Mareike Hirschfeld

Batrachochytrium dendrobatidis in amphibians of Cameroon, including first records for caecilians

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.

Evaluating mechanisms of diversification in a Guineo-Congolian tropical forest frog using demographic model selection

The accumulation of biodiversity in tropical forests can occur through multiple allopatric and parapatric models of diversification, including forest refugia, riverine barriers and ecological gradients. Considerable debate surrounds the major diversification process, particularly in the West African Lower Guinea forests, which contain a complex geographic arrangement of topographic features and historical refugia. We used genomic data to investigate alternative mechanisms of diversification in the Gaboon forest frog, Scotobleps gabonicus, by first identifying population structure and then performing demographic model selection and spatially explicit analyses. We found that a majority of population divergences are best explained by allopatric models consistent with the forest refugia hypothesis and involve divergence in isolation with subsequent expansion and gene flow. These population divergences occurred simultaneously and conform to predictions based on climatically stable regions inferred through ecological niche modelling. Although forest refugia played a prominent role in the intraspecific diversification of S. gabonicus, we also find evidence for potential interactions between landscape features and historical refugia, including major rivers and elevational barriers such as the Cameroonian Volcanic Line. We outline the advantages of using genomewide variation in a model‐testing framework to distinguish between alternative allopatric hypotheses, and the pitfalls of limited geographic and molecular sampling. Although phylogeographic patterns are often species‐specific and related to life‐history traits, additional comparative studies incorporating genomic data are necessary for separating shared historical processes from idiosyncratic responses to environmental, climatic and geological influences on diversification.

Dramatic Declines of Montane Frogs in a Central African Biodiversity Hotspot.

Amphibian populations are vanishing worldwide. Declines and extinctions of many populations have been attributed to chytridiomycosis, a disease induced by the pathogenic fungus Batrachochytrium dendrobatidis (Bd). In Africa, however, changes in amphibian assemblages were typically attributed to habitat change. We conducted a retrospective study utilizing field surveys from 2004–2012 of the anuran faunas on two mountains in western Cameroon, a hotspot of African amphibian diversity. The number of species detected was negatively influenced by year, habitat degradation, and elevation, and we detected a decline of certain species. Because another study in this region revealed an emergence of Bd in 2008, we screened additional recent field-collected samples and also pre-decline preserved museum specimens for the presence of Bd supporting emergence before 2008. When comparing the years before and after Bd detection, we found significantly diminished frog species richness and abundance on both mountains after Bd emergence. Our analyses suggest that this may be the first disease-driven community-level decline in anuran biodiversity in Central Africa. The disappearance of several species known to tolerate habitat degradation, and a trend of stronger declines at higher elevations, are consistent with Bd-induced declines in other regions. Not all species decreased; populations of some species remained constant, and others increased after the emergence of Bd. This variation might be explained by species-specific differences in infection probability. Increased habitat protection and Bd-mitigation strategies are needed for sustaining diverse amphibian communities such as those on Mt. Manengouba, which contains nearly half of Cameroon’s frog diversity.

What makes a successful species? Traits facilitating survival in altered tropical forests

BackgroundOngoing conversion, disturbance and fragmentation of tropical forests stress this ecosystem and cause the decline or disappearance of many species. Particular traits have been identified which indicate an increasing extinction risk of a species, but traits facilitating survival in altered habitats have mostly been neglected. Here we search for traits that make a species tolerant to disturbances, thus independent of pristine forests. We identify the fauna that have an increasing effect on the ecosystem and its functioning in our human-dominated landscapes.MethodsWe use a unique set of published data on the occurrences of 243 frog species in pristine and altered forests throughout the tropics. We established a forest dependency index with four levels, based on these occurrence data and applied Random Forest classification and binomial Generalized Linear Models to test whether species life history traits, ecological traits or range size influence the likelihood of a species to persist in disturbed habitats.ResultsOur results revealed that indirect developing species exhibiting a large range size and wide elevational distribution, being independent of streams, and inhabiting the leaf litter, cope best with modifications of their natural habitats.ConclusionThe traits identified in our study will likely persist in altered tropical forest systems and are comparable to those generally recognized for a low species extinction risk. Hence our findings will help to predict future frog communities in our human-dominated world.

The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae)

We describe and compare the tadpoles of nine Leptodactylodon species from Cameroon. The tadpoles of Leptodactylodon bicolor, L. mertensi, L. ovatus, L. perreti and L. ventrimarmoratus are herein reinvestigated, partly based on larger series than previously available. In addition we present first descriptions for the tadpoles of L. boulengeri, L. erythrogaster, L. ornatus, and L. cf. polyacanthus. The morphology of these exotrophic, lotic and neustonic tadpoles is discussed in comparison with other stream-dwelling tadpoles. Based on the assumed biology of these tadpoles, living in interstices of gravel or debris, the functioning of several special morphological features, in particular the funnel-mouth of Leptodactylodon tadpoles, are interpreted.

Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa

Frogs in the genus Amnirana (family Ranidae) are widely distributed across sub-Saharan Africa and present a model system for exploring the relationship between diversification and geography across the continent. Using multiple loci from the mitochondrial (16S) and nuclear genomes (DISP2, FICD, KIAA2013, REV3L), we generated a strongly supported species-level phylogeny that provides insights into the continental biogeography of African species of Amnirana, which form a monophyletic group within the genus. Species delimitation analyses suggest that there may be as many as seven additional species of Amnirana in Africa. The biogeographic history of Amnirana is marked by several dispersal and vicariance events, including dispersal from the Lower Guinean Forest into the Congo Basin. In addition, phylogeographic patterns within two widespread species, A. albolabris and A. galamensis, reveal undescribed cryptic diversity. Populations assigned to A. albolabris in western Africa are more closely related to A. fonensis and require recognition as a distinct species. Our analyses reveal that the Lower and Upper Guinean Forest regions served as important centers of interspecific and intraspecific diversifications for Amnirana.

Two new species of long-fingered frogs of the genus Cardioglossa (Anura: Arthroleptidae) from Central African rainforests

Abstract We describe two new frog species of Cardioglossa (Family Arthroleptidae) from Central Africa. The new species are found in the rainforests of western-central Democratic Republic of the Congo and the Republic of the Congo near the Gabonese border, respectively. We demonstrate that these species are morphologically and genetically distinct from each other and all other species of Cardioglossa. Both new species lack the dorsal hour-glass pattern present in many species of the genus, but they can be distinguished from each other and related species by distinctive colour patterns on their lateral surfaces and extremities. Both new species most closely resemble C. gratiosa, which occurs in the Atlantic coastal forests extending from Cameroon through Gabon. The new species can be differentiated from C. gratiosa by the absence of black transverse bars on all limbs or by distinctive lateral colouration. Analysis of mitochondrial ribosomal 16S DNA sequences reveals low to moderate levels (1.9–7.5%) of divergence between these new species and closely related species of Cardioglossa. The floodplains of the Congo and Ubangi Rivers may be important geographic barriers for many of these species. The occurrence of these two new lowland species in the Congo Basin reveals that the distribution and diversity of Cardioglossa in this region was underestimated. In addition, we elevate C. nigromaculata inornata to species-level status, based in part on newly available colour photographs from 1950 of specimens from the only known locality.

Sexual Dichromatism Drives Diversification Within a Major Radiation of African Amphibians

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 Afrobrachia 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 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 both micro- and macroevolutionary timescales.