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Frogs of the genus Platypelis are known to have their center of species richness in the mountain massifs of northern Madagascar. We here formally describe a new species of Platypelis from this region. Platypelis saikamavo sp. nov. has been known for over 20 years but its taxonomic identity has only recently been robustly clarified as a result of ongoing taxonomic revisions of other Platypelis species. The new species is known from rainforest on the Tsaratanàna Massif and Bemanevika, at elevations of 1000-1530 m above sea level. It is characterized by an isolated phylogenetic position and is genetically highly divergent, with an uncorrected pairwise distance ≥4.5% in the mitochondrial 16S rRNA gene to all other nominal species of Platypelis. It has a unique combination of morphological and chromatic character states such as a relatively large body size of 28-31 mm in snout-vent length, distinct superciliary tubercles, and bright yellow color around the forelimb insertion and on the posterior abdomen where this color extends in the form of flash marks onto the inguinal region.

The Lygodactylus tolampyae complex includes several deep genetic lineages of small diurnal geckos from the West and North West of Madagascar whose taxonomy is largely unsolved. We sequenced DNA fragments of one mitochondrial and four nuclear-encoded genes for up to 70 samples across the entire known range of these geckos. We find as many as 11 mitochondrial lineages differentiated by >4% pairwise distances in the 16S rRNA gene fragment, with >9% pairwise distance for the majority of lineage comparisons. Many of these lineages were concordantly differentiated in all of the nuclear-encoded genes without any haplotype sharing, despite the syntopic occurrence of some of them. We therefore hypothesize that the complex contains seven candidate species, but a comprehensive taxonomic resolution is complicated by various hindrances. These include incomplete sampling, with two lineages each known only from a single specimen, and one further lineage with no voucher specimens available for examination. Further hurdles are the probably lost holotype of L. tolampyae and its imprecise type locality, as well as the apparent lack of any morphological differentiation between the majority of the genetic lineages. Based on a survey of historical literature and the travel routes of the original collector, A. Grandidier, we conclude that the provenance of the holotype of L. tolampyae is likely in the wider Morondava area in the West and assign the sole candidate species from this area to this name. We then proceed to describe three species that represent separate genetic lineages for all markers studied: Lygodactylus morii sp. nov., a species common in Ankarafantsika National Park and several nearby sites in the North West; L. herilalai sp. nov., a species occurring in close syntopy with L. morii in Ankarafantsika without any signal of genetic admixture; and L. schwitzeri sp. nov. from Sahamalaza Peninsula in the North West. This leaves three more lineages without a name and with the need to gather additional samples, two from Namoroka National Park and one from other sites in the North West. We confirm the L. tolampyae complex to be an apparently rare example of truly cryptic reptile species in Madagascar, where even detailed morphological examination does not reveal morphological differences among lineages that are clearly evolutionarily independent and require recognition as distinct species due to their co-occurrence without admixture.

Many studies on reptiles have been conducted across Madagascar but some areas are poorly known in terms of the diversity of reptiles such as the Bobaomby Complex in the northern tip of Madagascar. In February and March 2018, we conducted a biodiversity survey within five sites. This biological survey is to collect scientific information for helping new protected creations. Three main methods were used including pitfall trap, visual and acoustic searching along the transect and refuge examination. In total, we recorded 42 species including 5 chameleons, 8 skinks, 11 geckos, 16 snakes and 2 blinds snake species. All recorded species are endemic to Madagascar except Hemidactylus frenatus , Ebenavia inunguis and Phelsuma abbotti . Rare species known only from a few specimens have been recorded in the Bobaomby Complex: Heteroliodon fohy , Pseudoxyrhopus ambreensis , and Madascincus arenicola . Thirteen species are classified as threatened on the IUCN Red List, of which three are Critically Endangered: Paracontias minimus , Madascincus arenicola , and Paroedura lohatsara ; three are Endangered: Heteroliodon fohy , Lycodryas inopinae , and Phisalixella variabilis ; and seven are Vulnerable: Brookesia ebenaui , Furcifer petteri , Blaesodactylus boivini , Uroplatus ebenaui , Uroplatus henkeli , Liophidium therezieni and Flexiseps ardouini . Our results reveal the importance of the Bobaomby Complex for conserving reptile diversity and highlight the need to protect it.

Speciation rates vary substantially across the tree of life. These rates should be linked to the rate at which population structure forms if a continuum between micro and macroevolutionary patterns exists. Previous studies examining the link between speciation rates and the degree of population formation in clades have been shown to be either correlated or uncorrelated depending on the group, but no study has yet examined the relationship between speciation rates and population structure in a young group that is constrained spatially to a single-island system. We examine this correlation in 109 gemsnakes (Pseudoxyrhophiidae) endemic to Madagascar and originating in the early Miocene, which helps control for extinction variation across time and space. We find no relationship between rates of speciation and the formation rates of population structure over space in 33 species of gemsnakes. Rates of speciation show low variation, yet population structure varies widely across species, indicating that speciation rates and population structure are disconnected. We suspect this is largely due to the persistence of some lineages not susceptible to extinction. Importantly, we discuss how delimiting populations versus species may contribute to problems understanding the continuum between shallow and deep evolutionary processes., Competing Interests: We have no competing interests., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Populations of phytotelmic frogs from northern Madagascar assigned to Guibemantis (Pandanusicola) pulcher are known to differ genetically from populations further south in the eastern rainforest belt of the island, but to date, their status has not been analyzed in depth. We combined molecular genetic data with an examination of color pattern to clarify the taxonomy of these frogs. DNA sequences of both mitochondrial and nuclear-encoded genes were consistently differentiated between the northern populations and those occurring further south. Uncorrected pairwise distance in the 16S rRNA gene was 3.7‒4.3% and thus at a level usually characterizing distinct frog species in Madagascar. Furthermore, the northern specimens were characterized by more and smaller purplish-brown spots on their green dorsal surface, and a less distinct brown patch on the flanks. Although fully conclusive evidence for the species status of the northern lineage from bioacoustic differences, sympatric occurrence or narrow hybrid zone is currently lacking, such species-level distinctness is currently the most likely hypothesis. We therefore name the northern populations as Guibemantis (Pandanusicola) pulcherrimus sp. nov. The new species is known from Makira (type locality) and Bemanevika, and specimens morphologically assignable to this taxon have also been recorded from Masoala, Marojejy and Anjanaharibe-Sud.

A fundamental assumption of evolutionary biology is that phylogeny follows a bifurcating process. However, hybrid speciation and introgression are becoming more widely documented in many groups. Hybrid inference studies have been historically limited to small sets of taxa, while exploration of the prevalence and trends of reticulation at deep time scales remains unexplored. We study the evolutionary history of an adaptive radiation of 109 gemsnakes in Madagascar (Pseudoxyrhophiinae) to identify potential instances of introgression. Using several network inference methods, we find 12 reticulation events within the 22-million-year evolutionary history of gemsnakes, producing 28% of the diversity for the group, including one reticulation that resulted in the diversification of an 18 species radiation. These reticulations are found at nodes with high gene tree discordance and occurred among parental lineages distributed along a north-south axis that share similar ecologies. Younger hybrids occupy intermediate contact zones between the parent lineages showing that post-speciation dispersal in this group has not eroded the spatial signatures of introgression. Reticulations accumulated consistently over time, despite drops in overall speciation rates during the Pleistocene. This suggests that while bifurcating speciation rates may decline as the result of species accumulation and environmental change, speciation by hybridization may be more robust to these processes., (© The Author(s) 2023. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE). All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

How environmental factors shape patterns of biotic diversity in tropical ecosystems is an active field of research, but studies examining the possibility of ecological speciation in terrestrial tropical ecosystems are scarce. We use the isolated rainforest herpetofauna on the Montagne d'Ambre (Amber Mountain) massif in northern Madagascar as a model to explore elevational divergence at the level of populations and communities. Based on intensive sampling and DNA barcoding of amphibians and reptiles along a transect ranging from ca. 470-1470 m above sea level (a.s.l.), we assessed a main peak in species richness at an elevation of ca. 1000 m a.s.l. with 41 species. The proportion of local endemics was highest (about 1/3) at elevations >1100 m a.s.l. Two species of chameleons ( Brookesia tuberculata, Calumma linotum ) and two species of frogs ( Mantidactylus bellyi, M. ambony ) studied in depth by newly developed microsatellite markers showed genetic divergence up the slope of the mountain, some quite strong, others very weak, but in each case with genetic breaks between 1100 and 1270 m a.s.l. Genetic clusters were found in transect sections significantly differing in bioclimate and herpetological community composition. A decrease in body size was detected in several species with increasing elevation. The studied rainforest amphibians and reptiles show concordant population genetic differentiation across elevation along with morphological and niche differentiation. Whether this parapatric or microallopatric differentiation will suffice for the completion of speciation is, however, unclear, and available phylogeographic evidence rather suggests that a complex interplay between ecological and allopatric divergence processes is involved in generating the extraordinary species diversity of Madagascar's biota. Our study reveals concordant patterns of diversification among main elevational bands, but suggests that these adaptational processes are only part of the complex of processes leading to species formation, among which geographical isolation is probably also important., Competing Interests: The authors declare no conflicts of interest., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The highly diverse snake superfamily Elapoidea is considered to be a classic example of ancient, rapid radiation. Such radiations are challenging to fully resolve phylogenetically, with the highly diverse Elapoidea a case in point. Previous attempts at inferring a phylogeny of elapoids produced highly incongruent estimates of their evolutionary relationships, often with very low statistical support. We sought to resolve this situation by sequencing over 4,500 ultraconserved element loci from multiple representatives of every elapoid family/subfamily level taxon and inferring their phylogenetic relationships with multiple methods. Concatenation and multispecies coalescent based species trees yielded largely congruent and well-supported topologies. Hypotheses of a hard polytomy were not retained for any deep branches. Our phylogenies recovered Cyclocoridae and Elapidae as diverging early within Elapoidea. The Afro-Malagasy radiation of elapoid snakes, classified as multiple subfamilies of an inclusive Lamprophiidae by some earlier authors, was found to be monophyletic in all analyses. The genus Micrelaps was consistently recovered as sister to Lamprophiidae. We establish a new family, Micrelapidae fam. nov., for Micrelaps and assign Brachyophis to this family based on cranial osteological synapomorphy. We estimate that Elapoidea originated in the early Eocene and rapidly diversified into all the major lineages during this epoch. Ecological opportunities presented by the post-Cretaceous-Paleogene mass extinction event may have promoted the explosive radiation of elapoid snakes., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Author(s). Published by Elsevier Inc. All rights reserved.)

In the processes that give rise to new species, changes first occur at the population level. But with the continuous nature of the divergence process, change in biological properties delimiting the shift from "individuals of divergent populations" towards "individuals of distinct species", as well as abiotic factors driving the change, remain largely ambivalent. Here we study diversification processes at the population level in a semi-aquatic frog, Mantidactylus (Brygoomantis) bellyi, across the diverse vegetation types of Montagne d'Ambre National Park (MANP), Madagascar. Genetic diversity was assessed with seven newly developed microsatellite markers as well as mitochondrial DNA sequences and concordance with patterns of ecological, morphological, and bioacoustic divergence evaluated. We found M. bellyi lacking mitochondrial differentiation within MANP, while microsatellite datasets partitioned them into three highly differentiated, geographically separated subpopulations (with indications for up to five subpopulations). The molecular grouping-primarily clustering individuals by geographic proximity-was coincident with differences in mean depth and width of waters, suggesting a possible role of fluvial characteristics in genetic exchange in this stream-breeding species. Genetic clustering not consistent with differences in call properties, except for dominant call frequencies under the two-subpopulations model. Morphological divergence was mostly consistent with the genetic clustering; subpopulations strongly differed by their snout-vent length, with individuals from high-elevation subpopulations smaller than those from populations below 1000 m above sea level. These results exemplify how mountains and environmental conditions might primarily shape genetic and morphological divergence in frog populations, without strongly affecting their calls., Competing Interests: The authors have declared that no competing interests exist.

The genus Scaphiophryne (Anura: Microhylidae) contains at least nine species that, together with their probable sister genus Paradoxophyla, form the Madagascar-endemic subfamily Scaphiophryninae. Scaphiophryne are robust burrowing frogs with explosive breeding behavior and are characterized by a unique larval morphology, being intermediate between that of filter-feeding and generalized tadpole types. Based on tadpole morphology, the nominal subgenus Scaphiophryne can be distinguished from the subgenus Pseudohemisus, a clade occurring in arid biomes with a largely unsolved taxonomy and nomenclature. Here we combine a comprehensive molecular phylogeny of all scaphiophrynines based on five mitochondrial and two nuclear genes with a re-examination of the name-bearing type specimens of the seven species-level nomina assignable to the subgenus Pseudohemisus. Our molecular phylogeny supports monophyly of the two subgenera and assigns the enigmatic S. matsoko to the subgenus Scaphiophryne. In this subgenus, our analysis confirms all currently accepted species as independent evolutionary lineages based on concordant differentiation in mitochondrial and nuclear genes, and provides evidence for a combination of low mitochondrial divergence, distinct nuclear divergence, and high nuclear heterozygosity. Yet, contrary to previous hypotheses, we do not find evidence of ongoing hybridization. In the subgenus Pseudohemisus we find support for three clearly delimited species-level lineages, one of which contains two deep conspecific lineages that warrant further study. To stabilize the taxonomy in this subgenus, and considering the very poor state of preservation of several type series and the heterogeneity of some of them, we here designate lectotypes for the nomina: Hemisus obscurus Grandidier, 1872; Calophrynus calcaratus Mocquard, 1895; Pseudohemisus verrucosus Angel, 1930; Pseudohemisus longimanus Angel, 1930; and Pseudohemisus longimanus var. melanopleura Angel, 1934. Taxonomically, we accept Scaphiophryne brevis (Boulenger, 1896) as a well-defined species; restrict S. calcarata (Mocquard, 1995) to the southern lineage of the S. calcarata complex; resurrect Hemisus obscurus Grandidier, 1872 (previously considered to be a dubious name or nomen inquirendum) as Scaphiophryne obscura to refer to the western and north-western lineage of the S. calcarata complex; consider Pseudohemisus verrucosus Angel, 1930 as junior synonym of Scaphiophryne brevis; and consider Pseudohemisus longimanus Angel, 1930 and Pseudohemisus longimanus var. melanopleura Angel, 1934 as junior synonyms of Scaphiophryne obscura. Finally, to accelerate taxonomic progress, we (i) outline a set of criteria by which such old and ambiguous nomina can be applied to known lineages, (ii) suggest that the International Code of Zoological Nomenclature should slightly broaden the conditions under which neotypes are to be designated, and (iii) propose to redouble discussions on the introduction of the concept of the epitype to zoology.

The northern part of Madagascar is well known for its high species diversity and endemism. Exceptional species richness is related to the existence of large forest blocks and mountain complexes. These areas shelter a diverse variety of habitats occupied by a wide diversity of species, including leaf-tailed geckos of the genus Uroplatus. Based on morphological and molecular evidence, we here formally name two evolutionary lineages as new species that previously had been considered as candidate species (Uroplatus spp. Ca3 and Ca4), both small-sized species of the Uroplatus ebenaui group. Genetically, both new species are related to U. finiavana with a genetic divergence (uncorrected pairwise distance) of 10.3-12.8% in a fragment of the mitochondrial 16S rRNA gene, and separated from each other by 10.3-10.7%. Uroplatus fangorn sp. nov. is described from the Marojejy massif and is also known from Andrevorevo, Lohanandroranga and Sorata; it is similar to U. ebenaui and U. fetsy but is distinguishable by its rather short tail and only partially black pigmented oral mucosa. Uroplatus fivehy sp. nov. is described from the Sorata Massif and is wider-ranging, occurring in an area from Marotandrano and Makira to Sorata, comprising Marojejy, Anjanaharibe-Sud, Ankitsika and Betaolana; it is morphologically similar to U. finiavana but distinguishable by tail size and shape. Both newly described species are found in rainforest from mid to high elevation and range respectively from 840-1417 m for U. fivehy sp nov. and 1300-1800 m for U. fangorn sp. nov. According to their respective geographical distributions, we propose to classify the two new species as Vulnerable under the IUCN Red List criteria, due to their occurrence in some forests outside of the protected area network plus the continuing decline of forest patches in the north of Madagascar.

Taxonomic progress is often hindered by intrinsic factors, such as morphologically cryptic species that require a broad suite of methods to distinguish, and extrinsic factors, such as uncertainties in the allocation of scientific names to species. These uncertainties can be due to a wide variety of factors, including old and poorly preserved type specimens (which contain only heavily degraded DNA or have lost important diagnostic characters), inappropriately chosen type specimens (e.g. juveniles without diagnostic characters) or poorly documented type specimens (with unprecise, incorrect, or missing locality data). Thanks to modern sequencing technologies it is now possible to overcome many such extrinsic factors by sequencing DNA from name-bearing type specimens of uncertain assignment and assigning these to known genetic lineages. Here, we apply this approach to frogs of the Mantidactylus ambreensis complex, which was recently shown to consist of two genetic lineages supported by concordant differentiation in mitochondrial and nuclear genes. These lineages co-occur on the Montagne d'Ambre Massif in northern Madagascar but appear to have diverged in allopatry. We use a recently published bait set based on three mitochondrial markers from all known Malagasy frog lineages to capture DNA sequences from the 127-year-old holotype of Mantidactylus ambreensis Mocquard, 1895. With the obtained sequences we are able to assign the name M. ambreensis to the lowland lineage, which is rather widespread in the rainforests of northern Madagascar, leaving the microendemic high-elevation lineage on Montagne d'Ambre in north Madagascar in need of description. We describe this species as Mantidactylus ambony sp. nov., differing from M. ambreensis in call parameters and a smaller body size. Thus, using target enrichment to obtain DNA sequence data from this old specimen, we were able to resolve the extrinsic (nomenclatural) hindrances to taxonomic resolution of this complex. We discuss the broad-scale versatility of this 'barcode fishing' approach, which can draw on the enormous success of global DNA barcoding initiatives to quickly and efficiently assign type specimens to lineages.

Madagascar's biota is characterized by a high degree of microendemism at different taxonomic levels, but how colonization and in-situ speciation contribute to the assembly of local species communities has rarely been studied on this island. Here we analyze the phylogenetic relationships of riparian frogs of the Mantidactylus ambreensis species complex, which is distributed in the north of Madagascar and was originally described from Montagne d'Ambre, an isolated mountain of volcanic origin, currently protected within Montagne d'Ambre National Park (MANP). Data from mitochondrial DNA, and phylogenomic data from FrogCap, a sequence capture method, independently confirm that this species complex is monophyletic within the subgenus Ochthomantis, and identify two main clades within it. These two clades are separated by 5.6-6.8% pairwise distance in the mitochondrial 16S rRNA gene and co-occur in MANP, with one distributed at high elevations (940-1375 m a.s.l.) and the other at lower elevations (535-1010 m a.s.l.), but show almost no haplotype sharing in the nuclear RAG1 gene. This occurrence in syntopy without admixture confirms them as independent evolutionary lineages that merit recognition as separate species, and we here refer to them as high-elevation (HE) and low-elevation (LE) lineage; they will warrant taxonomic assessment to confidently assign the name ambreensis to one or the other. Populations of the M. ambreensis complex from elsewhere in northern Madagascar all belong to the LE lineage, although they do occur over a larger elevational range than in Montagne d'Ambre (285-1040 m a.s.l.). Within LE there are several phylogroups (LE1-LE4) of moderately deep divergence (1.5-2.8% in 16S), but phylogroup LE4 that occurs in MANP has a deeply nested phylogenetic position, as recovered separately by mitochondrial and sequence capture datasets. This suggests that HE and LE did not diverge by a local fission of lower and upper populations, but instead arose through a more complex biogeographic scenario. The branching pattern of phylogroups LE1-LE4 shows a clear south-to-north phylogeographic pattern. We derive from these results a testable hypothesis of vicariant speciation that restricted the HE lineage to MANP and the LE candidate species to a climatic refugium further south, with subsequent northwards range expansion and secondary colonization of MANP by LE. These results provide an example for complex assembly of local microendemic amphibian faunas on Madagascar., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Processes leading to spectacular diversity of both form and species on islands have been well-documented under island biogeography theory, where distance from source and island size are key factors determining immigration and extinction resistance. But far less understood are the processes governing in situ diversification on the world's mega islands, where large and isolated land masses produced morphologically distinct radiations from related taxa on continental regions. Madagascar has long been recognized as a natural laboratory due to its isolation, lack of influence from adjacent continents, and diversification of spectacular vertebrate radiations. However, only a handful of studies have examined rate shifts of in situ diversification for this island. Here, we examine rates of diversification in the Malagasy snakes of the family Pseudoxyrhophiinae (gemsnakes) to understand if rates of speciation were initially high, enhanced by diversification into distinct biomes, and associated with key dentition traits. Using a genomic sequence-capture data set for 366 samples, we determine that all previously described and newly discovered species are delimitable and therefore useful candidates for understanding diversification trajectories through time. Our analysis detected no shifts in diversification rate between clades or changes in biome or dentition type. Remarkably, we demonstrate that rates of diversification of the gemsnake radiation, which originated in Madagascar during the early Miocene, remained steady throughout the Neogene. However, we do detect a significant slowdown in diversification during the Pleistocene. We also comment on the apparent paradox where most living species originated in the Pleistocene, despite diversification rates being substantially higher during the earlier 15 myr., (© The Author(s) 2019. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Previous studies on leaf-tailed geckos of the genus Uroplatus identified a lineage from the Ankarana karst massif in northern Madagascar as genetically highly divergent, but only fragmentary information was available on these geckos. Here, we provide an integrative analysis based on molecular and morphological data, including a newly collected specimen from this locality. Phylogenetic analysis placed the Ankarana lineage sister to U. ebenaui, but with a surprisingly high genetic divergence of over 19% uncorrected pairwise distance in the mitochondrial 16S rRNA gene. We formally describe the Ankarana lineage as Uroplatus fetsy sp. nov. and provide new information on the distribution and genetic differentiation of its sister species, Uroplatus ebenaui. Based on DNA sequences of newly examined samples this latter species is confirmed as widespread, ranging from low-elevation dry forests from Beanka in western Madagascar to Forêt d'Ambre and Analalava/Fanambana in northern Madagascar, and thus far has not been found in the Ankarana Massif, where U. fetsy occurs.

The Marojejy Massif in northern Madagascar is a constant source of herpetological surprises. Herein we describe a new species of leaf-mimicking leaf-tailed gecko, Uroplatus finaritra sp. nov., based on morphological and molecular phylogenetic evidence. This new species inhabits the rainforests of Marojejy National Park at low elevations and is morphologically similar to Uroplatus phantasticus, but differs by having a larger body size, relatively shorter tail, and dark-red pigmentation of the oral mucosa. Molecular data reveals a distance of 14.0-14.7% in the 16S rRNA gene to U. phantasticus, and haplotype sharing in the c-mos nuclear gene only with species of much shorter tails (i.e., with U. kelirambo and two undescribed candidate species). Similar to other members of the leaf-mimicking Uroplatus species in northern Madagascar, Uroplatus finaritra sp. nov. likely has a small geographic distribution, and is currently known only from the lowland rainforests of Marojejy. We discuss the value of the buccal mucosa colouration and tail dimensions of Uroplatus as taxonomic characters, and the potential drivers of their evolution. In order to reduce the risk of international trade under incorrect species names we suggest that exported CITES species should be obligatorily accompanied by information about their precide geographic origin within Madagascar.

Background: Few details are available on the consumption of ectoparasites, specifically bat flies (Diptera: Nycteribiidae and Streblidae), by their chiropteran hosts while grooming. Such details are important to document consumption rates of ectoparasites by their bat host provide details on the dynamics of host-parasite interactions. We present data on ectoparasite consumption rates for an endemic Malagasy fruit bat (Pteropodidae: Rousettus madagascariensis) occupying a cave day roost colony in northern Madagascar. Using quantified behavioral analyses, grooming and associated ingestion rates were measured from infrared videos taken in close proximity to day-roosting bats. The recorded individual bats could be visually identified to age (adult, juvenile) and sex (male, female), allowing analyses of the proportion of time these different classes allocated to consuming ectoparasites via auto-grooming (self) or allo-grooming (intraspecific) per 10 min video recording session. These figures could then be extrapolated to estimates of individual daily consumption rates., Results: Based on video recordings, adults spent significantly more time auto-grooming and allo-grooming than juveniles. The latter group was not observed consuming ectoparasites. Grooming rates and the average number of ectoparasites consumed per day did not differ between adult males and females. The mean extrapolated number consumed on a daily basis for individual adults was 37 ectoparasites. When these figures are overlaid on the estimated number of adult Rousettus occurring at the roost site during the dry season, the projected daily consumption rate was 57,905 ectoparasites., Conclusions: The details presented here represent the first quantified data on bat consumption rates of their ectoparasites, specifically dipterans. These results provide new insights in host-parasite predation dynamics. More research is needed to explore the mechanism zoonotic diseases isolated from bat flies might be transmitted to their bat hosts, specifically those pathogens that can be communicated via an oral route.

The rainforest biome of eastern Madagascar is renowned for its extraordinary biodiversity and restricted distribution ranges of many species, whereas the arid western region of the island is relatively species poor. We provide insight into the biogeography of western Madagascar by analyzing a multilocus phylogeographic dataset assembled for an amphibian, the widespread Malagasy bullfrog, Laliostoma labrosum. We find no cryptic species in L. labrosum (maximum 1.1% pairwise genetic distance between individuals in the 16S rRNA gene) attributable to considerable gene flow at the regional level as shown by genetic admixture in both mtDNA and three nuclear loci, especially in central Madagascar. Low breeding site fidelity, viewed as an adaptation to the unreliability of standing pools of freshwater in dry and seasonal environments, and a ubiquitous distribution within its range may underlie overall low genetic differentiation. Moreover, reductions in population size associated with periods of high aridity in western Madagascar may have purged DNA variation in this species. The mtDNA gene tree revealed seven major phylogroups within this species, five of which show mostly non-overlapping distributions. The nested positions of the northern and central mtDNA phylogroups imply a southwestern origin for all extant mtDNA lineages in L. labrosum. The current phylogeography of this species and paleo-distributions of major mtDNA lineages suggest five potential refugia in northern, western and southwestern Madagascar, likely the result of Pleistocene range fragmentation during drier and cooler climates. Lineage sorting in mtDNA and nuclear loci highlighted a main phylogeographic break between populations north and south of the Sambirano region, suggesting a role of the coastal Sambirano rainforest as a barrier to gene flow. Paleo-species distribution models and dispersal networks suggest that the persistence of some refugial populations was mainly determined by high population connectivity through space and time., (Copyright © 2015 Elsevier Inc. All rights reserved.)

We describe a new leaf-tailed gecko species of the Uroplatus ebenaui group from the eastern central rainforests of Madagascar, which had previously been considered as a confirmed candidate species. Our description of Uroplatus fiera sp. nov. relies on integrating evidence from molecular and morphological characters and is based on newly collected material from two localities. A phylogenetic analysis based on multiple mitochondrial DNA fragments places the new species as sister to a lineage of uncertain status (Uroplatus ebenaui [Ca8]), and the clade consisting of these two lineages is sister to a further undescribed candidate species (U. ebenaui [Ca1]). This entire clade is sister to U. phantasticus plus another candidate species. The new species differs from these close relatives, and all other congenerics, by strong differences in DNA sequences of mitochondrial genes (>8.5% uncorrected p-distance in 16S rDNA to all nominal species of the genus) and lacks shared alleles with any of the nominal species in the nuclear CMOS gene. From its closest relatives the new species further differs in its much smaller tail size (relative to U. phantasticus), and a narrower tail, fewer supralabials, and more toe lamellae (relative to U. ebenaui [Ca1]). Morphologically the new species is most similar to U. ebenaui but differs in its larger body size and unpigmented oral mucosa. Given its distribution in central eastern Madagascar, with records from near Fierenana and Ambatovy, its range overlaps with that of U. phantasticus. Based on examination of the U. phantasticus holotype, we confirm that this latter has a blackish pigmented oral mucosa as do those specimens typically attributed to this nomen, thereby confirming its distinctness from U. fiera sp. nov., in which the mucosa is unpigmented.

There are insufficient resources available to manage the world's existing protected area portfolio effectively, so the most important sites should be prioritised in investment decision-making. Sophisticated conservation planning and assessment tools developed to identify locations for new protected areas can provide an evidence base for such prioritisations, yet decision-makers in many countries lack the institutional support and necessary capacity to use the associated software. As such, simple heuristic approaches such as species richness or number of threatened species are generally adopted to inform prioritisation decisions. However, their performance has never been tested. Using the reptile fauna of Madagascar's dry forests as a case study, we evaluate the performance of four site prioritisation protocols used to rank the conservation value of 22 established and candidate protected areas. We compare the results to a benchmark produced by the widely-used systematic conservation planning software Zonation. The four indices scored sites on the basis of: i) species richness; ii) an index based on species' Red List status; iii) irreplaceability (a key metric in systematic conservation planning); and, iv) a novel Conservation Value Index (CVI), which incorporates species-level information on endemism, representation in the protected area system, tolerance of habitat degradation and hunting/collection pressure. Rankings produced by the four protocols were positively correlated to the results of Zonation, particularly amongst high-scoring sites, but CVI and Irreplaceability performed better than Species Richness and the Red List Index. Given the technological capacity constraints experienced by decision-makers in the developing world, our findings suggest that heuristic metrics can represent a useful alternative to more sophisticated analyses, especially when they integrate species-specific information related to extinction risk. However, this can require access to, and understanding of, more complex species data.

Lizards and snakes exhibit colour variation of adaptive value for thermoregulation, camouflage, predator avoidance, sexual selection and speciation. Furcifer pardalis, the panther chameleon, is one of the most spectacular reptilian endemic species in Madagascar, with pronounced sexual dimorphism and exceptionally large intraspecific variation in male coloration. We perform here an integrative analysis of molecular phylogeography and colour variation after collecting high-resolution colour photographs and blood samples from 324 F. pardalis individuals in locations spanning the whole species distribution. First, mitochondrial and nuclear DNA sequence analyses uncover strong genetic structure among geographically restricted haplogroups, revealing limited gene flow among populations. Bayesian coalescent modelling suggests that most of the mitochondrial haplogroups could be considered as separate species. Second, using a supervised multiclass support vector machine approach on five anatomical components, we identify patterns in 3D colour space that efficiently predict assignment of male individuals to mitochondrial haplogroups. We converted the results of this analysis into a simple visual classification key that can assist trade managers to avoid local population overharvesting., (© 2015 The Authors. Molecular Ecology Published by John Wiley & Sons Ltd.)

Although numerous studies have attempted to find single unifying mechanisms for generating Madagascar's unique flora and fauna, little consensus has been reached regarding the relative importance of climatic, geologic and ecological processes as catalysts of diversification of the region's unique biota. Rather, recent work has shown that both biological and physical drivers of diversification are best analysed in a case-by-case setting with attention focused on the ecological and life-history requirements of the specific phylogenetic lineage under investigation. Here, we utilize a comprehensive analytical approach to examine evolutionary drivers and elucidate the biogeographic history of Malagasy plated lizards (Zonosaurinae). Data from three genes are combined with fossil information to construct time-calibrated species trees for zonosaurines and their African relatives, which are used to test alternative diversification hypotheses. Methods are utilized for explicitly incorporating phylogenetic uncertainty into downstream analyses. Species distribution models are created for 14 of 19 currently recognized species, which are then used to estimate spatial patterns of species richness and endemicity. Spatially explicit analyses are employed to correlate patterns of diversity with both topographic heterogeneity and climatic stability through geologic time. We then use inferred geographic ranges to estimate the biogeographic history of zonosaurines within each of Madagascar's major biomes. Results suggest constant Neogene and Quaternary speciation with divergence from the African most recent common ancestor ~30 million years ago when oceanic currents and African rivers facilitated dispersal. Spatial patterns of diversity appear concentrated along coastal regions of northern and southern Madagascar. We find no relationship between either topographic heterogeneity or climatic stability and patterns of diversity. Ancestral state reconstructions suggest that western dry forests were important centres of origin with recent invasion into spiny and rain forest. These data highlight the power of combining multilocus phylogenetic and spatially explicit analyses for testing alternative diversification hypotheses within Madagascar's unique biota and more generally, particularly as applied to phylogenetically and biologically constrained systems., (© 2015 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2015 European Society For Evolutionary Biology.)

Elapotinus picteti Jan, 1862 is an enigmatic snake taxon that has been described without locality data. Genus and species, both based on the unique holotype, were considered to belong to the venomous African Aparallactinae for more than a century, but although this taxon was never rediscovered it was accepted as a valid species until present. To clarify the taxonomic status of E. picteti its characters were compared with literature and determination keys for the whole world. This literature survey and the subsequent study of type specimens revealed that the monotypic Elapotinus is a subjective senior synonym of the monotypic Malagasy pseudoxyrhophiine snake genus Exallodontophis Cadle, 1999 which is characterized by its unique aglyphous dentition. We therefore transfer Elapotinus from the lamprophiid subfamily Aparallactinae to the subfamily Pseudoxyrhophiinae. Furthermore, Elapotinus picteti strongly resembles the Malagasy species Exallodontophis albignaci (Domergue, 1984) in external morphology, coloration and dentition. As a consequence we consider Elapotinus picteti a subjective senior synonym of Exallodontophis albignaci.

Background: An understanding of the conservation status of Madagascar's endemic reptile species is needed to underpin conservation planning and priority setting in this global biodiversity hotspot, and to complement existing information on the island's mammals, birds and amphibians. We report here on the first systematic assessment of the extinction risk of endemic and native non-marine Malagasy snakes, lizards, turtles and tortoises., Methodology/principal Findings: Species range maps from The IUCN Red List of Threatened Species were analysed to determine patterns in the distribution of threatened reptile species. These data, in addition to information on threats, were used to identify priority areas and actions for conservation. Thirty-nine percent of the data-sufficient Malagasy reptiles in our analyses are threatened with extinction. Areas in the north, west and south-east were identified as having more threatened species than expected and are therefore conservation priorities. Habitat degradation caused by wood harvesting and non-timber crops was the most pervasive threat. The direct removal of reptiles for international trade and human consumption threatened relatively few species, but were the primary threats for tortoises. Nine threatened reptile species are endemic to recently created protected areas., Conclusions/significance: With a few alarming exceptions, the threatened endemic reptiles of Madagascar occur within the national network of protected areas, including some taxa that are only found in new protected areas. Threats to these species, however, operate inside and outside protected area boundaries. This analysis has identified priority sites for reptile conservation and completes the conservation assessment of terrestrial vertebrates in Madagascar which will facilitate conservation planning, monitoring and wise-decision making. In sharp contrast with the amphibians, there is significant reptile diversity and regional endemism in the southern and western regions of Madagascar and this study highlights the importance of these arid regions to conserving the island's biodiversity.

We analyzed the phylogenetic relationships of the Malagasy plated lizards in the family Gerrhosauridae based on DNA sequence fragments of four mitochondrial and five nuclear genes. Various clades were strongly supported by the concatenated data set and also recovered by separate analyses of mtDNA and nucDNA. In particular, two clades here named the Z. rufipes group (containing Z. bemaraha , Z. brygooi , Z. rufipes , Z. subunicolor , Z. tsingy and an undescribed candidate species from northern Madagascar) and the Z. ornatus group (containing Z. anelanelany , Z. laticaudatus , Z. karsteni , Z. ornatus , Z. quadrilineaus, and Z. trilineatus) were resolved with strong support. A third clade named the Z. madagascariensis group contains Z. madagascariensis with a nested Z. haraldmeieri; the status of that species requires further investigation. Tentatively we also include Z. aeneus in this species group although its phylogenetic relationships were poorly resolved. A fourth clade with less support included Z. boettgeri and Z. maximus. The phylogenetic position of the genus Tracheloptychus remains uncertain: whereas in the species tree it was recovered as the sister group to Zonosaurus, other methods indicated that it was nested within Zonosaurus, albeit alternative topologies were rejected with only marginal statistical support., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Understanding the remarkably high species diversity and levels of endemism found among Madagascar's flora and fauna has been the focus of many studies. One hypothesis that has received much attention proposes that Quaternary climate fluctuations spurred diversification. While spatial patterns of distribution and phylogenetic relationships can provide support for biogeographic predictions, temporal estimates of divergence are required to determine the fit of these geospatial patterns to climatic or biogeographic mechanisms. We use multilocus DNA sequence data to test whether divergence times among Malagasy iguanid lizards of the subfamily Oplurinae are compatible with a hypotheses of Pliocene-Pleistocene diversification. We estimate the oplurine species tree and associated divergence times under a relaxed-clock model. In addition, we examine the phylogeographic structure and population divergence times within two sister species of Oplurus primarily distributed in the north-west and south-west of Madagascar (Oplurus cuvieri and Oplurus cyclurus, respectively). We find that divergence events among oplurine lineages occurred in the Oligocene and Miocene and are thus far older and incompatible with the hypothesis that recent climate fluctuations are related to current species diversity. However, the timing of intraspecific divergences and spatial patterns of population genetic structure within O. cuvieri and O. cyclurus suggest a role for both intrinsic barriers and recent climate fluctuations at population-level divergences. Integrating information across spatial and temporal scales allows us to identify and better understand the mechanisms generating patterns diversity., (© 2012 Blackwell Publishing Ltd.)

The endemic plated lizards (Gerrhosauridae) of Madagascar are one of the most diverse groups of lizards on the island (19 species) and are found in all ecoregions. On an island that presents so many interesting biological questions, plated lizards are an ideal group for examining patterns of diversification due to their high (but tractable) diversity and wide distribution. To resolve the taxonomy and evolutionary relationships of Malagasy plated lizards, and to explore their biogeographic history, we have generated a molecular phylogeny based on >2kb of both mitochondrial (cob, 1142 bp) and nuclear (NT-3, 419 bp; c-mos, 542 bp) DNA sequences. Phylogenetic relationships and divergence-time estimates indicate Zonosaurinae are monophyletic, arising from a single colonization event, likely from Africa to Madagascar in the Paleogene (Paleocene/early Eocene), with subsequent radiation. Furthermore, our results reveal widespread mischaracterization of species delimitation and assignment based on phenotypic characteristics. Paraphyly of a number of zonosaurine species was strongly supported by our dataset, revealing not only instances of likely cryptic species (phenotypic conservatism) but also phenotypic/genotypic discordance in the delimitation of species (phenotypic variability). These results clearly demonstrate the complex history of biotic diversification on Madagascar and provide novel insight into biogeographic patterns on the island.

The biodiversity of Madagascar is extraordinarily distinctive, diverse, and endangered. It is therefore urgent that steps be taken to document, describe, interpret, and protect this exceptional biota. As a collaborative group of field and laboratory biologists, we employ a suite of methodological and analytical tools to investigate the vertebrate portion of Madagascar's fauna. Given that species are the fundamental unit of evolution, where micro- and macroevolutionary forces converge to generate biological diversity, a thorough understanding of species distribution and abundance is critical for understanding the evolutionary, ecological, and biogeographic forces that have shaped Malagasy vertebrate diversity. We illustrate the means by which we apply Mayr's "three basic tasks" of the systematist [Mayr, E. (1942) Systematics and the Origin of Species from the Viewpoint of a Zoologist (Harvard Univ. Press, Cambridge, MA)] to identify, classify, and study the organisms that together constitute Madagascar's vertebrate community. Using field inventory methods, specimen-based studies, and morphological and molecular analyses, we formulate hypotheses of species identity that then serve as the foundation for subsequent studies of biology and history. Our experience, as well as that of other investigators, has shown that much of the vertebrate species diversity in Madagascar is "cryptic" for both biological and practical reasons. Beyond issues of cryptic biological diversity, the resolution of species identity in Madagascar has been hampered because of a lack of vouchered comparative material at the population level. Through our activities, we are attempting to remedy these limitations while simultaneously enhancing research capacity in Madagascar.