Ongoing declines for the world's amphibians in the face of emerging threats.

Systematic assessments of species extinction risk at regular intervals are necessary for informing conservation action1,2. Ongoing developments in taxonomy, threatening processes and research further underscore the need for reassessment3,4. Here we report the findings of the second Global Amphibian Assessment, evaluating 8,011 species for the International Union for Conservation of Nature Red List of Threatened Species. We find that amphibians are the most threatened vertebrate class (40.7% of species are globally threatened). The updated Red List Index shows that the status of amphibians is deteriorating globally, particularly for salamanders and in the Neotropics. Disease and habitat loss drove 91% of status deteriorations between 1980 and 2004. Ongoing and projected climate change effects are now of increasing concern, driving 39% of status deteriorations since 2004, followed by habitat loss (37%). Although signs of species recoveries incentivize immediate conservation action, scaled-up investment is urgently needed to reverse the current trends.

A new forest frog of the genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Tirahanulap) from Leyte and Samar islands, eastern Philippines.

We describe a new species of frog of the genus Platymantis Günther (subgenus Tirahanulap), from the east-central regions of the Philippines. It belongs to the the previously-defined P. hazelae Group) based on morphological and bioacoustic datasets. The new species is phenotypically and ecologically most similar to members of Tirahanulap, an assemblage of small-bodied arboreal frogs inhabiting montane forests of the central and northern islands of the Philippine archipelago. The new species represents the first taxon in the Cloud Frog species known from the biogeographically unique Mindanao Pleistocene Aggregate Island Complex. Particularly susceptible to local extirpation following deforestation, all known species of Tirahanulap are important indicator species for environmental and conservation assessments, making this new species not only an exceptional addition to Philippine biodiversity but also an important symbol for conservation initiatives in the region.

A new Liopeltis Fitzinger, 1843 (Squamata: Colubridae) from Pulau Tioman, Peninsular Malaysia.

Liopeltis is a genus of poorly known, infrequently sampled species of colubrid snakes in tropical Asia. We collected a specimen of Liopeltis from Pulau Tioman, Peninsular Malaysia, that superficially resembled L. philippina, a rare species that is endemic to the Palawan Pleistocene Aggregate Island Complex, western Philippines. We analyzed morphological and mitochondrial DNA sequence data from the Pulau Tioman specimen and found distinct differences to L. philippina and all other congeners. On the basis of these corroborated lines of evidence, the Pulau Tioman specimen is described as a new species, L. tiomanica sp. nov. The new species occurs in sympatry with L. tricolor on Pulau Tioman, and our description of L. tiomanica sp. nov. brings the number of endemic amphibians and reptiles on Pulau Tioman to 12.

Niche shifts and environmental non-equilibrium undermine the usefulness of ecological niche models for invasion risk assessments.

Niche shifts and environmental non-equilibrium in invading alien species undermine niche-based predictions of alien species' potential distributions and, consequently, their usefulness for invasion risk assessments. Here, we compared the realized climatic niches of four alien amphibian species (Hylarana erythraea, Rhinella marina, Hoplobatrachus rugulosus, and Kaloula pulchra) in their native and Philippine-invaded ranges to investigate niche changes that have unfolded during their invasion and, with this, assessed the extent of niche conservatism and environmental equilibrium. We investigated how niche changes affected reciprocal transferability of ecological niche models (ENMs) calibrated using data from the species' native and Philippine-invaded ranges, and both ranges combined. We found varying levels of niche change across the species' realized climatic niches in the Philippines: climatic niche shift for H. rugulosus; niche conservatism for R. marina and K. pulchra; environmental non-equilibrium in the Philippine-invaded range for all species; and environmental non-equilibrium in the native range or adaptive changes post-introduction for all species except H. erythraea. Niche changes undermined the reciprocal transferability of ENMs calibrated using native and Philippine-invaded range data. Our paper highlights the difficulty of predicting potential distributions given niche shifts and environmental non-equilibrium; we suggest calibrating ENMs with data from species' combined native and invaded ranges, and to regularly reassess niche changes and recalibrate ENMs as species' invasions progress.

The curious case of the endemic freshwater crocodile Crocodylus mindorensis as incidental host of marine fish acanthocephalan.

We performed the first host-parasite survey of the Philippine crocodile, Crocodylus mindorensis, a critically endangered species for which ecological information is lacking. We collected by gastric lavage samples of the stomach contents of crocodiles (n = 10) residing at the Palawan Wildlife Rescue and Conservation Center in Puerto Princesa, Palawan, Philippines. The only parasite detected was an acanthocephalan, which was identified as Neorhadinorhynchus nudus (n = 68), a parasite typically found in the marine fish species consumed by three crocodile individuals. Given the known hosts of N. nudus, its parasitism of C. mindorensis in captivity is likely established by consumption of marine fish. Our findings have implications for the conservation management of C. mindorensis, particularly in terms of preventing introduction of parasites that could lead to development of infectious disease or alter the fitness of captive animals.

There is no evidence that Podoctidae carry eggs of their own species: Reply to Machado and Wolff (2017).

In our recent publication (Sharma et al., 2017), we tested the hypothesis that eggs attached to the legs of male Podoctidae (Opiliones, Laniatores) constituted a case of paternal care, using molecular sequence data in tandem with multiple sequence alignments to test the prediction that sequences of the eggs and the adults that carried them would indicate conspecific identity. We discovered that the sequences of the eggs belonged to spiders, and thus rejected the paternal care hypothesis for these species. Machado and Wolff (2017) recently critiqued our work, which they regarded as a non-critical interpretation and over-reliance on molecular sequence data, and defended the traditional argument that the eggs attached to podoctids are in fact harvestman eggs. Here we show that additional molecular sequence data also refute the identity of the eggs as conspecific harvestman eggs, using molecular cloning techniques to rule out contamination. We show that individual gene trees consistently and reliably place the egg and adult sequences in disparate parts of the tree topology. Phylogenetic methods consistently place all egg sequences within the order Araneae (spiders). We submit that evidence for the paternal care hypothesis based on behavioral, morphological, and natural history approaches is either absent or insufficient for concluding that the eggs of podoctids are conspecific.

A multilocus phylogeny of Podoctidae (Arachnida, Opiliones, Laniatores) and parametric shape analysis reveal the disutility of subfamilial nomenclature in armored harvestman systematics.

The taxonomy and systematics of the armored harvestmen (suborder Laniatores) are based on various sets of morphological characters pertaining to shape, armature, pedipalpal setation, and the number of articles of the walking leg tarsi. Few studies have tested the validity of these historical character systems in a comprehensive way, with reference to an independent data class, i.e., molecular sequence data. We examined as a test case the systematics of Podoctidae, a family distributed throughout the Indo-Pacific. We tested the validity of the three subfamilies of Podoctidae using a five-locus phylogeny, and examined the evolution of dorsal shape as a proxy for taxonomic utility, using parametric shape analysis. Here we show that two of the three subfamilies, Ibaloniinae and Podoctinae, are non-monophyletic, with the third subfamily, Erecananinae, recovered as non-monophyletic in a subset of analyses. Various genera were also recovered as non-monophyletic. As first steps toward revision of Podoctidae, the subfamilies Erecananinae Roewer, 1912 and Ibaloniinae Roewer, 1912 are synonymized with Podoctinae Roewer, 1912 new synonymies, thereby abolishing unsubstantiated subfamilial divisions within Podoctidae. We once again synonymize the genus Paralomanius Goodnight & Goodnight, 1948 with Lomanius Roewer, 1923 revalidated. We additionally show that eggs carried on the legs of male Podoctidae are not conspecific to the males, falsifying the hypothesis of paternal care in this group.

The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands.

We summarize all available amphibian and reptile species distribution data from the northeast Mindanao faunal region, including small islands associated with this subcenter of endemic vertebrate biodiversity. Together with all publicly available historical information from biodiversity repositories, we present new data from several major herpetological surveys, including recently conducted inventories on four major mountains of northeast Mindanao, and adjacent islands of Camiguin Sur, Dinagat, and Siargao. We present species accounts for all taxa, comment on unresolved taxonomic problems, and provide revisions to outdated IUCN conservation status assessments in cases where our new data significantly alter earlier classification status summaries. Together, our comprehensive analysis of this fauna suggests that the greater Mindanao faunal region possesses distinct subcenters of amphibian and reptile species diversity, and that until this area is revisited and its fauna and actually studied, with on-the-ground field work including targeted surveys of species distributions coupled to the study their natural history, our understanding of the diversity and conservation status of southern Philippine herpetological fauna will remain incomplete. Nevertheless, the northeast Mindanao geographical area (Caraga Region) appears to have the highest herpetological species diversity (at least 126 species) of any comparably-sized Philippine faunal subregion.

A new species of Pseudogekko (Squamata: Gekkonidae) from the Romblon Island Group, Central Philippines.

We describe a new species of lizard in the genus Pseudogekko from Sibuyan and Tablas islands in the Romblon Island Group of the central Philippines. The new species is diagnosed from other Philippine Pseudogekko by body size and shape, color pattern, and multiple differences in scale characteristics. Pseudogekko isapa sp. nov. has been collected only twice from leaves of shrubs in forested habitat on Sibuyan and Tablas islands. The distinctive new species of false gecko is undoubtedly endemic to this single, isolated island group. The fact that populations of such a distinctive new species of Pseudogekko has escaped notice of herpetologists on the reasonably well-studied and largely protected Sibuyan Island further emphasizes the secretive and forest-dependent habits of Philippine false geckos. These characteristics of their behavior and natural history render them difficult to study and challenge biologists' efforts to accurately assess their conservation status.

Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) II: a new species from the northern Philippines.

We describe a new digitless scincid lizard of the genus Brachymeles from northern Luzon and Camiguin Norte islands in the Philippines. This species belongs to the Brachymeles bonitae Complex, and both molecular and morphological data confirm that this species is distinct from all other congeners. Formerly considered to be a single widespread species, this group of species has been the focus of recent systematic reviews. Here we describe a new species in the B. bonitae Complex, recognized currently to constitute five species. Brachymeles ilocandia sp. nov. is the second digitless and the seventeenth non-pentadactyl species in genus. The description of this species brings the total number of species in the genus to 40, and provides new insight into unique distribution patterns of species of the northern Philippines.

Phylogeny of the island archipelago frog genus Sanguirana: Another endemic Philippine radiation that diversified 'Out-of-Palawan'.

Recent higher-level frog phylogenetic analyses have included a few members of the endemic Philippine frog genus Sanguirana. Although the monophyly of the group has never been disputed, the recent phylogenetically-supported inclusion of the Palawan Wood Frog (Sanguirana sanguinea) in this clade was highly unexpected. In addition, species boundaries and relationships remain unclear and new species continue to be discovered. We estimate the phylogeny for this endemic Philippine genus using two mitochondrial gene regions and six nuclear loci and complete sampling for all known species. We use a time-calibrated Bayesian estimate of phylogeny and model-testing approach to biogeographic inference to infer ancestral areas and probable means of diversification. These analyses identify Sanguirana as an additional clade for which the 'Out-of-Palawan' biogeographic scenario is unambiguously preferred. This study lends additional support to recent work suggesting that a substantial portion of Philippine vertebrate megadiversity originated via colonization of the archipelago from the Palawan microcontinent, with subsequent invasion of oceanic islands (e.g., range expansion over Huxley's Modification of Wallace's Line), numerous instances of overwater dispersal, and geographic radiation across the archipelago.

A new species of limestone karst inhabiting forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Lupacolus) from southern Luzon Island, Philippines.

We describe a new species of limestone karst dwelling forest frog of the genus Platymantis from the Quezon Protected Landscape in southeastern Luzon Island, Philippines. We assign Platymantis quezoni, sp. nov., to the diverse assemblage of terrestrial species in the Platymantis dorsalis Group, subgenus Lupacolus on the basis of its body size and proportions, only slightly expanded terminal discs of the fingers and toes, and its terrestrial microhabitat. The new species is distinguished from these and all other Philippine congeners by features of its external morphology, its restriction to a distinctive limestone karst microhabitat, and its advertisement call, which is unique among frogs of the family Ceratobatrachidae. Several distinguishing morphological characters include its moderate body size (22.1-33.9 mm SVL for 16 adult males and 32.4-39.7 mm SVL for five adult females), slightly expanded terminal discs of the fingers and toes, smooth skin with limited dermal tuberculation, and a dorsal color pattern of mottled tan to dark brown with black blotches. The new species is the sixth Philippine Platymantis known to occur exclusively on limestone karst substrates (previously known karst-obligate species include: P. bayani, P. biak, P. insulatus, P. paengi, and P. speleaus). Recently accelerated discovery of limestone karst anurans across the Philippines suggests that numerous additional species may await discovery on the hundreds of scattered karst formations throughout the archipelago. This possibility suggests that a major conservation priority in coming years will be to study, characterize, describe, and preserve the endemic species supported by this patchy, unique and imperiled type of forest ecosystem in the Philippines.

Redescription of the rare Philippine false gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species.

Recent investigations into the species diversity of false geckos (genus Pseudogekko Taylor) have revealed several cryptic species, highlighting the need for a more thorough understanding of diversity within this enigmatic genus of endemic Philippine geckos. Newly available genetic data reveal that two of the four currently recognized species are complexes of multiple deeply divergent evolutionary lineages. In this paper we evaluate species diversity in one of these complexes, P. brevipes Boettger, and describe one additional new species. For nearly a century, P. brevipes has been recognized as a single, "widespread" species with a geographic range spanning two major faunal regions and several island groups. Poor understanding of this species has persisted due to both limited sampling and its apparent rarity. We evaluate both morphological and genetic data to define species limits in P. brevipes, and find character-based evidence to justify the recognition of two unique evolutionary lineages, one of which we describe as a new species (P. atiorum sp. nov.). The species included in this study have allopatric distributions and differ from congeners by numerous diagnostic characters of external morphology, and therefore should be recognized as full species in accordance with lineage-based species concepts. This newly described species increases the total number of species of Pseudogekko to seven.

Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos (Reptilia: Squamata: Scincidae), with description of three new species.

We review the recent discovery of multiple populations of the enigmatic, semi-aquatic Sphenomorphus Group skink, Parvoscincus leucospilos Peters, and investigate the morphological and genetic diversity of isolated, allopatric populations of this unique skink. Our investigations support the recognition of four unique evolutionary lineages distributed across Luzon Island in the Philippines, three of which are herein described as new species (P. tikbalangi sp. nov., P. manananggalae sp. nov., and P. duwendorum sp. nov.). All four recognized species are genetically divergent in both mitochondrial and nuclear DNA sequences, and morphologically distinct. The description of three new Luzon Island endemic species adds to the growing body of literature suggesting that mechanisms driving the accumulation of vertebrate diversity in the Philippines may vary regionally across the archipelago. 

Conservation genetics of the Philippine tarsier: cryptic genetic variation restructures conservation priorities for an island archipelago primate.

Establishment of conservation priorities for primates is a particular concern in the island archipelagos of Southeast Asia, where rates of habitat destruction are among the highest in the world. Conservation programs require knowledge of taxonomic diversity to ensure success. The Philippine tarsier is a flagship species that promotes environmental awareness and a thriving ecotourism economy in the Philippines. However, assessment of its conservation status has been impeded by taxonomic uncertainty, a paucity of field studies, and a lack of vouchered specimens and genetic samples available for study in biodiversity repositories. Consequently, conservation priorities are unclear. In this study we use mitochondrial and nuclear DNA to empirically infer geographic partitioning of genetic variation and to identify evolutionarily distinct lineages for conservation action. The distribution of Philippine tarsier genetic diversity is neither congruent with expectations based on biogeographical patterns documented in other Philippine vertebrates, nor does it agree with the most recent Philippine tarsier taxonomic arrangement. We identify three principal evolutionary lineages that do not correspond to the currently recognized subspecies, highlight the discovery of a novel cryptic and range-restricted subcenter of genetic variation in an unanticipated part of the archipelago, and identify additional geographically structured genetic variation that should be the focus of future studies and conservation action. Conservation of this flagship species necessitates establishment of protected areas and targeted conservation programs within the range of each genetically distinct variant of the Philippine tarsier.

Archipelago colonization by ecologically dissimilar amphibians: evaluating the expectation of common evolutionary history of geographical diffusion in co-distributed rainforest tree frogs in islands of Southeast Asia.

Widespread, co-distributed species with limited relative dispersal abilities represent compelling focal taxa for comparative phylogeography. Forest vertebrates in island archipelagos often exhibit pronounced population structure resulting from limited dispersal abilities or capacity to overcome marine barriers to dispersal. The exceptionally diverse Old World tree frogs of the family Rhacophoridae have colonized the forested island archipelagos of Southeast Asia on multiple occasions, entering the islands of Indonesia and the Philippines via a "stepping stone" mode of dispersal along elongate island chains, separated by a series of marine channels. Here we evaluate the prediction that two tightly co-distributed Philippine rhacophorids colonized the archipelago during concomitant timescales and in the same, linear, "island-hopping" progression. We use a new multilocus dataset, utilize dense genetic sampling from the eastern arc of the Philippines, and we take a model-based phylogeographic approach to examining the two species for similar topological patterns of diversification, genetic structure, and timescales of diversification. Our results support some common mechanistic predictions (a general south-to-north polarity of colonization) but not others (timescale for colonization and manner and degree of lineage diversification), suggesting differing biogeographic scenarios of geographical diffusion through the archipelago and unique and idiosyncratic ecological capacities and evolutionary histories of each species.

The challenge of species delimitation at the extremes: diversification without morphological change in philippine sun skinks.

An accurate understanding of species diversity is essential to studies across a wide range of biological subdisciplines. However, delimiting species remains challenging in evolutionary radiations where morphological diversification is rapid and accompanied by little genetic differentiation or when genetic lineage divergence is not accompanied by morphological change. We investigate the utility of a variety of recently developed approaches to examine genetic and morphological diversity, and delimit species in a morphologically conserved group of Southeast Asian lizards. We find that species diversity is vastly underestimated in this unique evolutionary radiation, and find an extreme case where extensive genetic divergence among lineages has been accompanied by little to no differentiation in external morphology. Although we note that different conclusions can be drawn when species are delimited using molecular phylogenetics, coalescent-based methods, or morphological data, it is clear that the use of a pluralistic approach leads to a more comprehensive appraisal of biodiversity, and greater appreciation for processes of diversification in this biologically important geographic region. Similarly, our approach demonstrates how recently developed methodologies can be used to obtain robust estimates of species limits in "nonadaptive" or "cryptic" evolutionary radiations.

An adaptive radiation of frogs in a southeast Asian island archipelago.

Living amphibians exhibit a diversity of ecologies, life histories, and species-rich lineages that offers opportunities for studies of adaptive radiation. We characterize a diverse clade of frogs (Kaloula, Microhylidae) in the Philippine island archipelago as an example of an adaptive radiation into three primary habitat specialists or ecotypes. We use a novel phylogenetic estimate for this clade to evaluate the tempo of lineage accumulation and morphological diversification. Because species-level phylogenetic estimates for Philippine Kaloula are lacking, we employ dense population sampling to determine the appropriate evolutionary lineages for diversification analyses. We explicitly take phylogenetic uncertainty into account when calculating diversification and disparification statistics and fitting models of diversification. Following dispersal to the Philippines from Southeast Asia, Kaloula radiated rapidly into several well-supported clades. Morphological variation within Kaloula is partly explained by ecotype and accumulated at high levels during this radiation, including within ecotypes. We pinpoint an axis of morphospace related directly to climbing and digging behaviors and find patterns of phenotypic evolution suggestive of ecological opportunity with partitioning into distinct habitat specialists. We conclude by discussing the components of phenotypic diversity that are likely important in amphibian adaptive radiations.

The amphibians and reptiles of Luzon Island, Philippines, VIII: the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre Mountain Range.

We provide the first report on the herpetological biodiversity (amphibians and reptiles) of the northern Sierra Madre Mountain Range (Cagayan and Isabela provinces), northeast Luzon Island, Philippines. New data from extensive previously unpublished surveys in the Municipalities of Gonzaga, Gattaran, Lasam, Santa Ana, and Baggao (Cagayan Province), as well as fieldwork in the Municipalities of Cabagan, San Mariano, and Palanan (Isabela Province), combined with all available historical museum records, suggest this region is quite diverse. Our new data indicate that at least 101 species are present (29 amphibians, 30 lizards, 35 snakes, two freshwater turtles, three marine turtles, and two crocodilians) and now represented with well-documented records and/or voucher specimens, confirmed in institutional biodiversity repositories. A high percentage of Philippine endemic species constitute the local fauna (approximately 70%). The results of this and other recent studies signify that the herpetological diversity of the northern Philippines is far more diverse than previously imagined. Thirty-eight percent of our recorded species are associated with unresolved taxonomic issues (suspected new species or species complexes in need of taxonomic partitioning). This suggests that despite past and present efforts to comprehensively characterize the fauna, the herpetological biodiversity of the northern Philippines is still substantially underestimated and warranting of further study.