphies. Dorosuchus neoetus is included for the first time in a numerical cladistic analysis and is recovered as the sole sister taxon to Archosauria + Phytosauria. A monophyletic Euparkeriidae including D. neoetus and E. capensis is slightly less parsimonious. In addition, a mandible and pterygoid that were previously referred to D. neoetus subsequent to the original description of the species are also included separately within the phylogenetic analysis and are recovered within Archosauria, possibly raising questions as to their correct taxonomic referral. However, this phylogenetic placement is based primarily on the absence of palatal teeth, but the presence or absence of palatal teeth exhibits considerable homoplasy within Archosauriformes. Based on other aspects of their morphology, we do not reject the referral of these elements to D. neoetus.
[Show abstract][Hide abstract] ABSTRACT: AimThe persistence and stability of habitats through time are considered predictors of high levels of biodiversity in some environments. Long-term habitat persistence and stability may explain the species-rich, endemic forest fauna and flora of the Eastern Afromontane Biodiversity Region (EABR). Using complementary phylogenetic and biogeographical approaches, we examine evolutionary patterns in EABR brevicipitid frogs. Using these data, we test whether brevicipitid history reflects patterns of long-term forest persistence and/or stability across the EABR.LocationEast Africa.MethodsA dated phylogeny for brevicipitids was constructed using two nuclear and three mitochondrial markers. Alternative diversification models were used to determine signal for constant or varying net diversification rates. Using our dated tree, we identified areas of high phylogenetic diversity (PD), and inferred ancestral areas using likelihood and Bayesian approaches.ResultsBrevicipitids have a long history, with generic diversification among extant lineages pre-dating the Oligocene (> 33 Ma). Ancestral-area reconstructions indicate the presence of brevicipitids in the EABR since the Oligocene, and support a scenario of palaeoendemics surviving in EABR refugia. Ancestral-area reconstructions indicate that the central Eastern Arc Mountains (EAM) formed the initial centre of diversification of forest brevicipitids. Measures of PD show that diversity varies across the EABR but is highest in the EAM. Constant net diversification rate in brevicipitids is a significantly better fit than alternative, rate-variable models.Main conclusionsThe degree of persistence of forest habitats appears to be a contributing factor to the varying levels of diversity across the EABR in brevicipitids (and other organisms). In contrast to the Southern Highlands and Ethiopian Bale Mountains, the EAM stands out as an area that enabled the constant accumulation of brevicipitid species over a long period of time.
Journal of Biogeography 04/2014; · 4.86 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: According to current understanding, five lineages of amphibians, but no other tetrapods, are secondarily lungless and are believed to rely exclusively on cutaneous gas exchange. One explanation of the evolutionary loss of lungs interprets lunglessness as an adaptation to reduce buoyancy in fast-flowing aquatic environments, reasoning that excessive buoyancy in such an environment would cause organisms being swept away. While not uncontroversial, this hypothesis provides a plausible potential explanation of the evolution of lunglessness in four of the five lungless amphibian lineages. The exception is the most recently reported lungless lineage, the newly described Guyanan caecilian genus and species Caecilita iwokramae Wake & Donnelly, 2010, which is inconsistent with the reduced disadvantageous buoyancy hypothesis by virtue of it seemingly being terrestrial and having a terrestrial ancestry. Re-examination of the previously only known specimen of C. iwokramae and of recently collected additional material reveal that this species possesses a reasonably well-developed right lung and is a species of the pre-existing caecilian genus Microcaecilia Taylor, 1968. We therefore place Caecilita in the synonymy of Microcaecilia, and re-evaluate the plausibility of the reduced disadvantageous buoyancy hy- pothesis as a general explanation of the evolution of lunglessness.
[Show abstract][Hide abstract] ABSTRACT: The phylogenetic relationships of many taxa remain poorly known because of a lack of appropriate data and/or analyses. Despite substantial recent advances, amphibian phylogeny remains poorly resolved in many instances. The phylogenetic relationships of the Ethiopian endemic monotypic genus Ericabatrachus has been addressed thus far only with phenotypic data and remains contentious.
We obtained fresh samples of the now rare and Critically Endangered Ericabatrachus baleensis and generated DNA sequences for two mitochondrial and four nuclear genes. Analyses of these new data using de novo and constrained-tree phylogenetic reconstructions strongly support a close relationship between Ericabatrachus and Petropedetes, and allow us to reject previously proposed alternative hypotheses of a close relationship with cacosternines or Phrynobatrachus.
We discuss the implications of our results for the taxonomy, biogeography and conservation of E. baleensis, and suggest a two-tiered approach to the inclusion and analyses of new data in order to assess the phylogenetic relationships of previously unsampled taxa. Such approaches will be important in the future given the increasing availability of relevant mega-alignments and potential framework phylogenies.
[Show abstract][Hide abstract] ABSTRACT: We analyze mitochondrial genomes to reconstruct a robust phylogenetic framework for caecilian amphibians and use this to investigate life-history evolution within the group. Our study comprises 45 caecilian mitochondrial genomes (19 of them newly reported), representing all families and 27 of 32 currently recognized genera, including some for which molecular data had never been reported. Support for all relationships in the inferred phylogenetic tree is high to maximal, and topology tests reject all investigated alternatives, indicating an exceptionally robust molecular phylogenetic framework of caecilian evolution consistent with current morphology-based supraspecific classification. We used the mitogenomic phylogenetic framework to infer ancestral character states and to assess correlation among three life-history traits (free-living larvae, viviparity, specialized pre-adult or vernal teeth), each of which occurs only in some caecilian species. Our results provide evidence that an ancestor of the Seychelles caecilians abandoned direct development and re-evolved a free-living larval stage. This study yields insights into the concurrent evolution of direct development and of vernal teeth in an ancestor of Teresomata that likely gave rise to skin-feeding (maternal dermatophagy) behavior and subsequently enabled evolution of viviparity, with skin feeding possibly a homologous precursor of oviduct feeding in viviparous caecilians.
Molecular Phylogenetics and Evolution 01/2014; 73:177-189. · 4.07 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Anonymous nuclear markers were developed
for Seychelles caecilian amphibians. Using a previously
published bioinformatics pipeline (developed for Roche
454 data), 36 candidate anonymous nuclear loci (ANL) of
at least 180 bp length were identiﬁed from Illumina MiSeq
next generation sequencing data for ﬁve Seychelles species. We designed primer pairs for the 36 candidate ANL
and tested these by PCR and Sanger sequencing. Seven
ANL ampliﬁed and sequenced well for at least ﬁve of the
six nominal Seychelles caecilian species (in three genera),
and represent potentially useful markers for systematics
[Show abstract][Hide abstract] ABSTRACT: A new species of striped ichthyophiid caecilian, Ichthyophis multicolor sp. nov., is described on the basis of morphological and molecular data from a sample of 14 specimens from Ayeyarwady Region, Myanmar. The new species resembles superficially the Indian I. tricolor Annandale, 1909 in having both a pale lateral stripe and an adjacent dark ventrolateral stripe contrasting with a paler venter. It differs from I. tricolor in having many more annuli, and in many details of cranial osteology, and molecular data indicate that it is more closely related to other Southeast Asian Ichthyophis than to those of South Asia. The caecilian fauna of Myanmar is exceptionally poorly known but is likely to include chikilids as well as multiple species of Ichthyophis.
[Show abstract][Hide abstract] ABSTRACT: The hyperoliid frog Tachycnemis seychellensis, the only species of its genus, is endemic to the four largest granitic islands of the Seychelles archipelago and is reliant on freshwater bodies for reproduction. Its presence in the Seychelles is thought to be the product of a transoceanic dispersal, diverging from the genus Heterixalus, its closest living relative (currently endemic to Madagascar), between approximately 10-35Ma. A previous study documented substantial intraspecific morphological variation among island populations and also among populations within the largest island (Mahé). To assess intraspecific genetic variation and to infer the closest living relative(s) of T. seychellensis, DNA sequence data were generated for three mitochondrial and four nuclear markers. These data support a sister-group relationship between T. seychellensis and Heterixalus, with the divergence between the two occurring between approximately 11-19Ma based on cytb p-distances. Low levels of genetic variation were found among major mitochondrial haplotype clades of T. seychellensis (maximum 0.7% p-distance concatenated mtDNA), and samples from each of the islands (except La Digue) comprised multiple mitochondrial haplotype clades. Two nuclear genes (rag1 and tyr) showed no variation, and the other two (rho, pomc) lacked any notable geographic structuring, counter to patterns observed within presumably more vagile Seychelles taxa such as lizards. The low levels of genetic variation and phylogeographic structure support an interpretation that there is a single but morphologically highly variable species of Seychelles treefrog. The contrasting genetic and morphological intraspecific variation may be attributable to relatively recent admixture during low sea-level stands, ecophenotypic plasticity, local adaptation to different environmental conditions, and/or current and previously small population sizes. Low genetic diversity but substantial morphological variation is unusual within anurans.
Molecular Phylogenetics and Evolution 01/2014; 75:194-201. · 4.07 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Insights into morphological diversification can be obtained from the ways the species of a clade occupy morphospace. Projecting a phylogeny into morphospace provides estimates of evolutionary trajectories as lineages diversified information that can be used to infer the dynamics of evolutionary processes that produced patterns of morphospace occupation. We present here a large-scale investigation into evolution of morphological variation in the skull of caecilian amphibians, a major clade of vertebrates. Because caecilians are limbless, predominantly fossorial animals, diversification of their skull has occurred within a framework imposed by the functional demands of head-first burrowing. We examined cranial shape in 141 species, over half of known species, using X-ray computed tomography and geometric morphometrics. Mapping an existing phylogeny into the cranial morphospace to estimate the history of morphological change (phylomorphospace), we find a striking pattern: most species occupy distinct clusters in cranial morphospace that closely correspond to the main caecilian clades, and each cluster is separated by unoccupied morphospace. The empty spaces in shape space are unlikely to be caused entirely by extinction or incomplete sampling. The main caecilian clades have different amounts of morphological disparity, but neither clade age nor number of species account for this variation. Cranial shape variation is clearly linked to phyletic divergence, but there is also homoplasy, which is attributed to extrinsic factors associated with head-first digging: features of caecilian crania that have been previously argued to correlate with differential microhabitat use and burrowing ability, such as subterminal and terminal mouths, degree of temporal fenestration (stegokrotaphy/zygokrotaphy), and eyes covered by bone, have evolved and many combinations occur in modern species. We find evidence of morphological convergence in cranial shape, among species that have eyes covered by bone, resulting in a narrow bullet-shaped head. These results reveal a complex history, including early expansion of morphospace and both divergent and convergent evolution resulting in the diversity we observe today.
[Show abstract][Hide abstract] ABSTRACT: A taxonomic review of the monogeneric northeast Indian caecilian family Chikilidae is presented based on 64 specimens. Chikila fulleri (Alcock, 1904), known previously only from a single specimen collected more than 100 years ago, is rediagnosed and characterised based on recent collections. We describe three additional species new to science, Chikila alcocki sp. nov., Chikila darlong sp. nov., and Chikila gaiduwani sp. nov. This species-level taxonomy is consistent with mitochondrial DNA sequence data. A key to the species of Chikila is presented.
[Show abstract][Hide abstract] ABSTRACT: We describe the distinctive but previously overlooked shape and position of the fat bodies of some scolecomorphid caecilians. Anteriorly positioned, finger-like fat bodies projecting into the coelom from narrow bases in the genital mesenteries are found in both sexes of all species of the East African caecilian genus Scolecomorphus and are unknown elsewhere. We interpret this as a visceral synapomorphy of Scolecomorphus that may be related to viviparity in this genus.
African Journal of Herpetology 06/2013; 62(1):21-27. · 0.82 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Batrachochytrium dendrobatidis (Bd) is commonly termed the 'amphibian chytrid fungus' but thus far has been documented to be a pathogen of only batrachian amphibians (anurans and caudatans). It is not proven to infect the limbless, generally poorly known, and mostly soil-dwelling caecilians (Gymnophiona). We conducted the largest qPCR survey of Bd in caecilians to date, for more than 200 field-swabbed specimens from five countries in Africa and South America, representing nearly 20 species, 12 genera, and 8 families. Positive results were recovered for 58 specimens from Tanzania and Cameroon (4 families, 6 genera, 6+ species). Quantities of Bd were not exceptionally high, with genomic equivalent (GE) values of 0.052-17.339. In addition, we report the first evidence of lethal chytridiomycosis in caecilians. Mortality in captive (wild-caught, commercial pet trade) Geotrypetes seraphini was associated with GE scores similar to those we detected for field-swabbed, wild animals.
[Show abstract][Hide abstract] ABSTRACT: Amphibian chytrid fungus Batrachochytrium dendrobatidis (Bd) has been hypothesised to be an indigenous parasite of African amphibians. In Cameroon, however, previous surveys in one region (in the northwest) failed to detect this pathogen, despite the earliest African Bd having been recorded from a frog in eastern Cameroon, plus one recent record in the far southeast. To reconcile these contrasting results, we present survey data from 12 localities across 6 regions of Cameroon from anurans (n = 1052) and caecilians (n = 85) of ca. 108 species. Bd was detected in 124 amphibian hosts at 7 localities, including Mt. Oku, Mt. Cameroon, Mt. Manengouba and lowland localities in the centre and west of the country. None of the hosts were observed dead or dying. Infected amphibian hosts were not detected in other localities in the south and eastern rainforest belt. Infection occurred in both anurans and caecilians, making this the first reported case of infection in the latter order (Gymnophiona) of amphibians. There was no significant difference between prevalence and infection intensity in frogs and caecilians. We highlight the importance of taking into account the inhibition of diagnostic qPCR in studies on Bd, based on all Bd-positive hosts being undetected when screened without bovine serum albumin in the qPCR mix. The status of Bd as an indigenous, cosmopolitan amphibian parasite in Africa, including Cameroon, is supported by this work. Isolating and sequencing strains of Bd from Cameroon should now be a priority. Longitudinal host population monitoring will be required to determine the effects, if any, of the infection on amphibians in Cameroon.
Diseases of Aquatic Organisms 02/2013; 102(3):187-94. · 1.73 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Surveys of the potentially lethal amphibian chytrid fungus (Batrachochytrium dendrobatidis - Bd) in Africa are patchy, especially in some regions of high species endemicity. We present results of the first Bd surveys of wild amphibians in Ethiopia, for two upland regions on either side of the Rift Valley: the Bale Mountains and the Kaffa region. Surveys were opportunistic so that robust interpretation of the data is limited. Utilizing diagnostic qPCR assays, 51 out of 120 frogs (14 species in 10 genera) tested positive for Bd at altitudes of 1,620–3,225 m, across all genera and species, and all but two localities. Prevalence was not significantly different between the two regions or two years (2008, 2009) sampled. Prevalence and parasite load was higher in species with aquatic tadpoles than those with terrestrial early life-history stages, but these differences were not significant. Impacts of Bd infection were not investigated, but no dead or dying frogs were found. This is the first report of Bd in Ethiopia, a country in which approximately 40% of its more than 60 species are endemic. Declines have occurred in some frog species in some localities in Ethiopia, and although habitat degradation is a likely cause in at least some places, further studies of Bd in Ethiopia are required to understand if it is a threat.