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Albatrosses and other seabirds are generally highly philopatric, returning to natal colonies when they achieve breeding age. This is not universal, however, and cases of extraordinary vagrancy are rare. The Tristan Albatross (Diomedea dabbenena) breeds on Gough Island in the South Atlantic Ocean, with a small population on Inaccessible Island, Tristan da Cunha, ca 380 km away. In 2015, we observed an adult male albatross in Gonydale, Gough Island, which had been ringed on Ile de la Possession, Crozet Islands in 2009 when it was assumed to be an immature Wandering Albatross (D. exulans). We sequenced 1109 bp of the cytochrome b mitochondrial gene from this bird, and confirmed it to be a Tristan Albatross, meaning its presence on Crozet 6 years previous, and nearly 5000 km away, was a case of prospecting behaviour in a heterospecific colony. Given the challenges in identifying immatureDiomedeaalbatrosses, such dispersal events may be more common than thought previously.

Although Conraua goliath is well known as the largest living frog species, the diversity and evolution of the genus Conraua across sub‐Saharan Africa remain poorly understood. We present multilocus phylogenetic analyses of the six currently recognized species that provide insights into divergence times, biogeography, body size evolution and undescribed species. An analysis of divergence times demonstrates that crown‐group Conraua arose some time during the latest Oligocene to mid‐Miocene followed by divergence into major lineages in the mid‐Miocene that may reflect the fragmentation of widespread tropical forests in Africa that began at this time. We find three pairs of sister species, C. crassipes + C. beccarii, C. alleni + C. derooi and C. goliath + C. robusta, each of which diverged during the Miocene. These relationships reject phylogenetic hypotheses based solely on biogeography as the geographically peripheral C. beccarii from north‐eastern Africa is nested within western African species and the Central African species do not form a clade. Our species delimitation analyses provide support for undescribed species in C. alleni, C. beccarii and C. derooi, and possibly C. crassipes, suggesting that the current taxonomy substantially underestimates species diversity. There is no clear directional trend of either increasing or decreasing body size in Conraua and the three largest species do not form a clade. With a robust phylogenetic hypothesis in hand, further field‐based studies are needed to understand the evolution of morphology and life history in this charismatic African anuran clade.

The taxonomic validity of the East African reed frogs Hyperolius mitchelli Loveridge, 1953 and Hyperolius rubrovermiculatus Schiotz, 1975 has been questioned in the literature, with the latter havi...

The spiny-throated reed frog species group is a small radiation of Hyperolius frogs from East Africa. Unlike many members of the genus which have relatively wide distributions, these species tend to be small-range endemics found in montane and submontane forests. Recent discovery of a golden-hued frog with the clade-specific traits of spines on their gular discs prompted a morphological and genetic exploration of the distinctness of this new lineage and relationships to other members of the clade. Genetic (mitochondrial and nuclear loci) results resolved many sister-relationships, but deeper nodes in the phylogeny were poorly resolved. A reduced-representation genome-wide Single Nucleotide Polymorphism (SNP) dataset was able to fully resolve the phylogenetic relationships within this clade, placing this new lineage, here named after the mountain range in which is it found–H. ukaguruensis sp. nov., as an early diverging lineage within the group. This new species is distinct from all other spiny-throated reed frogs, necessitating further understanding as a single-mountain endemics vulnerable to habitat loss and potential decline. Morphometric analyses identify clear morphological characteristics that are distinct for the herein described species, most noticeably in that the eyes are significantly smaller than other members of the genus for which we have samples.

Using integrative approaches, a new large-bodied species of Phrynobatrachus is described from a series of 48 specimens from the montane forests of the West Usambara and North Pare Mountains of Tanz...

African amphibian diversity remains underestimated with many cryptic lineages awaiting formal description. An important hotspot of amphibian diversification is the Guineo-Congolian rainforest in Central Africa, its richness attributable to present day and ancestral range fragmentation through geological barriers, habitat expansion and contraction, and the presence of steep ecological gradients. The charismatic Nectophryne tree toads present an interesting case study for diversification in this region. The two formally described species comprising this genus show nearly identical geographic distributions extending across most of the Guineo-Congolian rainforest, but show little morphological disparity. Both species harbour extensive genetic diversity warranting taxonomic revisions, and interestingly, when comparing the subclades within each, the two species show remarkably parallel diversification histories, both in terms of timing of phylogenetic splits and their geographic distributions. This indicates that common processes may have shaped the evolutionary history of these lineages.

Streicher, Jeffrey W., Sadler, Rowland, Loader, Simon P. (2020): Amphibian taxonomy: early 21st century case studies. Journal of Natural History 54 (1-4): 1-13, DOI: 10.1080/00222933.2020.1777339, URL: http://dx.doi.org/10.1080/00222933.2020.1777339

Nothophryne Poynton, 1963 is a monotypic genus of frog, with the nominal species N. broadleyi found only on Mount Mulanje, in southern Malawi. Recent surveys in northern Mozambique, however, have u...

We present the first annotated amphibian checklist of Shimba Hills National Reserve (SHNR). The list comprises of 30 currently known amphibians (28 anurans and two caecilians), which includes 11 families and 15 genera. In addition, individual records per species, distribution in the reserve and brief remarks about the species are presented. The checklist is based on information from museum collections, field guides, unpublished reports and newly collected field data. We are able to confirm the presence of two Eastern Afromontane species in the SHNR: Scolecomorphus cf. vittatus and Callulina cf. kreffti. The latter has not been recorded since the original collection of a single specimen over 50 years ago. SHNR contains the highest number of amphibian species of any known locality in Kenya (about 30% of the country's total number); therefore it is of national conservation importance. Finally, we briefly discuss the biogeography of the SHNR and its connections to nearby biogeographic regions.

A new species of tree snake Dipsadoboa montisilva Branch, Conradie & Tolley sp. nov. (Serpentes: Colubridae) is described from the ‘sky islands’ of Mount Mabu and Mount Ribáuè in northern Mozambique. Features of scalation, colour, body form and habitat distinguish the new species from other Dipsadoboa. This is supported by a phylogenetic analysis using one mitochondrial marker (cytochrome b) that shows the new Mozambican species is divergent from other sampled Dipsadoboa, including D. flavida and D. aulica, the only congeners known to occur in Mozambique. Morphologically, the new Dipsadoboa forms part of the D. werneri-shrevei complex from east and southeast Africa, but differs in having higher subcaudal counts, a different temporal pattern and only two supralabials entering the orbit. Phylogenetically, it occurs in a clade with D. shrevei and D. werneri. The status of D. shrevei in East Africa is reassessed, particularly in terms of the poorly-known Dipsadoboa shrevei kageleri from northern Tanzania. It is morphologically well defined from D. shrevei shrevei and utilises a different habitat. Although based on limited genetic data, it appears to be well-defined from typical D. shrevei and is accordingly raised to specific status. The only Tanzanian record for typical D. shrevei from Mtene, Rondo Plateau in southeast Tanzania is well isolated from the species’ range to the west (e.g. Zambia, Angola) and the published scalation features, particularly ventral counts, do not fully accord with D. shrevei. The Rondo Plateau population is treated as Dipsadoboa incerta sedis, and because we return D. shrevei to its binomial status, we can no longer consider D. shrevei as occurring in Tanzania. Biogeographically, the Rondo Plateau population may have a stronger affinity to the new Mozambican species. The discovery of isolated populations of the new species in mid-altitude forest remnants on Mt Mabu and Mt Ribáuè emphasizes the high conservation importance of the Mozambique forest ‘sky islands’ from which numerous other endemic new species have been recently discovered. These species are impacted by ongoing habitat destruction through slash and burn clearing for subsistence agriculture.

The niche-filling process predicted by the “ecological opportunity” (EO) model is an often-invoked mechanism for generating exceptional diversity in island colonizers. Whether the same process governs lineage accumulation and trait disparity during continental colonization events is less clear. Here, we test this prediction by investigating the rate dynamics and trait evolution of one of Africa's most widespread amphibian colonizers, the true toads (Bufonidae). By reconstructing the most complete molecular phylogeny of African Bufonidae to date, we find that the diversification of lineages in Africa best conforms to a constant rate model throughout time and across subclades, with little support for EO. Evolutionary rates of life-history traits have similarly been constant over time. However, an analysis of generalists and specialists showed a shift toward higher speciation rates associated with habitat specialization. The overall lack of EO signal can be interpreted in a number of ways and we propose several explanations. Firstly, methodological issues might preclude the detection of EO. Secondly, colonizers might not experience true EO conditions and due to the size, ecological heterogeneity and age of landmasses, the diversification processes might be more complex. Thirdly, lower speciation rates of habitat generalists may have affected overall proliferation of lineages.

Narrow-mouthed frogs (Anura: Microhylidae) are globally distributed and molecular data suggest the rapid evolution of multiple subfamilies shortly after their origin. Despite recent progress, several subfamilial relationships remain unexplored using phylogenomic data. We analysed 1,796 nuclear ultraconserved elements, a total matrix of 400,664 nucleotides, from representatives of most microhylid subfamilies. Summary method species-tree and maximum likelihood analyses unambiguously supported Hoplophryninae as the earliest diverging microhylid and confirm Chaperininae as a junior synonym of Microhylinae. Given the emerging consensus that subfamilies from mainland Africa diverged early, microhylids have likely occupied the continent for more than 66 million years.

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, L. 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 toward 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.

Theory predicts that sexually dimorphic traits under strong sexual selection, particularly those involved with intersexual signaling, can accelerate speciation and produce bursts of diversification. Sexual dichromatism (sexual dimorphism in color) is widely used as a proxy for sexual selection and is associated with rapid diversification in several animal groups, yet studies using phylogenetic comparative methods to explicitly test for an association between sexual dichromatism and diversification have produced conflicting results. Sexual dichromatism is rare in frogs, but it is both striking and prevalent in African reed frogs, a major component of the diverse frog radiation termed Afrobatrachia. In contrast to most other vertebrates, reed frogs display female-biased dichromatism in which females undergo color transformation, often resulting in more ornate coloration in females than in males. We produce a robust phylogeny of Afrobatrachia to investigate the evolutionary origins of sexual dichromatism in this radiation and examine whether the presence of dichromatism is associated with increased rates of net diversification. We find that sexual dichromatism evolved once within hyperoliids and was followed by numerous independent reversals to monochromatism. We detect significant diversification rate heterogeneity in Afrobatrachia and find that sexually dichromatic lineages have double the average net diversification rate of monochromatic lineages. By conducting trait simulations on our empirical phylogeny, we demonstrate that our inference of trait-dependent diversification is robust. Although sexual dichromatism in hyperoliid frogs is linked to their rapid diversification and supports macroevolutionary predictions of speciation by sexual selection, the function of dichromatism in reed frogs remains unclear. We propose that reed frogs are a compelling system for studying the roles of natural and sexual selection on the evolution of sexual dichromatism across micro- and macroevolutionary timescales.

We present an account of the 909 globally threatened taxa (793 species, 74 subspecies, 42 varieties) of animals and plants in the Eastern Arc Mountains and Coastal Forests of Kenya and Tanzania and the sites in which they occur based upon a review of the 2015 IUCN Red List of Threatened Species. Results for animals are summarised by Class (Amphibia, Aves, Gastropoda, Insecta, Malacostraca, Mammalia, Reptilia) and presented for plants as a whole (Classes Bryopsida, Cycadopsida, Jungermanniopsida, Liliopsida, Lycopodiopsida, Magnoliopsida, Pinopsida, Polypodiopsida). We analyse the status of previously known and newly identified sites in which globally threatened biodiversity occurs and summarise the current state of research on the globally threatened and ecologically critical biodiversity of the EACF. We then provide recommendations for future research, environmental regulations, and management regimes based upon comprehensive and reliable data to ensure the continued survival of the EACF’s biodiversity within the context of sustainable resource utilisation.Keywords: biodiversity, conservation, endemism, hotspot, IUCN Red List

Populations of many frogs have declined alarmingly in recent years, placing nearly one third of the > 6,000 species under threat of extinction. Declines have been attributed largely to habitat loss, environmental degradation and/or infectious diseases such as chytridiomycosis. Many frogs undergo dramatic natural population fluctuations such that long-term data are required to determine population trends without undue influence of stochastic factors. We present long-term quantitative data (individuals encountered per person hour of searching) for four monotypic frog genera endemic to an Afromontane region of exceptional importance but growing conservation concern: one endemic to the Ethiopian highlands (Spinophrynoides osgoodi) and three endemic to the Bale Mountains (Altiphrynoides malcolmi, Balebreviceps hillmani, Ericabatrachus baleensis), collected during 15 field trips to the Bale Mountains between 1971 and 2009. Only a single confirmed sighting of S. osgoodi has been made since 1995. The other three species have also declined, at least locally. E. baleensis appears to have been extirpated at its type locality and at the same site B. hillmani has declined. These declines are in association with substantial habitat degradation caused by a growing human population. Chytrid fungus has been found on several frog species in Bale, although no dead or moribund frogs have been encountered. These results expose an urgent need for more amphibian surveys in the Bale Mountains. Additionally, we argue that detrimental human exploitation must be halted immediately in at least some parts of the Harenna Forest if a conservation crisis is to be averted.

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.

Aim: The 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. Location: East Africa. Methods: A 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. Results: Brevicipitids 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 conclusions: The 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. Fil: Loader, Simon P.. Universidad de Basilea; Suiza Fil: Ceccarelli, Fadia Sara. Universidad de Basilea; Suiza. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Menegon, Michele. Museo delle Scienze di Trento; Italia Fil: Howell, Kim M.. University of Dar es Salaam; Tanzania Fil: Kassahun, Roman. Ethiopian Wildlife Conservation Authority; Etiopía Fil: Mengistu, Abebe A.. Adama Science and Technology University; Etiopía Fil: Saber, Samy A.. Al-Azhar University; Egipto Fil: Gebresenbet, Fikirte. Oklahoma State University; Estados Unidos Fil: de Sa, Rafael. University of Richmond; Estados Unidos Fil: Davenport, Tim R. B.. Wildlife Conservation Society; Tanzania Fil: Larson, Joanna G.. Harvard University; Estados Unidos Fil: Müller, Hendrik. Universitat Jena; Alemania Fil: Wilkinson, Mark. Natural History Museum; Reino Unido Fil: Gower, David J.. Natural History Museum; Reino Unido

Little is known about reproductive trade-offs in African amphibians, but such data, particularly in the form of quantitative measurements, are a key for investigating life history evolution. Here we compile and analyze known data on African bufonids from published material and new data from preserved museum specimens, to investigate interspecific patterns of egg and clutch sizes variation. Our data is a composite of mixed sources, including ova data from dissected females and laid clutches from observations in the field. Our study shows that, as body size increases, clutch size increases but egg size decreases, and when correcting for body size, egg size is inversely correlated with clutch size. These parameter interactions however, are different for different reproductive modes. In free-swimming larval developing species, the same trends are recovered, but for lecithotrophic viviparous species no significant correlations could be recovered for clutch size and body size nor for the trade-off between clutch size and egg size, and egg size is positively related to body size. The egg size of Nimbaphrynoides occidentalis (Angel, 1943) is a clear outlier, which may be due to its matrotrophic viviparous reproduction. In addition, we observed no statistical difference between ova data collected from dissections and laid clutch data from field observations, which suggests that such a mixed dataset has utility in comparative analyses.

The 15th meeting of the African Amphibian Working Group celebrated the contributions of John Charles Poynton to African Herpetology. The meeting, in Trento, Italy, was timed to commemorate John's 81st birthday (in May 2012), and he presented a keynote speech that focused on a topic that has long fascinated him - the Afrotemperate amphibian fauna. His presentation is published here, in a modified form, for this special issue of African Journal of Herpetology (Poynton 2013). Other papers on the main themes of John's research are also included in this issue, which include biogeography (Loader et al. 2013; Schreiner et al. 2013), taxonomy (Cruz et al. 2013) and morphology (Wilkinson et al. 2013).

How evolutionary novelties evolve is a major question in evolutionary biology. It is widely accepted that changes in environmental conditions shift the position of selective optima, and advancements in phylogenetic comparative approaches allow the rigorous testing of such correlated transitions. A longstanding question in vertebrate biology has been the evolution of terrestrial life histories in amphibians and here, by investigating African bufonids, we test whether terrestrial modes of reproduction have evolved as adaptations to particular abiotic habitat parameters. We reconstruct and date the most complete species-level molecular phylogeny and estimate ancestral states for reproductive modes. By correlating continuous habitat measurements from remote sensing data and locality records with life-history transitions, we discover that terrestrial modes of reproduction, including viviparity evolved multiple times in this group, most often directly from fully aquatic modes. Terrestrial modes of reproduction are strongly correlated with steep terrain and low availability of accumulated water sources. Evolutionary transitions to terrestrial modes of reproduction occurred synchronously with or after transitions in habitat, and we, therefore, interpret terrestrial breeding as an adaptation to these abiotic conditions, rather than an exaptation that facilitated the colonization of montane habitats., Swiss National Science Fund (grant no. 31003A-133067 awarded to S.P.L. and P2BSP3_158846 to H.C.L.); Freiwillige Akademische Gesellschaft Basel; National Geographic Expedition Grant (CRE Grant #8532-08); Swiss Academy of Science (SCNAT); German Science Foundation (MU2914/2-1); Percy Sladen Memorial Fund (to D.J.G.).

Biodiversity research is becoming increasingly dependent on genomics, which allows the unprecedented digitization and understanding of the planet’s biological heritage. The use of genetic markers i.e. DNA barcoding, has proved to be a powerful tool in species identification. However, full exploitation of this approach is hampered by the high sequencing costs and the absence of equipped facilities in biodiversity-rich countries. In the present work, we developed a portable sequencing laboratory based on the portable DNA sequencer from Oxford Nanopore Technologies, the MinION. Complementary laboratory equipment and reagents were selected to be used in remote and tough environmental conditions. The performance of the MinION sequencer and the portable laboratory was tested for DNA barcoding in a mimicking tropical environment, as well as in a remote rainforest of Tanzania lacking electricity. Despite the relatively high sequencing error-rate of the MinION, the development of a suitable pipeline for data analysis allowed the accurate identification of different species of vertebrates including amphibians, reptiles and mammals. In situ sequencing of a wild frog allowed us to rapidly identify the species captured, thus confirming that effective DNA barcoding in the field is possible. These results open new perspectives for real-time-on-site DNA sequencing thus potentially increasing opportunities for the understanding of biodiversity in areas lacking conventional laboratory facilities.

Amphibian chytrid fungus Batrachochytrium dendrobatidis (Bd) has been hypothe- sised to be an indigenous parasite of African amphibians. In Cameroon, however, previous sur- veys 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 south- east. 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. Manen- gouba 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.

Based on morphology there are currently three described East African species of torrent frogs belonging to the tropical African genus Petropedetes (P. dutoiti P. martiennseni and P. yakusini). Taxonomic uncertainties are evident within this group: a putative new species reported from the Nguru Mountains and the species P. yakusini have a fragmented distribution across the Southern Tanzanian highlands. We conducted a molecular systematic study sampling populations of East African petropedetids occurring in the mountains of Tanzania (P. martiennseni and P. yakusini) to investigate species boundaries and biogeography. Data provide evidence for the recognition of well defined genetically distinct populations occurring in each mountain fragment (for P. yakusini) and further support for a new undescribed species from the Nguru Mountains. We examine phylogenetic patterns in relation to the biogeographic history of the region. © 2013 Herpetological Association of Africa.

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. © 2013 International Association for Ecology and Health.

Ecological niche models (ENMs) have been used in a wide range of ecological and evolutionary studies. In biogeographic studies these models have, among other things, helped in the discovery of new allopatric populations, and even new species. However, small sample sizes and questionable taxonomic delimitation can challenge models, often decreasing their accuracy. Herein we examine the sensitivity of ENMs to the addition of new, geographically isolated populations, and the impact of applying different taxonomic delimitations. The East African reed frog Hyperolius substriatus Ahl, 1931 was selected as a case study because it has been the subject of previous ENM predictions. Our results suggest that addition of new data and reanalysis of species lineages of H. substriatus improved our understanding of the evolutionary history of this group of frogs. ENMs provided robust predictions, even when some populations were deliberately excluded from the models. Splitting the lineages based on genetic relationships and analysing the ENMs separately provided insights about the biogeographical processes that led to the current distribution of H. substriatus.

Molecular analysis indicates that African material previously referred to Amietophrynus maculatus (Hallowell, 1854; now Sclerophrys maculata), is divisible into two distinct clades: a Western Clade from Cameroon westwards and an Eastern Clade from Central African Republic eastwards, and Uganda southwards to South Africa, extending to Angola-Namibia. Preliminary morphological and bioacoustic data support this division. The two clades are recognised here as two separate species. The Western species retains the name S. maculata, with Hallowell's designated type locality of Liberia. The Eastern Clade retains the name published by Mertens (1937), S. pusilla. It is noted that a type specimen of S. maculata cannot be traced and is presumed lost; the so-called syntypes in the Philadelphia Academy of Natural Sciences are not the material described by Hallowell. None of these have been designated as a neotype, consequently a specimen from Liberia in the collection of the Natural History Museum, London, is designated here as the neotype of S. maculata.

We characterise nine polymorphic microsatellites for the caecilian amphibian Boulengerula (cf.) uluguruensis. We found between five and 13 alleles per locus in 17 individuals from six sites across four Eastern Arc Mountain blocks and coastal forest in Tanzania. In the population with the largest sample size (Uluguru North, n = 8), two loci deviated from Hardy–Weinberg equilibrium. We also report cross-species utility of these markers in three other Boulengerula species (B. taitanus, B. niedeni, and B. boulengeri), and populations representing potentially undescribed species, approximately proportionate to their phylogenetic divergence from B. uluguruensis. The loci have the potential to quantify reproductive success (in view of skin feeding, a unique mode of parental care) and to determine the genetic structure of local populations, providing vital information for conservation studies of this endangered and little studied genus.

Amphibians of the Eastern Arc Mountains are poorly known, apart from those of the East Usambara, Uluguru, and Udzungwa Mountains. Here, we detail specimens collected in the northwestern Eastern Arc Mountains (West Usambara, South Pare, and North Pare) over the past 100 years. Although not comprehensive, the list is compiled from specimens held at institutions in Europe and the United States and in personal collections; it includes most of the specimens known from this area. From these specimens, we describe one new species of the frog genus Arthroleptis and discuss other specimens that may represent new species. We add substantially to the known diversity in these areas. Using the compiled species list, we assess species diversity and relate the differences to sampling bias and possible biogeographic differences among the mountains. The future preservation of amphibians across this area is of concern given the increasing loss of forest.

The Mahenge Mountains are located in the Eastern Arc Mountains, part of a global biodiversity hotspot. Few amphibian and reptile surveys have been conducted in this area, with only one study of the submontane forest zone. This previous study revealed a characteristic endemic Eastern Arc amphibian and reptile forest fauna. New studies outlined here extend the sampling time considerably and the geographical coverage (including Mselezi Forest Reserve). These studies reveal a remarkable diversity of amphibians and reptiles increasing known faunal diversity considerably, including several putative new species that await further taxonomic verification. Biogeographical analyses using Bray-Curtis indices show Mahenge clusters with the geographically close Udzungwa Mountains and Southern Highlands (non–Eastern Arc highland fragment including Mt. Rungwe). This clustering of areas suggests an interesting biogeographical history and deserves special attention in the future. In light of the remarkable amphibi...

Material ascribed to the genus Callulina from north-east Tanzania and south-east Kenya is assessed. Three newspecies of Callulina are described from the North (Callulina laphami sp. nov.) and South (Callulina shengenasp. nov. and Callulina stanleyi sp. nov.) Pare Mountains in Tanzania. The species are diagnosed based onmorphological, acoustic, and molecular data. A new key to the species of Callulina is provided. Based on aninterpretation of the International Union for Conservation of Nature (IUCN) red list, we suggest that the threespecies will qualify as critically endangered, because of their small distributions and the ongoing threat to theirhabitat. We reveal the high local endemism of Callulina in the northern part of the Eastern Arc Mountains, witheach species restricted to no more than one mountain (fragment) block. This high local endemism in Callulina isprobably widespread across the Eastern Arc, raising further conservation concern for this group of amphibians.Based on new molecular phylogenetic data for Callulina, we discuss biogeographical relationships among northeastTanzanian mountains, and evolutionary patterns in Eastern Arc breviciptids.

The type series of Nectophrynoides viviparus (Tornier, 1905) from the southern Tanzanian highlands has been rediscovered in the Museum für Naturkunde, Berlin. A re-description is given and a lectotype designated. Available topotypic material is included in our assessment of morphological variation in this population. With clarification of the species’ identity, nomenclatural confusions can be addressed. We reject the designation of an earlier neotype, and also of records attributed to the species north and east of the Southern Highlands.

A new species Callulina dawida is described from the Taita Hills, Kenya. It is distinguished from other members of the genus on the basis of the degree of digital expansion. The species further differs from other members of the genus based on molecular sequence comparisons and on its call. The morphological variation in the new species is described, including a comparison of internal and external characters and sexual dimorphism with other species of Callulina. The conservation status of the species, on the basis of its restricted distribution and land use changes in the area, is considered to be of high concern. An updated key of the species of Callulina is provided.

The external and musculoskeletal morphology of the head is described for an ontogenetic series of the scolecomorphid caecilian Scolecomorphus kirkii. The rostral region of foetuses and juveniles is expanded into large, posterolaterally pointing paraoral processes that are formed by the maxilla. Extraoral teeth are present on the underside of the rostrum and laterally on the paraoral processes. In the foetuses, teeth are covered by epidermal tissue. The endoskeletal part of the foetal skull is largely cartilaginous, but all of the dermal bones, with the exception of the squamosal, are present. The foetal chondrocranium is extensively developed and shows a peculiar, posterolateral process of the nasal capsule that is connected to the trabecula cranii by a transverse bar posterior to the choana, and extends further posterior beyond the level of the posterior end of the pila antotica. Only two mm. adductor mandibulae are present, together with two pterygoideus muscles that insert onto the lower jaw. The palatoquadrate and quadrate of foetuses and juveniles, respectively, are highly mobile. It is suggested that the derived head morphology of Scolecomorphus foetuses and juveniles is an adaptation to specialized postparitive feeding. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 491–504.

Specimens of Arthroleptis attaining a body length of nearly 58 mm have been collected above 1790 m a.s.l. on the Nguru Mountains, part of the Eastern Arc mountain chain of Tanzania. In body size this material slightly surpasses the other giant Eastern Arc high-altitude species A. tanneri and A. nikeae, and differs in the proportions of the foot, and patterning of the chin. The Nguru material is referred to a new species which qualifies as being critically endangered on account of its apparently lim- ited geographical range. A clade including the three giant Arthroleptis species seems likely, showing par- allels with several other isolated forest-dwelling Eastern Arc high-altitude anurans.

North-to-south clinal variation in morphology is described in Arthroleptis affinis of the Eastern Arc mountains of Tanzania. While northern populations are readily separable from other Arthroleptis species, southern populations are not. This complicates differential diagnosis. Despite this, it is concluded that in the present state of knowledge, it is more useful to include the clinal variation under the name A. affinis, rather than attempt to divide the species into groups that defy clear morphological definition.

The Kenyan caeciliid caecilian Boulengerula denhardtiNieden 1912 was considered to be a junior synonym of Schistometopum (Dermophis) gregorii (Boulenger, 1894) by Loveridge (1936). This synonymy has been accepted, implicitly or explicitly, by all subsequent workers. Re-examination of the holotype and only known specimen of B. denhardti confirmed Nieden's (1912) assessment that this form represents a distinctive species of Boulengerula. We resurrect B. denhardti from the synonymy of Schistometopum gregorii, rediagnose the species, redescribe the holotype, and provide an updated key to the species of Boulengerula. The phylogenetic relationships of the species of Boulengerula are examined on the basis of morphological data. It is concluded that the available morphological data are insufficient to resolve relationships with any confidence at this time. The caecilian fauna of the Tana River region is briefly discussed. Boulengerula denhardti (Gymnophiona: Caeciliidae) wurde von Nieden (1912) anhand eines einzigen Exemplars aus dem Gebiet des Tana Flusses im sudostlichen Kenia beschrieben. Im Zuge seiner eigenen Untersuchungen zur Gymnophionenfauna des Tana-Deltas uberfuhrte Loveridge (1936) B. denhardtii in die Synonymie von Schistometopum gregorii. Diese Synonymisierung erfolgte offensichtlich aufgrund einer Fehlinterpretation der publizierten Beschreibung. Unsere Untersuchung des Holotypus von B. denhardti zeigt, dass es sich hierbei um eine distinkte, valide Art handelt, welche wir hier neu beschreiben und neu diagnostizieren. Ein neuer Bestimmungsschlussel fur alle bekannten Arten von Boulengerula wird ebenfalls vorgestellt. Die Verwandtschaftsbeziehungen der einzelnen Boulengerula-Arten werden neu untersucht, wobei gezeigt wird, dass die bekannten morphologischen Merkmale zu deren Auflosung nicht ausreichen. Die Gymnophionenfauna des Tanagebietes wird kurz diskutiert.

Bergmann's rule predicts increasing body sizes at higher elevations. The elevational Rapoport's rule predicts an increase of elevational range size with higher elevations. Both rules have often been related to effects of temperature. Larger bodies allow more efficient heat preservation at lower temperature, explaining Bergmann's rule. Higher temperature variability may select for adaptations that allow increased range sizes, explaining Rapoport's rule. The generality of both rules has been challenged and evidence towards explanatory mechanisms has been equivocal. We investigated temperature and its variability as explanations for Bergmann's and Rapoport's rule in moths along an elevation gradient in Switzerland. In particular, we tested for relationships between elevation, temperature and body size across almost 300 species of Macrolepidoptera along a gradient from 600 to 2400 m a.s.l. The gradient was resampled throughout the vegetation season, which allowed assessing temperature effects independently from elevation. We controlled analyses for covariate traits of moths and their phylogeny. We found a positive relationship between body size and elevation, but no link with temperature. Furthermore, there was no positive link between average elevation and elevational range, but there was between temperature variability and elevational range. We conclude that mechanisms other than temperature can lead to increasing body sizes with elevation (supporting Bergmann's pattern, but not the mechanism). Contrary to that, data support the mechanism for Rapoport's rule: high temperature variability is associated with large ranges. However, because temperature variability is not necessarily increasing with elevation, it may not always lead to the geographic pattern predicted.

Multiple sources of data in combination are essential for species delimitation and classification of difficult taxonomic groups. Here we investigate a cicada taxon with unusual cryptic diversity and we attempt to resolve seemingly contradictory data sets. Cicada songs act as species-specific premating barriers and have been used extensively to reveal hidden taxonomic diversity in morphologically similar species. The Palaearctic Cicadetta montana species complex is an excellent example where distinct song patterns have disclosed multiple recently described species. Indeed, two taxa turned out to be especially diverse in that they form a “complex within the complex”: the Cicadetta cerdaniensis song group (four species studied previously) and Cicadetta brevipennis (examined in details here). Based on acoustic, morphological, molecular, ecological and spatial data sampled throughout their broad European distribution, we find that Cicadetta brevipennis s. l. comprises five lineages. The most distinct lineage is identified as Cicadetta petryi Schumacher, 1924, which we re-assign to the species level. Cicadetta brevipennis litoralis Puissant & Hertach ssp. n. and Cicadetta brevipennis hippolaidica Hertach ssp. n. are new to science. The latter hybridizes with Cicadetta brevipennis brevipennis Fieber, 1876 at a zone inferred from intermediate song patterns. The fifth lineage requires additional investigation. The C. cerdaniensis and the C. brevipennis song groups exhibit characteristic, clearly distinct basic song patterns that act as reproductive barriers. However, they remain completely intermixed in the Bayesian and maximum likelihood COI and COII mitochondrial DNA phylogenies. The closest relative of each of the four cerdaniensis group species is a brevipennis group taxon. In our favoured scenario the phylogenetic pairs originated in common Pleistocene glacial refuges where the taxa speciated and experienced sporadic inter-group hybridization leading to extensive introgression and mitochondrial capture.

An egg clutch of the Tanzanian brevicipitid frog, Probreviceps m. macrodactylus, is described for the first time. The single clutch consists of 32 eggs and 21 infertile, jelly-filled capsules, tightly bound together by a thin, tough outer layer. The infertile capsules are concentrated toward the top of the clutch. The eggs were found in a dry season, and were perhaps likely to hatch at the onset of the short rains. The specific identification is confirmed through analysis of mitochondrial DNA sequence data. Eggs are yolky and large, about 8 mm in diameter (including capsule), suggesting that P. m. macrodactylus has a direct mode of development. The embryos observed were in an early developmental stage and showed the early division of the brain and anlagen of three pairs of visceral arches. Brevicipitid reproduction and development are poorly known, but a review of the existing literature and parsimonious optimization of simple characters onto a phylogenetic framework suggest that direct developme...

The Eastern Arc Mountains are renown in Africa for high concentrations of endemic species of animals and plants. Thirteen separate mountain blocks comprise the Eastern Arc, supporting around 3300 km2 of sub-montane, montane and upper montane forest, less than 30% of the estimated original forested area. At least 96 vertebrate species are endemic, split as follows: 10 mammal, 19 bird, 29 reptile and 38 amphibian species. This includes four endemic or nearly endemic species of primate – the Sanje Mangabey, the Iringa Red Colobus, the Mountain Galago and the new Kipunji monkey that forms its own monotypic genus. A further 71 vertebrate species are near-endemic. At least 800 vascular plant species are endemic, almost 10% of these being trees. These endemics include the majority of the species of African violet –Saintpaulia, a well-known flowering plant in Western households. An additional 32 species of bryophytes are also endemic. Many hundreds of invertebrates are also likely to be endemic, with data for butterflies, millipedes and dragonflies indicating potential trends in importance. Seventy-one of the endemic or near-endemic vertebrates are threatened by extinction (8 critical, 27 endangered, 36 vulnerable), with an additional seven wide ranging threatened species. Hundreds of plant species are also threatened. Most Eastern Arc endemics are closed-forest specialists and comprise taxa with an ancient history and those of more recent origin, including some possessing ancient affinities with taxa from West Africa, Madagascar, and even South America and Southeast Asia. Mountain block prioritisation for biodiversity conservation shows that Udzungwas, East Usambaras and Ulugurus are the most important blocks, with other important blocks being the Ngurus and West Usambaras. Rankings are correlated closely with the area of remaining forest. Most of the remaining forest is found within nearly 150 Government Forest Reserves, with 106 of these managed nationally for water catchment, biodiversity and soil conservation and where forest exploitation is not allowed. Outside these areas most forest has been cleared, except in small village burial/sacred sites, a few Village Forest Reserves, and inaccessible areas. In most Eastern Arc Mountains the local populations have not encroached beyond the reserve boundaries to develop farms, but forest resources within the boundaries are used for fuel and building materials and some forests are heavily degraded. Fire is also a problem as it enters and destroys forests during the dry seasons. The future of the biodiversity on the Eastern Arc Mountains is closely tied to management policies and capacity of the Tanzania Forestry and Beekeeping Division, Tanzania National Parks Authority, and Kenya Forest Department. Supporting these agencies in their mandated job is an essential conservation investment over the longer term.

The phylogenetic relationships of the African mongrel frog genus Nothophryne are poorly understood. We provide the first molecular assessment of the phylogenetic position of, and diversity within, this monotypic genus from across its range-the Afromontane regions of Malawi and Mozambique. Our analysis using a two-tiered phylogenetic approach allowed us to place the genus in Pyxicephalidae. Within the family, Nothophryne grouped with Tomopterna, a hypothesis judged significantly better than alternative hypotheses proposed based on morphology. Our analyses of populations across the range of Nothophryne suggest the presence of several cryptic species, at least one species per mountain. Formal recognition of these species is pending but there is a major conservation concern for these narrowly distributed populations in an area impacted by major habitat change. The phylogenetic tree of pyxicephalids is used to examine evolution of life history, ancestral habitat, and biogeography of this group.

The niche-filling process predicted by the "ecological opportunity" (EO) model is an often-invoked mechanism for generating exceptional diversity in island colonizers. Whether the same process governs lineage accumulation and trait disparity during continental colonization events is less clear. Here, we test this prediction by investigating the rate dynamics and trait evolution of one of Africa's most widespread amphibian colonizers, the true toads (Bufonidae). By reconstructing the most complete molecular phylogeny of African Bufonidae to date, we find that the diversification of lineages in Africa best conforms to a constant rate model throughout time and across subclades, with little support for EO. Evolutionary rates of life-history traits have similarly been constant over time. However, an analysis of generalists and specialists showed a shift toward higher speciation rates associated with habitat specialization. The overall lack of EO signal can be interpreted in a number of ways and we propose several explanations. Firstly, methodological issues might preclude the detection of EO. Secondly, colonizers might not experience true EO conditions and due to the size, ecological heterogeneity and age of landmasses, the diversification processes might be more complex. Thirdly, lower speciation rates of habitat generalists may have affected overall proliferation of lineages.

Background Peripatric speciation and peripheral isolation have uncertain importance in species accumulation, and are largely overshadowed by assumed dominance of allopatric modes of speciation. Understanding the role of different speciation mechanisms within biodiversity hotspots is central to understanding the generation of biological diversity. Here, we use a phylogeographic analysis of the spiny-throated reed frogs and examine sister pairings with unbalanced current distributional ranges for characteristics of peripatric speciation. We further investigate whether forest/grassland mosaic adapted species are more likely created through peripatric speciation due to instability of this habitat type. Results We reconstructed a multi-locus molecular phylogeny of spiny-throated reed frogs which we then combined with comparative morphologic data to delimit species and analyze historical demographic change; identifying three new species. Three potential peripatric speciation events were identified along with one case of allopatric speciation. Peripatric speciation is supported through uneven potential and realized distributions and uneven population size estimates based on field collections. An associated climate shift was observed in most potentially peripatric splits. Morphological variation was highest in sexually dimorphic traits such as body size and gular shape, but this variation was not limited to peripatric species pairs as hypothesized. The potentially allopatric species pair showed no niche shifts and equivalent effective population sizes, ruling out peripatry in that speciation event. Two major ecological niche shifts were recovered within this radiation, possibly as adaptations to occupy areas of grassland that became more prevalent in the last 5 million years. Restricted and fluctuating grassland mosaics within forests might promote peripatric speciation in the Eastern Arc Biodiversity Hotspot (EABH). Conclusions In our case study, peripatric speciation appears to be an important driver of diversity within the EABH biodiversity hotspot, implying it could be a significant speciation mechanism in highly fragmented ecosystems. Extensive peripatric speciation in this montane archipelago may explain the abundance of discrete lineages within the limited area of the EABH, as inferred in remote island archipelagos. Future phylogenetic studies incorporating demographic and spatial analyses will clarify the role of peripatric speciation in creating biodiversity hotspots. Electronic supplementary material The online version of this article (doi:10.1186/s12862-015-0384-3) contains supplementary material, which is available to authorized users.