Sunda-Papuan keelback snakes (Serpentes: Natricidae: Tropidonophis Jan 1863) include 20 species distributed from the Philippines south-east through the Moluccas to New Guinea and Australia. Diversity of this insular snake lineage peaks on the island of New Guinea. Previous phylogenetic studies incorporating Tropidonophis have been limited to multi-locus Sanger-sequenced datasets with broad squamate or family-level focus. We used a targeted-sequence capture approach to sequence thousands of nuclear ultraconserved elements (UCEs) to construct the most comprehensive sequence-based phylogenetic hypothesis for this genus and estimate ancestral biogeography. Phylogenies indicate the genus is monophyletic given recent taxonomic reassignment of Rhabdophis spilogaster to Tropidonophis. All UCE phylogenies recovered a monophyletic Tropidonophis with reciprocally monophyletic Philippine and New Guinean clades. Divergence dating and ancestral range estimation suggest dispersal to New Guinea from the Philippines to have occurred during the Mid-Miocene via the Oceanic Arc Terranes. From Late Miocene into the Pliocene the genus experienced rapid diversification from orogeny of the New Guinean Central Cordillera from Oceanic Arc Terrane accretion on the northern boundary of the Sahul Shelf. Future collecting of missing taxa from the Moluccas and Indonesian Papua will better the understanding of non-volant faunal biogeography and diversification in this tectonically complex Pacific arena.

Skinks are the most diverse component of the reptile fauna in the mountains of New Guinea and many seemingly specialised high -elevation species remain undescribed. Here we describe two spectacular new gold -patterned skinks in the montane -specialist genus Papuascincus. Both species can be diagnosed from all congeners by their distinctive colouration, in addition to aspects of scalation and body size. One new species is mainly recorded from lower montane forest in karst habitats spanning more than five hundred kilometres along the southern edge of New Guinea's Central Cordillera and is likely to warrant an IUCN conservation status of Least Concern. The second new species has thus far only been recorded from cloud forest on the summit of Mt. Menawa in the North Coastal Ranges and we suggest it should be considered Data Deficient. However, if further survey work confirms a restricted distribution with little scope for upslope elevational retreat under future warming climates it will likely qualify for Endangered or Critically Endangered status.

Environmental factors, such as temperature, precipitation, and elevation, explain most of the variation in species richness at the global scale. Nevertheless, richness patterns may have different drivers across taxa and regions. To date, a comprehensive global examination of how various factors such as climate or topography drive patterns of species richness across all terrestrial vertebrates, using the same methods and predictors, has been lacking. Recent advances in species-distribution data allowed us to model and examine the richness pattern of all terrestrial tetrapods comprehensively. We tested the relationship between environmental and biogeographical variables and richness of amphibians (5983 species), birds (9630), mammals (5004), reptiles (8939), and tetrapods as a whole, globally, and across biogeographical realms. We studied the effects of climatic, ecological, and biogeographic drivers using generalized additive models. Richness patterns and their environmental associations varied among taxa and realms. Overall precipitation was the predominant richness predictor. However, temperature was more important in realms where both cold and warm conditions exist. In the Indomalayan realm, elevational range was very important. Richness patterns of mammals, birds, and amphibians were strongly related to precipitation whereas reptile richness was mostly associated with temperature. Our results support the universal importance of precipitation but also suggest that future global-scaled research should incorporate other relevant variables other than climate, such as elevational range, to gain a better understanding of the richness-environment relationship. By doing so, we can further advance our knowledge of the complex relationships between biodiversity and the environment. In this study we tested the relationship between environmental and biogeographical variables and richness of amphibians, birds, mammals, reptiles, and all tetrapods, globally, and across biogeographical realms, using the most up-to-date richness dataset. We found taxonomic and spatial variations, but about a common theme. Precipitation is largely the most influential predictor with the exception of the Nearctic (and the Palearctic to some extent) and reptiles. Elevational range is usually less important than climate, but it is highly influential in the Indomalaya realm.*image

Context Skinks comprise the dominant component of the terrestrial vertebrate fauna in Oceania, New Guinea, and Eastern Wallacea (ONGEW). However, knowledge of their diversity is incomplete, and their conservation needs are poorly understood. Aims To explore the diversity and threat status of the skinks of ONGEW and identify knowledge gaps and conservation needs. Methods We compiled a list of all skink species occurring in the region and their threat categories designated by the International Union for Conservation of Nature. We used available genetic sequences deposited in the National Center for Biotechnology Information’s GenBank to generate a phylogeny of the region’s skinks. We then assessed their diversity within geographical sub-divisions and compared to other reptile taxa in the region. Key results Approximately 300 species of skinks occur in ONGEW, making it the second largest global hotspot of skink diversity following Australia. Many phylogenetic relationships remain unresolved, and many species and genera are in need of taxonomic revision. One in five species are threatened with extinction, a higher proportion than almost all reptile families in the region. Conclusions ONGEW contain a large proportion of global skink diversity on <1% of the Earth’s landmass. Many are endemic and face risks such as habitat loss and invasive predators. Yet, little is known about them, and many species require taxonomic revision and threat level re-assessment. Implications The skinks of ONGEW are a diverse yet underexplored group of terrestrial vertebrates, with many species likely facing extreme risks in the near future. Further research is needed to understand the threats they face and how to protect them.

A bstract Studies of the Papuan region have provided fundamental insights into both the evolutionary processes generating its exceptional biodiversity, as well support for alternative hypotheses of geological history. Lying at the junction of five tectonic plates, this region has experienced a turbulent geological history that has not only produced towering mountains allowing elevational specialization, and island archipelagos of varying distance promoting vicariance, but also active margins where land masses have collided and been subsequently rifted apart creating a mosaic of intermixed terranes with vastly different geological histories which may influence the evolutionary history of its biota. Asterophryine frogs are a hyperdiverse clade representing half the world’s microhylid diversity (over 360 species) centered on New Guinea and its satellite islands. We show that vicariance facilitated by geological history, and not elevational specialization best explain this far and wide distribution of a clade that should have poor dispersal abilities. Thus, some of the predictions of island biogeography theory are supported if informed by geological history. We recovered a mainland tectonic unit, the East Papua Composite Terrane (EPCT), as the center of origin for Asterophryinae and no fewer than 71 instances of what appear to be long-distance dispersal events, 29 of which are between mainland regions, with 42 from the mainland to the islands, some presently as far as 200 km away from source populations over open ocean. Furthermore, we find strong support for a “Slow and Steady” hypothesis for the formation of the northern margin of New Guinea by many separate accretion events during the Miocene, over other major geological alternatives, consistent with the 20 M year age of the clade and arrival via the EPCT. In addition, the historical biogeography of our frogs strongly support an affiliation of the Louisiade Archipelago and Woodlark Island with the Owen Stanley Mountain range on the EPCT, and the recent proximity of the large New Britain island. Our results show that Asterophryinae did not have to repeatedly and independently disperse across and large ocean barriers to the offshore islands, but that the current distribution can be explained through vicariance and short-distance oceanic dispersal as historical land connections disappeared and islands slowly became separated from each other. We show that islands have a life history, undergoing changes in area through island-building and erosion, but also change in distance from other land masses, with consequent opportunities for dispersal, isolation, and cladogenesis of their biotas. More broadly, we can begin to see how the geological history of the Papuan region can result in the rapid accumulation and staggering number of extant species.

The data provided here are related to the article "Resolving the Deep Phylogeny: Implications for Early Adaptive Radiation, Cryptic, and Present-day Ecological Diversity of Papuan Microhylid Frogs" [1]. The dataset is based on 233 tissue samples of the subfamily Asteroprhyinae, with representatives from all recognized genera, in addition to three outgroup taxa. The sequence dataset contains over 2400 characters per sample for five genes: three nuclear (Seventh in Absentia (SIA), Brain Derived Neurotrophic Factor (BDNF), Sodium Calcium Exchange subunit-1 (NXC-1)), and two mitochondrial loci (Cytochrome oxidase b (CYTB), and NADH dehydrogenase subunit 4 (ND4)); and is 99% complete. New primers were designed for all loci and accession numbers for the raw sequence data are provided. The sequences are used with geological time calibrations to produce time-calibrated Bayesian inference (BI) and Maximum Likelihood (ML) phylogenetic reconstructions using BEAST2 and IQ-TREE. Lifestyle data (arboreal, scansorial, terrestrial, fossorial, semi-aquatic) were collected from the literature and field notes and used to infer ancestral character states for each lineage. Collection location and elevation data were used to verify sites where multiple species or candidate species co-occur. All sequence data, alignments, and associated metadata (voucher specimen number, species identification, type locality status, global positioning system [GPS] coordinates, elevation, site with species list, and lifestyle) as well as the code to produce all analyses and figures are provided.

Studies of the Papuan region have provided fundamental insights into the evolutionary processes generating its exceptional biodiversity, but the influence of geological processes merits further study. Lying at the junction of five tectonic plates, this region has experienced a turbulent geological history that has not only produced towering mountains allowing elevational specialization and island archipelagos with varying degrees of isolation promoting vicariance, but also active margins where land masses have collided and been subsequently rifted apart creating a mosaic of intermixed terranes with vastly different geological histories. Asterophryine frogs are a hyperdiverse clade representing half the world's microhylid diversity (over 360 species) centered on New Guinea and its satellite islands. We show that vicariance facilitated by geological history explains this far and wide distribution of a clade that should have poor dispersal abilities. We recovered a mainland tectonic unit, the East Papua Composite Terrane (EPCT), as the center of origin for Asterophryinae and no fewer than 71 instances of what appear to be long-distance dispersal events, 29 of which are between mainland regions, with 42 from the mainland to the islands, some presently as far as 200 km away from source populations over open ocean. Furthermore, we find strong support for a "Slow and Steady" hypothesis for the formation of the northern margin of New Guinea by many separate accretion events during the Miocene, over other major geological alternatives, consistent with the 20 M year age of the clade and arrival via the EPCT. In addition, the historical biogeography of our frogs strongly supports an affiliation of the Louisiade Archipelago and Woodlark Island with the Owen Stanley Range on the EPCT, and the recent proximity of the large New Britain Island. Our results show that Asterophryinae did not have to repeatedly and independently disperse across large ocean barriers to the offshore islands, against the predictions of island biogeography theory, but that the current distribution can be explained through vicariance and short-distance oceanic dispersal as historical land connections disappeared and islands slowly became separated from each other. We show that islands have a life history, changing in distance from other land masses, with consequent opportunities for dispersal, isolation, and cladogenesis of their biotas. More broadly, we can begin to see how the geological history of the Papuan region can result in the rapid accumulation and staggering number of extant species.

Comprehensive assessments of species' extinction risks have documented the extinction crisis(1) and underpinned strategies for reducing those risks(2). Global assessments reveal that, among tetrapods, 40.7% of amphibians, 25.4% of mammals and 13.6% of birds are threatened with extinction(3). Because global assessments have been lacking, reptiles have been omitted from conservation-prioritization analyses that encompass other tetrapods(4-7). Reptiles are unusually diverse in arid regions, suggesting that they may have different conservation needs(6). Here we provide a comprehensive extinction-risk assessment of reptiles and showthat at least 1,829 out of 10,196 species (21.1%) are threatened-confirming a previous extrapolation(8) and representing 15.6 billion years of phylogenetic diversity. Reptiles are threatened by the same major factorsthat threaten othertetrapods-agriculture, logging, urban development and invasive species-although the threat posed by climate change remains uncertain. Reptiles inhabiting forests, where these threats are strongest, are more threatened than those in arid habitats, contrary to our prediction. Birds, mammals and amphibians are unexpectedly good surrogates for the conservation of reptiles, although threatened reptiles with the smallest rangestend to be isolated from other threatened tetrapods. Although some reptiles-including most species of crocodiles and turtles-require urgent, targeted action to prevent extinctions, efforts to protect other tetrapods, such as habitat preservation and control of trade and invasive species, will probably also benefit many reptiles.

The microhylid frogs of the New Guinea region are the largest and most ecologically diverse subfamily (Asterophryinae) of one of the largest anuran families in the world and can live in communities of up to 20 species. While there has been recent progress in resolving the phylogenetic relationships of Asterophryinae, significant uncertainties remain, impeding further progress in understanding the evolution of microhabitat use, parental care, and life history variation in this group. In particular, the early divergences at the base of the tree remain unclear; as does the monophyly of some genera; and recent studies have discovered that species with wide geographic distribution are instead cryptic species complexes. In this study, we fortified geographic sampling of the largest previous phylogenetic effort by sequencing an additional 62 taxa and increased data quality and quantity by adding new layers of data vetting and by filling in previously incomplete loci to the five gene dataset (2 mitochondrial, 3 nuclear protein-coding genes) to obtain a dataset that is now 99% complete in over 2400 characters for 233 samples (205 taxa) of Asterophryinae and 3 outgroup taxa, and analyzed microhabitat use data for these taxa from field data and data collected from the literature. Importantly, our sampling includes complete community complements at 19 sites as well as representatives at over 80 sites across New Guinea and its offshore islands. We present a highly resolved molecular phylogeny which, for the first time, has over 95% of nodes supported (84% highly supported) whether using Maximum Likelihood or Bayesian Inference, allowing clarification of all genera (whether monophyletic or clearly not), their sister genera relationships, as well as an age estimate for the Asterophryinae at approximately 20MYA. Early generic diversification occurring between 17 and 12 MYA gave rise to a surprising diversity of about 18 genera as well as the 5 putative microhabitat types. Our tree reveals extensive cryptic diversity calling any widespread taxa into doubt, and clearly demonstrates that complex multispecies communities of Asterophryinae are ecologically diverse, are numerous, and of ancient origin across New Guinea. We discuss the implications of our phylogeny for explaining the explosive diversification of Asterophryinae as the result of adaptive radiation, niche conservatism, and non-adaptive radiation.

Identifying hotspots of biological diversity is a key step in conservation prioritisation. Melanesia—centred on the vast island of New Guinea—is increasingly recognised for its exceptionally species-rich and endemic biota. Here we show that Melanesia has the world’s most diverse insular amphibian fauna, with over 7% of recognised global frog species in less than 0.7% of the world’s land area, and over 97% of species endemic. We further estimate that nearly 200 additional candidate species have been discovered but remain unnamed, pointing to a total fauna in excess of 700 species. Nearly 60% of the Melanesian frog fauna is in a lineage of direct-developing microhylids characterised by smaller distributions than co-occurring frog families, suggesting lineage-specific high beta diversity is a key driver of Melanesian anuran megadiversity. A comprehensive conservation status assessment further highlights geographic concentrations of recently described range-restricted threatened taxa that warrant urgent conservation actions. Nonetheless, by world standards, the Melanesian frog fauna is relatively intact, with 6% of assessed species listed as threatened and no documented extinctions; and thus it provides an unparalleled opportunity to understand and conserve a megadiverse and relatively intact insular biota.

The data provided here is related to the article “Resolving the Deep Phylogeny: Implications for Early Adaptive Radiation, Cryptic, and Present-day Ecological Diversity of Papuan Microhylid Frogs” [1]. The dataset is based on 233 tissue samples of the subfamily Asteroprhyinae, with representatives from all recognized genera, in addition to three outgroup taxa. The sequence dataset contains over 2400 characters per sample for five genes (three nuclear loci: BDNF, NXC-1, and SIA; and two mitochondrial loci: ND4 and CYTB) and is 99% complete. New primers were designed for all loci and accession numbers for the raw sequence data are provided. The sequences are used with geological time calibrations to produce time-calibrated Bayesian inference and Maximum Likelihood phylogenetic reconstructions using BEAST2 and IQTREE. Lifestyle data (arboreal, scansorial, terrestrial, fossorial, semi-aquatic) were collected from the literature and field notes and used to infer ancestral character states for each lineage. Collection location data (GPS and elevation data) were used to verify sites where multiple species or candidate species co-occur. All sequence data, alignments, and associated metadata (voucher specimen number, species identification, type locality status, GPS, elevation, site with species list, and lifestyle) as well as the code to produce all analyses and figures are provided.

The skink genus Lobulia is endemic to New Guinea, the largest and highest tropical island in the world. Lobulia and its related genera represent an important component of the montane herpetofauna of New Guinea, but it remains understudied and poorly known. We here provide the first, large-scale, systematic revision of Lobulia, using molecular phylogenetic and morphological comparisons to assess the monophyly of the genus and the diversity of species within it. We find that Lobulia, as currently defined, is polyphyletic. The eight species currently assigned to it form three clades. Furthermore, many specimens from New Guinea of unknown specific affinity are genetically and morphologically distinct from each other. Based on these data, we re-diagnose Lobulia and two of its closely related genera, Prasinohaema and Papuascincus. We erect four new genera (Alpinoscincus gen. nov., Nubeoscincus gen. nov., Ornithuroscincus gen. nov. and Palaia gen. nov.) to address the problem of polyphyly and describe nine new species Lobulia fortis sp. nov., Lobulia huonensis sp. nov., Loublia marmorata sp. nov., Lobulia vogelkopensis sp. nov., Ornithuroscincus bengaun sp. nov., Ornithuroscincus inornatus sp. nov., Ornithuroscincus pterophilus sp. nov., Ornithuroscincus shearmani sp. nov. and Ornithuroscincus viridis sp. nov. We supplement this taxonomic revision by investigating the biogeographic history of Lobulia s.l. and find evidence for a large radiation in the accreted terranes of New Guinea, with multiple independent colonizations of montane habitats and subsequent recolonization of lowland habitats. Our study reinforces the uniqueness and richness of the montane herpetofauna of New Guinea and the importance of mountains to biodiversity in the Tropics.

Horizontal transfer (HT) of genes between multicellular animals, once thought to be extremely rare, is being more commonly detected, but its global geographic trend and transfer mechanism have not been investigated. We discovered a unique HT pattern of Bovine-B (BovB) LINE retrotransposons in vertebrates, with a bizarre transfer direction from predators (snakes) to their prey (frogs). At least 54 instances of BovB HT were detected, which we estimate to have occurred across time between 85 and 1.3 Ma. Using comprehensive transcontinental sampling, our study demonstrates that BovB HT is highly prevalent in one geographical region, Madagascar, suggesting important regional differences in the occurrence of HTs. We discovered parasite vectors that may plausibly transmit BovB and found that the proportion of BovB-positive parasites is also high in Madagascar where BovB thus might be physically transported by parasites to diverse vertebrates, potentially including humans. Remarkably, in two frog lineages, BovB HT occurred after migration from a non-HT area (Africa) to the HT hotspot (Madagascar). These results provide a novel perspective on how the prevalence of parasites influences the occurrence of HT in a region, similar to pathogens and their vectors in some endemic diseases.

1. Adaptations for efficient performance are expected to shape animal morphology based on selection for microhabitat use and ecological forces. The presence of competitor species is predicted to cause niches to contract and enhance trait divergence. Therefore, increased species richness is expected to lead to greater trait divergence, and to result in reduced overlap and similarity between morphologies of sympatric species. 2. We examined patterns of morphospace occupancy and partitioning in the skink fauna of New Guinea, the world's largest tropical island. Because skink species richness is largely decoupled from elevation in New Guinea, we could examine the effects of both factors (as proxies for competition and abiotic conditions), on morphospace occupancy and partitioning. 3. We measured 1,860 specimens from 79 species of skinks throughout Papua New Guinea, and examined their morphospace occupancy in a spatial context. We calculated, for each assemblage within equal-area cells, the volume of morphospace occupied by all skinks, the mean volume occupied per species, and the mean distance and overlap between all species pairs. We then examined whether these metrics are related to species richness and elevation. 4. Elevation is a stronger predictor of morphospace occupancy than species richness. As elevation increases, intraspecific variation decreases and morphologies become more similar to each other such that overall morphospace occupancy decreases. Highland skinks are, on average, smaller, thinner and shorter limbed than lowland species. 5. We hypothesise that harsh climates in the New Guinea highland habitats impose strong selection on skinks to occupy specific areas of morphospace that facilitate efficient thermoregulation in suboptimal thermal conditions. We conclude that the effect of competition on trait divergence on a community and assemblage scale is eclipsed by abiotic selection pressures in these harsh environments.

New Guinea, the world's largest and highest tropical island, has a rich but poorly known biota. Papuascincus is a genus of skinks endemic to New Guinea's mountain regions, comprising two wide-ranging species and two species known only from their type series. The phylogeny of the genus has never been examined and the relationships among its species - as well as between it and closely related taxa - are hitherto unknown. We performed the first large-scale molecular-phylogenetic study of Papuascincus, including sampling across the genus' range in Papua New Guinea. We sequenced three mitochondrial and two nuclear markers from 65 specimens of Papuascincus and reconstructed their phylogenetic relationships. We also performed species-delimitation analyses, estimated divergence times and ancestral biogeography, and examined body-size evolution within the genus. Papuascincus was strongly supported as monophyletic. It began radiating during the mid-Miocene in the area now comprising the Central Cordillera of New Guinea, then dispersed eastward colonising the Papuan Peninsula. We found evidence of extensive cryptic diversity within the genus, with between nine and 20 supported genetic lineages. These were estimated using three methods of species delimitation and predominantly occur in allopatry. Distribution and body-size divergence patterns indicated that character displacement in size took place during the evolutionary history of Papuascincus. We conclude that the genus requires comprehensive taxonomic revision and likely represents a species-rich lineage of montane skinks.

Morphology is expected to represent species' ecological niches, based on microhabitat and ecological selection pressures dictating morphological adaptations for efficient performance. However, the presence of competitor species is predicted to cause niches to contract. Therefore, an increase in species richness is expected to lead to narrower niches, and reduced overlap and distances between niches of different species. We tested these predictions on the skink fauna of New Guinea, the world's largest tropical island. We show that, while some morphospace metrics change predictably with species richness, elevation is a stronger predictor of morphospace occupancy. As elevation increases niches become narrower and closer to each other, and overall morphospace occupancy decreases. Highland skinks are, on average, smaller, thinner, and with shorter limbs than lowland species. We hypothesize that harsh climates in highland habitats impose strong selection on skinks to occupy specific areas of morphospace that facilitate efficient thermoregulation in sub-optimal thermal conditions.