Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea's Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs.

New Guinea has the most diverse insular frog fauna in the world, and rates of species discovery and description have increased rapidly in the last two decades. Pelodryadid treefrogs are the second most diverse family of anurans on the island but their taxonomy, relationships, and especially ecology remain poorly documented. Based on differences in morphology, advertisement calls (where available) and phylogenetic analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA, we describe five new species of small treefrogs from hill and lower montane forests in the high rainfall belt that straddles the southern versant of Papua New Guinea's Central Cordillera. Three of these species are known only from forest growing on karst substrates, adding to the growing number of herpetofauna species currently known only from the extensive karst habitats of Papua New Guinea's South-fold Mountains. We also describe the arboreal breeding strategies of two of the new species, and report obligate treehole (phytotelm) breeding in New Guinean frogs for the first time. The new phytotelm-breeding species has juveniles with colour and patterning that closely resemble bird droppings, suggesting defensive mimicry or masquerade. A preliminary phylogeny suggests that arboreal-breeding frogs do not form a monophyletic group and that arboreal breeding has evolved multiple times within the New Guinean pelodryadid radiation. A further striking feature of the phylogeny is poor support for most basal nodes in the most diverse radiation of Melanesian Pelodryadidae, suggesting rapid ecological diversification and speciation, potentially following colonisation from Australia and/or mountain uplift. These new taxa and observations highlight previously unrecognised ecological and reproductive diversity in the Melanesian Pelodryadidae.

Ecologically diverse island-associated lizard radiation shows idiosyncratic trait diversification shifts and homogenous speciation dynamics.

A key area of interest in evolutionary biology has been understanding the role of ecological opportunity in the formation of adaptive radiations, lineages where speciation and phenotypic diversification are driven by open ecological opportunity. Evolutionary theory posits that adaptive radiations should show initial bursts of ecomorphological diversification and rapid speciation, and that these two processes are correlated. Here, we investigate and contrast these predictions across ecomorphologically diverse continental (Australia) and insular (New Caledonia and New Zealand) radiations of diplodactyloid geckos. We test two key hypotheses: (a) that island colonization and the transition to novel niche-space has resulted in increased rates of speciation and trait diversification and (b) that rates of morphological diversification are correlated across multiple trait axes. Surprisingly, we find that speciation rate is homogenous and morphological diversification rates are idiosyncratic and uncorrelated with speciation rates. Tests of morphological integration suggests that while all traits coevolve, constraint may act differentially on individual axes. This accords with a growing number of studies indicating that ecologically diverse and species-rich radiations can show limited or no evidence of exceptional regime shifts in speciation dynamics or morphological diversification, especially in continental contexts.

Melanesia holds the world's most diverse and intact insular amphibian fauna.

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.

Cryptic extinction risk in a western Pacific lizard radiation.

Cryptic ecologies, the Wallacean Shortfall of undocumented species' geographical ranges and the Linnaean Shortfall of undescribed diversity, are all major barriers to conservation assessment. When these factors overlap with drivers of extinction risk, such as insular distributions, the number of threatened species in a region or clade may be underestimated, a situation we term 'cryptic extinction risk'. The genus Lepidodactylus is a diverse radiation of insular and arboreal geckos that occurs across the western Pacific. Previous work on Lepidodactylus showed evidence of evolutionary displacement around continental fringes, suggesting an inherent vulnerability to extinction from factors such as competition and predation. We sought to (1) comprehensively review status and threats, (2) estimate the number of undescribed species, and (3) estimate extinction risk in data deficient and candidate species, in Lepidodactylus. From our updated IUCN Red List assessment, 60% of the 58 recognized species are threatened (n = 15) or Data Deficient (n = 21), which is higher than reported for most other lizard groups. Species from the smaller and isolated Pacific islands are of greatest conservation concern, with most either threatened or Data Deficient, and all particularly vulnerable to invasive species. We estimated 32 undescribed candidate species and linear modelling predicted that an additional 18 species, among these and the data deficient species, are threatened with extinction. Focusing efforts to resolve the taxonomy and conservation status of key taxa, especially on small islands in the Pacific, is a high priority for conserving this remarkably diverse, yet poorly understood, lizard fauna. Our data highlight how cryptic ecologies and cryptic diversity combine and lead to significant underestimation of extinction risk. Supplementary Information: The online version contains supplementary material available at 10.1007/s10531-022-02412-x.

Diversity and phylogenetic relationships of haemosporidian and hemogregarine parasites in Australian lizards.

Apicomplexa is a large monophyletic phylum of unicellular, parasitic organisms. Reptiles are hosts to both haemosporidian (Haemosporida) and hemogregarine (Eucoccidiorida) apicomplexan blood parasites. Within reptiles our understanding of their diversity remains limited, with a paucity of information from Australia, despite a high diversity of squamates (snakes and lizards). We provide a preliminary assessment of haemosporidian and hemogregarine diversity occurring in lizards across northern tropical Australia, building on existing data with results from a microscopy and genetic assessment. We screened total of 233 blood slides using microscopy and detected hemogregarines in 25 geckos, 2 skinks and 1 agamid, while haemosporidians were detected in 13 geckos. DNA sequencing of 28 samples of the hemogregarine 18S rRNA (∼900 bp) nuclear gene revealed five lineages of Australian lizard hemogregarines within heteroxenous adeleids. We sequenced 10 samples of Haemosporida mtDNA (cytb & coI: ∼1313 bp) and phylogenetic analysis with 30 previously published sequences revealed that the Australian Haemosporida grouped within the Haemoproteidae but were not supported as a monophyletic clade. Our results demonstrate that there is significant undocumented evolutionary diversity in Australian lizard haemosporidian and hemogregarine parasites, with preliminary evidence of significantly higher infection rates in geckos.

Whos your daddy? On the identity and distribution of the paternal hybrid ancestor of the parthenogenetic gecko Lepidodactylus lugubris (Reptilia: Squamata: Gekkonidae).

The widespread parthenogenetic gecko Lepidodactylus lugubris is comprised of several clonal lineages, at least one of which has been known for some time to have originated from hybridization between its maternal ancestor, Lepidodactylus moestus, and a putatively undescribed paternal ancestor previously known only from remote islands in the Central Pacific. By integrating new genetic sequences from multiple studies on Lepidodactylus and incorporating new genetic sequences from previously sampled populations, we recovered a phylogenetic tree that shows a close genetic similarity between the generally hypothesized paternal hybrid ancestor and a recently described species from Maluku (Indonesia), Lepidodactylus pantai. Our results suggest that the paternal hybrid ancestor of at least one parthenogenetic clone of L. lugubris is conspecific with L. pantai and that the range of this species extends to Palau, the Caroline Islands, the Kei Islands, Wagabu, and potentially other small islands near New Guinea. Deeper genetic structure in the western (Palau, Maluku) versus eastern (eastern Melanesia, Micronesia, Polynesia) part of this species range suggests that the western populations likely dispersed via natural colonization, whereas the eastern populations may be the result of human-mediated dispersal. The potential taxonomic affinities and biogeographic history should be confirmed with further morphological and genetic analyses, including research on L. woodfordi from its type locality, which would have nomenclatural priority if found to be conspecific with L. pantai. We recommend referring to the wide-ranging sexual species as Lepidodactylus pantai until such a comparison can be made.

A new species of Cyrtodactylus (Squamata: Gekkonidae) from the northern versant of New Guineas Central Cordillera.

We describe a new species in the Cyrtodactylus loriae group from the northern versant of New Guineas Central Cordillera in far western Papua New Guinea. Cyrtodactylus hutchinsoni sp. nov. can be distinguished from related taxa by its large size, absence of serrate tubercles on the tail and ventrolateral fold, and further aspects of scalation and colouration. Genetic data suggest the new species is closely allied to C. serratus from the southern versant of the Central Cordillera of New Guinea, providing a potential example of a species pair isolated by Pliocene uplift of these mountains. The recognised diversity of Cyrtodactylus from New Guinea and other islands east of Lydekkers Line now stands at 30 species, of which 22 have been described in the last 20 years.

A return-on-investment approach for prioritization of rigorous taxonomic research needed to inform responses to the biodiversity crisis.

Global biodiversity loss is a profound consequence of human activity. Disturbingly, biodiversity loss is greater than realized because of the unknown number of undocumented species. Conservation fundamentally relies on taxonomic recognition of species, but only a fraction of biodiversity is described. Here, we provide a new quantitative approach for prioritizing rigorous taxonomic research for conservation. We implement this approach in a highly diverse vertebrate group-Australian lizards and snakes. Of 870 species assessed, we identified 282 (32.4%) with taxonomic uncertainty, of which 17.6% likely comprise undescribed species of conservation concern. We identify 24 species in need of immediate taxonomic attention to facilitate conservation. Using a broadly applicable return-on-investment framework, we demonstrate the importance of prioritizing the fundamental work of identifying species before they are lost.

The living heart: Climate gradients predict desert mountain endemism.

Mountain regions are centers of biodiversity endemism at a global scale but the role of arid-zone mountain ranges in shaping biodiversity patterns is poorly understood. Focusing on three guilds of taxa from a desert upland refugium in Australia, we sought to determine: (a) the relative extent to which climate, terrain or geological substrate predict endemism, and (b) whether patterns of endemism are complimentary across broad taxonomic guilds. We mapped regional endemism for plants, land snails, and vertebrates using combined Species Distribution Models (SDMs) for all endemic taxa (n = 82). We then modelled predictors of endemism using Generalised Additive Models (GAMs) and geology, terrain, and climate variables. We tested for the presence of inter- and intraguild hotspots of endemism. Many individual plant and land snail taxa were tightly linked with geology, corresponding to small distributions. Conversely, most vertebrate taxa were not constrained to specific geological substrates and occurred over larger areas. However, across all three guilds climate was the strongest predictor of regional endemism, particularly for plants wherein discrete hotspots of endemism were buffered from extreme summer temperatures. Land snail and vertebrate endemism peaked in areas with highest precipitation in the driest times of the year. Hotspots of endemism within each guild poorly predicted endemism in other guilds. We found an overarching signal that climatic gradients play a dominant role in the persistence of endemic taxa in an arid-zone mountain range system. An association with higher rainfall and cooler temperatures indicates that continuing trends toward hotter and drier climates may lead to range contractions in this, and potentially other, arid-zone mountain biotas. Contrasting patterns of endemism across guilds highlight the need to couple comprehensive regional planning for the protection of climate refugia, with targeted management of more localized and habitat specialist taxa.

A new species of large green treefrog (Litoria, Pelodryadidae) from Papua, Indonesia.

We describe a new species of large green treefrog from southern Papua Province, Indonesia. Litoria lubisi sp. nov. is placed in the L. infrafrenata Group based on its size, colouration and configuration of hand webbing, but differs from other members of this group in aspects of body size and proportions, extent of hand webbing, colouration and male advertisement call. Litoria lubisi sp. nov. occurs in close proximity to two other species in the L. infrafrenata Group, emphasising an until-recently overlooked pattern that multiple species of large, arboreal Litoria have overlapping distributions across much of the lowlands of southern New Guinea.

A new velvet gecko (Oedura: Diplodactylidae) from Groote Eylandt, Northern Territory.

Over the last decade, the combination of biological surveys, genetic diversity assessments and systematic research has revealed a growing number of previously unrecognised vertebrate species endemic to the Australian Monsoonal Tropics. Here we describe a new species of saxicoline velvet gecko in the Oedura marmorata complex from Groote Eylandt, a large island off the eastern edge of the Top End region of the Northern Territory. Oedura nesos sp. nov. differs from all congeners in combination of moderate size, and aspects of tail morphology and colouration. It has not been reported from the nearby mainland regions (eastern Arnhem Land) suggesting it may be an insular endemic, although further survey work is required to confirm this. While Groote Eylandt is recognised as a contemporary ecological refuge for declining mammal species of northern Australia, newly detected endemic species suggest it may also be of significance as an evolutionary refuge for many taxa, especially those associated with sandstone escarpments.

Oligocene divergence of frogmouth birds (Podargidae) across Wallace's Line.

Wallace's Line demarcates the transition between the differentiated regional faunas of Asia and Australia. However, while patterns of biotic differentiation across these two continental landmasses and the intervening island groups (Wallacea) have been extensively studied, patterns of long-term dispersal and diversification across this region are less well understood. Frogmouths (Aves: Podargidae) are a relictual family of large nocturnal birds represented by three extant genera occurring, respectively, in Asia, 'Sahul' (Australia and New Guinea) and the Solomon Islands, thus spanning Wallace's Line. We used new mitochondrial genomes from each of the extant frogmouth genera to estimate the timeline of frogmouth evolution and dispersal across Wallace's Line. Our results suggest that the three genera diverged and dispersed during the mid-Cenozoic between approximately 30 and 40 Mya. These divergences are among the oldest inferred for any trans-Wallacean vertebrate lineage. In addition, our results reveal that the monotypic Solomons frogmouth (Rigidipenna inexpectata) is one of the most phylogenetically divergent endemic bird lineages in the southwest Pacific. We suggest that the contemporary distribution of exceptionally deep divergences among extant frogmouth lineages may be explained by colonization of, and subsequent long-term persistence on, island arcs in the southwest Pacific during the Oligocene. These island arcs may have provided a pathway for biotic dispersal out of both Asia and Australia that preceded the formation of extensive emergent landmasses in Wallacea by at least 10 million years.

A new species of Lepidodactylus (Squamata: Gekkonidae) from the mountains of northeastern Papua New Guinea: older than the hills.

We describe a new species of Lepidodactylus with an unusual distribution across scattered localities in three isolated mountain ranges of northeastern New Guinea. It is a member of the Lepidodactylus pumilus group and can be distinguished from all other Melanesian Lepidodactylus by aspects of size, scalation, digital webbing, and coloration. Previously published genetic and morphological data indicate that the new species is most similar to Lepidodactylus magnus, but it diverged from this species and other close relatives in the mid-Miocene or earlier, potentially on islands of the former South Caroline Arc. Estimated divergence dates between the new species and its sister taxon suggest that cladogenesis occurred before the uplift of the mountains that they currently inhabit. Recent systematic work also emphasizes an apparent pattern of increasing body size with elevation in species from the Lepidodactylus pumilus group from northeastern New Guinea, with the largest species occurring in montane habitats where few or no other gecko species are known. Both lines of evidence are consistent with an existing hypothesis that the Lepidodactylus pumilus group is an old insular lineage within which contemporary species diversity and distributions have been strongly shaped by low ability to effectively compete against other aggressive geckos in species-rich lowland rainforests. The role that biotic interactions may have played in shaping tropical gecko communities along elevational gradients warrants further investigation.

A new species of Cyrtodactylus Gray (Gekkonidae: Squamata) from Manus Island, and extended description and range extension for Cyrtodactylus sermowaiensis (De Rooij).

Systematic investigations of vertebrate faunas from the islands of Melanesia are revealing high levels of endemism, dynamic biogeographic histories, and in some cases surprisingly long evolutionary histories of insularity. The bent-toed geckos in the Cyrtodactylus sermowaiensis Group mainly occur in northern New Guinea and nearby islands, however a further isolated population occurs on Manus Island in the Admiralty Archipelago approximately 300 km to the north of New Guinea. Here we first present a review of the genetic diversity, morphological variation and distribution of Cyrtodactylus sermowaiensis from northern New Guinea. Genetic structure and distributional records within Cyrtodactylus sermowaiensis broadly overlap with underlying Terranes in northern New Guinea, suggesting divergence on former islands that have been obscured by the infill and uplift of sedimentary basins since the late Pleistocene. Based on a combination of genetic and morphological differentiation we then describe the population from Manus Island as a new species, Cyrtodactylus crustulus sp. nov. This new species emphasises the high biological endemism and conservation significance of the Admiralty Islands, and especially Manus Island.

Crypsis and convergence: integrative taxonomic revision of the Gehyra australis group (Squamata: Gekkonidae) from northern Australia.

For over two decades, assessments of geographic variation in mtDNA and small numbers of nuclear loci have revealed morphologically similar, but genetically divergent, intraspecific lineages in lizards from around the world. Subsequent morphological analyses often find subtle corresponding diagnostic characters to support the distinctiveness of lineages, but occasionally do not. In recent years it has become increasingly possible to survey geographic variation by sequencing thousands of loci, enabling more rigorous assessment of species boundaries across morphologically similar lineages. Here we take this approach, adding new, geographically extensive SNP data to existing mtDNA and exon capture datasets for the Gehyra australis and G. koira species complexes of gecko from northern Australia. The combination of exon-based phylogenetics with dense spatial sampling of mitochondrial DNA sequencing, SNP-based tests for introgression at lineage boundaries and newly-collected morphological evidence supports the recognition of nine species, six of which are newly described here. Detection of discrete genetic clusters using new SNP data was especially convincing where candidate taxa were continuously sampled across their distributions up to and across geographic boundaries with analyses revealing no admixture. Some species defined herein appear to be truly cryptic, showing little, if any, diagnostic morphological variation. As these SNP-based approaches are progressively applied, and with all due conservatism, we can expect to see a substantial improvement in our ability to delineate and name cryptic species, especially in taxa for which previous approaches have struggled to resolve taxonomic boundaries.

A new species of Cyrtodactylus (Squamata: Gekkonidae) from central New Guinea.

We describe a new species of bent-toed gecko in the genus Cyrtodactylus from hill forest in Southern Highlands Province, Papua New Guinea. Cyrtodactylus manos sp. nov. can be distinguished from all congeners in New Guinea by its small size in combination with aspects of colouration and body and tail scalation. The new species adds to the growing number of vertebrate species known only from karstic mountains along the southern edge of New Guinea's Central Cordillera, suggesting that this region holds previously overlooked endemic karst-associated biota.

Two new species of treefrog (Pelodrydidae: Litoria) from southern New Guinea elucidated by DNA barcoding.

New Guinea is home to the world's most diverse insular frog biota, but only a small number of taxa have been included in genetically informed assessments of species diversity. Here we describe two new species of New Guinea treefrog in the genus Litoria that were first flagged during assessments of genetic diversity (DNA barcoding) and are currently only known from the holotypes. Litoria pterodactyla sp. nov. is a large green species in the Litoria graminea species complex from hill forests in Western Province, Papua New Guinea and is the third member of this group known from south of the Central Cordillera. Litoria vivissimia sp. nov. is a small, spike-nosed species from mid-montane forests on the Central Cordillera. It is morphologically very similar to Litoria pronimia, but occurs nearly 1000 m higher than any known locality for that species. More extensive genetically informed assessment of diversity in New Guinea frogs seems certain to reveal many more as-yet-unrecognised taxa in complexes of morphologically similar species.

Systematics of New Guinea treefrogs (Litoria: Pelodryadidae) with erectile rostral spikes: an extended description of Litoria pronimia and a new species from the Foja Mountains.

A small number of treefrog species (Litoria) from Melanesia are unusual amongst Anura in having distinctive fleshy rostral spikes. Here, we first present an extended description for Litoria pronimia Menzies, a small species that is widespread along the southern edge of the Central Cordillera of New Guinea, and in which males have a long and erectile rostral spike. Second, we describe Litoria pinocchio sp. nov. a new, morphologically similar, yet geographically disjunct species from the Foja Mountains in northern Papua Province, Indonesia. The new species differs from Litoria pronimia in aspects of body shape, proportions and colouration. A review of variation in the size, structure and degree of sexual dimorphism of the rostral spike across different species of Litoria suggests varying function including mate selection and camouflage.

Relicts and radiations: Phylogenomics of an Australasian lizard clade with east Gondwanan origins (Gekkota: Diplodactyloidea).

Australasia harbors very high squamate diversity and is a center of endemicity for a number of major lineages. However, despite this diversity, the diplodactyloid geckos of Australia, New Caledonia, and New Zealand (comprised of three endemic families and >200 species) are the only extant squamates with unequivocal Mesozoic origins in the region. Diplodactyloid geckos also exhibit notable phenotypic and ecological diversity, most strikingly illustrated by the functionally limbless pygopods. Here, we present the first phylogenomic analyses of the pattern and timing of diplodactyloid evolution, based on a dataset of more than 4000 ultraconserved elements (UCEs) from 180 species. These analyses fully resolve nearly all nodes, including a number of intergeneric relationships that have proven problematic in previous studies. The hypothesis that New Caledonia and New Zealand clades represent independent post-KT boundary colonization events of Tasmantis from Australian ancestors is confirmed. Phylogenetic relationships recovered here further highlight contrasting patterns of diversity, most strikingly between insular and/or morphologically highly derived clades that have diversified rapidly, as opposed to other species poor and phylogenetically divergent relictual lineages on mainland Australia. Our new timetree suggests slightly older branching times than previous analyses and does not find a mass extinction event in the early Cenozoic. Finally, our new phylogeny highlights caudal variation across the clade. Most strikingly, the distinctive leaf-tail morphology shown by one family may in fact be plesiomorphic.