Stephen J Richards

South Australian Museum · Herpetology

Despite recent advances in the systematics and taxonomy of the relictual family Scolebythidae, the type genus Scolebythus Evans, 1963 remained monospecific for more than sixty years. Here, I describe and illustrate Scolebythus bekilyensis sp. nov. from a female specimen collected in southern Madagascar. This discovery emphasizes on the endemism of...

We describe a new species of Cyrtodactylus from the northern lowlands and foothills of mainland New Guinea. Cyrtodactylus mamberamo sp. nov. is distinguished from all other Melanesian Cyrtodactylus except C. aaroni and C. mimikanus by the combination of moderate size (max SVL <100 mm), widened subcaudals, dorsal pattern of numerous narrow light ban...

We describe two new species of torrent-breeding Litoria Tschudi, 1838 from low-elevation hill-forest habitats on the southern fringe of Papua New Guinea’s Central Cordillera. One is currently known only from the Kikori River basin, and the other is known from the Kikori and adjacent Strickland River basins. The two new species can be distinguished...

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-...

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...

New Guinea is the largest tropical island in the world and hosts immense endemic biodiversity. However, our understanding of how the gradual emergence of the terrestrial ecosystems of the island over the last 40 Myr has generated this biological richness is hampered by poorly documented species diversity and distributions. Here, we address both the...

Five new species of Metagrion are described from Papua New Guinea: Metagrion annehueberae sp. nov. (holotype ♂ in RMNH), Metagrion pseudolamprostoma sp. nov. (holotype ♂ in SAMA), Metagrion furcatum sp. nov. (holotype ♂ in SAMA), Metagrion lopaui sp. nov. (holotype ♂ in RMNH), Metagrion toxopeus sp. nov. (holotype ♂ in SAMA). New distribution recor...

Rainforests on high tropical mountains are globally important species diversity hotspots. We studied amphibians along an extensive rainforest elevation gradient on Mt. Wilhelm (4509 m) in Papua New Guinea. We established eight sites at 500 m elevation increments between 200 and 3700 m a.s.l. and relate their community composition to the known speci...

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 Inter...

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 Inter...

We examined the morphology, colour patterns and genetic relationships of Nososticta populations allied to N. salomonis (Selys) from across Melanesia. Seven species-level taxa are recognised in the N. salomonis ‘complex’: N. africana (Schmidt), N. boonei sp. nov., N. chrismulleri Theischinger & Richards, N. hedigeri sp. nov., N. salomonis (Selys), N...

We describe two new species of moderate-sized (male body length 26.5–29.8 mm and 41.0 mm), predominantly green treefrogs in the genus Litoria from hill forest on the northern slopes of Papua New Guinea’s Central Cordillera. Phylogenetic analysis of mitochondrial ND4 nucleotide sequences shows that the first species is related to Litoria iris (Tyler...

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...

A new arboreal species of the microhylid genus Cophixalus Boettger, 1892 is described from montane rainforest on Papua New Guinea’s central cordillera. With a male SUL exceeding 44.0 mm, the new species is among the largest members of the genus; the only other Papuan species known to reach this size is C. riparius Zweifel, 1962. The new species dif...

We describe a new species of very small (male body length 22.0–22.8 mm), green treefrog in the pelodryadid genus Litoria Tschudi from lowland rainforest on the southern margin of Papua New Guinea's central cordillera. The new species is morphologically most similar to Litoria leucova, a small green treefrog known only from the north-flowing Sepik R...

The microhylid genus Oreophryne reaches its greatest diversity in the New Guinea region, where more than 60 species have been documented to date. Most Oreophryne are small (<30 mm SVL) and only three species, O. anthonyi, O. idenburgensis and O. inor-nata, exceed 40 mm SVL adult body size. Here we describe a fourth large species of Oreophryne that...

The mountains of New Guinea are home to species-rich but poorly understood communities of stream or torrent-breeding pelodryadid treefrogs. Here we describe a new species of moderately sized torrent-breeding Litoria from the mountains of Papua Province, Indonesia. The new species is most similar to Litoria dorsivena but differs from that species in...

The Islands of East Melanesia have a unique and highly endemic frog fauna derived entirely from overseas colonisation events. Within East Melanesia New Britain is a notable centre of frog diversity and endemism, with at least 15 endemic species, mostly in the ceratobatrachid genus Cornufer . Here we describe the first endemic pelodryadid treefrog f...

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 specie...

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 undere...

We describe a new species of the asterophryine microhylid genus Xenorhina from the lowlands of northwestern Papua New Guinea. It is a medium-sized species (SUL of two males 29.2 and 29.9 mm; of four females 29.9–33.0 mm) that can be distinguished from congeners by having a single short, triangular odontoid spike (palatal tooth) on each vomeropalati...

We describe six new species of the microhylid frog genus Xenorhina from the southern slopes of Papua New Guinea’s central cordillera and adjacent lowlands, based on a combination of morphological (including osteology) and bioacoustic features. All of the new species are fossorial or terrestrial inhabitants of tropical rainforest habitats and belong...

Australia and New Guinea (together referred to as Sahul) were linked by land for much of the late Tertiary and share many biotic elements. However, New Guinea is dominated by rainforest, and northern Australia by savannah. Resolving patterns of biotic interchange between these two regions is critical to understanding the expansion and contraction o...

We show that the Wotjulum frog, Litoria watjulumensis (Copland, 1957), comprises two deeply divergent mitochondrial DNA lineages that are also reciprocally monophyletic for a nuclear gene locus and have discrete distributions. The taxa are differentiated in multivariate analysis of shape but show no appreciable differences in colour and pattern. Th...

We describe a new species of groundsnake of the genus Stegonotus (Colubridae) from the Purari River basin in Gulf Province, Papua New Guinea. The new species can be most readily distinguished from all other New Guinean Stegonotus by its unique dorsal colour pattern which consists of a dark head and creamy-white anterior one third to two thirds of t...

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...

We describe a new species in the Australopapuan pelodryadid frog genus Litoria from upper hill forest (940 m a.s.l.) on the northern slopes of Papua New Guinea’s central cordillera. The new species is moderately small (male body length = 31.9–35.1 mm) and slender (head width/body length = 0.29–0.30), with extensive golden-yellow markings ventrally....

We describe a new species of gecko in the Lepidodactylus novaeguineae Group from Salawati Island, West Papua Province, Indonesia. The new species can be distinguished from all congeners by a unique combination of aspects of body size, shape, colouration, and scalation. The holotype and only known specimen is a mature male with a snout-vent length o...

We describe two new species in the microhylid frog genus Copiula from Western Province, Southern Highlands Province, and Gulf Province in southern Papua New Guinea based on morphological and acoustic data. Both species are medium-sized (between 27 – 30 mm SVL), terrestrial frogs with a predominantly beige-colored dorsum and a yellow ventrum. They c...

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...

New Guinea has been considered both as a refuge for mesic rainforest-associated lineages that contracted in response to the late Cenozoic aridification of Australia and as a centre of biotic diversification and radiation since the mid-Miocene or earlier. Here, we estimate the diversity and a phylogeny for the Australo-Papuan forest dragons (Sauria:...

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 nu...

The amphibian chytrid fungus Batrachochytrium dendrobatidis (chytrid) has caused the most widespread, disease‐induced declines and extinctions in vertebrates recorded to date. The largest climatically suitable landmass that may still be free of this fungus is New Guinea. The island is home to a sizeable proportion of the world's known frog species...

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...

A new species of the dicroglossid frog genus Limnonectes is described from recent and historical museum specimens collected in central and southern Laos and northeastern Thailand. Limnonectessavansp. nov. has males that bear a caruncle on top of the head, and most closely resembles L.dabanus from adjacent southern Vietnam and eastern Cambodia. Howe...

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 erectil...

Bironides ypsilon sp. nov. and Nannophlebia ballerina sp. nov. from PNG 331 Odonatologica 46(3/4) Abstract. Two new species of stream-dwelling libellulid dragonflies are described from Gulf Province, Papua New Guinea. Bironides ypsilon sp. nov. and Nannophlebia ballerina sp. nov. are both small black and yellowish green dragonflies that appear to b...

A new species of damselfly, Rhyacocnemis gassmanni sp. n. from Papua New Guinea, is described and illustrated from both sexes with notes on its habitat and habits. It represents the fourth species of an enigmatic genus, known from only a handful of specimens. The placement of the new species is problematical and is discussed. Introduction The zygop...

Two new species of the genus Idiocnemis Selys, 1886 from southern Papua New Guinea are described: Idiocnemis lakekamuensis sp. nov. from the Lakekamu Basin and I. milou sp. nov. from Lakekamu and the Kikori River lowlands. Males and females are illustrated and compared with other species of the Idiocnemis bidentata group. Both new species are known...

A series of photographs of the recently described Star Mountains Worm-eating Snake, Toxicocalamus ernstmayri O'Shea et al., 2015, taken at the Ok Tedi Mine in the Star Mountains, North Fly District, Western Province, Papua New Guinea, represents only the second record of this poorly-known species. Toxicocalamus ernstmayri was hitherto only known fr...

Two new microhylid frogs in the genera Choerophryne and Oreophryne are described from the Foja Mountains in Papua Province of Indonesia. Both are small species (males 15.9 - 18.5 mm snout-urostyle length [SUL] and 21.3 - 22.9 mm SUL respectively) that call from elevated positions on foliage in primary lower montane rainforest. The new Choerophryne...

Papua New Guinea is an independent nation that occupies the eastern half of New Guinea, the world's largest tropical island. New Guinea has an exceptionally high biodiversity, and a large proportion of its fauna and flora is found nowhere else on Earth. Because the PNG LNG project was built and operates in an area with high biodiversity values that...

The published repository for the holotype of Pseudagrion Jumipennis Polhemus, Michalski & Richards, 2008, is corrected to the South Australian Museum, Adelaide, Australia, not the 'Australian Museum of Natural History, Sydney' as stated in the original description. © 2018 Entomological Society of Queensland. All rights resrerved.

A new species of damsel fly, Nososticta moginae sp. n. from Gulf Province, Papua New Guinea, is described from both sexes and its affinities are discussed. It represents the 80th species of the genus and is currently known only from a single location in the Kikori River Basin, A description of the hitherto unknown female of the morphologically simi...

Regions with complex geological histories present a major challenge for scientists studying the processes that have shaped their biotas. The history of the vast and biologically rich tropical island of New Guinea is particularly complex and poorly resolved. Competing geological models propose New Guinea emerged as a substantial landmass either duri...

We describe a new species of Cornufer, subgenus Batrachylodes, from high-elevation forests of New Britain Island in the Bismarck Archipelago of Eastern Melanesia. The new species, Cornufer exedrus sp. nov., is a biogeographically disjunct member of the Batrachylodes clade, representing the first record of the subgenus from outside of the Solomon Ar...

To ensure that the name Papuargia stueberi luciedecknerae, described (2017) in Odonatologica 41: 283-291, is available, the type repository, inadequately presented in the original description, is stated along with a diagnosis of the subspecies.

The island of New Guinea has been identified as biologically megadiverse but many taxa are still poorly known. This is especially the case for many of the island’s snakes, which by their very nature can be difficult to collect and study. Here we examine the phylogenetic and phylogeographic structure of a poorly studied snake genus, Stegonotus, focu...

A second species of the hitherto monotypic New Guinean genus Papuargia Lief- tinck, 1938 and a new subspecies of P. stueberi Lieftinck, 1938 are described from Papua New Guinea. These are P. brevistigma sp. nov. and P. stueberi luciedecknerae ssp. nov. The char- acters which define the genus are discussed with special reference to the labium and pe...

We describe three new species of the microhylid frog genus Choerophryne from the mountains and foothills of southern and northeastern Papua New Guinea. All three species lack elongated snouts and all are arboreal, calling from elevated perch sites between ~1 and 10 m above the forest floor. Advertisement calls and habitat preferences are described...

Aims Mountain ranges in the tropics are characterised by high levels of localised endemism, often-aberrant evolutionary trajectories, and some of the world’s most diverse regional biotas. Here we investigate the evolution of montane endemism, ecology and body size in a clade of direct-developing frogs ( Choerophryne, Microhylidae) from New Guinea....

Sequence alignment

Appendix S3 Treefile for chronogram estimated in BEAST with ancestral states.

Appendix S1 Supplementary tables and figures. Specimen numbers, locality information and GenBank accession numbers for Choerophryne specimens included in analyses (Table S1); GenBank accession details for outgroup samples (Table S2); genetic distance data for species and candidate lineages (Table S3); and summary data on body, elevational distribut...

Appendix S2 BEAST input file for ancestral state analyses

Aims. Mountain ranges in the tropics are characterised by high levels of localised endemism, often-aberrant evolutionary trajectories, and some of the world’s most diverse regional biotas. Here we investigate the evolution of montane endemism, ecology and body size in a clade of direct-developing frogs ( Choerophryne, Microhylidae) from New Guinea....

Aims. Mountain ranges in the tropics are characterised by high levels of localised endemism, often-aberrant evolutionary trajectories, and some of the world’s most diverse regional biotas. Here we investigate the evolution of montane endemism, ecology and body size in a clade of direct-developing frogs ( Choerophryne, Microhylidae) from New Guinea....

Aims. Mountain ranges in the tropics are characterised by high levels of localised endemism, often-aberrant evolutionary trajectories, and some of the world’s most diverse regional biotas. Here we investigate the evolution of montane endemism, ecology and body size in a clade of direct-developing frogs ( Choerophryne, Microhylidae) from New Guinea....

The East Melanesian Islands have been a focal area for research into island biogeography and community ecology. However, previously undescribed and biogeographically significant new species endemic to this region continue to be discovered. Here we describe a phylogenetically distinct (~20% divergence at the mitochondrial ND2 gene) and biogeographic...

Gynacantha nuda sp. nov. is described based on a male from Southern Highlands Province, Papua New Guinea. It is the largest known species of the genus to be reported from New Guinea. Characters of the adult male are illustrated, the affinities of the new species are discussed, and a key is presented to males of the Gynacantha species recorded from...

Two new species of the microhylid genus Oreophryne are described on the basis of material collected from the Kikori River Basin in southern Papua New Guinea. Both species belong to the group of Oreophryne that have a ligamentous connection between the procoracoid and scapula. One of the new species is small (snout-urostyle length in males 19.7 - 23...

Idiocnemis schorri sp. nov. is described from the Hindenburg and Muller Ranges and the Kikori River Basin of southern Papua New Guinea. The new species differs from all congeners by, among other characters, a unique colour pattern on the thorax. Characters of males and females are illustrated and compared to those of similar species from the Idiocn...

We describe a new species of treefrog from northern Australia. Litoria bella sp. nov. is morphologically and genetically most similar to frogs in the L. gracilenta and L. chloris groups but is distinguished from all members in these groups by a combination of a moderately large male body size (34.5-41.8 mm SVL), near-immaculate green dorsum, orange...

We describe a striking new species of the microhylid frog genus Mantophryne from Woodlark Island in Milne Bay Province, Papua New Guinea. It is most similar to M. lateralis but is distinguished from that species by its more slender body, longer shanks, larger discs on the toes, and unique advertisement call. Most known specimens had, in life, a str...

Pseudagrion woodlarkensis sp. nov., a new damselfly species from Woodlark Island, Papua New Guinea, is described and male and female characters are illustrated. The new species differs from all regional congeners by having a predominantly yellow thorax with bold black stripes, and males can be clearly distinguished from all Papuan Pseudagrion speci...

The diverse biota of New Guinea includes many nominally widespread species that actually comprise multiple deeply divergent lineages with more localised histories of evolution. Here we investigate the systematics of the very large geckos of the Cyrtodactylus novaeguineae complex using molecular and morphological data. These data reveal two widespre...