Article

Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata: Colubridae) with descriptions of two new cryptic species from Central Africa

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Abstract

The genus Toxicodryas, historically included with the renowned Australasian cat-eyed snakes of the colubrid genus Boiga, currently includes two widespread species (T. blandingii and T. pulverulenta) in western, central, and eastern Africa. We leverage findings from a recent phylogenomic and historical demographic analysis of this genus (based on 2848-4471 Rad-seq loci from across the genome), with robust sampling from throughout the ranges of both species, to define two additional taxonomic units, with species boundaries corresponding to river barriers. Additional morphometric data from scores of examined museum specimens and literature records bolster the recognition of these two new cryptic species. We hypothesize that T. blandingii occurs west of the confluence of the Congo and Ubangi rivers, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of ventral scale counts in both sexes, additional significant differences in several scale counts, and lower venom toxicity. Toxicodryas pulverulenta occurs west of the Niger Delta in West Africa, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of subcaudal scale counts in both sexes. A review of published information regarding morphological variation, ecology, natural history, habitat, and venom is summarized for these four Toxicodryas species.

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Emerald Labyrinth is a scientist and adventurer’s chronicle of years exploring the rainforests of sub-Saharan Africa. The richly varied habitats of the Democratic Republic of the Congo offer a wealth of animal, plant, chemical, and medical discoveries. But the country also has a deeply troubled colonial past and a complicated political present. Author Eli Greenbaum is a leading expert in sub-Saharan herpetology—snakes, lizards, and frogs—who brings a sense of wonder to the question of how science works in the twenty-first century. Along the way he comes face to face with spitting cobras, silverback mountain gorillas, wild elephants, and the teenaged armies of AK-47-toting fighters engaged in the continent’s longest-running war. As a bellwether of the climate and biodiversity crises now facing the planet, the Congo holds the key to our planet’s future. Writing in the tradition of books like The Lost City of Z, Greenbaum seeks out the creatures struggling to survive in a war-torn, environmentally threatened country. Emerald Labyrinth is an extraordinary book about the enormous challenges and hard-won satisfactions of doing science in one of the least known, least hospitable places on earth.
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Aim: To delineate bioregions in tropical Africa and determine whether different plant growth forms (trees, terrestrial herbs, lianas and shrubs) display the same pattern of regionalization, diversity and endemism as the whole flora. Location: Tropical Africa (excl. Madagascar), from 20° N to 25° S. Taxon: Vascular plants. Methods: Analyses were based on occurrences of 24,719 vascular plant species distributed across tropical Africa extracted from the RAINBIO database. The majority of species (93%) were classified into four growth forms: terrestrial herbs, trees, shrubs and lianas. Biogeographical regions (bioregions) were delimited using a bipartite network clustering approach on the whole dataset and then separately for each growth form. Relationships among bioregions were investigated using non-metric multidimensional scaling ordination, flora nestedness and endemism patterns. Results: Analyses of the whole dataset identified 16 bioregions and 11 transition zones. These were congruent with most of the currently recognized phytogeographical classifications, and also highlighted previously under-recognized bioregions. Bioregion endemism rates were lower and species richness higher when compared to estimates from the White/Association pour l'Etude Taxonomique de la Flore d'Afrique Tropicale (AETFAT) classification. Analysed separately, plant growth forms showed contrasting geographical patterns. Bioregionalization was better resolved for closed forest types using trees and lianas and for open vegetation types using terrestrial herbs, while shrubs showed good discriminative power in all vegetation types. Main conclusions: We show that distribution patterns based on solely trees are not sufficient to define floristic bioregions in tropical Africa. Analyses of spatial patterns using different growth forms are complementary, likely reflecting different evolutionary processes and ecological relationships. The contribution of growth forms to delimit geographical floristic patterns across tropical Africa is of critical importance for land use planning and management, and for selecting priority conservation areas.
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Five problematic cases regarding the presence of reptile taxa on the island of São Tomé (Gulf of Guinea) are reviewed. These cases deal with 1) the past presence of crocodilians in the island, 2) the putative type locality of Dipsas pulverulenta Fischer, 1856, 3) the dubious presence of a mamba (genus Dendroaspis) on the island, 4) an erroneous reference to the presence of Gastropyxis smaragdina, and 5) two specimens (Psammophis lineatus and Psammophis elegans) from Carl Weiss collection, currently deposited in the Zoological Museum of Hamburg and wrongly labelled as being from São Tomé island. A provisional checklist to the terrestrial herpetofauna of the three oceanic islands of the Gulf of Guinea—São Tomé, Príncipe and Annobon—is presented in order to clarify the current numbers of the endemics of each island, as well as to review and present current research.
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The Cobra-Preta (black snake in Portuguese) of Sao Tomé Island in the Gulf of Guinea has historically been referred to as Naja (Boulengerina) melanoleuca (Squamata: Elapidae). Its presence on the island has been traditionally explained as an introduction from the mainland by Portuguese settlers, supposedly to control the rat population. This explanation has been widely accepted by local authorities and even international conservation agencies. The taxonomic identity of this snake has remained undisputed by all taxonomists who have published about it, with the exception of L. Capocaccia in 1961. Arguments supporting the human introduction hypothesis are weak and are contradicted by historical, morphological and molecular data. Further, the biogeographic history of the Gulf of Guinea oceanic islands and recent insights on the taxonomic identity and evolutionary history of other taxonomic groups occurring there suggest that the Cobra-Preta, in fact, represents a distinct lineage of the melanoleuca group, endemic to São Tomé. We here describe the Cobra Preta as a new species. The new species differs from N. (B.) melanoleuca, its sister species, by a distinct coloration ventral pattern and the type of contact of the sublingual scales. Data on the toxicology, distribution, ecology, folklore and conservation status of the new species are presented.
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We present new Gabonese locality records, ecological data or unpublished museum material for Kinixys erosa (Testudinidae), Cycloderma aubryi, Trionyx triunguis (Trionychidae), Agama picticauda (Agamidae), Chamaeleo dilepis (Chamaeleonidae), Calabaria reinhardtii (Boidae), Grayia ornata, Thrasops jacksonii, Toxicodryas blandingii (Colubridae), Naja a. annulata and N. melanoleuca (Elapidae), Psammophis cf. phillipsii (Lamprophiidae), Natriciteres fuliginoides and N. olivacea (Natricidae). We refer all Gabonese records of Gonionotophis capensis to G. savorgnani (Lamprophiidae). We provide the first report on Albert Schweitzer’s snake collection at Lambaréné Hospital Museum, part of the oldest natural history collection settled in Gabon. We add one new snake species record to Woleu-Ntem Province and one reptile species each to Minkébé and Pongara national parks.
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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 modeling. Though 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 genome-wide 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. This article is protected by copyright. All rights reserved.
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We present new Gabonese locality records for Gerrhosaurus nigrolineatus (Gerrhosauridae), Trachylepis maculilabris (Scincidae), Varanus ornatus (Varanidae), Calabaria reinhardtii (Boidae), Dipsadoboa viridis, Toxicodryas blandingii (Colubridae), Naja melanoleuca and N. nigricollis (Elapidae), Python sebae (Pythonidae), Atheris squamigera (orange morph), Bitis arietans, B. gabonica and B. nasicornis (Viperidae). We add two, one, two and one species to Moyen-Ogooué, Ngounié, Nyanga and Woleu-Ntem provinces’ reptile lists, respectively. We newly record two, one and one snake species for Mwagna National Park, Waka National Park and Bas Ogooué Ramsar Site, respectively. We report predation cases by a Naja melanoleuca on a Toxicodryas blandingii and by a Seba’s python on a domestic dog (Mammalia: Canidae: Canis lupus familiaris), respectively.
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We present new Gabonese locality records, ecological data or unpublished museum material for Crocodylus niloticus (Crocodylidae), Trionyx triunguis (Trionychidae), Agama lebretoni (Agamidae), Hemidactylus fasciatus and H. mabouia (Gekkonidae), Gastropholis echinata (Lacertidae), Trachylepis albilabris (Scincidae), Afrotyphlops angolensis (Typhlopidae), Dipsadoboa viridis, Hapsidophrys smaragdinus, Toxicodryas pulverulenta (Colubridae), Naja melanoleuca (Elapidae), Lamprophis olivaceus, Psammophis cf. phillipsii (Lamprophiidae), Natriciteres fuliginoides (Natricidae), Causus lichtensteinii and C. maculatus (Viperidae). We document predation cases by Hapsidophrys smaragdinus on Hemidactylus mabouia and Trachylepis albilabris, by Naja melanoleuca on Sclerophrys regularis (Anura: Bufonidae) and by Psammophis cf. phillipsii on Phrynobatrachus auritus (Anura: Phrynobatrachidae), and consumption of Arius latiscutatus (Siluriformes: Ariidae) and Tragelaphus spekii (Cetartiodactyla: Bovidae) by Crocodylus niloticus. We add one, two and one snake species, respectively, to Estuaire, Moyen-Ogooué and Nyanga provinces’ reptile lists. We add four reptile species to the list for Wonga-Wongué Presidential Reserve. We refer all records of Agama agama in Gabon to A. picticauda.
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The savannah and tropical forest biomes of Africa have a long history of expansion and contraction, and the recent and rapid spread of dry savannah habitats has influenced the spatial and temporal diversification of vertebrate taxa across this region. We used a combination of species tree and phylogeographic methods to describe the spatio-temporal changes through time and across space (= species diffusion) in a clade of seven West African lizard species in the Agama agama species group. A Bayesian species tree diffusion approach was used to compare the relative rates at which species ranges changed across the landscape. We found that some species have high diffusion rates characterized by significant movement in their range location and minor changes to their overall range size, whereas other species show little movement in their range centre with an exponential increase in range size. This discrepancy between the rates that range locations shift versus change in their relative area could be linked to populations tracking their preferred habitats through time. A continuous Bayesian phylogeography approach using a relaxed random walk model was used to estimate the timing and rate of population size change and geographic diffusion in A. picticauda, the single species in the group with an extensive African distribution from Mauritania to Ethiopia. The mean dispersal rate of A. picticauda increased dramatically throughout the Pleistocene, and a Bayesian skyride analysis supports exponential population growth over this same time period. A comparison of genetic diversity across different loci and species suggests that A. lebretoni experienced a mitochondrial selective sweep that has caused a deficit of variation at this locus in relation to nuclear loci.
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Background: With over 3,500 species encompassing a diverse range of morphologies and ecologies, snakes make up 36% of squamate diversity. Despite several attempts at estimating higher-level snake relationships and numerous assessments of generic- or species-level phylogenies, a large-scale species-level phylogeny solely focusing on snakes has not been completed. Here, we provide the largest-yet estimate of the snake tree of life using maximum likelihood on a supermatrix of 1745 taxa (1652 snake species + 7 outgroup taxa) and 9,523 base pairs from 10 loci (5 nuclear, 5 mitochondrial), including previously unsequenced genera (2) and species (61). Results: Increased taxon sampling resulted in a phylogeny with a new higher-level topology and corroborate many lower-level relationships, strengthened by high nodal support values (> 85%) down to the species level (73.69% of nodes). Although the majority of families and subfamilies were strongly supported as monophyletic with > 88% support values, some families and numerous genera were paraphyletic, primarily due to limited taxon and loci sampling leading to a sparse supermatrix and minimal sequence overlap between some closely-related taxa. With all rogue taxa and incertae sedis species eliminated, higher-level relationships and support values remained relatively unchanged, except in five problematic clades. Conclusion: Our analyses resulted in new topologies at higher- and lower-levels; resolved several previous topological issues; established novel paraphyletic affiliations; designated a new subfamily, Ahaetuliinae, for the genera Ahaetulla, Chrysopelea, Dendrelaphis, and Dryophiops; and appointed Hemerophis (Coluber) zebrinus to a new genus, Mopanveldophis. Although we provide insight into some distinguished problematic nodes, at the deeper phylogenetic scale, resolution of these nodes may require sampling of more slowly-evolving nuclear genes.
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Abstract The snake fauna of the Comoé National Park, lvory Coast: Supplements and prospect. We report on seven additional species and on further biological data of the hitherto known snake fauna of the Comoe National Park (CNP), Ivory Coast (West Africa). We have recorded 44 snake species within the study area so far. The occurrence of 25 further species, which appears to be possible, would attribute the CNP to hold one of Africa's most diverse snake communities. We analysed the snake fauna with respect to habitat selection, mode of living, and geographical distribution of the species, thereby comparing the composition of the snake community with other vertebrate taxa (Anura, Sauria, Mammalia) of the Comoé National Park. Key words: Serpentes; faunistics; Ivory Coast; Guinea savannah; habitats; distribution; diversity. Resume La faune des serpents du Parc National de la Comoé, Côte d’IÌvoire: Supplements et perspective. Nous rapportons sept autres espèces des serpents du Parc National de la Comoé (CNP), Côte d'Ivoire (Afrique de l'Ouest). Des informations sur la biologie de quelques espèces sont ajoutées. Actuellement, 44 espèces de serpents sont connues pour le CNP, mais l'existence d'autres 25 espèces est probable. Cela pourrait etablir le CNP comme une des regions avec la plus grande diversité de serpents de taute l 'Afrique. Nous analysons la faune des serpents de la CNP selon l 'habitat, la mode de vie et la distribution des espèces en Afrique et nous la comparons avec la faune des vertèbres de la region (Anura, Sauria, Mammalia). Mots-cles: Serpentes; communitée; Côte d'Ivoire; savane guinéenne; habitats; distribution géographique; diversité. Zusammenfassung Wir berichten von weiteren sieben Schlangennachweisen aus dem Comoe-Nationa!lpark (CNP), Elfenbeinküste (West-Afrika) und geben zusätzliche Informationen zur Biologie einiger Arten. Wir wiesen bislang 44 Schlangenarten im Untersuchungsgebiet nach, wobei das Vorkommen weiterer 25 Arten möglich erscheint. Der CNP wäre damit eine der an Schlangen­ arten vielfältigsten Regionen Afrikas. Wir analysieren die Schlangenfauna des CNP nach Habitat, Lebensweise und afrikanischer Verbreitung der Arten und stellen einen Vergleich mit der Fauna anderer Wirbeltiergruppen (Anura, Sauria, Mammalia) des Gebiets an. Schlagwörter: Serpentes; Faunistik ; Elfenbeinküste; Guinea-Savanne; Habitate; geogra­phische Verbreitung; Diversität.
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Aim The aim of this study was to understand the origin of the forest flora cur-rently found in the Dahomey Gap (DG), a 200-km-wide forest-savanna mosaicseparating the West African and Central African rain forest blocks. Morespecifically, using a widespread rain forest tree species, Distemonanthus ben-thamianus (Fabaceae), we will test the hypothesis that the DG populations areremnants of a population dating back from the African Humid Period of theHolocene, when West African and Central African rain forests were supposedly connected. Location Tropical forests of Upper Guinea (West Africa) and Lower Guinea(Atlantic Central Africa) and the forest-savanna mosaic of the DG extendingfrom eastern Ghana to Benin. Methods Four hundred and twenty-nine D. benthamianus samples from Westto Central Africa were genotyped with 11 nuclear microsatellite markers. Afterdetecting geographically coherent gene pools, their genetic diversity and differ-entiation were estimated and their demographic histories were inferred usingcoalescent simulations and approximate Bayesian computation (ABC) tests. Results Five parapatric gene pools were identified: three in Lower Guinea, onein Upper Guinea and one in the DG. ABC tests indicate that the DG gene poolprobably originates from the admixture of adjacent Upper and Lower Guineangene pools, with a higher contribution from Upper Guinea, at a timeframeconsistent with the early Holocene (around 13– 7 ka). The lower genetic diver-sity documented in the DG could result from a founder effect and/or from ademographic decline consistent with the Holocene climatic pejoration docu-mented around 3 ka. Main conclusions This phylogeographical study inferring the history of theDG populations of D. benthamianus is consistent with palaeovegetation datathat suggest that the forest flora of the DG might be essentially relicts of theearly Holocene period when the Guineo-Congolian forest reached its maximumgeographical distribution.
Article
We conducted a taxonomic revision of the arboreal snakes of the Boiga ceylonensis-group, all inhabiting the Indian subcontinent. Based on 15 characters, recorded from more than 100 specimens, representing eight recognized taxa, we conducted a multivariate morphometric analysis to redefine the taxonomic boundaries. We provide redescriptions of the nominate taxa in this group based on re-examination of the name-bearing types. The types of B. ceylonensis, B. beddomei, B. barnesii, B. dightoni, B. nuchalis and B. andamanensis are redescribed, where appropriate lectotypes are selected, illustrated and their nomenclature, synonymy and distribution are discussed. Status of the nominate taxon Boiga ceylonensis dakhunensis Deraniyagala, 1955 is fixed as an objective junior synonym of Dipsas nuchalis Günther, 1875. An updated key to the species of this group is presented.
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Comparative phylogeographic studies often support shared divergence times for co‐distributed species with similar life histories and habitat specializations. During the late Holocene, West Africa experienced aridification and the turnover of rain forest habitats into savannas. These fragmented rain forests harbor impressive numbers of endemic and threatened species. In this setting, populations of co‐distributed rain forest species are expected to have diverged simultaneously, whereas divergence events for species adapted to savanna and forest‐edge habitats should be absent or idiosyncratic. We conducted a Bayesian analysis of shared evolutionary events to test models of population divergence for 20 species of anurans (frogs) and squamates (lizards and snakes) that are distributed across the Dahomey Gap, a climate change induced savannah barrier responsible for fragmenting previously contiguous rain forests of Ghana into two regions: the Togo‐Volta Hills and the Southwestern Forests. A model of asynchronous diversification is supported for anurans and squamates, suggesting that drivers of diversification are not specifically related to ecological and life history associations with habitat types. Instead, the wide variability of genetic divergence histories exhibited by these species suggests that biodiversity in this region has been shaped by diversification events that extend beyond the Holocene. Comparisons of the genealogical divergence index (gdi), a measure of the genetic divergence between populations due to the combined effects of genetic isolation and gene flow, illustrate that these populations represent a broad sampling of the speciation continuum. This article is protected by copyright. All rights reserved
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In March and April 2018 we surveyed amphibians and reptiles in two Proposed Protected Areas (PPAs) in Liberia. In the Krahn-Bassa Proposed Protected Area (KBPPA) in eastern Liberia 36 species of amphibians and 13 species of reptiles were recorded. In the Foya Proposed Protected Area (FPPA) in western Liberia 39 species of amphibians and 10 species of reptiles were recorded. The encountered herpetological communities in both sites were typical for West African rainforests. However, some species indicated disturbances, in particular at the edges of the study areas, the surrounding villages and plantations, and old artisanal gold mining sites within forests. Of particular conservation interest was the discovery of a high percentage of typical rainforest specialists with ranges restricted to the western part of the Upper Guinea rainforest biodiversity hotspot. Outstanding discoveries in KBPPA were two new species of puddle frogs, and the first country record for the arboreal, parachuting lizard Holaspisguentheri . Remarkable records in FPPA comprise a new species of stiletto snake, a new puddle frog and records of various frog species typically breeding in undisturbed rainforest streams, such as Odontobatrachusnatator and Conrauaalleni . Both study areas comprise an important proportion of the remaining rainforests in the Upper Guinea forest zone. The new discoveries indicate that within this biogeographic area, southeastern and western Liberian rainforest may still hold various undiscovered species and species of conservation concern. Further surveys in KBPPA and FPPA and nearby forests should clarify the distribution and conservation status of the new taxa. This study also emphasizes that the western part of the Liberian forests comprise at least partly a herpetofauna which differs from that of the East of the country. The recorded threatened amphibian species are all specialized on relatively undisturbed rainforests and they all have only small geographic ranges. The remaining parts of undisturbed or little disturbed forests thus have high importance for the long-term survival of these species. In conclusion the study areas have a high conservation potential and should be urgently protected from any further forest loss degradation and uncontrolled hunting.
Article
We report on reptilian specimens collected in southern Sudan (currently the Republic of South Sudan) in 1978 and stored at the Zoologisches Forschungsmuseum Alexander Koenig, Bonn. Six species (one lizard, Leptosiaphos kili-mensis, and five snakes, Hapsidophrys lineatus, Thrasops jacksoni, Toxicodryas pulverulenta, Amblyodipsas unicolor, Atheris squamigera) are documented as new records for the fauna of South Sudan and are discussed in a biogeographi-cal context.
Book
East Africa is home to a remarkable assemblage of reptiles, from crocodiles and chameleons to turtles and tortoises, lizards, worm-lizards, and a stunning array of snakes. The region is a true herpetological hot-spot. This second edition, a fully revised version of the classic field guide to the region's reptiles, explores the full diversity of these animals. With updated text (well over 100 species have had their generic names changes since publication of the first edition), coverage of an additional 60 species, detailed maps and more than 600 new photographs, Field Guide to East African Reptiles includes every one of the 492 species in the region. All are described and mapped, with virtually every species accompanied by at least one colour photograph. Comprehensive and definitive, Field Guide to East African Reptiles is an essential tool for all naturalists, conservationists, educators, field workers, medical personnel and students in the region.
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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.