Australasian Journal of Herpetology

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online) ISSUEISSUE 44,44, PUBLISHEDPUBLISHED 55 JUNEJUNE 20202020 2 Australasian Journal of Herpetology Australasian Journal of Herpetology 44-46:1-192. Published 5 June 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

For the first time ever! An overdue review and reclassification of Australasian Tree Frogs (Amphibia: Anura: Pelodryadidae), including formal descriptions of 12 tribes, 11 subtribes, 34 genera, 26 subgenera, 62 species and 12 subspecies new to science.

LSIDURN:LSID:ZOOBANK.ORG:PUB:0A14E077-B8E2-444E-B036-620B311D358D

RAYMOND T. HOSER LSIDURN:LSID:ZOOBANK.ORG:AUTHOR:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 4 November 2019, Accepted 2 June 2020, Published 5 June 2020.

ABSTRACT For the past 200 years most, if not all Australian Tree Frogs have been treated as being in a single genus. For many years this was Hyla Laurenti, 1768, before the genus name Litoria Tschudi, 1838 was adopted by Cogger et al. (1983) and has been in general use by most herpetologists since, including Cogger (2014). Tyler and Davies (1978) divided the putative genus Litoria into 37 “species groups” and this type of classification has been used by numerous authors since, including most recently Menzies (2006) for the New Guinea species and Anstis (2014) for the Australian ones. By merging all previous molecular and morphological studies with systematic scientific methodology, including a review of all previous species and subspecies descriptions, including those of synonymised forms, all significant past papers on the group and inspection of thousands of specimens, this paper takes the logical next step of completing an overdue full family-wide reclassification of the Australiasian Tree Frogs in the Pelodryadidae. Besides resurrecting dozens of old and available names for genera and species when applicable, this paper also formally names 12 tribes, 11 subtribes, 34 genera, 26 subgenera, 62 species and 12 subspecies for the first time in accordance with the rules of the International Code of Zoological Nomenclature (Ride et al. 1999). It remains self-evident that the species-level diversity of the group remains underestimated, especially in New Guinea. In the face of ongoing rapid, exponential human population growth in Australia and New Guinea, habitat destruction, introduced pests and pathogens, the survival threat to many species within the Pelodryadidae has never been greater. This increases the urgency to formally identify relevant taxa in order to be able to conserve them. Keywords: Taxonomy; Tree Frog; Amphibia; nomenclature; Frog; Pelodryadidae; Australia; New Guinea; Hyla; Litoria; Ranoidea; Cyclorana; Chirodryas; Nyctimystes; Dryopsophus; Pelodryas; Drymomantis; Euscelis; Mitrolysis; Brendanura; Coggerdonia; Colleeneremia; Llewellynura; Mahonabatrachus; Mosleyia; Neophractops; Pengilleyia; Rawlinsonia; Saganura; Sandyrana; new tribe; Adelynhoserhyleini; Coggerdoniani; Cycloranini; Daraninanurini; Fiacumminganurini; Maxinehoserranini; Nyctimystini; Kumanjayiwalkerini; Pelodryanini; Pustulataranini; Saganurini; Wowranaini; new subtribes; Leucodigiranina; Ranoideina; Gedyerananina; Dryopsophina; Drymomantina; Badiohylina; Audaxurina; Rawlinsonina; Shireenhoserhylina; Salmocularanina; Sandyranina; new genus; Adelynhoserhylea; Jackyhoserhylea; Leucodigirana; Crottyrana; Gedyerana; Daraninanura; Fiacumminganurea; Maxinehoserranae; Angularanta; Bellarana; Fluvirana; Hopviridi; Incertanura; Inlustanura; Moechaeanura; Ornatanura; Nasuscuspis; Rotundaura; Variabilanura; Albogibba; Occultatahyla; Nigreosoculus; Badiohyla; Magnumoculus; Kumanjayiwalkerus; Audaxura; Brevicrusyla; Shireenhoserhylea; Summaviridis; Pustulatarana; Salmocularana; Paralitoria; Quasilitoria; Wowrana; new subgenus; Yikesanura; Paramitrolysis; Invisibiliaauris; Sandgroperanura; Amnisrana; Leucolatera; Ausverdarana; Vegrandihyla; Alliuma; Longuscrusanura; Naveosrana; Raucus; Scelerisqueanura; Aspercutis; Telaater; Sudesanura; Magnummanibus; Asperohyla; Ratiobrunneis; Webpede; Balatusrana; Emeraldhyla; Microlitoria; Ferelitoria; Vultusamolitoria; Parawowrana; new species; adelynhoserae; yikes; ernieswilei; jackyhoserae; flavoranae; leucodorsalinea; rosea; crottyi; sloppi; gedyei; cottoni; michaelsmythi; piloti; fiacummingae; timdalei; jarrodthomsoni; brettbarnetti; maxinehoserae; piersoni; chydaeus; communia; extentacrus; mukherjii; quaeinfernas; vulgarans; oxyeei; cuspis; fakfakensis; inluster; albatermacula; tritong; spica; leucopicturas; parscinereo; parsviridis; tomcottoni; ausviridis; celantur; northstradbrokensis; mondoensis; charlottae; doggettae; aspera; georgefloydi; ingens; kumanjayi; bogfrog; chunda; dunnyseat; watdat; wifi; ventrileuco; megaviridis; fukker; yehbwudda; chriswilliamsi; marionanstisae; pailsae; roypailsi; saxacola; mickpughi; mippughae; new subspecies; dorsaruber; occultatum; dumptrashensis; inornata; gippslandensis; leucopunctata; brunetus; toowoombaensis; tozerensis; davidtribei; dunphyi; serventyi.

Presented herein is a revision of the Pelodryadidae. A full list of recognized tribes, genera and species is presented at the end of this paper in alphabetical order, with page references for all descriptions and redescriptions of tribes, subtribes, genera and subgenera as well as for newly named species and subspecies. The paper is published in three volumes, of Australasian Journal of Herpetology, being Issues 44-46, all published on 5 June 2020.

INTRODUCTION For most of the past 200 years, most, if not all Australian per the doctrine of Kaiser et al. 2013). and New Guinea Tree Frogs have been treated as being However the unscientific edicts of Kaiser et al. were in a single genus. effectively ignored by Eipper and Rowland (2018). For many years this was Hyla Laurenti, 1768, before the Instead they heeded the advice of Hoser (2007) and genus name Litoria Tschudi, 1838 was adopted by published an amended classification of Australian Tree Cogger et al. (1983) and has been in general use by Frogs that was essentially similar to that of Wells and most herpetologists since, including Cogger (2014). Wellington (1985) by relying on recently published While a number of inveterate and obsessive name molecular phylogenies. grabbers from the 1800’s, including the usual suspects of While relying on the published phylogenies of Duellman Cope, Peters, Günther and Fitzinger put new genus et al. (2016) and Bell et al. (2017) that they drew their names on several species and without a shred of conclusions from and using the available names, scientific basis to do so, most of those names went into including many of those first proposed by Wells and synonymy. Wellington (1985), the Eipper and Rowland (2018 Because a number are objective synonyms of one taxonomy presented had obvious gaps and deficiencies, another, this is where a lot must remain. including genera they simply identified as “unnamed”. However two of the the more morphologically divergent Further to that, generic assignments made in line with species groups within the Australasian Tree Frog either Wells and Wellington (1985) or Duellman et al. radiation were given different genus names, namely (2016) at times simply did not correlate with the relevant Cyclorana Steindachner, 1867 and Nyctimystes published phylogenies. Stejneger, 1916 and both names have remained in use While I note that the published phylogenies of Frost et by herpetologists to the present day, including for al. (2006), Pyron et al. (2011), Duellman et al. (2016) and example by Menzies (2006), Venderduys (2012), Anstis Bell et al. (2017) have clearly confirmed most of the (2013), Cogger (2014) and Eipper and Rowland (2018). Wells and Wellington classification as correct, but not all In terms of the many dozens of other recognized species of it, these cited publications have highlighted a number of Australasian Tree Frogs (being over 200 on most of other well-defined and divergent lineages that by any species lists as of 2020), almost all have been simply reasonable analysis should be placed in their own, as yet lumped into the genus Litoria sensu Cogger et al. (1983) unnamed genera. by most authors since that date, without a shred of With this in mind a review of the relevant species, scientific justification. including all species groups from Australia and New As of 2020, this is the current state of play in terms of the Guinea was conducted. taxonomy of Australian tree frogs so far as most of the Specimens and literature were reviewed to find a superior Australian public is aware. taxonomy for the Australian and New Guinea Tree Frogs Unfortunately the same in fact applies to most of the in order to get a genus level taxonomy and nomenclature “scientific community”. that can be used by other herpetologists as a framework Tyler and Davies (1978) divided the putative genus Litoria for further research. into 37 “species groups”. Unnamed tribe and genus-level groupings identified in This was a reasonable start to the dismemberment of the the process, have been formally named in accordance genus Litoria and from that point on the formal naming of with the rules of the International Code of Zoological these species groups should have been completed within Nomenclature (Ride et al. 1999). a few short years. Obviously divergent forms are also formally named as After Tyler and Davies (1978), Wells and Wellington new species for the first time. This being a total of 62 (1985) decided to place obvious phylogenetically related species and 12 new subspecies. groups within this massive putative genus of ancient MATERIALS AND METHODS origins into obvious morphologically distinctive genera. These are inferred in both the abstract and introduction Acting ethically, these authors resurrected seven names and self evident in the descriptions that follow. from synonymy and erected another 11 putative genera An audit of all previously described species of for obvious well-known species groups for the first time. Australasian Tree Frogs in the family Pelodryadidae A group known as the Wolfgang Wüster gang of thieves Günther, 1858 was conducted. (see Hoser 2007, Hoser 2015a-f and 2019a-b) have In simple terms this included all species lumped in the successfully harassed most publishing herpetologists to genera Litoria Tschudi, 1838, Cyclorana Steindachner, ignore the taxonomy of Wells and Wellington and to 1867, Nyctimystes Stejneger, 1916, which has for most simply synonymise all genera into a greater Litoria (as Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 4 Australasian Journal of Herpetology publishing authors in the past 50 years been the sum 2016b), Barbour (1908, 1912, 1921), Barker and Grigg total of the family. (1977), Barker et al. (1995), Bell et al. (2012, 2017), In the period post-dating Wells and Wellington (1985), in Bevelander (2014), Boettger (1895, 1900), Böhme particular post-dating the paper of Duellman et al. (2016), (2014), Böhme and Bischoff (1984), Boulenger (1882, it has been increasingly difficult for authors to justify the 1883, 1887a, 1887b, 1892, 1896, 1897a, 1897b, 1898, use of just three genus names for all the Pelodryadidae 1905, 1911, 1912, 1914, 1915), Brongersma (1953), and some earlier names have been resurrected and used Brown (1952), Burns (2004), Burns and Crayn (2006), on online databases and publications. Burt and Burt (1932), Capocaccia (1957), Clyne (1969), However, as a rule, these published lists of names have Cogger (1966, 1979, 2014), Cogger et al. (1983), Condit not made sense as the species are often placed in (1964), Cope (1866, 1867), Copland (1957, 1960, 1951, genera in which the type species is not remotely related 1963a, 1963b), Courtice and Grigg (1975), Covacevich to the assigned species, giving further reason for a major (1974), Coventry (1970), Czechura et al. (1987), Daan in-depth review of the family, the results of which are and Hillenius (1966), Daudin (1802, 1803), Davies et al. published herein. (1983, 1986), De la Riva et al. (2014), Dennis and It has also been self evident that often the authors Cunningham (2006), De Vis (1884), Donnellan and assigning species to their supposedly authoritative Mahony (2004), Donnellan et al. (1999), Doughty (2011), “genus and species list” clearly have no idea about Doughty and Anstis (2007), Dubois (1984, 2005, 2007, classification and rules of nomenclature and invariably 2018), Dubois and Frétey (2016), Dubois et al. (2019), have assigned species to genera with little if any Duellman (1977, 1993), Duellman and Trueb (1986), connection to the originally described type form. Duellman et al. (2016), Duméril (1853), Duméril, and Bibron (1841), Duméril and Duméril (1851), Eipper Published species lists were checked against the (2012), Eipper and Rowland (2018), Faivovitch et al. relevant literature (e.g. Menzies 2006, Cogger et al. (2005), Fitzinger (1826, 1860), Fletcher (1898), Forcart 2013, Tyler 1968, Anstis 2013, etc) and online databases (1953), Frost (2013), Frost et al. (2006), Fry (1912, 1913, such as “Amphibia Web” and “Amphibian Species of the 1915), Gadow (1901), Gassó Miracle et al. (2007), World” hosted on the American Museum of Natural Gillespie (2001, 2002, 2004, 2010, 2011, 2012), Gillespie History website, with those listed species in this paper and Hollis (1996), Gillespie et al. (2015), Goldman et al. being treated as valid, either on the basis of prior (1969), Gray (1841, 1842, 1848), Guibé (1948), Günther descriptions in the scientific literature, or new (1858, 1863a, 1863b, 1864, 1867, 1873, 1876, 1897), descriptions herein. Günther (2003, 2006a, 2006b, 2006c, 2008, 2014), When possible, specimens of all relevant species Günther (2003, 2004a, 2004b, 2006a, 2006b, 2006c, (named and until now unnamed) were examined both live 2008, 2014), Günther and Richards (2000, 2005), Hiaso in the wild and via museum collections and their records, and Richards (2006), Hoser (1989, 2013b, 2014, 2015g, including all State and Territory Museums on mainland 2018a, 2018b, 2018c, 2018d, 2018e, 2020a, 2020b, Australia and some collections held outside of Australia 2020c, 2020d, 2020e), Hoskin (2007), Hosmer (1964), as well. Furthermore photos and information with Hunter (2012), Hunter and Smith (2013), Hunter et al. accurate locality data was also assessed, as was all (2011), Ingram and Corben (1990), Ingram et al. (1992, relevant previously published scientific literature and the 1993), James (1998) James and Moritz (2000), Johnston so-called grey literature in the form of popular mass- and Richards (1994), Keferstein (1867), Kraus (2007, market books, internet sites, blogs, photo-sharing sites 2009, 2010, 2012a, 2012b, 2013a, 2013b, 2018), Kraus and the like. and Allison (2004a, 2004b, 2009), Krell and Marshall Scrutinized in particular was taxonomic studies and (2017), Lamb (1911), Laurance et al. (1996), Lesson phylogenies that uncovered relationships between (1829), Leunis (1844), Liem (1974a, 1974b), Liem and species and species groups, although I note that a lot of Ingram (1977), Lönnberg (1900), Loveridge (1945, 1948, this is self-evident to someone who has spent a lifetime 1950), Macleay (1879), Mahony (2001), Main (1965), working on these animals. Martin et al. (1978), McDonald (1997), McDonald et al. Not all type material was inspected, however I was (2016), McDowell (1969), Méhely (1897), Menzies (1969, generally able to ascertain relevant details from the 1972, 1976, 1993, 2006, 2014a, 2014b), Menzies and published literature, including each and every original Johnston (2015), Menzies and Tippett (1976), Menzies description which I read (including foreign language (non and Tyler (2004), Menzies and Zug (1979), Menzies and English) ones which I had translated). Zweifel (1974, 1976), Menzies et al. (2009), Merrem Before a decision is made to name any new taxon at any (1820), Mertens (1922, 1930, 1964, 1967), Meyer (1874, level, including genus, subgenus, species or subspecies, 1887), Meyer and Agnew (2013), Minister for the reasonable steps must be taken to ensure that it is Environment, Commonwealth Government of Australia justified on all relevant grounds, including that it is (2017), Moore (1961), Murray and Hose (2005), Nieden morphologically, genetically and reproductively (1923), Noble (1931), Obst (1977), Ogilby (1890, 1907), sufficiently diverged to warrant the erection of a new Oken (1816), Oliver and Richards (2007), Oliver et al. taxonomic grouping over and above what is already (2007, 2008, 2019a, 2019b,), Parker (1936, 1938, 1940), available and in common usage. Péron (1807), Peters (1863, 1867, 1869, 1871, 1873a, Key literature relevant to the taxonomic and 1873b, 1874, 1877, 1878, 1880, 1882), Peters and Doria nomenclatural conclusions within this paper include (1878), Pyron and Wiens (2011), Ramsay (1878), Regan Ahl (1929, 1935), Andersson (1916), Anstis (2013), (2002), Rensch (1936), Reynolds (2007), Richards Anstis and Tyler (2005), Anstis et al. (1998, 2010, 2016a, (1992, 2001, 2002, 2005, 2007a, 2007b), Richards and

Alford (2005), Richards and Iskander (2001, 2006), RESULTS Richards and Johnston (1993), Richards and Oliver The published phylogenies of Frost et al. (2006), Pyron et (2006a, 2006b), Richards et al. (2006, 2009), Ride et al. al. (2011), Duellman et al. (2016) and Bell et al. (2017), (1999), Robinson (1993), Rosauer et al. (2009), Rowley clearly indicated that Australasian Tree Frogs should be and Cutajar (2018), Savage (1986), Schneider (1799), placed in their own family as done by Wells and Scott (1942), Shaw (1802), Shea and Kraus (2007), Shea Wellington (1985) and as done in the title of this paper. and Sadlier (1999), Sonnini de Manoncourt and Latreille I note that Pyron et al. (2011) referred to the relevant (1801), Spencer (1896, 1901), Steindachner (1867), group of genera as “Pelodryadinae”, which is almost the Stejneger (1916), Straughan (1969), Stuart et al. (2008), same thing and clearly indicates that the thinking of Wells Thomson et al. (1996), Tschudi (1838), Tyler (1962, and Wellington (1985), some 35 years ago, was not out 1963a, 1963b, 1964a, 1964b, 1964c, 1965, 1967 1968a, of whack with the peer reviewed scientific reality or 1968b, 1968c, 1969, 1971, 1985, 1992), Tyler and Anstis consensus as is often claimed by the Wolfgang Wüster (1975, 1983), Tyler and Davies (1977, 1978, 1979, 1983, gang of thieves, including via their war cry blog Kaiser et 1985, 1986), Tyler and Dobson (1973), Tyler and Knight al. (2013). (2009), Tyler and Martin (1977), Tyler and Parker (1972, The published phylogenies also indicated a large number 1974), Tyler et al. (1972, 1977, 1978a, 1978b, 1981, of divergent lineages, which in the main coincided with 1982, 1986, 1994), Van Beurden and McDonald (1980), the treatment by Wells and Wellington (1985). Vanderduys (2012), Van Kampen (1906, 1909, 1919, However the coincidence was not exact. 1923), Van Tuijl (1995), Vogt (1912), Wagler (1830), Two of the Wells and Wellington generic placements Wandolleck (1910, 1911), Watson et al. (1971, 1991), were not justified on the basis of the phylogeny of Pyron Weijola (2020), Wells and Wellington (1985), Werner and Weins (2011), but see below. (1898, 1901), White et al. (1994), White (1970), Wichmann (1912), Wiens et al. (2010), Withers (1993, Furthermore other unnamed lineages beyond those 1995, 1998), Woodruff (1972), Zweifel (1956, 1958, identified or named by Wells and Wellington (1985) were 1960, 1980, 1983) and sources cited therein (duplicitous clearly apparent. references not necessarily included). The phylogenies of Frost et al. (2006), Pyron et al. (2011), Duellman et al. (2016) and Bell et al. (2017) also As already mentioned, live and dead specimens as well had significant differences in parts (e.g. for the putative as available bone specimens, were examined as was species “Litoria inermis”), which immediately drew other relevant material, including past climate data for the attention to potential errors in their analysing of material. relevant regions, sea level depths, and other relevant Before assigning species to generic groups, it was information, the latter being importany aids in important to cross-check the phylogenies with the frogs establishing divergences between relevant forms and/or themselves to ensure that they matched and that likely gene flow between populations currently disjunct. divergences in published phylogenies reflected the It goes without saying that at the present time, even after physical differences as well. publication of this paper there remain numerous Before mentioning generic assignments for relevant undescribed species within the Australia and New species groups, I note that Cogger et al. (1983) had Guinea areas. published a synonyms list to that date and Wells and I also note that, notwithstanding the theft of relevant Wellington added 12 more names to the collection, all materials from this author in an illegal armed raid on 17 bar one of which were confirmed by myself as properly August 2011, which were not returned in breach of applied to unnamed species groups as determined undertakings to the court (Court of Appeal Victoria 2014 herein based on the phylogeny of Pyron and Weins and VCAT 2015), I have made a decision to publish this (2011) combined with that of Duellman et al. (2016). paper. I note that Wells and Wellington published at a time when This is in view of the conservation significance attached mtDNA and the like were not available to herpetologists to the formal recognition of unnamed taxa at all levels to more accurately determine relationships between and on the basis that further delays may in fact put these putative taxa. presently unnamed or potentially improperly assigned As an arbitrary line, I recognized as genera, all taxa at greater risk of extinction as outlined by Hoser phylogenetic groups that could reasonably be estimated (2019a, 2019b). to have diverged from nearest common ancestors, well This comment is made noting the extensive increase in over ten million years back. human population in Australia and New Guinea, with a This was all 12 of 12 Wells and Wellington groups and 8 conservative forecast of a four-fold increase in human previously named genera, as well as those newly named population in the next 100 years (from 25 million to 100 herein (see below). million) in Australia and an even more dramatic increase Because one of the genera named by Wells and in New Guinea (both sides) and the general Wellington (1985) was a duplicate name for another environmental destruction across the continental region earlier name, which they overlooked, the Wells and as documented by Hoser (1991), including low density Wellington name is simply synonymised. areas without a large permanent human population. When the divergence date number is close to 10 MYA, I I also note the abysmal environmental record of various have opted for subgenus-level division if the relevant Australian National, State and Local governments in the species are morphologically divergent from one another, relevant part of the Australasian region over the past 200 but in most cases not split the relevant genus. years as detailed by Hoser (1989, 1991, 1993 and 1996). It explains for example, why the species Hyla moorei Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 6 Australasian Journal of Herpetology

Copeland, 1957 is formally partly removed from Neophractops Wells and Wellington, 1985, type species Ranoidea Tschudi, 1838, type species Ranoidea Chiroleptes platycephalus Günther, 1873. jacksionensis Tschudi, 1838 = Rana aurea Lesson, 1831 Pengilleyia Wells and Wellington, 1985, type species and placed in the subgenus Sandgroperanura subgen. Litoria tyleri Martin, Watson, Gartside, Littlejohn and nov.. Loftus-Hills, 1979. Burns (2004) and Burns and Crayn (2006) used Rawlinsonia Wells and Wellington, 1985, type species molecular data to find that the two taxa diverged from one Hyla ewingi Duméril and Bibron, 1841. another 10 to 10.8 MYA and Duellman et al. (2016) found Sagunura Wells and Wellington, 1985, type species Hyla a divergence of 12.0 MYA for the same groups. burrowsi Scott, 1942. While two generic groupings of Wells and Wellington Sandyrana Wells and Wellington, 1985, type species (1985) associated with the genus Cyclorana Hyla infrafrenata Günther, 1867. Steindachner, 1867, type species Alytes australis Gray, I note in passing that just two of the genera erected by 1842, were not strongly supported by the phylogeny of Wells and Wellington in 1985 were even close to the 10 Pyron and Weins (2011) as being worthy of genus-level MYA divergence minimum I have arbitrarily set as separation, that of Deullman et al. (2016), with date defining generic assignments. calibrations placed both the relevant groups as diverging These are Neophractops Wells and Wellington, 1985, 11 and 12 MYA from nearest common ancestors and so type species Chiroleptes platycephalus Günther, 1873 they are also recognized herein as full genera. and Brendanura Wells and Wellington, 1985, type However the genus Brendanura Wells and Wellington, species Chiroleptes alboguttatus Günther, 1867, both 1985, type species Chiroleptes alboguttatus Günther, previously synonymised with Cyclorana Steindachner, 1867 was named in error as it is in fact an objective junior 1867. synonym of Mitrolysis Cope, 1889, with the same type The genus Euscelis Fitzinger, 1843 was not recognized species, so the earlier name is used instead. by Wells and Wellington (1985), those species being To make things abundantly clear, there is no cirsumtance placed by them within Litoria. within the rules of the International Code of Zoological (Other Wells and Wellington, 1985 genera were shown Nomenclature (Ride et al. 1999), that the name by Duellman et al. (2016) to have diverged from nearest Brendanura Wells and Wellington, 1985 can or should be other named genera well over 10 MYA). used. The species composition of these above genera is In summary the twenty extant genera (by name) in date generally as put by Wells and Wellington (1985) and order of priority recognized herein are as follows: Eipper and Rowland (2018), allowing for newly described Litoria Tschudi, 1838, type species L. freycineti Tschudi, forms, except where otherwise indicated in this paper and 1838. the synonymies as indicated. Ranoidea Tschudi, 1838, type species Ranoidea The species groups that are placed in each newly jacksionensis Tschudi, 1838 = Rana aurea Lesson, 1831. erected genus are often indicated in the original Dryopsophus Fitzinger, 1843, type species “Hyla citropa descriptions themselves even without reference to this Peron” = Hyla citropa Dümeril and Bibron, 1841. paper, although all genera are formally defined or Euscelis Fitzinger, 1843, type species Hyla lesueurii redefined herein for the pre-existing named groups. Dümeril and Bibron, 1841. This was absolutely necessary as, with very few Pelodryas Günther, 1858, type species Rana caerulea exceptions, these genera have been wholly redefined as White, 1790. per previous concepts in use, including those of Wells Cyclorana Steindachner, 1867, type species Alytes and Wellington. australis Gray, 1842. The published phylogenies, including for example Frost Chirodryas Keferstein, 1867, type species Chirodryas et al. (2006), Pyron et al. (2011), Duellman et al. (2016) raniformis Keferstein, 1867. and Bell et al. (2017), wholly corroborate the division of Drymomantis Peters, 1882, type species Drymomantis tribes, subtribes, genera and subgenera within this paper fallax Peters, 1882. and provide an added basis for divisions on the basis of Mitrolysis Cope, 1889, type species Chiroleptes calibrated divergence dates. alboguttatus Günther, 1867. Divergent forms formally named as new species herein Nyctimystes Stejneger, 1916, type species Nyctimantis for the first time are also placed in the relevant proper papua Boulenger, 1897. genus at the same time. In other words, for the first time ever and in a single Coggerdonia Wells and Wellington, 1985, type species paper, this paper formally describes each and every tribe, Hyla adelaidensis Gray, 1841. subtribe, genus and subgenus within the Pelodryadidae. Colleeneremia Wells and Wellington, 1985, type species In terms of the final taxonomic determinations, from Hyla rubella, Gray, 1842. which all relevant nomenclature follows, I note the Llewellynura Wells and Wellington, 1985, type species following important points. Hyla dorsalis microbelos Cogger, 1966. Species recognized herein are in the main widely Mahonabatrachus Wells and Wellington, 1985, type recognized and acknowledged and reflect those taxa species Hyla meriana Tyler, 1969. seen in the most recent regional field guides such as Mosleyia Wells and Wellington, 1985, type species Hyla Cogger (2014), Anstis (2013), Eipper and Rowland nannotis Andersson, 1916. (2018) or Menzies (2006), save for more recently

described and named forms which are also included from one another more than 20 MYA. herein. These somewhat arbitrary divisions are more The justification and basis for naming the new species conservative (ancient) than adopted by a number of other and subspecies is either self evident or alternatively recent authors, but even with this extreme conservatism, explained within the relevant descriptions. I was able to split the Pelodryadidae into 12 tribes and 11 I need to note that some of the newly named forms within subtribes. this paper are separated from other species with minimal The use of tribes in classification is important as it keeps morphological divergence, which on the surface may relevant species groups contained and is a superior appear unjustified. means of dealing with large groups of like genera within a I also note that a number of species resurrected from speciose family such as Pelodryadidae. synonymy without specific comment in this paper, appear The order of treatment of taxa in this paper is to have been resurrected from synonymy from forms they alphabetical, but by use of the tribe level of classification, are also often morphologically similar or near identical to. closely related genera and species are dealt with In both these situations, when they occur in this paper, an together in effectively monophyletic groups. added and critically important basis for the recognition of The use of tribes to divide specious families such as the these taxa is that in every case the relevant populations Pelodryadidae would be a major improvement in popular are split across biogeographical barriers of known books and field guides as it would greatly assist in putting antiquity, confirming the taxonomically distinct nature of accounts of like species together and not interrupted by each population and warranting species-level division. accounts for wholly dissimilar taxa. One such example that arises in this paper on a number Besides resurrecting dozens of old and available names of occasions are species pairs split across the central for genera and species when applicable, this paper also cordillera of New Guinea, which have evidently diverged formally names 12 tribes, 11 subtribes, 34 genera, 26 from one another for millions of years. subgenera, 62 species and 12 subspecies for the first Although the nature and antiquity of the relevant time in accordance with the rules of the International biogeographical barriers may not be spelt out in the Code of Zoological Nomenclature (Ride et al. 1999). relevant descriptions in this paper or resurrection of taxa FORMAL DESIGNATION OF A LECTOTYPE IN from synonymy in the species lists herein, they are self ACCORDANCE WITH THE INTERNATIONAL CODE OF evident when distributions of the relevant taxa are ZOOLOGICAL NOMENCLATURE. mapped out. Chiroleptes alboguttatus Günther, 1867 was originally The preceding is important as it needs to be made known described with two syntypes being from Bowen and Cape that all taxonomic decisions herein have been made with York in Queensland. a body of evidence supporting them and also been These specimens are themselves morphologically subjected to peer review from no less than four other divergent and so as to remove confusion as to which experts in the field of amphibian herpetology. taxon or specimen the name Chiroleptes alboguttatus Each and every species within this paper has been Günther, 1867 should be applied to, I herein make the scrutinized in detail and with the benefit of all relevant Cape York specimen (BMNH 1947.2.20.6-7) as the published material before being determined as valid and lectotype. at what level of classification, as well as their genus-level The syntype from Bowen in Queensland (BMNH assignment based on previously published phylogenies 1947.2.18.50-51) herein ceases to have nomenclatural and the morphological evidence from the species status. themselves. This action is taken in accordance with paragraph 6, and RELEVANT NUMBERS Article 74, including all of 74.1 and 74.73, of the Species were determined as being such based on the International Code of Zoological Nomenclature (Ride et usual criteria of reproductive isolation from nearest living al. 1999), including this being an “express statement of common ancestor and on the basis that this isolation is the taxonomic purpose of the designation”, this being to ancient. remove potential doubt as to which subspecies or When the duration of such isolation couldn’t be species should retain the name alboguttatus Günther, established with certainty of being more than 1.5 MYA, or 1867 on the basis of divergence, either morphological or reasonably assumed not to be, then obviously divergent molecular between the two syntype specimens. taxa were conservatively identified as subspecies. INFORMATION RELEVANT TO THE FORMAL Genera were arranged based on common morphology DESCRIPTIONS THAT FOLLOW and divergence. There is no conflict of interest in terms of this paper or As a rule, groups of species diverged well over 10 MYA the conclusions arrived at herein. were placed in their own genera, this being in line with Several people including anonymous peer reviewers who similar divisions for other groups of reptiles and frogs in revised the manuscript prior to publication are also recent reclassifications. thanked as are relevant staff at museums who made Species groups that were morphologically divergent, but specimens and records available in line with international with divergence around the 10 MYA mark were generally obligations. placed in subgenera. In terms of the following formal descriptions of tribes, At the higher level, tribes were generally determined to subtribes, genera, subgenera, species or subspecies, exist for genera that diverged from one another more spellings should not be altered in any way for any than 25 MYA and subtribes those genera that diverged Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 8 Australasian Journal of Herpetology purpose unless expressly and exclusively called for by species or subspecies is readily and consistently the rules governing Zoological Nomenclature as separable from other similar taxon or group as indicated administered by the International Commission of and that which until now the relevant newly named group Zoological Nomenclature. or form have been treated as being within. In the unlikely event two newly named tribes, subtribes, Delays in recognition of these unique taxa could genera, subgenera, species or subspecies are deemed jeopardise the long-term survival of these taxa as to be the same by a first reviser, then the name to be outlined by Hoser (2019a, 2019b) and sources cited used and retained is that which first appears in this paper therein. by way of page priority and as listed in the abstract Therefore attempts by taxonomic vandals like the keywords. Wolfgang Wüster gang via Kaiser (2012a, 2012b, 2013, Some material in descriptions for taxa may be repeated 2014a, 2014b) and Kaiser et al. (2013) (as frequently for other taxa or groups in this paper and this is amended) to unlawfully suppress the recognition of these necessary to ensure each fully complies with the taxa on the basis they have a personal dislike for the provisions of the International Code of Zoological person who formally named it should be resisted (Dubois Nomenclature (Fourth edition) (Ride et al. 1999) as et al. 2019). amended online since. Claims by the Wüster gang against this paper and the In terms of the subtribe or subgenus descriptions, there is descriptions herein will no doubt be no different to those usually no corresponding description for the nominate the gang have made previously, all of which were subtribe or subgenus (the nominate one being identified discredited long ago as outlined by Dubois et al. (2019), in the relevant description/s). Hoser, (2007, 2009, 2012a, 2012b, 2013a, 2015a-f, However these must be assumed as identified and by 2019a, 2019b) and sources cited therein. way of reverse diagnosis of the opposing subtribe/s or The order of treatment of all the Pelodryadidae within this subgenus/genera (within those descriptions), thereby paper is as follows. making the nominate subtribe or subgenus name also Species are dealt with in clades corresponding to tribes available, and as of the date of publication of this paper as outlined above. (5 June 2020) for nomenclatural purposes in accordance Under each tribe or subtribe heading are the genus and with the rules of the International Code of Zoological subgenus descriptions, followed by any relevant species- Nomenclature (Ride et al. 1999). level descriptions. That is the nominate subtribes are formally named in this Each tribe is dealt with alphabetically herein as are the paper, even when not expressly done so in the relevant groups within each. part. Each is designated herein as “new”. The same applies with regards to subgenera and However there have been some family level nominate subgenera. classifications in the past and the following names have Material downloaded from the internet and cited been used and are therefore available: anywhere in this paper was downloaded and checked Pelodryadidae Günther, 1858 most recently as of 22 May 2020, unless otherwise stated Cycloraninae Parker, 1940 (some downloads are later than 22 May 2020) and were Nyctimystinae Laurent, 1975 accurate in terms of the context cited herein as of that Because tribe is taken as being family-level classification, date. these three authors may be taken as name authorities for Unless otherwise stated explicitly, colour descriptions for the three relevant tribes formally assigned for the first frogs apply to living adult specimens of generally good time as tribes within this paper. health, as seen in normal daytime conditions and not At the end of the genus and species level descriptions under any form of stress by means such as excessive the 12 tribes and 11 subtribes are formally described and cool, heat, dehydration or abnormal skin reaction to named. Zoobank numbers for tribes and subtribes are chemical or other input. placed with their full descriptions and not in the earlier A general reference to “colour” is unless otherwise text of this paper. stated, referring to the dorsal and obvious colouration of The diagnostic information for the tribes wholly reflects the frog on the usually visible surfaces. that of component genera and this means that persons Unless otherwise stated, the following applies. Size only interested in species or genus-level classification measurements and ratios quoted herein are for normal need not read the formal tribe descriptions, but can adults of normal adult size. Where one number only is ascertain tribe placement from the relevant accounts given, this is the average measurement. Where two earlier in the paper. numbers are given in the form of a range, this means After the references cited list is a full species list for the “known range” based on previously measured and Pelodryadidae. recorded specimens. Acronyms used in descriptions in this paper, unless While numerous texts and references were consulted otherwise stated, are as follows: prior to publication of this paper, the criteria used to (SV) (S-V): Snout to vent length. The distance between separate the relevant species has already been spelt out the anterior tip of the snout and the superior margin of and/or is done so within each formal description and the cloacal aperture. does not rely on material within publications not explicitly (HL): Head length. Distance between the tip of the snout cited herein. and the posterior margin of the tympanum, including the Each newly named tribe, subtribe, genus, subgenus, Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 9

tympanic annulus. continue to do so. (HW): Head width. The greatest width of the head, The conservation situation for frogs in New Guinea and usually recorded at a level with the tympana. offshore islands is even more dire and again gives (E-N): Eye to naris distance. The distance between the justification and urgency for the naming of hitherto anterior margin of the eye and the posterior margin of the unnamed species in the Australasian region. naris. According to the website https://www.worldometers.info (IN): Internarial span: The minimum distance separating Papua New Guinea claimed a population of nearly 9 the nares. million people in 2020. (ED): Eye diameter: The distance between the anterior This is up from just over 2 million in 1955, more than a 4 and posterior corners. fold rise in 65 years. (TD): Tympanum diameter. In the vast majority of In that time Papua New Guinea has been converted from specimens of most species the tympanum is circular and largely untouched jungle to mainly heavily cleared and or the horizontal diameter, measured to the outer margins of generally vandalized habitat, with an ever decreasing the tympanic annulus, is taken as the standard amount of native wildlife. measurement. The pace of habitat destruction is getting faster, year on (TL): Tibia length. The length of the tibia measured from year. the convex surface of the knee to the tibio-tarsal joint, The destruction on the Indoensian side of New Guinea is with the knee held in the flexed position. of similar scale, but at the moment coming from a lower (RL/SVL): Rostral length/snout-vent length. population base. (HW/SVL): Head width measured as transverse distance In 1990 there were 385,509 people in the Indonesian between tympanum/snout-vent length from vent to tip of province of Irian Jaya. This has nearly tripled in 30 years snout. to be about 1 million in 2020. (EYE/SVL) (E/SV); Horizontal eye diameter/snout-vent Transnational companies clearing land for agriculture and length from vent to tip of snout. deforestation to satisfy insatiable global demand makes (TL/SVL) (TL/SV): Tibia length/snout-vent length from up for any lack of local people doing environmental vent to tip of snout. damage just in their quest to stay alive and satisfy daily needs. (TYM/SVL): Horizontal tympanum diameter/snout-vent length from vent to tip of snout. In any event, 1 million people in 2020 is likely to multiply to at least 4 million in 65 years and 16 million in 130 (4TD/SVL): Transverse diameter of toe four disc/snout- years and 72 million people in less than 200 years! vent length from vent to tip of snout. And this assumes no mass immigration from other even (3FD/SVL): Transverse diameter of finger three disc/ more overpopulated parts of the planet! snout-vent length from vent to tip. All in a land area of just 126,093 square kilometres. CONSERVATION OF RELEVANT SPECIES AND The ecological disaster evolving on the island of New GENERA Guinea over just a few human life spans is a disaster of In terms of the conservation outlook for the relevant biblical proportions. species and genera, the outlook is generally not good, as Even in 2020, in some areas near Port Moresby, Papua detailed in Hoser (1991), the comments being as relevant New Guinea, streams that 30 years ago were pristine and in 2020 as they were in 1991, if not more so. full of a diverse array of frogs are now nothing more than With a few exceptions, most species of frogs are open sewers taking run off from the homes of increasing regarded as being in serious decline and at risk of numbers of people living in third-world poverty and extinction, with primary blame being placed on the squalor. Australian government, State governments and likewise Humans are literally an ecological plague in both for Papua New Guinea and Indonesia. Australia and New Guinea and the non-stop population In particular via the actions of the State wildlife explosion must be arrested with urgency. departments and their steadfast refusal to enact proper The globalisation of trade has also globalized the spread captive breeding programs for the relevant taxa in any of pathogens that has already had devastating effects on meaningful way, this means that many species face an amphibian populations worldwide including in particular in inevitable path towards extinction, due to this direct New South Wales and Queensland, Australia (Hoser action and other human caused threats. 1991, Anstis 2013). The long term overpopulation of the Australia with feral humans (Saunders, 2019) does not auger well for the Some species with the Pelodryadidae have gone from long term survival of many of the relevant species in abundant and at “no extinction risk” to “rare” or “critically Australia! endangered” within two decades due to a deadly fungus and such calamities are more likely as the human impact In line with the Australian Federal Government’s “Big increases. Australia” policy, that being to increase the human population of 25 million (2020), from 13 million in around Put simply, all other “conservation” efforts pale into 1970, to over 100 million within 100 years “so that we can insignificance when tallied against the benefits of tell China what to do”, as stated by the former Prime stopping human population growth. Minister, Kevin Rudd in 2019 (Zaczek 2019), the human In the material that follows, there is generally no mention pressure on the relevant ecosystems has increased in of conservation aspects relevant to the given species or line with the human populations nearby and will clearly genera, but all the preceding is invariably relevant. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 10 Australasian Journal of Herpetology

ADELYNHOSERHYLEINI TRIBE NOV. yikes sp. nov. being until now treated as Australian ADEYLNHOSERHYLEA GEN. NOV. populations of A. eucnemis. LSIDurn:lsid:zoobank.org:act:67AA47F1-D600-47B8- Frogs within the subgenus Yikesanura subgen. nov. are 8096-E4D768EFBF8B readily separated from other species in the nominate Type species: Adelynhoserhylea adelynhoserae sp. nov. subgenus Adelynhoserhylea gen. nov. by having a large triangular skin flap on each heel and a call that is a series Diagnosis: Species of tree frogs within the genus of short growls, as opposed to a small skin flap in the Adelynhoserhylea gen. nov. are readily separated from form of a tubercle or short spine on each heel and a call all other Australasian Tree Frogs (Pelodryadidae) by the that ranges from a series of soft ticks to a faster-paced following unique suite of characters: Vomerine teeth series of “tocs” in the nominate subgenus. present. Fingers with conspicuous webbing, reaching at least as far as the base of the penultimate phalanx of the Species of tree frogs within the genus Adelynhoserhylea fourth finger. A conspicuous serrated ridge along the gen. nov. are readily separated from all other forearm and another along the hind edge of the foot. The Australasian Tree Frogs (Pelodryadidae) by the following morphologically similar genus Jackyhoserhylea gen. nov. unique suite of characters: Vomerine teeth present. with species confined to the New Guinea subregion is Fingers with conspicuous webbing, reaching at least as readily separated from Adelynhoserhylea gen. nov. by far as the base of the penultimate phalanx of the fourth having dermal fringes on limbs poorly defined, fingers finger. A conspicuous serrated ridge along the forearm only part-webbed and they do not not have a well- and another along the hind edge of the foot. The developed crenulated fold along the outer edge of the morphologically similar genus Jackyhoserhylea gen. nov. fore and hind-limbs, this always being either reduced or with species confined to the New Guinea subregion is absent. readily separated from Adelynhoserhylea subgen. nov. by having dermal fringes on limbs poorly defined, fingers Frogs within the subgenus Yikesanura subgen. nov. are only part-webbed and they do not not have a well- readily separated from other species in the nominate developed crenulated fold along the outer edge of the subgenus Adelynhoserhylea subgen. nov. by having a fore and hind-limbs, this always being either reduced or large triangular skin flap on each heel and a call that is a absent. series of short growls, as opposed to a small skin flap in the form of a tubercle or short spine on each heel and a Distribution: Frogs of the subgenus Yikesanura subgen. call that ranges from a series of soft ticks to a faster- nov. are confined to New Guinea and Cape York, paced series of “tocs” in the nominate subgenus. Australia. According to Duellman et al. (2016), the species within Etymology: When visiting an extended family of this genus diverged from their nearest living relatives in Muluridji, the native Aboriginal tribe from the Mount the genus Jackyhoserhylea gen. nov. 20.2 MYA. Carbine area of far north Queensland, Australia in 2017, I According to Duellman et al. (2016), the subgenus showed them a specimen of the species named herein Yikesanura subgen. nov. diverged from their nearest as Adelynhoserhylea (Yikesanura) yikes sp. nov. and all living relatives in the nominate subgenus the females in the group yelled out “Yikes” and backed Adelynhoserhylea subgen. nov. 17 MYA. away in fear. Distribution: Frogs of the genus Adelynhoserhylea gen. Hence the subgenus and species (yikes sp. nov.) names. nov. are confined to New Guinea and Cape York, Content: Adelynhoserhylea (Yikesanura) yikes sp. nov. Queensland, Australia. (type species); A. (Yikesanura) eucnemis (Lönnberg, Etymology: Adelynhoserhylea gen. nov. is named in 1900). honour of Adelyn Hoser, the eldest daughter of this ADELYNHOSERHYLEA (ADELYNHOSERHYLEA) author in recognition of over 21 years of services with ADELYNHOSERAE SP. NOV. Australia’s best reptiles shows, educating others about LSIDurn:lsid:zoobank.org:act:4954BEF5-386D-49BB- Australian wildlife and their conservation. B558-924C5113DE68 The latter part of the genus name reflects that the Holotype: A preserved male specimen in the South species are tree frogs. Australian Museum, Adelaide, South Australia, Australia, Content: Adelynhoserhylea adelynhoserae sp. nov. (type specimen number R41068 collected at Mount Lewis, in species); A. eucnemis (Lönnberg, 1900); A. exophthalmia far North Queensland, Australia, Latitude -16.58 S., (Tyler, Davies and Aplin, 1986); A. myola (Hoskin, 2007); Longitude 145.28 E. A. serrata (Andersson, 1916); A. yikes sp. nov.. This government-owned facility allows access to its YIKESANURA SUBGEN. NOV. holdings. LSIDurn:lsid:zoobank.org:act:3BD043A9-2198-403E- Paratypes: Three preserved specimens at the 8694-873EEAC963B9 Queensland Museum, Brisbane, Queensland, Australia, Type species: Adelynhoserhylea yikes sp. nov. as follows: Diagnosis: The subgenus Yikesanura subgen. nov. 1/ Specimen number J25175 collected at Mount Finnigan includes all species within the group of species National Park, Queensland, Australia, Latitude -15.8194 associated with the putative taxon originally described as S. Longitude 145.2806 E. Hyla eucnemis Lönnberg, 1900. 2/ Specimen number J27274 collected at Horans Gully, These include the Adelynhoserhylea eucnemis Mount Finnigan, Queensland, Australia, Latitude - (Lönnberg, 1900) species complex as well as the 15.8333 S., Longitude 145.2667 E. Australian species formally named within this paper as A. 3/ Specimen number J87041 collected at Little Stony

Creek, Grey Range, Queensland, Australia, Latitude - morphologically similar genus Jackyhoserhylea gen. nov. 15.854 S., Longitude 145.211 E. with species confined to the New Guinea subregion is Diagnosis: Until now the species Adelynhoserhylea readily separated from Adelynhoserhylea gen. nov. by adelynhoserae sp. nov. has been treated as a northern having dermal fringes on limbs poorly defined, fingers population of A. serrata (Andersson, 1916). Richards et only part-webbed and they do not not have a well- al. (2010) confirmed the presence of two species within developed crenulated fold along the outer edge of the “Litoria serrata” as recognized by them. The division of fore and hind-limbs, this always being either reduced or the north and south wet tropics populations is across the absent. well-known Black Mountain corridor or barrier (or Gap) as Frogs within the subgenus Yikesanura subgen. nov. are detailed in Hoser (2020c). readily separated from other species in the nominate The type locality/ies of A. serrata is Carrington (Malanda) subgenus Adelynhoserhylea subgen. nov. by having a and Atherton, both in the southern wet tropics of North large triangular skin flap on each heel and a call that is a Queensland. There are no available names for the north series of short growls, as opposed to a small skin flap in wet tropics population which is why it is formally named the form of a tubercle or short spine on each heel and a herein. call that ranges from a series of soft ticks to a faster- A. adelynhoserae sp. nov. would key out as “Litoria paced series of “tocs” in the nominate subgenus. serrata” in both Anstis (2013) and Cogger (2014). The A photos of A. serrata in life can be found in Vanderduys species A. adelynhoserae sp. nov. is readily separated (2012) on page 70 at bottom right, with a photo of A. from A. serrata by a significantly greater preponderance adelynhoserae sp. nov. on the same page on the left. of green on the body as compared to A. serrata which is Photos of both species in life can also be seen in Anstis generally a reddish-brown frog, even when green (2013) at page 306 with A. serrata on top right and A. markings are taken into consideration. adelynhoserae sp. nov. in the other two images. The two species are readily separated as follows: Dozens of images of both taxa can be found online at: Green pigment reaches the tip of the snout in A. http://www.flickr.com adelynhoserae sp. nov. but not in A. serrata. Distribution: A. adelynhoserae sp. nov. is an endemic By day, adult A. adelynhoserae sp. nov. have a well- from the northern wet tropics of far north Queensland, defined dark stripe running from the eye (very narrow at Australia, in a region of coastal ranges between Mount start, before abruptly widening), through the entire Lewis Lat. -16.583 S., Longitude 145.283 E. in the south tympanum and towards the axilla of the forelimb. By and Endeavour River, Cape York Peninsula, Latitude - contrast in A. serrata the line is usually absent, or if 15.47 S., Longitude 145.25 E. in the north. present ill defined or indistinct. A. serrata is therefore restricted to mountainous areas Adult A. serrata invariably have a distinct and noticeable between Cairns in the North and Mount Spec, Paluma blueish green upper iris area, which while present in A. Range in the south. adelynhoserae sp. nov. is somewhat indistinct in most, Etymology: The newly named species A. adelynhoserae but not all specimens. sp. nov. is named in honour of Adelyn Hoser, the eldest Scattered tubercles on the upper forearms of A. serrata daughter of this author in recognition of over 21 years of are somewhat pointed or jagged, versus slightly blunted services with Australia’s best reptiles shows, educating in A. adelynhoserae sp. nov.. The lower iris of A. serrata others about Australian wildlife and their conservation. is orange, versus brown in A. adelynhoserae sp. nov.. ADELYNHOSERHYLEA (YIKESANURA) YIKES SP. The tip of the snout of A. serrata is either dark or heavily NOV. peppered, versus light or not heavily peppered in A. LSIDurn:lsid:zoobank.org:act:DB7788A3-73F9-4A65- adelynhoserae sp. nov.. Dark markings or cross bands 9EDE-12AA33B90B36 occupy less than half the surface area of the upper Holotype: A preserved male specimen in the surface of the rear hind leg below the knee in A. Queensland Museum, Brisbane, Queensland, Australia, adelynhoserae sp. nov. versus more than half in A. specimen number J67706 collected at the Upper Peach serrata. Creek in the McIlwraith Range on Cape York, The lower flanks of A. serrata are yellowish in colour, Queensland, Australia, Latitude -13.73 S. Longitude versus whitish in A. adelynhoserae sp. nov.. 143.3361 E. Both A. serrata and A. adelynhoserae sp. nov. are readily This government-owned facility allows access to its separated from the other species in the genus holdings. Adelynhoserhylea gen. nov. by the call of the males Paratypes: Thirteen preserved specimens at the which is a slow to barely medium paced “tic”, versus a Queensland Museum, Brisbane, Queensland, Australia, very fast paced series of “tics” or “tocs” or series of short specimen numbers J60905, J67703, J67714, J70581, growls as in all the other species. J70588, J70599, J70600, J74444, J74447, J74452, Species of tree frogs within the genus Adelynhoserhylea J74454, J74457 and J86917 all from within 10 km of the gen. nov. are readily separated from all other type locality, being the Upper Peach Creek in the Australasian Tree Frogs (Pelodryadidae) by the following McIlwraith Range on Cape York, Queensland, Australia, unique suite of characters: Vomerine teeth present. Latitude -13.73 S. Longitude 143.3361 E. Fingers with conspicuous webbing, reaching at least as Diagnosis: Until now the species Adelynhoserhylea far as the base of the penultimate phalanx of the fourth (Yikesanura) yikes sp. nov. has been treated as an finger. A conspicuous serrated ridge along the forearm Australian population of the species A. eucnemis and another along the hind edge of the foot. The (Lönnberg, 1900) with a type locality in New Guinea, Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 12 Australasian Journal of Herpetology known in most texts as “Litoria eucnemis” (e.g. Cogger far as the base of the penultimate phalanx of the fourth 2014, Anstis 2013), or alternatively as “Ranoidea finger. A conspicuous serrated ridge along the forearm eucnemis “ (e.g. at https:// and another along the hind edge of the foot. The amphibiansoftheworld.amnh.org/Amphibia/Anura/ morphologically similar genus Jackyhoserhylea gen. nov. Pelodryadidae/Pelodryadinae/Ranoidea/Ranoidea- with species confined to the New Guinea subregion is eucnemis) or “Dryopsophus eucnemis” (Duellman et al. readily separated from Adelynhoserhylea subgen. nov. by 2016). having dermal fringes on limbs poorly defined, fingers With a divergence of more than 20 MYA from all three only part-webbed and they do not not have a well- above-named genera, based on the results of Duellman developed crenulated fold along the outer edge of the et al. 2016, it is clear that a separate genus was fore and hind-limbs, this always being either reduced or warranted for this putative taxon and nearest relatives absent. and hence the formal erection of Adelynhoserhylea gen. Distribution: As far is is known, this taxon is restricted to nov.. isolated pockets on Cape York Peninsula, far north A. yikes sp. nov. would key out as A. eucnemis in both Queensland, Australia. Cogger (2014) and Anstis (2013) and is morphologically Etymology: As for the subgenus Yikesanura subgen. very similar to that species. nov.. A. yikes sp. nov. is however readily separated from A. JACKYHOSERHYLEA GEN. NOV. eucnemis by having an iris that is dominantly orange in LSIDurn:lsid:zoobank.org:act:D57B03A7-9A0C-4BCD- colour, versus yellow-gold in A. eucnemis. The flared skin B47E-D31BC69567FA folds or tubercles on the lower hind feet and rear of front Type species: Hyla genimaculata Horst, 1883. limbs are extremely large and prominent in A. eucnemis Diagnosis: The genus Jackyhoserhylea gen. nov. versus only moderately so in A. yikes sp. nov.. The includes all the frogs in the Hyla genimaculata Horst, webbing between the first and second toe reaches the 1883 species complex. disc in A. yikes sp. nov. but not quite so in A. eucnemis. While colour of both species varies significantly, The putative species Hyla genimaculata Horst, 1883, has Australian A. yikes sp. nov. are invariably a mainly a been treated by most authors since as being within the yellow to yellow brown colour dorsally, whereas A. genus Litoria Tschudi, 1838 (e.g. Anstis 2013, Cogger eucnemis from New Guinea is dominantly a green 2014), although also sometimes placed within Ranoidea colouration on top with brown or beige spots, blotches, Tschudi, 1838 (e.g. https:// flecks or peppering. amphibiansoftheworld.amnh.org/Amphibia/Anura/ Pelodryadidae/Pelodryadinae/Ranoidea/Ranoidea- A. yikes sp. nov. is depicted in life in photos in genimaculata), or even Dryopsophus Fitzinger, 1843 (e.g. Vanderduys (2012) on page 39, bottom left, Anstis (2013) Duellman et al. 2016). on page 195 (all photos), Cogger (2014) on page 163 (both images) and Eipper (2018) on page 129 (top It is in fact so divergent from the types of each of those image). genera, that there is no logical alternative but to place the taxon within a separate genus. A. eucnemis is depicted in life in Menzies (1976) plate 5 at bottom right, Menzies (2016) colour photo 48 and According to Duellman et al. (2016), this putative species Richards et al. (2010), Figure 4A. has a 20.2 MYA divergence from its nearest relatives (being Adelynhoserhylea gen. nov.) and due to significant A. eucnemis (Lönnberg, 1900) is believed to be a species morphological divergence, it makes sense for it to be complex. Hyla rhacophorus Van Kampen, 1909, Type placed within a newly erected genus as done herein. locality: “Etna Bai”, Papua, New Guinea, Indonesia, and Nyctimystes loveridgei Neill, 1954 Type locality: “small That the putative species is composite has been known stream near Taburi, a native village about 2 miles for many years (e.g. Richards et al. 2010) and this paper southeast of Rouna Fall, [Southeast Peninsula,] Papua formally names two of them for the first time, meaning [New Guinea]” are available names for two populations. this newly erected genus currently has three species, although this number may well expand. However the molecular results of Richards et al. (2010) were ambiguous and so both are tentatively treated as Species of tree frogs within the genus Jackyhoserhylea synonyms herein. gen. nov. with species confined to the New Guinea subregion are readily separated from all other A. yikes sp. nov. and A. eucnemis, consisting the Australasian Tree Frogs (Pelodryadidae) by the following subgenus Yikesanura subgen. nov. are readily separated unique suite of characters: Vomerine teeth present. from other species in the nominate subgenus Fingers only partly webbed and not with conspicuous Adelynhoserhylea gen. nov. by having a large triangular webbing that would reach at least as far as the base of skin flap on each heel and a call that is a series of short the penultimate phalanx of the fourth finger. The serrated growls, as opposed to a small skin flap in the form of a ridge along the forearm and another along the hind edge tubercle or short spine on each heel and a call that of the foot is either reduced or absent, body length (head ranges from a series of soft ticks to a faster-paced series to end of back) is less than 52 mm. of “tocs” in the nominate subgenus. The genus Jackyhoserhylea gen. nov. is readily Species of tree frogs within the genus Adelynhoserhylea separated from the morphologically similar genus gen. nov. are readily separated from all other Adelynhoserhylea gen. nov. by having dermal fringes on Australasian Tree Frogs (Pelodryadidae) by the following limbs poorly defined, fingers only part-webbed and they unique suite of characters: Vomerine teeth present. do not not have a well-developed crenulated fold along Fingers with conspicuous webbing, reaching at least as

the outer edge of the fore and hind-limbs, this always from the eye to a point above the insertion of the being either reduced or absent. forelimbs. The upper margin of the tympanic annulus is Species of tree frogs within the genus Adelynhoserhylea occasionally hidden beneath this fold. On the posterior gen. nov. are readily separated from all other surfaces of the Australasian Tree Frogs (Pelodryadidae) by the following forearm and tarsus are single rows of triangular dermal unique suite of characters: Vomerine teeth present. appendages. These are acutely pointed on the forearm Fingers with conspicuous webbing, reaching at least as and obtusely pointed on the tarsus. There is a very small far as the base of the penultimate phalanx of the fourth triangular dermal appendage on the heel, and a few finger. A conspicuous serrated ridge along the forearm prominent tubercles around the anus. The throat is and another along the hind edge of the foot body length tubercular, and the chest, abdomen and the lower less than 72 mm. surface of the thighs granular. Frogs within the subgenus Yikesanura subgen. nov. are Males possess a sub-gular vocal sac and there is a small readily separated from other species in the nominate nuptial pad on the inner surface of the first finger. subgenus Adelynhoserhylea subgen. nov. by having a According to Duellman et al. (2016), the species within large triangular skin flap on each heel and a call that is a this genus (Jackyhoserhylea gen. nov.) diverged from series of short growls, as opposed to a small skin flap in their nearest living relatives in the genus the form of a tubercle or short spine on each heel and a Adelynhoserhylea gen. nov. 20.2 MYA. According to call that ranges from a series of soft ticks to a faster- Duellman et al. (2016), the subgenus Yikesanura paced series of “tocs” in the nominate subgenus. subgen. nov. diverged from their nearest living relatives In further detail, all three species, J. ernieswilei sp. nov., in the nominate subgenus Adelynhoserhylea subgen. J. jackyhoserae sp. nov. and J. genimaculata being the nov. 17 MYA. entirety of the genus Jackyhoserhylea gen. nov. are Distribution: New Guinea, including north and south of further separated from all other Australasian Tree Frogs the central cordillera as well as on nearby offshore (Pelodryadidae) by the following unique suite of islands, at least as far west as Pulau Gebe, the type characters: locality for J. genimaculata (Horst, 1883). A very high E-N/IN ratio (1.250-1.585), crenulated dermal Etymology: Jackyhoserhylea gen. nov. is named in folds on the posterior surfaces of the forearms and tarsus honour of Jacky Hoser, the youngest daughter of this and a very small, triangular dermal appendage on the author in recognition of over 18 years of services with heel. Adult males have a snout to vent length of 30-41 Australia’s best reptiles shows, educating others about mm and adult females 40-52 mm. Australian wildlife and their conservation. The head is flattened and longer than broad (HL/HW Content: Jackyhoserhylea genimaculata (Horst, 1883); J. 1.055-1.103), its length more than one-third of the snout ernieswilei sp. nov.; J. jackyhoserae sp. nov.. to vent length (HL/S-V 0.350-0.404). JACKYHOSERHYLEA ERNIESWILEI SP. NOV. The snout is high; when viewed from above and in profile LSIDurn:lsid:zoobank.org:act:A0559864-890B-498E- it is truncate or very slightly rounded. The nostrils are B844-3DB810C95C97 lateral, their distance from the end of the snout Holotype: A preserved specimen at the Museum of considerably less than that from the eye. Comparative Zoology, Harvard University, Cambridge, The distance between the eye and the naris is greater MA 02138, United States of America, specimen number than the internarial span (E-N/IN 1.250-1.585). The MCZ Herp A-109000 collected from Weiana, Gold canthus rostralis is prominent and slightly curved. The Province, Papua New Guinea, Latitude -6.7626811 S., eye is large, its diameter slightly greater than the Longitude 144.875 E. distance separating it from the nostril. The tympanum is This facility allows access to its holdings. visible, its diameter equivalent to from one-quarter to Paratypes: Seven preserved specimens at the Museum almost two-thirds of the eye diameter. The vomerine teeth of Comparative Zoology, Harvard University, Cambridge, are in two oval series situated between and extending MA 02138, United States of America, specimen number slightly below the level of the choanae. The tongue is MCZ Herp A-109001- A-109007 collected from Weiana, small and broadly cordiform, with a slightly indented Gulf Province, Papua New Guinea, Latitude -6.7626811 posterior border. S., Longitude 144.875 E. The fingers are long and slender with very narrow lateral Diagnosis: Jackyhoserhylea jackyhoserae sp. nov. from fringes. In decreasing order of length 3>4>2>1. The Madang and J. ernieswilei sp. nov. from the Huon webbing on the fourth finger reaches the sub-articular Peninsula south-east and including the Gulf Province of tubercle at the base of the penultimate phalanx. The Papua New Guinea, south of the central cordillera have terminal discs are moderate to conspicuous. until now been treated as populations of Jackyhoserhylea The hind limbs are long and slender with a TL/S-V ratio of genimaculata (Horst, 1883), with a type locality of Pulau 0.516-0.613. Toes in decreasing order of length Gebe and a range extending east through nearby islands 4>5=or>3>2>1. The fourth and first toes are webbed to along the northern side of New Guinea in Irian Jaya to the sub-articular tubercles at the base of the penultimate north-west Papua New Guinea. phalanx, and the remainder are webbed to the base of All three morphologically similar frogs would have been the terminal discs. identified as Hyla genimaculata as defined and The skin on the dorsal surfaces is smooth or very finely diagnosed by Tyler (1968). tubercular. There is a very narrow supra-tympanic fold extending Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 14 Australasian Journal of Herpetology

The three species are however separated from one 4>5=or>3>2>1. The fourth and first toes are webbed to another as follows: the sub-articular tubercles at the base of the penultimate J. genimaculata is a light brownish to beige coloured frog phalanx and the remainder are webbed to the base of the with minimal obvious markings on the body or head. The terminal discs. flanks and nearby underbelly are bright yellowish in The skin on the dorsal surfaces is smooth or very finely colour and this often runs onto the limbs. The serrated tubercular. There is a very narrow supra-tympanic fold edges at the back of the forearm and feet are moderate. extending from the eye to a point above the insertion of Upper iris is yellow and lower iris is orange. the forelimbs. The upper margin of the tympanic annulus J. jackyhoserae sp. nov. is a frog with a mottled dorsum is occasionally hidden beneath this fold. On the posterior being a mix of yellowish and brown, again with flanks and surfaces of the nearby underbelly being bright yellowish in colour and forearm and tarsus are single rows of triangular dermal often running onto the limbs. The serrated edges at the appendages. These are acutely pointed on the forearm back of the forearm and feet are reduced as compared to and obtusely pointed on the tarsus. There is a very small J. genimaculata. The iris is generally orange top and triangular dermal appendage on the heel and a few bottom with a slight blue tinge at the very edge of top and prominent tubercles around the anus. The throat is bottom. tubercular and the chest, abdomen and the lower surface J. ernieswilei sp. nov. has green and brown on the of the thighs granular. dorsum which becomes yellow on the flanks and whitish Males possess a sub-gular vocal sac and there is a small below. There are scattered tiny tubercles on the dorsal nuptial pad on the inner surface of the first finger. surface, including most of the time a distinct small But in summary, species of tree frogs within the genus pointed one on top of either eye. The top of the iris is Jackyhoserhylea gen. nov. with species confined to the yellow and the bottom brown. The serrated edges at the New Guinea subregion are readily separated from all back of the forearm and feet are expanded and the folds other Australasian Tree Frogs (Pelodryadidae) by the tend to merge. following unique suite of characters: Vomerine teeth J. jackyhoserae sp. nov. is separate to the other two present. Fingers only partly webbed and not with species in that the front toe pads are much wider than the conspicuous webbing that would reach at least as far as digits, versus only marginally so in each of the other the base of the penultimate phalanx of the fourth finger. species. The serrated ridge along the forearm and another along All three species, J. ernieswilei sp. nov., J. jackyhoserae the hind edge of the foot is either reduced or absent, sp. nov. and J. genimaculata are separated from all other body length (head to end of back) is less than 52 mm. Australasian Tree Frogs (Pelodryadidae) by the following The genus Jackyhoserhylea gen. nov. is readily unique suite of characters: separated from the morphologically similar genus A very high E-N/IN ratio (1.250-1.585), crenulated dermal Adelynhoserhylea gen. nov. by having dermal fringes on folds on the posterior surfaces of the forearms and tarsus limbs poorly defined, fingers only part-webbed and they and a very small, triangular dermal appendage on the do not not have a well-developed crenulated fold along heel. Adult males have a snout to vent length of 30-41 the outer edge of the fore and hind-limbs, this always mm and adult females 40-52 mm. being either reduced or absent. The head is flattened and longer than broad (HL/HW Distribution: J. ernieswilei sp. nov. is a species from the 1.055-1.103), its length more than one-third of the snout Gulf Province of Papua New Guinea and appears to be to vent length (HL/S-V 0.350-0.404). The snout is high; found elsewhere south of central cordillera of New when viewed from above and in Guinea as well as north of the cordillera in south-east profile it is truncate or very slightly rounded. The nostrils Papua New Guinea east of the Huon Peninsula. are lateral, their distance from the end of the snout Etymology: Named in honour of Ernest Swile of Cape considerably less than that from the eye. Town in South Africa in recognition for his services to The distance between the eye and the naris is greater wildlife conservation through assisting this author’s team than the internarial span (E-N/IN 1.250-1.585). The in our research activities on African wildlife, in particular canthus rostralis is prominent and slightly curved. The with the venomous snakes including Cobras and Vipers. eye is large, its diameter slightly greater than the JACKYHOSERHYLEA JACKYHOSERAE SP. NOV. distance separating it from the nostril. The tympanum is LSIDurn:lsid:zoobank.org:act:75774B36-DF46-4163- visible, its diameter equivalent to from one-quarter to A80B-3D76E6D651D7 almost two-thirds of the eye diameter. The vomerine teeth Holotype: A preserved male specimen at the Bernice P. are in two oval series situated between and extending Bishop Museum, Honolulu, Hawaii, USA, specimen slightly below the level of the choanae. The tongue is number BPBM 12422 (also in register as 12233) small and broadly cordiform, with a slightly indented collected from 1km south west of Markham Point, on the posterior border. Lae to Wau Road, Madang Province, Papua New The fingers are long and slender with very narrow lateral Guinea. fringes. In decreasing order of length 3>4>2>1. The This facility allows access to its holdings. webbing on the fourth finger reaches the sub-articular Diagnosis: Jackyhoserhylea jackyhoserae sp. nov. from tubercle at the base of the penultimate phalanx. The Madang and J. ernieswilei sp. nov. from the Huon terminal discs are moderate to conspicuous. Peninsula south-east and including the Gulf Province of The hind limbs are long and slender with a TL/S-V ratio of Papua New Guinea, south of the central cordillera have 0.516-0.613. Toes in decreasing order of length Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 15

until now been treated as populations of Jackyhoserhylea posterior border. genimaculata (Horst, 1883), with a type locality of Pulau The fingers are long and slender with very narrow lateral Gebe and a range extending east through nearby islands fringes. In decreasing order of length 3>4>2> 1. The along the northern side of New Guinea in Irian Jaya to webbing on the fourth finger reaches the sub-articular north-west Papua New Guinea. tubercle at the base of the penultimate phalanx. The All three morphologically similar frogs would have been terminal discs are moderate to conspicuous. identified as Hyla genimaculata as defined and The hind limbs are long and slender with a TL/S-V ratio of diagnosed by Tyler (1968). 0.516-0.613. Toes in decreasing order of length The three species are however separated from one 4>5=or>3>2>1. The fourth and first toes are webbed to another as follows: the sub-articular tubercles at the base of the penultimate J. genimaculata is a light brownish to beige coloured frog phalanx, and the remainder are webbed to the base of with minimal obvious markings on the body or head. The the terminal discs. flanks and nearby underbelly are bright yellowish in The skin on the dorsal surfaces is smooth or very finely colour and this often runs onto the limbs. The serrated tubercular. There is a very narrow supra-tympanic fold edges at the back of the forearm and feet are moderate. extending from the eye to a point above the insertion of Upper iris is yellow and lower iris is orange. the forelimbs. The upper margin of the tympanic annulus J. jackyhoserae sp. nov. is a frog with a mottled dorsum is occasionally hidden beneath this fold. On the posterior being a mix of yellowish and brown, again with flanks and surfaces of the nearby underbelly being bright yellowish in colour and forearm and tarsus are single rows of triangular dermal often running onto the limbs. The serrated edges at the appendages. These are acutely pointed on the forearm back of the forearm and feet are reduced as compared to and obtusely pointed on the tarsus. There is a very small J. genimaculata. The iris is generally orange top and triangular dermal appendage on the heel, and a few bottom with a slight blue tonge at the very edge of top prominent tubercles around the anus. The throat is and bottom. tubercular, and the chest, abdomen and the lower J. ernieswilei sp. nov. has green and brown on the surface of the thighs granular. dorsum which becomes yellow on the flanks and whitish Males possess a sub-gular vocal sac and there is a small below. There are scattered tiny tubercles on the dorsal nuptial pad on the inner surface of the first finger. surface, including most of the time a distinct small But in summary, species of tree frogs within the genus pointed one on top of either eye. The top of the iris is Jackyhoserhylea gen. nov. with species confined to the yellow and the bottom brown. The serrated edges at the New Guinea subregion are readily separated from all back of the forearm and feet are expanded and the folds other Australasian Tree Frogs (Pelodryadidae) by the tend to merge. following unique suite of characters: J. jackyhoserae sp. nov. is separate to the other two Vomerine teeth present. Fingers only partly webbed and species in that the front toe pads are much wider than the not with conspicuous webbing that would reach at least digits, versus only marginally so in each of the other as far as the base of the penultimate phalanx of the species. fourth finger. The serrated ridge along the forearm and All three species, J. ernieswilei sp. nov., J. jackyhoserae another along the hind edge of the foot is either reduced sp. nov. and J. genimaculata are separated from all other or absent, body length (head to end of back) is less than Australasian Tree Frogs (Pelodryadidae) by the following 52 mm. unique suite of characters: The genus Jackyhoserhylea gen. nov. is readily A very high E-N/IN ratio (1.250-1.585), crenulated dermal separated from the morphologically similar genus folds on the posterior surfaces of the forearms and tarsus Adelynhoserhylea gen. nov. by having dermal fringes on and a very small, triangular dermal appendage on the limbs poorly defined, fingers only part-webbed and they heel. Adult males have a snout to vent length of 30-41 do not not have a well-developed crenulated fold along mm and adult females 40-52 mm. the outer edge of the fore and hind-limbs, this always The head is flattened and longer than broad (HL/HW being either reduced or absent. 1.055-1.103), its length more than one-third of the snout Distribution: Jackyhoserhylea jackyhoserae sp. nov. is to vent length (HL/S-V 0.350-0.404). The snout is high; currently known only from Northern Papua New Guinea when viewed from above and in in the general region of Madang, generally west of the profile it is truncate or very slightly rounded. The nostrils Huon Peninsula. are lateral, their distance from the end of the snout Etymology: The species Jackyhoserhylea jackyhoserae considerably less than that from the eye. sp. nov. is named in honour of Jacky Hoser, the youngest The distance between the eye and the naris is greater daughter of this author in recognition of over 19 years of than the internarial span (E-N/IN 1.250-1.585). The services with Australia’s best reptiles shows, educating canthus rostralis is prominent and slightly curved. The others about Australian wildlife and their conservation. eye is large, its diameter slightly greater than the LEUCODIGIRANINA SUBTRIBE NOV. distance separating it from the nostril. The tympanum is LEUCODIGIRANA GEN. NOV. visible, its diameter equivalent to from one-quarter to LSIDurn:lsid:zoobank.org:act:919C4E4C-C489-44FE- almost two-thirds of the eye diameter. The vomerine teeth 9D13-E880D94935C1 are in two oval series situated between and extending Type species: Litoria andiirrmalin McDonald, 1997. slightly below the level of the choanae. The tongue is small and broadly cordiform, with a slightly indented Diagnosis: The monotypic species compring the entirety Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 16 Australasian Journal of Herpetology of the genus Leucodigirana gen. nov. is readily separated and barely wider than digits; fingers unwebbed; second from all other Australasian Tree Frogs (Pelodryadidae) by finger slightly longer than first; anterior head stripe is the following unique suite of characters: Brown to golden present, usually narrow but always continuous, but brown across most of the dorsum, the body and limbs sometimes ill-defined, not interrupted by a vertical bar in with numerous scattered dark or light brown spots and front of the eye; posterior head stripe is narrow, no more blotches, among which are usually smaller pale brown or than half as wide as and not enclosing the tympanum; cream spots and blotches. Venter is white and lower there is a moderate inner metatarsal tubercle and a small flanks greyish-white, often with brown under the throat. outer metatarsal tubercle is present; vomerine teeth Axilla and groin are flesh coloured. Hind side of thighs is present; groin is yellow and heavily blotched with black. mottled with pale and dark brown. Skin is smooth to Whistish ventrally with granular skin. No dorsolateral skin leathery above and granular on the venter. Vomerine fold. teeth are present and between the choanae. There is no Duellman et al. (2016) found that the nearest living pectoral fold. There is no enlarged tubercle or crenulated relative of this genus diverged from these species 23.5 ridge along the hind edge of the forearm. Finger and toe MYA. discs are large. Fingers lack webbing and toes are nearly Distribution: Wetter parts of the east coast of south-east completely webbed, with webbing reaching the base of Australia, extending, from Victoria, through New South the penultimate phalanx of the fourth toe. There is a Wales and south east Queensland and then to the wet prominent inner metatarsal tubercle and no outer one. tropics of north-east Queensland. Heel of adpressed hind limb goes well beyond the eye. Content: Euscelis lesueurii (Dümeril and Bibron, 1841) Tympanum is large and obvious, with a well-developed (type species); E. booroolongensis (Moore, 1961); E. supratympanic fold. Above this, there is a semidistinct jungguy (Donnellan and Mahony, 2004); E. wilcoxi beige coloured stripe, extending to form broken blotches (Günther, 1864). along a line running along the margin of the upper flank. EUSCELIS BOOROOLONGENSIS DORSARUBER This is not however in the form of a distinct, well marked SUBSP. NOV. whitish stripe. There is also no pale line along the posterior edge of the upper jaw, although this region of LSIDurn:lsid:zoobank.org:act:5C0EDE3D-FB06-4EA1- the head is usually a light bluey, purplish grey in colour. 8E38-D696B86423DC Second finger is much longer than the first, the tip of the Holotype: A preserved specimen at the National first finger goes no further than the base of the disc of the Museum of Victoria, Melbourne, Victoria, Australia, second finger when they are pressed togeather. Males specimen number D73149, collected from Killimicat get to 80 mm in body length and females 100 mm. Creek, New South Wales, Australia, Latitude -35.2 S., Duellman et al. (2016) found that the type species for this Longitude 148.33 E. This government-owned facility monotypic genus diverged from its nearest living relative allows access to its holdings. 23.5 MYA. Being morphologicaly divergent from the Paratypes: Two preserved specimens at the National nearest living relatives in the genus Euscelis Fitzinger, Museum of Victoria, Melbourne, Victoria, Australia, 1843, the case for the erection of the new genus specimen numbers D73109 and D73123, both collected Leucodigirana gen. nov. was compelling. from Bombowlee Creek, New South Wales, Australia, Distribution: Known only from the type locality, being the Latitude -35.27 S., Longitude 148.33 E. Melville Range at Cape Melville, Cape York Peninsula, far Diagnosis: The species E. booroolongensis (Moore, north Queensland. 1961), with a type locality of Ebor, on the eastern edge of Etymology: The name leucodigirana literally means the New England Tableland, New South Wales, Australia, white toed frog, in reference to the usual character state is found from the New England region of New South of adults, in particular with reference to the discs which Wales, south along the Great Dividing Range and nearby are a distinctive white colour. slopes to north-east Victoria. Within this range, specimens are known from three general areas, each “Rana” means frog. separated by well known biogeographical barriers. The Content: Leucodigirana andiirrmalin (McDonald, 1997) populations are as follows: (monotypic). 1/ The nominate form, found in the New Engand Region EUSCELIS FITZINGER, 1843 of New South Wales, generally bounded in the south by Type species: Hyla lesueurii Dümeril and Bibron, 1841. the Hunter Valley and north to near the Queensland, New Diagnosis: The genus Euscelis Fitzinger, 1843 is South Wales border, and: separated from all other Australasian Tree Frogs 2/ A second population in the general region south of the (Pelodryadidae) by the following unique suite of Hunter Valler intrusion, including elevated areas west of characters: Sydney, New South Wales, generally west of the Great Colouration is a pale fawn to dark brown above, usually Divide in a region bounded by Kandos in the north, immaculate, or with darker markings, ranging from flecks Orange in the West, Blackheath in the east and Taralga to blotches and including one that forms a transverse bar in the south, herein identified as the subspecies E. between the eyes; the tympanum has a pale rim; dorsal booroolongensis occultatum subsp. nov. and: surface generally smooth or sometimes leathery, with one 3/ The third population is found in a region generally east species having slight warts; webbing may reach the disc and west of the Australian Capital Territory, generally of the fifth toe, but usually not, and generally extending south of Tarago, New South Wales, into Victoria on the no more than halfway along the penultimate phalanx; western side of the Great Dividing Range, herein discs on fingers and toes are small and inconspicuous identified as E. booroolongensis dorsaruber subsp. nov..

All three populations would be keyed as E. https://www.flickr.com/photos/88708273@N03/ booroolongensis using the key in Cogger (2014), as 49657691666/ “Litoria booroolongensis” but are sufficiently and morphologically divergent to be formally identified as https://www.flickr.com/photos/137559394@N07/ subspecies as done herein. 45899029804/ E. booroolongensis dorsaruber subsp. nov. is the most and distinctive of the three subspecies and is readily https://www.flickr.com/photos/euprepiosaur/ separated from the other two by the following suite of 16184175046/ characters: A strongly dark reddish-brown dorsal and colouration, which is in stark contrast to the other two subspecies. E. booroolongensis occultatum subsp. nov. https://www.flickr.com/photos/137559394@N07/ is only slightly reddish in colour, being generally greyish 46570626562/ with an orange tinge, while E. booroolongensis Distribution: E. booroolongensis dorsaruber subsp. nov. booroolongensis is strongly yellowish. E. is found in a region generally east and west of the booroolongensis dorsaruber subsp. nov. is also Australian Capital Territory, generally south of Tarago, separated from the other two subspecies by having a New South Wales, into Victoria on the western side of the significant amount of dark pigment in the form of Great Dividing Range. blotches, spots or thick peppering on the upper surface of Etymology: The name “dorsaruber” is derived from Latin the lower forearms, versus minimal in the other two and effectively means red dorsum, in reflection of the subspecies. strong dark reddish-brown colouration of the relevant Both E. booroolongensis booroolongensis and E. frogs. booroolongensis occultatum subsp. nov. have a strong EUSCELIS BOOROOLONGENSIS OCCULTATUM yellow colour or tinge in the armpits and groin, versus SUBSP. NOV. white or only slightly yellow in E. booroolongensis LSIDurn:lsid:zoobank.org:act:A4BA98D3-7CBC-4499- dorsaruber subsp. nov.. A19A-41CCD36B7C27 The black peppering or other markings on the back of E. Holotype: A preserved specimen at the Australian booroolongensis dorsaruber subsp. nov. is distinct versus Museum, Sydney, New South Wales, Australia slight or semi-distinct in both E. booroolongensis specimen number R.85580, collected 2 miles from the booroolongensis and E. booroolongensis occultatum Hampton-Jenolan Caves Road on the Oberon Road, subsp. nov. about 8 km (5 miles) east of Oberon, New South Wales, E. booroolongensis occultatum subsp. nov. is separated Australia, Latitude -33.700 S., Longitude 150.016 E. from the other two subspecies by lacking the strong This government-owned facility allows access to its reddish-brown hue of E. booroolongensis dorsaruber holdings. subsp. nov. or the strong yellow hue of E. Paratype: A preserved specimen at the Australian booroolongensis booroolongensis. Museum, Sydney, New South Wales, Australia The orange dorsal warts on both E. booroolongensis specimen number R.85579, collected 2 miles from the occultatum subsp. nov. and E. booroolongensis Hampton-Jenolan Caves Road on the Oberon Road, booroolongensis strongly contrast with the lighter colour about 8 km (5 miles) east of Oberon, New South Wales, of the back, which is not the case in E. booroolongensis Australia, Latitude -33.700 S., Longitude 150.016 E. dorsaruber subsp. nov.. Diagnosis: The species E. booroolongensis (Moore, E. booroolongensis booroolongensis is readily separated 1961), with a type locality of Ebor, on the eastern edge of from both other subspecies by having a lower iris that is the New England Tableland, New South Wales, Australia, almost black, versus not so in the other two subspecies, is found from the New England region of New South as well as a having a strongly yellowish hue through the Wales, south along the Great Dividing Range and nearby dorsal colouration, dorsal warts of contrasting colour to slopes to north-east Victoria. Within this range, the skin on the back; a prominent orange bar between specimens are known from three general areas, each the front part of the eyes, and minimal dark pigment on separated by well known biogeographical barriers. The the upper surfaces of the lower forearms, or if present, it populations are as follows: is invariably very faded and indistinct. 1/ The nominate form, found in the New Engand Region E. booroolongensis booroolongensis in life is depicted in of New South Wales, generally bounded in the south by Anstis (2013) on page 154 left, Cogger (2014) at page the Hunter Valley and north to near the Queensland, New 152 and online at: South Wales border, and: https://www.flickr.com/photos/23031163@N03/ 2/ A second population in the general region south of the 24027547442/ Hunter Valler intrusion, including elevated areas west of E. booroolongensis occultatum subsp. nov. in life is Sydney, New South Wales, generally west of the Great depicted in Anstis (2013) on page 154 right and in Divide in a region bounded by Kandos in the north, Hansen and Crosby (2016) on pages 166, 167 and 168, Orange in the West, Blackheath in the east and Taralga and online at: in the south, herein identified as the subspecies E. https://www.flickr.com/photos/mattsummerville/ booroolongensis occultatum subsp. nov. and: 16396285454/ 3/ The third population is found in a region generally east Images of E. booroolongensis dorsaruber subsp. nov. and west of the Australian Capital Territory, generally can be found online at: south of Tarago, New South Wales, into Victoria on the Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 18 Australasian Journal of Herpetology western side of the Great Dividing Range, herein Images of E. booroolongensis dorsaruber subsp. nov. identified as E. booroolongensis dorsaruber subsp. nov.. can be found online at: All three populations would be keyed as E. https://www.flickr.com/photos/88708273@N03/ booroolongensis using the key in Cogger (2014), as 49657691666/ “Litoria booroolongensis” but are sufficiently and morphologically divergent to be formally identified as https://www.flickr.com/photos/137559394@N07/ subspecies as done herein. 45899029804/ E. booroolongensis dorsaruber subsp. nov. is the most and distinctive of the three subspecies and is readily https://www.flickr.com/photos/euprepiosaur/ separated from the other two by the following suite of 16184175046/ characters: A strongly dark reddish-brown dorsal and colouration, which is in stark contrast to the other two https://www.flickr.com/photos/137559394@N07/ subspecies. E. booroolongensis occultatum subsp. nov. 46570626562/ is only slightly reddish in colour, being generally greyish with an orange tinge, while E. booroolongensis Distribution: E. booroolongensis occultatum subsp. nov. booroolongensis is strongly yellowish. E. occurs in the general region south of the Hunter Valler booroolongensis dorsaruber subsp. nov. is also intrusion, including elevated areas west of Sydney, New separated from the other two subspecies by having a South Wales, generally west of the Great Divide in a significant amount of dark pigment in the form of region bounded by Kandos in the north, Orange in the blotches, spots or thick peppering on the upper surface of West, Blackheath in the east and Taralga in the south. the lower forearms, versus minimal in the other two Etymology: The name “occultatum” is derived from Latin subspecies. and effectively means hidden and in as much as this form Both E. booroolongensis booroolongensis and E. has been hidden in terms of taxonomy until now, the booroolongensis occultatum subsp. nov. have a strong name is appropriate. yellow colour or tinge in the armpits and groin, versus COGGERDONIANI TRIBE NOV. white or only slightly yellow in E. booroolongensis GENUS COGGERDONIA WELLS AND WELLINGTON, dorsaruber subsp. nov.. 1985. The black peppering or other markings on the back of E. Type species: Hyla adelaidensis Gray, 1841. booroolongensis dorsaruber subsp. nov. is distinct versus Diagnosis: Living frogs in the genus Coggerdonia Wells slight or semi-distinct in both E. booroolongensis and Wellington, 1985 are all readily separated from all booroolongensis and E. booroolongensis occultatum other Australasian Tree Frogs (Pelodryadidae) by the subsp. nov. following unique suite of characters: A slender frog being E. booroolongensis occultatum subsp. nov. is separated light brown, fawn or light green above, with dark brown from the other two subspecies by lacking the strong patches or flecks arranged in distinct longitudinal lines. reddish-brown hue of E. booroolongensis dorsaruber There is a dark brown to black stripe running from the tip subsp. nov. or the strong yellow hue of E. of the snout below the canthus to the eye, somewhat booroolongensis booroolongensis. triangular in shape as it widens towards the eye and then The orange dorsal warts on both E. booroolongensis remaining broad as it extends across the tympanum and occultatum subsp. nov. and E. booroolongensis beyond along the flank to the rear of the body. This dark booroolongensis strongly contrast with the lighter colour coloured stripe is bordered on the lower edge with a well- of the back, which is not the case in E. booroolongensis defined line of white, also running from the upper lip. dorsaruber subsp. nov.. Sometimes this stripe will break up into a series of E. booroolongensis booroolongensis is readily separated adjacent spots. The hind parts of the thighs are dark from both other subspecies by having a lower iris that is brown with orange or reddish spots. Belly is whitish to almost black, versus not so in the other two subspecies, light brown. Skin is smooth above and coarsely granular as well as a strongly yellowish hue through the dorsal below, except under the throat, which is also smooth. colouration, dorsal warts of contrasting colour to the skin Vomerine teeth are prominent between and behind the on the back; a prominent orange bar between the front choanae. There is a distinct pectoral fold. Finger and toe part of the eyes, and minimal dark pigment on the upper discs are small and not much wider than the digits. surfaces of the lower forearms, or if present, it is Fingers have basal webbing only and toes are about invariably very faded and indistinct. three quarters webbed. Inner metatarsal tubercle is large and there is no outer one. Tympanum is large and distinct E. booroolongensis booroolongensis in life is depicted in and the second finger is longer than the first. Adult size Anstis (2013) on page 154 left, Cogger (2014) at page 50 to 60 mm (derived from Cogger, 2014). 152 and online at: https://www.flickr.com/photos/23031163@N03/ According to Duellman et al. (2016), the single living 24027547442/ member of this genus diverged from its nearest living relatives 30.8 MYA. E. booroolongensis occultatum subsp. nov. in life is depicted in Anstis (2013) on page 154 right and in Distribution: Wetter parts of south-western Western Hansen and Crosby (2016) on pages 166, 167 and 168, Australia, Australia. and online at: Content: Coggerdonia adelaidensis (Gray, 1841) https://www.flickr.com/photos/mattsummerville/ (monotypic). 16396285454/

CYCLORANININI TRIBE NOV. GENUS CYCLORANA STEINDACHNER, 1867 Mitrolysis alboguttata dumptrashensis subsp. novnov.. Type species: Cyclorana novaehollandiae Steindachner, 1867. Diagnosis: The genus concept of Cyclorana Steindachner, 1867 herein is narrower than that of most extant publications (e.g. Cogger 2014), but wholly in line with the taxonomy formally proposed by Wells and Wellington (1985). While a bunch of liars, thieves and ratbags, known as the Wolfgang Wüster gang will claim my doing so is purely out of friendship to Wells and Wellington and in the absence of scientific evidence, the reverse is in fact the case. The evidence for accepting and using the Wells and Wellington taxonomy is effectively compelling. Moonie, Qld Duellman et al. (2016) found that the species within Cyclorana as defined by Wells and Wellington (1985) diverged from their nearest living relatives 13.4 MYA, these being those species placed within each of the The subgenus Paramitrolysis subgen. nov. with the type genera Neophractops Wells and Wellington (1985), with species of Cyclorana verrucosa Tyler and Martin, 1977 is type species Chiroleptes platycephalus Günther, 1873 readily separated from the nominate subgenus (of and Mitrolysis Cope, 1889 with type species Chiroleptes Mitrolysis) by having the following unique combination of alboguttatus Günther, 1867. characters: A blunt snout with nostril distinctly nearer to As this is genus-level divergence, it makes sense to the tip than to the eye; a blackish stripe on the side of the divide the relevant species accordingly and using existing head from the snout through the eye and distinct available names. tympanum to the forelimb; a dorsum with numerous skin The latter genus Mitrolysis Cope, 1889 was inadvertently folds or large tubercles, with either: 1/ Many being white- renamed by Wells and Wellington (1985) as tipped, highlighted by being surrounded by dark blackish Brendananura with the same type species and so it is an pigment as well as a dorsal colouration dominated by objective junior synonym. Hence the latter name is brilliant dark lime green or large patches of brilliant dark effectively unavailable according to the rules of the lime green, or 2/ A dorsum with numerous skin folds or International Code of Zoological Nomenclature (Ride et large tubercles not marked in any way and a dorsal al. 1999). colouration of beige, overlain with scattered and faded One divergent taxon within Mitrolysis is also herein light olive green patches or blotches in irregular fashion. placed in a new subgenus Paramitrolysis subgen. nov.. Duellman et al. (2016) found the species in the subgenus Species within the genera Cyclorana, Mitrolysis and Paramitrolysis subgen. nov. to have diverged from Neophractops are all readily separated from all other nearest congeners outside the subgenus by 12 MYA. Australasian Tree Frogs (Pelodryadidae) by the following The subgenus Invisibiliaauris subgen. nov. with the type unique suite of characters: They are heavily built, often species Cyclorana cryptotis Tyler and Martin, 1977 is rotund, frogs; Inner metatarsal tubercle is shovel-shaped; readily separated from the other two subgenera by no fronto-parietal foramen in adults; vomerine teeth are having a hidden ear, being covered by skin, in stark present and largely between the choanae; tongue large contrast to the other subgenera which have an obvious and oval-shaped; Pupil horizontal; tympanum distinct in and exposed tympanum. all but one species group (Invisibiliaauris subgen. nov.); Duellman et al. (2016) found that species within phlanges are simple and tips with tiny discs or none at Invisibiliaauris subgen. nov. diverged from their nearest all; first finger opposed to remainder and toes are living relatives (within the nominate subgenus Mitrolysis) webbed. 11.2 MYA. The genus Cyclorana is readily separated from the Distribution: Most of the top half of continental Australia. genera Mitrolysis and Neophractops by having a marked, Content: Cyclorana novaehollandiae Steindachner, 1867 straight dorsolateral skin fold and a very stout build; toes (type species); C. australis (Gray, 1842). one third webbed; hind side of thighs lacking white spots. GENUS MITROLYSIS COPE, 1889. The genus Neophractops is readily separated from the Type species: Chiroleptes alboguttatus Günther, 1867. genera Mitrolysis and Cyclorana by having no definite Diagnosis: Species within the genera Cyclorana, straight dorsolateral skin fold and toes that are three Mitrolysis and Neophractops are all readily separated quarters webbed. from all other Australasian Tree Frogs (Pelodryadidae) by The genus Mitrolysis is separated from the genera the following unique suite of characters: They are heavily Cyclorana and Neophractops by one or other of: built, often rotund, frogs; Inner metatarsal tubercle is 1/ Having no definite straight dorsolateral skin fold and shovel-shaped; no fronto-parietal foramen in adults; toes that are less than half webbed, or: vomerine teeth are present and largely between the 2/ Having a marked, straight dorsolateral skin fold and a choanae; tongue large and oval-shaped; Pupil horizontal; very slender build; toes half webbed; hind side of thighs tympanum distinct in all but one species group

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. has numerous white spots. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 20 Australasian Journal of Herpetology

(Invisibiliaauris subgen. nov.); phlanges are simple and specimen number R.54762, collected by Richard W. tips with tiny discs or none at all; first finger opposed to Wells under trash at the rubbish dump (tip) at remainder and toes are webbed. Collarenebri, New South Wales, Australia, Latitude - The genus Cyclorana is readily separated from the 29.550 S., Longitude 148.583 E. This government-owned genera Mitrolysis and Neophractops by having a marked, facility allows access to its holdings. straight dorsolateral skin fold and a very stout build; toes Paratype: A preserved specimen at the Australian one third webbed; hind side of thighs lacking white spots. Museum in Sydney, New South Wales, Australia, The genus Neophractops is readily separated from the specimen number R.33294 collected from Wee Waa, genera Mitrolysis and Cyclorana by having no definite New South Wales, Australia, Latitude -30.20 S., straight dorsolateral skin fold and toes that are three Longitude 149.333 E. quarters webbed. Diagnosis: The subspecies Mitrolysis alboguttata The genus Mitrolysis is separated from the genera dumptrashensis subsp. nov. is a morphologically distinct Cyclorana and Neophractops by one or other of: southern form of the species originally described as 1/ Having no definite straight dorsolateral skin fold and Chiroleptes alboguttatus Günther, 1867. toes that are less than half webbed, or: Chiroleptes alboguttatus Günther, 1867 was originally 2/ Having a marked, straight dorsolateral skin fold and a described with two syntypes being from Bowen and Cape very slender build; toes half webbed; hind side of thighs York in Queensland. has numerous white spots. While it is contended that neither of the syntypes The subgenus Paramitrolysis subgen. nov. with the type Günther, 1867 of are of the subspecies Mitrolysis species of Cyclorana verrucosa Tyler and Martin, 1977 is alboguttata dumptrashensis subsp. nov., they are readily separated from the nominate subgenus (of themselves both morphologically divergent and so to Mitrolysis) by having the following unique combination of remove confusion as to which taxon or specimen the characters: A blunt snout with nostril distinctly nearer to name Chiroleptes alboguttatus Günther, 1867 should be the tip than to the eye; a blackish stripe on the side of the applied to, I herein make the Cape York specimen head from the snout through the eye and distinct (BMNH 1947.2.20.6-7) as the lectotype. tympanum to the forelimb; a dorsum with numerous skin The syntype from Bowen in Queensland (BMNH folds or large tubercles, with either: 1/ Many being white- 1947.2.18.50-51) herein ceases to have nomenclatural tipped, highlighted by being surrounded by dark blackish status. pigment as well as a dorsal colouration dominated by This is done in accordance with paragraph 6, and Article brilliant dark lime green or large patches of brilliant dark 74, including all of 74.1 and 74.73, of the International lime green, or 2/ A dorsum with numerous skin folds or Code of Zoological Nomenclature (Ride et al. 1999), large tubercles not marked in any way and a dorsal including this being an “express statement of the colouration of beige, overlain with scattered and faded taxonomic purpose of the designation”, this being to light olive green patches or blotches in irregular fashion. remove potential doubt as to which subspecies or Duellman et al. (2016) found the species in the subgenus species should retain the name alboguttatus Günther, Paramitrolysis subgen. nov. to have diverged from 1867 on the basis of divergence, either morphological or nearest congeners outside the subgenus by 12 MYA. molecular between the two syntype specimens. The subgenus Invisibiliaauris subgen. nov. with the type Occurring generally in a region encompassing areas species Cyclorana cryptotis Tyler and Martin, 1977 is drained by the upper Darling River basin, Mitrolysis readily separated from the other two subgenera by alboguttata dumptrashensis subsp. nov. is readily having a hidden ear, being covered by skin, in stark separated from the nominate form of contrast to the other subgenera which have an obvious M. alboguttata on the basis of colour and morphology. and exposed tympanum. Duellman et al. (2016) found Adult M. alboguttata dumptrashensis subsp. nov. are that species within Invisibiliaauris subgen. nov. diverged generally beige in colour with a general absence of dark from their nearest living relatives (within nominate blackish markings on the body save for those on the subgenus Mitrolysis) 11.2 MYA. warts, bumps and prominent folds arranged both Distribution: Most parts of the top third of Australia. longitudinally down the dorsum and also irregularly Content: Mitrolysis alboguttata (Günther, 1867) (type scattered. By contrast M. alboguttata alboguttata has species); M. brevipes (Peters, 1871); M. cryptotis (Tyler considerable amounts of dark pigment scattered across and Martin, 1977); M. cultripes (Parker, 1940); M. the dorsum giving a colouration that is “Blackish ashy longipes (Tyler and Martin, 1977); M. maculosa (Tyler and above, indistinctly marbled with black” as stated by Martin, 1977); M. maini (Tyler and Martin, 1977); M. Günther (1867) in his original description. Furthermore manya (van Buerden and Macdonald, 1980); M. vagitus the dorsum of M. alboguttata alboguttata is fairly smooth (Tyler, Davies and Martin, 1981); M. verrucosa (Tyler and and without the massive folds and numerous warts and Martin, 1977). bumps seen in M. alboguttata dumptrashensis subsp. MITROLYSIS (MITROLYSIS) ALBOGUTTATA nov.. This black marbling or peppering on the upper body DUMPTRASHENSIS SUBSP. NOV. in the nominate subspecies is not seen in M. alboguttata dumptrashensis subsp. nov.. LSIDurn:lsid:zoobank.org:act:2A17B806-9DFA-4BC0- A second colour difference between the two froms is that B4BA-7E2231EC4756 of the vertebral stripe running from snout to rear. In M. Holotype: A preserved specimen at the Australian alboguttata alboguttata it is generally very thin and Museum in Sydney, New South Wales, Australia, whitish or occasionally light yellow in colour. By contrast,

in M. alboguttata dumptrashensis subsp. nov. it is either PARAMITROLYSIS SUBGEN. NOV. green or yellowish green and of moderate thickness as LSIDurn:lsid:zoobank.org:act:BB3D6237-CDCC-428F- opposed to being thin. BE99-78F8DA0F3981 Many adult specimens of M. alboguttata dumptrashensis Type species: Cyclorana verrucosa Tyler and Martin, subsp. nov. have large amounts of green on various parts 1977. of the body, but this is not consistent among all Diagnosis: The subgenus Paramitrolysis subgen. nov. specimens of the subspecies. with the type species of Cyclorana verrucosa Tyler and The significant difference between the two subspecies in Martin, 1977 is readily separated from the nominate the degree and intensity of the skin folds and warts on subgenus (of Mitrolysis) by having the following unique the upper body is noteworthy and implies significant combination of characters: A blunt snout with nostril divergence between the two forms. distinctly nearer to the tip than to the eye; a blackish Nominate M. alboguttata alboguttata is “smooth above; stripe on the side of the head from the snout through the hinder lower parts very finely granulated” as stated by eye and distinct tympanum to the forelimb; a dorsum with Günther (1867) in his original description. As already numerous skin folds or large tubercles, with either: noted this is not the case in M. alboguttata 1/ Many being white-tipped, highlighted by being dumptrashensis subsp. nov. and based on morphological surrounded by dark blackish pigment as well as a dorsal divergence alone, a strong case would be made for colouration dominated by brilliant dark lime green or large species-level recognition. patches of brilliant dark lime green, or: However in the absence of detail about specimens that 2/ A dorsum with numerous skin folds or large tubercles occur within the region the two subspecies may meet and not marked in any way and a dorsal colouration of beige, whether or not there is interbreeding between the two overlain with scattered and faded light olive green forms and any significant hybridisation / introgression, I patches or blotches in irregular fashion. have formally identified the divergent southern form as a Duellman et al. (2016) found the species in the subgenus subspecies instead of species. Paramitrolysis subgen. nov. to have diverged from In terms of the coastal Queensland specimens of M. nearest congeners outside the subgenus by 12 MYA. alboguttata alboguttata from the region of Mackay and The subgenus Invisibiliaauris subgen. nov. with the type north, all are of the smoother skinned form as opposed to species Cyclorana cryptotis Tyler and Martin, 1977 is the more warty specimens of M. alboguttata readily separated from the other two subgenera by dumptrashensis subsp. nov., and all have the dark ash having a hidden ear, being covered by skin, in stark coloured mottling on the dorsum as noted by Günther contrast to the other subgenera which have an obvious (1867) in his original description of two specimens from and exposed tympanum. this general region. However what becomes immediately Duellman et al. (2016) found that species within apparent when inspecting specimens is that those from Invisibiliaauris subgen. nov. diverged from their nearest the wet tropics zone commencing north of Townsville, living relatives (within the nominate subgenus Mitrolysis) have considerably greater amounts of dark pigment on 11.2 MYA. the body as compared to those from further south Species within the genera Cyclorana, Mitrolysis and (including Bowen, in Queensland), indicating divergence Neophractops are all readily separated from all other between these populations as well. Australasian Tree Frogs (Pelodryadidae) by the following It was on this basis that the lectoptype was selected from unique suite of characters: They are heavily built, often the two divergent syntypes of Günther (1867) in his rotund, frogs; Inner metatarsal tubercle is shovel-shaped; original description. no fronto-parietal foramen in adults; vomerine teeth are Colour photos of specimens of Mitrolysis alboguttata present and largely between the choanae; tongue large dumptrashensis subsp. nov. in life can be found on page and oval-shaped; Pupil horizontal; tympanum distinct in 134 of Cogger (2014) on bottom left and Anstis (2013) all but one species group (Invisibiliaauris subgen. nov.); page 87 at top left. phlanges are simple and tips with tiny discs or none at Colour photos of specimens of M. alboguttata alboguttata all; first finger opposed to remainder and toes are in life can be found in Anstis (2013) page 87 at top right. webbed. Distribution: The subspecies Mitrolysis alboguttata The genus Cyclorana is readily separated from the dumptrashensis subsp. nov. is found in north-west New genera Mitrolysis and Neophractops by having a marked, South Wales and south-west Queensland, generally in straight dorsolateral skin fold and a very stout build; toes the region encompassed by the upper Darling River one third webbed; hind side of thighs lacking white spots. system, in a region extending north to between The genus Neophractops is readily separated from the Augathella and Blackhall in central Queensland, after genera Mitrolysis and Cyclorana by having no definite which the nominate form of the species occurs in a more- straight dorsolateral skin fold and toes that are three or-less continuous distribution to include most of the rest quarters webbed. of Queensland, including most of Cape York, Queensland The genus Mitrolysis is separated from the genera and extending across the lower Gulf of Carpentaria to be Cyclorana and Neophractops by one or other of: found in the adjoining section of the Northern Territory. 1/ Having no definite straight dorsolateral skin fold and Etymology: Named after the collection site and locality toes that are less than half webbed, or: for the holotype, being under a piece of trash at the 2/ Having a marked, straight dorsolateral skin fold and a council rubbish dump at Collarenebri, New South Wales, very slender build; toes half webbed; hind side of thighs Australia.

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 22 Australasian Journal of Herpetology has numerous white spots. Mitrolysis verrucosa inornata subsp. nov. is readily Duellman et al. (2016) found that the species within separated from M. verrucosa verrucosa by having a Cyclorana as defined by Wells and Wellington (1985) dorsum with numerous skin folds or large tubercles not diverged from their nearest living relatives 13.4 MYA, marked in any way and a dorsal colouration of beige, these being those species placed within each of the overlain with scattered and faded light olive green genera Neophractops Wells and Wellington (1985), with patches or blotches in irregular fashion, as opposed to type species Chiroleptes platycephalus Günther, 1873 that described for M. verrucosa verrucosa above. and Mitrolysis Cope, 1889 with type species Chiroleptes The colouration of M. verrucosa verrucosa is brilliant, alboguttatus Günther, 1867. intense and well defined, versus dull and ill-defined in M. Distribution: Northern New South Wales and southern verrucosa inornata subsp. nov.. Queensland in inland areas usually away from the coast An image of M. verrucosa verrucosa in life can be found and ranges. in Cogger (2014) at page 142, or Anstis (2013) on page Content: Mitrolysis (Paramitrolysis) verrucosa (Tyler and 118 (two photos on right), Vandersuys (2012) on page 75, Martin, 1977). or Eipper and Rowland (2018) at page 113 bottom. Etymology: As a prefefix, “para” means beyond or Images of Mitrolysis verrucosa inornata subsp. nov. both distinct from. As the species in this subgenus are distinct adult and tadpole, can be found on page 119 of Anstis from those in the nominate subgenus, the name is (2013). appropriate and hence “Paramitrolysis”. Distribution: Mitrolysis verrucosa inornata subsp. nov. is MITROLYSIS (PARAMITROLYSIS) VERRUCOSA found in the general region of north-west New South INORNATA SUBSP. NOV. Wales, generally west of Dubbo and Moree, including LSIDurn:lsid:zoobank.org:act:9500B2FF-61B1-4A02- areas associated with the Macquarie, Bogan and Darling 9058-D628E2DE0DE7 River systems and potentially into Queensland north-west Holotype: A preserved specimen at the Australian of this zone. Museum in Sydney, New South Wales, Australia, M. verrucosa verrucosa (Tyler and Martin, 1977) is found specimen number: R.5149 collected at Wilcannia, New in the north-east of New South Wales, west of the coastal South Wales, Latitude -31.5590 S., Longitude 143.3785 ranges to about Warialda, extending north-west of here E. into southern Queensland, west at least as far as Roma This facility allows access to its holdings. and north to at least Kilcummin. Paratypes: Two preserved specimens at the Museum of Etymology: Named in reflection of the relatively inornate Comparative Zoology, Harvard University, Cambridge, colouration of this subspecies. Massachusetts, USA, specimen numbers MCZ 3585 and INVISIBILIAAURIS SUBGEN. NOV. MCZ 3586 collected at Wilcannia, New South Wales, LSIDurn:lsid:zoobank.org:act:04A5DE6A-5C22-4F0C- Latitude -31.5590 S., Longitude 143.3785 E. AC9A-DA14819EE4E8 Diagnosis: Mitrolysis verrucosa inornata subsp. nov. has Type species: Cyclorana cryptotis Tyler and Martin, until now been treated as a western population of M. 1977. verrucosa (Tyler and Martin, 1977), which it would key as Diagnosis: The subgenus Invisibiliaauris subgen. nov. in both Cogger (2014) and Anstis (2013). with the type species Cyclorana cryptotis Tyler and However Anstis (2013) flagged that this taxon was not the Martin, 1977 is readily separated from the other two same as nominate M. verrucosa verrucosa, referring to it subgenera of Mitrolysis Cope, 1889 and the genera as “Form 2”. Cyclorana Steindachner, 1867 and Neophractops Wells Anstis (2013) also hypothesised that Mitrolysis verrucosa and Wellington, 1985 by having a hidden ear, being inornata subsp. nov. may be a rough-skinned form of M. covered by skin, in stark contrast to the other subgenera cultripes (Parker, 1940) or alternatively be an and morphologically similar genera which have an intermediate between M. verrucosa and M. cultripes obvious and exposed tympanum. (Parker, 1940) and therefore an undescribed species. Duellman et al. (2016) found that species within The results of Duellman et al. (2016) effectively scuttled Invisibiliaauris subgen. nov. diverged from their nearest the first of these propositions by finding that M. cultripes living relatives (within the subgenus genus Mitrolysis) was not closely related to M. verrucosa, having diverged 11.2 MYA. some 11.2 MYA, meaning that the “Form 2” of Anstis was Species within the genera Cyclorana Steindachner, 1867, either a hitherto unnamed species or perhaps just a Mitrolysis and Neophractops Wells and Wellington, 1985 subspecies of M. verrucosa. Hence this formal are all readily separated from all other Australasian Tree description here of this subspecies, this level of Frogs (Pelodryadidae) by the following unique suite of distinction being based on the molecular results of Anstis characters: et al. (2016b) showing both forms as broadly conspecific. They are heavily built, often rotund, frogs; Inner M. verrucosa verrucosa is readily separated from M. metatarsal tubercle is shovel-shaped; no fronto-parietal verrucosa inornata subsp. nov. by having a dorsum with foramen in adults; vomerine teeth are present and largely numerous skin folds or large tubercles many being white- between the choanae; tongue large and oval-shaped; tipped, highlighted by being surrounded by dark blackish Pupil horizontal; tympanum distinct in all but one species pigment as well as a dorsal colouration dominated by group (Invisibiliaauris subgen. nov.); phlanges are simple brilliant dark lime green or large patches of brilliant dark and tips with tiny discs or none at all; first finger opposed lime green. to remainder and toes are webbed.

The genus Cyclorana is readily separated from the Australian Museum, Adelaide, South Australia, Australia, genera Mitrolysis and Neophractops by having a marked, specimen numbers R17957, R17963, R17964, R17965, straight dorsolateral skin fold and a very stout build; toes R17967, R17968, R17970, R29118 and R29121, from one third webbed; hind side of thighs lacking white spots. the same location as the holotype. The genus Neophractops is readily separated from the Diagnosis: The morphologically variable putative genera Mitrolysis and Cyclorana by having no definite species Mitrolysis cryptotis (Tyler and Martin, 1977) with straight dorsolateral skin fold and toes that are three a type locality of Daly Waters in the Northern Territory, quarters webbed. has been found across the drier parts of the tropical north The genus Mitrolysis is separated from the genera of Australia in an area stretching from near Derby in Cyclorana and Neophractops by one or other of: Western Australia in the west, across to drier parts of 1/ Having no definite straight dorsolateral skin fold and Cape York in far north Queensland. Populations in the toes that are less than half webbed, or: Kimberley division of Western Australia and those from 2/ Having a marked, straight dorsolateral skin fold and a Cape York and south-eastern parts of the Gulf of very slender build; toes half webbed; hind side of thighs Carpentaria are morphologically divergent from the has numerous white spots. nominate form and are therefore formally named as new species herein. The subgenus Paramitrolysis subgen. nov. with the type species of Cyclorana verrucosa Tyler and Martin, 1977 is All can be readily separated on the basis of colouration. readily separated from the nominate subgenus (of M. cryptotis from the Northern Territory is coloured as Mitrolysis) by having the following unique combination of follows. “The dorsal surface is pale grey suffused with characters: A blunt snout with nostril distinctly nearer to irregular darker markings” (Tyler and Martin (1977). The the tip than to the eye; a blackish stripe on the side of the markings are a rich purplish-orange-brown colour. Some head from the snout through the eye and distinct specimens of this species (including Tyler’s holotype) tympanum to the forelimb; a dorsum with numerous skin have a semi-distinct narrow white and broken mid-dorsal folds or large tubercles, with either: stripe running from snout to rear. There is a dark purplish 1/ Many being white-tipped, highlighted by being coloured stripe running from the front of the snout, surrounded by dark blackish pigment as well as a dorsal through the eye and beyond to the pit of the forearm. The colouration dominated by brilliant dark lime green or large stripe is distinct anterior to the eye and indistinct and patches of brilliant dark lime green, or: mottled after the eye. The lower flanks are whitish with 2/ A dorsum with numerous skin folds or large tubercles purple marbling. The iris is orange on top and purple not marked in any way and a dorsal colouration of beige, below. overlain with scattered and faded light olive green M. flavoranae sp. nov. from the Kimberley District of patches or blotches in irregular fashion. Western Australia is separated from M. cryptotis and M. Duellman et al. (2016) found the species in the subgenus leucodorsalinea sp. nov. by being a dominantly yellowish Paramitrolysis subgen. nov. to have diverged from coloured frog, as in the over-riding dorsal colouration, nearest congeners outside the subgenus by 12 MYA. with the irregular darker markings on the body being either brown (southern Kimberley) or orange (north-east Distribution: The drier tropics of Australia from near Kimberley) and arranged more-or-less in the linear Derby in Western Australia, across the Kimberley manner running down the body or flanks. Some Division of Western Australia, through the top end of the specimens of this species have a semi-distinct narrow Northern Territory, to the edge of the arid zone and white and broken mid-dorsal stripe running from snout to across the Gulf of Carpentaria to the east side of drier rear. parts of Cape York in north Queensland. There is a brown coloured stripe running from the front of Etymology: The subgenus name Invisibiliaauris is taken the snout, through the eye and beyond to the pit of the directly from the Latin words meaning invisible ear, in forearm. The stripe is indistinct and broken both anterior reflection of this character state in all relevant species, to the eye posterior to it. being unique to species within the entire genus Mitrolysis Cope, 1889. The iris is yellow on top and brown below. Content: Mitrolysis (Invisibiliaauris) cryptotis (Tyler and M. leucodorsalinea sp. nov. from Cape York and the Martin, 1977) (type species); M. (Invisibiliaauris) eastern Gulf of Carpentaria in Queensland is separated flavoranae sp. nov.; M. (Invisibiliaauris) leucodorsalinea from M. cryptotis and M. flavoranae sp. nov. by having a sp. nov.. prominent and well defined narrow white and broken mid- dorsal stripe running from snout to rear. While the dorsal MITROLYSIS (INVISIBILIAAURIS) FLAVORANAE SP. surface of all three species in the subgenus NOV. Invisibiliaauris subgen. nov. is covered with numerous LSIDurn:lsid:zoobank.org:act:4D2AF343-E2E0-45C6- densely aggregated and flattened tubercles, these are 8DF2-EC0B74D31B97 significantly larger and more prominent in M. Holotype: A preserved specimen at the South Australian leucodorsalinea sp. nov.. Furthermore in M. Museum, Adelaide, South Australia, Australia, specimen leucodorsalinea sp. nov., the largest tubercular number R17962, collected from near Derby (10-18 km protrusions on the anterior part of the dorsal surface south of the town), Western Australia, Australia, Latitude invariably have yellowish-white tips. In this species, these -17.38 S., Longitude 123.68 E. This government-owned tubercles tend to merge on the anterior dorsal surface to facility allows access to its holdings. form small folds running down the dorsum in a linear Paratypes: Nine preserved specimens at the South manner. The elongate (rectangular) folds seen in this

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 24 Australasian Journal of Herpetology species are not seen in the other two, which instead have markings are a rich purplish-orange-brown colour. Some more-or-less rounded tubercles only. The dorsal specimens of this species (including Tyler’s holotype) colouration is an indistinct pattern of beige, brown and have a semi-distinct narrow white and broken mid-dorsal yellow (tricolour), with a heavy overlay of orange across stripe running from snout to rear. There is a dark purplish most, but not all of the dorsum. The stripe from snout to coloured stripe running from the front of the snout, eye is brownish-grey in colour and prominent, but through the eye and beyond to the pit of the forearm. The indistinct and broken between the eye and the top of the stripe is distinct anterior to the eye and indistinct and forearm. The iris is purple-grey below and orange on top. mottled after the eye. The lower flanks are whitish with All of M. cryptotis (Tyler and Martin, 1977), M. flavoranae purple marbling. The iris is orange on top and purple sp. nov. and M. leucodorsalinea sp. nov. being the below. entirety of the subgenus Invisibiliaauris subgen. nov. can M. flavoranae sp. nov. from the Kimberley District of be readily separated from all other species within the Western Australia is separated from M. cryptotis and M. genus Mitrolysis Cope, 1889, and the morphologically leucodorsalinea sp. nov. by being a dominantly yellowish similar genera Cyclorana Steindachner, 1867 and coloured frog, as in the over-riding dorsal colouration, Neophractops Wells and Wellington, 1985 by having a with the irregular darker markings on the body being hidden ear, being covered by skin, in stark contrast to the either brown (southern Kimberley) or orange (north-east other subgenera and morphologically similar genera Kimberley) and arranged more-or-less in the linear which have an obvious and exposed tympanum. manner running down the body or flanks. Some M. cryptotis in life can be seen online at: specimens of this species have a semi-distinct narrow https://www.flickr.com/photos/88708273@N03/ white and broken mid-dorsal stripe running from snout to 49490850576/ rear. M. flavoranae sp. nov. in life can be seen in Anstis (2013) There is a brown coloured stripe running from the front of on page 94 (top right). the snout, through the eye and beyond to the pit of the M. leucodorsalinea sp. nov. in life can be seen in forearm. The stripe is indistinct and broken both anterior Vanderduys (2012) on page 33 (2 images). to the eye posterior to it. Distribution: M. flavoranae sp. nov. is believed to be The iris is yellow on top and brown below. confined to the Kimberley District of Western Australia. M. leucodorsalinea sp. nov. from Cape York and the Etymology: Taken from Latin the species name eastern Gulf of Carpentaria in Queensland is separated “flavoranae” literally means “yellow frog”. from M. cryptotis and M. flavoranae sp. nov. by having a MITROLYSIS (INVISIBILIAAURIS) prominent and well defined narrow white and broken mid- LEUCODORSALINEA SP. NOV. dorsal stripe running from snout to rear. While the dorsal LSIDurn:lsid:zoobank.org:act:386ACE8F-667B-4FD8- surface of all three species in the subgenus A1BD-B3134BD85BB6 Invisibiliaauris subgen. nov. is covered with numerous densely aggregated and flattened tubercles, these are Holotype: A preserved specimen at the Australian significantly larger and more prominent in M. Museum, Sydney, New South Wales, Australia, specimen leucodorsalinea sp. nov.. Furthermore in M. number R.149848 collected from 8.2km west of Wakooka leucodorsalinea sp. nov., the largest tubercular Outstation, Cape York, North Queensland, Australia, protrusions on the anterior part of the dorsal surface Latitude -14.583 S., Longitude 144.4978 E. invariably have yellowish-white tips. In this species, these This government-owned facility allows access to its tubercles tend to merge on the anterior dorsal surface to holdings. form small folds running down the dorsum in a linear Paratype: A preserved specimen at the Queensland manner. The elongate (rectangular) folds seen in this Museum, Brisbane, Queensland, Australia, specimen species are not seen in the other two, which instead have number J66538 collected from 8.2km west of Wakooka more-or-less rounded tubercles only. The dorsal Outstation, Cape York, North Queensland, Australia, colouration is an indistinct pattern of beige, brown and Latitude -14.583 S., Longitude 144.4978 E. yellow (tricolour), with a heavy overlay of orange across Diagnosis: The morphologically variable putative most, but not all of the dorsum. The stripe from snout to species Mitrolysis cryptotis (Tyler and Martin, 1977) with eye is brownish-grey in colour and prominent, but a type locality of Daly Waters in the Northern Territory, indistinct and broken between the eye and the top of the has been found across the drier parts of the tropical north forearm. The iris is purple-grey below and orange on top. of Australia in an area stretching from near Derby in All of M. cryptotis (Tyler and Martin, 1977), M. flavoranae Western Australia in the west, across to drier parts of sp. nov. and M. leucodorsalinea sp. nov. being the Cape York in far north Queensland. Populations in the entirety of the subgenus Invisibiliaauris subgen. nov. can Kimberley division of Western Australia and those from be readily separated from all other species within the Cape York and south-eastern parts of the Gulf of genus Mitrolysis Cope, 1889, and the morphologically Carpentaria are morphologically divergent from the similar genera Cyclorana Steindachner, 1867 and nominate form and are therefore formally named as new Neophractops Wells and Wellington, 1985 by having a species herein. hidden ear, being covered by skin, in stark contrast to the All can be readily separated on the basis of colouration. other subgenera and morphologically similar genera M. cryptotis from the Northern Territory is coloured as which have an obvious and exposed tympanum. follows. “The dorsal surface is pale grey suffused with M. cryptotis in life can be seen online at: irregular darker markings” (Tyler and Martin (1977). The https://www.flickr.com/photos/88708273@N03/ 49490850576/ Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 25

M. flavoranae sp. nov. in life can be seen in Anstis (2013) Duellman et al. (2016) found that the species in the on page 94 (top right). Neophractops to have diverged from nearest living M. leucodorsalinea sp. nov. in life can be seen in species 13.4 MYA. Vanderduys (2012) on page 33 (2 images). Until recently all populations of the type species for this Distribution: M. leucodorsalinea sp. nov. is believed to genus, namely N. platycephalus, across Australia have be confined to Cape York and the eastern Gulf of been treated as a single species. Having caught and Carpentaria in Queensland. inspected specimens of this putative species in all Etymology: Taken from Latin the species name mainland states that they occur, I learnt decades ago that “leucodorsalinea” literally means “white-mid dorsal line” in this was clearly not the case. The The Western reflection of the trait of this species. Australian form was recently described and named as GENUS NEOPHRACTOPS WELLS AND “Clyclorana occidentalis Anstis, Price, Roberts, Catalano, WELLINGTON, 1985. Hines, Doughty and Donnellan 2016”. Type species: Chiroleptes platycephalus Günther, 1873. An unnamed form from northern Australia is formally Diagnosis: The genus Neophractops Wells and named in this paper. Wellington, 1985 is readily separated from the genera The type form is from Bourke in north-west New South Mitrolysis and Cyclorana by having no definite straight Wales. Cyclorana slevini Loveridge, 1950, was dorsolateral skin fold and toes that are three quarters resurrected from synonymy by Wells and Wellington webbed. (1985) in the mistaken belief that the type for Chiroletes platycephalus was from central Australia. They stated this Species within the genera Cyclorana, Mitrolysis and belief in their paper, believing that the Central Australian Neophractops are all readily separated from all other population was specifically distinct from the western New Australasian Tree Frogs (Pelodryadidae) by the following South Wales / southern Queensland one. While their unique suite of characters: They are heavily built, often taxonomy was in fact correct, their nomenclature was not. rotund, frogs; Inner metatarsal tubercle is shovel-shaped; no fronto-parietal foramen in adults; vomerine teeth are Cyclorana slevini is therefore relegated back to the present and largely between the choanae; tongue large synonym of N. platycephalus and the name is and oval-shaped; Pupil horizontal; tympanum distinct in unavailable for any other unnamed populations of the all but one species group (Invisibiliaauris subgen. nov.); putative species. phlanges are simple and tips with tiny discs or none at Distribution: Arid parts of all mainland Australian states all; first finger opposed to remainder and toes are (including the Northern Territory), excluding Victoria. webbed. Content: Neophractops platycephala (Günther, 1873) The genus Cyclorana is readily separated from the (type species); N. occidentalis (Anstis, Price, Roberts, genera Mitrolysis and Neophractops by having a marked, Catalano, Hines, Doughty and Donnellan, 2016); N. straight dorsolateral skin fold and a very stout build; toes rosea sp. nov.. one third webbed; hind side of thighs lacking white spots. NEOPHRACTOPS ROSEA SP. NOV. The genus Mitrolysis is separated from the genera LSIDurn:lsid:zoobank.org:act:7A3777FD-71F4-4FE4- Cyclorana and Neophractops by one or other of: A7A1-BFBA40F6800E 1/ Having no definite straight dorsolateral skin fold and Holotype: A preserved specimen at the Northern toes that are less than half webbed, or: Territory Art Gallery and Museum, Darwin, Northern 2/ Having a marked, straight dorsolateral skin fold and a Territory, Australia, specimen number R09674 collected very slender build; toes half webbed; hind side of thighs from 8 km south of Dunmarra, Northern Territory, has numerous white spots. Australia, Latitude -16.75 S., Longitude 133.75 E. The subgenus Paramitrolysis subgen. nov. with the type This government-owned facility allows access to its species of Cyclorana verrucosa Tyler and Martin, 1977 is holdings. readily separated from the nominate subgenus (of Paratypes: 1/ A preserved specimen at the Northern Mitrolysis) by having the following unique combination of Territory Art Gallery and Museum, Darwin, Northern characters: A blunt snout with nostril distinctly nearer to Territory, Australia, specimen number R09715 collected the tip than to the eye; a blackish stripe on the side of the at number 26 bore at Alroy Downs Station, Northern head from the snout through the eye and distinct Territory, Australia, Latitude -19.1 S., Longitude 136.067 tympanum to the forelimb; a dorsum with numerous skin E. folds or large tubercles, with either: 1/ Many being white- 2/ A preserved specimen at the Northern Territory Art tipped, highlighted by being surrounded by dark blackish Gallery and Museum, Darwin, Northern Territory, pigment as well as a dorsal colouration dominated by Australia, specimen number R30140 collected from brilliant dark lime green or large patches of brilliant dark Morphett Creek, Northern Territory, Australia, Latitude - lime green, or 2/ A dorsum with numerous skin folds or 18.883 S., Longitude 134.083 E. large tubercles not marked in any way and a dorsal Diagnosis: Until recently, all three species Neophractops colouration of beige, overlain with scattered and faded platycephala (Günther, 1873), N. rosea sp. nov. and N. light olive green patches or blotches in irregular fashion. occidentalis (Anstis, Price, Roberts, Catalano, Hines, Duellman et al. (2016) found the species in the subgenus Doughty and Donnellan 2016) have been treated as Paramitrolysis subgen. nov. to have diverged from populations of the same putative species N. nearest living congeners outside the subgenus by 11.2 platycephala. MYA. However, it was no secret among herpetologists that

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 26 Australasian Journal of Herpetology more than one species had been lumped under the shape, of moderate to large size. single name. As far back as 1985, Wells and Wellington Adult N. occidentalis are are separated from the other (1985), who at the time had conducted considerable two species as follows: The frog is of a uniform yellowish fieldwork on frogs across all Australian states wrote “We to dark brown colour, with marbling on the mid to lower believe that this is a species complex”. flanks. Warts on the upper surface are numerous and The same view was echoed by Tyler (1992) and again by merge to form a series of elongated skin folds, arranged Anstis (2013) who on pages 112 to 115 provided detailed in an indistinct but linear manner down the dorsum. The descriptions of each of three identified forms, including iris is orangeish yellow on top and bluey-purple to aqua colour photos of adults, tadpoles and newly below. The upper surfaces of the limbs have darker metamorphosed frogs. marbling on the otherwise brown skin, versus flecks or These three obvious species were identified as follows: spots on light coloured limbs on the other two species. 1/ “Eastern form” from inland New South Wales, The skin fold above the tympanum is prominent in N. Queensland, Southern Northern Territory and north-east occidentalis but only moderately so in the other two South Australia. This is in fact the nominate form of species. Neophractops platycephala, placed by Anstis in the Adult females of N. platycephala and N. rosea sp. nov. genus Cyclorana Steindachner, 1867. grow to 72 mm total length, versus 110 mm for N. 2/ “Northern Form”, herein formally named Neophractops occidentalis. This makes adult female N. occidentalis on rosea sp. nov. from the Northern Territory in a zone average more than double the weight of the other two between the Barkly Tableland in the east and the species. Northern Territory / Western Australian border in the Photos of N. platycephala in life can be found in Eipper west, in a zone bound by the tropical savannahs to the (2012) on page 81 and Robinson (1993) on page 76. north and the arid deserts to the south. A photo of N. rosea sp. nov. in life can be found in Eipper 3/ “Central to Western Form”, formally named by Anstis, and Rowland (2018) page 112, top photo. Price, Roberts, Catalano, Hines, Doughty and Donnellan, A photo of N. occidentalis in life can be found in Tyler, 2016 as Neophractops occidentalis from most of Western Smith and Johnstone (1994) on plate 7, image 1. Australia except for the northern third and far south, with Distribution: N. rosea sp. nov. is found in the Northern a distribution extending into the far south-west of the Territory, in a zone between the Barkly Tableland in the Northern Territory. east and the Northern Territory / Western Australian Because it was untenable that the relevant forms were border in the west, in a zone bound by the tropical unnamed, they are formally diagnosed herein. savannahs to the north and the arid deserts to the south. Anstis (2013) provides excellent photos on pages 112 Etymology: The species name “rosea” comes from the and 113 of typical adults of each species (one photo of Latin word for pink, noting that specimens of this species each), which clearly show obvious differences between usually have a pinkish hue. the three species and as lead author does much the CROTTYANURA GEN. NOV. same in Anstis et al. (2016). Anstis (2013) on page 115 provides comparative photos LSIDurn:lsid:zoobank.org:act:65B73B1E-867E-46CE- of typical metamorphosed frogs of each species (one 89E7-255574B1B615 photo of each), which clearly show obvious differences Type species: Crottyanura crottyi sp. nov. (this paper). between the three species. Diagnosis: Until now, the best known species in this Anstis (2013) on page 114 provides comparative photos genus (Crottyanura gen. nov.), originally described as of typical large tadpoles of each species (one photo of Chiroleptes dahlii Boulenger, 1896, has been generally each), which clearly show obvious differences between known as Litoria dahlii. However the molecular the three species. phylogenies of Pyron and Weins (2011) and Duellman et Adult N. platycephala are are separated from the other al. (2016), have clearly shown that species not to be two species as follows: It is dorsally a greenish-grey frog closely related to any other Australasian Tree Frogs with either a general hue of that colour, or alternatively (Pelodryadidae). indistinct blotches, flecks or markings incorporating both In the absence of any available generic name for the colours to give a greenish-grey appearance. On the sides species, it was necessary to erect a new genus for the of the back are a limited number of well spaced and tiny species as is done here. raised and pointed tubercles arranged in a somewhat However it has long been known to me that putative linear manner. The limbs are greyish or greenish in Chiroleptes dahlii Boulenger, 1896 is in fact composed of colour with closely spaced distinct small purple flecks. two related but different species. The iris is orangeish yellow. These two taxa from northern Australia are Adult N. rosea sp. nov. are are separated from the other morphologically divergent, geographically allopatric, two species as follows: The frog is grey infused with pink divided by a biogeographic barrier of known antiquity and in dorsal colour, sometimes ranging to be an immaculate therefore are two separate species by any reasonable grey-pink with broad pink areas, including indistinct interpretation. stripes from nostril to eye and then through ear, past The type species for this new genus is the newly named upper armpit to the upper flank, with or without diffuse species Crottyanurua crottyi sp. nov.. slightly darker flecks or mottling on the dorsum. The iris is The genus Crottyanura gen. nov. consists of two species, a light yellow colour. On the dorsum, but mainly the sides namely C. dahlii (Boulenger, 1896), with a type locality of are a number of raised warts of irregular but rounded Daly River, Northern Territory and found in the general

region of the type locality, being the western half of the metatarsal tubercle is not shovel-shaped. top end of the Northern Territory and into immediately It should be noted that the phylogeny for the Bell Frog adjacent north-west Western Australia, as well as the frog group of species by Burns and Crayn (2006) used C. newly named species Crottyanurua crottyi sp. nov. from crotty sp. nov. as their samples for C. dahlii. the eastern side of the Gulf of Carpentaria and the Duellman et al. (2016) found their sample of putative C. western side of Cape York in Queensland. dahlii had an 18 MYA divergence from its nearest living Both species are readily separated from other relative, that being the species within the genus Australasian Tree Frogs (Pelodryadidae) by the following Cyclorana sensu lato and not the so-called Bell frogs in diagnosis. the genera Ranoidea and Chirodryas, which they found Ranoidea Tschudi, 1838 as defined herein includes only had a 22 MYA divergence. the Bell Frog group of species, excluding the species In terms of divergences of the populations of each associated with the Chirodryas raniformis Keferstein, species within this genus, this can be inferred in the 1867 species group, herein placed in the resurrected absence of molecular data. genus Chirodryas Keferstein, 1867 and the two tropical The populations are in separate drainage systems which species Chiroleptes dahlii Boulenger, 1896, and remained disconnected even in times of glacial minima. It Crottyanurua crottyi sp. nov. herein placed in the new is noted that extant distributions and museum collection genus Crottyanura gen. nov.. locations of both species are strongly correlated with river All species within Ranoidea, Chirodryas and Crottyanura drainages, this being connected with the riverine swamp gen. nov. as defined herein, while morphologically similar, and floodplain dwelling nature of the relevant species. are sufficiently divergent from one another to warrant C. dahlii with a centre of distribution around the Daly and beling treated as separate genera. Victoria River systems occupies a zone that drained west All species within the genera Ranoidea, Chirodryas and of the western Australian Papuan landbridge in the last Crottyanura gen. nov. are readily separated from all other glacial minima, thereby preventing eastern mixture of Australasian Tree Frogs (Pelodryadidae) by the following populations from the Gulf of Carpentaria that occupied suite of characters: watersheds that drained into the Arafura Sea, thereby Vomerine teeth are present; fingers are free or webbed confirming ancient divergence of each group. only at the base; first finger is longer than, equal to or Distribution: C. dahlii (Boulenger, 1896), with a type only slightly shorter than the second finger; no outer locality of Daly River, Northern Territory is found in the metatarsal tubercle. general region of the type locality, being the western half The genera Ranoidea and Chirodryas are separated from of the top end of the Northern Territory and into the genus Crottyanura gen. nov. by having flanks that are immediately adjacent north-west Western Australia. The strongly granular below a distinct, glandular dorso-lateral newly named species Crottyanurua crottyi sp. nov. occurs skin fold with densely packed, large, rounded, usually on the eastern side of the Gulf of Carpentaria within pale coloured granules, contrasting with the smooth Queensland and the western side of Cape York in far tubercular skin on the back above the skin fold. north Queensland, Australia. In contrast Crottyanura gen. nov. has flanks that are Etymology: “Crotty”, was the abbreviated name of a now smooth or with a few scattered granules, not contrasting deceased Great Dane cross Rottweiller dog that with the skin on the back. protected this author’s scientific research facility for Ranoidea species are readily separated from species nearly 13 years. It is appropriate that a genus and within the genus Chirodryas by having either a smooth species (formally described below) are formally named in dorsum with few if any rounded warts or tubercles, or if his honour. His full name was “Crotalus”, being the present in any number or size, are arranged in regular generic name for a group of large venomous North longitudinal rows near the vertebral line. American Pit Viper snakes with a rattle on the tail. By contrast Chirodryas species have large warts on the Content: Crottyanura crottyi sp. nov. (type species); C. back that are irregularly scattered and not configured in dahlii (Boulenger, 1896). regular longitudinal rows. CROTTYANURA CROTTYI SP. NOV. The two subgenera within Ranoidea are separated as LSIDurn:lsid:zoobank.org:act:12A08B9C-7BFD-44EA- follows: Subgenus Ranoidea has a generally smooth B630-2E352B503950 dorsum with at most a few scattered, low tubercles. By Holotype: A preserved specimen at the South Australian contrast subgenus Sandgroperanura subgen. nov. has Museum, Adelaide, South Australia, Australia, specimen obvious flat warts on the back that are arranged in number R67895 collected from near Normanton, regular longitudinal rows near the vertebral line. Queensland, Australia, Latitude -17.6611 S., Longitude The build of Crottyanura gen. nov. while solid, is slender 141.1039 E. when compared to both Ranoidea and Chirodryas. The This government-owned facility allows access to its species in Crottyanura gen. nov. are further diagnosed by holdings. being olive green with brownish tinge above and a Paratypes: 1/ Two preserved specimens at the South distinctive light mid-vertebral line (in C dahlii but not often Australian Museum, Adelaide, South Australia, Australia, in C. crottyi sp. nov.); having the hind side of the thighs specimen numbers R67892 and R67894 collected from mottled, marbled or spotted with white (in C dahlii but not near Normanton, Queensland, Australia, Latitude - in C. crotty sp. nov.); a finely granular dorsum; smooth 17.6611 S., Longitude 141.1039 E. white-coloured venter; no dorsolateral skin fold; fully 2/ A preserved specimen at the Queensland Museum, webbed toes; no outer metatarsal tubercle and inner

Brisbane, Queensland, Australia, specimen number 1867 species group, herein placed in the resurected J66143, collected from Rutland Plains, Frenchs Lagoon, genus Chirodryas Keferstein, 1867 and the two tropical Cape York, Queensland, Australia, Latitude -15.8 S., species Chiroleptes dahlii Boulenger, 1896, and Longitude 141.5 E. Crottyanurua crottyi sp. nov. herein placed in the new Diagnosis: It is astounding that the species Crottyanura genus Crottyanura gen. nov.. crottyi sp. nov. has been unquestionably treated by all All species within Ranoidea, Chirodryas and Crottyanura Australian herpetologists as an eastern population of C. gen. nov. as defined herein, while morphologically similar, dahlii (Boulenger, 1896). Any comparative inspection of are sufficiently divergent from one another to warrant specimens from both widely separated areas of beling treated as separate genera. distribution of the two species (north western NT for C. All species within the genera Ranoidea, Chirodryas and dahlii and the eastern Gulf of Carpentaria region of Crottyanura gen. nov. are readily separated from all other Queensland for Crottyanura crottyi sp. nov.) clearly Australasian Tree Frogs (Pelodryadidae) by the following shows two separate species are involved. Hence the suite of characters: formal naming of C. crottyi sp. nov. herein. Vomerine teeth are present; fingers are free or webbed The species C. dahlii has a dorsal pattern reminiscent of only at the base; first finger is longer than, equal to or Ranoidea aurea (Lesson 1829) from south-east Australia. only slightly shorter than the second finger; no outer In C. dahlii the dorsal colour is greenish with well-defined metatarsal tubercle. largeish brown blotches covering the back and thighs. The genera Ranoidea and Chirodryas are separated from There is invariably a reasonably thick and well defined the genus Crottyanura gen. nov. by having flanks that are mid-dorsal stripe running from snout to rear. The hind strongly granular below a distinct, glandular dorso-lateral side of the thighs is prominently spotted, mottled or skin fold with densely packed, large, rounded, usually marbled with white which invariably has a well defined pale coloured granules, contrasting with the smooth dark coloured outer edge bordering the white parts. tubercular skin on the back above the skin fold. By contrast C. crottyi sp. nov. has a dorsal colouration of In contrast Crottyanura gen. nov. has flanks that are well defined green markings over a light grey to beige smooth or with a few scattered granules, not contrasting background, the green markings being of small size, with the skin on the back. jagged and irregular edged and close spaced, giving a Ranoidea species are readily separated from species very different view to that seen in C. dahlii. Most within the genus Chirodryas by having either a smooth specimens of C. crottyi sp. nov. lack any mid-dorsal line dorsum with few if any rounded warts or tubercles, or if runing down the midline from snout to rear and those that present in any number or size, are arranged in regular have such a line, have one that is indistinct and not longitudinal rows near the vertebral line. white, but rather simply green, brown or greyish, in line By contrast Chirodryas species have large warts on the with the dominant dorsal colouration. The hind side of the back that are irregularly scattered and not configured in thighs are not prominently spotted, mottled or marbled regular longitudinal rows. with white which invariably have a well defined dark coloured outer edge bordering the white parts. The two subgenera within Ranoidea are separated as follows: Subgenus Ranoidea has a generally smooth The upper surfaces of the hind limbs in C. crottyi sp. nov. dorsum with at most a few scattered, low tubercles. By have well defined dark patches, flecks or spots that contrast subgenus Sandgroperanura subgen. nov. has contrast and are obvious on an otherwise light coloured obvious flat warts on the back that are arranged in background. There are no similar markings and regular longitudinal rows near the vertebral line. configuration on the upper surfaces of the hind limbs in C. dahlii. The build of Crottyanura gen. nov. while solid, is slender when compared to both Ranoidea and Chirodryas. The Photos of C. dahlii in life can be found in Anstis (2013) on species in Crottyanura gen. nov. are further diagnosed by pages 180-182 and Cogger (2014) on page 160 bottom being olive green with brownish tinge above and a left. distinctive light mid-vertebral line (in C dahlii but not often Photos of C. crottyi sp. nov. in life can be found in in C. crottyi sp. nov.); having the hind side of the thighs Vanderduys (2012) on page 35. mottled, marbled or spotted with white (in C dahlii but not The genus Crottyanura gen. nov. consists of two species, in C. crotty sp. nov.); a finely granular dorsum; smooth namely C. dahlii (Boulenger, 1896), with a type locality of white-coloured venter; no dorsolateral skin fold; fully Daly River, Northern Territory and found in the general webbed toes; no outer metatarsal tubercle and inner region of the type locality, being the western half of the metatarsal tubercle is not shovel-shaped. top end of the Northern Territory and into immediately Distribution: The newly named species Crottyanurua adjacent north-west Western Australia, as well as the crottyi sp. nov. occurs on the eastern side of the Gulf of newly named species Crottyanurua crottyi sp. nov. from Carpentaria within Queensland and the western side of the eastern side of the Gulf of Carpentaria and the Cape York in far north Queensland, Australia. C. dahlii western side of Cape York in Queensland. (Boulenger, 1896), with a type locality of Daly River, Both species are readily separated from other Northern Territory is found in the general region of the Australasian Tree Frogs (Pelodryadidae) by the following type locality, being the western half of the top end of the diagnosis. Northern Territory and into immediately adjacent north- Ranoidea Tschudi, 1838 as defined herein includes only west Western Australia. the Bell Frog group of species, excluding the species Etymology: “Crotty”, was the abbreviated name of a now associated with the Chirodryas raniformis Keferstein, deceased Great Dane cross Rottweiller dog that Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 29

protected this author’s scientific research facility for SANDGROPERANURA SUBGEN. NOV. nearly 13 years. It is appropriate that a genus and LSIDurn:lsid:zoobank.org:act:B5658E8E-49A5-450A- species (formally described above) are formally named in B908-EA91ED60ACF6 his honour. Type species: Hyla aurea cyclorhynchus Boulenger, His full name was “Crotalus”, being the generic name for 1882. a group of large venomous North American Pit Viper Diagnosis: The two subgenera within Ranoidea Tschudi, snakes with a rattle on the tail. 1838 are separated as follows: Nominate subgenus RANOIDEINA SUBTRIBE NOV. Ranoidea has a generally smooth dorsum with at most a RANOIDEA TSCHUDI, 1838 few scattered, low tubercles. Type species: Ranoidea jacksionensis Tschudi, 1838 = By contrast subgenus Sandgroperanura subgen. nov. Rana aurea Lesson, 1831. has obvious flat warts on the back that are arranged in Diagnosis: Ranoidea Tschudi, 1838 as defined herein regular longitudinal rows near the vertebral line. includes only the Bell Frog group of species, excluding Duellman et al. (2016) found that the two recognized the species associated with the Chirodryas raniformis species in this subgenus, from Western Australia Keferstein, 1867 species group, herein placed in the diverged from one another 1.1 MYA. genus Chirodryas Keferstein, 1867 and the two tropical They also found that these two species diverged from species Chiroleptes dahlii Boulenger, 1896, and their nearest living congener, R. aurea from eastern Crottyanurua crottyi sp. nov. herein placed in the new Australia some 12 MYA, confirming that genus level genus Crottyanura gen. nov.. recognition of the two divergent, western Australian taxa All species within Ranoidea, Chirodryas and Crottyanura is warranted. gen. nov. as defined herein, while morphologically similar, Ranoidea Tschudi, 1838 as defined herein includes only are sufficiently divergent from one another to warrant the Bell Frog group of species, excluding the species beling treated as separate genera. associated with the Chirodryas raniformis Keferstein, All species within the genera Ranoidea, Chirodryas and 1867 species group, herein placed in the resurrected Crottyanura gen. nov. are readily separated from all other genus Chirodryas Keferstein, 1867 and the two tropical Australasian Tree Frogs (Pelodryadidae) by the following species Chiroleptes dahlii Boulenger, 1896, and suite of characters: Crottyanurua crottyi sp. nov. herein placed in the new Vomerine teeth are present; fingers are free or webbed genus Crottyanura gen. nov.. only at the base; first finger is longer than, equal to or All species within Ranoidea, Chirodryas and Crottyanura only slightly shorter than the second finger; no outer gen. nov. as defined herein, while morphologically similar, metatarsal tubercle. are sufficiently divergent from one another to warrant The genera Ranoidea and Chirodryas are separated from beling treated as separate genera. the genus Crottyanura gen. nov. by having flanks that are All species within the genera Ranoidea, Chirodryas and strongly granular below a distinct, glandular dorso-lateral Crottyanura gen. nov. are readily separated from all other skin fold with densely packed, large, rounded, usually Australasian Tree Frogs (Pelodryadidae) by the following pale coloured granules, contrasting with the smooth suite of characters: tubercular skin on the back above the skin fold. In Vomerine teeth are present; fingers are free or webbed contrast Crottyanura gen. nov. has flanks that are smooth only at the base; first finger is longer than, equal to or or with a few scattered granules, not contrasting with the only slightly shorter than the second finger; no outer skin on the back. metatarsal tubercle. Ranoidea species are readily separated from species The genera Ranoidea and Chirodryas are separated from within the genus Chirodryas by having either a smooth the genus Crottyanura gen. nov. by having flanks that are dorsum with few if any rounded warts or tubercles, or if strongly granular below a distinct, glandular dorso-lateral present in any number or size, are arranged in regular skin fold with densely packed, large, rounded, usually longitudinal rows near the vertebral line. pale coloured granules, contrasting with the smooth By contrast Chirodryas species have large warts on the tubercular skin on the back above the skin fold. In back that are irregularly scattered and not configured in contrast Crottyanura gen. nov. has flanks that are smooth regular longitudinal rows. or with a few scattered granules, not contrasting with the The two subgenera within Ranoidea are separated as skin on the back. follows: Subgenus Ranoidea has a generally smooth Ranoidea species are readily separated from species dorsum with at most a few scattered, low tubercles. within the genus Chirodryas by having either a smooth By contrast subgenus Sandgroperanura subgen. nov. dorsum with few if any rounded warts or tubercles, or if has obvious flat warts on the back that are arranged in present in any number or size, are arranged in regular regular longitudinal rows near the vertebral line. longitudinal rows near the vertebral line. Distribution: Coastal New South Wales (Australia) and By contrast Chirodryas species have large warts on the immediately adjacent parts of north-east Victoria back that are irregularly scattered and not configured in (subgenus Ranoidea), or wetter parts of south-west regular longitudinal rows. Australia (subgenus Sandgroperanura subgen. nov.). Distribution: Sandgroperanura subgen. nov. occurs in Content: Ranoidea aurea (Lesson, 1831) (type species); wetter parts of south-west Australia. The nominate R. cyclorhyncha (Boulenger, 1892); R. moorei (Copland, subgenus Ranoidea is found in coastal New South Wales 1957). (Australia) and immediately adjacent parts of north-east

Etymology: Named in reflection of where the frogs live, from the coast and immediately adjacent ranges. as in south-western Australia, being a generally sandy Content: Chirodryas raniformis (Keferstein, 1867) (type region, where the local people are referred to as “sand species); C. castanea (Steindachner, 1867); C. sloppi sp. gropers” in reflection of their alleged habits of groping in nov.. the sand. The “anura” part of the name refers to the CHIRODRYAS SLOPPI SP. NOV. genus being of frogs. LSIDurn:lsid:zoobank.org:act:9D4BD424-D271-4B67- Content: Sandgroperanura cyclorhyncha (Boulenger, AA83-4E344AD05D9F 1892) (type species); S. moorei (Copland, 1957). Holotype: A preserved female specimen at the National CHIRODRYAS KEFERSTEIN, 1867 Museum of Victoria, Melbourne, Victoria, Australia, Type species: Chirodryas raniformis Keferstein, 1867. specimen number D19286, collected 16 km south of Diagnosis: Ranoidea Tschudi, 1838 as defined herein Boorowa, New South Wales, Australia, Latitude -34.6 S., includes only the Bell Frog group of species, excluding Longitude 148.73 E. the species associated with the Chirodryas raniformis This government-owned facility allows access to its Keferstein, 1867 species group, herein placed in the holdings. resurrected genus Chirodryas Keferstein, 1867 and the Paratype: A preserved specimen at the National two tropical species Chiroleptes dahlii Boulenger, 1896, Museum of Victoria, Melbourne, Victoria, Australia, and Crottyanurua crottyi sp. nov. herein placed in the new specimen number D19494, collected 11.2 km south of genus Crottyanura gen. nov.. Tumut, New South Wales, Latitude -35.4 S., Longitude All species within Ranoidea, Chirodryas and Crottyanura 148.22 E. gen. nov. as defined herein, while morphologically similar, Diagnosis: Until now, Chirodryas sloppi sp. nov. has are sufficiently divergent from one another to warrant been treated as a southern population of C. castanea beling treated as separate genera. (Steindachner, 1867), with an unknown type locality, but All species within the genera Ranoidea, Chirodryas and of the same form and colouration as “Litoria flavipunctata Crottyanura gen. nov. are readily separated from all other Courtice and Grigg, 1975”, with a type locality of Guyra in Australasian Tree Frogs (Pelodryadidae) by the following the New England region of New South Wales (NSW), suite of characters: Australia, as detailed by Thomson et al. (1996). Vomerine teeth are present; fingers are free or webbed Thomson et al. (1996) established that C. castanea only at the base; first finger is longer than, equal to or (Steindachner, 1867) and “Litoria flavipunctata Courtice only slightly shorter than the second finger; no outer and Grigg, 1975” are one and the same species. metatarsal tubercle. They also established that the southern NSW and The genera Ranoidea and Chirodryas are separated from Australian Capital Territory (ACT) population of putative the genus Crottyanura gen. nov. by having flanks that are C. castanea are significantly different morphologically strongly granular below a distinct, glandular dorso-lateral and on this basis, the species is herein formally skin fold with densely packed, large, rounded, usually described as a new species. pale coloured granules, contrasting with the smooth Herein identified as Chirodryas sloppi sp. nov. it would tubercular skin on the back above the skin fold. In until now have been identified as C. castanea using the contrast Crottyanura gen. nov. has flanks that are smooth relevant keys in Cogger (2014) or Anstis (2013). or with a few scattered granules, not contrasting with the The three species of “Bell Frog” within the genus skin on the back. Chirodryas Keferstein, 1867 are separated from Ranoidea species are readily separated from species morphologically similar species by having large warts on within the genus Chirodryas by having either a smooth the back that are irregularly scattered and not configured dorsum with few if any rounded warts or tubercles, or if in regular longitudinal rows. present in any number or size, are arranged in regular The species Chirodryas sloppi sp. nov. and C. castanea longitudinal rows near the vertebral line. are both separated from congener C. raniformis By contrast Chirodryas species have large warts on the (Keferstein, 1867) and the sometimes sympatric and back that are irregularly scattered and not configured in morphologically similar species Ranoidea aurea (Lesson, regular longitudinal rows. 1831) by having a black and yellow marbled pattern The two subgenera within Ranoidea are separated as present on the ventral surface of the legs; toes fully follows: Subgenus Ranoidea has a generally smooth webbed; warty dorsum; large black-ringed yellow spots in dorsum with at most a few scattered, low tubercles. By the inguinal region and a series of yellow spots on the contrast subgenus Sandgroperanura subgen. nov. has hind edge of the thigh. obvious flat warts on the back that are arranged in By contrast congener C. raniformis is diagnosed by regular longitudinal rows near the vertebral line. having the ventral surface of the thighs immaculate and Duellman et al. (2016) found that the species within cream coloured; toes slightly less than fully webbed; Chirodryas diverged from their nearest living relatives warty dorsum; no large yellow spots in the inguinal region (Ranoidea) some 15.3 MYA supporting the contention and if spots are present on the posterior of the thigh, they that genus-level division and recognition is warranted. tend to be randomly arranged. Distribution: The three species within Chirodryas are all The sometimes sympatric and morphologically similar confined to Victoria, Tasmania, far south-east South species Ranoidea aurea (Lesson, 1831) is in turn Australia and cooler parts of eastern New South Wales separate from all species in Chirodryas by having (NSW), in NSW generally in higher altitude areas away expanded toe discs; toes half to three-quarter webbed;

no warts on the dorsum or if so, they are small and strongly granular below a distinct, glandular dorso-lateral pimple like only and spots are absent from the inguinal skin fold with densely packed, large, rounded, usually region, vent and posterior edge of the thigh. pale coloured granules, contrasting with the smooth Chirodryas sloppi sp. nov. from the New South Wales tubercular skin on the back above the skin fold. In (NSW) southern highlands and adjacent western slopes contrast Crottyanura gen. nov. has flanks that are smooth region is separated from C. castanea from the New or with a few scattered granules, not contrasting with the England Tableland region of NSW as follows. skin on the back. Although the black and yellow marbling on the ventral Ranoidea species are readily separated from species surface of the thigh is present in both species, the extent within the genus Chirodryas by having either a smooth and definition of these markings is different. In C. dorsum with few if any rounded warts or tubercles, or if castanea the marbling extends to the present in any number or size, are arranged in regular latero-ventral surface; whilst in C. sloppi sp. nov. it stops longitudinal rows near the vertebral line. at the inguinal area. Furthermore, the intensity or By contrast Chirodryas species have large warts on the definition of this patterning is sharp and clear in C. back that are irregularly scattered and not configured in castanea and not in C. sloppi sp. nov.. regular longitudinal rows. C. sloppi sp. nov. lacks white or yellow spots on the feet The two subgenera within Ranoidea are separated as as seen in C. castanea. follows: Subgenus Ranoidea has a generally smooth C. sloppi sp. nov. is generally a deep emerald-green dorsum with at most a few scattered, low tubercles. By colour when compared to the lime green colouration of C. contrast subgenus Sandgroperanura subgen. nov. has castanea. obvious flat warts on the back that are arranged in Most adult C. sloppi sp. nov. have a distinctive vertebral regular longitudinal rows near the vertebral line. stripe. Duellman et al. (2016) found that the species within C. sloppi sp. nov. in life are depicted on page 155 of Chirodryas diverged from their nearest living relatives Cogger (2014), Anstis (2013) on page 164 (two photos) (Ranoidea) some 15.3 MYA supporting the contention and Eipper and Rowland (2018) on page 121 (top). that genus-level division and recognition is warranted. O. castanea in life are depicted in Barker and Grigg Distribution: C. sloppi sp. nov. is found in the Southern (1977) on page 83 (top) and Tyler (1992) on page 19 Tablelands region of New South Wales, Australia and (top). immediately adjacent colder parts of the western slopes, in a region generally bound by Holbrook, in southern New In habit, I have observed specimens usually alone in fast South Wales in the south-west, Mount Clear, south of flowing stream environments in grazing country. They are Canberra in the ACT in the south-east, Oberon / Jenolan usually detected by day as they make a “plop” sound as Caves, NSW in the north-east and Orange, NSW in the they jump into the water as one approaches, having been northwest. The species O. castanea (Steindachner, 1867) resting in the open on a perch of some sort. Typically the is found in the New England region of NSW, specifically frog remains underwater until the threat passes (or it is known from an area bounded by Armidale in the south, caught). Cobbadah in the west, Inverell in the north-west and At night in heavy rain in warmer months, specimens are between Guyra and Glen Innes in the north-east, with the found crossing roads. On such nights, this species is not vast majority of records being from within 30 km of found in the large numbers of other sympatric forms of Guyra. frog. Etymology: Slop (AKA Slopp) was the name of the Ranoidea Tschudi, 1838 as defined herein includes only author’s Great Dane dog (aged 8 in 2020) that protected the Bell Frog group of species, excluding the species this author’s scientific research facility from thieves for 8 associated with the Chirodryas raniformis Keferstein, years. 1867 species group, herein placed in the resurrected genus Chirodryas Keferstein, 1867 and the two tropical It is appropriate that a species (formally described above) species Chiroleptes dahlii Boulenger, 1896, and is formally named in his honour. Crottyanurua crottyi sp. nov. herein placed in the new GEDYERANINA SUBTRIBE NOV. genus Crottyanura gen. nov.. GEDYERANA GEN. NOV. All species within Ranoidea, Chirodryas and Crottyanura LSIDurn:lsid:zoobank.org:act:BF631A5E-E1A7-4E2C- gen. nov. as defined herein, while morphologically similar, BCBC-B4C35564AC57 are sufficiently divergent from one another to warrant Type species: Gedyerana gedyei sp. nov. beling treated as separate genera. Diagnosis: The genus Gedyerana gen. nov. includes the All species within the genera Ranoidea, Chirodryas and species originally described as Hyla dayi Günther, 1897 Crottyanura gen. nov. are readily separated from all other and a closely related species herein formally described Australasian Tree Frogs (Pelodryadidae) by the following as Gedyerana gedyei sp. nov.. suite of characters: Until now, both had been treated as the same putative Vomerine teeth are present; fingers are free or webbed species, known in most texts as Nyctimystes dayi only at the base; first finger is longer than, equal to or (Günther, 1897) as seen in Cogger (2014) or alternatively only slightly shorter than the second finger; no outer “Litoria dayi” in recent texts such as Anstis (2013) and metatarsal tubercle. Eipper and Rowland (2018). The genera Ranoidea and Chirodryas are separated from Numerous molecular phylogenies, including that of the genus Crottyanura gen. nov. by having flanks that are Duellman et al. (2016) found this putative species to have

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 32 Australasian Journal of Herpetology diverged from the type species of the genus Nyctimystes Queensland Museum, Brisbane, Queensland, Australia, Stejneger, 1916, type species Nyctimantis papua specimen numbers J136326-9 collected from Cape Boulenger, 1897, 41 MYA. Tribulation, North Queensland, Australia, Latitude - With a divergence of that magnitude, clearly “Hyla dayi” 16.0878 S., Longitude 145.4548 E. needs to be assigned to a separate genus. Diagnosis: Until now Gedyerana gedyei sp. nov. has Mosleyia Wells and Wellington, 1985 is the closest match been treated as a northern population of G. dayi according to Duellman et al. (2016), but having a (Günther, 1897), known in most texts under the genus divergence of 21 MYA and with species that are very names of Nyctimystes Stejneger, 1916 type species morphologically divergent. The next nearest match is Nyctimantis papua Boulenger, 1897, or Litoria Tschudi, 26.1 MYA for several other generic groupings. 1838 type species Litoria freycineti Tschudi, 1838. The With a minimum divergence of 21 MYA from nearest two species are herein treated as the entire content of available genus group, it is clear that a new genus the genus Gedyerana gen. nov.. needed to be erected for this species group and hence Both species within the genus Gedyerana gen. nov. the erection of Gedyerana gen. nov. herein. would be diagnosed as either “Litoria dayi” in Anstis Gedyerana gen. nov. is readily separated from all other (2013) or “Nyctimystes dayi” in Cogger (2014). Australasian Tree Frogs (Pelodryadidae) by being the These frogs have large protruding eyes, with a dark iris, a only Australasian Tree Frog species having a lower broad head and a slender body. Males get to about 45 eyelid with a characteristic reticulatum or palpebral mm in body length, but females are considerably larger venation of fine, pigmented lines and a horizontally and get to about 60 mm in length. elliptical pupil. All other species within this region with a Their tadpoles have large sucker mouths. Dorsal colour lower eyelid with a characteristic reticulatum or palpebral ranges from grey, brown, yellow, orange or red, with or venation of fine, pigmented lines have a vertically without mottling, spots or flecks and with or without elliptical pupil, including species within Nyctimystes discrete white or cream blotches or ocelli on the upper Stejneger, 1916 sensu lato, as defined by other authors surfaces. Snout is moderately rounded or acuminate including Cogger (2014) at page 197 under the heading (tending to be pointed) and the genus occurs in rainforest “genus Nyctimystes Stejneger, 1916”. stream habitats. Both species within the genus Gedyerana gen. nov. Both species within Gedyerana gen. nov. are readily would be diagnosed as either “Litoria dayi” in Anstis separated from all other Australasian Tree Frogs (2013) or “Nyctimystes dayi” in Cogger (2014). (Pelodryadidae) by being the only Australasian Tree Frog These frogs have large protruding eyes, with a dark iris, a species having a lower eyelid with a characteristic broad head and a slender body. Males get to about 45 reticulatum or palpebral venation of fine, pigmented lines mm in body length, but females are considerably larger and a horizontally elliptical pupil. All other species within and get to about 60 mm in length. Their tadpoles have this region with a lower eyelid with a characteristic large sucker mouths. Dorsal colour ranges from grey, reticulatum or palpebral venation of fine, pigmented lines brown, yellow, orange or red, with or without mottling, have a vertically elliptical pupil, including species within spots or flecks and with or without discrete white or Nyctimystes Stejneger, 1916 sensu lato, as defined by cream blotches or ocelli on the upper surfaces. Snout is other authors including Cogger (2014) at page 197 under moderately rounded or acuminate (tending to be pointed) the heading “genus Nyctimystes Stejneger, 1916”. and the genus occurs in rainforest stream habitats. Gedyerana gedyei sp. nov. is readily separated from G. Distribution: Gedyerana gen. nov. is endemic to the wet dayi by having a snout that is acuminate (tending to be tropics region of far North Queensland. For more detail pointed) and a dorsal colouration being generally see the description of Gedyerana gedyei sp. nov. in this creamish, light yellow or light grey, versus having a snout paper. being moderately rounded and a dorsal colouration being Etymology: The genus is named in honour of Andrew generally brown, orange or reddish in G. dayi. Gedye of Cairns, Queensland, Australia, in recognition of Gedyerana gedyei sp. nov. tend to lack the white dorsal his services to herpetology in Australia, in particular spots and ocelli that are commonly seen in specimens of through his breeding of rare and threatened species of G. dayi. Spotting or marks on G. gedyei sp. nov. if pythons and elapids (snakes). present tend to be dark or blackish in colour. Content: Gedyerana gedyei sp. nov. (type species); G. There are a number of available synonyms for putative G. dayi (Günther, 1897). dayi, being as follows: GEDYERANA GEDYEI SP. NOV. Hyla dayi Gunther 1897 with a type locality of Bartle LSIDurn:lsid:zoobank.org:act:EEBA7E33-3788-4440- Frere Mountains, North East Queensland. AAC7-49D7FBE245CA Hyla tympanocryptis Andersson (1916) from Malanda on Holotype: A preserved specimen at the Queensland the Atherton Tableland near Mount Bartle Frere, North Museum, Brisbane, Queensland, Australia, specimen East Queensland. number J136323 collected from Cape Tribulation, North Nyctimystes hosmeri Tyler, 1964 from Tully Falls on the Queensland, Australia, Latitude -16.0878 S., Longitude southern edge of the Atherton Tablelands, North East 145.4548 E. Queensland. This government-owned facility allows access to its Nyctimystes vestigea Tyler, 1964 from Mount Bartle holdings. Frere, North East Queensland. Paratypes: Four preserved specimens at the All names and specimens apply to frogs from the

southern wet tropics, in a region south of the so-called within the genus Mosleyia Wells and Wellington, 1985 as Black Mountain Corridor (or Gap) as detailed in Hoser defined by Wells and Wellington (1985), a diagnosis (2020c). adopted wholly within this paper, diverged from their As there was no available name in the literature for the nearest living relative 21 MYA, that species group being northern specimens of putative “Hyla dayi” the species the genus Gedyerana gen. nov. formally described Gedyerana gedyei sp. nov. had to be erected based on elsewhere in this paper. material from north of the Black Mountain Corridor (or The genus Mosleyia is readily separated from all other Gap) as detailed in Hoser (2020c). Australasian Tree Frogs (Pelodryadidae) by one or other Gedyerana gedyei sp. nov. in life, and showing the of the two unique suites of characters: diagnostic features outlined above, can be found online 1/ Vomerine teeth prominent and largely between the at: choanae. No pectoral fold; A series of small enlarged https://www.flickr.com/photos/outstarwild/49663732508/ tubercles along the hind edge of the forearm forming a and low but distinct crenulated ridge; fingers with strong basal https://www.flickr.com/photos/akashsherping/ webbing; toes nearly fully webbed; a prominent inner 36614786851/ metatarsal tubercle and a small outer one. Heel of and adpressed hind limb reaches to eye or beyond; first finger https://www.flickr.com/photos/126237772@N07/ much smaller and shorter than the second and when 36120089595/ pressed together the tip of the first finger reaches no further than the base of the disc of the second finger; and tympanum indistinct; no pale stripe along the https://www.flickr.com/photos/shaneblackfnq/ supratympanic ridge or along posterior edge of upper 14996849032/ jaw; adult snout-vent length 65 mm; dorsal pattern is and mostly grey or dull green, almost blackish and consists of https://www.flickr.com/photos/shaneblackfnq/ a series of relatively narrow dark lines and spots forming 15494532365/ a fine, continuous reticulum over the back, head and G. dayi in life is depicted in Barker and Grigg (1977) on limbs. The flanks have a bluish metal sheen. White page 43 (both images) and page 84 bottom, Vanderduys below, commonly with brown on the throat; axilla and (2012) on page 36 (all images), Anstis (2013) on pages groin flesh coloured; hind side of thighs are dark brown. 186-188 (all images) and Cogger (2014) on page 197 Skin leathery, finely granular, or with numerous small (both images) and online at: scattered warts. Granular below (M. nannotis (Andersson, https://www.flickr.com/photos/edwardevans/ 1916), M. cottoni sp. nov. M. lorica (Davies and 48935973742/ McDonald, 1979)) (nominate subgenus), or: and 2/ Vomerine teeth conspicuous in two rows on a line with https://www.flickr.com/photos/edwardevans/ the hind edge of the choanae; with or without pectoral 31847701407/ fold; fingers with conspicuous webbing, reaching at least Distribution: Gedyerana gedyei sp. nov. is found in the as far as the base of the penultimate phalanx of the northern wet tropics of far north Queensland. fourth finger; hind edge of forearm is smooth or with at More specifically it is known from within a region bound most a few, low, discontinuous tubercles; hind edge of by Bushy Creek, near Julatten, Latitude -16.6 S., foot is smooth; hind side of thighs more or less uniform Longitude 145.3333 E. in the south, O’Keefe Creek, on and without black and yellow markings or marbling; Big Tableland, Latitude -15.7 S., Longitude 145.25 E. in colour brown to blackish above, never green, with or the north and on the Windsor Tableland, Latitude - without a broad, darker vertebral patch and other 16.2083 S., Longitude 144.8778 E. in the west, based on contrasting markings; a dark dorsal patch, if present, is holdings within the Queensland Museum, Brisbane, inconspicuous and commencing from a line joining the Australia. centre of each eye; tympanum covered by skin (adult size is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. G. dayi is found in the southern wet tropics of far north nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola Queensland. (Liem, 1974)) (subgenus Amnisrana subgen. nov.). More specifically this is, Mount Spec, (north of According to Duellman et al. (2016) species within the Townsville), Latitude -18.95 S., Longitude 146.1833 E, in subgenus Amnisrana subgen. nov. diverged from those the south, Upper Barron River, immediately west of in the subgenus Mosleyia 12.1 MYA. Cairns, Latitude -17.00 S., Longitude 145.4333 E. in the north and Charmillan Creek, Tully Falls Road, 13 km Due to the significant morphological divergence between south of Ravenshoe, Latitude -17.7 S., Longitude the often sympatric forms, subgenus level recognition of 145.5167 E. in the west. the latter group was deemed essential. Etymology: The new species is named in honour of Distribution: Restricted to the wet tropics of North-east Andrew Gedye of Cairns, Queensland, Australia, in Queensland, Australia. recognition of his services to herpetology in Australia, in Content: Mosleyia nannotis (Andersson, 1916) (type particular through his breeding of rare and threatened species); M. cottoni sp. nov.; M. lorica (Davies and species of pythons and elapids (snakes). McDonald, 1979); M. michaelsmythi sp. nov.; M. pilloti sp. MOSLEYIA WELLS AND WELLINGTON, 1985. nov.; M. nyakalensis (Liem, 1974); M. rheocola (Liem, 1974). Type species: Hyla nannotis Andersson, 1916. Diagnosis: Duellman et al. (2016) found that the species Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 34 Australasian Journal of Herpetology

AMNISRANA SUBGEN. NOV. Etymology: “Amnis” in Latin means stream and “Rana” LSIDurn:lsid:zoobank.org:act:06568884-50B6-4A86- means frog, hence the genus name Amnisrana or 8282-DF689642F1F6 “stream frog”. Type species: Litoria rheocolus Liem, 1974. Content: M. (Amnisrana) rheocola (Liem, 1974) (type Diagnosis: Duellman et al. (2016) found that the species species); M. (Amnisrana) michaelsmythi sp. nov.; M. within the genus Mosleyia Wells and Wellington, 1985 as (Amnisrana) pilloti sp. nov.; M. (Amnisrana) nyakalensis defined by Wells and Wellington (1985), a diagnosis (Liem, 1974). adopted wholly within this formal description, diverged MOSLEYIA (MOSLEYIA) COTTONI SP. NOV. from their nearest living relative 21 MYA, that species LSIDurn:lsid:zoobank.org:act:EFC50F6C-85A5-4FB6- group being the genus Gedyerana gen. nov. formally 98BA-C1B7DC0F712F described elsewhere in this paper. Holotype: A preserved specimen at the Queensland The genus Mosleyia is readily separated from all other Museum, Brisbane, Queensland, Australia, specimen Australasian Tree Frogs (Pelodryadidae) by one or other number J5570, collected from Mount Spurgeon, North of the two unique suites of characters: Queensland, Australia, Latitude 16.4327 S., Longitude 1/ Vomerine teeth conspicuous in two rows on a line with 145.2042 E. the hind edge of the choanae; with or without pectoral This government-owned facility allows access to its fold; fingers with conspicuous webbing, reaching at least holdings. as far as the base of the penultimate phalanx of the Paratypes: Three preserved specimens at the fourth finger; hind edge of forearm is smooth or with at Queensland Museum, Brisbane, Queensland, Australia, most a few, low, discontinuous tubercles; hind edge of specimen numbers J17861-3 collected from Parrot foot is smooth; hind side of thighs more or less uniform Creek, 50 km south of Cooktown, North Queensland, and without black and yellow markings or marbling; Australia, Latitude -15.8 S., Longitude 145.25 E. colour brown to blackish above, never green, with or Diagnosis: Until now, Mosleyia cottoni sp. nov. has been without a broad, darker vertebral patch and other treated as a northern outlier population of Mosleyia contrasting markings; a dark dorsal patch, if present, is nannotis (Andersson, 1916), more commonly known as inconspicuous and commencing from a line joining the “Litoria nannotis” as defined by Liem (1974), Anstis centre of each eye; tympanum covered by skin (adult size (2013) and Cogger (2014). is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. The two species are readily separated as follows: nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola Tympanum is distinct in M. cottoni sp. nov., as opposed (Liem, 1974)) (subgenus Amnisrana subgen. nov.) (this to barely visible in M. nannotis. being a formal diagnosis for this subgenus), or: The toes of M. nannotis are light in colour, but with 2/ Vomerine teeth prominent and largely between the obvious dark centres, versus light in colour and without choanae. No pectoral fold; A series of small enlarged obvious dark centres in M. cottoni sp. nov.. tubercles along the hind edge of the forearm forming a Colouration of specimens varies considerably both low but distinct crenulated ridge; fingers with strong basal between and within localities and sub-populations and webbing; toes nearly fully webbed; a prominent inner with sex, age and time of day so it is difficult to quantify metatarsal tubercle and a small outer one. Heel of differences, save for the general observation that dorsal adpressed hind limb reaches to eye or beyond; first finger pattern in the form of indistinct mottling, specks and much smaller and shorter than the second and when flecks is more dense and intense in M. nannotis than in pressed together the tip of the first finger reaches no M. cottoni sp. nov.. further than the base of the disc of the second finger; Both these species within the nominate subgenus tympanum indistinct; no pale stripe along the Mosleyia, Wells and Wellington, 1985, are separated supratympanic ridge or along posterior edge of upper from the four species in the other subgenus (Amnisrana jaw; adult snout-vent length 65 mm; dorsal pattern is subgen. nov.) by the following suite of characters: Snout mostly grey or dull green, almost blackish and consists of is shorter than the eye diameter; snout bluntly rounded a series of relatively narrow dark lines and spots forming but the head in general is not; loreal region strongly a fine, continuous reticulum over the back, head and concave; prepollex enlarged; nuptial pad large with limbs. The flanks have a bluish metal sheen. White coarse spinules; single outer metacarpal tubercle; adults below, commonly with brown on the throat; axilla and grow to more than 40 mm in body length. IN/EN ratio less groin flesh coloured; hind side of thighs are dark brown. than 0-970; web on first toe does not extend beyond the Skin leathery, finely granular, or with numerous small proximal subarticular tubercle; ventral surfaces of scattered warts. Granular below (M. nannotis (Andersson, posterior portion of body are not cream with a reddish 1916), M. cottoni sp. nov. and M. lorica (Davies and brown or pinkish red tinge and an absence of a dense McDonald, 1979)) (nominate subgenus). large black nupital spines around the base of the thumb. According to Duellman et al. (2016) species within the The third species in the subgenus Mosleyia, M. lorica subgenus Amnisrana subgen. nov. diverged from those (Davies and McDonald, 1979) is separated from the other in the subgenus Mosleyia 12.1 MYA. two species by its far smaller adult size (37 mm in body Due to the significant morphological divergence between length or less, vs 65 mm in body length in the other two the often sympatric forms, subgenus level recognition of species) and the fact that males have dense large black the latter group was deemed eminently sensible. nupital spines around the base of the thumb. Distribution: Restricted to the wet tropics of North-east The genus Mosleyia is readily separated from all other Queensland, Australia.

Australasian Tree Frogs (Pelodryadidae) by one or other Etymology: Named in honour of Thomas Cotton, of the two unique suites of characters: formerly of Ringwood, Victoria who for many years played 1/ Vomerine teeth prominent and largely between the an important role in the wildlife conservation efforts of the choanae. No pectoral fold; A series of small enlarged team this author works with. tubercles along the hind edge of the forearm forming a This includes in fieldwork across Australia as well as a low but distinct crenulated ridge; fingers with strong basal displayer of reptiles with Snakebusters: Australia’s best webbing; toes nearly fully webbed; a prominent inner reptiles shows, being the only hands-on reptile metatarsal tubercle and a small outer one. Heel of exhibitions in Australia that let people hold the animals. adpressed hind limb reaches to eye or beyond; first finger MOSLEYIA (AMNISRANA) MICHAELSMYTHI SP. NOV. much smaller and shorter than the second and when LSIDurn:lsid:zoobank.org:act:B04C0456-EFB5-4246- pressed together the tip of the first finger reaches no 9559-BD26BF63FE44 further than the base of the disc of the second finger; Holotype: A preserved male specimen at the tympanum indistinct; no pale stripe along the Queensland Museum, Brisbane, Queensland, Australia, supratympanic ridge or along posterior edge of upper specimen number J29447 collected 16.7 km from the jaw; adult snout-vent length 65 mm; dorsal pattern is Mossman-Mount Molloy Road on the Mount Lewis Road, mostly grey or dull green, almost blackish and consists of far north Queensland, Australia, Latitude -16.5833 S., a series of relatively narrow dark lines and spots forming Longitude 145.25 E. a fine, continuous reticulum over the back, head and This government-owned facility allows access to its limbs. The flanks have a bluish metal sheen. White holdings. below, commonly with brown on the throat; axilla and groin flesh coloured; hind side of thighs are dark brown. Paratypes: Three preserved specimens at the Skin leathery, finely granular, or with numerous small Queensland Museum, Brisbane, Queensland, Australia, scattered warts. Granular below (M. nannotis (Andersson, specimen numbers J29455 and J29446 collected 16.7 1916), M. cottoni sp. nov. and M. lorica (Davies and km from the Mossman-Mount Molloy Road on the Mount McDonald, 1979)) (nominate subgenus), or: Lewis Road, far north Queensland, Australia, Latitude - 16.5833 S., Longitude 145.25 E and specimen number 2/ Vomerine teeth conspicuous in two rows on a line with J52942 collected from Mount Lewis, Queensland, the hind edge of the choanae; with or without pectoral Australia, Latitude -16.5736 S., Longitude 145.2633 E. fold; fingers with conspicuous webbing, reaching at least as far as the base of the penultimate phalanx of the Diagnosis: Until now, Mosleyia michaelsmythi sp. nov. fourth finger; hind edge of forearm is smooth or with at has been treated as a northern outlier population of most a few, low, discontinuous tubercles; hind edge of Mosleyia nyakalensis (Liem, 1974), more commonly foot is smooth; hind side of thighs more or less uniform known as “Litoria nyakalensis” as defined by Liem and without black and yellow markings or marbling; (1974), Anstis (2013) and Cogger (2014). colour brown to blackish above, never green, with or The two species are readily separated as follows: In life without a broad, darker vertebral patch and other adult M. nyakalensis are a uniform slate colour to greyish contrasting markings; a dark dorsal patch, if present, is brown dorsally, sometimes with irregular dark markings in inconspicuous and commencing from a line joining the the form of spots of mottling. By contrast M. centre of each eye; tympanum covered by skin (adult size michaelsmythi sp. nov. is a markedly lighter coloured frog is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. being cream, yellow or beige dorsally, sometimes with nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola irregular dark markings in the form of spots or mottling. (Liem, 1974)) (subgenus Amnisrana subgen. nov.). M. nyakalensis of both sexes has a brown iris. By According to Duellman et al. (2016) species within the contrast M. michaelsmythi sp. nov. males have an subgenus Amnisrana subgen. nov. diverged from those orangeish-red iris in males and in females it is beige on in the subgenus Mosleyia 12.1 MYA. Due to the top and greyish below. significant morphological divergence between the often In M. nyakalensis the ventral surfaces of the posterior sympatric forms, subgenus level recognition of the latter portion of body is cream with reddish brown overlay, group was deemed essential. versus pinkish-red in M. michaelsmythi sp. nov.. M. cottoni sp. nov. in life is depicted in Anstis (2013) on In M. nyakalensis on the fingers, the web extends to one- page 253 at top right and P. 245 at top left. third down the length of the proximal phalanx on outer M. nannotis in life is depicted in Vanderduys (2012) on margin of the third finger, but does not extend this far in page 56 (two images) and Cogger (2014) on page 176 at M. michaelsmythi sp. nov.. bottom. M. michaelsmythi sp. nov. in life is depicted in Anstis Distribution: M. cottoni sp. nov. is restricted to the (2013) on pages 266 and 267 (all photos), including northern wet tropics of North-east Queensland, Australia adults of both sexes and also tadpoles. A photo of this in a region bound by Hunter Creek, Brookland Nature species in life is also seen in Vanderduys (2012) on page Reserve (adjacent to Mount Lewis), about 60 km north of 60. Cairns, Latitude -16.6205 S., Longitude 145.3064 E, and Both M. michaelsmythi sp. nov. and M. nyakalensis are Mungumby Creek, about 25 km south of Cooktown in Far separated from all other species in the genus Mosleyia North Queensland, Latitude -15.700 S., Longitude Wells and Wellington, 1985 by the following unique suite 145.250 E, bounded to the east by the Pacific Ocean, dry of characters: Head rounded; IN/EN ratio more than 0- habitats to the west and drier lowlands immediately north 970; web on first toe on or just beyond the proximal and south. subarticular tubercle; ventral surfaces of posterior portion

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 36 Australasian Journal of Herpetology of body cream with reddish brown overlay or pinkish-red populations to safeguard the two species (at the time and an absence of dense large black nupital spines treated as a single species) can be blamed for the around the base of the thumb. presumed extinction of the two species, as similarly noted The genus Mosleyia is readily separated from all other in comments in Hoser (1991). Australasian Tree Frogs (Pelodryadidae) by one or other The cause of decline is believed to be an introduced of the two unique suites of characters: pathogen in the form of a fungus (Anstis 2013). 1/ Vomerine teeth conspicuous in two rows on a line with Etymology: Named in honour of Michael Smyth, formerly the hind edge of the choanae; with or without pectoral of Ringwood, Victoria who for many years played an fold; fingers with conspicuous webbing, reaching at least important role in the wildlife conservation efforts of the as far as the base of the penultimate phalanx of the team this author works with. This includes in fieldwork fourth finger; hind edge of forearm is smooth or with at across Australia as well as a displayer of reptiles with most a few, low, discontinuous tubercles; hind edge of Snakebusters: Australia’s best reptiles shows, being the foot is smooth; hind side of thighs more or less uniform only hands-on reptile exhibitions in Australia that let and without black and yellow markings or marbling; people hold the animals. colour brown to blackish above, never green, with or MOSLEYIA (AMNISRANA) PILLOTI SP. NOV. without a broad, darker vertebral patch and other LSIDurn:lsid:zoobank.org:act:4EC060C6-0B7C-476E- contrasting markings; a dark dorsal patch, if present, is AC25-BC7E2FCF85AC inconspicuous and commencing from a line joining the Holotype: A preserved female specimen at the centre of each eye; tympanum covered by skin (adult size Queensland Museum, Brisbane, Queensland, Australia, is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. specimen number J22643 collected from Cape nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola Tribulation, North Queensland, Australia, Latitude - (Liem, 1974)) (subgenus Amnisrana subgen. nov.) (this 16.0878 S., Longitude 145.4548 E. being a formal diagnosis for this subgenus), or: This government-owned facility allows access to its 2/ Vomerine teeth prominent and largely between the holdings. choanae. No pectoral fold; A series of small enlarged Paratypes: Two preserved female specimens at the tubercles along the hind edge of the forearm forming a Queensland Museum, Brisbane, Queensland, Australia, low but distinct crenulated ridge; fingers with strong basal specimen number J22644 collected from the type locality webbing; toes nearly fully webbed; a prominent inner above and specimen number J22645 collected from metatarsal tubercle and a small outer one. Heel of Shiptons Flat, 50 km south of Cooktown, North adpressed hind limb reaches to eye or beyond; first finger Queensland, Australia, Latitude -15.8000 S., Longitude much smaller and shorter than the second and when 145.2500 E. pressed together the tip of the first finger reaches no Diagnosis: Until now, Mosleyia pilloti sp. nov. has been further than the base of the disc of the second finger; treated as a northern outlier population of Mosleyia tympanum indistinct; no pale stripe along the rheocola (Liem, 1974), more commonly known as supratympanic ridge or along posterior edge of upper “Litoria rheocola” as defined by Liem (1974), Anstis jaw; adult snout-vent length 65 mm; dorsal pattern is (2013) and Cogger (2014). mostly grey or dull green, almost blackish and consists of The two species are readily separated as follows: Resting a series of relatively narrow dark lines and spots forming male M. rheocola show an obvious pale crescent shaped a fine, continuous reticulum over the back, head and mark on the upper flank. This is either abent or very limbs. The flanks have a bluish metal sheen. White indistinct in M. pilloti sp. nov.. below, commonly with brown on the throat; axilla and groin flesh coloured; hind side of thighs are dark brown. While both species posess small scattered spinose Skin leathery, finely granular, or with numerous small tubercles on the dorsum and uppel limbs, these are scattered warts. Granular below (M. nannotis (Andersson, prominent in M. pilloti sp. nov. versus reduced and 1916), M. cottoni sp. nov. and M. lorica (Davies and blunted in form in M. rheocola.Tubercles on the lower McDonald, 1979)) (nominate subgenus). flanks of M. rheocola are numerous, obvious and raised, versus not so in M. pilloti sp. nov.. According to Duellman et al. (2016) species within the subgenus Amnisrana subgen. nov. diverged from those As mid-sized pre-metamophosing tadpoles, M. pilloti sp. in the subgenus Mosleyia 12.1 MYA. nov. have a dorsum that is light grey in colour, with bold and distinctive dark brownish-black markings. At the Distribution: Mosleyia michaelsmythi sp. nov. is same life stage M. rheocola tadpoles are with a yellowish restricted to the northern wet tropics region of Australia dorsum which has a thick peppering of orange and with a range extending north from the southern side of brown, especially posterior to the eyes and anterior to the Mount Lewis, far north Queensland, Latitude -16.5903 S., tail. Comparative photos of tadpoles of both species are Longitude 145.2750 E. and south of Mungumby Creek, in Anstis (2013) on page 295 with M. pilloti sp. nov. 25 km south of Cooktown, far north Queensland, Latitude identified as “Carbine Tableland, Qld” and M. rheocola -15.72 S., Longitude 145.25 E. identified as “Henrietta Creek, Palmerston National Park, Conservation: Neither M. michaelsmythi sp. nov. or M. Qld”. nannotis have been seen alive in the wild since about Both M. piloti and M. rheocola are separated from all 1990 (Hero and Fickling, 1994) and the two species may other species in the genus Mosleyia Wells and already be extinct. Wellington, 1985 by the following unique suite of The reckless Queensland Government policy of characters: Head not rounded; IN/EN ratio less than 0- prohibiting the establishment of captive breeding 970; web on first toe not beyond proximal subarticular

tubercle; webbing on outer margin of third finger reaches on page 294 middle and bottom right and in Cogger base of penultimate phalanx; ventral surfaces of posterior (2014) on page 186 at top right. portion of body not cream with reddish brown or pinkish M. rheocola in life is depicted in Vanderduys (2012) on red tinge; snout longer than eye diameter; snout bluntly page 67, bottom left and Anstis (2013) on page 294 top rounded; loreal region not strongly concave; prepollex right. normal and not enlarged; nuptial pad small, spinules fine Distribution: Mosleyia piloti sp. nov. is restricted to the and not large with coarse spinules; forearm of males northern wet tropics region of Australia with a range normal, not robust; single outer metacarpal tubercle on extending north of Port Douglas, far north Queensland base of palm and an elongated inner metacarpal tubercle Latitude 16.4836 S., Longitude 145.4653 E. and south of on the proximo-ventral inner surface of the metacarpal of Cooktown in far North Queensland (northern limit of first finger; Fingers moderately webbed: between first and range being near the small village of Rossville, Latitude second fingers free of web; web present between second 15.7005 S., Longitude 145.2542 E.). and third and third and fourth fingers; adult less than 40 Conservation: A major decline in numbers in the late mm in body length. 1980’s appears to have reversed in some (mainly lower The genus Mosleyia is readily separated from all other elevation) areas. However the species must be deemed Australasian Tree Frogs (Pelodryadidae) by one or other at high risk of extinction. of the two unique suites of characters: Refer to the relevant comments in Hoser (1991, 2019a, 1/ Vomerine teeth conspicuous in two rows on a line with 2019b). the hind edge of the choanae; with or without pectoral Etymology: Named in honour of Christian Pillot, formerly fold; fingers with conspicuous webbing, reaching at least of Ringwood, Victoria who for many years played an as far as the base of the penultimate phalanx of the important role in the wildlife conservation efforts of the fourth finger; hind edge of forearm is smooth or with at team this author works with. This includes in fieldwork most a few, low, discontinuous tubercles; hind edge of across Australia as well as a displayer of reptiles with foot is smooth; hind side of thighs more or less uniform Snakebusters: Australia’s best reptiles shows, being the and without black and yellow markings or marbling; only hands-on reptile exhibitions in Australia that let colour brown to blackish above, never green, with or people hold the animals. without a broad, darker vertebral patch and other DARANINANURINI TRIBE NOV. contrasting markings; a dark dorsal patch, if present, is inconspicuous and commencing from a line joining the DARANINANURA GEN. NOV. centre of each eye; tympanum covered by skin (adult size LSIDurn:lsid:zoobank.org:act:E6C1197B-B538-443D- is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. 996F-9AA74B74A1BE nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola Type species: Litoria brevipalmata Tyler, Martin and (Liem, 1974)) (subgenus Amnisrana subgen. nov.) (this Watson, 1972. being a formal diagnosis for this subgenus), or: Diagnosis: The genus Daraninanura gen. nov. 2/ Vomerine teeth prominent and largely between the monotypic for the type species D. brevipalmata (Tyler, choanae. No pectoral fold; A series of small enlarged Martin and Watson, 1972) is readily separated from all tubercles along the hind edge of the forearm forming a other Australasian Tree Frogs (Pelodryadidae) by the low but distinct crenulated ridge; fingers with strong basal following unique suite of characters: Rich brown to webbing; toes nearly fully webbed; a prominent inner chocolate brown above, occasionally with scattered small metatarsal tubercle and a small outer one. Heel of black flecks. There is a wide canthal stripe running from adpressed hind limb reaches to eye or beyond; first finger snout to eye, continuing past the eye as a wide black much smaller and shorter than the second and when band, almost over-writing the standard (for frogs) sized pressed together the tip of the first finger reaches no tympanum, continuing to the flank and sometimes further than the base of the disc of the second finger; bordered above with white or yellow. The upper lip has a tympanum indistinct; no pale stripe along the narrow white or yellow stripe, narrowly edged below with supratympanic ridge or along posterior edge of upper brown, which continues as a glandular stripe from the jaw; adult snout-vent length 65 mm; dorsal pattern is angle of the mouth to the base of the forearm. The lower mostly grey or dull green, almost blackish and consists of flanks are yellowish with scattered black spots, flecks or a series of relatively narrow dark lines and spots forming peppering. Groin is green or blue green. There are no red a fine, continuous reticulum over the back, head and or orange spots on the hind side of the thighs. There is a limbs. The flanks have a bluish metal sheen. White dark stripe along the front edge of the hindlimb. Venter is below, commonly with brown on the throat; axilla and white to light yellow. Top of iris is silver to gold in colour. groin flesh coloured; hind side of thighs are dark brown. Skin is smooth to slightly leathery above and coarsely Skin leathery, finely granular, or with numerous small granular below. Snout is rounded in shape. Vomerine scattered warts. Granular below (M. nannotis (Andersson, teeth are prominent between the choanae. There is no 1916), M. cottoni sp. nov. and M. lorica (Davies and pectoral fold. Finger and toe discs are of medium size, McDonald, 1979)) (nominate subgenus). fingers are unwebbed and toes about one third webbed. According to Duellman et al. (2016) species within the There is a prominent inner metatarsal tubercle and an subgenus Amnisrana subgen. nov. diverged from those indistinct small outer tubercle. The second finger is in the subgenus Mosleyia 12.1 MYA. longer than the first, the first finger being so short that Mosleyia pilloti sp. nov. in life is depicted in Vanderduys when pressed together with the second, it reaches no (2012) on page 67, bottom right as well as Anstis (2013) further than the base of the disc of the second. Duellman et al. (2016) found the type and only species in Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 38 Australasian Journal of Herpetology the genus Daraninanura gen. nov. to have diverged from genus named Fiacumminganurea gen. nov.. its nearest living relative 35.1 MYA, necessitating the The three morphologically similar species within transfer of this species to a new genus and the genus to Fiacumminganurea gen. nov. are readily separated from a new tribe as done within this paper. all other Australasian Tree Frogs (Pelodryadidae) by the Photos of Daraninanura brevipalmata (Tyler, Martin and following unique suite of characters: A colouration that is Watson, 1972), can be found in Cogger (2014) on page grey to green above, with irregular darker mottling, 153 (two images), Vanderduys (2012) on page 27, Eipper marbling or flecks, the latter often forming irregular cross- and Rowland (2018) on page 119 at top and Anstis bands on the limbs. (2013) on pages 156 (right side), 157 and 158. Ventral surface white or yellow or white becoming yellow Distribution: Known only from wet sclerophyll forests of towards the rear. Lower and concealed surface of limbs the north coast of New South Wales, north from about are yellowish. Skin is somewhat leathery, with a few tiny Gosford and into the wetter parts of south-east whitish tubercles on the back becoming numerous on the Queensland, Australia. sides. Skin below is granular. Small but prominent Etymology: Named in honour of Dara Nin of Ringwood, vomerine teeth are located mostly behind the choanae. A Victoria, Australia, who for many years has been a pectoral fold is indistinct. Finger and toes discs are member of the team doing Reptile Parties and Reptile moderate, being only a little wider than the digits. Fingers Shows, with Snakebusters: Australia’s best reptiles with distinct basal webbing and the toes are fully webbed. shows. He has helped to educate Australians about There is a small but prominent inner metatarsal tubercle, wildlife and wildlife conservation, with Australia’s only no outer one. Tympanum is indistinct. Second finger is hands on reptile shows that let people hold the animals. larger than the first; adult size to 45 mm in lenth. Content: Daraninanura brevipalmata (Tyler, Martin and The tadpole is free-swimming, elongated and flattened, Watson, 1972) (monotypic). and reaches a total length of 40 mm prior to FIACUMMINGANURINI TRIBE NOV. metamorphosis. The body is dark brown to black above, FIACUMMINGANURA GEN. NOV. with fine silver chromatophores extending onto the flanks. Darker spots may be present on the dorsal surface, while LSIDurn:lsid:zoobank.org:act:91ADD2D0-9D5C-4882- the ventral surface is darkly pigmented. The tail fin and 90B7-277A74B6E7B5 muscle are covered with fine melanophores. The tail is Type species: Fiacumminganura fiacummingae sp. nov. moderately thick and has a rounded tip. The eyes are (this paper). dorso-lateral, and the mouth is ventral. The oral disc is Diagnosis: Until now the single putative species within large relative to closely-related species, and the oral this genus, has been known under various names. papillae have a wide anterior gap. There are two rows of It was originally described as Hyla maculata Spencer, anterior labial teeth and three posterior rows (Hero et al. 1901, later transferred to the genus Litoria Tschudi, 1838 1995; Anstis 2013). by Cogger et al. (1983). Dubois (1984) assigned a In terms of morphologically similar and potentially replacement name Litoria spenceri to the same taxon, as sympatric species the warty back of Fiacumminganurea the species name maculata had been earlier applied to a gen. nov. distinguishes species in this genus from the different species (preoccupied by Hyla maculata Gray, morphologically similar species Dryopsophus nudidigita 1830). The putative taxon “Litoria spenceri”, has been (Copland, 1962) and its lack of a distinct tympanum kept in that genus by most authors since. See for distinguishes it from Dryopsophus citropa (Dümeril and example Tyler (1992), Anstis (2013), Cogger (2014) and Bibron, 1841). Eipper and Rowland (2018). Distribution: Eastern Victoria and immediately adjacent Contrary to this has been a realisation by many southern New South Wales, on both sides of the Great herpetologists that the continued placement of Dividing Range. morphologically and genetically divergent species within Etymology: The genus and type species are named in the single putative genus Litoria is untenable. honour of Fia Cumming of Canberra, ACT, Australia, in Ranoidea spenceri (Dubois, 1984) (Dubois and Frétey, recognition for her services to wildlife conservation in 2016), was one such attempt to place this divergent Australia as detailed in Hoser (1993) and Hoser (1996). taxon in another genus. Duellman et al. (2016) placed Content: Fiacumminganurea fiacummingae sp. nov. putative “Litoria spenceri” within the genus Dryopsophus (type species); F. spenceri (Dubois, 1984); F. timdalei sp. Fitzinger, 1843, with the type spcies Hyla citropa Dümeril nov. and Bibron, 1841. FIACUMMINGANUREA FIACUMMINGAE SP. NOV. In line with Pyron and Weins (2011), Duellman et al. LSIDurn:lsid:zoobank.org:act:93A202C8-5C46-41A5- (2016) found that the divergent “Litoria spenceri” was 93F3-0486501B4B43 most closely related to the species group including Dryopsophus citropa. However there was still a 22.3 MYA Holotype: A preserved specimen at the National divergence between the two groups of species. Museum of Victoria, Melbourne, Victoria, Australia, specimen number D73096 collected at Bogong Creek, Hence the erection of a new genus to accommodate Kosciuszko National Park, New South Wales, Australia, putative “Litoria spenceri” as done herein. Latitude -36.1 S., Longitude 148.4 E. Investigations have also shown that putative “Litoria spenceri” in fact consists of three divergent species and This government-owned facility allows access to its the two unnamed forms are formally named herein, one holdings. of them being assigned the type species for this new Paratypes: Fifteen preserved specimens at the National

Museum of Victoria, Melbourne, Victoria, Australia, F. fiacummingae sp. nov. is found in the Upper Murray specimen numbers D72995, D72996, D72997, D72998, Basin, comprising the West Kiewa, Mitta Mitta, Indi and D72999, D73060, D73061, D73062, D73063, D73064, Bundarra Rivers as well as Buffalo, Snowy, Lightning, D73065, D73097, D73098, D73099 and D73100 all Wheeler and Bogong Creeks. collected at Bogong Creek, Kosciuszko National Park, F. timdalei sp. nov. previously known as the so-called New South Wales, Australia, Latitude -36.1 S., Longitude “Green Form” of “Litoria spenceri” is found in the upper 148.4 E. Goulburn River basin including the Goulburn, Taponga, Diagnosis: That there are three highly divergent species Big, Black, North Jamieson and Howqua Rivers as well until now all placed within the single species described as as Snake, White and Still Creeks. Litoria spenceri (Dubois, 1984), herein placed in the new The three species are readily separated from one genus Fiacumminganurea gen. nov. has been known for another on the basis of colour of adult specimens. some time (Minister for the Environment, Commonwealth F. spenceri is olive grey above, blotched with darker Government of Australia. 2017). markings, with the same on the upper surfaces of the All three populations (species) have declined sharply due limbs. The flanks are generally light, but with prominent to the effects of Batrachochytrium dendrobatidis large dark spots or blotches. (Gillespie et al. 2015) and therefore with reference to There is a purply-grey line running from the snout, Hoser (2019a, 2019b) it is of critical importance that all through the top of the eye and down to the top of the three species be formally identified and named forelimb, where it terminates. Below this line, the snout is immediately so that all can have proper conservation generally green. The top of the head is prominently management plans enacted. spotted. With reference to Hoser (2020a), it is also important that F. timdalei sp. nov. is generally lime green above no all three species (AKA divergent genetic lineages) be markings on the upper surfaces of the limbs save for kept intact and not otherwise be destroyed by well- limited peppering on the front limbs. Unlike both other meaning but unwise translocations of specimens. species it also has a distinctive yellow-gold line The type form of L. spenceri and Hyla maculata Spencer, (occasionally whitish) running from the snout, through the 1901 are both based on a holotype specimen at the nostril, above the eye and down to the upper arm, where National Museum of Victoria, in Melbourne, Australia, it widens and fades on the anterior flank. There is a specimen number D8498 allegedly from “Powong [= blackish-grey border to the lower edge of this line on the Poowong], Victoria”, Australia. snout and to the back of the head. The flanks are whitish To date this locality information has not been questioned with purply-grey mottling. in the literature, but clearly the location is in error. F. fiacummingae sp. nov. is generally an olive grey colour The Poowong area in West Gippsland is relatively flat above, like in F. spenceri, but is separated from that and devoid of the mountain stream habitats occupied by species by having no obvious contrasting dark blotches the putative species. on the upper body or upper surfaces of the limbs. Furthermore this site is disconnected from all known Any markings on the body are indistinct and consist of populations by way of drainage systems and potential numerous tiny darker spots and patches, giving the frog a migration routes. peppered appearance. Limbs are greenish to grey and No further specimens have been found there since the have indistinct lighter markings in the form of spots, lodgement of the holotype and none would be expected peppering or patches. to be found there either. F. fiacummingae sp. nov. is also characterised by a Inspection of the said specimen and also the original distinctive lime-green semicircle under the eye, which descriptions of Spencer (1901) and Dubois (1984) spreads in less intensity to the nearby upper lip, posterior confirms it is of the nearest geographically proximal to the nostril. lineage, being that from the south-side of the Great There is no obvious spotting on the upper surface of the Dividing Range, currently known from the Wongungarra head in this species as seen in F. spenceri. River drainage, being part of the Mitchell River system in Both F. fiacummingae sp. nov. and F. timdalei sp. nov. eastern Victoria. Hence it is that form that must be have larger tubercles or warts on the upper surface of the treated as nominate F. spenceri. body, being generally tubercular across the entire upper Even allowing for human-created declines in the last 250 body than is seen in F. spenceri. years, the nearest potential drainage system for the taxon Comparative illustrative photos of F. timdalei sp. nov. and to occur near the the alleged type locality would be those F. spenceri can be seen (in that order) on pages 36 and of the Latrobe Valley which like the Mitchell River drain 37 of Hero et al. (1991). into Victoria’s East Gippsland Lakes region, the likely The three morphologically similar species within collection point still being a watershed on the south-side Fiacumminganurea gen. nov. are readily separated from of the Great Dividing Range. all other Australasian Tree Frogs (Pelodryadidae) by the Nominate F. spenceri is therefore treated as the form following unique suite of characters: A colouration that is currently only known with certainty from the Wongungarra grey to green above, with irregular darker mottling, River system in eastern Victoria and at its widest those marbling or flecks, the latter often forming irregular cross- from south of the Great Dividing Range. bands on the limbs. Ventral surface white or yellow or All other forms previously treated as that taxon are herein white becoming yellow towards the rear. Lower and described as new species. concealed surface of limbs are yellowish. Skin is somewhat leathery, with a few tiny whitish tubercles on Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 40 Australasian Journal of Herpetology the back becoming numerous on the sides. Skin below is This government-owned facility allows access to its granular. Small but prominent vomerine teeth are located holdings. mostly behind the choanae. A pectoral fold is indistinct. Paratypes: Two preserved specimens at the South Finger and toes discs are moderate, being only a little Australian Museum, Adelaide, South Australia, Australia, wider than the digits. Fingers with distinct basal webbing specimen numbers R46682 and R46684, collected from and the toes are fully webbed. There is a small but Richies Hut at the Howkwa River, Victoria, Australia, prominent inner metatarsal tubercle, no outer one. Latitude -37.1944 S., Longitude 146.475 E. Tympanum is indistinct. Second finger is larger than the Diagnosis: That there are three highly divergent species first; adult size to 45 mm in lenth. until now all placed within the single species described as The tadpole is free-swimming, elongated and flattened, Litoria spenceri (Dubois, 1984), herein placed in the new and reaches a total length of 40 mm prior to genus Fiacumminganurea gen. nov. has been known for metamorphosis. The body is dark brown to black above, some time (Minister for the Environment, Commonwealth with fine silver chromatophores extending onto the flanks. Government of Australia. 2017). Darker spots may be present on the dorsal surface, while All three populations (species) have declined sharply due the ventral surface is darkly pigmented. The tail fin and to the effects of Batrachochytrium dendrobatidis muscle are covered with fine melanophores. The tail is (Gillespie et al. 2015) and therefore with reference to moderately thick and has a rounded tip. The eyes are Hoser (2019a, 2019b) it is of critical importance that all dorso-lateral and the mouth is ventral. The oral disc is three species be formally identified and named large relative to closely-related species, and the oral immediately so that all can have proper conservation papillae have a wide anterior gap. There are two rows of management plans enacted. anterior labial teeth and three posterior rows (Hero et al. With reference to Hoser (1995) and Hoser (2020a), it is 1995; Anstis 2013). also important that all three species (AKA divergent In terms of morphologically similar and potentially genetic lineages) be kept intact and not otherwise be sympatric species the warty back of Fiacumminganurea destroyed by well-meaning but unwise translocations of gen. nov. distinguishes species in this genus from the specimens. morphologically similar species Dryopsophus nudidigita The type form of L. spenceri and Hyla maculata Spencer, (Copland, 1962) and its lack of a distinct tympanum 1901 are both based on a holotype specimen at the distinguishes it from Dryopsophus citropa (Dümeril and National Museum of Victoria, in Melbourne, Australia, Bibron, 1841). specimen number D8498 allegedly from “Powong [= F. fiacummingae sp. nov. in life can be found in Cogger Poowong], Victoria”, Australia. (2014) at page 189 and Anstis (2013), on page 309, top To date this locality information has not been questioned left and second down from top on right and again on in the literature, but clearly the location is in error. page 310 (tadpoles). The Poowong area in west Gippsland is relatively flat and F. timdalei sp. nov. in life can be found in Hero et al. devoid of the mountain stream habitats occupied by the (1991) on page 36, Anstis (2013) on page 309, top right, putative species. as well as in Eipper and Rowland (2018) on page 152 Furthermore this site is disconnected from all known bottom. populations by way of drainage systems and potential F. spenceri in life can be found in Hero et al. (1991) on migration routes. page 37. No further specimens have been found there since the Photos of all three species can be found online on photo- lodgement of the holotype and none would be expected sharing sites such as “www.flickr.com”, many not to be found there either. identifiable by location, but easily distinguished from one Inspection of the said specimen and also the original another on the basis of the diagnostic characters given descriptions of Spencer (1901) and Dubois (1984) above. confirms it is of the nearest geographically proximal Distribution: F. fiacummingae sp. nov. is found in the lineage, being that from the south-side of the Great Upper Murray Basin, comprising the West Kiewa, Mitta Dividing Range, currently known from the Wongungarra Mitta, Indi and Bundarra Rivers as well as Buffalo, River drainage, being part of the Mitchell River system in Snowy, Lightning, Wheeler and Bogong Creeks. eastern Victoria. Etymology: This species and the genus it is placed in Hence it is that form that must be treated as nominate F. are named in honour of Fia Cumming of Canberra, ACT, spenceri. Australia, in recognition for her services to wildlife Even allowing for human-created declines in the last 250 conservation in Australia as detailed in Hoser (1993) and years, the nearest potential drainage system for the taxon Hoser (1996). to occur near the the alleged type locality would be those FIACUMMINGANUREA TIMDALEI SP. NOV. of the Latrobe Valley which like the Mitchell River drain LSIDurn:lsid:zoobank.org:act:19E954EC-4E8F-4D6F- into Victoria’s East Gippsland Lakes region, the likely 9E58-4285D59BB988 collection point still being a watershed on the south-side Holotype: A preserved male specimen at the National of the Great Dividing Range. Museum of Victoria, Melbourne, Victoria, Australia, Nominate F. spenceri is therefore treated as the form specimen number D48831 collected from Bindaree Hut, currently only known with certainty from the Wongungarra Howqua River, 9.5 km West of Mount Howitt, Victoria, River system in eastern Victoria or at its widest being Australia, Latitude -37.18 S., Longitude 146.53 E. from south of the Great Dividing Range.

All other forms previously treated as that taxon are herein surface white or yellow or white becoming yellow towards described as new species. the rear. Lower and concealed surface of limbs are F. fiacummingae sp. nov. is found in the Upper Murray yellowish. Skin is somewhat leathery, with a few tiny Basin, comprising the West Kiewa, Mitta Mitta, Indi and whitish tubercles on the back becoming numerous on the Bundarra Rivers as well as Buffalo, Snowy, Lightning, sides. Skin below is granular. Small but prominent Wheeler and Bogong Creeks. vomerine teeth are located mostly behind the choanae. A F. timdalei sp. nov. previously known as the so-called pectoral fold is indistinct. Finger and toes discs are “Green Form” of “Litoria spenceri” is found in the upper moderate, being only a little wider than the digits. Fingers Goulburn River basin including the Goulburn, Taponga, with distinct basal webbing and the toes are fully webbed. Big, Black, North Jamieson and Howqua Rivers as well There is a small but prominent inner metatarsal tubercle, as Snake, White and Still Creeks. no outer one. Tympanum is indistinct. Second finger is The three species are readily separated from one larger than the first; adult size to 45 mm in lenth. another on the basis of colour of adult specimens. The tadpole is free-swimming, elongated and flattened, F. spenceri is olive grey above, blotched with darker and reaches a total length of 40 mm prior to markings, with the same on the upper surfaces of the metamorphosis. The body is dark brown to black above, limbs. The flanks are generally light, but with prominent with fine silver chromatophores extending onto the flanks. large dark spots or blotches. Darker spots may be present on the dorsal surface, while the ventral surface is darkly pigmented. The tail fin and There is a purply-grey line running from the snout, muscle are covered with fine melanophores. The tail is through the top of the eye and down to the top of the moderately thick and has a rounded tip. The eyes are forelimb, where it terminates. Below this line, the snout is dorso-lateral and the mouth is ventral. generally green. The top of the head is prominently spotted. The oral disc is large relative to closely-related species, and the oral papillae have a wide anterior gap. There are F. timdalei sp. nov. is generally lime green above no two rows of anterior labial teeth and three posterior rows markings on the upper surfaces of the limbs save for (Hero et al. 1995; Anstis 2013). limited peppering on the front limbs. Unlike both other species it also has a distinctive yellow-gold line In terms of morphologically similar and potentially (occasionally whitish) running from the snout, through the sympatric species the warty back of Fiacumminganurea nostril, above the eye and down to the upper arm, where gen. nov. distinguishes species in this genus from the it widens and fades on the anterior flank. There is a morphologically similar species Dryopsophus nudidigita blackish-grey border to the lower edge of this line on the (Copland, 1962) and its lack of a distinct tympanum snout and to the back of the head. The flanks are whitish distinguishes it from Dryopsophus citropa (Dümeril and with purply-grey mottling. Bibron, 1841). F. fiacummingae sp. nov. is generally an olive grey colour F. fiacummingae sp. nov. in life can be found in Cogger above, like in F. spenceri, but is separated from that (2014) at page 189 and Anstis (2013), on page 309, top species by having no obvious contrasting dark blotches left and second down from top on right and again on on the upper body or upper surfaces of the limbs. page 310 (tadpoles). Any markings on the body are indistinct and consist of F. timdalei sp. nov. in life can be found in Hero et al. numerous tiny darker spots and patches, giving the frog a (1991) on page 36, Anstis (2013) on page 309, top right, peppered appearance. as well as in Eipper and Rowland (2018) on page 152 bottom. Limbs are greenish to grey and have indistinct lighter markings in the form of spots, peppering or patches. F. spenceri in life can be found in Hero et al. (1991) on page 37. F. fiacummingae sp. nov. is also characterised by a distinctive lime-green semicircle under the eye, which Photos of all three species can be found online on photo- spreads in less intensity to the nearby upper lip, posterior sharing sites such as “www.flickr.com”, many not to the nostril. identifiable by location, but easily distinguished from one another on the basis of the diagnostic characters given There is no obvious spotting on the upper surface of the above. head in this species as seen in F. spenceri. Distribution: F. timdalei sp. nov. previously known as the Both F. fiacummingae sp. nov. and F. timdalei sp. nov. so-called “Green Form” of “Litoria spenceri” is found in have larger tubercles or warts on the upper surface of the the upper Goulburn River basin including the Goulburn, body, being generally tubercular across the entire upper Taponga, Big, Black, North Jamieson and Howqua Rivers body than is seen in F. spenceri. as well as Snake, White and Still Creeks. Comparative illustrative photos of F. timdalei sp. nov. and F. spenceri can be seen (in that order) on pages 36 and Etymology: F. timdalei sp. nov. is named in honour of 37 of Hero et al. 1991. Tim Dale of Warrandyte, Victoria, in recognition of his special connection with the drainages in the area this The three morphologically similar species within species occurs, including those that flow off Mount Buller, Fiacumminganurea gen. nov. are readily separated from in which he has crashed when riding on his snowboard all other Australasian Tree Frogs (Pelodryadidae) by the too many times, sometimes after hitting a grazing following unique suite of characters: wombat Vombatus ursinus (Shaw, 1800) at very high A colouration that is grey to green above, with irregular speed. darker mottling, marbling or flecks, the latter often forming irregular cross-bands on the limbs. Ventral

DRYOPSOPHINA SUBTRIBE NOV. LEUCOLATERA SUBGEN. NOV. DRYOPSOPHUS FITZINGER, 1843 LSIDurn:lsid:zoobank.org:act:82F4DFED-E14C-4FB3- Type species: Hyla citropa Dümeril and Bibron, 1841. A5FE-4D52206C6E0B Diagnosis: Frogs in the genus Dryopsophus Fitzinger, Type species: Litoria subglandulosa Tyler and Anstis, 1843 as defined herein, are readily separated from all 1983. other Australasian Tree Frogs (Pelodryadidae) by the Diagnosis: The two recognized species in the subgenus following unique suite of characters: Leucolatera subgen. nov. can be separated from all other Vomerine teeth present and the hind edge is behind the Australasian Tree Frogs (Pelodryadidae) by the following choanae; fingers are free or only webbed at the base; the unique suite of characters: first finger is much shorter and smaller than the second Vomerine teeth present and the hind edge is behind the and when pressed together the tip of the first finger choanae; fingers are free or only webbed at the base; the reaches no further than the base of the disc of the first finger is much shorter and smaller than the second second finger; the heel of the adpressed hindlimb and when pressed together the tip of the first finger reaches to the eye or beyond; and one or other of the reaches no further than the base of the disc of the following three suites of characters: second finger; the heel of the adpressed hindlimb 1/ There is a narrow light line along at least the posterior reaches to the eye or beyond; there is a narrow light line edge of the upper jaw, from below the eye to the along at least the posterior edge of the upper jaw, from glandular region behind the angle of the mouth; the below the eye to the glandular region behind the angle of tympanum is distinct; the tympanic annulus is clearly the mouth; the tympanum is indistinct; the tympanic visible (subgenus Dryopsophus), or: annulus is indistinct. 2/ There is a narrow light line along at least the posterior In turn each of the other subgenera within Dryopsophus edge of the upper jaw, from below the eye to the are separated from the subgenus Leucolatera subgen. glandular region behind the angle of the mouth; the nov. by one or other of the following suites of characters: tympanum is indistinct; the tympanic annulus is indistinct, 1/ There is a narrow light line along at least the posterior (subgenus Leucolatera subgen. nov.), or: edge of the upper jaw, from below the eye to the 3/ There is no light line along the posterior edge of the glandular region behind the angle of the mouth; the upper jaw; there is a narrow light green, white or gold tympanum is distinct; the tympanic annulus is clearly stripe, sometimes with indefinite edges, but always visible (subgenus Dryopsophus), or: bordered below with dark brown or black along the supra- 2/ There is no light line along the posterior edge of the tympanic ridge; fingers have barely a trace of webbing upper jaw; there is a narrow light green, white or gold and the white or gold supratympanic streak is sharp stripe, sometimes with indefinite edges, but always edged (subgenus Ausverdarana subgen. nov.) bordered below with dark brown or black along the supra- The nominate subgenus Dryopsophus Fitzinger, 1843 tympanic ridge; fingers have barely a trace of webbing only includes the type species D. citropa (Dümeril and and the white or gold supratympanic streak is sharp Bibron, 1841), which is herein split into two edged (subgenus Ausverdarana subgen. nov.). morphologically divergent and apparently allopatric Duellman et al. (2016) found that the species in this subspecies. It is distributed from the lower north coast of subgenus diverged from the other two subgenera about New South South Wales, southwards along the coast and 15.3 MYA. nearby ranges south to north-east Victoria. Distribution: The coast and ranges from just north According to Duellman et al. (2016), the assemblage of (within 50 km) of the Queensland / New South Wales frogs within the genus Dryopsophus, diverged from its border, extending south for about 500 km in a straight nearest living relatives, being species within line into northern New South Wales, Australia. Fiacumminganurea gen. nov. (this paper), about 22.3 Etymology: “Leuco” in Latin means white and so the MYA. name Leucolatera refers to the white typically found on Distribution: South-eastern Australia from about the flanks (lateral surfaces AKA “Latera” in Latin) of the Kroombit Tops, Queensland in the North, generally along relevant frog species. wetter regions near the coast and adjacent ranges to Content: Dryopsophus (Leucolatera) subglandulosa north-east Victoria. (Tyler and Anstis, 1983) (type species); D. (Leucolatera) Content: Dryopsophus citropa (Dümeril and Bibron, daviesae (Mahony, Knowles, Foster and Donnellan, 1841) (type species); D. barringtonensis (Copland, 1957); 2001). D. daviesae (Mahony, Knowles, Foster and Donnellan, AUSVERDARANA SUBGEN. NOV. 2001); D. jarrodthomsoni sp. nov. (Warwick, Qld); D. LSIDurn:lsid:zoobank.org:act:30C68D75-FF13-439E- kroombitensis (Hoskin, Hines, Meyer, Clarke and 918F-EB1CE72F9612 Cunningham, 2013); D. nudidigita (Copland, 1962); D. Type species: Hyla phyllochroa Günther, 1863. pearsoniana (Copland, 1961); D. phyllochroa Günther, Diagnosis: Frogs in the subgenus Ausverdarana 1863 (type species); D. piperata (Tyler and Davies, subgen. nov. as defined herein, are readily separated 1985); D. subglandulosa (Tyler and Anstis, 1983). from all other Australasian Tree Frogs (Pelodryadidae) by the following unique suite of characters: Vomerine teeth present and the hind edge is behind the choanae; fingers are free or only webbed at the base; the first finger is much shorter and smaller than the second

and when pressed together the tip of the first finger Diagnosis: Until now, the taxon Dryopsophus citropa reaches no further than the base of the disc of the gippslandensis subsp. nov. has been treated by all second finger; the heel of the adpressed hindlimb herpetologists as the southernmost population of the reaches to the eye or beyond; there is no light line along species D. citropa (Dümeril and Bibron, 1841) and no the posterior edge of the upper jaw; there is a narrow authors I am aware of have ever speculated of light green, white or gold stripe, sometimes with indefinite differences between the Victorian form and type form edges, but always bordered below with dark brown or (type locality of Sydney, New South Wales). black along the supra-tympanic ridge; fingers have barely However having collected many hundreds of specimens a trace of webbing and the white or gold supratympanic over a period exceeding 50 years, both in and around the streak is sharp edged. Sydney region as well as in north-east Victoria, I have In turn the other two subgenera within Dryopsophus been aware of sufficient morphological differences Fitzinger, 1843 are separated from Ausverdarana between the populations to warrant taxonomic division subgen. nov. by one or other of the following suites of since at least the 1990’s. characters: Both subspecies would be identified as D. citropa (under 1/ There is a narrow light line along at least the posterior the name “Litoria citropa”) using the key in Cogger edge of the upper jaw, from below the eye to the (2014). glandular region behind the angle of the mouth; the D. citropa gippslandensis subsp. nov. is essentially tympanum is distinct; the tympanic annulus is clearly confined to north-east Victoria and immediately adjacent visible (subgenus Dryopsophus), or: parts of south-east New South Wales, north to 2/ There is a narrow light line along at least the posterior Merimbula. edge of the upper jaw, from below the eye to the After an apparent break in the distribution of the species glandular region behind the angle of the mouth; the (of about 20 km straight line measurement or 30 km by tympanum is indistinct; the tympanic annulus is indistinct, road), D. citropa citropa is found north from about Bega (subgenus Leucolatera subgen. nov.). in New South Wales to the region of the Hunter Valley, Duellman et al. (2016) found that the species in this north of Sydney and includes the form described as Hyla subgenus diverged from the closest related living jenolanensis Copland, 1957. congener in the nominate subgenus by 14.5 MYA. The two subspecies are readily separated from one Distribution: South-eastern Australia from about another by the following defining characters: Kroombit Tops, Queensland in the North, generally along D. citropa citropa from the type region in New South wetter regions near the coast and adjacent ranges to Wales have tight spaced dark flecks all over the upper north-east Victoria. body and well defined markings on the front and back Etymology: The subgenus name is derived as follows: legs. There is obvious patches of bright lime green on the “Aus” is an abbreviation of Australia, being from where lower arm and feet. There is also lots of lime green these frogs occur. “Verda” is Latin for green, being the anterior to eye on the snout. The lime green markings on dominant dorsal colouration of most species in the genus the lower sides of the back and groin are obvious, well- as adults, most of the time. defined and brilliant. “Rana” is Latin for frog and so the full translation of the By contrast, D. citropa gippslandensis subsp. nov. have name Ausverdarana means “Green Australian Frog”. well spaced dark flecks on the upper body and poorly Content: Dryopsophus (Ausverdarana) phyllochroa defined markings on the front and back legs. As a rule, (Günther, 1863) (type species); D. (Ausverdarana) there is no green on the lower arm or feet (except in barringtonensis (Copland, 1957); D. (Ausverdarana) aberrant specimens). There is also minimal green jarrodthomsoni sp. nov.; D. (Ausverdarana) kroombitensis anterior to the eye on the snout. The green on the lower (Hoskin, Hines, Meyer, Clarke and Cunningham, 2013); sides of the back and groin is faded or absent. D. (Ausverdarana) nudidigita (Copland, 1962); D. Frogs in the genus Dryopsophus Fitzinger, 1843 as (Ausverdarana) pearsoniana (Copland, 1961); D. defined herein, are readily separated from all other (Ausverdarana) piperata (Tyler and Davies, 1985). Australasian Tree Frogs (Pelodryadidae) by the following DRYOPSOPHUS (DRYOPSOPHUS) CITROPA unique suite of characters: GIPPSLANDENSIS SUBSP. NOV. Vomerine teeth present and the hind edge is behind the LSIDurn:lsid:zoobank.org:act:AEEB16B9-31AB-4480- choanae; fingers are free or only webbed at the base; the B9B8-BBEF4916AB03 first finger is much shorter and smaller than the second Holotype: A preserved specimen at the National and when pressed together the tip of the first finger Museum of Victoria, Melbourne, Victoria, Australia, reaches no further than the base of the disc of the specimen number D66034 collected at a small creek at second finger; the heel of the adpressed hindlimb Sandpatch Track, East Gippsland, Victoria, Latitude - reaches to the eye or beyond; and one or other of the 37.59 S., Longitude 149.553 E. following three suites of characters: This government-owned facility allows access to its 1/ There is a narrow light line along at least the posterior holdings. edge of the upper jaw, from below the eye to the Paratype: A preserved specimen at the National glandular region behind the angle of the mouth; the Museum of Victoria, Melbourne, Victoria, Australia, tympanum is distinct; the tympanic annulus is clearly specimen number D56458 collected at the Tambo River, visible (subgenus Dryopsophus , being monotypic for D. at a bridge on Collins Road, East Gippsland, Victoria, citropa), or: Australia, Latitude -37.55 S., Longitude 147.85 E. 2/ There is a narrow light line along at least the posterior Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 44 Australasian Journal of Herpetology edge of the upper jaw, from below the eye to the distinct black spots running across the back. glandular region behind the angle of the mouth; the In D. jarrodthomsoni sp. nov. there is a well-defined dark tympanum is indistinct; the tympanic annulus is indistinct, brown bar running from the snout, through the eye, (subgenus Leucolatera subgen. nov.), or: across all or most of the tympanum and past the top of 3/ There is no light line along the posterior edge of the the forelimb, with a thin white boundary on top. upper jaw; there is a narrow light green, white or gold Significantly this bar does not narrow as it passes stripe, sometimes with indefinite edges, but always through the tympanum, which contrasts with the situation bordered below with dark brown or black along the supra- in D. pearsoniana. In that species the bar narrows tympanic ridge; fingers have barely a trace of webbing significantly to become a thin line as it passes over the and the white or gold supratympanic streak is sharp tympanum. edged (subgenus Ausverdarana subgen. nov.). The toes of D. jarrodthomsoni sp. nov. are yellow, versus In areas where either of these subspecies of D. citropa green or brown in D. pearsoniana. occur I have found D. citropa gippslandensis subsp. nov. D. pearsoniana has obvious moderately distinct blotches in significantly greater numbers, although as rule D. or markings on the limbs, versus none in D. citropa of either subspecies are never seen in the jarrodthomsoni sp. nov.. massive numbers that other sympatric frogs in the same Numerous photos of both D. pearsoniana and D. locations commonly occur. jarrodthomsoni sp. nov. in life can be found at the website Distribution: D. citropa gippslandensis subsp. nov. is http://www.flickr.com essentially confined to north-east Victoria and Distribution: D. jarrodthomsoni sp. nov. is presently immediately adjacent parts of south-east New South known from the ranges and nearby areas west of the Wales, north to Merimbula (Latitude 36.8875 S., Gold Coast/Tweed Heads area, generally in the zone of Longitude 149.9059 E.). the headwaters of the Condamine River and other nearby Etymology: The subspecies name gippslandensis, refers drainages, in particular to the south in New South Wales to where this taxon occurs. as well headwaters of some east draining watercourses DRYOPSOPHUS (AUSVERDARANA) as well. JARRODTHOMSONI SP. NOV. Etymology: Named in in honour of Jarrod Thomson of LSIDurn:lsid:zoobank.org:act:8E210C37-8242-4035- Croydon North in Victoria, Australia, for his assistances in 98D5-1D88E2FE40DE maintaining the infrastructure at the world-class reptile Holotype: A preserved specimen at the Queensland captive breeding facility held at the Snakebusters Museum in Brisbane, Queensland, Australia, specimen Reptiles shows business address in Victoria, Australia. number: J26361 collected at Farm Creek, via Warwick, KUMANJAYIWALKERINI TRIBE NOV. Queensland, Australia, Latitude -28.3 S., Longitude 152.2 KUMANJAYIWALKERUS GEN. NOV. E. LSIDurn:lsid:zoobank.org:act:666B94F4-985E-4E91- This government-owned facility allows access to its 8473-F7E6926C7646 holdings. Type species: Kumanjayiwalkerus kumanjayi sp. nov. Paratypes: 1/ A preserved specimen at the Queensland Diagnosis: The genera Pengilleyia Wells and Museum in Brisbane, Queensland, Australia, specimen Wellington, 1985 as defined within this paper, and number: J51998, collected at Girraween National Park, Kumanjayiwalkerus gen. nov. are as a pair, both readily Queensland, Australia, Latitude -28.83 S., Longitude separated from all other Australasian Tree Frogs 151.9371 E. (Pelodryadidae) by the following unique suite of 2/ A preserved male specimen at the Queensland characters: Museum in Brisbane, Queensland, Australia, specimen Vomerine teeth present; fingers with conspicuous number: J17328 collected at Scenic Rim, Cunninghams webbing reaching at least as far as the base of the Gap, Queensland, Australia, Latitude -28.05 S., penultimate phalanx of the fourth finger; hind edge of Longitude 152.4 E. forearm is smooth, or with at most a few low, Diagnosis: Dryopsophus jarrodthomsoni sp. nov. has discontinuous tubercles; hind edge of foot is smooth; until now been treated as a population of hind side of thighs with contrasting black and yellow bars D. phyllochroa (Günther, 1863) or more recently D. or marbling, at least dorsally. pearsoniana (Copland, 1961) as defined by Anstis (2013) The genus Pengilleyia Wells and Wellington, 1985 is or keyed out by Cogger (2014), however the molecular readily separated from the genus Kumanjayiwalkerus evidence of Donnellan et al. (1999) indicated potential gen. nov. by having a back that is either very warty or species level divergence for the population. moderately warty, versus virtually smooth or with well D. jarrodthomsoni sp. nov. would key out as D. scattered small pointed tubercles on an otherwise pearsoniana in Cogger (2014) as a means of separating smooth body in Kumanjayiwalkerus gen. nov.. both species from all other Australasian tree frogs. Furthermore species within Pengilleyia invariably have D. jarrodthomsoni sp. nov. is readily separated from its green spots, flecks or blotches on the back versus none nearest congener D. pearsoniana, by colouration. in Kumanjayiwalkerus gen. nov.. D. jarrodthomsoni sp. nov. is a light brownish to light Kumanjayiwalkerus gen. nov.. has a strongly contrasting green colour dorsally with no flecks or only tiny scattered reddish-brown upper iris, with grey below, versus either blackish flecks. weakly contrasting reddish-brown upper iris or the iris being grey all over in Pengilleyia. By contrast D. pearsoniana has either a few, or many

Duellman et al. (2016) found that the species within each of Pengilleyia and Kumanjayiwalkerus gen. nov. diverged Kumanjayiwalkerus kumanjayi sp. nov. from one another 16.7 MYA and these two genera in turn diverged from their nearest living relatives 23.2 MYA. Distribution: Tropical Australia, extending down the east coast of Queensland as far as the wetter south-east of that State and the Fly River region of Southern Papua New Guinea. Etymology: The genus Kumanjayiwalkerus gen. nov. is named in honour Kumanjayi Walker, another indigenous Australian victim of an execution by a racist white Australian police officer in the Northern Territory, being one of over 400 such victims in Australia since 1990. Kunanurra, Western Australia. The 19 year old was shot and killed by NT Police Officer Zachary Rolfe in November 2019. KUMANJAYIWALKERUS KUMANJAYI SP. NOV. The world is full of memorials and species named in LSIDurn:lsid:zoobank.org:act:9D99A68E-3A87-4AAE- honour mass murders, thieves and despots and It is B0AA-B7B31078D521 appropriate that victims of these people should also be Holotype: A preserved specimen at the Western honoured. Australian Museum, Perth, Western Australia, Australia, Indigenous Australians were subjected to a British specimen number R50585, collected at the Drysdale Genocide that was pro-rata significantly more destructive River National Park, Kimberley District, Western than the holocaust caused by the German Nazi regime to Australia, Australia, Latitude -14.7667 S., Longitude Jews and others. The British attempts to rewrite the 127.0833 E. factual historical record of Australia was far worse than This government-owned facility allows access to its that of the Nazi Propaganda Ministry, including that holdings. Australia was “uninhabited” prior to 1770 when Captain Paratypes: 14 preserved specimens at the Western James Cook “discovered” the place, but these facts are Australian Museum, Perth, Western Australia, Australia, not well known, even to many Australians. specimen numbers, R50377, R50382, R50581, R50582, Also not well known outside Australia is the endemic R50583, R50805, R50806, R131681, R173564, corruption of Australian police forces in the modern era. R173578, R173579, R173580, R164955 and R164956, It is hoped that by naming of a widespread and common collected at the Drysdale River National Park, Kimberley species of frog in honour of a victim of a police muder in District, Western Australia, Australia, Latitude -14.7667 Australia that attention is not only drawn to the victim of S., Longitude 127.0833 E. the crime, but also other victims and a hope that by doing Diagnosis: Kumanjayiwalkerus kumanjayi sp. nov. has so, the crime and corruption that is endemic in Australian until now been treated as merely the western population police forces is eventually stopped (see also etymology of K. rothii (De Vis, 1884), better known as “Litoria rothii”, for Nyctimystes (Asperohyla) georgefloydi sp. nov. earlier with a type locality of Mackay in North Queensland, in this paper). Australia. Notable is that George Floyd was killed by police on 26 However the possibility of there being at least two May 2020 in Minnesota, USA. species within putative “Litoria rothii”, has been known for The videos of the killing made by bystanders were shared decades. Hoser (1989) on page 41 had photos of the online, sparking widespread protests in over 30 cities and western and eastern forms of this species and Anstis worldwide. 2013 on page 297 stated “it is possible there may be Two days later, on 28 May 2020, Victorian Police officers more than one species involved. pulled over a motorist on the busy Monash Freeway at Kumanjayiwalkerus kumanjayi sp. nov. is the form Dandenong North. They got out of their police car and encompassing the Kimberley District of Western proceeded to execute the 53 year old family man by Australia and the top end of the Northern Territory, shooting him dead in broad daylight in full view of other including the western side of the Gulf of Carpentaria. K. drivers, the entire incident being captured on dashcams rothii of the type form occupies most parts of and even State Government-owned freeway CCTV Queensland, except the driest areas and the south, as cameras. Unlike in Minnesota, USA, the Victorian Police well as southern New Guinea. successfully hijacked every video made and none was Both species would key out as “Litoria rothii” in either made public or shared on social media. Anstis (2013) or Cogger (2014). As a result there were no protests in Australia arising from the incident. The two species are readily separated from one another Police Union boss, Wayne Gatt (remember the Gatt as follows: name … and for the wrong reasons), told the media that K. rothii has obvious black spots and blobs above the the execution was wholly justified. arm pit, on the rear flank, groin and inner and rear hind Content: Kumanjayiwalkerus kumanjayi (type species); leg. In contrast these areas of black are either absent, K. rothii (De Vis, 1884). heavily reduced as to be inconspicuous or shaded as grey. Where the black spotting and marks are present in K.

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 46 Australasian Journal of Herpetology rothii, K. kumanjayi sp. nov. has light bluish-purple flash discontinuous tubercles; hind edge of foot is smooth; colours instead. hind side of thighs with contrasting black and yellow bars Some north Queensland specimens of K. rothii may also or marbling, at least dorsally. have light bluish-purple in these inner areas, but in these The genus Pengilleyia Wells and Wellington, 1985 is specimens, the light bluish-purple areas are always well readily separated from the genus Kumanjayiwalkerus bounded by thick black outlines (not seen in K. gen. nov. by having a back that is either very warty or Kumanjayi sp. nov). moderately warty, versus virtually smooth or with well K. rothii has yellow and black mottling along the inside scattered small pointed tubercles on an otherwise edge of the foot, versus light bluish-purple and yellow smooth body in Kumanjayiwalkerus gen. nov.. mottling in K. kumanjayi sp. nov.. Furthermore species within Pengilleyia invariably have Black spotting extends along the lower flank, anterior to green spots, flecks or blotches on the back versus none the hind limb in K. rothii, whereas this is not the case in in Kumanjayiwalkerus gen. nov.. K. kumanjayi sp. nov.. The upper lip of K. rothii is whitish, Kumanjayiwalkerus gen. nov.. has a strongly contrasting versus yellow-brown in K. kumanjayi sp. nov.. reddish-brown upper iris, with grey below, versus either Both species K. kumanjayi sp. nov. and K. rothii are weakly contrasting reddish-brown upper iris or the iris further defined as follows: A medim-sized, somewhat being grey all over in Pengilleyia. slender tree frog. Has a slender hind limb. Tongue oval, Duellman et al. (2016) found that the species within each with its free hind edge rather deeply emarginate. of Pengilleyia and Kumanjayiwalkerus gen. nov. diverged Vomerine teeth in two small groups between the from one another 16.7 MYA and these two genera in turn choanee. Choanae rather large and angular. Head small. diverged from their nearest living relatives 23.2 MYA. Snout subacute, longer than orbit or interorbit. Nostril Distribution: Kumanjayiwalkerus kumanjayi sp. nov. is much nearer the tip of the snout than to the eye. Loreal found from the Kimberley District of Western Australia region shelving is rather concave. Tympanum distinct, and the top end of the Northern Territory, to the western two-thirds of orbit. Fingers half-webbed, but fringed to the side of the Gulf of Carpentaria. K. rothii of the type form discs; discs about two-thirds of tympanum in size. Toes occupies most parts of Queensland, except the driest entirely webbed, with small discs. On protraction of the areas and the south, as well as southern New Guinea. hind foot, the ankle reaches between the eye and the Etymology: The species Kumanjayiwalkerus kumanjayi nostril. There is no distinct tarsal fold. There is a faint fold sp. nov. is named in honour Kumanjayi Walker, another over the wrist. Males to 40 mm, and females to 60 mm in indigenous Australian victim of an execution by a racist body length. Colour variable, but is usually lead grey, white Australian police officer in the Northern Territory olive or reddish brown, uniform or mottled with darker (NT), being one of over 400 such victims in Australia patches. Colour and intensity changes significantly with since 1990. time of day, temperature, activity and other factors. The The 19 year old was shot and killed by NT Police Officer the flash markings of the arm pits, groin and legs are Zachary Rolfe in November 2019. outlined above, but invariably include yellow and one or The world is full of memorials and species named in other of black or bluish-light-purple, or sometimes both. honour mass murderers, thieves and despots and it is Diagnostic of both species is the distinctive iris of the appropriate that victims of these people should also be eye, which has a rediish-orange upper orbit and grey honoured occasionally. lower half. The spelling of the species name is deliberate and Hoser (1989) at page 41 has comparative photos of both should NOT be changed. While it is convention for an “i” K. kumanjayi sp. nov. and K. rothii in life, with K. to be added to a male patronym name, in this case I have kumanjayi sp. nov. on middle right and K. rothii on bottom chosen to dispense with this so that the species name right. Anstis (2013) has comparative photos of both K. more accurately reflects the person it is named in honour kumanjayi sp. nov. and K. rothii in life, with K. kumanjayi of, especially noting that the taxon is common and sp. nov. on page 297 two photos on right and K. rothii on widespread and it is hoped that people remember the page 298 on top left. Vanderduys (2012) has a photo of exact person the frog is named after and not be K. rothii in life, from near the type locality on page 68 otherwise confused into thinking the person’s name in bottom left and also bottom right. Eipper (2012) at page fact ended with two “i’s” noting that in this case, this is 113 bottom also has a photo of typical K. rothii in life, as done for the benefit of lay people and not those familiar does Eipper and Rowland (2018) on page 150 (both with the minor details of scientific nomenclature of living photos), as does Cogger (2014) at page 187 bottom. animals. Clyne (1969) has a photo of K. rothii in life, on page 35 at top. Furthermore this nomenclature decision was made after discussions with people within the Yuendemu Aboriginal The genera Pengilleyia Wells and Wellington, 1985 as Community. defined within this paper, and Kumanjayiwalkerus gen. nov. are as a pair, both readily separated from all other PENGILLEYIA WELLS AND WELLINGTON, 1985 Australasian Tree Frogs (Pelodryadidae) by the following Type species: Litoria tyleri Martin, Watson, Gartside, unique suite of characters: Littlejohn and Loftus-Hills, 1979. Vomerine teeth present; fingers with conspicuous Diagnosis: Pengilleyia Wells and Wellington, 1985 as webbing reaching at least as far as the base of the defined by them was literally a “dogs breakfast” of widely penultimate phalanx of the fourth finger; hind edge of divergent species placed into a single putative genus. forearm is smooth, or with at most a few low, This creation by them, does stand out as one of their

more significant mess-ups and proves that even weakly contrasting reddish-brown upper iris or iris being competant herpetologists with many years experience grey all over in Pengilleyia. can get things really wrong. Distribution: Pengilleyia Wells and Wellington, 1985 Notwithstanding this, the name is available in terms of occurs in Eastern Australia, New Guinea, the Moluccas the rules of the International Code of Zoological and lesser Sundas in Indonesia. Nomenclature (Ride et al. 1999) and all preceding Content: Pengilleyia tyleri (Martin, Watson, Gartside, editions. To that extent, all that matters is the Littlejohn and Loftus-Hills, 1979) (type species); P. phylogenetic placement of the type species and where it amboinensis (Horst, 1883); P. darlingtoni (Loveridge, relates to other putative genera. 1945); P. everetti (Boulenger, 1897); P. peronii (Tschudi, The so-called “Litoria peroni ” (Tschudi, 1838) species 1838); P. obtusirostris (Meyer, 1875). group, includes the species originally described as Litoria AUDAXURINA SUBTRIBE NOV. tyleri Martin, Watson, Gartside, Littlejohn and Loftus-Hills, AUDAXURA GEN. NOV. 1979, and as this species group did not have an available LSIDurn:lsid:zoobank.org:act:8864D28E-5477-46B5- genus name prior to 1985, the Wells and Wellington 842E-9D9F521D6AD0 name Pengilleyia, is that which must be used and Type species: Hyla (Litoria) congenita Peters and Doria, attached to the species group. 1878. If one were to include the “Hyla rothii De Vis, 1884” Diagnosis: The genera Colleeneremia Wells and species group into the genus Pengilleyia as done by Wellington, 1985 and Audaxura gen. nov. are all readily Wells and Wellington (1985), also otherwise then not separated from all other Australasian Tree Frogs assignable to any nearer group, one finds a 23.2 MYA (Pelodryadidae) by the following unique suite of divergence from their nearest living (genus assigned) characters: Grey, brown or fawn above, usually with a relatives according to Duellman et al. (2016). Those broad, darker vertebral band, bounded on either side by relevant species are within the genera Audaxura gen. a lighter brown zone. nov. and Colleeneremia Wells and Wellington, 1985, which each diverged from one another 17 MYA according There is a blackish stripe along the side of the head, to Duellman et al. (2016). continuing behind the eye, over the base of the forelimb and along the side of the body almost to the groin. Dorsal Because the Hyla rothii De Vis, 1884 species group surface of the body and limbs is flecked with dark brown diverged from others within Pengilleyia as outlined or black and sometimes small to medium sized patches above, being the so-called “Litoria peroni ” (Tschudi, of darker pigment. Hind side of thighs brown with fine 1838) species group, approximately 16.7 MYA according white spots. to Duellman et al. (2016), that species group (treated as two species herein) is placed within a newly named Groin is usually lemon-yellow. Ventral surface white, genus Kumanjayiwalkerus gen. nov. which is defined and cream or yellowish. Dorsal surface usually smooth or with diagnosed previously in this paper. numerous tiny granules above and coarsely granular below. Vomerine teeth almost entirely behind the The genera Pengilleyia Wells and Wellington, 1985 and choanae. Pectoral fold prominent. Finger and toe discs Kumanjayiwalkerus gen. nov. are as a pair, both readily large. Toes about two thirds webbed. Inner metatarsal separated from all other Australasian Tree Frogs tubercle moderate and elongated, outer tubercle is small (Pelodryadidae) by the following unique suite of and rounded. Tympanum is large, rounded and distinct. characters: Second finger longer than first. Average adult size 35 mm Vomerine teeth present; fingers with conspicuous in length. webbing reaching at least as far as the base of the Species within the genus Colleeneremia are separated penultimate phalanx of the fourth finger; hind edge of from the morphologically similar species within the genus forearm is smooth, or with at most a few low, Audaxura gen. nov., their closest living relatives as discontinuous tubercles; hind edge of foot is smooth; follows: Colleeneremia species always exhibits a very hind side of thighs with contrasting black and yellow bars broad, dark stripe on the side of the head and body. This or marbling, at least dorsally. is not found in species within Audaxura gen. nov.. The genus Pengilleyia Wells and Wellington, 1985 is Furthermore Audaxura gen. nov. have pale regular or readily separated from the genus Kumanjayiwalkerus irregular stripes or patches on the dorsal surface of the gen. nov. by having a back that is either very warty or body not exhibited in the same configuration or form in moderately warty, versus virtually smooth in any Colleeneremia species. Colleeneremia have shorter Kumanjayiwalkerus gen. nov.. limbs than Audaxura gen. nov.. The TL/S-V ratios The genus Pengilleyia Wells and Wellington, 1985 is provides a means of distinguishing the genera, this being readily separated from the genus Kumanjayiwalkerus 0.335-0.432 for Colleeneremia and 0.477-0.520 for gen. nov. by having a back that is either very warty or Audaxura gen. nov.. moderately warty, versus virtually smooth or with well Within Colleeneremia the two subgenera are separated scattered small pointed tubercles on an otherwise as follows: smooth body in Kumanjayiwalkerus gen. nov.. 1/ Fingers with only rudimentary webbing, being the Furthermore species within Pengilleyia invariably have subgenus Colleeneremia, or: green spots, flecks or blotches on the back versus none in Kumanjayiwalkerus gen. nov.. 2/ With conspicuous webbing on the fingers, reaching at least as far as the base of the penultimate phalanx of the Kumanjayiwalkerus gen. nov. has a strongly contrasting fourth finger being the subgenus Balatusrana subgen. reddish-brown upper iris, with grey below, versus either nov.. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 48 Australasian Journal of Herpetology

Brevicrusyla gen. nov. includes two species from the prominent. Finger and toe discs large. Toes about two Indonesian side of New Guinea that are morphologically thirds webbed. Inner metatarsal tubercle moderate and similar to species within the genera Colleeneremia Wells elongated, outer tubercle is small and rounded. and Wellington, 1985 and Audaxura gen. nov. as detailed Tympanum is large, rounded and distinct. Second finger above, but are readily separated from them in that adults longer than first. Average adult size 35 mm in length. of Brevicrusyla gen. nov. while having a smooth dorsum Species within the genus Colleeneremia are separated on the body like the other two genera, instead has from the morphologically similar species within the genus tubercles on the head, that is not seen in the other two Audaxura gen. nov., their closest living relatives as genera. Tadpoles of Brevicrusyla gen. nov. have a long follows: Colleeneremia species always exhibits a very muscular tail, with narrow dorsal and ventral fins, versus broad, dark stripe on the side of the head and body. This a relatively short tail, with broad fins in species within is not found in species within Audaxura gen. nov.. Audaxura gen. nov. and Colleeneremia. Furthermore Audaxura gen. nov. have pale regular or Duellman et al. (2016) found that the genera irregular stripes or patches on the dorsal surface of the Colleeneremia and Audaxura gen. nov. diverged from body not exhibited in the same configuration or form in one another 17 MYA, with a similar divergence indicated any Colleeneremia species. Colleeneremia have shorter by Pyron and Weins (2011). limbs than Audaxura gen. nov.. The TL/S-V ratios Distribution: Audaxura gen. nov. is found in New provides a means of distinguishing the genera, this being Guinea, immediately offshore Islands to the south and 0.335-0.432 for Colleeneremia and 0.477-0.520 for south-west and north-east as well as the nearby Audaxura gen. nov.. Moluccan Islands. Within Colleeneremia the two subgenera are separated Etymology: Audax in Latin means bold, and by frog as follows: standards this is an invasive genus, with specimens 1/ Fingers with only rudimentary webbing, being the commonly entering sites of human habitation in lowland subgenus Colleeneremia, or: areas. “ura” is an abbreviation of “anura” = frog and 2/ With conspicuous webbing on the fingers, reaching at hence the name Audaxura gen. nov.. least as far as the base of the penultimate phalanx of the Content: Audaxura congenita (Peters and Doria, 1878) fourth finger being the subgenus Balatusrana subgen. (type species); A. capitula (Tyler, 1968); A. pygmaea nov.. (Meyer, 1875); A. quadrilineata (Tyler and Parker, 1974). Tyler and Davis (1978) originally placed their species BREVICRUSYLA GEN. NOV. “Hyla wisselensis” in a group with “Hyla rubella” now of LSIDurn:lsid:zoobank.org:act:D6264CA2-F412-46DC- the genus Colleeneremia . Tyler and Davis (1983) placed A481-2FC0CE7962F8 both “Hyla wisselensis” and “Litoria umbonata” in a new Type species: Hyla wisselensis Tyler, 1968. group of species including just these two. Diagnosis: Brevicrusyla gen. nov. includes two species Significant morphological divergence at both larval and from the Indonesian side of New Guinea that are adult stages of the life cycle support this contention and morphologically similar to species within the genera the creation of the genus Brevicrusyla gen. nov. Colleeneremia Wells and Wellington, 1985 and Audaxura Distribution: Known only from the Wissel Lakes area for gen. nov., but readily separated from them in that adults B. wisselensis (Tyler, 1968) and the Baliem River Valley of Brevicrusyla gen. nov. while having a smooth dorsum of West Papua, Indonesia B. umbonata (Tyler and Davies on the body like the other two genera, instead has 1983). tubercles on the head, that is not seen in the other two Etymology: The genus name is a slightly abbreviated genera. form of Latin for short (brevis), leg (Crus) and tree frog as Tadpoles of Brevicrusyla gen. nov. have a long muscular in “Hyla”, combined to form the word “Brevicrusyla”. This tail, with narrow dorsal and ventral fins, versus a is a reflection of the relatively short hind limbs of the relatively short tail, with broad fins in species within species. Audaxura gen. nov. and Colleeneremia. Content: Brevicrusyla wisselensis (Tyler, 1968) (type The genera Colleeneremia Wells and Wellington, 1985, species); B. umbonata (Tyler and Davies 1983). Brevicrusyla gen. nov. and Audaxura gen. nov. are all GENUS COLLEENEREMIA WELLS AND readily separated from all other Australasian Tree Frogs WELLINGTON, 1985 (Pelodryadidae) by the following unique suite of Type species: Hyla rubella Gray, 1842. characters: Grey, brown or fawn above, usually with a Diagnosis: The genus Colleeneremia Wells and broad, darker vertebral band, bounded on either side by Wellington, 1985 as defined herein is different to the a lighter brown zone. There is a blackish stripe along the original diagnosis and definition of Wells and Wellington, side of the head, continuing behind the eye, over the 1985. base of the forelimb and along the side of the body Wells and Wellington (1985) treated the genus as almost to the groin. Dorsal surface of the body and limbs monotypic for the taxon Hyla rubella, Gray, 1842, which is flecked with dark brown or black and sometimes small they correctly stated was “Believed to be a complex of to medium sized patches of darker pigment. Hind side of several undescribed species.” To the extent that the thighs brown with fine white spots. Groin is usually taxon “Litoria electrica” Ingram and Corben, 1990 was lemon-yellow. Ventral surface white, cream or yellowish. defined by Ingram and Corben (1990) as a species Dorsal surface usually smooth or with numerous tiny previously treated as putative C. rubella (Gray, 1842), granules above and coarsely granular below. Vomerine Wells and Wellington (1985) have been vindicated well teeth almost entirely behind the choanae. Pectoral fold

before the publication of this paper. of a new genus for the L. congenita group as the only Confirming the position of Wells and Wellington (1985) logical situation for those species. That genus is called was Anstis (2013) who at page 301 also stated she Audaxura gen. nov.. A similar genus-level division is thought C. rubella was a species complex. done for the genus Pengilleyia Wells and Wellington, Eipper and Rowland (2018) at page 151 stated, 1985 within this paper. “Probably a species complex”. In accordance with the preceding, the genus Even the ultra-conservative Harold Cogger in Cogger Colleeneremia Wells and Wellington, 1985 are all readily (2014) stated of C. rubella, separated from all other Australasian Tree Frogs “Almost certainly composite”. (Pelodryadidae) by the following unique suite of Mention of the three above authorities is so that I am not characters: Grey, brown or fawn above, usually with a to be accused of merely accepting Wells and Wellington broad, darker vertebral band, bounded on either side by taxonomy and nomenclature on the basis of alleged a lighter brown zone. There is a blackish stripe along the friendship of the pair as repeatedly stated by the side of the head, continuing behind the eye, over the Wolfgang Wüster gang. base of the forelimb and along the side of the body Instead the taxonomy herein and nomenclature that almost to the groin. Dorsal surface of the body and limbs follows is solely as result of body of scientific evidence is flecked with dark brown or black and sometimes small that has also passed peer review (notably unlike Kaiser to medium sized patches of darker pigment. Hind side of et al. 2013). thighs is brown with fine white spots. Groin is usually lemon-yellow. Ventral surface white, cream or yellowish. In terms of the widespread and variable putative taxon Dorsal surface usually smooth or with numerous tiny known as C. rubella it is herein split into six well defined granules above and coarsely granular below. Vomerine species all separated by well known biogeograpcial teeth almost entirely behind the choanae. Pectoral fold barriers of known antiquity. All populations are prominent. Finger and toe discs large. Toes about two morphologically distinct and also appear to be allopatric. thirds webbed. Inner metatarsal tubercle moderate and Hence to this extent the original diagnosis of Wells and elongated, outer tubercle is small and rounded. Wellington (1985) is confirmed. Tympanum is large, rounded and distinct. Second finger In terms of divergence, Duellman et al. (2016) found that longer than first. Average adult size 35 mm in length. the genus Colleeneremia as defined by Wells and As well as one or other of the following characters: Wellington, had a 14 million year divergence from its 1/ Fingers with only rudimentary webbing, being the nearest relative outside, being Hyla dentata Keferstein, subgenus Colleeneremia, or: 1868, which they placed in their newly erected genus Rawlinsonia Wells and Wellington, 1985, which had a 2/ With conspicuous webbing on the fingers, reaching at type species of Hyla ewingi Duméril and Bibron, 1841, least as far as the base of the penultimate phalanx of the which in fact is a different phylogentic grouping. fourth finger being the subgenus Balatusrana subgen. Transfer of Hyla dentata Keferstein, 1868 to nov.. Colleeneremia retains the monophyly of Rawlinsonia Species within the genus Colleeneremia are separated (which according to Duellman et al. (2016) has a from the morphologically similar species within the genus divergence of 24.7 MYA from nearest living relatives, Audaxura gen. nov., their closest living relatives as confirming the good sense in erecting that genus). follows: Colleeneremia species always exhibits a very The preceding transfer gives the genus Colleeneremia a broad, dark stripe on the side of the head and body. This divergence of 17 MYA from its nearest living relative, is not found in species within Audaxura gen. nov.. according to Duellman et al. (2016). Furthermore Audaxura gen. nov. have pale regular or irregular stripes or patches on the dorsal surface of the That nearest living relative species, is Hyla (Litoria) body not exhibited in the same configuration or form in congenita Peters and Doria, 1878 of southern New any Colleeneremia species. Colleeneremia have shorter Guinea and the associated group of species. limbs than Audaxura gen. nov.. The TL/S-V ratios Current taxonomy preceding this paper, has all species in provides a means of distinguishing the genera, this being the L. congenita group placed in the genus Litoria 0.335-0.432 for Colleeneremia and 0.477-0.520 for Tschudi, 1838. Audaxura gen. nov.. This is clearly untenable, because according to Duellman Brevicrusyla gen. nov. includes two species from the et al. (2016) the type group for that genus and all relevant Indonesian side of New Guinea that are morphologically members of the species group have a 31.5 MYA similar to species within the genera Colleeneremia Wells divergence from Colleeneremia. and Wellington, 1985 and Audaxura gen. nov. as detailed The only solution to the problem of placement of L. above, but are readily separated from them in that adults congenita group of species is therefore one or other of of Brevicrusyla gen. nov. while having a smooth dorsum wholly subsuming them within Colleeneremia, erecting a on the body like the other two genera, instead has subgenus for the L. congenita group, or alternatively tubercles on the head, that is not seen in the other two erecting a new genus for the group. genera. Tadpoles of Brevicrusyla gen. nov. have a long Based on the divergence of 17 MYA between the species muscular tail, with narrow dorsal and ventral fins, versus groups and a view that the species diversity of both a relatively short tail, with broad fins in species within species groups are in fact underestimated in terms of Audaxura gen. nov. and Colleeneremia. species diversity (even after the publication of formal Distribution: Most of continental Australia except for the descriptions of new species in this paper), I see erection far south, as well as southern New Guinea.

Content: Colleeneremia rubella (Gray, 1842) (type genera. Tadpoles of Brevicrusyla gen. nov. have a long species); C. bogfrog sp. nov.; C. chunda sp. nov.; C. muscular tail, with narrow dorsal and ventral fins, versus dentata (Keferstein, 1868); C. dunnyseat sp. nov.; C. a relatively short tail, with broad fins in species within electrica (Ingram and Corben, 1990); C. watdat sp. nov.; Audaxura gen. nov. and Colleeneremia. C. wifi sp. nov.; Distribution: East coast of New South Wales, including BALATUSRANA SUBGEN. NOV. wetter parts of nearby south-east Queensland and also LSIDurn:lsid:zoobank.org:act:24C7D6E6-2F46-417E- far north-east Victoria. 8CD2-8CAE4B59463E Content: Colleeneremia (Balatusrana) dentata Type species: Hyla dentata Keferstein, 1868. (Keferstein, 1868) (treated herein as monotypic, but Diagnosis: The subgenus Balatusrana subgen. nov. is including at least one subspecies). separated from the nominate subgenus Colleeneremia COLLEENEREMIA BOGFROG SP. NOV. Wells and Wellington, 1985 by having conspicuous LSIDurn:lsid:zoobank.org:act:BE7164BB-D292-48A8- webbing on the fingers, reaching at least as far as the BBE6-5711983E6192 base of the penultimate phalanx of the fourth finger. Holotype: A preserved specimen at the Northern In the nominate subgenus Colleeneremia the fingers Territory Art Gallery and Museum, Darwin, Northern have only rudimentary webbing. Territory, Australia, specimen number R30936, collected In addition to the character states just mentioned for each at Jay Creek, central Australia, Northern Territory, subgenus, both subgenera, comprising the totality of the Australia, Latitude -23.8 S., Longitude 133.05 E. genus Colleeneremia are separated from all other This government-owned facility allows access to its Australasian Tree Frogs (Pelodryadidae) by the following holdings. unique suite of characters: Paratypes: Five preserved specimens at the Northern Grey, brown or fawn above, usually with a broad, darker Territory Art Gallery and Museum, Darwin, Northern vertebral band, bounded on either side by a lighter brown Territory, Australia, specimen numbers R30934, R30935, zone. There is a blackish stripe along the side of the R30827, R30828 and R15422, all collected from the type head, continuing behind the eye, over the base of the locality (above). forelimb and along the side of the body almost to the Diagnosis: The five species, Colleeneremia watdat sp. groin. Dorsal surface of the body and limbs is flecked nov. from the Pilbara, Gascoyne and Murchison regions with dark brown or black and sometimes small to medium of western Western Australia, C. bogfrog sp. nov. from sized patches of darker pigment. Hind side of thighs the Macdonnell Ranges of Central Australia, C. brown with fine white spots. Groin is usually lemon- dunnyseat sp. nov. from the Flinders Ranges of South yellow. Ventral surface white, cream or yellowish. Dorsal Australia, C. wifi from western New South Wales and surface usually smooth or with numerous tiny granules southern Queensland and C. chunda from the Cape York above and coarsely granular below. Vomerine teeth region in far north Queensland and probably southern almost entirely behind the choanae. Pectoral fold New Guinea have until now all been treated as prominent. Finger and toe discs large. Toes about two populations of putative C. rubella (Keferstein, 1868), thirds webbed. Inner metatarsal tubercle moderate and herein confined to the dry tropics of the Northern Territory elongated, outer tubercle is small and rounded. and the Kimberley district of Western Australia. Tympanum is large, rounded and distinct. Second finger Another morphologically similar species C. electrica longer than first. Average adult size 35 mm in length. (Ingram and Corben, 1990), was prior to 1990 also Species within the genus Colleeneremia are separated treated as a regional population of C. rubella. from the morphologically similar species within the genus The seven species comprise the total of the subgenus Audaxura gen. nov., their closest living relatives (with a Colleeneremia Wells and Wellington, 1985 as defined in divergence of 17 MYA according to Duellman et al. 2016) this paper. as follows: Colleeneremia species always exhibits a very While these species within Colleeneremia have diverse broad, dark stripe on the side of the head and body. This habitat choices, their distribution does coincide with is not found in species within Audaxura gen. nov.. biogeographical barriers generally affecting saxicoline Furthermore Audaxura gen. nov. have pale regular or genera of Australian fauna. irregular stripes or patches on the dorsal surface of the This is including for example species of Varanid lizards body not exhibited in the same configuration or form in (Hoser 2013, 2014, 2015g, 2018b-e), Ring-tailed any Colleeneremia species. Colleeneremia have shorter Dragons in the genus Ctenophorus Fitzinger, 1843 limbs than Audaxura gen. nov.. The TL/S-V ratios (Hoser 2020c) other lizards within the genus Egernia provides a means of distinguishing the genera, this being Gray, 1838 sensu lato (Hoser 2018a) and also Petrogale 0.335-0.432 for Colleeneremia and 0.477-0.520 for Gray, 1837 wallabies (Hoser, 2020b). Audaxura gen. nov.. The seven aforementioned species are separated from Brevicrusyla gen. nov. includes two species from the one another as follows: Indonesian side of New Guinea that are morphologically The type form of C. rubella and C. chunda sp. nov. similar to species within the genera Colleeneremia Wells conforms to the genus description for Colleeneremia. and Wellington, 1985 and Audaxura gen. nov. as detailed above, but are readily separated from them in that adults They can be further diagnosed as follows: Compared to of Brevicrusyla gen. nov. while having a smooth dorsum all other species in the genus, these two species have on the body like the other two genera, instead has relatively undeveloped lateral digital fringes and a tubercles on the head, that is not seen in the other two relatively narrow head (medium in cross-section), versus

broad (broad in cross-section) and well-developed lateral high density of spotting and it being relatively evenly digital fringes in all other species. C. chunda sp. nov. is spaced on the contrasting lighter background, so the readily separated from C. rubella by the obvious bright dorsum appears spotted, as opposed to one with yellow feet, well demarcated from the grey of the digits, blotches as may seen in some of the other species. versus yellowish feet and digits without a well defined Yellow in armpits and groin is often absent or when boundary in C. rubella. present not as brilliant, or prominent as in other species. C. chunda sp. nov. is characterised by significant C. wifi sp. nov. is readily separated from the other six speckling on the upper dorsum and a well-defined white species by being chocolate brown on the dorsum and upper labial area below the eye, versus little specking on gun metal grey on the flanks. The dorsum is irregularly, the dorsum of C. rubella and ill-defined lighter areas on but heavily peppered, with the same gun-metal grey the the upper labial region. result being these patches of peppering forming semi- C. chunda sp. nov. has well defined areas of yellow in the distinct areas of grey surrounded by otherwise mainly groin, versus ill-defined and spreading up the lower flank brown pigment. The upper surfaces of the limbs are the in C. rubella. same chocolate brown colour as the body, but with even Both C. chunda sp. nov. and C. rubella have either heavier grey peppering. The labial area is also generally smooth, or very minutely granular skin on the upper grey and the top of the snout is brown, with either no surface. peppering or minimal peppering that tends to be faded. C. chunda sp. nov. is further separated from all other Toes may be brown or grey (in a single specimen) species by having a poorly formed and semi-distinct mid- depending on which are peppered heavily and which are vertebral line. not. The upper iris is a dull orange brown in this species, versus brilliant red or orange in all the other six. C. chunda sp. nov. and C. rubella further differ from the other species by having a distinctively bitonal upper C. rubella in life can be seen in Anstis (2013) on page surface, with a distinct broad grey or red-brown band 301 in the two right hand photos. down the middle of the back on a beige or fawn C. chunda sp. nov. in life can be seen in Anstis (2013) on background. page 302, on right (bottom image) and in Vanderduys C. electrica (Ingram and Corben, 1990) is separated from (2012) on page 69 bottom. the other six species by having three distinctive dark bars C. electrica in life can be seen in Anstis (2013) on page or blotches on the back, a dorsum that is obviously 192, two right hand photos, and Cogger (2014) on page tubercular and the yellow patches in the groin and arm 162 (two photos). pits are bold and distinctive and made more so by being C. bogfrog sp. nov. in life can be seen in Anstis (2013) on edged with dark brownish-grey bordering, not seen in any page 302 on left at top. other species. C. watdat sp. nov. in life can be seen online at: C. bogfrog sp. nov. is separated from the other six https://www.flickr.com/photos/27026445@N06/ species by having a pinkish red colour on the dorsum 31419696523/ and flanks with pinkish white and very light grey marbling. and: The dorsum has smooth or leathery skin, but without https://www.flickr.com/photos/warpedtime/2245649035/ obvious raised tubercles. and: Speckling is minimal and where present, usually on the https://www.flickr.com/photos/27026445@N06/ snout and rear, it is still scattered, minimal and very faded 32155803631/ as to be indistinct. There are also irregular purple-grey C. dunnyseat sp. nov. in life can be seen in Anstis (2013) patches running down two longitudinal lines on the upper on page 302 at top right image. dorsum, not near the mid-line and these too are faded. C. wifi sp. nov. in life can be seen in Hoser (1989) on C. watdat sp. nov. is similar in dorsal patterning to C. page 42 in the top photo and Cogger (2014) on page bogfrog sp. nov. except that there is scattered flecking 188, top left photo. across all the body, as opposed to more being at front of The subgenus Balatusrana subgen. nov. is separated snout and rear. C. watdat sp. nov. is also instead a mud- from the nominate subgenus Colleeneremia Wells and brownish coloured frog (not pinkish-red) and also has a Wellington, 1985 (being all seven species separated by strongly granulated dorsum and to a greater degree than the descriptions herein) by having conspicuous webbing any other species in the complex. Yellow in the groin and on the fingers, reaching at least as far as the base of the armpits is either non-existent or minimal. penultimate phalanx of the fourth finger. C. watdat sp. nov. is further characterised by having a In the nominate subgenus Colleeneremia the fingers well defined dark brown stripe running from snout, have only rudimentary webbing. through eye and ear over back leg to flank. The line is In addition to the character states just mentioned for each well defined, contrasting with the lighter brown surfaces subgenus, both subgenera, comprising the totality of the adjacent and clearly runs across the entire tympanum. genus Colleeneremia are separated from all other C. dunnyseat sp. nov. is readily separated from all other Australasian Tree Frogs (Pelodryadidae) by the following six species by having a light greyish brown, almost unique suite of characters: Grey, brown or fawn above, tending to white coloured dorsum with distinctive dark usually with a broad, darker vertebral band, bounded on brownish-black spots and mottling spread relatively either side by a lighter brown zone. There is a blackish evenly across the dorsum. Limbs are also creamish in stripe along the side of the head, continuing behind the colour with obvious darker flecks and mottling. This taxon eye, over the base of the forelimb and along the side of comes across as a uniquely speckled frog, due to the Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 52 Australasian Journal of Herpetology the body almost to the groin. Dorsal surface of the body COLLEENEREMIA CHUNDA SP. NOV. and limbs is flecked with dark brown or black and LSIDurn:lsid:zoobank.org:act:AF3507CE-3294-4777- sometimes small to medium sized patches of darker AA7A-6B38ACDD474E pigment. Hind side of thighs brown with fine white spots. Holotype: A preserved specimen at the Queensland Groin is usually lemon-yellow. Ventral surface white, Museum, Brisbane, Queensland, Australia, specimen cream or yellowish. Dorsal surface usually smooth or with number J16011 collected at Tolga, Atherton Tablelands, numerous tiny granules above and coarsely granular Queensland, Australia, Latitude -17.2167S., Longitude below. Vomerine teeth almost entirely behind the 145.4833 E. choanae. Pectoral fold prominent. This government-owned facility allows access to its Finger and toe discs large. Toes about two thirds holdings. webbed. Inner metatarsal tubercle moderate and Paratypes: Five preserved specimens at the Queensland elongated, outer tubercle is small and rounded. Museum, Brisbane, Queensland, Australia, specimen Tympanum is large, rounded and distinct. Second finger numbers J79439, J79472, J81038, J55732 and J55723 longer than first. Average adult size 35 mm in length. collected from near Tolga, Atherton Tablelands, Species within the genus Colleeneremia are separated Queensland, Australia, Latitude -17.2167 S., Longitude from the morphologically similar species within the genus 145.4833 E. Audaxura gen. nov., their closest living relatives (with a Diagnosis: The five species, Colleeneremia watdat sp. divergence of 17 MYA according to Duellman et al. 2016) nov. from the Pilbara, Gascoyne and Murchison regions as follows: Colleeneremia species always exhibits a very of western Western Australia, C. bogfrog sp. nov. from broad, dark stripe on the side of the head and body. This the Macdonnell Ranges of Central Australia, C. is not found in species within Audaxura gen. nov.. dunnyseat sp. nov. from the Flinders Ranges of South Furthermore Audaxura gen. nov. have pale regular or Australia, C. wifi from western New South Wales and irregular stripes or patches on the dorsal surface of the southern Queensland and C. chunda from the Cape York body not exhibited in the same configuration or form in region in far north Queensland have until now all been any Colleeneremia species. Colleeneremia have shorter treated as populations of putative C. rubella (Keferstein, limbs than Audaxura gen. nov.. The TL/S-V ratios 1868), herein confined to the dry tropics of the Northern provides a means of distinguishing the genera, this being Territory and the Kimberley district of Western Australia. 0.335-0.432 for Colleeneremia and 0.477-0.520 for Another morphologically similar species C. electrica Audaxura gen. nov.. (Ingram and Corben, 1990), was prior to 1990 also Brevicrusyla gen. nov. includes two species from the treated as a regional population of C. rubella. Indonesian side of New Guinea that are morphologically The seven species comprise the total of the subgenus similar to species within the genera Colleeneremia Wells Colleeneremia Wells and Wellington, 1985 as defined in and Wellington, 1985 and Audaxura gen. nov. as detailed this paper. above, but are readily separated from them in that adults While these species within Colleeneremia have diverse of Brevicrusyla gen. nov. while having a smooth dorsum habitat choices, their distribution does coincide with on the body like the other two genera, instead has biogeographical barriers generally affecting saxicoline tubercles on the head, that is not seen in the other two genera of Australian fauna. genera. This is including for example species of Varanid lizards Tadpoles of Brevicrusyla gen. nov. have a long muscular (Hoser 2013, 2014, 2015g, 2018b-e), Ring-tailed tail, with narrow dorsal and ventral fins, versus a Dragons in the genus Ctenophorus Fitzinger, 1843 relatively short tail, with broad fins in species within (Hoser 2020c) other lizards within the genus Egernia Audaxura gen. nov. and Colleeneremia. Gray, 1838 sensu lato (Hoser 2018a) and also Petrogale Distribution: Colleeneremia bogfrog sp. nov. occurs in Gray, 1837 wallabies (Hoser, 2020b). the Macdonnell Ranges of Central Australia, Northern The seven aforementioned species are separated from Territory, Australia. one another as follows: Etymology: In 1978 I was staying with at a camp of The type form of C. rubella and C. chunda sp. nov. Arrernte people near Alice Springs, Northern Territory, conforms to the genus description for Colleeneremia. which is where these aboriginal people live. They can be further diagnosed as follows: Compared to I asked them what they called this taxon of frog and they all other species in the genus, these two species have told me they were the “bog frog”, because whenever they relatively undeveloped lateral digital fringes and a got to do a bog, they see these frogs. relatively narrow head (medium in cross-section), versus To confirm the point, that evening, I was led to an outside broad (broad in cross-section) and well-developed lateral toilet block and in the toilet cubicle were about six of digital fringes in all other species. C. chunda sp. nov. is these frogs on the floor and toilet seat. readily separated from C. rubella by the obvious bright Hence the somewhat unusual scientific name for this yellow feet, well demarcated from the grey of the digits, species. versus yellowish feet and digits without a well defined The name will also serve to draw attention to an boundary in C. rubella. otherwise locally abundant and potentially ignored C. chunda sp. nov. is characterised by significant component of Australia’s native wildlife. speckling on the upper dorsum and a well-defined white upper labial area below the eye, versus little specking on the dorsum of C. rubella and ill-defined lighter areas on the upper labial region. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 53

C. chunda sp. nov. has well defined areas of yellow in the distinct areas of grey surrounded by otherwise mainly groin, versus ill-defined and spreading up the lower flank brown pigment. The upper surfaces of the limbs are the in C. rubella. same chocolate brown colour as the body, but with even Both C. chunda sp. nov. and C. rubella have either heavier grey peppering. The labial area is also generally smooth, or very minutely granular skin on the upper grey and the top of the snout is brown, with either no surface. peppering or minimal peppering that tends to be faded. C. chunda sp. nov. is further separated from all other Toes may be brown or grey (in a single specimen) species by having a poorly formed and semi-distinct mid- depending on which are peppered heavily and which are vertebral line. not. The upper iris is a dull orange brown in this species, C. chunda sp. nov. and C. rubella further differ from the versus brilliant red or orange in all the other six. other species by having a distinctively bitonal upper C. rubella in life can be seen in Anstis (2013) on page surface, with a distinct broad grey or red-brown band 301 in the two right hand photos. down the middle of the back on a beige or fawn C. chunda sp. nov. in life can be seen in Anstis (2013) on background. page 302, on right (bottom image) and in Vanderduys C. electrica (Ingram and Corben, 1990) is separated from (2012) on page 69 bottom. the other six species by having three distinctive dark bars C. electrica in life can be seen in Anstis (2013) on page or blotches on the back, a dorsum that is obviously 192, two right hand photos, and Cogger (2014) on page tubercular and the yellow patches in the groin and arm 162 (two photos). pits are bold and distinctive and made more so by being C. bogfrog sp. nov. in life can be seen in Anstis (2013) on edged with dark brownish-grey bordering, not seen in any page 302 on left at top. other species. C. watdat sp. nov. in life can be seen online at: C. bogfrog sp. nov. is separated from the other six https://www.flickr.com/photos/27026445@N06/ species by having a pinkish red colour on the dorsum 31419696523/ and flanks with pinkish white and very light grey marbling. and: The dorsum has smooth or leathery skin, but without https://www.flickr.com/photos/warpedtime/2245649035/ obvious raised tubercles. Speckling is minimal and where and: present, usually on the snout and rear, it is still scattered, https://www.flickr.com/photos/27026445@N06/ minimal and very faded as to be indistinct. There are 32155803631/ also irregular purple-grey patches running down two C. dunnyseat sp. nov. in life can be seen in Anstis (2013) longitudinal lines on the upper dorsum, not near the mid- on page 302 at top right image. line and these too are faded. C. wifi sp. nov. in life can be seen in Hoser (1989) on C. watdat sp. nov. is similar in dorsal patterning to C. page 42 in the top photo and Cogger (2014) on page bogfrog sp. nov. except that there is scattered flecking 188, top left photo. across all the body, as opposed to more being at front of snout and rear. C. watdat sp. nov. is also instead a mud- The subgenus Balatusrana subgen. nov. is separated brownish coloured frog (not pinkish-red) and also has a from the nominate subgenus Colleeneremia Wells and strongly granulated dorsum and to a greater degree than Wellington, 1985 (being all seven species separated by any other species in the complex. Yellow in the groin and the descriptions herein) by having conspicuous webbing armpits is either non-existent or minimal. on the fingers, reaching at least as far as the base of the penultimate phalanx of the fourth finger. C. watdat sp. nov. is further characterised by having a well defined dark brown stripe running from snout, In the nominate subgenus Colleeneremia the fingers through eye and ear over back leg to flank. The line is have only rudimentary webbing. well defined, contrasting with the lighter brown surfaces In addition to the character states just mentioned for each adjacent and clearly runs across the entire tympanum. subgenus, both subgenera, comprising the totality of the C. dunnyseat sp. nov. is readily separated from all other genus Colleeneremia are separated from all other six species by having a light greyish brown, almost Australasian Tree Frogs (Pelodryadidae) by the following tending to white coloured dorsum with distinctive dark unique suite of characters: Grey, brown or fawn above, brownish-black spots and mottling spread relatively usually with a broad, darker vertebral band, bounded on evenly across the dorsum. Limbs are also creamish in either side by a lighter brown zone. There is a blackish colour with obvious darker flecks and mottling. This taxon stripe along the side of the head, continuing behind the comes across as a uniquely speckled frog, due to the eye, over the base of the forelimb and along the side of high density of spotting and it being relatively evenly the body almost to the groin. Dorsal surface of the body spaced on the contrasting lighter background, so the and limbs is flecked with dark brown or black and dorsum appears spotted, as opposed to one with sometimes small to medium sized patches of darker blotches as may seen in some of the other species. pigment. Hind side of thighs brown with fine white spots. Yellow in armpits and groin is often absent or when Groin is usually lemon-yellow. Ventral surface white, present not as brilliant, or prominent as in other species. cream or yellowish. Dorsal surface usually smooth or with numerous tiny granules above and coarsely granular C. wifi sp. nov. is readily separated from the other six below. Vomerine teeth almost entirely behind the species by being chocolate brown on the dorsum and choanae. Pectoral fold prominent. Finger and toe discs gun metal grey on the flanks. The dorsum is irregularly, large. Toes about two thirds webbed. Inner metatarsal but heavily peppered, with the same gun-metal grey the tubercle moderate and elongated, outer tubercle is small result being these patches of peppering forming semi-

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 54 Australasian Journal of Herpetology and rounded. Tympanum is large, rounded and distinct. COLLEENEREMIA DUNNYSEAT SP. NOV. Second finger longer than first. Average adult size 35 mm LSIDurn:lsid:zoobank.org:act:CEFE393B-3B11-4669- in length. AC4E-8BFB0EB58650 Species within the genus Colleeneremia are separated Holotype: A preserved specimen at the South Australian from the morphologically similar species within the genus Museum, Adelaide, South Australia, Australia, specimen Audaxura gen. nov., their closest living relatives (with a number R7364, collected at Leigh Creek, South divergence of 17 MYA according to Duellman et al. 2016) Australia, Australia, Latitude -30.48 S., Longitude 138.42 as follows: Colleeneremia species always exhibits a very E. broad, dark stripe on the side of the head and body. This This government-owned facility allows access to its is not found in species within Audaxura gen. nov.. holdings. Furthermore Audaxura gen. nov. have pale regular or Paratypes: 39 preserved specimens at the South irregular stripes or patches on the dorsal surface of the Australian Museum, Adelaide, South Australia, Australia, body not exhibited in the same configuration or form in specimen numbers R7365- R7394, R7396- R7398 and any Colleeneremia species. Colleeneremia have shorter R7400- R7405, also all collected at Leigh Creek, South limbs than Audaxura gen. nov.. The TL/S-V ratios Australia, Australia, Latitude -30.48 S., Longitude 138.42 provides a means of distinguishing the genera, this being E. 0.335-0.432 for Colleeneremia and 0.477-0.520 for Diagnosis: The five species, Colleeneremia watdat sp. Audaxura gen. nov.. nov. from the Pilbara, Gascoyne and Murchison regions Brevicrusyla gen. nov. includes two species from the of western Western Australia, C. bogfrog sp. nov. from Indonesian side of New Guinea that are morphologically the Macdonnell Ranges of Central Australia, C. similar to species within the genera Colleeneremia Wells dunnyseat sp. nov. from the Flinders Ranges of South and Wellington, 1985 and Audaxura gen. nov. as detailed Australia, C. wifi from western New South Wales and above, but are readily separated from them in that adults southern Queensland and C. chunda from the Cape York of Brevicrusyla gen. nov. while having a smooth dorsum region in far north Queensland have until now all been on the body like the other two genera, instead has treated as populations of putative C. rubella (Keferstein, tubercles on the head, that is not seen in the other two 1868), herein confined to the dry tropics of the Northern genera. Tadpoles of Brevicrusyla gen. nov. have a long Territory and the Kimberley district of Western Australia. muscular tail, with narrow dorsal and ventral fins, versus Another morphologically similar species C. electrica a relatively short tail, with broad fins in species within (Ingram and Corben, 1990), was prior to 1990 also Audaxura gen. nov. and Colleeneremia. treated as a regional population of C. rubella. Distribution: C. chunda sp. nov. is generally found on The seven species comprise the total of the subgenus Cape York in far north Queensland including nearby Colleeneremia Wells and Wellington, 1985 as defined in areas to there, including the wet tropics of far north this paper. Queensland, being the coastal region north of Mount While these species within Colleeneremia have diverse Spec (near Townsville, Queensland). habitat choices, their distribution does coincide with Etymology: Chunda is Australian slang for “Watch biogeographical barriers generally affecting saxicoline Under” and this is the word commonly used by residents genera of Australian fauna. in North Queensland when they encounter this species of This is including for example species of Varanid lizards frog on floors of toilet blocks at camping grounds, (Hoser 2013, 2014, 2015g, 2018b-e), Ring-tailed aboriginal missions, “illegal alien” detention centres and Dragons in the genus Ctenophorus Fitzinger, 1843 other Australian Federal Government concentration (Hoser 2020c) other lizards within the genus Egernia camps and prisons. The Chunda cry is heard by adults Gray, 1838 sensu lato (Hoser 2018a) and also Petrogale and kids as they scream to others not to tread on the Gray, 1837 wallabies (Hoser, 2020b). small frogs as they walk into a toilet block and can be The seven aforementioned species are separated from heard across North Queensland camping grounds on a one another as follows: nightly basis. Scared Australian females also often yell out “Chunda Fuck!”, but it was determined to remove the The type form of C. rubella and C. chunda sp. nov. word “Fuck” from the etymology and name, as it may conforms to the genus description for Colleeneremia. cause offence and would therefore be contrary to the They can be further diagnosed as follows: Compared to voluntary recommendations of the International Code of all other species in the genus, these two species have Zoological Nomenclature (Ride et al. 1999). relatively undeveloped lateral digital fringes and a Chunda is also a word used by British convicts who were relatively narrow head (medium in cross-section), versus forced onto boats sent to Australia to help in the broad (broad in cross-section) and well-developed lateral extermination campaign against the local Aboriginal digital fringes in all other species. C. chunda sp. nov. is People (see etymology for Kumanjayiwalkerus kumanjayi readily separated from C. rubella by the obvious bright sp. nov.). After getting sea sick, the convict would lean yellow feet, well demarcated from the grey of the digits, over the edge of the boat to vomit and would yell out versus yellowish feet and digits without a well defined “Watch under” … “Chunda” to anyone below. The word boundary in C. rubella. Chunda then became an established Australian term for C. chunda sp. nov. is characterised by significant vomit, although there is no connection whatsoever speckling on the upper dorsum and a well-defined white between this species of frog and humans vomiting, upper labial area below the eye, versus little specking on although drunk Australians often vomit in toilet blocks at the dorsum of C. rubella and ill-defined lighter areas on camping grounds. the upper labial region. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 55

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 56 Australasian Journal of Herpetology and rounded. Tympanum is large, rounded and distinct. COLLEENEREMIA WATDAT SP. NOV. Second finger longer than first. Average adult size 35 mm LSIDurn:lsid:zoobank.org:act:2AF9CC4A-09B4-4F28- in length. A5AC-4C5132C47287 Species within the genus Colleeneremia are separated Holotype: A preserved specimen at the Western from the morphologically similar species within the genus Australian Museum, Perth, Western Australia, Australia, Audaxura gen. nov., their closest living relatives (with a specimen number R62379 collected at Mount Narryer divergence of 17 MYA according to Duellman et al. 2016) Station, Western Australia, Australia, Latitude -26.3500 as follows: Colleeneremia species always exhibits a very S., Longitude 115.5500 E. broad, dark stripe on the side of the head and body. This This government-owned facilility allows access to its is not found in species within Audaxura gen. nov.. holdings. Furthermore Audaxura gen. nov. have pale regular or Paratypes: Six preserved specimens at the Western irregular stripes or patches on the dorsal surface of the Australian Museum, Perth, Western Australia, Australia, body not exhibited in the same configuration or form in specimen numbers R62374, R62375, R62376, R62377, any Colleeneremia species. Colleeneremia have shorter R62378 and R62380 all collected at Mount Narryer limbs than Audaxura gen. nov.. The TL/S-V ratios Station, Western Australia, Australia, Latitude -26.3500 provides a means of distinguishing the genera, this being S., Longitude 115.5500 E. 0.335-0.432 for Colleeneremia and 0.477-0.520 for Diagnosis: The five species, Colleeneremia watdat sp. Audaxura gen. nov.. nov. from the Pilbara, Gascoyne and Murchison regions Brevicrusyla gen. nov. includes two species from the of western Western Australia, C. bogfrog sp. nov. from Indonesian side of New Guinea that are morphologically the Macdonnell Ranges of Central Australia, C. similar to species within the genera Colleeneremia Wells dunnyseat sp. nov. from the Flinders Ranges of South and Wellington, 1985 and Audaxura gen. nov. as detailed Australia, C. wifi from western New South Wales and above, but are readily separated from them in that adults southern Queensland and C. chunda from the Cape York of Brevicrusyla gen. nov. while having a smooth dorsum region in far north Queensland have until now all been on the body like the other two genera, instead has treated as populations of putative C. rubella (Keferstein, tubercles on the head, that is not seen in the other two 1868), herein confined to the dry tropics of the Northern genera. Tadpoles of Brevicrusyla gen. nov. have a long Territory and the Kimberley district of Western Australia. muscular tail, with narrow dorsal and ventral fins, versus Another morphologically similar species C. electrica a relatively short tail, with broad fins in species within (Ingram and Corben, 1990), was prior to 1990 also Audaxura gen. nov. and Colleeneremia. treated as a regional population of C. rubella. Distribution: Colleeneremia dunnyseat sp. nov. occurs The seven species comprise the total of the subgenus at the Flinders Ranges of South Australia, Australia. Colleeneremia Wells and Wellington, 1985 as defined in Etymology: In 2019 I spent a week at Leigh Creek in this paper. South Australia, to educate local miners about snake While these species within Colleeneremia have diverse handling and snake safety where I managed to kill yet habitat choices, their distribution does coincide with another motor vehicle, that one being a Toyota Rav4. biogeographical barriers generally affecting saxicoline I also spent considerable time surveying the local fauna genera of Australian fauna. including C. dunnyseat sp. nov.. This is including for example species of Varanid lizards While at Leigh Creek I tried to find one or more members (Hoser 2013, 2014, 2015g, 2018b-e), Ring-tailed of the local Adnyamathanha Aboriginal Tribe in order to Dragons in the genus Ctenophorus Fitzinger, 1843 find out the local tribal name for this species of frog. (Hoser 2020c) other lizards within the genus Egernia Unfortunately pretty much the entire tribe had been killed Gray, 1838 sensu lato (Hoser 2018a) and also Petrogale off a hundred years prior as part of the British Empire Gray, 1837 wallabies (Hoser, 2020d). genocide of the Australian Aboriginals. The seven aforementioned species are separated from This was done so that the British King could steal the one another as follows: entire country and pillage it as they saw fit. The type form of C. rubella and C. chunda sp. nov. There were some Adnyamathanha still hiding in the hills conforms to the genus description for Colleeneremia. behind the Leigh Creek reservoir, occasionally emerging They can be further diagnosed as follows: Compared to to grab some Freshwater Tortoises in the dam lake. all other species in the genus, these two species have But they ran when I approached them for fear of being relatively undeveloped lateral digital fringes and a shot. After all my skin was “white” and they knew white relatively narrow head (medium in cross-section), versus people carried and used guns against them (see broad (broad in cross-section) and well-developed lateral etymology for Kumanjayiwalkerus kumanjayi sp. nov.). digital fringes in all other species. C. chunda sp. nov. is The locals in the Leigh Creek mining camp called the readily separated from C. rubella by the obvious bright relevant frogs “dunny seat frogs” as they regularly yellow feet, well demarcated from the grey of the digits, secreted themselves under the toilet seat. versus yellowish feet and digits without a well defined Dunny is an Australian word for toilet. Men and women boundary in C. rubella. would find them by day when they lifted a toilet seat and C. chunda sp. nov. is characterised by significant saw then balled up as they slept. speckling on the upper dorsum and a well-defined white Hence the scientific name “dunnyseat”. upper labial area below the eye, versus little specking on the dorsum of C. rubella and ill-defined lighter areas on the upper labial region. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 57

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 58 Australasian Journal of Herpetology and rounded. Tympanum is large, rounded and distinct. Territory and the Kimberley district of Western Australia. Second finger longer than first. Average adult size 35 mm Another morphologically similar species C. electrica in length. (Ingram and Corben, 1990), was prior to 1990 also Species within the genus Colleeneremia are separated treated as a regional population of C. rubella. from the morphologically similar species within the genus The seven species comprise the total of the subgenus Audaxura gen. nov., their closest living relatives (with a Colleeneremia Wells and Wellington, 1985 as defined in divergence of 17 MYA according to Duellman et al. 2016) this paper. as follows: Colleeneremia species always exhibits a very While these species within Colleeneremia have diverse broad, dark stripe on the side of the head and body. This habitat choices, their distribution does coincide with is not found in species within Audaxura gen. nov.. biogeographical barriers generally affecting saxicoline Furthermore Audaxura gen. nov. have pale regular or genera of Australian fauna. irregular stripes or patches on the dorsal surface of the This is including for example species of Varanid lizards body not exhibited in the same configuration or form in (Hoser 2013, 2014, 2015g, 2018b-e), Ring-tailed any Colleeneremia species. Colleeneremia have shorter Dragons in the genus Ctenophorus Fitzinger, 1843 limbs than Audaxura gen. nov.. The TL/S-V ratios (Hoser 2020c) other lizards within the genus Egernia provides a means of distinguishing the genera, this being Gray, 1838 sensu lato (Hoser 2018a) and also Petrogale 0.335-0.432 for Colleeneremia and 0.477-0.520 for Gray, 1837 wallabies (Hoser, 2020b). Audaxura gen. nov.. The seven aforementioned species are separated from Brevicrusyla gen. nov. includes two species from the one another as follows: Indonesian side of New Guinea that are morphologically The type form of C. rubella and C. chunda sp. nov. similar to species within the genera Colleeneremia Wells conforms to the genus description for Colleeneremia. and Wellington, 1985 and Audaxura gen. nov. as detailed They can be further diagnosed as follows: Compared to above, but are readily separated from them in that adults all other species in the genus, these two species have of Brevicrusyla gen. nov. while having a smooth dorsum relatively undeveloped lateral digital fringes and a on the body like the other two genera, instead has relatively narrow head (medium in cross-section), versus tubercles on the head, that is not seen in the other two broad (broad in cross-section) and well-developed lateral genera. Tadpoles of Brevicrusyla gen. nov. have a long digital fringes in all other species. C. chunda sp. nov. is muscular tail, with narrow dorsal and ventral fins, versus readily separated from C. rubella by the obvious bright a relatively short tail, with broad fins in species within yellow feet, well demarcated from the grey of the digits, Audaxura gen. nov. and Colleeneremia. versus yellowish feet and digits without a well defined Distribution: Colleeneremia watdat sp. nov. occurs in boundary in C. rubella. the Pilbara, Gascoyne and Murchison regions of western C. chunda sp. nov. is characterised by significant Western Australia, Australia. speckling on the upper dorsum and a well-defined white Etymology: In 1981, I was in the wilderness of Western upper labial area below the eye, versus little specking on Australia with some Yamatji Aboriginal elders sharing a the dorsum of C. rubella and ill-defined lighter areas on few drinks. When I pulled a specimen of this species out the upper labial region. of a bag he exclaimed “watdat”, so I took that as the local C. chunda sp. nov. has well defined areas of yellow in the word for this taxon. groin, versus ill-defined and spreading up the lower flank COLLEENEREMIA WIFI SP. NOV. in C. rubella. LSIDurn:lsid:zoobank.org:act:7734BCBF-314E-4954- Both C. chunda sp. nov. and C. rubella have either BEBB-D01E4405958D smooth, or very minutely granular skin on the upper Holotype: A preserved specimen at the Australian surface. Museum, Sydney, New South Wales, Australia, specimen C. chunda sp. nov. is further separated from all other number R.43573 collected at Walgett, New South Wales, species by having a poorly formed and semi-distinct mid- Australia, Latitude -29.416 S., Longitude 147.566 E. vertebral line. This government-owned facility allows access to its C. chunda sp. nov. and C. rubella further differ from the holdings. other species by having a distinctively bitonal upper Paratypes: Five preserved specimens at the Australian surface, with a distinct broad grey or red-brown band Museum, Sydney, New South Wales, Australia, specimen down the middle of the back on a beige or fawn numbers R.43574, R.43575, R.43576, R.45511 and background. R.45512 all collected at Walgett, New South Wales, C. electrica (Ingram and Corben, 1990) is separated from Australia, Latitude -29.416 S., Longitude 147.566 E. the other six species by having three distinctive dark bars Diagnosis: The five species, Colleeneremia watdat sp. or blotches on the back, a dorsum that is obviously nov. from the Pilbara, Gascoyne and Murchison regions tubercular and the yellow patches in the groin and arm of western Western Australia, C. bogfrog sp. nov. from pits are bold and distinctive and made more so by being the Macdonnell Ranges of Central Australia, C. edged with dark brownish-grey bordering, not seen in any dunnyseat sp. nov. from the Flinders Ranges of South other species. Australia, C. wifi from western New South Wales and C. bogfrog sp. nov. is separated from the other six southern Queensland and C. chunda from the Cape York species by having a pinkish red colour on the dorsum region in far north Queensland have until now all been and flanks with pinkish white and very light grey marbling. treated as populations of putative C. rubella (Keferstein, The dorsum has smooth or leathery skin, but without 1868), herein confined to the dry tropics of the Northern

obvious raised tubercles. Speckling is minimal and where present, usually on the snout and rear, it is still scattered, Colleeneremia wifi sp. nov. minimal and very faded as to be indistinct. There are also irregular purple-grey patches running down two longitudinal lines on the upper dorsum, not near the midline and these too are faded. C. watdat sp. nov. is similar in dorsal patterning to C. bogfrog sp. nov. except that there is scattered flecking across all the body, as opposed to more being at front of snout and rear. C. watdat sp. nov. is also instead a mud- brownish coloured frog (not pinkish-red) and also has a strongly granulated dorsum and to a greater degree than any other species in the complex. Yellow in the groin and Moonie, Queensland armpits is either non-existent or minimal. C. watdat sp. nov. is further characterised by having a well defined dark brown stripe running from snout, and: through eye and ear over back leg to flank. The line is https://www.flickr.com/photos/27026445@N06/ well defined, contrasting with the lighter brown surfaces 32155803631/ adjacent and clearly runs across the entire tympanum. C. dunnyseat sp. nov. in life can be seen in Anstis (2013) C. dunnyseat sp. nov. is readily separated from all other on page 302 at top right image. six species by having a light greyish brown, almost tending to white coloured dorsum with distinctive dark C. wifi sp. nov. in life can be seen in Hoser (1989) on brownish-black spots and mottling spread relatively page 42 in the top photo and Cogger (2014) on page evenly across the dorsum. Limbs are also creamish in 188, top left photo. colour with obvious darker flecks and mottling. This taxon The subgenus Balatusrana subgen. nov. is separated comes across as a uniquely speckled frog, due to the from the nominate subgenus Colleeneremia Wells and high density of spotting and it being relatively evenly Wellington, 1985 (being all seven species separated by spaced on the contrasting lighter background, so the the descriptions herein) by having conspicuous webbing dorsum appears spotted, as opposed to one with on the fingers, reaching at least as far as the base of the blotches as may seen in some of the other species. penultimate phalanx of the fourth finger. Yellow in armpits and groin is often absent or when In the nominate subgenus Colleeneremia the fingers present not as brilliant, or prominent as in other species. have only rudimentary webbing. C. wifi sp. nov. is readily separated from the other six In addition to the character states just mentioned for each species by being chocolate brown on the dorsum and subgenus, both subgenera, comprising the totality of the gun metal grey on the flanks. The dorsum is irregularly, genus Colleeneremia are separated from all other but heavily peppered, with the same gun-metal grey the Australasian Tree Frogs (Pelodryadidae) by the following result being these patches of peppering forming semi- unique suite of characters: Grey, brown or fawn above, distinct areas of grey surrounded by otherwise mainly usually with a broad, darker vertebral band, bounded on brown pigment. The upper surfaces of the limbs are the either side by a lighter brown zone. There is a blackish same chocolate brown colour as the body, but with even stripe along the side of the head, continuing behind the heavier grey peppering. The labial area is also generally eye, over the base of the forelimb and along the side of grey and the top of the snout is brown, with either no the body almost to the groin. Dorsal surface of the body peppering or minimal peppering that tends to be faded. and limbs is flecked with dark brown or black and Toes may be brown or grey (in a single specimen) sometimes small to medium sized patches of darker depending on which are peppered heavily and which are pigment. Hind side of thighs brown with fine white spots. not. The upper iris is a dull orange brown in this species, Groin is usually lemon-yellow. Ventral surface white, versus brilliant red or orange in all the other six. cream or yellowish. Dorsal surface usually smooth or with C. rubella in life can be seen in Anstis (2013) on page numerous tiny granules above and coarsely granular 301 in the two right hand photos. below. Vomerine teeth almost entirely behind the C. chunda sp. nov. in life can be seen in Anstis (2013) on choanae. Pectoral fold prominent. Finger and toe discs page 302, on right (bottom image) and in Vanderduys large. Toes about two thirds webbed. Inner metatarsal (2012) on page 69 bottom. tubercle moderate and elongated, outer tubercle is small and rounded. Tympanum is large, rounded and distinct. C. electrica in life can be seen in Anstis (2013) on page Second finger longer than first. Average adult size 35 mm 192, two right hand photos, and Cogger (2014) on page in length. 162 (two photos). Species within the genus Colleeneremia are separated C. bogfrog sp. nov. in life can be seen in Anstis (2013) on from the morphologically similar species within the genus page 302 on left at top. Audaxura gen. nov., their closest living relatives (with a C. watdat sp. nov. in life can be seen online at: divergence of 17 MYA according to Duellman et al. 2016) https://www.flickr.com/photos/27026445@N06/ as follows: Colleeneremia species always exhibits a very 31419696523/ broad, dark stripe on the side of the head and body. This and: is not found in species within Audaxura gen. nov.. https://www.flickr.com/photos/warpedtime/2245649035/ Furthermore Audaxura gen. nov. have pale regular or Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 60 Australasian Journal of Herpetology irregular stripes or patches on the dorsal surface of the Diagnosis: The south-east Queensland subspecies body not exhibited in the same configuration or form in Colleeneremia dentata toowoombaensis subsp. nov. is any Colleeneremia species. Colleeneremia have shorter similar in most respects to nominate C. dentata limbs than Audaxura gen. nov.. The TL/S-V ratios (Keferstein, 1868) from New South Wales. provides a means of distinguishing the genera, this being Both would key out as “Litoria dentata” in Anstis (2013) 0.335-0.432 for Colleeneremia and 0.477-0.520 for and Cogger (2014). Audaxura gen. nov.. C. dentata toowoombaensis subsp. nov. is readily Brevicrusyla gen. nov. includes two species from the separated from C. dentata dentata as follows: Indonesian side of New Guinea that are morphologically Both have a similar dorsal pattern incorporating an similar to species within the genera Colleeneremia Wells irregularly shaped broad mid vertebral stripe that is dark and Wellington, 1985 and Audaxura gen. nov. as detailed in colour, with lighter surfaces on the sides of the dorsum above, but are readily separated from them in that adults and upper flank.all also have yellow in the armpits, groin of Brevicrusyla gen. nov. while having a smooth dorsum and yellowish toes. on the body like the other two genera, instead has However C. dentata toowoombaensis subsp. nov. has a tubercles on the head, that is not seen in the other two significantly greater contrast between dark and light on genera. Tadpoles of Brevicrusyla gen. nov. have a long the dorsum (resting by day) than seen in C. dentata muscular tail, with narrow dorsal and ventral fins, versus dentata and the stripe from snout, through eye and a relatively short tail, with broad fins in species within tympanum to upper anterior flank is dark-grey to black Audaxura gen. nov. and Colleeneremia. and distinct, versus light grey or brown and ranging from Distribution: C. wifi sp. nov. is generally found in the distinct to semi-distinct. Perhaps the most obvious region of the Darling River basin in western New South diference between the two subspecies is the that the toes Wales and inland southern Queensland, with a of C. dentata toowoombaensis subsp. nov. are a brilliant distribution towards the coast north of the south-east yellow colour, strongly contrasting with the grey or brown Queensland subtropical wet zone, but not including Cape feet, versus at most slightly yellowish and with minimal York or nearby areas to there, including the wet tropics of contrast between the non-yellow feet in C. dentata far north Queensland. dentata. Etymology: In 1992, I was speaking with a well-known The subgenus Balatusrana subgen. nov. herein treated Aboriginal Elder and ex underworld figure, Steve Gordon, as monotypic for C. dentata is separated from the also a commissioner with the Australian Federal nominate subgenus Colleeneremia Wells and Wellington, Government’s Aboriginal and Torres Strait Islander 1985 (herein treated as seven species as outlined in this Commission (ATSIC). He liked the job as ATSIC paper) by having conspicuous webbing on the fingers, Commissioner as in his own words “I get paid heaps to reaching at least as far as the base of the penultimate do bugger all!” He was from Brewarrina in New South phalanx of the fourth finger. Wales, which is where C. wifi sp. nov. occurs and I asked In the nominate subgenus Colleeneremia the fingers him about these and other frogs in the area. I did this by have only rudimentary webbing. showing him the relevant pictures in the book Hoser In addition to the character states just mentioned for each (1989). He called the relevant species the “wife ee” frog subgenus, both subgenera, comprising the totality of the and when I asked why, he said it was because of the genus Colleeneremia are separated from all other bleating call of the species, which sounded like a wife Australasian Tree Frogs (Pelodryadidae) by the following yelling at you. Hence the species name “wifi”. It does not unique suite of characters: Grey, brown or fawn above, have a connection with the well known “wifi” of more usually with a broad, darker vertebral band, bounded on recent wireless internet fame. For more about other either side by a lighter brown zone. There is a blackish relevant to herpetology conversations with Steve Gordon, stripe along the side of the head, continuing behind the refer to Hoser (1993) and Hoser (1996). eye, over the base of the forelimb and along the side of COLLEENEREMIA (BALATUSRANA) DENTATA the body almost to the groin. Dorsal surface of the body TOOWOOMBAENSIS SUBSP. NOV. and limbs is flecked with dark brown or black and LSIDurn:lsid:zoobank.org:act:9E5884CC-18ED-40E3- sometimes small to medium sized patches of darker B022-5AEB01982952 pigment. Hind side of thighs brown with fine white spots. Holotype: A preserved male specimen at the Groin is usually lemon-yellow. Ventral surface white, Queensland Museum, Brisbane, Queensland, Australia, cream or yellowish. Dorsal surface usually smooth or with specimen number J93230 collected from the Esk- numerous tiny granules above and coarsely granular Hampton Road, between Perseverance Hall Road and below. Vomerine teeth almost entirely behind the Clive Road, Perseverance, Queensland, Australia, choanae. Pectoral fold prominent. Finger and toe discs Latitude -27.3733 S., Longitude 152.1081 E. large. Toes about two thirds webbed. Inner metatarsal This government-owned facility allows access to its tubercle moderate and elongated, outer tubercle is small holdings. and rounded. Tympanum is large, rounded and distinct. Paratype: A preserved specimen at the Queensland Second finger longer than first. Average adult size 35 mm Museum, Brisbane, Queensland, Australia, specimen in length. number J93231 collected from the Esk-Hampton Road, Species within the genus Colleeneremia are separated between Perseverance Hall Road and Clive Road, from the morphologically similar species within the genus Perseverance, Queensland, Australia, Latitude -27.3733 Audaxura gen. nov., their closest living relatives (with a S., Longitude 152.1081 E. divergence of 17 MYA according to Duellman et al. 2016)

as follows: Colleeneremia species always exhibits a very Distribution: Wetter parts of south-east Australia, broad, dark stripe on the side of the head and body. This ranging from South Australia, through Victoria and is not found in species within Audaxura gen. nov.. Tasmania, along the New South Wales coast to south- Furthermore Audaxura gen. nov. have pale regular or east Queensland, with outlier populations at Eungella, irregular stripes or patches on the dorsal surface of the west of Mackay and the Atherton Tableland and adjacent body not exhibited in the same configuration or form in mountains in the southern wet tropics of North any Colleeneremia species. Colleeneremia have shorter Queensland. limbs than Audaxura gen. nov.. The TL/S-V ratios Content: Rawlinsonia ewingi (Duméril and Bibron, 1841) provides a means of distinguishing the genera, this being (type species); R. alpina (Fry, 1915); R. corbeni Wells 0.335-0.432 for Colleeneremia and 0.477-0.520 for and Wellington, 1985; R. ventrileuco sp. nov.; R. Audaxura gen. nov.. jervisensis (Duméril and Bibron, 1851); R. littlejohni The nominate form of C. dentata dentata in life is (White, Whitford and Mahoney, 1994); R. paraewingi depicted in Cogger (2014) on page 161. (Wilson, Loftus-Hills and Littlejohn, 1971); R. revelata C. dentata toowoombaensis subsp. nov. in life is depicted (Ingram, Corben and Hosmer, 1982); R. verreauxii in Vanderduys (2012) on page 37. (Duméril, 1853). Brevicrusyla gen. nov. includes two species from the RAWLINSONIA VENTRILEUCO SP. NOV. Indonesian side of New Guinea that are morphologically LSIDurn:lsid:zoobank.org:act:B00A719B-60AD-4BAB- similar to species within the genera Colleeneremia Wells B092-0CE78100283F and Wellington, 1985 and Audaxura gen. nov. as detailed Holotype: A preserved specimen in the Queensland above, but are readily separated from them in that adults Museum, Brisbane, Queensland, Australia, specimen of Brevicrusyla gen. nov. while having a smooth dorsum number J35087 collected at Mount William Road, 18 km on the body like the other two genera, instead has from Dalrymple Heights, near Eungella, near Mackay, tubercles on the head, that is not seen in the other two Queensland, Australia, Latitude -21.03 S., Longitude genera. Tadpoles of Brevicrusyla gen. nov. have a long 148.6 E. muscular tail, with narrow dorsal and ventral fins, versus This government-owned facility allows access to its a relatively short tail, with broad fins in species within holdings. Audaxura gen. nov. and Colleeneremia. Paratypes: Four preserved specimens at the Distribution: C. dentata toowoombaensis subsp. nov. is Queensland Museum, Brisbane, Queensland, Australia, believed to be restricted to South-east Queensland, specimen numbers J35105-6 and J35115-6 all collected generally north of the border ranges area on the New at Mount William Road, 18 km from Dalrymple Heights, South Wales and Queensland border. Nominate C. near Eungella, near Mackay, Queensland, Australia, dentata is believed to occupy the rest of the range for this Latitude -21.03 S., Longitude 148.6 E. species and subgenus, being the coast and nearby Diagnosis: R. ventrileuco sp. nov. has until now been ranges of New South Wales, along its entire length and treated as an isolated population of R. revelata (Ingram, into far north-east Victoria. Corben and Hosmer, 1982), with a type locality of Etymology: The subspecies C. dentata toowoombaensis “O’Reillys (28.14' S, 153.08' E), Lamington Plateau, SE. subsp. nov. is named in reflection of the type locality for Queensland”, Australia, as has the species R. corbeni the taxon. Wells and Wellington, 1985. RAWLINSONINA SUBTRIBE NOV. All three would be identified as R. revelata, called “Litoria RAWLINSONIA WELLS AND WELLINGTON, 1985 revelata” in either Anstis (2013) or Cogger (2014). Type species: Hyla ewingi Duméril and Bibron, 1841. All three have similar morphology and call, although all Diagnosis: The nine known living species within the are widely disjunct in terms of distribution and clearly genus Rawlinsonia Wells and Wellington, 1985 (one evolving as separate species. They are also formally described within this paper) are readily morphologically divergent. separated from all other Australasian Tree Frogs While divergence between the three populations is (Pelodryadidae) by the following unique suite of relatively recent in geological terms, it still appropriate characters: smallish frogs usually about 35 mm in body that each be recognized as separate species, as length. Fingers are free or only webbed at the base. First opposed to subspecies. finger is much shorter and smaller than the second, when I also note that while Anstis (2013) treats all populations pressed together the tip of the first finger reaches no as a single species, she also makes comments indicating further than the base of the disc of the second finger. in her view the likelihood that there is more than one Hind edge of vomerine teeth are between the choanae. species being called “Litoria revelata” and to that extent Usually one or a pair of semidistinct mid dorsal patches, details obvious differences between forms. darker than the ground colour, extending from a line The three species are readily separated from one joining the centre of the eyes; white stripe below eye, if another as follows: present does not extend beyond the anterior edge of the base of the forelimb. By distribution, Rawlinsonia revelata from coastal New South Wales, immediately adjacent ranges and nearby Duellman et al. (2016) found that the living species in this wetter parts of south-east Queensland is separated from genus diverged from their nearest living relatives 23.2 R. corbeni from the southern wet tropics of far north MYA, giving support to the erection of this genus by Wells Queensland and R. ventrileuco sp. nov. from the and Wellington, 1985 and this is before one considers Eungella uplands region near Mackay in north eastern the significant morphological divergence of the species Queensland. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. group. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 62 Australasian Journal of Herpetology

The three species are turn also separated from one from nearest living relatives according to Duellman et al. another as follows: R. revelata and R. ventrileuco sp. nov. (2016), genus-level recognition of this group of species is have a few small black spots in the groin, versus an obvious taxonomic judgement. prominent black blotches in the groin and red-orange on The only surprising thing is that this judgement has not the back of the thighs as seen in R. corbeni. been sooner, and hence the erection of a new genus R. ventrileuco sp. nov. and R. corbeni are in turn both herein. separated from R. revelata by having small pointed I note that the nearest living relatives (from within the tubercles running in a distinct line down the upper New Guinea area) are also in what have until now been surface of the forearm, versus mainly scattered tubercles ungrouped species and they too are placed in new on the upper surface of the forearm in R. corbeni. genera, all of which (as a group) are most closely related Scattered, raised tubercles on the upper surface of the to Maxinehoserranae gen. nov. and not any other hind limb are prominent in both R. revelata and R. species. corbeni but not in R. ventrileuco sp. nov.. Species within Maxinehoserranae gen. nov. are The upper iris of both R. corbeni and R. ventrileuco sp. separated from all other Australasian (Australian and nov. is noticeably lighter than the lower, versus not so in New Guinea) Tree Frogs (Pelodryadidae) by the following R. revelata. suite of characters: No vomerine teeth (Australian The upper forearms of both R. corbeni and R. ventrileuco species) or in two small patches between the choanae sp. nov. is dark brown, versus light brown or yellow in R. (New Guinea species); dorsal colour is usually green, or revelata. occasionally fawn or a mixture of green and bronze; The transverse row of usually about four tubercles dorsal surface has a broad vertebral band of bronze between the eyes is found in all three species. bordered on either side by green; in terms of flecks or The three species R. corbeni, R. ventrileuco sp. nov. and blotches, there are at most a few dark flecks on the R. revelata are in turn separated from all other species dorsal surface; dorsal surface is smooth; at least one within the genus Rawlinsonia Wells and Wellington, 1985 strong pectoral fold; no tubercles above the eye; brown by having finger and toe discs that are conspicuously head streak present; tympanum brown; internarial broader than the digits; conspicuous black spots or distance/eye-naris distance ratio is less than 1.0. blotches on the groin and a mid-dorsal patch divided Frogs within the subgenus Vegrandihyla subgen. nov. are along at least part of its length, including between the separated from the nominate subgenus eyes. Maxinehoserranae subgen. nov. by having concealed All species within the genus Rawlinsonia Wells and surfaces of the legs being bright red in colour (in life), Wellington, 1985 are readily separated from all other versus blue-black, brown, yellow to orange in colour in all Australasian Tree Frogs (Pelodryadidae) by the following other species. unique suite of characters: smallish frogs usually about Vegrandihyla subgen. nov. are further separated from 35 mm in body length. Fingers are free or only webbed at New Guinea species within Maxinehoserranae subgen. the base. First finger is much shorter and smaller than nov. by their non-overlapping HL/HW ratios, 1.027-1.189 the second, when pressed together the tip of the first versus 1.243-1.254. finger reaches no further than the base of the disc of the In terms of the Australian species in Maxinehoserranae second finger. Hind edge of vomerine teeth are between subgen. nov. the magnitude of the difference is less, the choanae. Usually one or a pair of semidistinct mid being a mean HL/HW for Vegrandihyla subgen. nov. dorsal patches, darker than the ground colour, extending 1.04, versus 1.10 in the the Australian species in from a line joining the centre of the eyes; white stripe Maxinehoserranae subgen. nov.. below eye, if present does not extend beyond the anterior Distribution: Found in New Guinea, including islands edge of the base of the forelimb. north, south and east, as well as tropical northern Distribution: R. ventrileuco sp. nov. is known only from Australia. the vicinity of Eungella, near Mackay in north-east Etymology: The new genus is named in honour of Queensland and is believed to be isolated to this Maxine Hoser of Margate a loyal subject (minion) the relatively wet uplands region. United Kingdom of England, Scotland, Wales, Northern Etymology: The name “ventrileuco” refers to the whitish Island, Gibralta, the Falkland Islands and formerly venter of this species of frog in the adult form. including Hong Kong and other colonies, in recognition of MAXINEHOSERRANINI TRIBE NOV. her services to the author in herpetology. MAXINEHOSERRANAE GEN. NOV. Content: Maxinehoserranae bicolor (Gray, 1842) (type LSIDurn:lsid:zoobank.org:act:B603EFB7-4D33-4BDF- species); M. albolabris (Wandolleck, 1911); M. bibonius 86C7-9552414B2127 (Kraus and Allison, 2014); M. brettbarnetti sp. nov.; M. Type species: Eucnemis bicolor Gray, 1842. chloristona (Menzies, Richards and Tyler, 2008); M. contrastens (Tyler, 1968); M. eurynastes (Menzies, Diagnosis: Known in many books as the “Litoria bicolor Richards and Tyler, 2008); M. lodesdema (Menzies, complex” (e.g. Menzies 2006), this distinctive group of Richards and Tyler, 2008); M. maxinehoserae sp. nov.; M. frogs has long been recognized as distinct within the mystax (Van Kampen, 1906); M. piersoni sp. nov.; M. genus Litoria Tschudi, 1838 as defined by Menzies viranula (Menzies, Richards and Tyler, 2008). (2006), Cogger (2014), Anstis (2013) Eipper and Rowland (2018), Vanderduys (2012) and most other contemporary authors. With a divergence of 21.5 MYA

VEGRANDIHYLA SUBGEN. NOV. including the Kimberley Division of Western Australia, the LSIDurn:lsid:zoobank.org:act:5E1A91FF-2426-4DCA- top end of the Northern Territory and Cape york in B8DA-EDEB534437E3 Queensland, as well as most parts of New Guinea and Type species: Hyla contrastens Tyler, 1968. islands to the north, east and west of New Guinea. Diagnosis: Frogs within the subgenus Vegrandihyla Various species have been “split off” with Menzies, subgen. nov. are separated from the nominate subgenus Richards and Tyler (2008) formally restricting Maxinehoserranae subgen. nov. by having concealed Maxinehoserranae bicolor (Oken, 1816) (as Litoria surfaces of the legs being bright red in colour (in life), bicolor) to Northern Australia, with a view that the Cape versus blue-black, brown, yellow to orange in colour in all York population was closest to their New Guinea species, other species. but not conspecific with them. Vegrandihyla subgen. nov. are further separated from They did not put a name to that taxon and as of 2019, New Guinea species within Maxinehoserranae subgen. this taxon remained unnamed. nov. by their non-overlapping HL/HW ratios, 1.027-1.189 While restricting M. bicolor to the top end of the Northern versus 1.243-1.254. In terms of the Australian species in Territory and Western Australia, Menzies, Richards and Maxinehoserranae subgen. nov. the magnitude of the Tyler (2008) and no one since appears to have ever difference is less, being a mean HL/HW for Vegrandihyla countenanced the possibility that there may be more than subgen. nov. 1.04, versus 1.10 in the the Australian one species in north-west Australia, save for material to species in Maxinehoserranae subgen. nov.. this effect in a thesis by James (1998). Species within Maxinehoserranae gen. nov. are However over more than 3 decades of active fieldwork in separated from all other Australasian (Australian and the relevant region, it was always apparent that those New Guinea) Tree Frogs (Pelodryadidae) by the following specimens from the West Kimberley were radically suite of characters: No vomerine teeth (Australian different to those from near Darwin and were therefore a species) or in two small patches between the choanae separate species. (New Guinea species); dorsal colour is usually green, or As that one was also unnamed as of 2019, it too is occasionally fawn or a mixture of green and bronze; described herein. dorsal surface has a broad vertebral band of bronze All three species, namely L. bicolor from the Northern bordered on either side by green; in terms of flecks or Territory, M. maxinehoserae sp. nov. from the Kimberley blotches, there are at most a few dark flecks on the District of Western Australia and M. brettbarnetti sp. nov. dorsal surface; dorsal surface is smooth; at least one from Cape York in Queensland would key out as M. strong pectoral fold; no tubercles above the eye; brown bicolor in either Cogger (2014) or Anstis (2013). head streak present; tympanum brown; internarial The three Australian species are readily separated from distance/eye-naris distance ratio is less than 1.0. those in the genus outside Australia (New Guinea and Distribution: Found in highlands of Papua New Guinea, nearby offshore islands) by their call, which in the being mountains between Kundiawa and Wau-Bulolo at Australian species is a very distinctive short rolling around 1200-1500 m elevation. sound, or rasp, which accelerates slightly at the end of the sequence. Etymology: The subgenus name Vegrandihyla in Latin means “small” and “Hyla”, with Hyla known as a tree frog The three Australian species are separated from one and hence “small tree frog”, which accurately describes another as follows: the species in the subgenus. Adult male M. brettbarnetti sp. nov. and M. bicolor both Content: Maxinehoserranae (Vegrandihyla) contrastens have a well-defined broad band running from the back of (Tyler, 1968) (monotypic). the eye along the side of the back towards the groin. This is either green, or yellow with a stong greenish tinge in M. MAXINEHOSERRANAE (MAXINEHOSERRANAE) brettbarnetti sp. nov., versus yellow in M. bicolor. Below BRETTBARNETTI SP. NOV. this line, the border is brown in M. brettbarnetti sp. nov. LSIDurn:lsid:zoobank.org:act:96665B4E-56F6-45AA- versus brown, purple or grey in M. bicolor. 9CE3-7D8B889F0BEA The forelimbs of both sexes of adult M. brettbarnetti sp. Holotype: A preserved specimen at the South Australian nov. are always heavily peppered with dark brown Museum, Adelaide, South Australia, Australia, specimen pigment, versus not so in M. bicolor which have light number R12264, collected from 5 miles west of forelimbs. Cooktown, North Queensland, Australia, Latitude -15.45 The flanks of both sexes of adult M. brettbarnetti sp. nov. S., Longitude 145.17 E. has a moderate amount of scattered black peppering, This government-owned facility allows access to its versus none or very little in M. bicolour. holdings. In terms of further separation of the two species Menzies, Paratypes: Five preserved specimens at the Queensland Richards and Tyler (2008) stated M. bicolor are larger Museum, specimen numbers: J23245, J23246, J23247, than Queensland M. brettbarnetti sp. nov., with mean J23248 and J23249 collected from Marina Plains, North snout-vent 25.0 mm +/-0.98 versus 23.1 mm +/-1.58 mm Queensland, Australia, Latitude -14.55 S., Longitude in M. brettbarnetti sp. nov., Queensland M. brettbarnetti 143.8667 S. sp. nov., have larger heads, mean HL/HB 0.34 +/- 0.017 Diagnosis: Until now, the putative species originally versus 0.33 +/- 0.014 in M. bicolor, but there are no described as Hyla bicolor Oken, 1816, with a type locality differences in head proportions. of Port Essington, Northern Territory, has been treated as occurring across the northern wet tropics of Australia, ... CONTINUES IN AJH ISSUE 45

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192.

Hoser, R. T. 2020. For the first time ever! An overdue Availablereview and reclassification online of atAustralasian www.herp.net Tree Frogs (Amphibia: Anura: Pelodryadidae), including formal Copyright-descriptions Kotabi of Publishing12 tribes, 11 subtribes, 34- Allgenera, rights 26 reserved subgenera, 62 species and 12 subspecies new to science. Australasian Journal of Herpetology 44-46:1-192. AustralasianAustralasian JournalJournal ofof HerpetologyHerpetology

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online) ISSUES 44-46, PUBLISHED 5 JUNE 2020 Australasian Journal of Herpetology

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

... Continued from AJH Issue 44 ... Etymology: Named in honour of Brett Barnett of Ardeer, M. brettbarnetti sp. nov. have more widely placed nostrils, Victoria, Australia in recognition of his significant mean EN/IN 0.99 +/- 0.01 versus 1.06 +/- 0.01 in M. contributions to herpetology over many decades. bicolor.. A combination of two metrics, i.e. HB and HL/HB, MAXINEHOSERRANAE (MAXINEHOSERRANAE) provides better, but not absolute, separation MAXINEHOSERAE SP. NOV. (paraphrased here). M. maxinehoserae sp. nov. is by far LSIDurn:lsid:zoobank.org:act:C3873D43-BDD9-4F22- the most distinctive Australian species in the genus A5EB-0D5A72AAC722 Maxinehoserranae gen. nov.. It is readily separated from Holotype: A preserved specimen at the Western both other species as follows: Australian Museum, Australia, specimen number M. maxinehoserae sp. nov. has a distinctive, bright lime R47475, collected at Lake Gilbert, Beverley Springs green dorsum with thin well-defined chocolate brown line Station, near Charnley River, Prince Regent District, running from tip of snout to eye, after which the line Kimberley Division of Western Australia, Australia, continues as a broad stripe, over-writing the entire Latitude -16.5667 S., Longitude 125.2667 E. tympanum and going along the mid flank to the groin. On This government-owned facility allows access to its the flank of the body, the line is distinctive and well holdings. defined. It is bordered on top by the lime green dorsum Paratypes: Nine preserved specimens at the Western and below by a thick white band, bordered below with Australian Museum, specimen numbers R47476, brown, that then fades to the white underside. R47477, R47478, R47479, R47480, R47481, R47482, The chocolate coloured line from snout, through nostril to R47483 and R47484 all collected at Lake Gilbert, eye is bordered on top by a thin gold line in turn bordered Beverley Springs Station, near Charnley River, Prince by the lime green on top of the head and body. Regent District, Kimberley Division of Western Australia, This thin gold line upper border is not seen in the other Australia, Latitude -16.5667 S., Longitude 125.2667 E. two species. The limbs of Maxinehoserranae gen. nov. Diagnosis: Until now, the putative species originally have scattered brown spots and the dorsum of this described as Hyla bicolor Oken, 1816, with a type locality species is moderately granular. of Port Essington, Northern Territory, has been treated as M. brettbarnetti sp. nov. also has a moderately granular occurring across the northern wet tropics of Australia, dorsum, whereas M. bicolor has a smooth dorsum. including the Kimberley Division of Western Australia, the Specimens of M. brettbarnetti sp. nov. that have a top end of the Northern Territory and Cape york in brownish coloured line running from the ear to the groin Queensland, as well as most parts of New Guinea and do not have either the rich dark chocolate colour seen in islands to the north, east and west of New Guinea. M. maxinehoserae sp. nov. or the sharp well defined Various species have been “split off” with Menzies, upper edge on the dorsum as seen in M. maxinehoserae Richards and Tyler (2008) formally restricting sp. nov.. Maxinehoserranae bicolor (Oken, 1816) (as Litoria The upper dorsum of M. maxinehoserae sp. nov. does bicolor) to Northern Australia, with a view that the Cape not have the somewhat faded two tone colouration seen York population was closest to their New Guinea species, in both M. brettbarnetti sp. nov. and M. bicolor. but not conspecific with them. They did not put a name to that taxon and as of 2019, Images of M. maxinehoserae sp. nov. in life can be seen this taxon remained unnamed. in Tyler, Smith and Johnstone (1994) on plate 27 at While restricting M. bicolor to the top end of the Northern bottom and online at: Territory and Western Australia, Menzies, Richards and https://www.flickr.com/photos/euprepiosaur/ Tyler (2008) and no one since appears to have ever 22190269488/in/photolist-885upC-zNSW2f-z9rhah countenanced the possibility that there may be more than and: one species in north-west Australia, save for material to https://www.flickr.com/photos/euprepiosaur/ this effect in a thesis by James (1998). 21755237474/in/photolist-885upC-zNSW2f-z9rhah/ However over more than 3 decades of active fieldwork in and: the relevant region, it was always apparent that those https://www.flickr.com/photos/liquidghoul/4674588858/in/ specimens from the West Kimberley were radically photolist-885upC-zNSW2f-z9rhah/ different to those from near Darwin and were therefore a Images of M. brettbarnetti sp. nov. in life can be seen in separate species. Anstis (2013) on page 150 at bottom and page 151 at top As that one was also unnamed as of 2019, it too is left and page 153 top, in Vanderduys (2012) on page 26 described herein. in all images and Tyler and Davies (1986) in plate 20. All three species, namely L. bicolor from the Northern An image of M. bicolor in life can be seen in Anstis Territory, M. maxinehoserae sp. nov. from the Kimberley (2013) on page 150 at top. District of Western Australia and M. brettbarnetti sp. nov. Distribution: M. brettbarnetti sp. nov. is restricted to from Cape York in Queensland would key out as M. Cape York in Queensland. M. bicolor is restricted to the bicolor in either Cogger (2014) or Anstis (2013). area spanning the environs of Darwin, across the top of The three Australian species are readily separated from the Northern Territory, including offshore islands and the those in the genus outside Australia (New Guinea and western side of the Gulf of Carpentaria. nearby offshore islands) by their call, which in the Australian species is a very distinctive short rolling sound, or rasp, which accelerates slightly at the end of the sequence as well as an absence of vomerine teeth,

or very tiny ones, versus presence in New Guinea The upper dorsum of M. maxinehoserae sp. nov. does species. not have the somewhat faded two tone colouration seen The three Australian species are separated from one in both M. brettbarnetti sp. nov. and M. bicolor. another as follows: Images of M. maxinehoserae sp. nov. in life can be seen Adult male M. brettbarnetti sp. nov. and M. bicolor both in Tyler, Smith and Johnstone (1994) on plate 27 at have a well-defined broad band running from the back of bottom and online at: the eye along the side of the back towards the groin. This https://www.flickr.com/photos/euprepiosaur/ is either green, or yellow with a stong greenish tinge in M. 22190269488/in/photolist-885upC-zNSW2f-z9rhah brettbarnetti sp. nov., versus yellow in M. bicolor. Below and: this line, the border is brown in M. brettbarnetti sp. nov. https://www.flickr.com/photos/euprepiosaur/ versus brown, purple or grey in M. bicolor. 21755237474/in/photolist-885upC-zNSW2f-z9rhah/ The forelimbs of both sexes of adult M. brettbarnetti sp. and: nov. are always heavily peppered with dark brown https://www.flickr.com/photos/liquidghoul/4674588858/in/ pigment, versus not so in M. bicolor which have light photolist-885upC-zNSW2f-z9rhah/ forelimbs. Images of M. brettbarnetti sp. nov. in life can be seen in The flanks of both sexes of adult M. brettbarnetti sp. nov. Anstis (2013) on page 150 at bottom and page 151 at top has a moderate amount of scattered black peppering, left and page 153 top, in Vanderduys (2012) on page 26 versus none or very little in M. bicolour. in all images and Tyler and Davies (1986) in plate 20. In terms of further separation of the two species Menzies, An image of M. bicolor in life can be seen in Anstis Richards and Tyler (2008) stated M. bicolor are larger (2013) on page 150 at top. than Queensland M. brettbarnetti sp. nov., with mean Distribution: M. maxinehoserae sp. nov. is found snout-vent 25.0 mm +/-0.98 versus 23.1 mm +/-1.58 mm throughout the Kimberley Division of Western Australia in M. brettbarnetti sp. nov., Queensland M. brettbarnetti and into nearby parts of far north-west Western Australia, sp. nov., have larger heads, mean HL/HB 0.34 +/- 0.017 in the region generally south of the Daly River (inclusive). versus 0.33 +/- 0.014 in M. bicolor, but there are no M. bicolor is restricted to the area spanning the environs differences in head proportions. M. brettbarnetti sp. nov. of Darwin, across the top of the Northern Territory, have more widely placed nostrils, mean EN/IN 0.99 +/- including offshore islands and the western side of the 0.01 versus 1.06 +/- 0.01 in M. bicolor, but none of these Gulf of Carpentaria. characters will provide Etymology: the new species is named in honour of absolute separation as all measurements and ratios Maxine Hoser of Margate a loyal subject (minion) the show extensive overlap and so are of little practical value. United Kingdom of England, Scotland, Wales, Northern A combination of two metrics, i.e. HB and HL/HB, Island, Gibralta, the Falkland Islands and formerly provides better, but not absolute, separation including Hong Kong and other colonies, in recognition of (paraphrased here). her services to the author in herpetology. M. maxinehoserae sp. nov. is by far the most distinctive MAXINEHOSERRANAE (MAXINEHOSERRANAE) Australian species in the genus Maxinehoserranae gen. PIERSONI SP. NOV. nov.. It is readily separated from both other species as follows: LSIDurn:lsid:zoobank.org:act:10C131B1-4741-4046- 9702-4A782CFEC91F M. maxinehoserae sp. nov. has a distinctive, bright lime green dorsum with thin well-defined chocolate brown line Holotype: A preserved specimen in the University of running from tip of snout to eye, after which the line Papua New Guinea, Port Moresby, Papua New Guinea, continues as a broad stripe, over-writing the entire specimen number UP5276, collected from Yala River, tympanum and going along the mid flank to the groin. On Seram, Indonesia, Latitude 3.28 S., Longitude 128.99 E. the flank of the body, the line is distinctive and well This facility allows access to its holdings. defined. It is bordered on top by the lime green dorsum Paratypes: Four preserved specimens in the University and below by a thick white band, bordered below with of Papua New Guinea, Port Moresby, Papua New brown, that then fades to the white underside. Guinea, specimen numbers UP5271-74, collected from The chocolate coloured line from snout, through nostril to Yala River, Seram, Indonesia, Latitude 3.28 S., Longitude eye is bordered on top by a thin gold line in turn bordered 128.99 E. by the lime green on top of the head and body. Diagnosis: Maxinehoserranae piersoni sp. nov. from the This thin gold line upper border is not seen in the other island of Seram has until now been variously treated as two species. M. bicolor (Gray, 1842) (in most texts placed in the genus Litoria Tschudi, 1838) and herein restricted to the top end The limbs of Maxinehoserranae gen. nov. have scattered of the Northern Territory, Australia, or more recently as a brown spots and the dorsum of this species is moderately western population of the species formally named Litoria granular. eurynastes Menzies et al. (2008) with a type locality of M. brettbarnetti sp. nov. also has a moderately granular Madang, in the north of Papua New Guinea. dorsum, whereas M. bicolor has a smooth dorsum. However this taxon (M. piersoni sp. nov.) is neither and Specimens of M. brettbarnetti sp. nov. that have a herein formally described as a new species. brownish coloured line running from the ear to the groin While it would be diagnosed as Maxinehoserranae do not have either the rich dark chocolate colour seen in eurynastes (Menzies et al., 2008) using the diagnostic M. maxinehoserae sp. nov. or the sharp well defined information within Menzies et al. (2018) the two species upper edge on the dorsum as seen in M. maxinehoserae Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. sp. nov.. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 68 Australasian Journal of Herpetology are readily separated on the basis of the colouration of sp. nov. is 28 mm, versus 29 mm for the Yamur Lake and the concealed surfaces of the thighs. In M. piersoni sp. Siewa River forms, being of no significant difference. nov. these surfaces are vaguely mottled brown and not For the record M. viranula is separated from M. piersoni distinct, versus concealed surfaces of thighs with sparse sp. nov. by being a significantly smaller species to moderate sprinkling of blue-black. maximum snout-vent of 23.5 mm and HBF 26.4 mm. M. piersoni sp. nov. is further separated by having a M. viranula is further separated from M. piersoni sp. nov. generally variable but dull olive green coloured dorsum, by having a green dorsum with mid-dorsal bronze band; versus immaculate yellow green to bright green in M. narrow dark canthal stripe continuing through eye and eurynastes. tympanum, but becoming M. piersoni sp. nov. is further diagnosed by a pale yellow indistinct between green dorsum and white ventrum; band running from the upper lip to the mid-body and upper lip white, colour continuing below eye and vague dark dorsolateral stripe (versus bronze in M. tympanum before merging with ventral colour; male eurynastes); and males with a pale yellow venter; throat throat pale yellow, ventrum elsewhere white; groin and deep yellow and gold iris. concealed surfaces of thigh dark brown (versus an Both M. piersoni sp. nov. and M. eurynastes are indistinct mottled brown in M. piersoni sp. nov.). separated from all other species within M. viranula was illustrated in life by Menzies (2006) in Maxinehoserranae gen. nov. by the following quite of plate 44 as “Litoria bicolor”. characters: A larger species of Maxinehoserranae gen. M. piersoni sp. nov. was illustrated by Menzies (2006) in nov. with adult snout-vent 25.7-31.5 mm, HBF 27.3-33.0 plate 46 as “Litoria cf. bicolor, Yala River, Seram”. mm; snout rounded and projecting in profile, somewhat Based on the original description of Hyla albolabris angular from above; lores slightly sloping, flat; canthus Wandolleck, 1911 and relevant account in Tyler and rostralis rounded, straight; nostrils lateral, slightly visible Davies (1978), this taxon appears to fall within the genus from above; eyes large, tympanum large, distinct, except Maxinehoserranae gen. nov., but that placement is of extreme upper margin; shallow postorbital fold fades course tentative. before axilla; outer fingers about half webbed, others Tyler and Davies (1978) placed the taxon in its own webbed only at base; simple, pale brown nuptial pad; all species group, but provided no evidence or justification fingers with discs, larger than on the toes; toes fully for this quite drastic action. In key characters, M. webbed, except fourth with terminal phalanx free, all with albolabris clearly matches species within discs; dorsal skin finely granular; ventral surface more Maxinehoserranae gen. nov. (AKA “the bicolor group”) coarsely; throat wrinkled; vomerine teeth in two small including finger and toe webbing, adult size and colour, patches between the choanae and otherwise as for the both dorsal and ventral. On this basis, it is highly likely genus. that this taxon is in fact conspecific with M. viranula, or Menzies et al. (2008) placed the Seram population alternatively related to M. contrastens (Tyler, 1968) and described herein as M. piersoni sp. nov. within their possibily conspecific that that. If either situation is species M. eurynastes on the basis of biogeographic ultimately found to be correct, then M. albolabris will be proximity to New Guinea and the associated fact that the senior name and the other taxon name will move into other New Guinea frogs occur there, namely the its synonymy in accordance with the rules of the identified (by them) species of “Litoria amboinensis, L. International Code of Zoological Nomenclature (Ride et infrafrenata, Platymantis papuensis”. al. 1999). In doing so, they chose to ignore the morphological Distribution: Known definitively only from the southern divergence between their type form of M. eurynastes parts of the island of Seram, Indonesia, however this from Alexishafen, Madang Province Papua New Guinea taxon presumably occurs in all suitable lowland parts of and this new species from Seram, which is highlighted the island, and nearby islands. Specimens from Yamur here to separate the two taxa. Lake and Siewa River in the neck of the Bird’s Head of It should also be noted that biogeographically, Seram New Guinea, (south running drainages) are associates with southern New Guinea fauna and not that morphologically similar to M. piersoni sp. nov. and are of the north, of which their species M. eurynastes is. tentatively included within this species. Based on biogeography, the closest match to the Seram Etymology: Named in honour of Charles Pierson of population (identified herein as M. piersoni sp. nov.) Moss Vale, New South Wales, Australia, publisher of would logically be their southern New Guinea species, M. numerous books on Australian wildlife, including Hoser viranula (Menzies et al., 2008), with a type locality of (1989, 1991 and 1993) in recognition of his major Wegamu Camp, Bensbach River, Western Province, contributions to wildlife conservation in Australia and Papua New Guinea. assisting. However Menzies et al., 2008 identify a raft of His efforts were also critical in forcing the Australian morphological differences separating those two forms government to revoke draconian and anti-conservation and hence the only logical conclusion is that M. piersoni wildlife laws that were enforced at gunpoint in Australia sp. nov. must therefore be a separate species. for two decades from the 1970’s to the 1990’s as detailed The population of frogs from Yamur Lake and Siewa in Hoser (1993 and 1996). Those laws which, Person’s River in the neck of the Bird’s Head of New Guinea, efforts finally had revoked did in that 20 year period (south running drainages) are morphologically similar to cause the extinctions of several species including frogs M. piersoni sp. nov. and are tentatively included within and at least one dragon species (Tympanocryptis this species. The quoted mean snout-vent for M. piersoni pinguicolla Mitchell, 1948) (Hoser 2019a, 2019b).

ANGULARANTA GEN. NOV. lobulated anteriorly in retaining a double condition and LSIDurn:lsid:zoobank.org:act:5BEECB54-ABB0-444B- does not extend between the nasals. The frontoparietal B17E-36C69FBF7105 fontanelle is large and ovoid. The squamosal has a short Type species: Hyla (Litoria) arfakiana Peters and Doria, zygomatic ramus and slightly longer otic ramus. The 1878. quadratojugal is not developed. The pars facialis is Diagnosis: The genus Angularanta gen. nov. includes shallow and the short posterior process does not most of the better-known small, medium or large sized articulate with the maxillary process of the nasal. The mountain stream dwelling tree frogs of New Guinea as alary processes of the premaxillaries are well developed, well as a number of larger but morphologically similar bifurcated at their extremities, widely separated medially lowland species. and perpendicular to the pars dentalis. The palatine processes are well developed and do not articulate with The described species in the genus Angularanta gen. each other medially. The prevomers are reduced. The nov. are most easily separated from all other Australasian sacral diapophyses are broadly expanded and the ilia (Australian and New Guinea) Tree Frogs (Pelodryadidae) extend one third along their length. No flange is present by separation of each of the relevant subgenera. on the third metacarpal and the intercalary structures are Frogs in the nominate subgenus Angularanta subgen. long and cartilaginous. Alary processes of the hyoid plate nov. are separated from all other Australasian (Australian are lacking. The adductor mandibulae externus and New Guinea) Tree Frogs (Pelodryadidae) by one of superficialis is absent. the two following suites of characters: Frogs within the subgenus Scelerisqueanura subgen. 1/ Small to medium sized, stream-breeding species with nov. are separated from all other Australasian (Australian unwebbed or slightly webbed, long fingers, large finger and New Guinea) Tree Frogs (Pelodryadidae) by the discs and fully webbed toes. Intercalary structures are following suite of characters: Small (adult males 25.8- broad or elongate and ossified. Straight canthus rostralis. 30.3 mm) frogs. Dorsum is chocolate brown, with or The hyoid plate lacks alary processes. The eggs are without paler patches. Short, narrow fringed, half-webbed large and unpigmented, or alternatively: fingers and webbed toes. The finger lengths are as 2/ Medium to large species wlth long and variously follows 3>4>2>1. The webbing between the third and webbed fingers, long hindlimbs, pigmented or fourth fingers extends to a point slightly below the paired unpigmented bones and very highly variable dorsal subarticular tubercles at the base of the penultimate coloration. The intercalary structures are small and phalanx on the fourth toe. Broadly spaced nares (E-N/IN cartilaginous. The hyoid plate bears pedunculate alary 0.657-0.758). No vomerine teeth. The head is slightly processes. The ova are small and pigmented. longer than broad (HL/HW 1.031), its length equivalent to Frogs in the subgenus Alliuma subgen. nov. are more than one-third of the snout to vent length (HL/S-V separated from all other Australasian (Australian and 0.356). The snout is not prominent; abrupt and truncate New Guinea) Tree Frogs (Pelodryadidae) by the following when viewed from above and very slightly rounded in suite of characters: They are medium-sized montane tree profile. The nostrils are lateral, their distance from the frogs with spiniform tubercles on the hindlimbs, but no end of the snout slightly less than that from the eye. The spiniform tubercles on the body, a green and brown distance between the eye and the naris is less than the blotched dorsum, and yellow colouration on the hidden internarial span (E-N/IN 0.694). The canthus rostralis is surfaces of the thighs.The ventral surface is variegated well defined and very slightly curved. The eye is large with dark pigments. Moderate to extensive finger and conspicuous, its diameter greater than the eye to webbing. Vocal slits present in males; strongly curved naris distance. canthus rostralis. The tympanum is covered with skin and very small, its Frogs in the subgenus Naveosrana subgen. nov. are diameter equivalent to one-third of the eye diameter, separated from all other Australasian (Australian and separated from the eye by a distance greater than its own New Guinea) Tree Frogs (Pelodryadidae) by the following diameter. The tongue is broadly cordiform with weakly suite of characters: A small (under 50 mm snout-vent in indented posterior border. both sexes) grey or brown species of frog with broad half- The terminal discs are prominent. Long and slender webbed fingers bearing large discs, and having a hindlimbs with a TL/S-V ratio of 0.584. strongly curved canthus rostralis. The intercalary Toe lengths 4>5=3>2>1. The webbing of all toes except structures are elongate and cartilaginous. The hyoid plate for the fourth reaches the base of the discs, while on the lacks alary processes. Ova are small and unpigmented. fourth toes the webbing reaches the subarticular tubercle Further diagnostic characters of this subgenus are as at the base of the penultimate phalanx and continues to follows: The dorsum varies from grey to dark brown with the disc via a narrow fringe. The dorsal and lateral pale markings including light pigment over the site of the surfaces of the body are finely pitted and striated. There cutaneous blood vessels. The snout is gently rounded is an inconspicuous supratympanic fold. The throat and and the canthus rostralis sharply curved. The fingers chest are smooth. Abdomen and nearby halves of the have broad discs and are approximately half webbed. ventral surface of the thighs are coarsely granular. There The toes are webbed to the base of the discs. Tubercles is a small pigmented nuptial pad at the base of the first occur commonly on the upper eyelids, below the anus, finger. Vocal sac openings are exceptionally long, on the back of the thighs and on the heels. extending from the base of the tongue to the angles of The cranial elements are reduced. The nasals are small the jaw. and very widely separated medially. They do not Ventrally the frogs are a pale creamish colour, stippled articulate with the sphenethmoid, which appears to be with dark brown on the throat. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 70 Australasian Journal of Herpetology

Frogs within the subgenus Longuscrusanura subgen. Etymology: The new genus name Angularanta comes nov. are separated from all other Australasian (Australian from the Latin words meaning sharp snout, or sharp front, and New Guinea) Tree Frogs (Pelodryadidae) by the in reference to the generally pointed snouts of most of following suite of characters: Small size with male these species. maximum length 23 mm, female 28 mm. Fingers are Content: Angularanta arfakiana (Peters and Doria, 1878) slightly fringed and slightly less than half webbed and of (type species); L. mukherjii sp. nov.; A. becki (Loveridge, moderate length. The toes are fully webbed and the 1943); A. brongersmai (Loveridge, 1945); A. bulmeri hindlimbs are particularly long, with a ratio of tibia length (Tyler, 1968); A. chydaeus sp. nov.; A. communia sp. to snout-vent length of 0.59-0.68. nov.; A. dorsivena (Tyler, 1968); A. extentacrus sp. nov.; The dorsum is greyish (in preservative) and marked with A. flavescens (Kraus and Allison, 2004); A. fuscula pale, cryptic markings resembling lichens. (Oliver and Richards, 2007); A. impura (Peters and Doria, The snout is short and very slightly pointed. 1878); A. longicrus (Boulenger, 1911); L. louisiadensis Ovarian eggs are unpigmented. Vomerine teeth are (Tyler, 1968); A. lutea (Boulenger, 1887); A. macki present in some individuals and absent in others. (Richards, 2001); A. milneana (Loveridge, 1945); A. The snout is prominent, slightly projecting and obtusely napaea (Tyler, 1968); A. oenicolen (Menzies and Zweifel, pointed when viewed from above and slightly projecting 1974); A. oxyeei sp. nov.; A. pratti (Boulenger, 1911); A. in profile. The nostrils are more lateral than superior, their quaeinfernas sp. nov.; A. solomonis (Vogt, 1912); A. distance from the end of the snout approximately two spartacus (Richards and Oliver, 2006); A. spinifera (Tyler, thirds that from the eye. The distance from the eye and 1968); A. thesaurensis (Peters, 1877); A. vulgarans sp. the naris is less than the internarial span. The canthus nov.; A. wollastoni (Boulenger, 1914). rostralis is well defined and strongly rounded. The eyes ALLIUMA SUBGEN. NOV. are large and prominent the eye diameter being greater LSIDurn:lsid:zoobank.org:act:C23E103A-0397-4910- than the eye to naris distance. A0FB-30A3216FC26F The tympanum is covered with skin, only the inferior half Type species: Litoria spartacus Richards and Oliver, of the annulus is visible. The tympanum diameter 2006. approximates the equivalent of two fifths of the eye Diagnosis: Frogs in the subgenus Alliuma subgen. nov. diameter, separated from the eye by a distance greater (being a subgenus within the genus Angularanta subgen. than its own diameter. The tongue shape is somewhat nov.) are separated from all other Australasian distorted and roughly cordiform in shape with a very (Australian and New Guinea) Tree Frogs (Pelodryadidae) slightly indented posterior margin. by the following suite of characters: They are medium- The two species Angularanta impura and A. oxyeei sp. sized montane tree frogs with spiniform tubercles on the nov. constitute the entirety of the subgenus Raucus hindlimbs, but no spiniform tubercles on the body, a subgen. nov. and are separated from all other species green and brown blotched dorsum and yellow colouration within Angularanta gen. nov. and all other Australasian on the hidden surfaces of the thighs.The ventral surface (Australian and New Guinea) Tree Frogs (Pelodryadidae) is variegated with dark pigments. Moderate to extensive by the following suite of characters: unpigmented bones; finger webbing. Vocal slits present in males; strongly single subarticular tubercle on fourth finger; reduced toe curved canthus rostralis. webbing; reduced hand webbing (versus moderate to See the detailed description for the genus Angularanta extensive in one or both in all species of Angularanta gen. nov. in this paper for details of the separation of the gen. nov.), narrow lateral fringes on fingers; dark brown other subgenera from all other Australasian (Australian to blackish chin in males; dark brown to reddish brown and New Guinea) Tree Frogs (Pelodryadidae). dorsum; dark brown canthal stripe; upper-lip may be According to Duellman et al. (2016), the subgenus white, with a very thin white line bordered by black or Alliuma subgen. nov. diverged from the nominate brown; concealed surfaces of thighs are brown with subgenus Angularanta subgen. nov. 10.8 MYA. yellow spots; yellow to orange iris, sometimes with a According to Duellman et al. (2016), the genus green upper margin; ventrum white; slightly pointed snout Angularanta gen. nov. diverged from its nearest living when viewed from above, side on or below; a raucous relatives in the divergent genus Bellarana gen. nov. 12.7 call; IN/SV 0.072-0.08, TY/SV 0.067-0.075, HW/SV 0.34- MYA, in turn diverged as a pair from their next nearest 0.3, HL/SV 0.34-0.3, EN/IN 1.1-1.2. living relatives in the two genera Hopviridi gen. nov. and According to Duellman et al. (2016), the genus Ornatanura gen. nov. 13.5 MYA and all diverged 21.5 Angularanta gen. nov. diverged from its nearest living MYA from the most closely related living species in relatives in the divergent genus Bellarana gen. nov. 12.7 previously named genera. MYA, in turn diverged as a pair from their next nearest Distribution: Known only from a few scattered locations living relatives in the two genera Hopviridi gen. nov. and around watercourses at 800-1500 metres in altitude in Ornatanura gen. nov. 13.5 MYA and all diverged 21.5 the central highlands region of central New Guinea, only MYA from the most closely related living species in in association with the main central cordillera. previously named genera. Etymology: Alliuma is derived from the Latin word Distribution: Frogs in this genus are found in all parts of “alium” meaning different, reflecting the flanges on the New Guinea, nearby offshore Islands, including those to frog, giving it a different appearance to a lot of other the north (New Britain, New Ireland), with some species frogs. The name spelling of this genus with the addition also occurring as far afield as most major islands within of the letter “a” is deliberate as it avoids the genus being the Solomon Islands. a homonym with the plant genus Allium Linnaeus, 1753

and the extra “l” is added for similar reasons. subgen. nov.) are separated from all other Australasian Content: Angularanta (Alliuma) spartacus (Richards and (Australian and New Guinea) Tree Frogs (Pelodryadidae) Oliver, 2006) (type species); A. (Alliuma) macki by the following suite of characters: A small (under 50 (Richards, 2001); A. (Alliuma) spinifera (Tyler, 1968). mm snout-vent in both sexes) grey or brown species of LONGUSCRUSANURA SUBGEN. NOV. frog with broad half-webbed fingers bearing large discs LSIDurn:lsid:zoobank.org:act:83CEADF8-E6D4-421E- and having a strongly curved canthus rostralis. The B5E9-7AA0124924B6 intercalary structures are elongate and cartilaginous. The hyoid plate lacks alary processes. Ova are small and Type species: Hyla napaea Tyler, 1968. unpigmented. Further diagnostic characters of this Diagnosis: Frogs within the subgenus Longuscrusanura subgenus are as follows: The dorsum varies from grey to subgen. nov. (being a subgenus within the genus dark brown with pale markings including light pigment Angularanta subgen. nov.) are separated from all other over the site of the cutaneous blood vessels. The snout is Australasian (Australian and New Guinea) Tree Frogs gently rounded and the canthus rostralis sharply curved. (Pelodryadidae) by the following suite of characters: The fingers have broad discs and are approximately half Small size with male maximum length 23 mm, female 28 webbed. The toes are webbed to the base of the discs. mm. Fingers are slightly fringed and slightly less than half Tubercles occur commonly on the upper eyelids, below webbed and of moderate length. the anus, on the back of the thighs and on the heels. The toes are fully webbed and the hindlimbs are The cranial elements are reduced. The nasals are small particularly long, with a ratio of tibia length to snout-vent and very widely separated medially. They do not length of 0.59-0.68. articulate with the sphenethmoid, which appears to be The dorsum is greyish (in preservative) and marked with lobulated anteriorly in retaining a double condition, and pale, cryptic markings resembling lichens. does not extend between the nasals. The frontoparietal The snout is short and very slightly pointed. fontanelle is large and ovoid. The squamosal has a short Ovarian eggs are unpigmented. zygomatic ramus and slightly longer otic ramus. The Vomerine teeth are present in some individuals and quadratojugal is not developed. The pars facialis is absent in others. shallow and the short posterior process does not The snout is prominent, slightly projecting and obtusely articulate with the maxillary process of the nasal. The pointed when viewed from above and slightly projecting alary processes of the premaxillaries are well developed, in profile. The nostrils are more lateral than superior, their bifurcated at their extremities, widely separated medially distance from the end of the snout approximately two and perpendicular to the pars dentalis. The palatine thirds that from the eye. The distance from the eye and processes are well developed and do not articulate with the naris is less than the internarial span. The canthus each other medially. The prevomers are reduced. The rostralis is well defined and strongly rounded. The eyes sacral diapophyses are broadly expanded and the ilia are large and prominent, the eye diameter being greater extend one third along their length. No flange is present than the eye to naris distance. on the third metacarpal and the intercalary structures are long and cartilaginous. Alary processes of the hyoid plate The tympanum is covered with skin, only the inferior half are lacking. The adductor mandibulae externus of the annulus is visible. The tympanum diameter superficialis is absent. approximates the equivalent of two fifths of the eye diameter, separated from the eye by a distance greater See the detailed description for the genus Angularanta than its own diameter. gen. nov. in this paper for details of the separation of the other subgenera from all other Australasian (Australian The tongue shape is somewhat distorted and roughly and New Guinea) Tree Frogs (Pelodryadidae). cordiform in shape with a very slightly indented posterior margin. Distribution: Known only from locations around watercourses in the central highlands region of central See the detailed description for the genus Angularanta New Guinea, only in association with the main central gen. nov. in this paper for details of the separation of the cordillera mainly in the vicinity of the upper Fly and Sepik other subgenera from all other Australasian (Australian Rivers and slightly west of there in the Star Mountains as and New Guinea) Tree Frogs (Pelodryadidae). well as the Darewa River headwaters in West Papua, Distribution: Known only from the type locality in the although there are specimens from the Ramu River, Snow Mountains and from the Wapoga River catchments Eastern Highlands as well. 300 km to the west, New Guinea, Papua, Indonesia. Reports of (putative) species in this genus from the Arfak Etymology: The new subgenus name is taken from Latin Mountains further east are not confirmed, but if true, are and literally means long legged frog, in direct reflection of probably of a different species. the sole species in the subgenus. Etymology: Taken from Latin, the words “naveos” means Content: Angularanta (Longuscrusanura) napaea (Tyler, flecked and “rana” means frog and so the new genus 1968) (monotypic). name “Naveosrana” literally means flecked frog, which is NAVEOSRANA SUBGEN. NOV. the usual colouration of relevant species. LSIDurn:lsid:zoobank.org:act:89B00246-658A-4EAF- Content: Angularanta (Naveosrana) dorsivena (Tyler, AA19-7E09E4284B1C 1968) (type species); A. (Naveosrana) fuscula (Oliver and Type species: Hyla dorsivena Tyler, 1968. Richards, 2007). Diagnosis: Frogs in the subgenus Naveosrana subgen. nov. (being a subgenus within the genus Angularanta

RAUCUS SUBGEN. NOV. A. impura in life is depicted on plate 51 of Menzies LSIDurn:lsid:zoobank.org:act:1DEED3E3-61E6-49B4- (2006). B560-8BB44C0F6D64 A. oxyeei sp. nov. is depicted in life on plate 50 of Type species: Hyla (Litoria) impura Peters and Doria, Menzies (2006). 1878. Distribution: The two species within Raucus subgen. Diagnosis: The two species Angularanta impura and A. nov. are only definitvely known from the south east of oxyeei sp. nov. constitute the entirety of the subgenus New Guinea in the Central and Milne Bay Provinces of Raucus subgen. nov. and are separated from all other Papua New Guinea. species within Angularanta gen. nov. and all other Etymology: The new subgenus name Raucus is taken Australasian (Australian and New Guinea) Tree Frogs directly from the Latin word meaning raucous in English, (Pelodryadidae) by the following suite of characters: meaning “making or constituting a disturbingly harsh and unpigmented bones; single subarticular tubercle on loud noise”, which reflects the nature of the mating call of fourth finger; reduced toe webbing; reduced hand males in both species. webbing (versus moderate to extensive in one or both in Content: Angularanta (Raucus) impura (Peters and all other species of Angularanta gen. nov.), narrow lateral Doria, 1878) (type species); A. (Raucus) oxyeei sp. nov.. fringes on fingers; dark brown to blackish chin in males; SCLERISQUEANURA SUBGEN. NOV. dark brown to reddish brown dorsum; dark brown canthal LSIDurn:lsid:zoobank.org:act:C1D3835F-B3D2-4A6A- stripe; upper-lip may be white, with a very thin white line 857F-DF8C3F9C3A2F bordered by black or brown; concealed surfaces of thighs Type species: Hyla louisiadensis Tyler, 1968. are brown with yellow spots; yellow to orange iris, Diagnosis: Frogs within the subgenus Scelerisqueanura sometimes with a green upper margin; ventrum white; subgen. nov. (being a subgenus within the genus slightly pointed snout when viewed from above, side-on Angularanta subgen. nov.) are separated from all other or below; a raucous call; IN/SV 0.072-0.08, TY/SV 0.067- Australasian (Australian and New Guinea) Tree Frogs 0.075, HW/SV 0.34-0.3, HL/SV 0.34-0.3, EN/IN 1.1-1.2. (Pelodryadidae) by the following suite of characters: Until now Angularanta oxyeei sp. nov.. has been treated Small (adult males 25.8-30.3 mm) frogs. Dorsum is as the far south-eastern population of A. impura (Peters chocolate brown, with or without paler patches. Short, and Doria, 1878) with a type locality of Yule Island, north- narrow fringed, half-webbed fingers and webbed toes. west of Port Moresby, in Central Province, Papua New The finger lengths are as follows 3>4>2>1. The webbing Guinea, being the form of the genus from that area. between the third and fourth fingers extends to a point Angularanta oxyeei sp. nov. is the similar and closely slightly below the paired subarticular tubercles at the related species from the far east of the Milne Bay region base of the penultimate phalanx on the fourth toe. in Papua New Guinea. Broadly spaced nares (E-N/IN 0.657-0.758). No vomerine Angularanta oxyeei sp. nov. is readily separated from A. teeth. The head is slightly longer than broad (HL/HW impura by the following suite of characters: Dorsum 1.031), its length equivalent to more than one-third of the smooth and flanks either smooth or only very slightly snout to vent length (HL/S-V 0.356). The snout is not granular; a chocolate brown dorsum with a significant prominent; abrupt and truncate when viewed from above amount of distinctive and well-defined blackish markings and very slightly rounded in profile. The nostrils are and mottling, including on the upper surfaces of the limbs lateral, their distance from the end of the snout slightly as well as the posterior part of the upper surface of the less than that from the eye. The distance between the head. Anterior to the eyes, the upper surface of the snout eye and the naris is less than the internarial span (E-N/IN is immaculate or near immaculate chocolate brown, but 0.694). the sides of the snout below the eye and anterior to them The canthus rostralis is well defined and very slightly is wholly blackish brown in colour, there being a well curved. The eye is large and conspicuous, its diameter defined border at the upper margin where the colours greater than the eye to naris distance. The tympanum is meet to form a line from tip of snout to top of eye along a covered with skin and very small, its diameter equivalent well-defined ridge. The lower margin at the jawline is in to one-third of the eye diameter, separated from the eye turn bordered by a thin and well defined white line by a distance greater than its own diameter. The tongue running the entire length of the mouth. is broadly cordiform with weakly indented posterior The upper surfaces of the front legs are well marked with border. chocolate brown and blackish blotches tending to form The terminal discs are prominent. Long and slender cross bands. Markings on the upper surfaces of the back hindlimbs with a TL/S-V ratio of 0.584. legs are well defined but do not form any obvious pattern. Toe lengths 4>5=3>2>1. The webbing of all toes except By contrast, A. impura is separated from A. oxyeei sp. for the fourth reaches the base of the discs, while on the nov. by the following suite of characters: slightly granular fourth toes the webbing reaches the subarticular tubercle dorsum, becoming more granular on the flanks; a at the base of the penultimate phalanx and continues to medium brownish dorsum with minimal markings, being the disc via a narrow fringe. The dorsal and lateral indistinct grey mottling or peppering mainly between the surfaces of the body are finely pitted and striated. There eyes and on the lower back. The upper lip and below the is an inconspicuous supratympanic fold. The throat and eye is also mainly brown with indistinct blackish markings chest are smooth. Abdomen and nearby halves of the on the upper surfaces of the limbs. The iris is light ventral surface of the thighs are coarsely granular. orangeish in colour. There is no obvious thin white line There is a small pigmented nuptial pad at the base of the demarcating the upper lip. first finger. Vocal sac openings are exceptionally long,

extending from the base of the tongue to the angles of different call than the type form of A. thesaurensis. the jaw. A. quaeinfernas sp. nov. is separated from the other Ventrally the frogs are a pale creamish colour, stippled three species in that the yellow colouration of the with dark brown on the throat. undersurfaces extends onto the tympanum (versus not so See the detailed description for the genus Angularanta in the other three species) and the posterior part of the gen. nov. in this paper for details of the separation of the middle iris is orange in colour. other subgenera from all other Australasian (Australian Kraus and Allison (2014a) also noted that A. and New Guinea) Tree Frogs (Pelodryadidae). quaeinfernas sp. nov. (identified by them as a potential Distribution: Known only from Sudest (Tagula) and new species) has a different call than the type form of A. Rossell Islands in the Louisiade Islands, in the Milne Bay thesaurensis. Province, south-east of the main landmass of New A. thesaurensis is separated from the other three species Guinea, Papua New Guinea. by its larger adult size with a maximum snout vent of 67 Etymology: The new subgenus name taken from Latin mm in females and 50 mm in males, versus 59 mm in literally means chocolate (brown) coloured frog in females and 45 mm in males in the other three species, reflection of the usual colour of the dorsum. an orange iris and a narrow well-defined ring of greenish- Content: Angularanta (Scelerisqueanura) louisiadensis yellow around the eye. (Tyler, 1968), including one subspecies. A. vulgarans sp. nov. is separated from A. chydaeus sp. ANGULARANTA (ANGULARANTA) CHYDAEUS SP. nov. by having an orange iris, versus yellow with red NOV. peppering or obvious red venation in A. chydaeus sp. LSIDurn:lsid:zoobank.org:act:6B3CE863-7862-41D8- nov.. 9093-ADE406B22687 The flanks of A. vulgarans sp. nov. and A. thesaurensis Holotype: A preserved specimen in the National are yellowish in colour, with no markings other than some Museum of Natural History; Smithsonian Institution; indistinct darker, greyish marbling. In A. chydaeus sp. Washington, DC, USA, specimen number Amphibians nov. and A. quaeinfernas sp. nov. the flanks are also and Reptiles 195538 collected from north of Brown River yellow in colour but punctuated by well defined dark Road, between Laloki River and Brown River, Central brown spots, flecks or peppering. The hind legs of both A. Province, Papua New Guinea, Latitude -9.2916 S., chydaeus sp. nov. and A. quaeinfernas sp. nov. have well Longitude 147.227 E. defined black spots, blotches or marks on them, versus either none or indistinct at best in both A. vulgarans sp. This facility allows access to its holdings. nov. and A. thesaurensis. Paratype: A preserved specimen at the Museum of A. vulgarans sp. nov. and A. thesaurensis have smooth Comparative Zoology, Harvard University in Cambridge, skin above the eyelid, versus a few tiny tubercles above Massachusetts, USA, specimen number MCZ Herp A- the eyelid in A. chydaeus sp. nov. and A. quaeinfernas 90396 collected at 2 km west of Sogeri, 3 km north of sp. nov.. Bisianumu, Central Province, Papua New Guinea, Latitude -9.3968 S., Longitude 147.4016 E. A photo of A. chydaeus sp. nov. in life can be found on plate 53 in Menzies (2006). Diagnosis: The three species A. chydaeus sp. nov. from the Central Province of Papua New Guinea, in the Port The four species A. chydaeus sp. nov., A. quaeinfernas Moresby region, A. quaeinfernas sp. nov. from the West sp. nov., A. vulgarans sp. nov. and A. thesaurensis until Sepik region of Papua New Guinea including the Bewani now treated as the single putative taxon A. thesaurensis Mountains and Angularanta vulgarans sp. nov. from can all be separated from other species within the genus Guadalcanal, Solomon Islands, have all until now been Angularanta gen. nov. by the following suite of treated as populations of the widespread species A. characters: Reduced webbing of the fingers and green thesaurensis (Peters, 1877), originally described as “Hyla bones. Adult maximum snout vent of 67 mm in females thesaurensis Peters, 1877” with a type locality of and 50 mm in males (smaller in all of the three species Treasury Island, Solomon Islands. formally named herein). The four species are separated from one another as The head is flattened and longer than broad (HL/HW follows: 1.023-1.200), its length greater than one-third of the snout to vent length (HL/S-V 0.340-0.373). The snout is Both A. vulgarans sp. nov. and A. chydaeus sp. nov. are prominent, when viewed from above and in profile it is separated from A. quaeinfernas sp. nov. and A. slightly rounded. thesaurensis by their shorter legs with a TL/S-V of less than 0.55, versus greater than this in the latter two The nostrils are more lateral than superior, their distance species. from the end of the snout approximately half that from the eye. The distance between the eye and the naris is very A. vulgarans sp. nov. is separated from the other three much greater than the internarial span (E-N/IN 1.257- species by the chin having a distinctive mottlling with 1.485). brown, which in effect also separates A. chydaeus sp. nov. from the other three species in that it, A. chydaeus The canthus rostralis is curved and slightly defined. The sp. nov., lacks distinctive brown mottling on the chin and eye is large and prominent, its diameter less than, equal also has shorter legs than both A. quaeinfernas sp. nov. to, or greater than the distance separating it from and A. thesaurensis. the nostril. The tympanum is visible, its diameter Kraus and Allison (2014a) noted that A. chydaeus sp. equivalent to two-thirds of the eye diameter. The nov. (identified by them as a potential new species) has a vomerine teeth are in two large, oblique series between the upper margins of the choanae. The tongue is broadly Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 74 Australasian Journal of Herpetology oval with a slightly indented posterior border. ANGULARANTA (ANGULARANTA) COMMUNIA SP. The fingers are long and slender with large terminal discs NOV. and very narrow lateral fringes, in decreasing order of LSIDurn:lsid:zoobank.org:act:830EEE3B-4E23-4A5E- length 3>4>2>1. The webbing between the fingers is A2C1-7D0F8B769684 scant, just reaching the sub-articular tubercle at the base Holotype: A preserved specimen at the Australian of the penultimate phalanx of the fourth. Museum, Sydney, New South Wales, Australia, specimen The length of the hind limbs is highly variable, differing number R.137991 collected at Param Village, Milne Bay markedly between species and within species. The range Province, Papua New Guinea, Latitude -9.966 S., of TL/S-V is from 0.528-0.629. Toes in decreasing order Longitude 149.483 E. of length 4>5 =3>2>1. The webbing reaches the base of This government-owned facility allows access to its the disc on the fifth toe and the sub-articular tubercle at holdings. the base of the penultimate phalanx on the fourth. Paratype: A preserved specimen at the Australian The skin on the dorsal surface is smooth. In some Museum, Sydney, New South Wales, Australia, specimen specimens the upper eyelids are slightly granular. A row number R.137945 collected at the Munimum Village, of tubercles on the posterior surface of the forearm vary Agaun area, Milne Bay Province, Papua New Guinea, from well developed to being scarcely detectable. Latitude -9.883 S., Longitude 149.383 E. There is a very weak supra-tympanic fold extending from Diagnosis: Angularanta communia sp. nov. has until now the posterior corner of the eye to a point above the been treated as a far south-eastern population of insertion of the forearm. A very small portion of the Angularanta arfakiana (Peters and Doria, 1878) (type superior margin of the tympanic annulus is usually hidden species for the genus Angularanta gen. nov.). beneath the supra-tympanic fold. There are a few They are however readily separated on the basis of flattened tubercles on the throat. The chest is smooth colouration. and the abdomen and lower surfaces of the thighs Adult A. arfakiana have an olive grey or olive brown coarsely granular. dorsum, vaguely mottled with or without an hour-glass The male possesses a sub-gular vocal sac and an mak. Concealed parts of thighs are yellowish-purplish exceptionally elongate nuptial pad on the first finger. spotted yellow. The dorsum merges into a plain whitish The ground colouration of the dorsal surfaces is usually ventrum and the iris is speckled fawn in colour. Scattered pale grey, buff, brown or cream with or without lighter or tubercles on upper hind leg are small. Tubercles above darker markings. the eye are tiny. The lighter markings sometimes consist of three pale By contrast Adult A. communia sp. nov. are light brown yellow longitudinal stripes situated mid-dorsally and with an hour-glass mark. Snout is lighter or with a pale dorso-laterally. The darker markings are commonly in the bar between the eyelids. Lores are dark, the colour form of irregularly shaped brown or black spots. A white extending to include the tympanum. Flanks are speckled (in A. chydaeus sp. nov.) or yellow (in A. vulgarans sp. dark and light brown and green. Concealed surfaces of nov., A. thesaurensis, and A. quaeinfernas sp. nov.) bar the thighs are dark brown, speckled white. Ventrum is is present above the upper lip directly beneath the eye of white and spotted with grey-brown. Scattered tubercles most specimens. The backs of the thighs are brown with on upper hind leg are medium to large. Tubercles above or without numerous, very small, circular white or cream eye obvious. spots. The white spots on the backs of the thighs being Both A. communia sp. nov. and A. arfakiana are readily most common in the two Solomon Islands species. The separated from all other species in the genus bones are green. Pigmentation is not confined to the Angularanta gen. nov. by the following suite of periosteum but is present throughout the bone and characters: A prominent pointed snout with a sharp persists in specimens preserved in formalin for at least canthus rostralis, and a conspicuous, straight supra- thirty years (according to Tyler, 1978). tympanic fold. Large sub-anal tubercles are exhibited by In the two Solomon Islands species the three pale to all specimens and conical heel tubercles are usually bright yellow longitudinal stripes, found on most if not all present, ranging in size from tiny to large, depending on specimens, are united by a narrow trans-ocular bar. species. The S-V length of adult males varies from See the detailed description for the genus Angularanta approximately 35 mm to 45 mm and from 45 mm to 65 gen. nov. in this paper for details of the separation of the mm in females. subgenus Angularanta subgen. nov. and other subgenera The head is flattened, being longer than broad (HL/HW within Angularanta gen. nov. from all other Australasian 1.077-1.152), its length slightly more than one-third of the (Australian and New Guinea) Tree Frogs snout to vent length (HL/S-V 0.340-0.373). The snout is (Pelodryadidae). projecting; when viewed from above its shape may be Distribution: Angularanta chydaeus sp. nov. is belived to acutely pointed (males) or obtusely pointed (most large be restricted to the region generally around the Central females); in profile the snout is acutely pointed in both Province of Papua New Guinea including the outskirts of sexes. The nostrils are lateral, their distance from the tip the PNG capital Port Moresby. of the snout slightly less than that from the eye. The Etymology: The new species name “chydaeus” is taken distance between the eye and the naris is less than the from the same Latin word meaning common or ordinary, internarial span (E-N/IN 0.600-0.760). The canthus as this species is both common where found and rostralis is very slightly curved and extremely sharply ordinary in general appearance. As a frog species there defined. The loreal region is concave and oblique. The is nothing particularly unusual about its appearance. eye is large, its diameter greater than the distance Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 75

separating it from the nostril. The tympanum is visible, its This government owned facility allows access to its diameter equivalent to one-third to one-half of the eye holdings. diameter. The vomerine teeth are in two, short oblique Paratypes: Two preserved specimens at the South series situated between the rounded choanae. The Australian Museum, Adelaide, South Australia, Australia, tongue is cordiform with a slightly indented posterior specimen numbers R9154 and R9429, collected from border. The fingers are long and slender and lack lateral “Camp 2” at Pio River, Gulf Province, Papua New fringes; in decreasing order of length 3>4>2>1; Guinea. unwebbed or with a trace of webbing at the base of the Diagnosis: Until now Angularanta extentacrus sp. nov. third and fourth fingers. The terminal discs are extremely from the central highlands of New Guinea has been prominent. The hind limbs are slender and there is treated as an eastern population of A. longicrus considerable variation in the TL/S-V ratio (range 0.554- (Boulenger, 1911) with syntypes type localities of Fakfak 0.712, mean 0.645). The extreme individuals in most and Wendessi, both on the Vogelkop Peninsula (Bird’s samples differ by approximately 10 percent. Toes in Head), Indonesian New Guinea. decreasing order of length 4>5>3>2>1. The webbing A. extentacrus sp. nov. is however readily separated from between the outer and fourth toes extends to the sub- A. longicrus by having pigment on the dorsal surface of articular tubercle at the base of the penultimate phalanx the thigh (versus unpigmented) and in lacking the white or to approximately half way up the penultimate phalanx bar below the eye (versus a short white bar below the eye on the fifth, whilst the penultimate phalanx of the fourth extending posteriorly to the angle of the jaws in A. may be free or webbed to slightly above the sub-articular longicrus). tubercle. The skin on the dorsal surfaces is roughened. Both A. extentacrus sp. nov. and A. longicrus are readily Conical tubercles are distributed as follows: on the upper separated from all other species in the genus eyelid (usually confined to the posterior half), where they Angularanta gen. nov. by the following unique suite of may be tiny to large, depending on species and characters: Exceptionally long hind limbs (TL/S-V 0.623- individual, and on the heel and beneath the vent. Throat 0.628), widely spaced nares (E-N/IN 0.579-0.616) and lightly granular, abdomen and lower femur coarsely uniform dorsal colouration. In more detail, adult snout to granular. There is a conspicuous, straight supra-tympanic vent lengths are approximately 31.8 mm (female), 27.4 fold extending from the eye to the shoulder, an irregular, mm (male). The head is flattened and as long as broad, slightly raised tubercular ridge above the vent and rows its length equivalent to slightly less than or more than of small tubercles on the posterior surface of the forearm one-third of the snout to vent length (HL/S-V 0.321- and the tarsus. The canthus rostralis and the skin fold 0.358). The snout is not prominent; rounded when viewed behind the eye are usually darker than the surrounding from above and in profile. The nostrils are more lateral areas and may be surrounded by a narrow white line. than superior, their distance from the end of the snout The pre-anal fold is white and the backs of the thighs are about one-half that from the eye. The distance between red or dull brown. Males possess a sub-gular vocal sac. the eye and the naris is less than the internarial span (E- See the detailed description for the genus Angularanta N/IN 0.579-0.616). The canthus rostralis is slightly gen. nov. in this paper for details of the separation of the defined and gently curved. The eye is large and subgenus Angularanta subgen. nov. and other subgenera conspicuous, its diameter equivalent to one and one-half within Angularanta gen. nov. from all other Australasian the eye to naris distance. The tympanum is visible, its (Australian and New Guinea) Tree Frogs diameter equivalent to slightly less than one-half the eye (Pelodryadidae). diameter. The vomerine teeth are in two very small, Distribution: A. communia sp. nov. is currently known circular series on a line directly between the choanae. only from the vicinity of the type locality and the Owen The tongue is oval with a weakly indented posterior Stanley Ranges in south-east New Guinea. A. arfakiana margin. The fingers are short and equipped with narrow as a species complex, including those species lateral fringes; in decreasing order of length 3>4>2>1. resurrected from synonymy in this paper are found in The webbing between the third and fourth fingers most other parts of New Guinea from the Central reaches the sub-articular tubercle at the base of the Highlands and west to include the Bird’s Head region of penultimate phalanx on the fourth. Irian Jaya, Indonesia. The hind limbs are extremely long with a TL/S-V ratio of Etymology: The new species name “communia” comes 0.623-0.628. Toes in decreasing order of length from the Latin word communia which means common, 4>5>3>2>1. The webbng of all toes except the fourth general, ordinary, universal, public, usual, in reflection of reaches the base of the discs. On the fourth toe the the fact that these frogs are common and “ordinary” to webbing reaches the sub-articular tubercle at the base of people where they naturally occur. the penultimate phalanx. The dorsal and lateral surfaces ANGULARANTA (ANGULARANTA) EXTENTACRUS of the body are smooth. There is an inconspicuous, SP. NOV. flattened supra-tympanic fold. The throat and chest are LSIDurn:lsid:zoobank.org:act:6102BFFE-7AAA-4114- smooth, and the abdomen and lower surface of the A300-A848EC162992 thighs are granular. Holotype: A preserved specimen at the South Australian In life, the dorsal surface is green and unmarked. In Museum, Adelaide, South Australia, Australia, specimen preservative the dorsal surface of the body is dull blue number R6482 collected at Kawolabib in the Star and unmarked. The limbs with the exception of the Mountains, Papua New Guinea. thighs, are a similar colour. The thighs are unpigmented. In A. longicrus there is a short white bar below the eye

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 76 Australasian Journal of Herpetology extending posteriorly to the angle of the jaws, but this is between the eye and the naris is very much less than the absent in A. extentacrus sp. nov.. The ventral surfaces of internarial span (E-N/IN 0.605-0.641). The canthus the body and limbs are white to cream. rostralis is slightly defined and gently rounded. The eye is See the detailed description for the genus Angularanta not prominent, its diameter is considerably greater than gen. nov. in this paper for details of the separation of the the eye to naris distance. The tympanum is covered with subgenus Angularanta subgen. nov. and other subgenera skin and the annulus only slightly defined. The tympanum within Angularanta gen. nov. from all other Australasian diameter is equivalent to one-third of the eye diameter; (Australian and New Guinea) Tree Frogs separated from the eye by a distance equivalent to half (Pelodryadidae). its own diameter. Vomerine teeth may or may not be Distribution: Angularanta extentacrus sp. nov. occurs in present, depending on the individual frog. When present the central highlands of New Guinea. A. longicrus they are small and round and situated near the upper (Boulenger, 1911) is apparently confined to the Vogelkop margin of the choana. The choanae are very widely Peninsula (Bird’s Head), Indonesian New Guinea. spaced. The tongue is small and broadly cordiform with a Etymology: The new species name is taken from the slightly indented posterior border. The fingers are very Latin words “extent” meaning stretched and “crus” long and slender with narrow lateral fringes, in meaning leg, thereby meaning stretched leg, in reflection decreasing order of length 3>4>2>1. There is no webbing of the relative length of the leg in this species. between the fingers. The terminal discs are prominent. The hind limbs are long and slender (TL/S-V 0.590- ANGULARANTA (ANGULARANTA) MUKHERJII SP. 0.665). Toes in decreasing order of length 4>5=3>2>1. NOV. The webbing between the toes reaches the base of the LSIDurn:lsid:zoobank.org:act:C669E69D-8BC3-4145- disc on the fifth toe and slightly below the sub-articular 861E-57CD08DE4CA7 tubercle at the base of the penultimate phalanx on the Holotype: A preserved specimen at the South Australian fourth toe. There is a small depressed inner metatarsal Museum, Adelaide, South Australia, Australia, specimen tubercle. The dorsal surface of the head and body is number R70092 collected from Townsville Drill Station at smooth. There is a very weak supra-tympanic fold the headwaters of the Ok Tedi River, Western Province, extending from the posterior corner of the eye to above Papua New Guinea, Latitude -5.15972 S., Longitude the insertion of the forearm. Above and at the sides of the 141.1666 E. anus is a row of tubercles. The throat and chest are This government-owned facility allows access to its smooth, and the abdomen and lower surface of the holdings. thighs coarsely granular. Diagnosis: Until now Angularanta mukherjii sp. nov. from There is a nuptial pad on the first finger. the Fly River basin has been regarded as a southern There is a short black, horizontal line beneath the eye population of Angularanta bulmeri (Tyler, 1968), with a and a smaller marking beneath it on the margin of the type locality of upper Aunjung Valley, Schrader upper jaw. The anterior and posterior surfaces of the Mountains, Eastern Mountains, Madang Province, Papua thigh are pale yellow and the anal region pale brown. The New Guinea. throat is white, finely stippled with palegreen or otherwise A. mukherjii sp. nov. is separated from A. bulmeri by a simply pale green or grey. The chest and lower surfaces lack of a well-defined black lateral stripe extending from of the thighs are dull yellow or green and the abdomen the naris to a point at least midway along the body, cream. On the abdomen and lower surfaces of the thighs instead the stripe becoming indistinct on the body in A. are small indistinct spots of pale brown. The plantar bulmeri as well as numerous small tubercles on the surface is a very dark brown and there is a very fine white upper surface of the hind legs, versus relatively few in A. line at the margin of the plantar surface and the lateral bulmeri. greenish grey colouration. The proximal half of the A. mukherjii sp. nov. and A. bulmeri are readily separated posterior surface of the thigh is usually dull greyish- from all other species in the subgenus Angularanta greenish-brown. subgen. nov. by the following suite of characters: small See the detailed description for the genus Angularanta size (males 29-34 mm), broadly spaced nares (E-N/IN gen. nov. in this paper for details of the separation of the 0.605-0.641) and distinctive colouration. The dorsal subgenus Angularanta subgen. nov. and other subgenera surface is a dark greenish grey, being somewhat mottled within Angularanta gen. nov. from all other Australasian and with white spots or flecks (often on tubercles). There (Australian and New Guinea) Tree Frogs is a broad, black lateral stripe extending from the naris (Pelodryadidae). through the tympanum and posterior to this to a point at Distribution: The species A. mukherjii sp. nov. is only least midway along the body in A. bulmeri or merging known from the type locality. A. bulmeri is found in with the body colour past the ear in A. mukherjii sp. nov.. uplands of headwaters draining north of the main The head is high and the head is consistently longer than cordillera. broad with a range 1.030-1.160, and the head length less Etymology: The new species name “mukherjii” is named than or greater than one-third of the snout to vent length, in honour of Melbourne, Victoria, Australia barrister its length equivalent to slightly more than one-third of the Gautam Mukherji, in recognition for his services to wildlife snout to vent length (HL/S-V 0.366). The snout is abrupt, conservation, defending Raymond Hoser against broadly rounded when viewed from above, projecting and unlawful attacks by trademark infringing thieves and almost pointed in profile. The nostrils are more lateral others working to attack the wildlife conservation cause than superior, their distance from the end of the snout for their own financial self-gratification. less than one-half that from the eye. The distance

ANGULARANTA (ANGULARANTA) QUAEINFERNAS yellow in colour but punctuated by well defined dark SP. NOV. brown spots, flecks or peppering. The hind legs of both A. LSIDurn:lsid:zoobank.org:act:474462DF-1FA2-4723- chydaeus sp. nov. and A. quaeinfernas sp. nov. have well ABDC-AB4FC2D3CD5F defined black spots, blotches or marks on them, versus Holotype: A preserved specimen at the Australian either none or indistinct at best in both A. vulgarans sp. Museum, Sydney, New South Wales, Australia, specimen nov. and A. thesaurensis. number R.135574 collected at Imonda Village, West A. vulgarans sp. nov. and A. thesaurensis have smooth Sepik Province, Papua New Guinea, Latitude 3.20 S., skin above the eyelid, versus a few tiny tubercles above Longitude 141.10 E. the eyelid in A. chydaeus sp. nov. and A. quaeinfernas This government-owned facility allows access to its sp. nov.. holdings. The four species A. chydaeus sp. nov., A. quaeinfernas Diagnosis: The three species A. quaeinfernas sp. nov. sp. nov., A. vulgarans sp. nov. and A. thesaurensis until from the West Sepik region of Papua New Guinea now treated as the single putative taxon A. thesaurensis including the Bewani Mountains, Angularanta vulgarans can all be separated from other species within the genus sp. nov. from Guadalcanal, Solomon Islands and A. Angularanta gen. nov. by the following suite of chydaeus sp. nov. from the Central Province of Papua characters: Reduced webbing of the fingers and green New Guinea, in the Port Moresby region, have all until bones. Adult maximum snout vent of 67 mm in females now been treated as populations of the widespread and 50 mm in males (smaller in all of the three species species A. thesaurensis (Peters, 1877), originally formally named herein). The head is flattened and longer described as “Hyla thesaurensis Peters, 1877” with a than broad (HL/HW 1.023-1.200), its length greater than type locality of Treasury Island, Solomon Islands. one-third of the snout to vent length (HL/S-V 0.340- The four species are separated from one another as 0.373). The snout is prominent, when viewed from above follows: and in profile it is slightly rounded. The nostrils are more lateral than superior, their distance from the end of the Both A. vulgarans sp. nov. and A. chydaeus sp. nov. are snout approximately half that from the eye. The distance separated from A. quaeinfernas sp. nov. and A. between the eye and the naris is very much greater than thesaurensis by their shorter legs with a TL/S-V of less the internarial span (E-N/IN 1.257-1.485). The canthus than 0.55, versus greater than this in the latter two rostralis is curved and slightly defined. The eye is large species. and prominent, its diameter less than, equal to, or greater A. vulgarans sp. nov. is separated from the other three than the distance separating it from the nostril. The species by the chin having a distinctive mottlling with tympanum is visible, its diameter equivalent to two-thirds brown, which in effect also separates A. chydaeus sp. of the eye diameter. The vomerine teeth are in two large, nov. from the other three species in that it, A. chydaeus oblique series between the upper margins of the sp. nov., lacks distinctive brown mottling on the chin and choanae. The tongue is broadly oval with a slightly also has shorter legs than both A. quaeinfernas sp. nov. indented posterior border. and A. thesaurensis. The fingers are long and slender with large terminal discs Kraus and Allison (2014a) noted that A. chydaeus sp. and very narrow lateral fringes. In decreasing order of nov. (identified by them as a potential new species) has a length 3>4>2>1. The webbing between the fingers is different call than the type form of A. thesaurensis. scant, just reaching the sub-articular tubercle at the base A. quaeinfernas sp. nov. is separated from the other of the penultimate phalanx of the fourth. three species in that the yellow colouration of the The length of the hind limbs is highly variable, differing undersurfaces extends onto the tympanum (versus not so markedly between species and within species. The range in the other three species) and the posterior part of the of TL/S-V is from 0.528-0.629. Toes in decreasing order middle iris is orange in colour. of length 4>5 =3>2>1. The webbing reaches the base of Kraus and Allison (2014a) also noted that A. the disc on the fifth toe and the sub-articular tubercle at quaeinfernas sp. nov. (identified by them as a potential the base of the penultimate phalanx on the fourth. new species) has a different call than the type form of A. The skin on the dorsal surface is smooth. In some thesaurensis. specimens the upper eyelids are slightly granular. A row A. thesaurensis is separated from the other three species of tubercles on the posterior surface of the forearm vary by its larger adult size with a maximum snout vent of 67 from well developed to being scarcely detectable. mm in females and 50 mm in males, versus 59 mm in There is a very weak supra-tympanic fold extending from females and 45 mm in males in the other three species, the posterior corner of the eye to a point above the an orange iris and a narrow well-defined ring of greenish- insertion of the forearm. A very small portion of the yellow around the eye. superior margin of the tympanic annulus is usually hidden A. vulgarans sp. nov. is separated from A. chydaeus sp. beneath the supra-tympanic fold. There are a few nov. by having an orange iris, versus yellow with red flattened tubercles on the throat. The chest is smooth peppering or obvious red venation in A. chydaeus sp. and the abdomen and lower surfaces of the thighs nov.. coarsely granular. The flanks of A. vulgarans sp. nov. and A. thesaurensis The male possesses a sub-gular vocal sac and an are yellowish in colour, with no markings other than some exceptionally elongate nuptial pad on the first finger. indistinct darker, greyish marbling. In A. chydaeus sp. The ground colouration of the dorsal surfaces is usually nov. and A. quaeinfernas sp. nov. the flanks are also pale grey, buff, brown or cream with or without lighter or

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 78 Australasian Journal of Herpetology darker markings. Solomon Islands. The lighter markings sometimes consist of three pale Diagnosis: The three species Angularanta vulgarans sp. yellow longitudinal stripes situated mid-dorsally and nov. from Guadalcanal, Solomon Islands, A. dorso-laterally. The darker markings are commonly in the quaeinfernas sp. nov. from the West Sepik region of form of irregularly shaped brown or black spots. A white Papua New Guinea including the Bewani Mountains and (in A. chydaeus sp. nov.) or yellow (in A. vulgarans sp. A. chydaeus sp. nov. from the Central Province of Papua nov., A. thesaurensis, and A. quaeinfernas sp. nov.) bar New Guinea, in the Port Moresby region, have all until is present above the upper lip directly beneath the eye of now been treated as populations of the widespread most specimens. The backs of the thighs are brown with species A. thesaurensis (Peters, 1877), originally or without numerous, very small, circular white or cream described as “Hyla thesaurensis Peters, 1877” with a spots. The white spots on the backs of the thighs being type locality of Treasury Island, Solomon Islands. most common in the two Solomon Islands species. The The four species are separated from one another as bones are green. Pigmentation is not confined to the follows: periosteum but is present throughout the bone and Both A. vulgarans sp. nov. and A. chydaeus sp. nov. are persists in specimens preserved in formalin for at least separated from A. quaeinfernas sp. nov. and A. thirty years (according to Tyler, 1978). thesaurensis by their shorter legs with a TL/S-V of less In the two Solomon Islands species the three pale to than 0.55, versus greater than this in the latter two bright yellow longitudinal stripes, found on most if not all species. specimens, are united by a narrow trans-ocular bar. A. vulgarans sp. nov. is separated from the other three See the detailed description for the genus Angularanta species by the chin having a distinctive mottlling with gen. nov. in this paper for details of the separation of the brown, which in effect also separates A. chydaeus sp. subgenus Angularanta subgen. nov. and other subgenera nov. from the other three species in that it, A. chydaeus within Angularanta gen. nov. from all other Australasian sp. nov., lacks distinctive brown mottling on the chin and (Australian and New Guinea) Tree Frogs also has shorter legs than both A. quaeinfernas sp. nov. (Pelodryadidae). and A. thesaurensis. Distribution: Angularanta quaeinfernas sp. nov. is Kraus and Allison (2014a) noted that A. chydaeus sp. belived to be restricted to northern Papua New Guinea nov. (identified by them as a potential new species) has a and nearby Irian Jaya, including the Bewani Mountains, different call than the type form of A. thesaurensis. in areas generally proximate to the type locality in the A. quaeinfernas sp. nov. is separated from the other West Sepik Province and west of the Huon Peninsula. three species in that the yellow colouration of the Etymology: The new species name “quaeinfernas” is undersurfaces extends onto the tympanum (versus not so taken from the Latin words meaning lowland dwelling, in the other three species) and the posterior part of the which is apt, as this species, while being found in some middle iris is orange in colour. hilly locations is not found in the higher altitude regions. Kraus and Allison (2014a) also noted that A. ANGULARANTA (ANGULARANTA) VULGARANS SP. quaeinfernas sp. nov. (identified by them as a potential NOV. new species) has a different call than the type form of A. LSIDurn:lsid:zoobank.org:act:9C4268DD-45F0-4B3D- thesaurensis. A251-CD423D9088F3 A. thesaurensis is separated from the other three species Holotype: A preserved specimen at the Australian by its larger adult size with a maximum snout vent of 67 Museum, Sydney, New South Wales, Australia, specimen mm in females and 50 mm in males, versus 59 mm in number R.90277 collected at Mount Austin, Guadalcanal, females and 45 mm in males in the other three species, Solomon Islands, Latitude -9.29 S., Longitude 160.00 E. an orange iris and a narrow well-defined ring of greenish- This government owned facility allows access to its yellow around the eye. holdings. A. vulgarans sp. nov. is separated from A. chydaeus sp. Paratypes: 1/ A preserved specimen at the Australian nov. by having an orange iris, versus yellow with red Museum, Sydney, New South Wales, Australia, specimen peppering or obvious red venation in A. chydaeus sp. number R.136387 collected from Mount Austen, nov.. Guadalcanal, Solomon Islands, Latitude -9.29 S., The flanks of A. vulgarans sp. nov. and A. thesaurensis Longitude 159.58 E. are yellowish in colour, with no markings other than some 2/ A preserved specimen at the Australian Museum, indistinct darker, greyish marbling. In A. chydaeus sp. Sydney, New South Wales, Australia, specimen number nov. and A. quaeinfernas sp. nov. the flanks are also R.137027.001 collected from Makarakomburu (South yellow in colour but punctuated by well defined dark Slope), Guadalcanal, Solomon Islands, Latitude -9.45 S., brown spots, flecks or peppering. Longitude 160.00 E. The hind legs of both A. chydaeus sp. nov. and A. 3/ Five preserved specimens at the American Museum of quaeinfernas sp. nov. have well defined black spots, Natural History, Manhattan, New York City, USA, blotches or marks on them, versus either none or specimen numbers AMNH 52173 and 52176-79 collected indistinct at best in both A. vulgarans sp. nov. and A. from Guadalcanal, Solomon Islands. thesaurensis. 4/ Two preserved specimens at the California Academy A. vulgarans sp. nov. and A. thesaurensis have smooth of Sciences, San Francisco, California, USA, specimen skin above the eyelid, versus a few tiny tubercles above numbers CAS. 49952-3 collected from Guadalcanal, the eyelid in A. chydaeus sp. nov. and A. quaeinfernas sp. nov.. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 79

The four species A. chydaeus sp. nov., A. quaeinfernas spots. The white spots on the backs of the thighs being sp. nov., A. vulgarans sp. nov. and A. thesaurensis until most common in the two Solomon Islands species. The now treated as the single putative taxon A. thesaurensis bones are green. Pigmentation is not confined to the can all be separated from other species within the genus periosteum but is present throughout the bone and Angularanta gen. nov. by the following suite of persists in specimens preserved in formalin for at least characters: Reduced webbing of the fingers and green thirty years (according to Tyler, 1978). bones. Adult maximum snout vent of 67 mm in females In the two Solomon Islands species the three pale to and 50 mm in males (smaller in all of the three species bright yellow longitudinal stripes, found on most if not all formally named herein). The head is flattened and longer specimens, are united by a narrow trans-ocular bar. than broad (HL/HW 1.023-1.200), its length greater than See the detailed description for the genus Angularanta one-third of the snout to vent length (HL/S-V 0.340- gen. nov. in this paper for details of the separation of the 0.373). The snout is prominent, when viewed from above subgenus Angularanta subgen. nov. and other subgenera and in profile it is slightly rounded. The nostrils are more within Angularanta gen. nov. from all other Australasian lateral than superior, their distance from the end of the (Australian and New Guinea) Tree Frogs snout approximately half that from the eye. The distance (Pelodryadidae). between the eye and the naris is very much greater than Distribution: Angularanta vulgarans sp. nov. occurs on the internarial span (E-N/IN 1.257-1.485). The canthus Guadalcanal, Solomon Islands. It is assumed that the rostralis is curved and slightly defined. The eye is large morphologically similar species A. thesaurensis is found and prominent, its diameter less than, equal to, or greater on the other main islands to the north and west, at least than the distance separating it from the nostril. The as far as Bougainville. tympanum is visible, its diameter equivalent to two-thirds Etymology: The new species name vulgarans is taken of the eye diameter. The vomerine teeth are in two large, from the Latin word “vulgarins” meaning common, or oblique series between the upper margins of the ordinary, which is a fair account of this species of frog choanae. The tongue is broadly oval with a slightly among local people where these frogs occur. The name indented posterior border. “vulgarans” is a deliberate misspelling and should not be The fingers are long and slender with large terminal discs amended as the spelling is chosen to avoid a potential and very narrow lateral fringes. In decreasing order of homonym with another species. length 3>4>2>1. The webbing between the fingers is ANGULARANTA (ANGULARANTA) LUTEA scant, just reaching the sub-articular tubercle at the base LEUCOPUNCTATA SUBSP. NOV. of the penultimate phalanx of the fourth. LSIDurn:lsid:zoobank.org:act:79031E4B-473C-4C65- The length of the hind limbs is highly variable, differing BC6E-02238BAC1C59 markedly between species and within species. The range of TL/S-V is from 0.528-0.629. Toes in decreasing order Holotype: A preserved specimen at the of length 4>5 =3>2>1. The webbing reaches the base of Naturhistorisches Museum, Basle, Switzerland, specimen the disc on the fifth toe and the sub-articular tubercle at number NMB 4274 collected from New Georgia, Solomon the base of the penultimate phalanx on the fourth. Islands. The skin on the dorsal surface is smooth. In some This facility allows access to its holdings. specimens the upper eyelids are slightly granular. A row Paratypes: 1/ A preserved specimen at the of tubercles on the posterior surface of the forearm vary Naturhistorisches Museum, Basle, Switzerland, specimen from well developed to being scarcely detectable. number NMB 4275 collected from New Georgia, There is a very weak supra-tympanic fold extending from Solomon Islands, and: the posterior corner of the eye to a point above the 2/ A preserved specimen at the Field Museum of Natural insertion of the forearm. A very small portion of the History, Chicago, Illinois, USA, specimen number FMNH superior margin of the tympanic annulus is usually hidden Amphibians and Reptiles 190313 collected from New beneath the supra-tympanic fold. There are a few Georgia, Solomon Islands. flattened tubercles on the throat. The chest is smooth Diagnosis: The subspecies A. lutea leucopunctata and the abdomen and lower surfaces of the thighs subsp. nov. is separated from the nominate form of A. coarsely granular. lutea lutea (Boulenger, 1887) by having a slightly The male possesses a sub-gular vocal sac and an granular dorsal surface (versus smooth), few if any white exceptionally elongate nuptial pad on the first finger. spots on the head between the eyes or anterior to them The ground colouration of the dorsal surfaces is usually (versus many) and an obvious orangeish upper iris, pale grey, buff, brown or cream with or without lighter or versus not so in A. lutea lutea. darker markings. The taxon A. lutea leucopunctata subsp. nov. has been conservatively formally named as a subspecies in the The lighter markings sometimes consist of three pale absence of molecular evidence. However in light of the yellow longitudinal stripes situated mid-dorsally and biogeograpical barrier separating the New Georgia group dorso-laterally. The darker markings are commonly in the of islands from the other populations of A. lutea in the form of irregularly shaped brown or black spots. A white form of a deep sea barrier present even in the height of (in A. chydaeus sp. nov.) or yellow (in A. vulgarans sp. the most recent glacial maxima, it is likely that the New nov., A. thesaurensis, and A. quaeinfernas sp. nov.) bar Georgia frogs may in fact be divergent at the species is present above the upper lip directly beneath the eye of level. most specimens. The backs of the thighs are brown with or without numerous, very small, circular white or cream The species A. lutea has been treated by some past authors as a synonym of A. thesaurensis (Peters, 1877). Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 80 Australasian Journal of Herpetology

However it is readily separated from that species by Etymology: The new subspecies name “leucopunctata” having a very flattened head; disc of third finger generally is taken from the Latin words meaning “white spots” or larger than the tympanum and discs of fingers not or “white points”, in reflection of the tiny white dots scattered scarcely broader than the subtending digit which is on the dorsum of the species. bordered by a wide flange of skin. By contrast A. ANGULARANTA (RAUCUS) OXYEEI SP. NOV. thesaurensis has a head that is only slightly flattened; LSIDurn:lsid:zoobank.org:act:9105FBA3-99CE-4072- disc of third finger generally smaller than the tympanum 8A81-906E48D8A696 and discs of fingers (except the inner one) much broader Holotype: A preserved specimen at the Bernice P. than the subtending digit which is not bordered by a wide Bishop Museum, Honolulu, Hawaii, USA, specimen flange of skin. number: BPBM 4101 collected from Alotau, Milne Bay A. lutea are separated from all other Australasian Province, Papua New Guinea (PNG), Latitude -10.3157 (Australian and New Guinea) Tree Frogs (Pelodryadidae) S., Longitude 150.4588 E. by the following suite of characters: Head slightly less This facility allows access to its holdings. broad to as broad as long, its breadth about 1/3 the Paratypes: Five preserved specimens at the Bernice P. length from snout to vent; snout round-pointed; eye Bishop Museum, Honolulu, Hawaii, USA, being BPBM moderate, its diameter about 1/3 the breadth of the head; 4102-04 collected from Alotau, Milne Bay Province, tympanum round, its diameter 1/2 or slightly more than Papua New Guinea, Latitude -10.3157 S., Longitude that of the eye; loreal region strongly oblique, concave; 150.4588 E. and BPBM 15194-95 collected from Pini canthus rostralis rounded, rather indistinct; interorbital Range at 300 metres elevation, Milne Bay Province, space broader than upper eyelid; vomerine teeth in two Papua New Guinea, Latitude -10.25 S., Longitude 150.18 transverse patches between the choanae and almost in E. contact medially; tongue broadly oval and but feebly indented at the mid-point of the posterior margin. Diagnosis: Until now Angularanta oxyeei sp. nov.. has been treated as the far south-eastern population of A. Forelimb well developed; finger tips strongly depressed impura (Peters and Doria, 1878) with a type locality of with large discs; discs broader than long (except for the Yule Island, north-west of Port Moresby, in Central inner finger), but scarcely broader than the subtending Province, Papua New Guinea, being the form of the digits as measured to include the flanges of skin on the genus from that area. lateral margins; fingers more extensively webbed than in the sympatric species A. thesaurensis (Peters, 1877), Angularanta oxyeei sp. nov. is the similar and closely which has minimal hand webbing, the third finger in A. related species from the far east of the Milne Bay region lutea being webbed to the subdistal tubercle on the in Papua New Guinea. outside and to a point between this and the basal Angularanta oxyeei sp. nov. is readily separated from A. tubercle on the inside, the second almost to the distal impura by the following suite of characters: Dorsum tubercle on the outside; subarticular tubercles small, low, smooth and flanks either smooth or only very slightly transversely elongate; metacarpal tubercles indistinct; granular; a chocolate brown dorsum with a significant hindlimb long, length of tibia about 3/5 the length from amount of distinctive and well-defined blackish markings snout to vent; discs of toes smaller than those of the and mottling, including on the upper surfaces of the limbs fingers. as well as the posterior part of the upper surface of the head. Anterior to the eyes, the upper surface of the snout Subarticular tubercles small, more strongly protrudent is immaculate or near immaculate chocolate brown, but than those of the hands; inner metatarsal tubercle narrow the sides of the snout below the eye and anterior to them and elliptical, its length less than its distance from the is wholly blackish brown in colour, there being a well basal tubercle of the inner toe; outer absent; toes defined border at the upper margin where the colours webbed almost to the discs except for the fourth; skin meet to form a line from tip of snout to top of eye along a smooth or granular dorsally except for the flat granules of well-defined ridge. the lower surfaces of the thighs and abdomen. The lower margin at the jawline is in turn bordered by a Two penultimate subarticular tubercles on finger four thin and well defined white line running the entire length (versus one in A. thesaurensis and all other species most of the mouth. similar to it, including the three formally named in this paper, formerly treated as populations of A. The upper surfaces of the front legs are well marked with thesaurensis). chocolate brown and blackish blotches tending to form cross bands. Markings on the upper surfaces of the back Colour of the dorsum is uniformly green save for legs are well defined but do not form any obvious pattern. numerous scattered white (usually, or rarely yellow) tiny By contrast, A. impura is separated from A. oxyeei sp. spots on the dorsum. Whiteish or yellow ventrally. nov. by the following suite of characters: slightly granular Concealed areas of limbs yellow. dorsum, becoming more granular on the flanks; a See the detailed description for the genus Angularanta medium brownish dorsum with minimal markings, being gen. nov. in this paper for details of the separation of the indistinct grey mottling or peppering mainly between the subgenus Angularanta subgen. nov. and other subgenera eyes and on the lower back. The upper lip and below the within Angularanta gen. nov. from all other Australasian eye is also mainly brown with indistinct blackish markings (Australian / New Guinea) Tree Frogs (Pelodryadidae). on the upper surfaces of the limbs. The iris is light Distribution: A. lutea leucopunctata subsp. nov. is orangeish in colour. There is no obvious thin white line restricted to the New Georgia group of islands within the demarcating the upper lip. A. impura in life is depicted on Solomon Islands. plate 51 of Menzies (2006).

A. oxyeei sp. nov. is depicted in life on plate 50 of tectonic movements of land and sea beds and factors Menzies (2006). that cannot be determined with precision. The two preceding species are separated from all other Significantly, Weijola et al. (2020) found late Pleistocene species within Angularanta gen. nov. and all other divergence between populations of putative Mangrove Australasian (Australian and New Guinea) Tree Frogs Monitors Euprepiosaurus indicus (Daudin, 1802) from (Pelodryadidae) by the following suite of characters: Rossell Island and Sudest Island. unpigmented bones; single subarticular tubercle on I note that an estuarine dwelling species, E. indicus fourth finger; reduced toe webbing; reduced hand would be far more able to breach any gap between the webbing (versus moderate to extensive in one or both in two nearby islands than a salt water vulnerable species all species of Angularanta gen. nov.), narrow lateral like a tree frog. fringes on fingers; dark brown to blackish chin in males; Those same authors named a species “Varanus bennetti dark brown to reddish brown dorsum; dark brown canthal Weijola et al., 2020” based on a similar late Pleistocene stripe; upper-lip may be white, with a very thin white line divergence between their new putative species and bordered by black or brown; concealed surfaces of thighs populations of the previously described “Varanus are brown with yellow spots; yellow to orange iris, tsukamotoi Kishida, 1929”. sometimes with a green upper margin; ventrum white; Fred Kraus a man well known for science grant rorting slightly pointed snout when viewed from above, side on and for engaging in taxonomic vandalism, this being the or below; a raucous call; IN/SV 0.072-0.08, TY/SV 0.067- recklessly renaming of species previously properly 0.075, HW/SV 0.34-0.3, HL/SV 0.34-0.3, EN/IN 1.1-1.2. described and named by others, did in Kraus (2009) Distribution: Angularanta oxyeei sp. nov. is known only name Toxicocalamus mintoni from Sudest Island as a from the far east of the Milne Bay region in Papua New distinct species from the near identical Toxicocalamus Guinea. holopelturus McDowell, 1969 from the adjacent Rossel Etymology: The new species is named in honour of Island based on a single scalation difference on the head “Oxy”, being the abbreviated name of a now deceased of the single specimen he had on hand. Great Dane dog that protected this author’s scientific This was the frontal fused with the supraoculars, which research facility for 8 years. It is appropriate that a may well have been an unusual trait for the specimen as species is formally named in his honour. His full name opposed to a species specific diagnostic character. was “Oxyuranus”, being the genus name for a group of In light of the two recent preceding cases of species-level large venomous Australian elapid snakes. The spelling of recognition of similarly divergent populations, I view late the species name with the extra two letter “e’s” is Pleistocene divergence being a depth of divergence deliberate and should not be changed. more appropriate for subspecies-level differentiation at ANGULARANTA (SCELERISQUEANURA) the taxonomic level and hence describe A. louisiadensis LOUSIADENSIS BRUNETUS SUBSP. NOV. brunetus subsp. nov. as a new subspecies accordingly. LSIDurn:lsid:zoobank.org:act:2ECB4A36-4D24-4EF0- A. louisiadensis brunetus subsp. nov. is readily separated B7A8-CABAE66B511D from nominate A. louisiadensis louisiadensis by having Holotype: A preserved specimen in the American relatively shorter hindlimbs. Comparative measurements Museum of Natural History, Manhattan, New York City, are TL/S-V range of 0.511-0.575, versus 0.584 in A. USA, specimen number 60069 collected at Mount Riu, louisiadensis louisiadensis. Sudest Island, Louisiadae Archipelago, Milne Bay A. louisiadensis brunetus subsp. nov. has a dark Province, Papua New Guinea. chocolate brown dorsum, lateral surfaces and limbs This facility allows access to its holdings. versus one that is dark, but with obscure, faint irregular Paratypes: Nine preserved specimens in the American patches in A. louisiadensis louisiadensis. Museum of Natural History, Manhattan, New York City, Both subspecies of A. louisiadensis, comprising the USA, specimen numbers 60070-60078 collected at entirety of the subgenus Scelerisqueanura subgen. nov. Mount Riu, Sudest Island, Louisiadae Archipelago, Milne are readily separated from all other Australasian Bay Province, Papua New Guinea. (Australian and New Guinea) Tree Frogs (Pelodryadidae) Diagnosis: The subspecies Angularanta by the following suite of characters: Small (adult males (Scelerisqueanura) louisiadensis brunetus subsp. nov. 25.8-30.3 mm) frogs. Dorsum is chocolate brown, with or from Sudest Island (AKA Tagula Island) has until now without paler patches. Short, narrow fringed, half-webbed been treated as wholly conspecific with A. louisiadensis fingers and webbed toes. The finger lengths are as (Tyler, 1968) from nearby Rossell Island, with the type follows 3>4>2>1. The webbing between the third and material here for A. brunetus subsp. nov. treated as fourth fingers extends to a point slightly below the paired paratypes of A. louisiadensis by Tyler (1968). subarticular tubercles at the base of the penultimate However both populations are consistently phalanx on the fourth toe. Broadly spaced nares (E-N/IN morphologically divergent, are clearly evolving separately 0.657-0.758). No vomerine teeth. and can be reasonably expected to have diverged most The head is slightly longer than broad (HL/HW 1.031), its recently at the end of the Pleistocene (11.7 K years ago). length equivalent to more than one-third of the snout to A check of sea level depths (to the 120 m level), shows a vent length (HL/S-V 0.356). The snout is not prominent; it likely narrow connection between Sudest Island and is abrupt and truncate when viewed from above and very Rossell Island, or alternatively a gap of less than 1 km. slightly rounded in profile. The nostrils are lateral, their The ambiguity in determination relates to positions of distance from the end of the snout slightly less than that fringing reefs, sediment accumulation or removal, from the eye. The distance between the eye and the naris Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 82 Australasian Journal of Herpetology is less than the internarial span (E-N/IN 0.694). The more or less than the internarial span (average E-N/IN canthus rostralis is well defined and very slightly curved. 0.509-0.884). The canthus rostralis may be curved, The eye is large and conspicuous, its diameter greater slightly defined or straight. The eye is moderate to large, than the eye to naris distance. The tympanum is covered its diameter greater than the distance separating it from with skin and very small, its diameter equivalent to one- the nostril. The tympanum is visible, its diameter third of the eye diameter, separated from the eye by a equivalent to one-third to one-quarter of the eye distance greater than its own diameter. The tongue is diameter. The vomerine teeth are in two short oblique broadly cordiform with weakly indented posterior border. series situated between the choanae. The tongue is The terminal discs are prominent. Long and slender cordiform wih a slightly indented posterior border. hindlimbs with a TL/S-V ratio of 0.584. The fingers are moderately long and are equipped with Toe lengths 4>5=3>2>1. The webbing of all toes except narrow lateral fringes; in decreasing order of length for the fourth reaches the base of the discs, while on the 3>4>2>1; the webbing is not extensive, not reaching the fourth toes the webbing reaches the subarticular tubercle sub-articular tubercle at the base of the penultimate at the base of the penultimate phalanx and continues to phalanx on the fourth finger. The terminal discs are the disc via a narrow fringe. The dorsal and lateral moderate. surfaces of the body are finely pitted and striated. There The hind limbs are long with a moderate to high TL/S-V is an inconspicuous supratympanic fold. The throat and ratio (0.524-0.689). Toes in decreasing order of length chest are smooth. Abdomen and nearby halves of the 4>3>or= 5>2>1. The toes are webbed to the discs with ventral surface of the thighs are coarsely granular. There the exception of the fourth which is webbed to the sub- is a small pigmented nuptial pad at the base of the first articular tubercle at the base of the penultimate phalanx. finger. Vocal sac openings are exceptionally long, The skin on the dorsal surfaces is smooth or at most extending from the base of the tongue to the angles of slightly tubercular with very small and widely scattered the jaw. tubercles distributed evenly across the upper body and Ventrally the frogs are a pale creamish colour, stippled upper surfaces of the limbs. The throat is smooth and the with dark brown on the throat. chest, abdomen and lower femora granular. There is a See the detailed description for the genus Angularanta prominent, curved supra-tympanic fold extending from gen. nov. in this paper for details of the separation of the the posterior corner of the eye to the shoulder. On the other subgenera within Angularanta subgen. nov. from all posterior surface of the forearm are tubercles which are other Australasian (Australian and New Guinea) Tree widely separated from one another, in juxtaposition in a Frogs (Pelodryadidae). distinct row, or replaced by a continuous fold. There is a Distribution: A. louisiadensis brunetus subsp. nov. is poorly defined dermal ridge on the posterior surface of confined to Sudest Island in the Louisiade Archipelago, the heel and tarsus. Milne Bay Province of Papua New Guinea. The colouration is very highly variable, including at times, Conservation: As an evolutionary divergent taxon within strong sexual dimorphism in many locations. a confined geographical range, A. louisiadensis brunetus Two species synonymised with Bellarana angiana subsp. nov. is always at high risk of extinction by way of (Boulenger, 1915), namely, B. mintima (Tyler, 1963), human activities or their consequences. The highest risks based on an exceptionally dark variant from south of the are probably from introduced species of vertebrate or central cordillera (Mintima, Chimbu Region, at 6,000 feet, pathogen of some form not previously encountered by Central Highlands, Papua New Guinea. Latitude -5.57 S., the species and for which it may have no reasonable Longitude 144.54 E.), and B. angularis (Loveridge, 1945) defence. from, Mount Wilhelm, 5000-8000 feet, Bismarck Range, Etymology: A. louisiadensis brunetus subsp. nov. is Madang Division, Papua New Guinea, are herein named in reflection of its generally brown dorsum (from resurrected. the Latin word brunneis). This is done on the basis that all three described taxa are BELLARANA GEN. NOV. from different parts of New Guinea to the type locality of LSIDurn:lsid:zoobank.org:act:AF9FC11C-932C-48AB- B. angiana (Arfak Mountains, Vogelkop Peninsula, West 95CB-9F5C3308917D Papua, Indonesia), separated by wide zones of unsuitable habitat and/or well-known biogeographical Type species: Hyla angiana Boulenger, 1915. barriers, combined with the known divergence of the B. Diagnosis: The five described species in the genus angiana lineage being about 9.5 MYA from nearest living Bellarana gen. nov. are readily separated from all other relatives indicating presence of more than one species. Australasian (Australian and New Guinea) Tree Frogs At this stage however, all these species cannot be (Pelodryadidae) by the following suite of characters: The properly separated on the basis of morphology or colour, head is high and usually broader than long (average HL/ or at least this has not yet been done. HW 0.878-1.012), its length usually less than one-third of Within most populations of putative B. angiana, B. the snout to vent length (average HL/S-V 0.300-0.347). mintima and B. angularis it is possible to separate the The snout is not prominent; when viewed from above it is majority of specimens in collections into at least four evenly rounded or obtusely angular; in profile it is main colour groups, with the remainder representing so- rounded or acutely angular. The nostrils are more lateral called intergrades beween the main colour variants. than superior, their distance from the end of the snout The characteristic features of each of the most common less than that from the eye. variant (a term of convenience coined by Tyler 1978) and The distance between the eye and the naris is either the names applied to them are as follows:

1/ Type Variant (as figured by Boulenger, 1915) and Content: Bellarana angiana (Boulenger, 1915) (type depicted on plate 66 of Menzies (2006). The dorsal species); B. angularis (Loveridge, 1945); B. megalops surface is pale green with or without a few indistinct black (Richards and Iskandar, 2006); B. micromembrana (Tyler, spots or marbling. There is a dark (usually black) patch at 1963); B. mintima (Tyler, 1963). the end of the snout, dividing on a level with the nares FLUVIRANA GEN. NOV. and continuing as a broad stripe along the canthus LSIDurn:lsid:zoobank.org:act:4F7D63C9-585D-4862- rostralis to the eye and continuing behind the eye as a BE99-7E1146CBD195 short stripe to the insertion of the forearm. There are Type species: Litoria rara Günther and Richards, 2005. brilliant, white longitudinal stripes on the posterior surface Diagnosis: Fluvirana gen. nov. are a genus of stream- of the forearm, heel and tarsus, and a broad, curved, dwelling frogs only known from the north-west of New white supra-anal bar. The lateral surfaces of the body are Guinea (Indonesia) separated from all other similar dull crimson marked with large, irregular white spots. The species in New Guinea (including those sympatric in ventral surfaces of the body are a paler crimson with northern New Guinea) by having a stout build, only white spots which are particularly dense on the throat. slightly pointed snout from above or below, also being 2/ Brown Variant as depicted in plate 67 of Menzies slightly truncate; distinctive finger webbing on otherwise (2006). A uniform brown dorsally and laterally. The only short, thick fingers, an absence of distinctive markings on markings are a triangular, pale blue patch behind the eye the dorsum; brownish or grey dorsum; limited markings and a white supra-anal bar. The ventral surfaces vary on upper limbs and usually in the form of scattered flecks from white to a very pale brown. or indistinct bars or spots; white to whitish underparts; 3/ So-called Mintima variant. The dorsal and lateral dorsal skin that is either smooth or only moderately surfaces are uniformly a very deep slate and lack granular or tuberculate; short thick limbs which are markings of any kind. The ventral surfaces are pale slate. greyish in colour and spotted with white; relatively dull 4/ Dull variant. Basically similar to the type variant but concealed areas of limbs and a moderate tympanic fold with extensive black markings on the dorsal and lateral that covers the top section of the otherwise exposed surfaces. The white limb and supra-anal markings are tympanum. absent or replaced by poorly defined pale grey patches. Distribution: As far as is known, the three species in this The ventral surfaces are dull grey. genus are confined to north-west New Guinea 5/ So-called Intergrades. Many specimens exhibit large (Indonesia), being known only from Nabire and Waponga white spots on the dorsal surfaces and some the hour- River, Irian Jaya, Indonesia. glass shaped marking on the posterior portion of the Etymology: The new genus name Fluvirana is taken head and anterior portion of the body (a marking from the Latin words “Fluvi” for stream and “rana” for frog, common to other New Guinea tree frog species). These as these are stream dwelling species. markings are not seen in any of the other variants Content: Fluvirana rara (Günther and Richards, 2005) described above. (type species); F. rivicola (Günther and Richards, 2005); B. micromembrana (Tyler, 1963) with a distribution also F. scabra (Günther and Richards, 2005). centered on the central cordillera of New Guinea is HOPVIRIDI GEN. NOV. readily separated from B. angiana (and those species LSIDurn:lsid:zoobank.org:act:FA913E50-A9B1-4667- recently synonymised with it) by having a larger eye, the 9B6A-88E99AD95623 eye is larger than the internarial span and the canthus Type species: Hyla leucova Tyler, 1968. rostralis is strongly curved (particularly in females), versus not curved in B. angiana. Diagnosis: The species in the genus Hopviridi gen. nov. are readily separated from all other Australasian In turn the species B. megalops (Richards and Iskandar, (Australian and New Guinea) Tree Frogs (Pelodryadidae) 2006) is separated from B. micromembrana by its smaller by the following suite of characters: Small, montane size (average male SVL: 24.6-27.5 mm versus 31.7-35.5 species. They are uniformly green, sometimes lacking mm in B. micromembrana), exceptionally large and flash markings on the undersides of limbs or alternatively prominent eyes and conical tubercles on the dorsum. pink with yellow spots in H. leucova (Tyler, 1968) or According to Duellman et al. (2016), the genus Bellarana otherwise bright orange in H. chloronota (Boulenger, gen. nov. diverged from its nearest living relatives in the 1911). Short, slightly fringed, half-webbed fingers, fully divergent genus Angularanta gen. nov. 12.7 MYA, in turn webbed toes and eggs unpigmented, 2 mm in diameter diverged as a pair from their next nearest living relatives ova. Maximum size of females is 30 mm snout-vent. The in the two genera Hopviridi gen. nov. and Ornatanura snout is short with a low internarial span. Vomerine teeth gen. nov. 13.5 MYA and all diverged 21.5 MYA from the are absent. most closely related living species in previously named According to Duellman et al. (2016), the genus Hopviridi genera. gen. nov. diverged from its nearest living relatives in the Distribution: The genus Bellarana gen. nov. is found divergent genus Ornatanura gen. nov. 12.1 MYA. along the main cordillera of New Guinea, from west to These in turn diverged as a pair from their next nearest East and including outlier mountain ranges such as the living relatives in the genera Bellarana gen. nov. and Foja Mountains. Angularanta gen. nov. 13.5 MYA. Etymology: Bella in Latin means “cute” and Rana means All these genera diverged 21.5 MYA from the most “frog” and these are “cute frogs”, hence the new genus closely related living species in previously named genera. name Bellarana.

Distribution: Known only from the upper Sepik River usually wider than toe discs and with distinct circum- system in Papua New Guinea (H. leucova) and the Arfak marginal grooves. Nuptial pads dark brown, roughly tear- Mountains in Irian Jaya (H. chloronota). Alleged records shaped with point of tear oriented in posteroventral from other areas to date (e.g. Upper Fly River, in PNG) direction. Indistinct unpigmented bifid subarticular have invariably turned out to be other, not particularly tubercles usually visible at the base of penultimate similar species. phalanx on all fingers, further indistinct unpigmented Etymology: These animals “hop” because they are frogs subarticular tubercles in series on finger four, indistinct and they are green in colour (viridi in Latin) and so the proximal metacarpal tubercles at base of finger one, and new genus name Hopviridi is a direct combination of small distal metacarpal tubercles at base of three and these. four. Content: Hopviridi leucova (Tyler, 1968) (type species); Toes moderately long, relative lengths 4>3>5>2>1. All H. chloronota (Boulenger, 1911). digits with extensive fleshy, opaque webbing, basal INCERTANURA GEN. NOV. between 1 and 2, extending to anterior end of LSIDurn:lsid:zoobank.org:act:26D001AA-6F36-4E1D- penultimate phalanx on distal edge of 2 and 3 and AB1B-BC898078DE6A proximal edge of five, to halfway along penultimate phalanx on proximal edge of three and four, and base of Type species: Incertanura fakfakensis sp. nov. penultimate tubercle on distal edge of four. Terminal discs Diagnosis: The genus Incertanura gen. nov. are slightly expanded to not really expanded and otherwise, separated from all other Australasian (Australian and narrower than finger discs and with distinct circum- New Guinea) Tree Frogs (Pelodryadidae) by the following marginal grooves. Indistinct unpigmented subarticular suite of characters: tubercles on penultimate phalanx of all toes, single on 1- A long (up to at least 2.8 mm) erectile rostral spike in 3, bifid on 4 and 5. No other tubercles clearly apparent. males that is circular in cross section; relatively small size Small, indistinct unpigmented metatarsal tubercle at base (SVL up to 30.5 mm); slender build (HW/SVL 0.22-0.26); of 1. tibia approximately half length of the body (TL/SVL 0.049- In preservative, ground colour of all dorsal surfaces is 0.053); moderately large eyes (EYE/SVL 0.10-0.12); medium brown, with extensive blueish green and darker moderately small tympanum (TYM/SVL 0.42-0.53); small, brown flecking, maculations, vermiculations and/or rounded and often green tubercles extending across the blotches across all dorsal surfaces, blueish pattern mid-dorsum in life; brown to yellowish-brown dorsal elements sometimes coalesce into three indistinct colouration with, or without, distinct green transverse transverse blotches, overall darker pigmentation bands; and orange colouration in concealed areas of the elements tend to be densest on body and less thighs and axilla. concentrated and more finely reticulated towards distal The five species within Incertanura gen. nov. are further extremities of limbs. defined as follows: Body very slender and elongate, tibia Rostral spike is usually light brown, with extensive darker approximately half length of body; head wider than body brown flecking at base, tending towards unpatterned at in dorsal profile, clearly distinct from neck. Snout rounded tip. White or yellow patches often, but not always, in dorsal view and truncate in lateral view. Rostral spike present on either side of head, extending from in males extending from tip of upper jaw; mean spike posterolateral edge of eye, below tympanum and around length varies among populations and species from axilla. Venter predominantly buff, internal organs different localities. Canthus rostralis moderately well sometimes visible, throat sometimes offwhite, and/or with defined, slightly curved; loreal region concave. Nares two clusters of dark brown maculations laterally, tubercles closer to tip of snout (excluding spike) than to eye, around vent and sometimes those along outer edge of oriented laterally. Eyes moderately large, protruding in limbs, off-white. both dorsal and lateral views; pupil horizontal. Upper jaw Appearance in life is as follows: Dorsal base colouration protruding marginally beyond lower jaw. Tympanum small is light to dark brown, with variable amounts of green and with distinct to indistinct annulus, bordered dorsally by a darker brown spotting, mottling or banding on the body, fleshy supratympanic fold extending to the superior edge limbs and head. Dorsal and lateral tubercles on torso and of the insertion of the upper arm. Choanae small and limbs often, but not always, green and contrasting circular, situated close to anterior and lateral edge of against the brown base colouration. Head brown, often palate; no vomerine teeth visible; tongue fleshy and ovoid with extensive green pigmentation, usually including a and usually with a slightly indented posterior edge. Vocal ring around the eyes and more variably a green slits present in males. Dorsal skin tuberculate; ventral transverse band between the eyes. Dense clusters of skin finely to coarsely granular on throat, abdomen and dark-brown to almost black maculations extend to a tibia; remaining ventral surfaces of limbs smooth; variable degree across the limbs, lateral portions of torso additional tubercles also present in clusters around the and in patches across the back. Off-white patches vent and to a variable extent on sides of ankles and sometimes present below tympanum and on exposed upper forearms. surfaces of lower hindlimbs. Iris pattern complex; base Fingers with relative lengths 3>4>2>1; fleshy, whitish to colouration is usually light brownish, with extensive translucent webbing between all digits, forming a narrow darker brown vermiculations; rim of pupil orange. Hidden basal strip between 1 and 2, extending to disc on distal regions of axilla and groin orange (adapted from Oliver et edge of 2 and proximal edge of 4 and to penultimate al. 2019a). phalanx on both sides on 3. Terminal finger discs Frogs treated by Oliver et al. (2019a) as outlier expanded or narrower (depending on species) and populations of Incertanura pronimia (Menzies, 1993), Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 85

which they placed in the genus Litoria Tschudi, 1838 from morphological differences to the earlier described taxon I. the Fakfak Mountains and near Timika, both being in Irian pronimia. Jaya are within this paper formally named as new With this in mind, it is important that this description of I. species, namely I. fakfakensis sp. nov.and I. cuspis sp. cuspis sp. nov. and the other newly named species, I. nov. based on geographical an morphological divergence fakfakensis sp. nov. set out what separates all five named of these forms with one another and all other known species within the genus from one another. species in the genus. Each of the five relevant species are separated from one According to Duellman et al. (2016), Incertanura gen. another by the following unique suites of meristic and nov. diverged from their nearest living relatives 14.6 MYA, other characters: being other newly named genera (within this paper) and I. pronimia can be readily separated from the other four at least 21.5 MYA diverged from nearest living relatives in species by the following unique suite of (average) previously named genera. meristic characters: rostral spike length/snout-vent length Distribution: Known only from mid-montane regions in from vent to tip of snout minus the spike if present (RL/ New Guinea, including at least one satellite Mountain SVL) of 0.067 (a figure much higher than for I. Range to the north of the main cordillera (Foja fakfakensis sp. nov.and I. cuspis sp. nov. and also much Mountains) and another to the west (Fakfak) lower than for I. pinocchio), head width measured as Etymology: In Latin “Incertis” means variable and transverse distance between tympana/snout-vent length “anura” means frog and hence the effective name from vent to tip of snout minus the spike if present (HW/ “variable frog”, which is a fair description of the variability SVL) of 0.24, horizontal eye diameter/snout-vent length of dorsal colour patterning in these frogs, even within a from vent to tip of snout minus the spike if present (EYE/ given population of a given species and of the same sex. SVL) 0.12, tibia length/snout-vent length from vent to tip Content: Incertanura pronimia (Menzies, 1993) (type of snout minus the spike if present (TL/SVL) 0.51, species); I. cuspis sp. nov.; I. fakfakensis sp. nov.; I. horizontal tympanum diameter/snout-vent length from pinocchio (Oliver, Günther and Richards, 2019); I. vent to tip of snout minus the spike if present (TYM/SVL) vivissimia (Oliver, Richards and Donnellan, 2019). 0.047, transverse diameter of toe 4 disc/snout-vent INCERTANURA CUSPIS SP. NOV. length from vent to tip of snout minus the spike if present LSIDurn:lsid:zoobank.org:act:A586FF1C-BB98-4C47- (4TD/SVL) 0.045 and transverse diameter of finger 3 B83E-AEDFCA6EDAEA disc/ snout-vent length from vent to tip of snout minus the spike if present (3FD/SVL) 0.048. Holotype: A preserved adult specimen at the Museum Zoologie Bogor, Indonesia, specimen numbers, MZB I. cuspis sp. nov. can be readily separated from the other amphibians 13775 collected from the Timika area, four species by the following unique suite of (average) Papua Province, Indonesia, Latitude -4.371 S., Longitude meristic characters: RL/SVL (rostral spike length/snout- 136.270 E. vent length from vent to tip of snout minus the spike if present) 0.034 (a figure much smaller than for all other This facility allows access to its holdings. species), HW/SVL 0.25, EYE/SVL 0.12, TL/SVL 0.51, Paratypes: Five preserved adult specimens at the TYM/SVL 0.047, 4TD/SVL (transverse diameter of toe 4 Museum Zoologie Bogor, Indonesia, specimen numbers, disc/snout-vent length from vent to tip of snout minus the MZB amphibians 13776-13380 collected from the Timika spike if present) 0.034 and 3FD/SVL (transverse area, Papua Province, Indonesia, Latitude -4.371 S., diameter of finger 3 disc/ snout-vent length from vent to Longitude 136.270 E. tip of snout minus the spike if present) 0.042, meaning Diagnosis: All species within Incertanura gen. nov. are this species has much smaller hand and toe pads than all morphologically similar. All can be separated from all other species, beingn indistinct versus distinct I the other other Tree Frogs (Pelodryadidae) by the following suite of species. characters: A long (up to at least 2.8 mm) erectile rostral I. fakfakensis sp. nov. can be readily separated from the spike in males that is circular in cross section; relatively other four species by the following unique suite of small size (SVL up to 30.5 mm); slender build (HW/SVL (average) meristic characters: RL/SVL 0.052, HW/SVL 0.22-0.26); tibia approximately half length of the body 0.25, EYE/SVL 0.11, TL/SVL (tibia length/snout-vent (TL/SVL 0.049-0.053); moderately large eyes (EYE/SVL length from vent to tip of snout minus the spike if present) 0.10-0.12); moderately small tympanum (TYM/SVL 0.42- 0.52 (a figure that is higher than for all other species), 0.53); small, rounded and often green tubercles TYM/SVL (horizontal tympanum diameter/snout-vent extending across the mid-dorsum in life; brown to length from vent to tip of snout minus the spike if present) yellowish-brown dorsal colouration with, or without, 0.043 (a figure lower than for all other species), 4TD/SVL distinct green transverse bands; and orange colouration 0.045, 3FD/SVL 0.049. in concealed areas of the thighs and axilla. I. pinocchio can be readily separated from the other four The three previously described species within the genus, species by the following unique suite of (average) are Incertanura pronimia (Menzies, 1993), with a type meristic characters: RL/SVL 0.086, HW/SVL 0.26, EYE/ locality of near Ok Ma, Western Province of Papua New SVL 0.12, tibia length/snout-vent length from vent to tip of Guinea, I. pinocchio (Oliver, Günther and Richards, 2019) snout minus the spike if present (TL/SVL) 0.48 (a number with a type locality of Foja Mountains, Papua Province, lower than for all other species), horizontal tympanum Indonesia and I. vivissimia (Oliver, Richards and diameter/snout-vent length from vent to tip of snout Donnellan, 2019), with type locality of Hides Ridge, Hela minus the spike if present (TYM/SVL) 0.055 (a number Province, Papua New Guinea. The latter two were higher than for all other species), 4TD/SVL 0.045, 3FD/ defined and diagnosed with direct reference to their

SVL .051. that is circular in cross section; relatively small size (SVL I. vivissimia can be readily separated from the other up to 30.5 mm); slender build (HW/SVL 0.22-0.26); tibia (preceding) four species by the following unique suite of approximately half length of the body (TL/SVL 0.049- characters: TL/SVL 0.52, TYM/SVL 0.049, extensive 0.053); moderately large eyes (EYE/SVL 0.10-0.12); olive-green tubercles across the dorsum and white moderately small tympanum (TYM/SVL 0.42-0.53); small, crenulations; tubercles along the outer edge of the ankle rounded and often green tubercles extending across the (vs absent). dorsal and ventral edges of the lateral mid-dorsum in life; brown to yellowish-brown dorsal flanges on the shanks with extensive white flecking and colouration with, or without, distinct green transverse tubercles and rugose skin on the ventral surface of the bands; and orange colouration in concealed areas of the thigh extending more than half the distance from the thighs and axilla. cloaca to the knee (vs extending less than half way in the The three previously described species within the genus, other species). I. vivissimia also has prominent and larger are Incertanura pronimia (Menzies, 1993), with a type blue (green in life) blotches on the dorsal surface of locality of near Ok Ma, Western Province of Papua New upper arms, forearms, thighs and shanks (vs mottled light Guinea, I. pinocchio (Oliver, Günther and Richards, 2019) and dark), more extensive white flecking on the dorsal with a type locality of Foja Mountains, Papua Province, and ventral edges of lateral flanges on the shank, and Indonesia and I. vivissimia (Oliver, Richards and only sparse dark-brown maculations on the digits and Donnellan, 2019), with type locality of Hides Ridge, Hela ankles (versus dense in the other species) (modified from Province, Papua New Guinea. The latter two were Oliver et al. 2019a). defined and diagnosed with direct reference to their Adult male I. cuspis sp. nov., I. pinocchio and I. morphological differences to the earlier described taxon I. fakfakensis sp. nov. also have noticeably thicker pronimia. forearms than the other two species in the genus. With this in mind, it is important that this description of I. A photo of the type form of I. pronimia in life can be found fakfakensis sp. nov. and the other newly named species, in Oliver et al. (2019a) on page 337 (two top images). I. cuspis sp. nov. set out what separates all five named A photo of I. cuspis sp. nov. in life can be found in Oliver species within the genus from one another. et al. (2019a) on page 337 middle right. Each of the five relevant species are separated from one A photo of I. pinocchio in life can be found in Oliver et al. another by the following unique suites of meristic and (2019a) on page 434, five photos on page. other characters: A photo of I. vivissimia in life can be found in Oliver et al. I. pronimia can be readily separated from the other four (2019b) on page 447 in the top image. species by the following unique suite of (average) All five species are known from relatively few collected meristic characters: rostral spike length/snout-vent length specimens in relatively undisturbed locations, which are from vent to tip of snout minus the spike if present (RL/ increasingly under threat as deforestation and agriculture SVL) of 0.067 (a figure much higher than for I. take hold in most parts of New Guinea, as a direct fakfakensis sp. nov.and I. cuspis sp. nov. and also much consequence of exploding human populations in New lower than for I. pinocchio), head width measured as Guinea and globally. transverse distance between tympana/snout-vent length from vent to tip of snout minus the spike if present (HW/ Distribution: I. cuspis sp. nov. is only known from the SVL) of 0.24, horizontal eye diameter/snout-vent length general area that the six type specimens were collected, from vent to tip of snout minus the spike if present (EYE/ being the Timika area, Papua Province, Indonesia. SVL) 0.12, tibia length/snout-vent length from vent to tip Etymology: The name cuspis in Latin, means “point” or of snout minus the spike if present (TL/SVL) 0.51, “cone” with reference to the frontal appendage of males horizontal tympanum diameter/snout-vent length from of this species. vent to tip of snout minus the spike if present (TYM/SVL) INCERTANURA FAKFAKENSIS SP. NOV. 0.047, transverse diameter of toe 4 disc/snout-vent LSIDurn:lsid:zoobank.org:act:2F7210FA-0ED5-499A- length from vent to tip of snout minus the spike if present 9660-4AAEC291BE7E (4TD/SVL) 0.045 and transverse diameter of finger 3 Holotype: A preserved adult specimen at the Museum disc/ snout-vent length from vent to tip of snout minus the für Naturkunde, Berlin, Germany, specimen number ZMB spike if present (3FD/SVL) 0.048. 85620 collected from Fakfak Mountains, Bomberai I. cuspis sp. nov. can be readily separated from the other Peninsula, Papua Barat Province, Indonesia Latitude - four species by the following unique suite of (average) 2.780 S., Longitude 132.270 E. meristic characters: RL/SVL (rostral spike length/snout- This facility allows access to its holdings. vent length from vent to tip of snout minus the spike if Paratype: A preserved adult specimen at the Museum für present) 0.034 (a figure much smaller than for all other Naturkunde, Berlin, Germany, specimen number ZMB species), HW/SVL 0.25, EYE/SVL 0.12, TL/SVL 0.51, 85621 collected from Fakfak Mountains, Bomberai TYM/SVL 0.047, 4TD/SVL (transverse diameter of toe 4 Peninsula, Papua Barat Province, Indonesia Latitude - disc/snout-vent length from vent to tip of snout minus the 2.780 S., Longitude 132.270 E. spike if present) 0.034 and 3FD/SVL (transverse Diagnosis: All species within Incertanura gen. nov. are diameter of finger 3 disc/snout-vent length from vent to morphologically similar. tip of snout minus the spike if present) 0.042, meaning this species has much smaller hand and toe pads than all All can be separated from all other Tree Frogs other species, beingn indistinct versus distinct in the (Pelodryadidae) by the following suite of characters: A other species. long (up to at least 2.8 mm) erectile rostral spike in males

I. fakfakensis sp. nov. can be readily separated from the INLUSTANURA GEN. NOV. other four species by the following unique suite of LSIDurn:lsid:zoobank.org:act:006C107D-4BEC-41A5- (average) meristic characters: RL/SVL 0.052, HW/SVL BD7B-D02627566E13 0.25, EYE/SVL 0.11, TL/SVL (tibia length/snout-vent Type species: Hyla multiplica Tyler, 1964. length from vent to tip of snout minus the spike if present) Diagnosis: Frogs of the genus Inlustanura gen. nov. are 0.52 (a figure that is higher than for all other species), separated from all other Australasian (Australian and TYM/SVL (horizontal tympanum diameter/snout-vent New Guinea) Tree Frogs (Pelodryadidae) by the following length from vent to tip of snout minus the spike if present) suite of characters: 0.043 (a figure lower than for all other species), 4TD/SVL The known species are characterised by fully webbed 0.045, 3FD/SVL 0.049. outer fingers, a large rounded head with inconspicuous to I. pinocchio can be readily separated from the other four large eyes (depending on the species), and distinctive species by the following unique suite of (average) well developed dermal folds on the posterior surface of meristic characters: RL/SVL 0.086, HW/SVL 0.26, EYE/ the forearm, anus, tibia, tarsus and fifth toe. However, in SVL 0.12, tibia length/snout-vent length from vent to tip of at least one species the development of the fold on the snout minus the spike if present (TL/SVL) 0.48 (a number tarsus may be reduced. These frogs have a distinctive lower than for all other species), horizontal tympanum dorsal colouration of green intersperced with yellow spots diameter/snout-vent length from vent to tip of snout in life (in preservative a distinctive colouration in which minus the spike if present (TYM/SVL) 0.055 (a number blue and violet are the predominant pigments), with higher than for all other species), 4TD/SVL 0.045, 3FD/ yellow spots absent in at least some specimens of one SVL .051. species. These frogs are moderately sized species. I. vivissimia can be readily separated from the other Females attain a maximum snout to vent length of 35-48 (preceding) four species by the following unique suite of mm and males 39-42 mm. characters: TL/SVL 0.52, TYM/SVL 0.049, extensive According to Duellman et al. (2016), the species within olive-green tubercles across the dorsum and white Inlustanura gen. nov. diverged from their nearest living crenulations; tubercles along the outer edge of the ankle relatives 13.4 MYA, being the newly named (within this (vs absent). dorsal and ventral edges of the lateral paper) genus Variabilanura gen. nov. and are at least flanges on the shanks with extensive white flecking and 21.5 MYA diverged from nearest living relatives in tubercles and rugose skin on the ventral surface of the previously named genera. thigh extending more than half the distance from the Distribution: Known from scattered locations in the cloaca to the knee (vs extending less than half way in the ranges areas of New Guinea both north and south of the other species). I. vivissimia also has prominent and larger main cordillera and in both the Papua New Guinea (PNG) blue (green in life) blotches on the dorsal surface of and Indonesian sides. upper arms, forearms, thighs and shanks (vs mottled light Etymology: The Latin word inluster means “bright and dark), more extensive white flecking on the dorsal coloured” and by adding “anura” for frog, the full genus and ventral edges of lateral flanges on the shank, and name means “bright coloured frog”. only sparse dark-brown maculations on the digits and ankles (versus dense in the other species) (modified from Content: Inlustanura multiplica (Tyler, 1964) (type Oliver et al. 2019a). species); I. gasconi (Richards, Oliver, Krey and Tjaturadi, 2009); I. inluster sp. nov. Adult male I. cuspis sp. nov., I. pinocchio and I. fakfakensis sp. nov. also have noticeably thicker INLUSTANURA INLUSTER SP. NOV. forearms than the other two species in the genus. LSIDurn:lsid:zoobank.org:act:D1535947-0F68-4274- A photo of the type form of I. pronimia in life can be found BD76-65FDD2CB230C in Oliver et al. (2019a) on page 337 (two top images). Holotype: A preserved specimen at the South Australian A photo of I. cuspis sp. nov. in life can be found in Oliver Museum, Adelaide, South Australia, Australia, specimen et al. (2019a) on page 337 middle right. number R34360, collected from Magidobo, Southern A photo of I. pinocchio in life can be found in Oliver et al. Highlands Province, Papua New Guinea, Latitude -6.11 (2019a) on page 434, five photos on page. S., Longitude 142.46 E. A photo of I. vivissimia in life can be found in Oliver et al. This government-owned facility allows access to its (2019b) on page 447 in the top image. holdings. All five species are known from relatively few collected Diagnosis: The species Inlustanura inluster sp. nov. from specimens in relatively undisturbed locations, which are the Southern Highlands Province, Papua New Guinea, increasingly under threat as deforestation and agriculture conforms to the genus diagnosis for the genus take hold in New Guinea, as a direct consequence of Inlustanura gen. nov. (this paper). It is morphologically exploding human populations in New Guinea and most similar to the species I. multiplica (Tyler, 1964), globally. originally described as Hyla multiplica Tyler, 1964. However I. inluster sp. nov. is separated from that Distribution: I. fakfakensis sp. nov. is only known from species by lacking any obvious sign of dorsal yellow the type locality in the Fakfak Mountains of Bomberai spots (in life), has a distinctive white marking along the Peninsula, Papua Barat Province, Indonesia and is posterior-ventral edge of the eye (not seen in I. probably confined to this range due to the absence of multiplica), and has reduced development of a dermal suitable habitat in nearby areas (being flat lands). flange on the tarsus, versus well-developed in both I. Etymology: The name fakfakensis is taken from where multiplica and the morphologically similar I. gasconi this species is known to occur. (Richards, Oliver, Krey and Tjaturadi, 2009). Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 88 Australasian Journal of Herpetology

I. gasconi is from the Foja Mountains, Irian Jaya, are coarsely granular. There is a short row of tubercles on the separated from both I. inluster sp. nov. and I. multiplica outer surface of the fourth finger. A more conspicuous by in lacking extensive bright blue, or orange mixed with dermal ridge extends along the outer surface of the bright blue markings on the groin, venter and lateral tarsus and fifth toe. There are numerous tubercles below surfaces, and instead having bright orange groin, thigh the anus and two extremely prominent femoral tubercles. and axillary colouration as well as having noticeably The supra-tympanic fold is inconspicuous. larger eyes (average EYE/SVL 0.13 versus 0.10). Predominantly green in life and may be marked with gold All three preceding species, consisting the entirety of the and black. The intercalary structures are elongate and genus Inlustanura gen. nov. are separated from all other ossified. The hyoid plate lacks alary processes. The ova Australasian (Australian and New Guinea) Tree Frogs are small and pigmented (see below). (Pelodryadidae) by the following suite of characters: The The dorsum is green, stippled with black and known species are characterised by fully webbed outer occasionally marked quite extensively with gold. The fingers, a large rounded head with inconspicuous to large fingers are short and slender, with about one third eyes (depending on the species), and distinctive well webbing; the toes are almost fully webbed. The snout is developed dermal folds on the posterior surface of the slightly rounded in profile. forearm, anus, tibia, tarsus and fifth toe. Although in at The cranial elements are poorly developed. The nasals least one species the development of the fold on the are widely separated medially and do not articulate with tarsus may be reduced. These frogs have a distinctive the sphenethmoid. The sphenethmoid remains double in dorsal colouration of green intersperced with yellow spots the adult. The frontoparietal foramen is large and ovoid. in life (in preservative a distinctive colouration in which The squamosals have short zygomatic rami and very blue and violet are the predominant pigments), with much longer otic rami. The pars facialis of the maxillary is dorsal yellow spots absent in at least some specimens of shallow and the posterior process does not make contact one species. These frogs are moderately sized species. with the maxillary process of the nasal. The alary Females attain a maximum snout to vent length of 35-48 processes of the premaxillary are broad, widely mm and males 39-42 mm. separated medially, and perpendicular to the pars According to Duellman et al. (2016), the species within dentalis. The palatine processes are well developed and Inlustanura gen. nov. diverged from their nearest living do not articulate with each other medially. The prevomers relatives 13.4 MYA, being the newly named (within this are very much reduced. paper) genus Variabilanura gen. nov. and are at least A phlange is present on the distal surface of the third 21.5 MYA diverged from nearest living relatives in metacarpal. The sacral diapophyses are broadly dilated previously named genera. and the intercalary structures are elongate and ossified. Distribution: The species Inlustanura inluster sp. nov. is There are no alary processes on the hyoid plate. The only known from the type locality in the Southern adductor mandibulae externus superficialis is absent. Highlands Province of Papua New Guinea. The ova are small and pigmented (brown animal pole) Etymology: The Latin word inluster means “bright and laid in stagnant marshes. The mean ovidiameters coloured” and is an accurate diagnosis of the colour in are 1.2-1.7 mm. Tadpoles have moderately developed life of this species, noting however that it is well fins and the labial teeth comprise 2 upper and 3 lower camoflagued in its natural habitat. rows (Tyler 1963; Menzies 1972). The chromosome MOECHAENURA GEN. NOV. number is 26. LSIDurn:lsid:zoobank.org:act:BFE0BAB6-ABE1-411D- Moechaeanura gen. nov. is further separated from a B244-7A3A54067C3F number of morphologically similar New Guinea species Type species: Hyperolius nigropunctatus Meyer, 1874. within the genus Variabilanura gen. nov. by its very Diagnosis: Species within Moechaeanura gen. nov. are different reproductive mode. Moechaeanura gen. nov. morphologically similar to those species in the genus species have small brown pigmented eggs (average ova Variabilanura gen. nov., with some differences between size of 1.2-1.7 mm) that are deposited in water, while the two detailed in this description. species in the genus Variabilanura gen. nov. attach a Moechaeanura gen. nov. are separated from all other mass of large green eggs with an average ova size of 2.5 Australasian (Australian and New Guinea) Tree Frogs mm to vegetation overhanging slow moving streams. (Pelodryadidae) by the following suite of characters: They In Variabilanura gen. nov. the ova are laid in groups of are small to medium species (male maximum snout-vent from 4 to 37 (mean 14) on the leaves of trees length 34 mm, female maximum snout-vent length 38 mm overhanging water and around the stems of vegetation at with short, three webbed fingers and almost fully webbed the edge of the water. toes. The ova are surrounded by a very large mass of clear The webbing of the fourth toe extends to a point midway albumen. A period of approximately fourteen days is between the disc and the sub-articular tubercle at the spent within the spawn clump, and the tadpole emerges base of the penultimate phalanx, (versus reaches the possessing internal gills and capable of coordinated sub-articular tubercle at the base of the penultimate movements (Tyler 1978). phalanx on the fourth finger in the genus Variabilanura Frogs within the subgenus Aspercutis subgen. nov. gen. nov.). The skin on the dorsal surfaces of the head, (within Moechaeanura gen. nov.) conform to the above body and limbs is minutely roughened. diagnosis for the genus Moechaeanura gen. nov., except The throat and chest are smooth except for a few for their more extensive finger webbing, highly flattened tubercles. The abdomen and lower femora are tuberculate dorsum (versus smooth to slightly tuberculate Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 89

in the nominate subgenus), distinctively crenulated ridges The abdomen and lower femora are coarsely granular. following the outer surfaces of the lower limbs, silvery- There is a short row of tubercles on the outer surface of gold and heavily veined iris and fully truncate snout the fourth finger. A more conspicuous dermal ridge (versus normal to slightly truncate in the nominate extends along the outer surface of the tarsus and fifth subgenus) which as a full diagnosis separates them from toe. There are numerous tubercles below the anus and all other New Guinea frogs. two extremely prominent femoral tubercles. The supra- Frogs within the subgenus Telaater subgen. nov. are tympanic fold is inconspicuous. separated from all other New Guinea tree frogs by having Predominantly green in life and may be marked with gold full black webbing on hands and feet, the venter with and black. The intercalary structures are elongate and extensive areas of black, white and yellow and a ossified. The hyoid plate lacks alary processes. The ova transparent periphery on the tympanic membrane are small and pigmented (see below). combined with a small adult size of less than 30 mm. the The dorsum is green, stippled with black and placement of these species within a subgenus within a occasionally marked quite extensively with gold. The greater Moechaeanura gen. nov. is tentative and fingers are short and slender, with about one third elevation to full genus may be required as further webbing; the toes are almost fully webbed. The snout is evidence is obtained. slightly rounded in profile. According to Duellman et al. (2016), the genus The cranial elements are poorly developed. The nasals Moechaeanura gen. nov. diverged from its nearest living are widely separated medially and do not articulate with relatives, the morphologically distinctive Nasuscuspis the sphenethmoid. The sphenethmoid remains double in gen. nov. 13.4 MYA, and the morphologically similar the adult. The frontoparietal foramen is large and ovoid. genera Incertanura gen. nov., Inlustanura subgen. nov. The squamosals have short zygomatic rami and very and Variabilanura gen. nov. 17.8 MYA and with each of much longer otic rami. The pars facialis of the maxillary is the latter three genera diverging from one another at shallow and the posterior process does not make contact least 13.4 MYA. with the maxillary process of the nasal. The alary Distribution: New Guinea including offshore islands to processes of the premaxillary are broad, widely the east and west at least as far as Halmahera. separated medially, and perpendicular to the pars Etymology: In Latin “Moechaeanura” means drab frog, dentalis. The palatine processes are well developed and which is true in the context of the colouration of a number do not articulate with each other medially. The prevomers of species in this genus. are very much reduced. Content: Moechaeanura nigropunctatus (Meyer, 1874) A phlange is present on the distal surface of the third (type species); M. albatermacula sp. nov.; M. biakensis metacarpal. The sacral diapophyses are broadly dilated, (Günther, 2006); M. christianbergmanni (Günther, 2008); and the intercalary structures are elongate and ossified. M. richardsi (Dennis and Cunningham, 2006); M. rubrops There are no alary processes on the hyoid plate. The (Kraus and Allison, 2004); M. singadanae (Richards, adductor mandibulae externus superficialis is absent. 2005); M. spica sp. nov.; M. tritong sp. nov.; M. umarensis The ova are small and pigmented (brown animal pole) (Günther, 2008); M. verae (Günther, 2004); M. and laid in stagnant marshes. The mean ovidiameters vocivincens (Menzies, 1972); M. wapogaensis (Richards are 1.2-1.7 mm. Tadpoles have moderately developed and Iskandar, 2001). fins and the labial teeth comprise 2 upper and 3 lower ASPERCUTIS SUBGEN. NOV. rows (Tyler 1963; Menzies 1972). The chromosome LSIDurn:lsid:zoobank.org:act:57D4B7F9-DD7E-4E98- number is 26. 8496-50B075E95CD4 Moechaeanura gen. nov. is further separated from a Type species: Litoria verae Günther, 2004. number of morphologically similar New Guinea species Diagnosis: Species within Moechaeanura gen. nov. are within the genus Variabilanura gen. nov. by its very morphologically similar to those species in the genus different reproductive mode. Moechaeanura gen. nov. Variabilanura gen. nov., with some differences between species have small brown pigmented eggs (average ova the two detailed in this description. size of 1.2-1.7 mm) that are deposited in water, while species in the genus Variabilanura gen. nov. attach a Moechaeanura gen. nov. are separated from all other mass of large green eggs with an average ova size of 2.5 Australasian (Australian and New Guinea) Tree Frogs mm to vegetation overhanging slow moving streams. (Pelodryadidae) by the following suite of characters: In Variabilanura gen. nov. the ova are laid in groups of They are small to medium species (male maximum from 4 to 37 (mean 14) on the leaves of trees snout-vent length 34 mm, female maximum snout-vent overhanging water, and around the stems of vegetation at length 38 mm with short, three webbed fingers and the edge of the water. The ova are surrounded by a very almost fully webbed toes. large mass of clear albumen. A period of approximately The webbing of the fourth toe extends to a point midway fourteen days is spent within the spawn clump, and the between the disc and the sub-articular tubercle at the tadpole emerges possessing internal gills and capable of base of the penultimate phalanx, (versus reaches the coordinated movements (Tyler 1978). sub-articular tubercle at the base of the penultimate Frogs within the subgenus Aspercutis subgen. nov. phalanx on the fourth finger in the genus Variabilanura (within Moechaeanura gen. nov.) conform to the above gen. nov.). The skin on the dorsal surfaces of the head, diagnosis for the genus Moechaeanura gen. nov., except body and limbs is minutely roughened. The throat and for their more extensive finger webbing, highly chest are smooth except for a few flattened tubercles. tuberculate dorsum (versus smooth to slightly tuberculate

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 90 Australasian Journal of Herpetology in the nominate subgenus), distinctively crenulated ridges abdomen and lower femora are coarsely granular. There following the outer surfaces of the lower limbs, silvery- is a short row of tubercles on the outer surface of the gold and heavily veined iris and fully truncate snout fourth finger. A more conspicuous dermal ridge extends (versus normal to slightly truncate in the nominate along the outer surface of the tarsus and fifth toe. There subgenus) which as a full diagnosis separates them from are numerous tubercles below the anus and two all other New Guinea frogs. extremely prominent femoral tubercles. The supra- Frogs within the subgenus Telaater subgen. nov. are tympanic fold is inconspicuous. separated from all other New Guinea tree frogs by having Predominantly green in life and may be marked with gold full black webbing on hands and feet, the venter with and black. The intercalary structures are elongate and extensive areas of black, white and yellow and a ossified. The hyoid plate lacks alary processes. The ova transparent periphery on the tympanic membrane are small and pigmented (see below). combined with a small adult size of less than 30 mm. the The dorsum is green, stippled with black and placement of these species within a subgenus within a occasionally marked quite extensively with gold. The greater Moechaeanura gen. nov. is tentative and fingers are short and slender, with about one third elevation to full genus may be required as further webbing; the toes are almost fully webbed. The snout is evidence is obtained. slightly rounded in profile. According to Duellman et al. (2016), the genus The cranial elements are poorly developed. The nasals Moechaeanura gen. nov. diverged from its nearest living are widely separated medially and do not articulate with relatives, the morphologically distinctive Nasuscuspis the sphenethmoid. The sphenethmoid remains double in gen. nov. 13.4 MYA, and the morphologically similar the adult. The frontoparietal foramen is large and ovoid. genera Incertanura gen. nov., Inlustanura subgen. nov. The squamosals have short zygomatic rami and very and Variabilanura gen. nov. 17.8 MYA and with each of much longer otic rami. The pars facialis of the maxillary is the latter three genera diverging from one another at shallow and the posterior process does not make contact least 13.4 MYA. with the maxillary process of the nasal. The alary Distribution: The two species in this subgenus are only processes of the premaxillary are broad, widely known from their type localities being, Wandamen separated medially, and perpendicular to the pars Peninsula, near the Birds Head of Irian Jaya, Indonesia dentalis. The palatine processes are well developed and and Huon Peninsula, northeastern Papua New Guinea. do not articulate with each other medially. The prevomers Etymology: The subgenus name “Aspercutis”comes are very much reduced. from the Latin words meaning rough skinned. The “a” at A phlange is present on the distal surface of the third the end of “Aspera” has been removed from this genus metacarpal. The sacral diapophyses are broadly dilated, name to prevent potential risk of creating a homonym and the intercalary structures are elongate and ossified. and so the spelling is deliberate. There are no alary processes on the hyoid plate. The Content: Moechaeanura (Aspercutis) verae (Günther, adductor mandibulae externus superficialis is absent. 2004) (type species); M. (Aspercutis) singadanae The ova are small and pigmented (brown animal pole) (Richards, 2005). and laid in stagnant marshes. The mean ovidiameters TELAATER SUBGEN. NOV. are 1.2-1.7 mm. Tadpoles have moderately developed LSIDurn:lsid:zoobank.org:act:028D0D83-90D3-4BDB- fins and the labial teeth comprise 2 upper and 3 lower 8BC4-12AE2518DA32 rows (Tyler 1963; Menzies 1972). The chromosome Type species: Litoria richardsi Dennis and Cunningham, number is 26. 2006. Moechaeanura gen. nov. is further separated from a Diagnosis: Species within Moechaeanura gen. nov. are number of morphologically similar New Guinea species morphologically similar to those species in the genus within the genus Variabilanura gen. nov. by its very Variabilanura gen. nov., with some differences between different reproductive mode. Moechaeanura gen. nov. the two detailed in this description. species have small brown pigmented eggs (average ova size of 1.2-1.7 mm) that are deposited in water, while Moechaeanura gen. nov. are separated from all other species in the genus Variabilanura gen. nov. attach a Australasian (Australian and New Guinea) Tree Frogs mass of large green eggs with an average ova size of 2.5 (Pelodryadidae) by the following suite of characters: They mm to vegetation overhanging slow moving streams. are small to medium species (male maximum snout-vent length 34 mm, female maximum snout-vent length 38 mm In Variabilanura gen. nov. the ova are laid in groups of with short, three webbed fingers and almost fully webbed from 4 to 37 (mean 14) on the leaves of trees toes. overhanging water, and around the stems of vegetation at the edge of the water. The webbing of the fourth toe extends to a point midway between the disc and the sub-articular tubercle at the The ova are surrounded by a very large mass of clear base of the penultimate phalanx, (versus reaches the albumen. A period of approximately fourteen days is sub-articular tubercle at the base of the penultimate spent within the spawn clump, and the tadpole emerges phalanx on the fourth finger in the genus Variabilanura possessing internal gills and capable of coordinated gen. nov.). movements (Tyler 1978). The skin on the dorsal surfaces of the head, body and Frogs within the subgenus Aspercutis subgen. nov. limbs is minutely roughened. The throat and chest are (within Moechaeanura gen. nov.) conform to the above smooth except for a few flattened tubercles. The diagnosis for the genus Moechaeanura gen. nov., except for their more extensive finger webbing, highly

tuberculate dorsum (versus smooth to slightly tuberculate limbs and concealed surfaces of the limbs. The dark in the nominate subgenus), distinctively crenulated ridges patch at the jaw angles. following the outer surfaces of the lower limbs, silvery- M. albatermacula sp. nov. has a dorsum that is a gold and heavily veined iris and fully truncate snout lichenous grey-brown with ill-defined black spots; lower (versus normal to slightly truncate in the nominate limbs are white with dark black cross bars; concealed subgenus) which as a full diagnosis separates them from surfaces of the thighs and posterior of the belly are a all other New Guinea frogs. brilliant golden yellow. Frogs within the subgenus Telaater subgen. nov. are M. nigropunctatus is readily separated from the other two separated from all other New Guinea tree frogs by having species by being grey-green to grey-brown dorsally with full black webbing on hands and feet, the venter with well-defined green or brown spotting or marks, especially extensive areas of black, white and yellow and a on the limbs, hands and feet; concealed surfaces of the transparent periphery on the tympanic membrane thighs, groins and axillae are brilliant orange; ventral combined with a small adult size of less than 30 mm. the surface white; lateral sides of throat lightly or heavily placement of these species within a subgenus within a spotted black; iris is gold and with heavy black veins. greater Moechaeanura gen. nov. is tentative and M. nigropunctatus, M. tritong sp. nov. and M. elevation to full genus may be required as further albatermacula sp. nov. are readily separated from all evidence is obtained. other Tree Frogs in New Guinea by the following suite of According to Duellman et al. (2016), the genus characters: The head is moderately flattened and longer Moechaeanura gen. nov. diverged from its nearest living than broad ( HL/HW 1.033-1.077), its length slightly or relatives, the morphologically distinctive Nasuscuspis considerably more than one third of the snout to vent gen. nov. 13.4 MYA, and the morphologically similar length (HL/S-V 0.334-0.366). The snout is not prominent; genera Incertanura gen. nov., Inlustanura subgen. nov. when viewed from above and in profile it is gently and Variabilanura gen. nov. 17.8 MYA and with each of rounded. the latter three genera diverging from one another at The nostrils are lateral, their distance from the tip of the least 13.4 MYA. snout less than that from the eye. The canthus rostralis is Distribution: Known only from the type localities of the curved and only slightly defined. The eye is large and type specimens of each species, being the upper prominent, its diameter greater than the distance reaches of the Mamberano Drainage, Irian Jaya, separating it from the nostril. The tympanum is covered Indonesia and the Upper Fly River drainage in Western with skin, its diameter equivalent to from one-quarter to Province, Papua New Guinea. one-half of the eye diameter. The vomerine teeth are Etymology: The subgenus name “Telaater” comes from reduced to a single or pair of teeth near each choana and the Latin words “web” and “black” in reference to the are not elevated. The tongue is broadly cordiform with a distinctive black webbing on the feet of both species. slightly indented posterior margin. The fingers are short Content: Moechaeanura (Telaater) richardsi (Dennis and and equipped with fairly prominent lateral fringes and Cunningham, 2006); M. (Telaater) spica sp. nov.. paired sub-articular tubercles; in decreasing order of MOECHAEANURA (MOECHAEANURA) TRITONG SP. length 3>4>2>1. The webbing reaches midway up the NOV. terminal phalanx on the fourth finger. The terminal discs LSIDurn:lsid:zoobank.org:act:1D81F542-71BB-4ECE- are prominent. 941D-5D68113AF63B The hind limbs are moderate with a TL/S-V ratio of 0.526- Holotype: A preserved specimen at the South Australian 0.566. Toes in decreasing order of length 4>5 >3>2>1. Museum, Adelaide, South Australia, Australia, specimen The webbing reaches the base of the disc of all toes, number R65036, collected from the Western Province, except the fourth where it extends to a point midway Papua New Guinea, Latitude -5.7292 S., Longitude between the disc and the sub-articular tubercle at the 142.2633 E. This government-owned facility allows base of the penultimate phalanx. access to its holdings. The skin on the dorsal surfaces of the head, body and Diagnosis: Moechaeanura (Moechaeanura) tritong sp. limbs is minutely roughened. The throat and chest are nov. and M. albatermacula sp. nov. have until now been smooth but for a few flattened tubercles. The abdomen treated as a Papua New Guinea populations of M. and lower femora are coarsely granular. There is a short nigropunctatus (Meyer, 1874), with a type locality of row of tubercles on the outer surface of the fourth finger. Japen Island, West Papua, Indonesia, better known to A more conspicuous dermal ridge extends along the most people as Litoria nigropunctatus (Meyer, 1874). outer surface of the tarsus and fifth toe. There are numerous tubercles below the anus and two extremely M. tritong sp. nov. and M. albatermacula sp. nov. would prominent femoral tubercles. The supra-tympanic fold is key out as M. nigropunctatus in Tyler (1968) in terms of inconspicuous. separation from all other New Guinea species of Tree Frogs. However M. tritong sp. nov. from Western Distribution: M. tritong sp. nov. is known only definitively Province, Papua New Guinea and M. albatermacula sp. from the type locality, south of the main central cordillera nov. from the northern foothills of the Owen Stanley in the Western Province of Papua New Guinea. Range, Papua New Guinea are readily separated from Etymology: In Latin, Trito means unmarked and so the M. nigropunctatus as follows: letters NG, which are an abbreviation for New Guinea are M. tritong sp. nov. has a grey-green dorsum without added to the name to to make the word tritong to mean marks, except for a yellow patch above the axilla. Ventral “unmarked from New Guinea”, in reflection to the most surface is white except for the intense yellow below the common dorsal colour state for the species and where it Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. comes from. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 92 Australasian Journal of Herpetology

MOECHAEANURA (MOECHAEANURA) prominent, its diameter greater than the distance ALBATERMACULA SP. NOV. separating it from the nostril. The tympanum is covered LSIDurn:lsid:zoobank.org:act:D65F2F2C-3972-4363- with skin, its diameter equivalent to from one-quarter to 8E2A-0BC145B18795 one-half of the eye diameter. The vomerine teeth are Holotype: A preserved specimen at the National reduced to a single or pair of teeth near each choana and Museum of Natural History, Smithsonian Institution are not elevated. The tongue is broadly cordiform with a (USNM), Washington, DC, United States of America, slightly indented posterior margin. The fingers are short Amphibians and Reptiles collection specimen number and equipped with fairly prominent lateral fringes and 121216 collected from Dobadura, Northern Province, paired sub-articular tubercles; in decreasing order of Papua New Guinea, Latitude -8.7708 S., Longitude length 3>4>2>1. The webbing reaches midway up the 148.375 E. terminal phalanx on the fourth finger. This facility allows access to its holdings. The terminal discs are prominent. Paratype: A preserved specimen at the National The hind limbs are moderate with a TL/S-V ratio of 0.526- Museum of Natural History, Smithsonian Institution 0.566. Toes in decreasing order of length 4>5 >3>2>1. (USNM), Washington, DC, United States of America, The webbing reaches the base of the disc of all toes, Amphibians and Reptiles collection specimen number except the fourth where it extends to a point midway 269421, collected from 7 miles north of Kokoda, Northern between the disc and the sub-articular tubercle at the Province, Papua New Guinea, Latitude -8.7785 S., base of the penultimate phalanx. Longitude 147.741. The skin on the dorsal surfaces of the head, body and Diagnosis: Moechaeanura (Moechaeanura) limbs is minutely roughened. The throat and chest are albatermacula sp. nov. and M. tritong sp. nov. have until smooth but for a few flattened tubercles. The abdomen now been treated as a Papua New Guinea populations of and lower femora are coarsely granular. There is a short M. nigropunctatus (Meyer, 1874), with a type locality of row of tubercles on the outer surface of the fourth finger. Japen Island, West Papua, Indonesia, better known to A more conspicuous dermal ridge extends along the most people as Litoria nigropunctatus (Meyer, 1874). outer surface of the tarsus and fifth toe. There are M. tritong sp. nov. and M. albatermacula sp. nov. would numerous tubercles below the anus and two extremely key out as M. nigropunctatus in Tyler (1968) in terms of prominent femoral tubercles. The supra-tympanic fold is separation from all other New Guinea species of Tree inconspicuous. Frogs. However M. tritong sp. nov. from Western Distribution: M. albatermacula sp. nov. is only Province, Papua New Guinea and M. albatermacula sp. definitively known from the Northern Province of Papua nov. from the northern foothills of the Owen Stanley New Guinea near where the holotype and paratype were Range, Papua New Guinea are readily separated from caught. M. nigropunctatus as follows: Etymology: In Latin “alba ater macula” literally means M. tritong sp. nov. has a grey-green dorsum without pale black spots and so the name albatermacula literally marks, except for a yellow patch above the axilla. Ventral means that same thing and refers to the dorsal surface is white except for the intense yellow below the colouration of the species. limbs and concealed surfaces of the limbs. The dark MOECHAEANURA (TELEATER) SPICA SP. NOV. patch at the jaw angles. LSIDurn:lsid:zoobank.org:act:B5929C08-060F-44E5- M. albatermacula sp. nov. has a dorsum that is a BA4A-84AFF569BEFD lichenous grey-brown with ill-defined black spots; lower Holotype: A preserved adult male specimen at the limbs are white with dark black cross bars; concealed Museum Zoologie Bogor, Indonesia, specimen number surfaces of the thighs and posterior of the belly are a MZB Amphibians 11823 caught in forest adjacent to Tin brilliant golden yellow. River, Mamberamo Drainage, Papua, Indonesia, Latitude M. nigropunctatus is readily separated from the other two -3.1730 S., Longitude 134.3453 E. species by being grey-green to grey-brown dorsally with This facility allows access to its holdings. well-defined green or brown spotting or marks, especially Diagnosis: The holotype of this species Moechaeanura on the limbs, hands and feet; concealed surfaces of the spica sp. nov. was the paratype for the similar species thighs, groins and axillae are brilliant orange; ventral originally described as Litoria richardsi Dennis and surface white; lateral sides of throat lightly or heavily Cunningham, 2006, but now placed in the genus spotted black; iris is gold and with heavy black veins. Moechaeanura gen. nov.. Their holotype was an adult M. nigropunctatus, M. tritong sp. nov. and M. female collected at 5.5 km west of Tabubil township, albatermacula sp. nov. are readily separated from all Western Province, Papua New Guinea, Latitude -5.1730 other Tree Frogs in New Guinea by the following suite of S., Longitude 141.1155 E. characters: The head is moderately flattened and longer Dennis and Cunningham (2006) concluded that they than broad ( HL/HW 1.033-1.077), its length slightly or were of opposite sexes of the same species, but this is considerably more than one third of the snout to vent not the contention here. length (HL/S-V 0.334-0.366). The snout is not prominent; Both specimens were morphologically distinct from one when viewed from above and in profile it is gently another and the noting that closely related species do not rounded. show significant sexual dimporphism to anywhere near The nostrils are lateral, their distance from the tip of the the extent seen between the two relevant specimens, one snout less than that from the eye. The canthus rostralis is is forced to conclude that they are in fact separate curved and only slightly defined. The eye is large and Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 93

species. Furthermore, the two specimens come from by the following suites of characters: These species are different bio-regions, with one being from north of the characterised by their small size (males 23.4-30.0 mm, central New Guinea cordillera and the other from the females 27.4-35.4 mm), moderate to relatively long limbs south. This also leads to a reasonable inference that the (average TL/S-V 0.522-0.604) and broadly spaced nares two frogs would be of different species, even in the (average E-N/IN 0.611-0.818). absence of strong morphological divergence. In The colouration is highly variable. combination these two elements are compelling evidence The dorsal surface of the head, body and limbs may for two species being involved and not just one. range from light sandy grey, or brown to dark grey and is Moechaeanura spica sp. nov. is readily separated from darkened by the presence of very dense, minute, dark M. richardsi (Dennis and Cunningham, 2006) by having brown to black stippling. little if any webbing between the first and second fingers, The anterior portion of the head bordered laterally by the versus webbing that goes to the first finger disc and canthus rostralis, and posteriorly to a line between the nearly the second finger disc in M. richardsi. anterior portions of the upper eyelids is an immaculate The dorsum of M. spica sp. nov. is a light grey colour pale green, although in some species this may be overlain with scattered yellowish brown irregular reduced to be brown or grey with some lime green spots markings, with a limited number of chocolate brown spots or markings. and blotches, mainly on the flanks. There are areas of The head is longer than broad (average HL/HW 1.132), bright yellow in the groin. Forearms have few if any dark its length equivalent to more than one-third of the snout brown or black markings. The cross-bands on the upper to vent length (average HL/S-V 0.350). The snout is hind legs are thick, brown in colour and mainly complete. evenly rounded when viewed from above and projects The white tipped tubercles on the sides of body and to a slightly in profile. The nostrils are more lateral than lesser extent elsewhere are of moderate size. The superior, their distance from the tip of the snout less than tympanum is mainly brown. The iris is pink and brown. that from the eye. By contrast M. richardsi has a relatively dark blue-grey The distance between the eye and the naris is less than dorsum, with irregular black markings forming cross the internarial span (average E-N/IN 0.727). The canthus bands across the body and also and huge areas of bright rostralis is well defined and distinctly curved. yellow in the groin. Forearms have numerous black The eye is prominent and bulges above the head, its markings. The cross-bands on the upper hind legs are diameter greater than the eye to naris distance and less thin, black in colour, mainly broken and mainly than the internarial span. The superior one-quarter to incomplete. The white tipped tubercles on the sides of one-third of the tympanum is hidden beneath the supra- body and to a lesser extent elsewhere are large and tympanic fold. The diameter of the tympanum is prominent. The tympanum is mainly blue. The iris is equivalent to less than one third of the eye diameter to yellow and grey. slightly more than one-third of the eye diameter. The Both M. spica sp. nov. and M. richardsi constituting the vomerine teeth are in two small oblique series between entirety of the subgenus Telaater subgen. nov. are the choanae. The tongue is almost circular and lacks a separated from all other Tree Frogs in New Guinea and posterior indentation. Australia by a combination of black webbing on the feet, The fingers are long and lack lateral fringes; in venter with large areas of black, white and yellow, a small decreasing order of length 3>4>2>1. There is a small size being less than 30 mm snout vent length in both vestige of webbing between the third and fourth fingers. sexes and a transparent periphery on the tympanic The terminal discs are not prominent, with the degree of membrane. expansion varying slightly between species. Areas where both M. spica sp. nov. and M. richardsi are The hind limbs are relatively long with an average TL/S-V known to occur in are regions of severe habitat ratio of 0.558. Toes in decreasing order of length degradation and explosive human population growth and 4>5=3>2>1. On the fifth toe, the webbing extends two- further research on these species should be conducted thirds up the penultimate phalanx of the fifth and to the as a matter of urgency. base of the penultimate phalanx on the fourth. Colour photos of both M. spica sp. nov. and M. richardsi The skin on the dorsal surface of the head, body and in life are depicted in Dennis and Cunningham (2006) on limbs is either smooth with numerous small, spaced page 66 (M. spica sp. nov. in photo “B”). apart, scattered tubercles all over, ranging down to Distribution: M. spica sp. nov. is known only from the smooth all over but for a few small and only slightly type locality at Tin River, Mamberamo Drainage, Papua, developed tubercles on the scapular region. Indonesia. The throat, abdomen and lower surface of the thighs are Etymology: The name “spica” means “spikey” in Latin in very weakly granular. There is a row of small tubercles reference to the tubercles on the upper flanks and other extending posteriorly from the angle of the jaws and a parts of the frog’s body. patch of similar sized tubercles beneath the anus. ORNATANURA GEN. NOV. Of the tubercles at the angles of the jaws those anterior LSIDurn:lsid:zoobank.org:act:C857380D-A567-4F6E- to the tympanum are green, grey or white, whilst those A3C5-298B91B032A8 posterior to it are usually white. Type species: Hyla modica Tyler, 1968. The scapular tubercles are usually green, and the lateral Diagnosis: The species in the genus Ornatanura gen. surfaces of the body between the axilla and the groin are nov. are readily separated from all other Australasian one or other of being, 1/ Liberally spotted with white, 2/ (Australian and New Guinea) Tree Frogs (Pelodryadidae) Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 94 Australasian Journal of Herpetology

Grey marks or spots on a cream background, 3/ Back The four preceding species are separated from one marks in the form of large spots or blotches, circled by another as follows. white and on an otherwise light lime green background. The species Ornatanura parsviridis sp. nov. from Madang The ventral surfaces are creamish with less dense is similar in most respects to O. modica (Tyler, 1968), but stippling than appears on the dorsum and although is readily separated from that taxon as well as O. dense stippling occurs in some specimens. parscinereo sp. nov. and O. leucopicturas sp. nov. by the The greatest density of groups of chromatophores on the following suite of colouration character and morphology ventral surface appear on the throat and particularly traits: Flanks are light greenish in colour overlain with towards the labial margins, with more on the upper than about 6 large black spots or blotches (sometimes lower margin and greatest intensity between eye and ear, merged), in turn edged with thick white lines, dorsal although the exact configuration varies between surface of the hindlimbs are medium brown with indistinct specimens and species. dark brown to black crossbands. Upper forearms are Males vary in size from 23.4 mm to 30.0 mm and females barred with white on a dark background. Small blunt from 27.4-35.4mm. The head length is consistently tubercles scattered across all parts of the dorsum and longer than broad (average HL/HW 1.028-1.155) and the limbs, and only a small percentage are white tipped. The head length varies from one-third to considerably more tympanum is less than a third of the diameter of the eye. than one-third of the snout to vent length, the complete By contrast O. modica from the higher elevation areas in HL/S-V range the region bound by the Star Mountain, West Papua and being 0.333-0.404. The E-N/IN range is 0.611-0.818 and the Eastern Highlands Province in Papua New Guinea, the eye diameter is consistently smaller than the mainly from upper reaches of south flowing drainages, internarial span. The average TL/S-V range is 0.522- has flanks that are greenish in colour and liberally 0.604. spotted with white. Markings on the upper forearms are The diploid chromosome number is 26. indistinct and blotches on the upper surfaces of the hind limbs do not form indistinct or obvious bands. The skin on The morphologically similar species Bellarana the dorsal surface of the head, body and limbs is smooth micromembrana (Tyler, 1963) from Madang, is most but for a few small and only slightly developed tubercles readily separated from the species within Ornatanura on the scapular region. The tympanum is slightly more gen. nov. by its distinctive orange upper iris, versus than a third of the diameter of the eye. yellowish in Ornatanura gen. nov. species. The species O. parscinereo sp. nov. from Utikini Village, According to Duellman et al. (2016), the genus At Waa River in the Tembagapura Area, West Papua is Ornatanura gen. nov. diverged from its nearest living separated from O. parsviridis sp. nov., O. modica and O. relatives in the divergent genus Hopviridi gen. nov. 12.1 leucopicturas by having a dorsum with numerous MYA, in turn diverged as a pair from their next nearest scattered and large tubercles that are the same colour as living relatives in the genera Bellarana gen. nov. and the skin surrounding them. There are no distinctive Angularanta gen. nov. 13.5 MYA and all diverged 21.5 markings on upper surfaces of limbs, green on the MYA from the most closely related living species in dorsum is restricted to the upper surface of the head previously named genera. anterior to the eyes, which also has some dark peppering Distribution: Species of Ornatanura gen. nov. are on it and similarly peppered green on the upper surfaces abundant in lower montane forests in geologically more of the arms, save for scattered green spots elsewhere. ancient ranges and uplands of New Guinea from the far The tympanum is less than a third of the diameter of the south-east to far north-west. eye. Etymology: The etymology of the new genus Ornatanura The species O. leucopicturas from central Province, is simply a reference in Latin to the fact that the species Papua New Guinea is readily separated from in this genus are ornate frogs. O. parsviridis sp. nov., O. parscinereo sp. nov., O. modica Content: Ornatanura modica (Tyler, 1968) (type speces); and O. leucopicturas by having a brownish grey dosum O. parsviridis sp. nov.; O. parscinereo sp. nov.; O. with numerous scattered black spots, as well as leucopicturas sp. nov. numerous areas of green, generally arranged in one or ORNATANURA LEUCOPICTURAS SP. NOV. more irregular blotches forming a sort of longitudinal LSIDurn:lsid:zoobank.org:act:CE695639-BD80-4EF6- stripe, rich immaculate green on the upper snout anterior B1A1-B34D57C22B6D to the eyes; large black spots and blotches from the Holotype: A preserved specimen at the Australian dorsum spread to the flanks, where their size generally Museum, Sydney, New South Wales, Australia, specimen increases but number decreases. There are scattered number R.23566 collected from Wharton Range, Central small tubercles all over the body, with a noticeable Province, Papua New Guinea, Latitude -8.33 S., general absence on the head, save for an exceptionally Longitude 147.15 E. large series of tubercles above each eye. Most are white This government-owned facility allows access to their or light tipped. holdings. O. parsviridis sp. nov. in life is depicted in Menzies (2006) Diagnosis: The three species O. leucopicturas sp. nov., on plate 75. O. parscinereo sp. nov. and O. parsviridis sp. nov. are all The four preceding species, being the entirety of the similar in most respects to O. modica (Tyler, 1968) and genus Hopviridi gen. nov. are readily separated from all like O. modica conform to the genus diagnosis for other Australasian (Australian and New Guinea) Tree Ornatanura gen. nov.. Frogs (Pelodryadidae) by the following suite of

characters: These species are characterised by their white and on an otherwise light lime green background. small size (males 23.4-30.0 mm, females 27.4-35.4 mm), The ventral surfaces are creamish with less dense moderate to relatively long limbs (average TL/S-V 0.522- stippling than appears on the dorsum and although 0.604) and broadly spaced nares (average E-N/IN 0.611- dense stippling occurs in some specimens. 0.818). The colouration is highly variable. The greatest density of groups of chromatophores on the The dorsal surface of the head, body and limbs may ventral surface appear on the throat and particularly range from light sandy grey, or brown to dark grey and is towards the labial margins, with more on the upper than darkened by the presence of very dense, minute, dark lower margin and greatest intensity between eye and ear, brown to black stippling. The although the exact configuration varies between anterior portion of the head bordered laterally by the specimens and species. canthus rostralis, and posteriorly to a line between the Males vary in size from 23.4 mm to 30.0 mm and females anterior portions of the upper eyelids is an immaculate from 27.4-35.4mm. The head length is consistently pale green, although in some species this may be longer than broad (average HL/HW 1.028-1.155) and the reduced to be brown or grey with some lime green spots head length varies from one-third to considerably more or markings. than one-third of the snout to vent length, the complete The head is longer than broad (average HL/HW 1.132), HL/S-V range its length equivalent to more than one-third of the snout being 0.333-0.404. The E-N/IN range is 0.611-0.818 and to vent length (average HL/S-V 0.350). The snout is the eye diameter is consistently smaller than the evenly rounded when viewed from above and projects internarial span. The average TL/S-V range is 0.522- slightly in profile. The nostrils are more lateral than 0.604. superior, their distance from the tip of the snout less than The diploid chromosome number is 26. that from the eye. The morphologically similar species Bellarana The distance between the eye and the naris is less than micromembrana (Tyler, 1963) from Madang, is most the internarial span (average E-N/IN 0.727). The canthus readily separated from the species within Ornatanura rostralis is well defined and distinctly curved. gen. nov. by its distinctive orange upper iris, versus The eye is prominent and bulges above the head, its yellowish in Ornatanura gen. nov. species. diameter greater than the eye to naris distance and less Distribution: O. leucopicturas sp. nov. is known only than the internarial span. The superior one-quarter to from hilly areas in the Central Province of New Guinea, one-third of the tympanum is hidden beneath the supra- associated with the headwaters of south-flowing tympanic fold. The diameter of the tympanum is drainages. equivalent to less than one third of the eye diameter to Etymology: The new species name is taken directly from slightly more than one-third of the eye diameter. The the Latin words “leuco picturas” which means “white vomerine teeth are in two small oblique series between markings” which accurately describes the dorsal the choanae. The tongue is almost circular and lacks a colouration of the labial region and flanks of this species. posterior indentation. ORNATANURA PARSCINEREO SP. NOV. The fingers are long and lack lateral fringes; in LSIDurn:lsid:zoobank.org:act:F8ACAFF8-E95B-483B- decreasing order of length 3>4>2>1. There is a small 8EBC-A266B291CD88 vestige of webbing between the third and fourth fingers. The terminal discs are not prominent, with the degree of Holotype: A preserved specimen at the Australian expansion varying slightly between species. Museum, Sydney, New South Wales, Australia, specimen number R.144550 collected from Utikini Village, At Waa The hind limbs are relatively long with an average TL/S-V River, Tembagapura Area, West Papua, Indonesia, ratio of 0.558. Toes in decreasing order of length Latitude -4.24 S., Longitude 137.08 E. 4>5=3>2>1. On the fifth toe, the webbing extends two- thirds up the penultimate phalanx of the fifth and to the This government-owned facility allows access to their base of the penultimate phalanx on the fourth. holdings. The skin on the dorsal surface of the head, body and Diagnosis: The three species O. parscinereo sp. nov., O. limbs is either smooth with numerous small, spaced parsviridis sp. nov. and O. leucopicturas sp. nov. are all apart, scattered tubercles all over, ranging down to similar in most respects to O. modica (Tyler, 1968) and smooth all over but for a few small and only slightly like O. modica conform to the genus diagnosis for developed tubercles on the scapular region. Ornatanura gen. nov.. The throat, abdomen and lower surface of the thighs are The four preceding species are separated from one very weakly granular. There is a row of small tubercles another as follows. extending posteriorly from the angle of the jaws, and a The species Ornatanura parsviridis sp. nov. from Madang patch of similar sized tubercles beneath the anus. is similar in most respects to O. modica (Tyler, 1968), but Of the tubercles at the angles of the jaws those anterior is readily separated from that taxon as well as O. to the tympanum are green, grey or white, whilst those parscinereo sp. nov. and O. leucopicturas sp. nov. by the posterior to it are usually white. following suite of colouration character and morphology traits: Flanks are light greenish in colour overlain with The scapular tubercles are usually green, and the lateral about 6 large black spots or blotches (sometimes surfaces of the body between the axilla and the groin are merged), in turn edged with thick white lines, dorsal one or other of being, 1/ Liberally spotted with white, 2/ surface of the hindlimbs are medium brown with indistinct Grey marks or spots on a cream background, 3/ Back dark brown to black crossbands. Upper forearms are marks in the form of large spots or blotches, circled by Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 96 Australasian Journal of Herpetology barred with white on a dark background. Small blunt species this may be reduced to be brown or grey with tubercles scattered across all parts of the dorsum and some lime green spots or markings. limbs, and only a small percentage are white tipped. The The head is longer than broad (average HL/HW 1.132), tympanum is less than a third of the diameter of the eye. its length equivalent to more than one-third of the snout By contrast O. modica from the higher elevation areas in to vent length (average HL/S-V 0.350). The snout is the region bound by the Star Mountain, West Papua and evenly rounded when viewed from above and projects the Eastern Highlands Province in Papua New Guinea, slightly in profile. The nostrils are more lateral than mainly from upper reaches of south flowing drainages, superior, their distance from the tip of the snout less than has flanks that are greenish in colour and liberally that from the eye. spotted with white. Markings on the upper forearms are The distance between the eye and the naris is less than indistinct and blotches on the upper surfaces of the hind the internarial span (average E-N/IN 0.727). The canthus limbs do not form indistinct or obvious bands. The skin on rostralis is well defined and distinctly curved. the dorsal surface of the head, body and limbs is smooth The eye is prominent and bulges above the head, its but for a few small and only slightly developed tubercles diameter greater than the eye to naris distance and less on the scapular region. The tympanum is slightly more than the internarial span. The superior one-quarter to than a third of the diameter of the eye. one-third of the tympanum is hidden beneath the supra- The species O. parscinereo sp. nov. from Utikini Village, tympanic fold. The diameter of the tympanum is At Waa River in the Tembagapura Area, West Papua is equivalent to less than one third of the eye diameter to separated from O. parsviridis sp. nov., O. modica and O. slightly more than one-third of the eye diameter. The leucopicturas by having a dorsum with numerous vomerine teeth are in two small oblique series between scattered and large tubercles that are the same colour as the choanae. The tongue is almost circular and lacks a the skin surrounding them. There are no distinctive posterior indentation. markings on upper surfaces of limbs, green on the The fingers are long and lack lateral fringes; in dorsum is restricted to the upper surface of the head decreasing order of length 3>4>2>1. There is a small anterior to the eyes, which also has some dark peppering vestige of webbing between the third and fourth fingers. on it and similarly peppered green on the upper surfaces The terminal discs are not prominent, with the degree of of the arms, save for scattered green spots elsewhere. expansion varying slightly between species. The tympanum is less than a third of the diameter of the The hind limbs are relatively long with an average TL/S-V eye. ratio of 0.558. Toes in decreasing order of length The species O. leucopicturas from central Province, 4>5=3>2>1. On the fifth toe, the webbing extends two- Papua New Guinea is readily separated from thirds up the penultimate phalanx of the fifth and to the O. parsviridis sp. nov., O. parscinereo sp. nov., O. modica base of the penultimate phalanx on the fourth. and O. leucopicturas by having a brownish grey dosum The skin on the dorsal surface of the head, body and with numerous scattered black spots, as well as limbs is either smooth with numerous small, spaced numerous areas of green, generally arranged in one or apart, scattered tubercles all over, ranging down to more irregular blotches forming a sort of longitudinal smooth all over but for a few small and only slightly stripe, rich immaculate green on the upper snout anterior developed tubercles on the scapular region. to the eyes; large black spots and blotches from the The throat, abdomen and lower surface of the thighs are dorsum spread to the flanks, where their size generally very weakly granular. There is a row of small tubercles increases but number decreases. There are scattered extending posteriorly from the angle of the jaws, and a small tubercles all over the body, with a noticeable patch of similar sized tubercles beneath the anus. general absence on the head, save for an exceptionally Of the tubercles at the angles of the jaws those anterior large series of tubercles above each eye. Most are white to the tympanum are green, grey or white, whilst those or light tipped. posterior to it are usually white. O. parsviridis sp. nov. in life is depicted in Menzies (2006) The scapular tubercles are usually green, and the lateral on plate 75. surfaces of the body between the axilla and the groin are The four preceding species, being the entirety of the one or other of being 1/ Liberally spotted with white, 2/ genus Hopviridi gen. nov. are readily separated from all Grey marks or spots on a cream background, 3/ Back other Australasian (Australian and New Guinea) Tree marks in the form of large spots or blotches, circled by Frogs (Pelodryadidae) by the following suite of white and on an otherwise light lime green background. characters: These species are characterised by their The ventral surfaces are creamish with less dense small size (males 23.4-30.0 mm, females 27.4-35.4 mm), stippling than appears on the dorsum and although moderate to relatively long limbs (average TL/S-V 0.522- dense stippling occurs in some specimens. 0.604) and broadly spaced nares (average E-N/IN 0.611- 0.818). The colouration is highly variable. The greatest density of groups of chromatophores on the ventral surface appear on the throat and particularly The dorsal surface of the head, body and limbs may towards the labial margins, with more on the upper than range from light sandy grey, or brown to dark grey and is lower margin and greatest intensity between eye and ear, darkened by the presence of very dense, minute, dark although the exact configuration varies between brown to black stippling. The anterior portion of the head specimens and species. bordered laterally by the canthus rostralis, and posteriorly to a line between the anterior portions of the upper Males vary in size from 23.4 mm to 30.0 mm and females eyelids is an immaculate pale green, although in some from 27.4-35.4mm. The head length is consistently

longer than broad (average HL/HW 1.028-1.155) and the but for a few small and only slightly developed tubercles head length varies from one-third to considerably more on the scapular region. The tympanum is slightly more than one-third of the snout to vent length, the complete than a third of the diameter of the eye. HL/S-V range The species O. parscinereo sp. nov. from Utikini Village, being 0.333-0.404. The E-N/IN range is 0.611-0.818 and At Waa River in the Tembagapura Area, West Papua is the eye diameter is consistently smaller than the separated from O. parsviridis sp. nov., O. modica and O. internarial span. The average TL/S-V range is 0.522- leucopicturas by having a dorsum with numerous 0.604. scattered and large tubercles that are the same colour as The diploid chromosome number is 26. the skin surrounding them. There are no distinctive The morphologically similar species Bellarana markings on upper surfaces of limbs, green on the micromembrana (Tyler, 1963) from Madang, is most dorsum is restricted to the upper surface of the head readily separated from the species within Ornatanura anterior to the eyes, which also has some dark peppering gen. nov. by its distinctive orange upper iris, versus on it and similarly peppered green on the upper surfaces yellowish in Ornatanura gen. nov. species. of the arms, save for scattered green spots elsewhere. Distribution: O. parscinereo sp. nov. is known only from The tympanum is less than a third of the diameter of the hilly areas west of the headwaters of the Lorentz River in eye. West Papua, Indonesia. The species O. leucopicturas from central Province, Etymology: The new species name is taken directly from Papua New Guinea is readily separated from the Latin words “pars cinereo” which means “partly grey” O. parsviridis sp. nov., O. parscinereo sp. nov., O. modica which accurately describes the dorsal colouration of this and O. leucopicturas by having a brownish grey dosum species. with numerous scattered black spots, as well as ORNATANURA PARSVIRIDUS SP. NOV. numerous areas of green, generally arranged in one or more irregular blotches forming a sort of longitudinal LSIDurn:lsid:zoobank.org:act:D0BEEEDF-C48E-4A81- stripe, rich immaculate green on the upper snout anterior BC95-DDB678CC7DE7 to the eyes; large black spots and blotches from the Holotype: A preserved specimen at the Australian dorsum spread to the flanks, where their size generally Museum, Sydney, New South Wales, Australia, specimen increases but number decreases. There are scattered number R.31051 collected from Wau, on the Eddie Creek small tubercles all over the body, with a noticeable Road, Morobe Province, Papua New Guinea, Latitude - general absence on the head, save for an exceptionally 7.24 S., Longitude 146.40 E. large series of tubercles above each eye. Most are white This government-owned facility allows access to their or light tipped. holdings. O. parsviridis sp. nov. in life is depicted in Menzies (2006) Diagnosis: The three species O. parsviridis sp. nov., O. on plate 75. parscinereo sp. nov. and O. leucopicturas sp. nov. are all The four preceding species, being the entirety of the similar in most respects to O. modica (Tyler, 1968) and genus Hopviridi gen. nov. are readily separated from all like O. modica conform to the genus diagnosis for other Australasian (Australian and New Guinea) Tree Ornatanura gen. nov.. Frogs (Pelodryadidae) by the following suite of The four preceding species are separated from one characters: These species are characterised by their another as follows. small size (males 23.4-30.0 mm, females 27.4-35.4 mm), The species Ornatanura parsviridis sp. nov. from Madang moderate to relatively long limbs (average TL/S-V 0.522- is similar in most respects to O. modica (Tyler, 1968), but 0.604) and broadly spaced nares (average E-N/IN 0.611- is readily separated from that taxon as well as O. 0.818). The colouration is highly variable. parscinereo sp. nov. and O. leucopicturas sp. nov. by the The dorsal surface of the head, body and limbs may following suite of colouration character and morphology range from light sandy grey, or brown to dark grey and is traits: Flanks are light greenish in colour overlain with darkened by the presence of very dense, minute, dark about 6 large black spots or blotches (sometimes brown to black stippling. The anterior portion of the head merged), in turn edged with thick white lines, dorsal bordered laterally by the canthus rostralis, and posteriorly surface of the hindlimbs are medium brown with indistinct to a line between the anterior portions of the upper dark brown to black crossbands. Upper forearms are eyelids is an immaculate pale green, although in some barred with white on a dark background. Small blunt species this may be reduced to be brown or grey with tubercles scattered across all parts of the dorsum and some lime green spots or markings. limbs, and only a small percentage are white tipped. The The head is longer than broad (average HL/HW 1.132), tympanum is less than a third of the diameter of the eye. its length equivalent to more than one-third of the snout By contrast O. modica from the higher elevation areas in to vent length (average HL/S-V 0.350). The snout is the region bound by the Star Mountain, West Papua and evenly rounded when viewed from above and projects the Eastern Highlands Province in Papua New Guinea, slightly in profile. The nostrils are more lateral than mainly from upper reaches of south flowing drainages, superior, their distance from the tip of the snout less than has flanks that are greenish in colour and liberally that from the eye. spotted with white. Markings on the upper forearms are The distance between the eye and the naris is less than indistinct and blotches on the upper surfaces of the hind the internarial span (average E-N/IN 0.727). The canthus limbs do not form indistinct or obvious bands. The skin on rostralis is well defined and distinctly curved. the dorsal surface of the head, body and limbs is smooth The eye is prominent and bulges above the head, its

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 98 Australasian Journal of Herpetology diameter greater than the eye to naris distance and less yellowish in Ornatanura gen. nov. species. than the internarial span. The superior one-quarter to Distribution: O. parsviridis sp. nov. is presently only one-third of the tympanum is hidden beneath the supra- known from Madang, Papua New Guinea. tympanic fold. The diameter of the tympanum is Etymology: The new species name is taken directly from equivalent to less than one third of the eye diameter to the Latin words “pars” meaning some, or partly and the slightly more than one-third of the eye diameter. The word “viridis” which means green. This reflects the reality vomerine teeth are in two small oblique series between that most specimens of these frogs have some green in the choanae. The tongue is almost circular and lacks a their colour, but they are never wholly green. posterior indentation. NASUSCUSPIS GEN. NOV. The fingers are long and lack lateral fringes; in LSIDurn:lsid:zoobank.org:act:ABB692F7-77D0-47C0- decreasing order of length 3>4>2>1. There is a small 86BE-9EC2A41BEC8A vestige of webbing between the third and fourth fingers. Type species: Hyla prora Menzies, 1969. The terminal discs are not prominent, with the degree of expansion varying slightly between species. Diagnosis: Species within the genus Nasuscuspis gen. nov. are separated from all other Australasian (Australian The hind limbs are relatively long with an average TL/S-V and New Guinea) Tree Frogs (Pelodryadidae) by the ratio of 0.558. Toes in decreasing order of length following suite of characters: They are montane species, 4>5=3>2>1. On the fifth toe, the webbing extends two- growing to a maximum of 50 mm snout vent length in thirds up the penultimate phalanx of the fifth and to the females and males to 45 mm. The dorsum is a mixture of base of the penultimate phalanx on the fourth. greys, greenish greys and ochres. The fingers and toes The skin on the dorsal surface of the head, body and are extensively webbed. There are highly developed limbs is either smooth with numerous small, spaced dermal appendages including crenulated ridges on the apart, scattered tubercles all over, ranging down to posterior surfaces of the radius and tarsus, prominent smooth all over but for a few small and only slightly dermal folds above the vent; a row of very large tubercles developed tubercles on the scapular region. on the undersurface of the mandible and an elongate The throat, abdomen and lower surface of the thighs are rostral, dermal spike always in males and females very weakly granular. There is a row of small tubercles although in some reduced form in females of some extending posteriorly from the angle of the jaws, and a species. patch of similar sized tubercles beneath the anus. Duellman et al. (2016) found that species in the genus Of the tubercles at the angles of the jaws those anterior Nasuscuspis gen. nov. diverged from nearest living to the tympanum are green, grey or white, whilst those relatives 13.4 MYA, being members of the genus posterior to it are usually white. Moechaeanura gen. nov.. In turn both these genera as a The scapular tubercles are usually green, and the lateral pair diverged from the morphologically similar genera surfaces of the body between the axilla and the groin are Incertanura gen. nov., Inlustanura subgen. nov. and one or other of being 1/ Liberally spotted with white, 2/ Variabilanura gen. nov. 17.8 MYA and with each of the Grey marks or spots on a cream background, 3/ Back latter three genera diverging from one another at least marks in the form of large spots or blotches, circled by 13.4 MYA. white and on an otherwise light lime green background. The species name “Litoria rostandi Kraus, 2007” is an The ventral surfaces are creamish with less dense illegally coined junior synonym of “Litoria hilli Hiaso and stippling than appears on the dorsum and although Richards, 2006 improperly created by the serial dense stippling occurs in some specimens. taxonomic vandal Fred Kraus, best known for scamming The greatest density of groups of chromatophores on the millions of dollars in US Government research grants that ventral surface appear on the throat and particularly he uses for personal self-gratification and then justifies towards the labial margins, with more on the upper than via the repeated stealing of work from other others in is lower margin and greatest intensity between eye and ear, acts of taxonomic vandalism. although the exact configuration varies between While US Government authorites are aware of his highly specimens and species. illegal activities, he continues (in 2020) to operate with Males vary in size from 23.4 mm to 30.0 mm and females corrupt protection from within the US government, giving from 27.4-35.4mm. The head length is consistently him immunity from prosecution in the same way that longer than broad (average HL/HW 1.028-1.155) and the police officers in the USA can kill unarmed black people head length varies from one-third to considerably more without risk of being charged and convicted fo such than one-third of the snout to vent length, the complete crimes. HL/S-V range Distribution: Nasuscuspis gen. nov. species are known being 0.333-0.404. The E-N/IN range is 0.611-0.818 and from scattered highland locations in various parts of the eye diameter is consistently smaller than the Papua New Guinea and Irian Jaya, Indonesia. internarial span. The average TL/S-V range is 0.522- Etymology: In Latin, the genus name “Nasuscuspis” 0.604. means “nose pointed” as fits the case for adult males and The diploid chromosome number is 26. many adult females in the genus. The morphologically similar species Bellarana Content: Nasuscuspis prora (Menzies, 1969) (type micromembrana (Tyler, 1963) from Madang, is most species); N. chrisdahli (Richards, 2007); N. readily separated from the species within Ornatanura humboldtorum (Günther, 2006); N. hilli (Hiaso and gen. nov. by its distinctive orange upper iris, versus Richards, 2006).

ROTUNDAURA GEN. NOV. much smaller adult size of less than 25 mm adult body LSIDurn:lsid:zoobank.org:act:264E5827-AA91-421E- length. 91D2-F56901121E9E Species of Rotundaura gen. nov. are separated from the Type species: Hyla jeudii Werner, 1901. genus Summaviridis gen. nov. by the tympanum being Diagnosis: Known only from the type specimen collected fully exposed and round, versus the upper surface being in the north of New Guinea, this species (and genus) do cut at the rear by a well-defined and prominent skin fold not conform to any other in New Guinea or Australia and forming a straight line and a significantly blunter snout in is therefore placed in a monotypic genus Rotundaura Rotundaura gen. nov. versus pointed, when both are gen. nov.. viewed in profile side-on. Rotundaura gen. nov. is separated from all other Distribution: The holotype and sole known Australasian (Australian and New Guinea) Tree Frogs representative of this genus is believed to be from (Pelodryadidae) by the following suite of characters: northern New Guinea somewhere in the vicinity of A very high E-N/IN ratio (1.435), short limbs with a low Madang (Tyler, 1978). TL/S-V ratio (0.482), wholly unwebbed fingers and Etymology: The name Rotundaura is derived from Latin incompletely webbed toes characterise this species. The and refers to the visible round ear (tympanum) in the sole head is long and flattened and longer than broad (HL/HW species in the genus. 1.073), its length more than one-third of the snout to vent Content: Rotundaura jeudii (Werner, 1901). length (HL/S-V 0.355). The snout is rather prominent, VARIABILANURA GEN. NOV. gently rounded when viewed from above and strongly LSIDurn:lsid:zoobank.org:act:573588FF-F254-4979- rounded in profile. The nostrils are more superior than AD95-1D9F134A7B34 lateral, their distance from the end of the snout about Type species: Hyla iris Tyler, 1962. one-half that from the eye and separated from each other Diagnosis: The genus Variabilanura gen. nov. are by a distance which is equivalent to approximately two- separated from all other Australasian (Australian and thirds of the eye to naris distance (E-N/IN 1.435). The New Guinea) Tree Frogs (Pelodryadidae) by the following canthus rostralis is straight and inconspicuous and the suite of characters: They are small montane species loreal region oblique. The eye is small, its diameter less (males 24.3-34.1 mm; females 30.1-43 mm in SV). In life than the eye to naris distance. The tympanum is visible, they are green, usually with yellow spots or markings (in its diameter equivalent to two-thirds of the eye diameter preservative the dorsal surface is blue, usually with and separated from the eye by a distance equivalent to darker and occasionally lighter markings). There is approximately one-half its own diameter. The vomerine usually a deep violet, orange, yellow or red patch in the teeth are in two obliquely oval series in juxtaposition on groin and similarly coloured markings occur on the axilla, the midline between the small, obliquely oval choanae. posterior surface of the thighs, tibia and tarsus. The tongue is small, triangular and very feebly indented The head is as broad as long or longer than broad (HL/ on its posterior border. The fingers are long, slender and HW 0.910-1.192), its length less than or greater than unwebbed. The terminal discs are large and the sub- one-third of the snout to vent length (HL/S-V 0.328- articular tubercles prominent. The fingers in decreasing 0.384). The shape of the snout is highly variable, being order of length 3>4>2>1. The hind limbs are short with a from prominent to inconspicuous and strongly or only TL/S-V ratio of 0.482. The toes are incompletely webbed, slightly rounded when viewed from above and in profile. the webbing reaching the base of the penultimate The nostrils are lateral in specimens with prominent phalanx on the fourth toe and half-way up the penultimate snouts and more lateral than superior in specimens with phalanx of the fifth. Toes in decreasing order of length inconspicuous snouts. The nares are very much nearer 4>5=3>2>1. The skin on the dorsal surfaces of the head the eye than the tip of the snout in the latter individuals and body is smooth but for a few flattened tubercles on and almost equidistant in the former. The distance the back. The throat is smooth and the chest, abdomen between the eye and the naris is less than the internarial and back of the thighs are coarsely granular. There is an span (E-N/IN 0.667-0.965). The canthus rostralis is inconspicuous supratympanic fold barely obscuring any curved and slightly or well defined. The eye is large and of the tympanum and a pronounced skin fold across the prominent, its diameter greater than the distance chest. There is neither a tarsal ridge nor a dermal separating it from the nostril. The tympanum is covered appendage on the heel. Dimensions: Of gravid female with skin, its diameter equivalent to from one-third to holotype are snout-vent 33.2 mm; tibia length 15.8 mm; slightly less than one-half the eye diameter. Vomerine head length 11.8 mm; head width 11.0 mm; eye to naris teeth are present in the majority of specimens and distance 3.3 mm; internarial span 2.3 mm; eye diameter confined to small circular series on slight elevations 3.1 mm; tympanum diameter 2.0 mm. The dorsal between the choanae. The tongue is broadly cordiform surfaces of the head and body and limbs are a very pale with a slightly indented posterior margin. The fingers are brown with indistinct darker and lighter marbling. Marbling short and equipped with very slight lateral fringes; in is present and most conspicuous on the dorsal surface of decreasing order of length 3>4>2>1. The webbing the thigh and tibia. reaches the sub-articular tubercle at the base of the The ventral surfaces of the body and limbs are dull cream penultimate phalanx on the fourth finger. The terminal with an irregular brown patch on the throat (modified from discs are prominent. The hind limbs are variable with a Tyler 1978). TL/S-V ratio of 0.485-0.587. Toes in decreasing order of Specimens of the morphologically similar Llewellynura length 4>5>3>2>1. The webbing between the toes Wells and Wellington, 1985 are readily separated by their reaches the base of the disc of all toes, except the fourth

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 100 Australasian Journal of Herpetology where it reaches the tubercle at the base of the band on upper lip, snout to axilla; raised tubercles on fore penultimate phalanx. The skin on the dorsal surface of limb white; ventral surfaces pure white. the head, body and limbs is minutely roughened. The According to Duellman et al. (2016), the species within throat and chest are slightly granular. There are a row of Variabilanura gen. nov. diverged from their nearest living tubercles on the posterior surface of the forearm, and relatives 13.4 MYA, being the newly named (within this conspicuous tubercles below the anus. Femoral tubercles paper) genus Inlustanura gen. nov. and are at least 21.5 are usually present. The supra-tympanic fold is MYA diverged from nearest living relatives in previously inconspicuous. In preservative and in life, dull orange named genera. patches may extend on to the dorsum in the axillary According to Duellman et al. (2016), the subgenus region. Sudesanura subgen. nov. diverged from the nominate The lateral surfaces of the body are frequently heavily genus 12 MYA. pigmented with black or deep yellow (in life, being violet The type species for Sudesanura subgen. nov., being in preservative), marked with large white patches. There “Litoria havina Menzies, 1993” as currently understood by is invariably a yellow, orange, red or violet patch in the most authors, is in fact a number of morphologically groin (in life) and the same colouration may be present in similar and closely related species. the axilla and on the posterior surface of the tibia and Distribution: New Guinea mainly in the mountainous tarsus. There is a white patch beneath the eye extending areas and adjacent foothills. to the angle of the jaws or on the lateral surfaces of the Etymology: Species within this genus can be quite body. The ventral surface of the body is cream to white variable in colouration, including depending on time of (white in subgenus Sudesanura subgen. nov.), with or day and with variation in body temperature and they are without sparse blue stippling on the throat near the angle frogs, so therefore the genus name Variabilanura reflects of the jaws. The lower surface of the hind limbs is both. creamish yellow or dull yellow. These species are often beautifully pigmented in life being marked with cream, Content: Variabilanura iris (Tyler, 1962) (type species); orange, yellow, green, violet and black (Tyler, 1962). V. tomcottoni sp. nov.; V. havina (Menzies, 1993); V. majikthise (Johnston and Richards, 1994); L. mucro The ovum is pale green in colour and has a diameter of (Menzies, 1993); L. mareku (Günther 2008); L. ollauro approximately 2.5 mm. The ova are laid in groups of from (Menzies, 1993). 4 to 37 (mean 14) on the leaves of trees overhanging water, and around the stems of vegetation at the edge of SUDESANURA SUBGEN. NOV. the water. LSIDurn:lsid:zoobank.org:act:4A1CB6B5-C8FF-47EC- The ova are surrounded by a very large mass of clear B94D-D22439DB3242 albumen. A period of approximately fourteen days is Type species: Litoria havina Menzies, 1993. spent within the spawn clump, and the tadpole emerges Diagnosis: Species in the subgenus Sudesanura possessing internal gills and capable of coordinated subgen. nov. (type species “Litoria havina Menzies, movements. The mouth is anterior in position, and 1993”) are separated from those in the nominate surrounded by a band of papillae on the inferior and subgenus (Variabilanura subgen. nov.) by having a red lateral borders. There are two upper and three lower rows patch in the groin, versus a deep violet, orange or yellow of labial teeth. patch in the groin in the nominate subgenus. Species in the subgenus Sudesanura subgen. nov. (type The genus Variabilanura gen. nov. are separated from all species “Litoria havina Menzies, 1993”) are separated other Australasian (Australian and New Guinea) Tree from those in the nominate subgenus by having a red Frogs (Pelodryadidae) by the following suite of patch in the groin, versus a deep violet, orange or yellow characters: They are small montane species (males 24.3- patch in the groin in the nominate subgenus. 34.1 mm; females 30.1-43 mm). In life they are green, Species within Sudesanura subgen. nov. are further usually with yellow spots or markings (in preservative the defined as follows: Small (SV max. 30.05 mm); head dorsal surface is blue, usually with darker and narrow (average HL/HW 1.08); Head always longer than occasionally lighter markings). There is usually a deep wide (average HL/HW>1.0) except in females, which lack violet, orange, yellow or red patch in the groin and the rostral spike, where HL=HW; canthus rounded, similarly coloured markings occur on the axilla, posterior concave, lores oblique, nostrils more or less lateral, surface of the thighs, tibia and tarsus. The head is as widely placed (average EN/IN 0.63 in males and EN/IN broad as long or longer than broad (HL/HW 0.910-1.192), 0.54 in females); snout with a prominent pointed rostral its length less than or greater than one-third of the snout spike; vomerine teeth absent; eye large (EY/SV 0.11); to vent length (HL/S-V 0.328-0.384). The shape of the tympanum visible, upper margin covered by skin fold. snout is highly variable, being from prominent to Fore limb with indistinct row of raised tubercles down inconspicuous and strongly or only slightly rounded when outer side; hind limb without heel lappets or other dermal viewed from above and in profile. The nostrils are lateral appendages; fingers half-webbed, toes fully webbed; in specimens with prominent snouts and more lateral subarticular tubercles poorly developed. Legs always than superior in specimens with inconspicuous snouts. long (average TL/SV>0.54). Dorsum usually immaculate, The nares are very much nearer the eye than the tip of bright pale green, yellow or fawn brown reduced to a very the snout in the latter individuals and almost equidistant narrow band on the thighs and ceasing at ankle and in the former. wrist, leaving hands and feet virtually colourless; The distance between the eye and the naris is less than concealed thighs and axillae bright cherry-red; white the internarial span (E-N/IN 0.667-0.965). The canthus

rostralis is curved and slightly or well defined. The eye is defined as follows: Small (SV max. 30.05 mm); head large and prominent, its diameter greater than the narrow (average HL/HW 1.08); Head always longer than distance separating it from the nostril. The tympanum is wide (average HL/HW>1.0) except in females, which lack covered with skin, its diameter equivalent to from one- the rostral spike, where HL=HW; canthus rounded, third to slightly less than one-half the eye diameter. concave, lores oblique, nostrils more or less lateral, Vomerine teeth are present in the majority of specimens, widely placed (average EN/IN 0.63 in males and EN/IN and confined to small circular series on slight elevations 0.54 in females); snout with a prominent pointed rostral between the choanae. The tongue is broadly cordiform spike; vomerine teeth absent; eye large (EY/SV 0.11); with a slightly indented posterior margin. The fingers are tympanum visible, upper margin covered by skin fold. short and equipped with very slight lateral fringes; in Fore limb with indistinct row of raised tubercles down decreasing order of length 3>4>2>1. The webbing outer side; hind limb without heel lappets or other dermal reaches the sub-articular tubercle at the base of the appendages; fingers half-webbed, toes fully webbed; penultimate phalanx on the fourth finger. The terminal subarticular tubercles poorly developed. Legs always discs are prominent. The hind limbs are variable with a long (average TL/SV>0.54). Dorsum usually immaculate, TL/S-V ratio of 0.485-0.587. Toes in decreasing order of bright pale green, yellow or fawn brown reduced to a very length 4>5>3>2>1. The webbing between the toes narrow band on the thighs and ceasing at ankle and reaches the base of the disc of all toes, except the fourth wrist, leaving hands and feet virtually colourless; where it reaches the tubercle at the base of the concealed thighs and axillae bright cherry-red; white penultimate phalanx. The skin on the dorsal surface of band on upper lip, snout to axilla; raised tubercles on fore the head, body and limbs is minutely roughened. The limb white; ventral surfaces pure white. throat and chest are slightly granular. There are a row of According to Duellman et al. (2016), the species within tubercles on the posterior surface of the forearm, and Variabilanura gen. nov. diverged from their nearest living conspicuous tubercles below the anus. Femoral tubercles relatives 13.4 MYA, being the newly named (within this are usually present. The supra-tympanic fold is paper) genus Inlustanura gen. nov. and are at least 21.5 inconspicuous. In preservative and in life, dull orange MYA diverged from nearest living relatives in previously patches may extend on to the dorsum in the axillary named genera. region. According to Duellman et al. (2016), the subgenus The lateral surfaces of the body are frequently heavily Sudesanura subgen. nov. diverged from the nominate pigmented with black or deep yellow (in life, being violet genus 12 MYA. in preservative), marked with large white patches. There The type species for Sudesanura subgen. nov., being is invariably a yellow, orange, red or violet patch in the “Litoria havina Menzies, 1993” as currently understood by groin (in life) and the same colouration may be present in most authors, is in fact a number of morphologically the axilla and on the posterior surface of the tibia and similar and closely related species. tarsus. There is a white patch beneath the eye extending From photos of the various regional forms alone this to the angle of the jaws or on the lateral surfaces of the contention is clearly supported. body. The ventral surface of the body is cream to white Due to the extreme urgency in which species of frog need (white in subgenus Sudesanura subgen. nov.), with or to be formally identified and named in the New Guinea without sparse blue stippling on the throat near the angle region, as in before human population growth and of the jaws. The lower surface of the hind limbs is associated issues cause their demise and extinction, I creamish yellow or dull yellow. These species are often seriously considered naming two outlier forms of putative beautifully pigmented in life being marked with cream, V. havina as new species herein on the basis of orange, yellow, green, violet and black (Tyler, 1962). previously published photos. The ovum is pale green in colour and has a diameter of Heeding the advice of Krell and Marshall (2017), I have approximately 2.5 mm. refrained from doing so. The ova are laid in groups of from 4 to 37 (mean 14) on However I take this opportunity to urge those who do the leaves of trees overhanging water, and around the have access to specimens of these frogs, to immediately stems of vegetation at the edge of the water. obtain specimens and lodge them with a State or The ova are surrounded by a very large mass of clear National Museum and then formally describe them so albumen. that proper conservation management plans can be A period of approximately fourteen days is spent within enacted. the spawn clump, and the tadpole emerges possessing Refer also to the statements in Hoser (2019a, 2019b). internal gills and capable of coordinated movements. The Distribution: Sudesanura subgen. nov. species are mouth is anterior in position, and surrounded by a band found in New Guinea mainly in the mountainous areas of papillae on the inferior and lateral borders. and adjacent foothills. There are two upper and three lower rows of labial teeth. Etymology: Sudes in Latin means “spike”, while “anura” As already mentioned, species in the subgenus means frog and therefore the new genus name Sudesanura subgen. nov. (type species “Litoria havina Sudesanura literally means spike frog, in reference to the Menzies, 1993”) are separated from those in the rostral appendage in males. nominate subgenus by having a red patch in the groin, Content: Variabilanura (Sudesanura) havina (Menzies, versus a deep violet, orange or yellow patch in the groin 1993) (type species) (treated herein as monotypic, but in in the nominate subgenus. fact a group of species). Species within Sudesanura subgen. nov. are further

VARIABILANURA (VARIABILANURA) TOMCOTTONI body between dorsal green and ventral white. SP. NOV. The genus Variabilanura gen. nov. are separated from all LSIDurn:lsid:zoobank.org:act:BB6A8CEF-84A4-40CB- other Australasian (Australian and New Guinea) Tree 9679-626F97E9DCFD Frogs (Pelodryadidae) by the following suite of Holotype: A preserved specimen at the Museum of characters: They are small montane species (males 24.3- Natural History, London, United Kingdom, specimen 34.1 mm; females 30.1-43 mm). In life they are green, number 1980.630-634 collected at the banks of Lake usually with yellow spots or markings (in preservative the Trist, Kuper Range, Morobe Province, Papua New dorsal surface is blue, usually with darker and Guinea. occasionally lighter markings). There is usually a deep This facility allows access to its holdings. violet, orange, yellow or red patch in the groin and Paratype: A preserved specimen at the Museum of similarly coloured markings occur on the axilla, posterior Natural History, London, United Kingdom, specimen surface of the thighs, tibia and tarsus. The head is as number 1980.635-637 collected at the banks of Lake broad as long or longer than broad (HL/HW 0.910-1.192), Trist, Kuper Range, Morobe Province, Papua New its length less than or greater than one-third of the snout Guinea. to vent length (HL/S-V 0.328-0.384). The shape of the snout is highly variable, being from prominent to Diagnosis: Variabilanura tomcottoni sp. nov. has until inconspicuous and strongly or only slightly rounded when now been treated as an eastern population of V. iris viewed from above and in profile. The nostrils are lateral (Tyler, 1962), but can be separated from that species by in specimens with prominent snouts and more lateral having generally unmarked and translucent toes in adults than superior in specimens with inconspicuous snouts. and a slightly larger tympanum, versus marked toes in The nares are very much nearer the eye than the tip of adult V. iris and more reduced tympanum. Outside of the snout in the latter individuals and almost equidistant these differences, there are general differences between in the former. The distance between the eye and the naris the forms, including a greater preponderance of mainly is less than the internarial span (E-N/IN 0.667-0.965). brown and green coloured specimens in V. tomcottoni sp. The canthus rostralis is curved and slightly or well nov. versus mainly with green and not necessarily brown defined. The eye is large and prominent, its diameter in V. iris, but in both species the variation between greater than the distance separating it from the nostril. specimens and various colour morphs within a single site The tympanum is covered with skin, its diameter is significant. equivalent to from one-third to slightly less than one-half That both populations are also reproductively isolated the eye diameter. Vomerine teeth are present in the across the Wau-Bulolo area and likely to have been so majority of specimens and confined to small circular for an extended geological time, gives me no hesitation in series on slight elevations between the choanae. The describing V. tomcottoni sp. nov. as a new species being tongue is broadly cordiform with a slightly indented different to the type form of V. iris from the south-side of posterior margin. The fingers are short and equipped with the main central cordillera. very slight lateral fringes; in decreasing order of length Both V. tomcottoni sp. nov. and V. iris are readily 3>4>2>1. The webbing reaches the sub-articular tubercle separated from all other species in the genus at the base of the penultimate phalanx on the fourth Variabilanura gen. nov. by the following unique suite of finger. The terminal discs are prominent. The hind limbs characters: Small species (male average SV 24-36 mm, are variable with a TL/S-V ratio of 0.485-0.587. Toes in female average SV 34-43 mm); legs short to long (TL/SV decreasing order of length 4>5>3>2>1. The webbing 0.46-0.63); head about as broad as long (HL/HW 0.91- between the toes reaches the base of the disc of all toes, 1.06); nostrils moderately widely spaced (EN/IN 0.77- except the fourth where it reaches the tubercle at the 0.97); snout truncate to slightly pointed in profile; eye base of the penultimate phalanx. The skin on the dorsal large; tympanum usually visible but small. surface of the head, body and limbs is minutely Dorsum is green, green-and-brown or entirely brown and roughened. The throat and chest are slightly granular. always mottled with black or darker green. Occasionally There are a row of tubercles on the posterior surface of the body is with scattered white spots and a white lateral the forearm and conspicuous tubercles below the anus. stripe commencing below the eye. In life it appears that Femoral tubercles are usually present. The supra- no two examples are exactly alike but they can be divided tympanic fold is inconspicuous. In preservative and in life, into four main morphotypes: (a) dark green, with indistinct dull orange patches may extend on to the dorsum in the black reticulation; (b) pale green, with distinct fine or axillary region. coarse black reticulation; (c) pale green, with distinct fine The lateral surfaces of the body are frequently heavily or coarse black reticulation, but also the green has pigmented with black or deep yellow (in life, being violet patches of brown; and (d) mostly brown, with distinct fine in preservative), marked with large white patches. There or coarse black reticulation. The ventral surface is white, is invariably a yellow, orange, red or violet patch in the tending to creamy posteriorily and on the hind limbs. The groin (in life) and the same colouration may be present in concealed parts of the thighs are basically purple- the axilla and on the posterior surface of the tibia and blotched white, or white and red, or white and pale blue, tarsus. There is a white patch beneath the eye extending these colours sometimes meeting below abdomen and to the angle of the jaws or on the lateral surfaces of the thighs. body. The ventral surface of the body is cream to white There is invariably a violet patch in the groin, and often in (white in subgenus Sudesanura subgen. nov.), with or the axilla as well, and this may be interrupted with blue or without sparse blue stippling on the throat near the angle white spots or the patch may extend along the side of the of the jaws. The lower surface of the hind limbs is

creamish yellow or dull yellow. These species are often shows that let people hold the animals and a significant beautifully pigmented in life being marked with cream, amount of other critically important wildlife conservation orange, yellow, green, violet and black (Tyler, 1962). work. Another species of tree frog named in honour of The ovum is pale green in colour and has a diameter of Tom Cotton has the nomen “cottoni”. approximately 2.5 mm. The ova are laid in groups of from DRYMONTANTINA SUBTRIBE NOV. 4 to 37 (mean 14) on the leaves of trees overhanging DRYOMANTIS PETERS, 1882 water, and around the stems of vegetation at the edge of Type species: Drymomantis fallax Peters, 1882. the water. The ova are surrounded by a very large mass Diagnosis: Frogs in the genus Drymomantis Peters, of clear albumen. A period of approximately fourteen 1882 are all readily separated from all other Australasian days is spent within the spawn clump, and the tadpole Tree Frogs (Pelodryadidae) by the following unique suite emerges possessing internal gills and capable of of characters: These are small elongated and agile frogs coordinated movements. The mouth is anterior in averaging about 25 mm in body length with moderately position, and surrounded by a band of papillae on the pointed snouts, varying somewhat between species; inferior and lateral borders. There are two upper and brown head streak present or absent; dorsal surface is three lower rows of labial teeth. uniform green or fawn, sometimes with darker flecks and Species in the subgenus Sudesanura subgen. nov. (type with at most a paler vertebral zone; there is not a broad species “Litoria havina Menzies, 1993”) are separated vertebral band of bronze bordered on either side with from those in the nominate subgenus by having a red green; skin is smooth above; finely granular on the throat patch in the groin, versus a deep violet, orange or yellow and coarsely granular below; at least one strong pectoral patch in the groin in the nominate subgenus. fold, no dorsolateral fold and no tubercles above each Species within Sudesanura subgen. nov. are further eye; fingers webbed at the base and toes moderately defined as follows: Small (SV max. 30.05 mm); head webbed. A moderate oval inner and small rounded outer narrow (average HL/HW 1.08); Head always longer than metatarsal tubercle; vomerine teeth absent, or if present, wide (average HL/HW>1.0) except in females, which lack the hind edge of vomerine teeth are between the the rostral spike, where HL=HW; canthus rounded, choanae; first finger is much smaller than the second concave, lores oblique, nostrils more or less lateral, when pressed together, the top of the first finger reaching widely placed (average EN/IN 0.63 in males and EN/IN no further than the base of the disc of the second finger. 0.54 in females); snout with a prominent pointed rostral According to Duellman et al. (2016), the members of this spike; vomerine teeth absent; eye large (EY/SV 0.11); genus diverged from their nearest living relatives 21.5 tympanum visible, upper margin covered by skin fold. MYA. Fore limb with indistinct row of raised tubercles down Distribution: East coast of Australia. outer side; hind limb without heel lappets or other dermal Content: Drymomantis fallax Peters, 1882 (type appendages; fingers half-webbed, toes fully webbed; species); D. ausviridis sp. nov.; D. celantur sp. nov.; D. subarticular tubercles poorly developed. Legs always cooloolensis (Liem, 1974); D. glauerti (Copland, 1957); long (average TL/SV>0.54). Dorsum usually immaculate, D. northdtradbrokeensis sp. nov.; D. olongburensis (Liem bright pale green, yellow or fawn brown reduced to a very and Ingram, 1977). narrow band on the thighs and ceasing at ankle and wrist, leaving hands and feet virtually colourless; DRYMOMANTIS AUSVIRIDIS SP. NOV. concealed thighs and axillae bright cherry-red; white LSIDurn:lsid:zoobank.org:act:FC390EC9-13B7-4C17- band on upper lip, snout to axilla; raised tubercles on fore AFBE-AA22A6DB27AE limb white; ventral surfaces pure white. Holotype: A preserved specimen in the Queensland According to Duellman et al. (2016), the species within Museum, Brisbane, Queensland, Australia, specimen Variabilanura gen. nov. diverged from their nearest living number J54722 collected at Cairns, Queensland, relatives 13.4 MYA, being the newly named (within this Australia, Latitude 16.9186 S., Longitude 145.7781 E. paper) genus Inlustanura gen. nov. and are at least 21.5 This government-owned facility allows access to its MYA diverged from nearest living relatives in previously holdings. named genera. Paratypes: Four preserved specimens at the According to Duellman et al. (2016), the subgenus Queensland Museum, Brisbane, Queensland, Australia, Sudesanura subgen. nov. diverged from the nominate specimen numbers J54723, J54727, J54730 and J54733 genus 12 MYA. all collected at Cairns, Queensland, Australia, Latitude The type species for Sudesanura subgen. nov., being 16.9186 S., Longitide 145.7781 E. “Litoria havina Menzies, 1993” as currently understood by Diagnosis: The putative species defined as “Litoria fallax most authors, is in fact a number of morphologically (Peters, 1882)” as defined and separated from other similar and closely related species. species by Cogger (2014) and Anstis (2013) is found Distribution: V. tomcottoni sp. nov. is known only from along the coast and ranges from southern New South the type locality being the banks of Lake Trist, Kuper Wales along the east Australian coast to the northern Range, Morobe Province, Papua New Guinea. edge of the wet tropics in far north Queensland. However as far back as 1985, Wells and Wellington (1985) Etymology: The new species name “tomcottoni” is correctly resurrected from synonymy the taxon named in honour of Tom Cotton, formerly of Ringwood in Drymomantis glauerti (Copland, 1957) (in this form) from Victoria in recognition of his many years work with the the synonymy of Litoria fallax (Peters, 1882). team at Snakebusters and Australia’s best reptiles At the same time, Wells and Wellington resurrected from educating people with Australia’s only hands on reptile

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 104 Australasian Journal of Herpetology synonymy of Litoria Tschudi, 1838 the genus relevant frogs from north of Cairns along the coast to Drymomantis Peters, 1882, as proposed for the species southern New South Wales must agree with the “fallax”. conclusions of Wells and Wellington (1985) with regards To make it clear, D. fallax has a type locality of Bowen, to the relevant frogs. Mackay and Rockhampton, while D. glauerti has a type I note herein that according to Duellman et al. (2016) the locality of Colo, north-west of Sydney, New South Wales. relevant species in the genus Drymomantis Peters, 1882 Wells and Wellington (1985) have been consistently being the lineage of Drymomantis fallax Peters, 1882 and lampooned and vilified by the Wolfgang Wüster gang of close relatives, diverged from their nearest living relatives thieves ever since 1985 (see for example a recent hate assigned to other genera by 21.5 MYA, confiming the post by Wolfgang Wüster on the ICZN internet list server correct genus-level assignment by Wells and Wellington dated 6 May 2020, posted at 7:44 AM, UK time). In that (1985). post, he yet again implores people to ignore the The name Drymomantis was a creation of Wilhelm Karl taxonomy and nomenclature of Wells and Wellington Hartwich (or Hartwig) Peters, a prominent German (1985) and all other publications by them and to ignore or herpetologist from the 1800’s, not of Wells and steal from the works of all other people the Wüster gang Wellington (1985), as they only resurrected the name declare as not being within their gang. from synonymy, but the Wüster gang were blissfully Wüster and his his cohort harass and threaten others unaware of this minor detail and so forced everyone else who dare to use the Wells and Wellington taxonomy on the planet not to use the name as part of their “mafia” including so far as committing acts of violence against style attack on Wells and Wellington. them and telephone death threats to kill. See for example I however will not be cowed or intimidated by these anti- the threat to kill phone call by police-protected criminal scientists and therefore correctly assign the relevant Crystal Kniese made on 10 Feb 2019. This call was frogs to the genus Drymomantis. made from a blocked phone number, which was tape At the species level of classification, Wells and recorded and has been posted online at: http:// Wellington were also correct. www.smuggled.com/Gatt-Girlfriend-11-2-19-11-43- James and Moritz (2000), published a molecular 17_AM.m4a phylogeny for the putative species from the entire known The Wüster gang have been involved in attacks on range and confirmed the Wells and Wellington private property, including for example a failed attempt to classification. burn down this author’s research facility with an They found two major genetic lineages for the putative incendiary attack, instigation of illegal and violent police taxon, including an estimated 6 MYA divergence between raids on properties and other criminal actions. the coastal New South Wales and east Queensland frogs This activity, described by Lynn Raw of the ICZN list (11.1-12.5 per cent sequence divergence of mtDNA), server in 2020 as “Mafia” style misconduct has been thereby confirming species level division between the two successful in that most herpetologists have in the past 35 populations. years been bludgeoned into not using the Wells and Furthermore James and Moritz (2000) found that the Wellington taxonomy or nomenclature, even when it is population from north of the Burdekin Gap, north clearly correct. Queensland and Kroombit Tops south-west of Gladstone, Hal Cogger, author of numerous major works on Queensland, had populations with a 5 percent sequence Australian herpetofauna has even openly stated he often divergence (= 2.5 MYA), also warranting species-level uses the Wüster gang taxonomy and nomenclature, recognition. knowing it to be illegal, unscientific and just plain “wrong” Those two species are therefore formally recognized and so that “I don’t get getting threatening phone calls from named herein. them at strange hours of the night”, as stated by him in a The four species, until now usually treated all as recorded phone call in 2018 (author’s phone Drymomantis fallax Peters, 1882 are readily separated automatically records and archives all phone calls). from their nearest relatives and congeners, D. These actions by Wüster and his gang are not science, cooloolensis (Liem, 1974); the similar D. but rather a form of anarachy instead. northstradbrokensis sp. nov. and D. olongburensis (Liem In terms of the taxonomic actions of Wells and Wellington and Ingram, 1977) by morphology. Same applies in terms taken with regards to the putative species Drymomantis of the morphologically similar species most widely known fallax Peters, 1882, the science is clearly on the side of as “Litoria bicolor (Gray, 1842)” (e.g. Cogger, 2014), but Wells and Wellington. These two men made their herein placed in the new genus Maxinehoserranae gen. taxonomic judgement on the basis of some decades of nov.. field experience with the putative taxon from most areas it Duellman et al. (2016) found a divergence of the so- was known to occur and with good reason thought the called bicolor species group of 21.5 MYA from the genus Queensland animals were different from those on the Drymomantis making separate genus-level recognition central coast of New South Wales, as explicitly stated by an obvious judgement. them on page 5 of their paper Wells and Wellington The morphologically similar species Drymomantis fallax (1985). Peters, 1882 from coastal Queensland south of the Unlike the Wüster gang, who have never stepped foot Burdekin River, North Queensland, just into north-east into a swamp in Eastern Australia to search for or inspect New South Wales and excluding Kroombit Tops, D. the relevant taxa, I have and on many occasions across a ausviridis sp. nov. from the wet tropics of North 50 year time span, having caught thousands of the Queensland, D. celantur sp. nov. from Kroombit Tops,

south-west of Gladstone, Queensland and D. glauerti taxon from the species D. fallax Peters, 1882 found south (Copland, 1957) from New South Wales and just into of there along the Queensland coast to the NSW south-east Queensland are all readily separated from the Queensland border south of which the similar species D. species D. cooloolensis (Liem, 1974), D. glauerti (Copland, 1957) occurs, further down the coast northstradbrokensis sp. nov., D. olongburensis (Liem and and adjacent ranges to southern New South Wales. Ingram, 1977) and Maxinehoserranae bicolor (Gray, This means D. ausviridis sp. nov. is found along the 1842) including associated similar species formerly Queensland coast and ranges from Townsville, north to included within M. bicolor by the following unique include the wet tropics region, but not including drier combination of characters: parts of Cape York further north. Dorsal colour uniformly green or brownish (not obviously Etymology: Named in refection of where the frog is bicoloured); web of first toe reaches base of disc; found (Australia) and the dorsal colour, usually being a posterior of thighs orange; IN/EN and HW/HL ratio is less bright emerald green (hence “aus” and “viridis” = than 1.000; no dark brown spots or reticulations of any “ausviridis”). form and no purplish-brown femoral streak bordering the DRYMOMANTIS CELANTUR SP. NOV. dorsal green colour of the thigh (this colour configuration LSIDurn:lsid:zoobank.org:act:230275D2-ACA8-4803- as seen in D. cooloolensis (Liem, 1974) and D. AA4B-CE6B0CF65211 northstradbrokensis sp. nov.); no extremely pointed snout Holotype: A preserved adult male specimen at the with a distinct, slightly upturned rostrum, with upper jaw Queensland Museum, Brisbane, Queensland, Australia, protruding rostrum (as seen in D. oblongburensis). specimen number J86217 collected at Kroombit Creek at While all of D. fallax, D. ausviridis sp. nov., D. celantur the crossing of Kroombit Forest Drive, Kroombit Tops sp. nov. and D. glauerti are morphologically similar and National Park, south-west of Gladstone, Queensland, also variable in colouration within a single population, Australia, Latitude -24.3842 S., Longitude 151.0014 E. there are consistent differences in terms of character This government-owned facility allows access to its suites that separate all from one another. holdings. The four species are separated from one another as Paratype: A preserved specimen at the Queensland follows: Museum, Brisbane, Queensland, Australia, specimen D. fallax has a brown iris, with a slight reddish tinge, the number J42175 collected at Kroombit Tops, south-west of dark brown line running from nostril to eye is bounded on Gladstone, Queensland, Australia, Latitude -24.3667 S., both sides by a light brown border; scattered raised Longitude 151.0167 E. tubercles on the forelimbs are obvious. Diagnosis: The putative species defined as “Litoria fallax D. glauerti has a brown iris, numerous indistinct tubercles (Peters, 1882)” as defined and separated from other on the forelimbs and the line running from nostril to eye is species by Cogger (2014) and Anstis (2013) is found not bordered by a lighter brown border and is otherwise along the coast and ranges from southern New South well defined and sharp edged. Wales along the east Australian coast to the northern D. ausviridis sp. nov. has a red iris and tiny white spots edge of the wet tropics in far north Queensland. on the upper hind limbs. The line between the nostril and That putative taxon is treated herein as four the eye has indistinct upper and lower morphologically similar species. boundaries.Tubercles on the forelimbs are so small as to The four species, until now usually treated all as be effectively invisible. Drymomantis fallax Peters, 1882 are readily separated D. celantur sp. nov. has scattered tiny raised white from their nearest relatives and congeners, D. tubercles on the front limbs, an ill-defined line from nostril cooloolensis (Liem, 1974); the similar D. to eye, a brown iris and often has indistinct blotches or northstradbrokensis sp. nov. and D. olongburensis (Liem flecks arranged longitudinally down the back. The white and Ingram, 1977) by morphology. Same applies in terms line commencing on the upper-lip below the eye, that of the morphologically similar species group most widely runs across the forelimb and along the lower side of the known as “Litoria bicolor (Gray, 1842)” (e.g. Cogger, flank is particularly well defined. 2014), but herein placed in the new genus D. ausviridis sp. nov. in life is depicted in Vanderduys Maxinehoserranae gen. nov.. (2012) on page 40 at bottom left and Anstis (2013) on Duellman et al. (2016) found a divergence of the so- page 200 on the right (2 images). called bicolor species group of 21.5 MYA from the genus D. fallax in life is depicted in Vanderduys (2012) on page Drymomantis making separate genus-level recognition 40 at bottom right. an obvious judgement. D. glauerti in life is depicted in Cogger (2014) on page The morphologically similar species Drymomantis fallax 164 bottom right and in Anstis (2013) on page 203 top Peters, 1882 from coastal Queensland south of the left (2 images). Burdekin River, North Queensland, just into north-east Photos of all of D. ausviridis sp. nov., D. fallax, D. glauerti New South Wales and excluding Kroombit Tops, D. and D. celantur sp. nov. can be found at http:// ausviridis sp. nov. from the wet tropics of North www.flickr.com Queensland, D. celantur sp. nov. from Kroombit Tops, By doing a search for “Litoria fallax”. south-west of Gladstone, Queensland and D. glauerti Distribution: D. ausviridis sp. nov. occurs on Cape York (Copland, 1957) from New South Wales and just into in Queensland, generally north of the Burdekin River gap, south-east Queensland are all readily separated from the a well defined biogeographical barrier separating this species D. cooloolensis (Liem, 1974), D.

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 106 Australasian Journal of Herpetology northstradbrokensis sp. nov., D. olongburensis (Liem and DRYMOMANTIS NORTHSTRADBROKENSIS SP. NOV. Ingram, 1977) and Maxinehoserranae bicolor (Gray, LSIDurn:lsid:zoobank.org:act:9DE222E5-C9E8-430E- 1842) by the following unique combination of characters: AF71-3C6BD6C0F2CF Dorsal colour uniformly green or brownish (not obviously Holotype: A preserved specimen at the Queensland bicoloured); web of 1st toe reaches base of disc; Museum, Brisbane, Queensland, Australia, specimen posterior of thighs orange; internarial/eye-naris distance number J27569 collected at Tortoise Lagoon, North and head-width/head-length ratio is less than 1; no dark Stradbroke Island, Queensland, Australia, Latitude - brown spots or reticulations of any form and no purplish- 27.5183 S., Longitude 153.4728 E. brown femoral streak bordering the dorsal green colour of This government-owned facility allows access to its the thigh (this colour configuration as seen in D. holdings. cooloolensis (Liem, 1974) and D. northstradbrokensis sp. Paratypes: 1/ A preserved specimen at the Queensland nov.); no extremely pointed snout with a distinct, slightly Museum, Brisbane, Queensland, Australia, specimen upturned rostrum, with upper jaw protruding rostrum (as number J27558 collected at Blue Lake, North Stradbroke seen in D. oblongburensis). Island, Queensland, Australia, Latitude -27.5303 S., While all of D. fallax, D. ausviridis sp. nov., D. celantur Longitude 153.4764 E. 2/ A preserved specimen at the sp. nov. and D. glauerti are morphologically similar and Queensland Museum, Brisbane, Queensland, Australia, also variable in colouration within a single population, specimen number J81495, collected at Swallow Lagoon, there are consistent differences in terms of character North Stradbroke Island, Queensland, Australia, Latitude suites that separate all from one another. The four -27.4989 S., Longitude 153.4547 E. species are separated from one another as follows: Diagnosis: Until now, Drymomantis northstradbrokensis D. fallax has a brown iris, with a slight reddish tinge, the sp. nov. from North Stradbroke Island, east of Brisbane in dark brown line running from nostril to eye is bounded on Queensland, Australia, has been treated as a southern both sides by a light brown border; scattered raised population of D. cooloolensis (Liem and Ingram, 1977), tubercles on the forelimbs are obvious. with a type locality of Cooloola, Queensland, Australia. D. glauerti has a brown iris, numerous indistinct tubercles However the two species are morphologically divergent on the forelimbs and the line running from nostril to eye is and this has been known for some time and hence it not bordered by a lighter brown border and is otherwise beingf ormally named herein. well defined and sharp edged. D. cooloolensis is restricted to Cooloola and nearby D. ausviridis sp. nov. has a red iris and tiny white spots Fraser Island, Queensland (Cogger, 2014), while D. on the upper hind limbs. The line between the nostril and northstradbrokensis sp. nov. is known only from North the eye has indistinct upper and lower boundaries. Stradbroke Island, East of Brisbane, Queensland. Tubercles on the forelimbs are so small as to be Both species are separated from all other species within effectively invisible. the genera Drymomantis Peters, 1882 and D. celantur sp. nov. has scattered tiny raised white the morphologically similar Maxinehoserranae gen. nov. tubercles on the front limbs, an ill-defined line from nostril (known as the “Litoria bicolor group” to most to eye, a brown iris and often has indistinct blotches or herpetologists as of 2020) by the following unique suite of flecks arranged longitudinally down the back. The white characters: The green dorsum has numerous brown line commencing on the upper-lip below the eye, that spots and reticulations (versus none or very few in all runs across the forelimb and along the lower side of the morphologically similar Australian species); absence of a flank is particularly well defined. brown or grey head streak between nostril and eye D. ausviridis sp. nov. in life is depicted in Vanderduys (present in all morphologically similar Australian species); (2012) on page 40 at bottom left and Anstis (2013) on the tympanum is always green; there is a purplish-brown page 200 on the right (2 images). streak along the upper thigh (absent or obscure in all D. fallax in life is depicted in Vanderduys (2012) on page morphologically similar Australian species); internarial/ 40 at bottom right. eye-naris distance and head-width/head-length ratio is D. glauerti in life is depicted in Cogger (2014) on page less than 1 (versus less than 1 in all morphologically 164 bottom right and in Anstis (2013) on page 203 top similar Australian species). left (2 images). Adult D. northstradbrokensis sp. nov. are readily Photos of all of D. ausviridis sp. nov., D. fallax, D. glauerti separated from adult D. cooloolensis by having an and D. celantur sp. nov. can be found at http:// orange iris (versus grey/brown in D. cooloolensis) and a www.flickr.com dorsum where the spots or flecks are ill-defined and By doing a search for “Litoria fallax”. subdued, versus obvious and distinct in D. cooloolensis. Distribution: Currently D. celantur sp. nov. is only known In D. northstradbrokensis sp. nov. the white line from the from Kroombit Tops National Park in Queensland, back of the upper-lip over the forelimb to the lower flank Australia and based on an absence of Museum voucher is obvious and distinct, versus indistinct and often broken specimens of these frogs from the periphery of this in D. cooloolensis. highland reserve, it is most probably restricted to this Photos of both D. northstradbrokensis sp. nov. and D. area. A similar situation exists at Kroombit Tops for other cooloolensis, side-by-side in life can be found in Cogger climatically confined taxa. (2014), page 158, with D. northstradbrokensis sp. nov. on Etymology: Celantur in Latin means concealed and this right, Vanderduys (2012) on page 31 with D. species has been effectively concealed from science for northstradbrokensis sp. nov. on left and in Anstis (2013) on page 173, with D. northstradbrokensis sp. nov. on left. longer than it should have been. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 107

NYCTIMYSTINI TRIBE NOV. gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. NYCTIMYSTES STEJNEGER, 1916. and Albogibba gen. nov.) by having sparse, broken Type species: Nyctimantis papua Boulenger, 1897. vertically-oriented palpebral venation across the Diagnosis: The genus Nyctimystes Stejneger, 1916 has nictitating membrane and in having adult males without in the past 3 decades undergone significant revision and vocal slits. alteration by various authors. Frogs in the subgenus Magnummanibus subgen. nov. are Zweifel (1958) defined the genus as follows: “The genus separated from those in the nominate subgenus Nyctimystes (family Hylidae) is defined as including those Nyctimystes and the subgenus Asperohyla subgen. nov. species that combine the following characteristics: pupil by their possession of very long, prominent heel lappets. forming a vertical slit when closed, lower eyelid with Frogs in the subgenus Asperohyla subgen. nov. are pigmented venation, feet without elongate or opposable separated from the other two subgenera by having a first digits.” completely hidden tympanum and a dorsal skin Cogger (2014) had a similar diagnosis for the genus roughened by tiny, conical asperities rather than by stating “Species of Nyctimystes are distinguished by their variable sized and more rounded irregularities as seen in large dark eyes, which have vertically elliptical pupils and some other New Guinea tree frog species; iris is brown in a lower eyelid with a characteristic reticulum or palpebral life. venation of fine pigmented lines. There is usually a Species within Occultatahyla gen. nov. are separated distinct, crenulated skin fold along the hind edge of the from the other five genera (Nyctimystes, Nigreosoculus forearm and the foot, the heel often having a small flap of gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. skin. Males are usually considerably smaller than and Albogibba gen. nov.) by large size (adult females 50 females.” to 84 mm. in snout to vent length); internarial distance With the preceding diagnosis in mind, Cogger (2014) distinctly smaller than distance from eye to naris (E-N/IN removed the Australian species from Nyctimystes on the averages 1.34 (range 1.2-1.5)); the head is broad, basis of recent phylogenetic studies and the fact that flattish, with depressed snout, closely spaced nares, and their pupils were in fact horizontally elliptical, in line with oblique loreal region. Palpebral venation is in oblique other Australian tree frogs. lines (more vertical than horizontal and with few horizontal interconnections), with a few horizontal With a divergence of 19.9 MYA from their nearest living interconnections. A dermal fold passes from the posterior relatives (according to Duellman et al. 2016), those corner of the eye over the upper edge of the tympanum species have been formally transferred to the newly and down to the insertion of the forelimb, sometimes erected genus Gedyerana gen. nov.. becoming obscure when leaving the tympanum. The However the preceding diagnosis applies to all of the six outer fingers are approximately half-webbed, the third following genera, all relevant species of which until now, and fifth toes webbed to the disc. A light-coloured dermal were included in the single genus Nyctimystes. ridge or series of tubercles extends from the elbow to the The other five newly erected genera are Occultatahyla disc of the fourth finger. A similar but less distinct ridge or gen. nov., Nigreosoculus gen. nov., Magnumoculus gen. line of tubercles occupies the outer edge of the tarsus nov., Badiohyla gen. nov. and Albogibba gen. nov.. and fifth toe. A very small heel tubercle is present. The The diagnosis of all six genera, including of a narrower skin of the dorsum is smooth to very slightly granular, that group of species now treated as the entirety of the genus of the venter is always coarsely granulate. Slit-like, vocal Nyctimystes and the other five genera which are formally sac openings are usually present in the floor of the named within this paper for the first time is given below. mouth. A small patch of very fine horny tubercles is found Duellman et al. (2016) expanded the genus Nyctimystes on the first finger of males. The iris is dark or black in to include a clade of frogs with a divergence of 41 MYA colour. from nearest living relatives, thereby including lineages of Exceptional to the preceding is the subgenus Webpede frogs previously included in the genus “Litoria Tschudi, subgen. nov. which conforms to the above genus 1838. However such deep divergence is more diagnosis, save for the following differences: The heel is appropriate for tribe level in classification and not genus without a tubercle, or in rare cases has a very small one. level. Hence the break up as indicated herein and/or The skin is smooth above, granular below. A weak, wavy throughout this paper in it’s full state. fold is present on the outer surface of the forearm. The Species of Nyctimystes, Occultatahyla gen. nov., head is broader than long; the canthus rostralis is not Nigreosoculus gen. nov., Magnumoculus gen. nov., distinct; the loreal region is oblique; Unique to this Badiohyla gen. nov. and Albogibba gen. nov. are subgenus is that the palpebral venation is distinct, but is separated from all other Australasian (Australian and reduced to individual pigment spots and thin, meandering New Guinea) Tree Frogs (Pelodryadidae) by the following lines, although the lines are still mainly oriented largely in suite of characters: Large dark eyes, which have a vertical direction, with few horizontal interconnections. vertically elliptical pupils and a lower eyelid with a Species within Nigreosoculus gen. nov. are separated characteristic reticulum or palpebral venation of fine from the other five genera (Nyctimystes, Occultatahyla pigmented lines. There is usually a distinct, crenulated gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. skin fold along the hind edge of the forearm and the foot, and Albogibba gen. nov.) by the following suite of the heel often having a small flap of skin. Males are characters: Vocal slits present; a very small heel tubercle; usually considerably smaller than females. basal webbing on the hand; exposed tympanum; vertical Nyctimystes Stejneger, 1916 are separated from the lines of palpebral reticulum oriented obliquely and with other five genera (Occultatahyla gen. nov., Nigreosoculus Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 108 Australasian Journal of Herpetology relatively few horizontal cross-connections; pale-tan to vent length); of a uniform dorsal coloration (green in life, near black iris; rear of thighs barred/mottled with brown, purple in alcohol-preserved specimens); palpebral caramel, or blue-gray. venation is a reticulum without obvious orientation and in Species within Magnumoculus gen. nov. are separated the nominate subgenus of this genus, the male bears a from the other five genera (Nyctimystes, Nigreosoculus spine-like process on the anterior surface of the proximal gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. part of the humerus (upper arm) (nominate subgenus and Albogibba gen. nov.) by the following suite of Albogibba subgen. nov.), or: characters: The head is broad (HL/HW <1.0), the snout 2/ As above, but male SVL of up to 80 mm; not including high and blunt, the canthus rostralis distinct, broadly a spine-like process on the anterior surface of the spaced nostrils (EN/IN ratio <1), and the loreal region proximal part of the humerus (upper arm) in males and nearly vertical. A strong, slightly curved, supratympanic with a unique dorsal pattern of black vermiculations on fold begins at the posterior corner of the eye and the body and limbs, with unmarked greyish lower flanks disappears above the insertion of the forelimb. The (Ratiobrunneis subgen. nov.). tympanum is small but distinct. The hands have little According to Duellman et al. (2016), each of the six webbing; the outer fingers are about one-quarter webbed. preceding genera had the following divergences from one The vomerine teeth are in two patches between the another. internal nares, nearly on a line connecting the posterior Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its margins of the nares. The skin is minutely granular nearest living relatives in other genera. above, coarser beneath. There is no tubercle on the heel. The subgenus Magnummanibus subgen. nov. diverged A nearly straight, very slightly crenulated fold of skin is 11.5 MYA from its nearest living relatives in the nominate present on the outer surface of the forearm. A moderate subgenus. body size (40-60 mm S-V), predominantly brown Occultatahyla gen. nov. diverged 16.7 MYA from its colouration, an eyelid venation composed of a network nearest living relatives in other genera. with numerous horizontal connections with relatively few vertical interconnections. There is decoration on the Nigreosoculus gen. nov. diverged 14.8 MYA from its forearm and tarsus in the form of rows of (often white- nearest living relatives in other genera. tipped) tubercles. Magnumoculus gen. nov. diverged 13 MYA from its Species within Badiohyla gen. nov. are separated from nearest living relatives, being Badiohyla gen. nov.. the other five genera (Nyctimystes, Nigreosoculus gen. Badiohyla gen. nov. diverged 13 MYA from its nearest nov., Occultatahyla gen. nov., Magnumoculus gen. nov. living relatives being Magnumoculus gen. nov.. and Albogibba gen. nov.) by having a unique “reticulated” In turn both Magnumoculus gen. nov. and Badiohyla gen. palpebral venation, half-webbed fingers and a size in nov. as a pair diverged from their nearest living relatives males with SVL to maximum of 50 mm to 100 mm 22 MYA being Nyctimystes and other genera. depending on the species. They are further defined as Albogibba gen. nov. diverged 16.7 MYA from its nearest follows: The snout is relatively short, blunt and high (E-N/ living relatives in other genera. IN 0.94), with distinct canthus rostralis and oblique loreal The subgenus Ratiobrunneis subgen. nov. diverged 10 region. The vomerine teeth are in two patches between MYA from its nearest living relatives in the nominate the posterior edges of the choanae. subgenus. The palpebral venation forms a heavy reticulum. The Distribution: New Guinea including nearby offshore tympanum is distinct and is separated from the eye by islands on the continental shelf. approximately the diameter of the tympanum. A fold of Content: Nyctimystes papua (Boulenger, 1897) (type skin passes from the posterior corner of the eye, over the species); N. aspera sp. nov.; N. charlottae sp. nov.; N. upper edge of the tympanum and down, becoming doggettae sp. nov.; N. disruptus Tyler, 1963; N. indistinct above the insertion of the forelimb. The skin of georgefloydi sp. nov.; N. mondoensis sp. nov.; N. the dorsum is minutely roughened, that of the venter oktediensis Richards and Johnston, 1993; N. pulchra coarsely granular. A row of tubercles is present along the (Wandolleck, 1910); N. trachydermis Zweifel, 1983; N. outer edge of the forearm and there is a similar row on tyleri (Zweifel, 1983). the tarsus. There is only a slight suggestion of a heel MAGNUMMANIBUS SUBGEN. NOV. tubercle. The outer fingers are approximately half- LSIDurn:lsid:zoobank.org:act:084E0A6D-0B8C-4521- webbed, the toes, except the fourth toe, are webbed to 8139-4E53BA85F70B the disc. Type species: Hyla pulchra Wandolleck, 1910. The body and head are light brown to purple dorsally, Diagnosis: Species of Nyctimystes, Occultatahyla gen. either with irregular dark brown blotches or markings nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., absent. The legs have a similar ground color, with or Badiohyla gen. nov. and Albogibba gen. nov. are without irregular darker spots and bands present on the separated from all other Australasian (Australian and tibia. New Guinea) Tree Frogs (Pelodryadidae) by the following Species within Albogibba gen. nov. are separated from suite of characters: Large dark eyes, which have the other five genera (Nyctimystes, Nigreosoculus gen. vertically elliptical pupils and a lower eyelid with a nov., Occultatahyla gen. nov., Magnumoculus gen. nov. characteristic reticulum or palpebral venation of fine and Badiohyla gen. nov.) by one or other of the following pigmented lines. unique suites of characters: There is usually a distinct, crenulated skin fold along the 1/ Being a large species (males to 100 mm. in snout to hind edge of the forearm and the foot, the heel often

having a small flap of skin. Males are usually by their possession of very long, prominent heel lappets. considerably smaller than females. Frogs in the subgenus Asperohyla subgen. nov. are Nyctimystes Stejneger, 1916 are separated from the separated from the other two subgenera by having a other five genera (Occultatahyla gen. nov., Nigreosoculus completely hidden tympanum and a dorsal skin gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. roughened by tiny, conical asperities rather than by and Albogibba gen. nov.) by having sparse, broken variable sized and more rounded irregularities as seen in vertically-oriented palpebral venation across the some other New Guinea tree frog species; iris is brown in nictitating membrane and in having adult males without life. vocal slits. For further detail see the diagnosis of the genus Frogs in the subgenus Magnummanibus subgen. nov. are Nyctimystes above. separated from those in the nominate subgenus Distribution: New Guinea from the Irian Jaya border, Nyctimystes and the subgenus Asperohyla subgen. nov. along the central ranges and nearby mountain ranges by their possession of very long, prominent heel lappets. east to Milne Bay on both sides of the central cordillera. Frogs in the subgenus Asperohyla subgen. nov. are Etymology: The subgenus name “Asperohyla” refers in separated from the other two subgenera by having a Latin to the relevant species having a relative rough to completely hidden tympanum and a dorsal skin touch skin on the dorsal surface and the fact that the said roughened by tiny, conical asperities rather than by frogs are also tree frogs. variable sized and more rounded irregularities as seen in Content: Nyctimystes (Asperohyla) trachydermis Zweifel, some other New Guinea tree frog species; iris is brown in 1983 (type species); N. (Asperohyla) aspera sp. nov.; N. life. (Asperohyla) georgefloydi sp. nov.. For further detail see the diagnosis of the genus NYCTIMYSTES (NYCTIMYSTES) MONDOENSIS SP. Nyctimystes above. NOV. Frogs within Magnummanibus subgen. nov. are further LSIDurn:lsid:zoobank.org:act:72AF2EBA-F8B9-46B2- diagnosed and separated from other morphologically 844C-3B6F88E0400B similar species as follows: Moderately large 60 mm in Holotype: A preserved specimen at the Museum of snout-vent length frogs. Head very broad and flat; Comparative Zoology, Harvard University in Cambridge, palpebral venation is a network of gold or dark brown Massachusetts, USA, specimen number MCZ A 21816 lines with vertical-oblique orientation and frequent collected at Mondo, elevation 5000 feet (1524 m.) on the horizontal cross connections. Very long, prominent heel north slope of Mt. Tafa, Papua New Guinea, Latitude - lappets. Dorsum varies in coliur and pattern, but is 8.6333 S., Longitude 147.1833 E. usually brownish. Venter is usually yellowish and mottled This facility allows access to its holdings. or speckled with darker pigment all over. Iris is dark to very dark brown. Paratypes: Three preserved specimens at the Museum of Comparative Zoology, Harvard University in Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its Cambridge, Massachusetts, USA, specimen numbers nearest living relatives in other genera. MCZ A 21817-21820 collected at Mondo, elevation 5000 The subgenus Magnummanibus subgen. nov. diverged feet (1524 m.) on the north slope of Mt. Tafa, Papua New 11.5 MYA from its nearest living relatives in the nominate Guinea, Latitude -8.6333 S., Longitude 147.1833 E. subgenus. Diagnosis: Nyctimystes mondoensis sp. nov. is similar in Distribution: Both north and south of the main cordillera most respects to N. papua (Boulenger, 1897) and would in New Guinea. key out as that species as seen in Zweifel (1958). Both Etymology: The name Magnummanibus is derived from species are separated from all other species in the genus the Latin meaning, which is “large hand” and is an Nyctimystes Stejneger, 1916 by having palpebral obvious characteristic of the subgenus. venation that is relatively weak and disconnected, Content: Nyctimystes (Magnummanibus) pulchra sometimes virtually absent (versus distinct in all other (Wandolleck, 1910) (type species); N. (Magnummanibus) species); outer fingers with only a basal web or not much charlottae sp. nov.; N. (Magnummanibus) doggettae sp. greater and male without vocal-sac openings and adult nov. size usually between 50-60 mm in snout-vent length. ASPEROHYLA SUBGEN. NOV. Nyctimystes mondoensis sp. nov. is separated from N. LSIDurn:lsid:zoobank.org:act:C7F57438-81FF-45EA- papua by having a semi-distinct pigmentation, with BB77-635BAACE1219 diagonal lines, occasionally interconnected, over three- Type species: Nyctimystes trachydermis Zweifel, 1983. quarters of the palpebral area (versus almost absent in Diagnosis: Nyctimystes Stejneger, 1916 are separated N. papua), moderate finger webbing (versus little in N. from the other five genera (Occultatahyla gen. nov., papua and a lot in the related species N. disruptus Tyler, Nigreosoculus gen. nov., Magnumoculus gen. nov., 1963 and N. oktediensis Richards and Johnston, 1993) Badiohyla gen. nov. and Albogibba gen. nov.) by having relatively shorter legged than N. papua and a greater EN sparse, broken vertically-oriented palpebral venation distance (over 5.8 mm) than in N. papua (below 5.7 mm). across the nictitating membrane and in having adult Distribution: Nyctimystes mondoensis sp. nov. is known males without vocal slits. only from the type locality, but probably occurs more Frogs in the subgenus Magnummanibus subgen. nov. are widely on the northern side of the Owen Stanley Range in separated from those in the nominate subgenus south-east New Guinea. Nyctimystes and the subgenus Asperohyla subgen. nov. Etymology: Named in reflection of the type locality.

NYCTIMYSTES (MAGNUMMANIBUS) CHARLOTTAE blue, with dark reticulum; entire ventral surface is SP. NOV. yellowish, mottled or speckled dark all over. The iris is a LSIDurn:lsid:zoobank.org:act:37FE756E-47A1-4ACC- very dark brown. 9A10-21AFD84CCE49 N. doggettae sp. nov. is chocolate brown above with Holotype: A preserved specimen at the Museum of irregular indistinct blotches of lighter brown breaking up Comparative Zoology, Harvard University in Cambridge, the darker dorsum. At the rear of the back on the dorsum Massachusetts, USA, specimen number MCZ A-110420, are areas of light yellowish-brown, more or less in the collected at Olsobip at an elevation of 1,500 feet, central region of the rear of the back. The thin dark and Western Province, Papua New Guinea, Latitude -5.3895 light bands on the upper surfaces of the upper and lower S., Longitude 141.5153 E. hind legs are very well defined (as opposed to being ill- This facility allows access to its holdings. defined or absent in the other two species). Prominent in Paratypes: 1/ A preserved specimen at the Museum of this species are scattered white tipped tubercles on the Comparative Zoology, Harvard University in Cambridge, back, which are either absent, or small and ill-defined in Massachusetts, USA, specimen number MCZ A-95510 the other two species. Iris is a very dark brown. collected at Imigabip, at an elevation of 4,200 feet, Nyctimystes Stejneger, 1916 are separated from the Western Province, Papua New Guinea, Latitude -5.2898 other five genera (Occultatahyla gen. nov., Nigreosoculus S., Longitude 141.5018 E. gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. 2/ A preserved specimen at the Museum of Comparative and Albogibba gen. nov.) by having sparse, broken Zoology, Harvard University in Cambridge, vertically-oriented palpebral venation across the Massachusetts, USA, specimen number MCZ A-110442, nictitating membrane and in having adult males without collected at Wangbin at an elevation of 4,800 feet, vocal slits. Western Province, Papua New Guinea, Latitude -5.2518 Frogs in the subgenus Asperohyla subgen. nov. are S., Longitude 141.2412 E. separated from the other two subgenera 3/ A preserved specimen at the Museum of Comparative (Magnummanibus subgen. nov. and Nyctimystes) and by Zoology, Harvard University in Cambridge, having a completely hidden tympanum and a dorsal skin Massachusetts, USA, specimen number MCZ A-87137 roughened by tiny, conical asperities rather than by collected at Kavorabip, elevation 5,000 feet, Western variable sized and more rounded irregularities as seen in Province, Papua New Guinea, Latitude -5.1301 S., some other New Guinea tree frog species; iris is brown in Longitude 141.1148 E. life. Diagnosis: Nyctimystes (Magnummanibus) charlottae Species of Nyctimystes, Occultatahyla gen. nov., sp. nov. and N. doggettae sp. nov. have both until now Nigreosoculus gen. nov., Magnumoculus gen. nov., been treated as populations of the species N. pulchra Badiohyla gen. nov. and Albogibba gen. nov. are (Wandolleck, 1910). separated from all other Australasian (Australian and New Guinea) Tree Frogs (Pelodryadidae) by the following All would until now have been identified as that species suite of characters: Large dark eyes, which have via the information given in Menzies (2006). vertically elliptical pupils and a lower eyelid with a These three species, constituting all frogs in the characteristic reticulum or palpebral venation of fine subgenus Magnummanibus subgen. nov. are separated pigmented lines. from those in the nominate subgenus Nyctimystes and There is usually a distinct, crenulated skin fold along the the subgenus Asperohyla subgen. nov. by their hind edge of the forearm and the foot, the heel often possession of very long, prominent heel lappets. having a small flap of skin. Males are usually Males grow to 70 mm in body length and females to 80 considerably smaller than females. mm. The body is slender and the head broad and flat with Distribution: N. charlottae sp. nov. is found in high large foreward facing eyes. The venation on the eyelid elevation regions south of the central cordillera in New forms a network of dark brown lines oriented in a vertical/ Guinea, including in both Papua New Guinea and oblique direction. Indonesia. The three species N. charlottae sp. nov. from south of the Etymology: The species is named in honour of Charlotte central cordillera in New Guinea, N. doggettae sp. nov. Doggett of Ringwood East, Victoria, this species being from the Milne Bay region of Papua New Guinea and N. named after her first name, in recognition of her services pulchra from Madang and nearby parts of the north of to wildlife conservation in assisting the team at Australia’s New Guinea are all readily separated from one another best reptiles shows and Snakebusters. on the basis of colour. NYCTIMYSTES (MAGNUMMANIBUS) DOGGETTAE SP. N. pulchra has a dorsum that is plain medium to dark NOV. brown with whitish or yellowish lichen-like spots or mottled tan to medium dark brown; flanks bluish mottled LSIDurn:lsid:zoobank.org:act:57D5C8C1-66AB-423A- with darker or yellowish brown, speckled black; 8CC2-834654F48FFB concealed surfaces of thighs bluish, mottled brown; Holotype: A preserved specimen at the Bernice P. ventral surface yellowish, mottled or speckled dark all Bishop Museum, Honolulu, Hawaii, USA, specimen over. Iris is dark brown. number BPBM Herp 11032, collected at Iana River, 3.8 N. charlottae sp. nov. is brown or olive brown above, with km West of Agaun, Papua New Guinea, Latitude -9.876 orange blotches; flanks mottled bluish or yellowish, S., Longitude 149.352 E. speckled black; concealed surfaces of thighs greenish This facility allows access to its holdings.

Paratypes: 1/ A preserved specimen at the Bernice P. separated from the other two subgenera Bishop Museum, Honolulu, Hawaii, USA, specimen (Magnummanibus subgen. nov. and Nyctimystes) and by number BPBM Herp 17045 collected at Uga River, Papua having a completely hidden tympanum and a dorsal skin New Guinea, Latitude -10.0245 S., Longitude 149.59 E. roughened by tiny, conical asperities rather than by 2/ A preserved specimen at the Bernice P. Bishop variable sized and more rounded irregularities as seen in Museum, Honolulu, Hawaii, USA, specimen number some other New Guinea tree frog species; iris is brown in BPBM Herp 39561 collected at Binigua, Papua New life. Guinea, Latitude -9.7071. S., Longitude 149.25 E. Species of Nyctimystes, Occultatahyla gen. nov., Diagnosis: Nyctimystes (Magnummanibus) doggettae Nigreosoculus gen. nov., Magnumoculus gen. nov., sp. nov. and N. charlottae sp. nov. have both until now Badiohyla gen. nov. and Albogibba gen. nov. are been treated as populations of the species N. pulchra separated from all other Australasian (Australian and (Wandolleck, 1910). New Guinea) Tree Frogs (Pelodryadidae) by the following All would until now have been identified as that species suite of characters: Large dark eyes, which have via the information given in Menzies (2006). vertically elliptical pupils and a lower eyelid with a These three species, constituting all frogs in the characteristic reticulum or palpebral venation of fine subgenus Magnummanibus subgen. nov. are separated pigmented lines. There is usually a distinct, crenulated from those in the nominate subgenus Nyctimystes and skin fold along the hind edge of the forearm and the foot, the subgenus Asperohyla subgen. nov. by their the heel often having a small flap of skin. Males are possession of very long, prominent heel lappets. usually considerably smaller than females. Males grow to 70 mm in body length and females to 80 Distribution: N. doggettae sp. nov. is found in the Milne mm. The body is slender and the head broad and flat with Bay region of Papua New Guinea. large foreward facing eyes. The venation on the eyelid Etymology: The species is named in honour of Charlotte forms a network of dark brown lines oriented in a vertical/ Doggett of Ringwood East, Victoria in recognition of her oblique direction. services to wildlife conservation in assisting the team at The three species N. doggettae sp. nov. from the Milne Australia’s best reptiles shows and Snakebusters. In this Bay region of Papua New Guinea, N. charlottae sp. nov. case the species name is taken from her last name (N. from south of the central cordillera in New Guinea and N. doggettae sp. nov.) as opposed to the first, which was the pulchra from Madang and nearby parts of the north of case for the previous species (N. charlottae sp. nov.) also New Guinea are all readily separated from one another named in her honour. on the basis of colour. NYCTIMYSTES (ASPEROHYLA) ASPERA SP. NOV. N. pulchra has a dorsum that is plain medium to dark LSIDurn:lsid:zoobank.org:act:E9912B48-3433-4D33- brown with whitish or yellowish lichen-like spots or A6D8-D918AE430260 mottled tan to medium dark brown; flanks bluish mottled Holotype: A preserved specimen at the American with darker or yellowish brown, speckled black; Museum of Natural History, Manhattan, New York, USA, concealed surfaces of thighs bluish, mottled brown; herpetology collection, specimen number 103184 ventral surface yellowish, mottled or speckled dark all collected at Nipa, Southern Highlands Province, Papua over. Iris is dark brown. New Guinea, Latitude 6.1272 S., Longitude 143.4203 E. N. doggettae sp. nov. is chocolate brown above with This facility allows access to its holdings. irregular indistinct blotches of lighter brown breaking up Paratypes. Four preserved specimens at the American the darker dorsum. At the rear of the back on the dorsum Museum of Natural History, Manhattan, New York, USA, are areas of light yellowish-brown, more or less in the herpetology collection, specimen number 103185-8 central region of the rear of the back. The thin dark and collected at Nipa, Southern Highlands Province, Papua light bands on the upper surfaces of the upper and lower New Guinea, Latitude 6.1272 S., Longitude 143.4203 E. hind legs are very well defined (as opposed to being ill- Diagnosis: Both species, Nyctimystes (Asperohyla) defined or absent in the other two species). Prominent in aspera sp. nov. and N. georgefloydi sp. nov. have until this species are scattered white tipped tubercles on the now been treated as populations of N. trachydermis back, which are either absent, or small and ill-defined in Zweifel, 1983 and as a trio form the entirety of the the other two species. Iris is a very dark brown. subgenus Asperohyla subgen. nov.. N. charlottae sp. nov. is brown or olive brown above, with The three species N. trachydermis Zweifel, 1983 from the orange blotches; flanks mottled bluish or yellowish, Bowutu Mountains in Morobe Province, Papua New speckled black; concealed surfaces of thighs greenish Guinea, N. aspera sp. nov. from the Southern Highlands blue, with dark reticulum; entire ventral surface is Province of Papua New Guinea and N. georgefloydi sp. yellowish, mottled or speckled dark all over. The iris is a nov. from south-flowing watersheds in the Owen Stanley very dark brown. Range, Central Province of Papua New Guinea can all be Nyctimystes Stejneger, 1916 are separated from the separated as follows. other five genera (Occultatahyla gen. nov., Nigreosoculus N. trachydermis has a dorsal ground color that is brown gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. in life, with a pale tan area on the top of the snout, and Albogibba gen. nov.) by having sparse, broken continuous along the outer edge of the eyelids with an vertically-oriented palpebral venation across the elongate patch of the same color on the right side and nictitating membrane and in having adult males without slightly separated from a similar, shorter patch on the left. vocal slits. Smaller, irregularly shaped areas of the same light color Frogs in the subgenus Asperohyla subgen. nov. are are present on the hind limbs and lower back. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 112 Australasian Journal of Herpetology

The upper lip is edged with yellowish tan, but with the separated as follows. upper edge of that color ill defined. N. trachydermis has a dorsal ground color that is brown The dorsal patterning just described may be prominent or in life, with a pale tan area on the top of the snout, indistinct. continuous along the outer edge of the eyelids with an The iris is reddish brown, with the conspicuous vertical elongate patch of the same color on pupil narrowly edged in yellow. The supratympanic ridge the right side and slightly separated from a similar, is moderately well-defined. Tubercles are significantly shorter patch on the left. Smaller, irregularly shaped enlarged on the lower flanks. areas of the same light color are present on the hind N. aspera sp. nov. is similar in most respects to N. limbs and lower back. The upper lip is edged with trachydermis but differs in not having significantly yellowish tan, but with the upper edge of that color ill enlarged tubercles on the lower flanks and prominent defined. whitish markings on the upper toes. The dorsal patterning just described may be prominent or N. georgefloydi sp. nov. is similar in most respects to N. indistinct. trachydermis and N. aspera sp. nov. but differs in not The iris is reddish brown, with the conspicuous vertical having significantly enlarged tubercles on the lower pupil narrowly edged in yellow. The supratympanic ridge flanks, no prominent whitish markings on the upper toes, is moderately well-defined. Tubercles are significantly a dorsum that is generally unmarked in any way and a enlarged on the lower flanks. distinctive dark brown colour with strong green peppering N. aspera sp. nov. is similar in most respects to N. all over, giving it an obvious dark greenish hue; no trachydermis but differs in not having significantly markings on snout or upper lip and a very strongly enlarged tubercles on the lower flanks, prominent whitish developed supratympanic ridge. markings on the upper toes and a very strongly In N. trachydermis and N. aspera sp. nov. the back of developed supratympanic ridge. forearm tubercles are large, prominent and closely N. georgefloydi sp. nov. is similar in most respects to N. spaced, sometimes forming a fold or ridge, whereas in N. trachydermis and N. aspera sp. nov. but differs in not georgefloydi sp. nov. the tubercles are well spaced and having significantly enlarged tubercles on the lower smaller. flanks, no prominent whitish markings on the upper toes, N. trachydermis in life is depicted in Zweifel (1983) at fig. a dorsum that is generally unmarked in any way and a 10. distinctive dark brown colour with strong green peppering N. georgefloydi sp. nov. in life is depicted in Menzies all over, giving it an obvious dark greenish hue; no (2006) at plate 96. markings on snout or upper lip and a very strongly Distribution: N. aspera sp. nov. is known from the developed supratympanic ridge. Southern Highlands Province of Papua New Guinea. In N. trachydermis and N. aspera sp. nov. the back of Etymology: The species name “Aspera” in Latin refers to forearm tubercles are large, prominent and closely the species being rough to touch. spaced, sometimes forming a fold or ridge, whereas in N. NYCTIMYSTES (ASPEROHYLA) GEORGEFLOYDI SP. georgefloydi sp. nov. the tubercles are well spaced and NOV. smaller. LSIDurn:lsid:zoobank.org:act:3711B439-F2E6-4FA8- N. trachydermis in life is depicted in Zweifel (1983) at fig. B210-3E6556B9FAE1 10. Holotype: A preserved specimen at the Bernice P. N. georgefloydi sp. nov. in life is depicted in Menzies Bishop Museum, Honolulu, Hawaii, USA, specimen (2006) at plate 96. number BPBM Herp 19446 collected at Mount Obree, Distribution: N. georgefloydi sp. nov. is known from Central Province, Papua New Guinea, Latitude -9.46014 south-flowing watersheds in the Owen Stanley Range, S., Longitude 148.03 S. Central Province of Papua New Guinea. This facility allows access to its holdings. Etymology: The species is named in honour of the late Paratypes: 8 preserved specimens at the Bernice P. George Floyd, a victim of an unprovoked attack by four Bishop Museum, Honolulu, Hawaii, USA, specimen thug police officers at Powderhorn, in downtown number BPBM Herp 18181, 19447, 19448, 19450, Minneapolis, USA, on 25 May 2020. 19451, 19453, 19455 and 19457 all from Mount Obree, Floyd was bashed by four Police Officers, Derek Central Province, Papua New Guinea, Latitude -9.46014 Chauvin, Thomas Kane, Tou Thao and J. Alexander S., Longitude 148.03 S. Jueng. The attack was apparently led by Derek Chauvin who in the final moments put his knee on top of Floyd’s Diagnosis: Both species, Nyctimystes (Asperohyla) neck and killed him by blocking his airways by georgefloydi sp. nov. and N. aspera sp. nov. have until maintaining the blockage for at least 8 minutes (Bungard now been treated as populations of N. trachydermis et al. 2020). Zweifel, 1983 and as a trio form the entirety of the subgenus Asperohyla subgen. nov.. The viciously subdued Floyd, was lying handcuffed, face- down on the bitumen of the city street. The three species N. trachydermis Zweifel, 1983 from the Bowutu Mountains in Morobe Province, Papua New There is no doubt Chauvin would have got away with the Guinea, N. aspera sp. nov. from the Southern Highlands murder were it not for the fact that his vicious attack was Province of Papua New Guinea and N. georgefloydi sp. filmed by members of the public from at least three nov. from south-flowing watersheds in the Owen Stanley different angles. Range, Central Province of Papua New Guinea can all be After the videos were posted on the internet, public

protests globally caused the Minneapolis Police committed against a fellow human being, but most department to stand the four officers down, but within certainly not the only crime being perpetrated by police days, the police were blaming the victim, alleging the and associated corrupt government employees. death was caused by pre-existing medical conditions and The United States in law has a “death penalty” and there not the police actions. as never a better case in which to apply this penalty for a Floyd died after Derek Chauvin put his weight on top of crime than arising from the murder of the unarmed and Floyd’s neck for more than eight minutes causing his handcuffed George Floyd by Derek Chauvin. death. Two other police were also on top of the Chauvin, had been subject of at least 18 previous handcuffed Floyd face-down with their body weight on his complaints against him since 2001 and all were lower body causing further pain and suffering, while a systematically whitewashed by the Minneapolis Police third, heavily armed officer made sure that no members Department. of the public intervened to stop the vicious attack. Had the law been properly applied in the first place, During this time, Floyd repeatedly asked the thug police George Floyd would not have been murdered by Derek officer Chauvin to get off him and at least fifteen times Chauvin. Floyd said “I can’t breath”. In terms of the 4 corrupt police who killed Chauvin, the One bystander told officers they needed to let him corruption did not stop there. breathe. Another yelled at them to check the man’s pulse The response of the Minneapolis Police Department was (Forliti and Baenen 2020). They were herded away from also coordinated to minimize damage to their murderer the scene by one of the police officers. cops from the outset. Chauvin only removed his body from on top of Floyd after Following a massive public uprising that caused over 30 it was clear Floyd was non-responsive and dead. police cars to be destroyed and a major Minneapolis Prior to this part of the attack being filmed by bystanders Police Station torched and burnt to the ground, the from both sides of the car, as well as two CCTV cameras Mineapolis Police charged Derek Chauvin with “third that also caught footage of the police bashing Floyd on degree murder”. The other three co-murderers were not the back seat of the police car, after which the attack was charged! continued with Floyd being removed from the police car In fact they were quietly shifted out of town with full police with his face slammed onto the hard road surface. protection. What a viewing of the video shows is that Chauvin was This degree of murder is a low-level of murder in US law. not just a single rogue cop. It borders on accidental murder, whereas first degree is a The four officers were working as a well organised team, planned killing. all knowing their well-defined roles and working together. However in line with such matters there is also little doubt Two were full body weight down holding handcuffed Floyd that the case will be sabotaged and Chauvin will walk onto the bitumen, without which, Chauvin would not have free in the same way that police who got caught on video been in a position to be able to put his foot on a fully bashing Rodney King in Los Angeles 3 March 1991. restrained Floyd to suffocate and kill him. The fourth This will be treated as a green light for more like-minded police officer as part of the team aggressively pushed, law-enforcement people to conduct similar acts of shoved and threatened concerned bystanders who brutality, lies, deception and murder in the future to wanted to assist the clearly severely injured and dying destroy the lives of yet more innocent victims. Floyd. Indication of the final result came when the Minnesota Derek Chauvin has also used his proceeds of crime to Police department released their “official” autopsy report amass a huge property portfolio spanning the United which claimed that Floyd had in fact died of “Coronavirus” States from California to Florida, which at this stage and not the bashing, with his death at the time of the seems safe and not likely to be seized on behalf of his bashing being nothing more than an unfortunate many victims. coincidence. State police were guarding his assets from So when you have 4 bad cops and a department with demonstrators to ensure none were attacked. 10,000 other cops who do nothing to stop the corruption, Floyd is one of millions of victims of police violence and lies and violence, you have an entire police department brutality and his death sparked massive demonstrations that is corrupt! That is 10,004 bad cops! of solidarity across the United States in over 30 major Meanwhile the USA President, Donald Trump got onto cities and other parts of the world including London, “Twitter” and tweeted to State Governors to get police to Berlin, Sydney and Melbourne. start shooting protesters, an instruction many actually Police buildings in California, Minnesota and Washington followed, adding a few dozen more people to the USA were torched as were countless police cars. death toll of people killed and maimed by their police, The government-controlled media in the United States already pumped up with over 100,000 coronavirus deaths and Australia, including the Murdoch Press and CNN, in the USA at the time Floyd was murdered. tried to paint the protests as black people versus white In the USA victims of police violence and corruption now people (see for example Bolt 2020), but this was never have the chants “I can’t breathe” and “George Floyd, say the case. his name!” Most of the protestors were in fact white people! Australian victims of police violence and corruption say The protest was against police corruption, police brutality, “lest we forget” which was a line borrowed from Nazi police lies and dishonesty and the destruction of innocent Holocaust survivors. See also the etymology for people’s lives, with murder being the most serious crime Kumanjayiwalkerus kumanjayi sp. nov. in this paper.

ALBOGIBBA GEN. NOV. Planusrususpes subgen. nov. which conforms to the LSIDurn:lsid:zoobank.org:act:CAA18748-21DC-4A71- above genus diagnosis, save for the following 859D-567F6548FB8D differences: The heel is without a tubercle, or in rare Type species: Hyla humeralis Boulenger, 1912. cases has a very small one. The skin is smooth above, Diagnosis: Species of Nyctimystes, Occultatahyla gen. granular below. A weak, wavy fold is present on the outer nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., surface of the forearm. The head is broader than long; Badiohyla gen. nov. and Albogibba gen. nov. are the canthus rostralis is not distinct; the loreal region is separated from all other Australasian (Australian and oblique; Unique to this subgenus is that the palpebral New Guinea) Tree Frogs (Pelodryadidae) by the following venation is distinct, but is reduced to individual pigment suite of characters: Large dark eyes, which have spots and thin, meandering lines, although the lines are vertically elliptical pupils and a lower eyelid with a still mainly oriented largely in a vertical direction, with few characteristic reticulum or palpebral venation of fine horizontal interconnections. pigmented lines. There is usually a distinct, crenulated Species within Nigreosoculus gen. nov. are separated skin fold along the hind edge of the forearm and the foot, from the other five genera (Nyctimystes, Occultatahyla the heel often having a small flap of skin. Males are gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. usually considerably smaller than females. and Albogibba gen. nov.) by the following suite of Nyctimystes Stejneger, 1916 as defined in this paper, are characters: Vocal slits present; a very small heel tubercle; separated from the other five genera (Occultatahyla gen. basal webbing on the hand; exposed tympanum; vertical nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., lines of palpebral reticulum oriented obliquely and with Badiohyla gen. nov. and Albogibba gen. nov.) by having relatively few horizontal cross-connections; pale-tan to sparse, broken vertically-oriented palpebral venation near black iris; rear of thighs barred/mottled with brown, across the nictitating membrane and in having adult caramel, or blue-gray. males without vocal slits. Species within Magnumoculus gen. nov. are separated Frogs in the subgenus Magnummanibus subgen. nov. are from the other five genera (Nyctimystes, Nigreosoculus separated from those in the nominate subgenus gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. Nyctimystes and the subgenus Asperohyla subgen. nov. and Albogibba gen. nov.) by the following suite of by their possession of very long, prominent heel lappets. characters: The head is broad (HL/HW <1.0), the snout high and blunt, the canthus rostralis distinct, broadly Frogs in the subgenus Asperohyla subgen. nov. are spaced nostrils (EN/IN ratio <1), and the loreal region separated from the other two subgenera by having a nearly vertical. A strong, slightly curved, supratympanic completely hidden tympanum and a dorsal skin fold begins at the posterior corner of the eye and roughened by tiny, conical asperities rather than by disappears above the insertion of the forelimb. The variable sized and more rounded irregularities as seen in tympanum is small but distinct. The hands have little some other New Guinea tree frog species; iris is brown in webbing; the outer fingers are about one-quarter webbed. life. The vomerine teeth are in two patches between the Species within Occultatahyla gen. nov. are separated internal nares, nearly on a line connecting the posterior from the other five genera (Nyctimystes, Nigreosoculus margins of the nares. The skin is minutely granular gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. above, coarser beneath. There is no tubercle on the heel. and Albogibba gen. nov.) by large size (adult females 50 A nearly straight, very slightly crenulated fold of skin is to 84 mm. in snout to vent length); internarial distance present on the outer surface of the forearm. A moderate distinctly smaller than distance from eye to naris (E-N/IN body size (40-60 mm S-V), predominantly brown averages 1.34 (range 1.2-1.5)); the head is broad, colouration, an eyelid venation composed of a network flattish, with depressed snout, closely spaced nares, and with numerous horizontal connections with relatively few oblique loreal region. Palpebral venation is in oblique vertical interconnections. There is decoration on the lines (more vertical than horizontal and with few forearm and tarsus in the form of rows of (often white- horizontal interconnections), with a few horizontal tipped) tubercles. interconnections. Species within Badiohyla gen. nov. are separated from A dermal fold passes from the posterior corner of the eye the other five genera (Nyctimystes, Nigreosoculus gen. over the upper edge of the tympanum and down to the nov., Occultatahyla gen. nov., Magnumoculus gen. nov. insertion of the forelimb, sometimes becoming obscure and Albogibba gen. nov.) by having a unique “reticulated” when leaving the tympanum. The outer fingers are palpebral venation, half-webbed fingers and a size in approximately half-webbed, the third and fifth toes males with SVL to maximum of 50 mm to 100 mm webbed to the disc. A light-coloured dermal ridge or depending on the species. They are further defined as series of tubercles extends from the elbow to the disc of follows: The snout is relatively short, blunt and high (E-N/ the fourth finger. A similar but less distinct ridge or line of IN 0.94), with distinct canthus rostralis and oblique loreal tubercles occupies the outer edge of the tarsus and fifth region. The vomerine teeth are in two patches between toe. A very small heel tubercle is present. The skin of the the posterior edges of the choanae. dorsum is smooth to very slightly granular, that of the The palpebral venation forms a heavy reticulum. The venter is always coarsely granulate. Slit-like, vocal sac tympanum is distinct and is separated from the eye by openings are usually present in the floor of the mouth. A approximately the diameter of the tympanum. A fold of small patch of very fine horny tubercles is found on the skin passes from the posterior corner of the eye, over the first finger of males. The iris is dark or black in colour. upper edge of the tympanum and down, becoming Exceptional to the preceding is the subgenus indistinct above the insertion of the forelimb. The skin of Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 115

the dorsum is minutely roughened, that of the venter Content: Albogibba humeralis (Boulenger, 1912) (type coarsely granular. A row of tubercles is present along the species); A. granti (Boulenger, 1914); A. ingens sp. nov.; outer edge of the forearm and there is a similar row on A. zweifeli (Tyler, 1967). the tarsus. There is only a slight suggestion of a heel RATIOBRUNNEIS SUBGEN. NOV. tubercle. The outer fingers are approximately half- LSIDurn:lsid:zoobank.org:act:AFD879FD-2C28-4032- webbed, the toes, except the fourth toe, are webbed to 9975-342A10E36B55 the disc. The body and head are light brown to purple Type species: Nyctimystes zweifeli Tyler, 1967. dorsally, either with irregular dark brown blotches or Diagnosis: Species of Nyctimystes, Occultatahyla gen. markings absent. The legs have a similar ground color, nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., with or without irregular darker spots and bands present Badiohyla gen. nov. and Albogibba gen. nov. are on the tibia. separated from all other Australasian (Australian and Species within Albogibba gen. nov. are separated from New Guinea) Tree Frogs (Pelodryadidae) by the following the other five genera (Nyctimystes, Nigreosoculus gen. suite of characters: nov., Occultatahyla gen. nov., Magnumoculus gen. nov. Large dark eyes, which have vertically elliptical pupils and Badiohyla gen. nov.) by one or other of the following and a lower eyelid with a characteristic reticulum or unique suites of characters: palpebral venation of fine pigmented lines. There is 1/ Being a large species (males to 100 mm. in snout to usually a distinct, crenulated skin fold along the hind vent length); of a uniform dorsal coloration (green in life, edge of the forearm and the foot, the heel often having a purple in alcohol-preserved specimens); palpebral small flap of skin. Males are usually considerably smaller venation is a reticulum without obvious orientation and in than females. the nominate subgenus of this genus, the male bears a spine-like process on the anterior surface of the proximal Nyctimystes Stejneger, 1916 as defined in this paper, are part of the humerus (upper arm) (nominate subgenus separated from the other five genera (Occultatahyla gen. Albogibba subgen. nov.), or: nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. and Albogibba gen. nov.) by having 2/ As above, but male SVL of up to 80 mm; not including sparse, broken vertically-oriented palpebral venation a spine-like process on the anterior surface of the across the nictitating membrane and in having adult proximal part of the humerus (upper arm) in males and males without vocal slits. with a unique dorsal pattern of black vermiculations on the body and limbs, with unmarked greyish lower flanks Frogs in the subgenus Magnummanibus subgen. nov. are (Ratiobrunneis subgen. nov.). separated from those in the nominate subgenus Nyctimystes and the subgenus Asperohyla subgen. nov. According to Duellman et al. (2016), each of the six by their possession of very long, prominent heel lappets. preceding genera had the following divergences from one another. Frogs in the subgenus Asperohyla subgen. nov. are separated from the other two subgenera by having a Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its completely hidden tympanum and a dorsal skin nearest living relatives in other genera. roughened by tiny, conical asperities rather than by The subgenus Magnummanibus subgen. nov. diverged variable sized and more rounded irregularities as seen in 11.5 MYA from its nearest living relatives in the nominate some other New Guinea tree frog species; iris is brown in subgenus. life. Occultatahyla gen. nov. diverged 16.7 MYA from its Species within Occultatahyla gen. nov. are separated nearest living relatives in other genera. from the other five genera (Nyctimystes, Nigreosoculus Nigreosoculus gen. nov. diverged 14.8 MYA from its gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. nearest living relatives in other genera. and Albogibba gen. nov.) by large size (adult females 50 Magnumoculus gen. nov. diverged 13 MYA from its to 84 mm. in snout to vent length); internarial distance nearest living relatives, being Badiohyla gen. nov.. distinctly smaller than distance from eye to naris (E-N/IN Badiohyla gen. nov. diverged 13 MYA from its nearest averages 1.34 (range 1.2-1.5)); the head is broad, living relatives being Magnumoculus gen. nov.. flattish, with depressed snout, closely spaced nares, and In turn both Magnumoculus gen. nov. and Badiohyla gen. oblique loreal region. Palpebral venation is in oblique nov. as a pair diverged from their nearest living relatives lines (more vertical than horizontal and with few 22 MYA being Nyctimystes and other genera. horizontal interconnections), with a few horizontal Albogibba gen. nov. diverged 16.7 MYA from its nearest interconnections. A dermal fold passes from the posterior living relatives in other genera. corner of the eye over the upper edge of the tympanum The subgenus Ratiobrunneis subgen. nov. diverged 10 and down to the insertion of the forelimb, sometimes MYA from its nearest living relatives in the nominate becoming obscure when leaving the tympanum. The subgenus. outer fingers are approximately half-webbed, the third and fifth toes webbed to the disc. A light-coloured dermal Distribution: Distributed on the northern and southern ridge or series of tubercles extends from the elbow to the flanks of the central mountain ranges of New Guinea, disc of the fourth finger. A similar but less distinct ridge or from approximately 138 deg E. to 147 deg E., New line of tubercles occupies the outer edge of the tarsus Guinea, usually at 600-1700 metres elevation. and fifth toe. A very small heel tubercle is present. Etymology: “Albogibba” in Latin means “white lumps”, The skin of the dorsum is smooth to very slightly granular, with the small raised white tipped tubercles or lumps on that of the venter is always coarsely granulate. Slit-like, the lower flanks being characteristic of the type species. vocal sac openings are usually present in the floor of the

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 116 Australasian Journal of Herpetology mouth. A small patch of very fine horny tubercles is found tympanum is distinct and is separated from the eye by on the first finger of males. The iris is dark or black in approximately the diameter of the tympanum. A fold of colour. skin passes from the posterior corner of the eye, over the Exceptional to the preceding is the subgenus upper edge of the tympanum and down, becoming Planusrususpes subgen. nov. which conforms to the indistinct above the insertion of the forelimb. The skin of above genus diagnosis, save for the following the dorsum is minutely roughened, that of the venter differences: The heel is without a tubercle, or in rare coarsely granular. A row of tubercles is present along the cases has a very small one. The skin is smooth above, outer edge of the forearm and there is a similar row on granular below. A weak, wavy fold is present on the outer the tarsus. There is only a slight suggestion of a heel surface of the forearm. The head is broader than long; tubercle. The outer fingers are approximately half- the canthus rostralis is not distinct; the loreal region is webbed, the toes, except the fourth toe, are webbed to oblique; Unique to this subgenus is that the palpebral the disc. The body and head are light brown to purple venation is distinct, but is reduced to individual pigment dorsally, either with irregular dark brown blotches or spots and thin, meandering lines, although the lines are markings absent. The legs have a similar ground color, still mainly oriented largely in a vertical direction, with few with or without irregular darker spots and bands present horizontal interconnections. on the tibia. Species within Nigreosoculus gen. nov. are separated Species within Albogibba gen. nov. are separated from from the other five genera (Nyctimystes, Occultatahyla the other five genera (Nyctimystes, Nigreosoculus gen. gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. nov., Occultatahyla gen. nov., Magnumoculus gen. nov. and Albogibba gen. nov.) by the following suite of and Badiohyla gen. nov.) by one or other of the following characters: Vocal slits present; a very small heel tubercle; unique suites of characters: basal webbing on the hand; exposed tympanum; vertical 1/ Being a large species (males to 100 mm. in snout to lines of palpebral reticulum oriented obliquely and with vent length); of a uniform dorsal coloration (green in life, relatively few horizontal cross-connections; pale-tan to purple in alcohol-preserved specimens); palpebral near black iris; rear of thighs barred/mottled with brown, venation is a reticulum without obvious orientation and in caramel, or blue-gray. the nominate subgenus of this genus, the male bears a Species within Magnumoculus gen. nov. are separated spine-like process on the anterior surface of the proximal from the other five genera (Nyctimystes, Nigreosoculus part of the humerus (upper arm) (nominate subgenus gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. Albogibba subgen. nov.), or: and Albogibba gen. nov.) by the following suite of 2/ As above, but male SVL of up to 80 mm; not including characters: The head is broad (HL/HW <1.0), the snout a spine-like process on the anterior surface of the high and blunt, the canthus rostralis distinct, broadly proximal part of the humerus (upper arm) in males and spaced nostrils (EN/IN ratio <1), and the loreal region with a unique dorsal pattern of black vermiculations on nearly vertical. A strong, slightly curved, supratympanic the body and limbs, with unmarked greyish lower flanks fold begins at the posterior corner of the eye and (Ratiobrunneis subgen. nov.). disappears above the insertion of the forelimb. The According to Duellman et al. (2016), each of the six tympanum is small but distinct. The hands have little preceding genera had the following divergences from one webbing; the outer fingers are about one-quarter webbed. another. The vomerine teeth are in two patches between the Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its internal nares, nearly on a line connecting the posterior nearest living relatives in other genera. margins of the nares. The skin is minutely granular The subgenus Magnummanibus subgen. nov. diverged above, coarser beneath. There is no tubercle on the heel. 11.5 MYA from its nearest living relatives in the nominate A nearly straight, very slightly crenulated fold of skin is subgenus. present on the outer surface of the forearm. A moderate Occultatahyla gen. nov. diverged 16.7 MYA from its body size (40-60 mm S-V), predominantly brown nearest living relatives in other genera. colouration, an eyelid venation composed of a network Nigreosoculus gen. nov. diverged 14.8 MYA from its with numerous horizontal connections with relatively few nearest living relatives in other genera. vertical interconnections. There is decoration on the forearm and tarsus in the form of rows of (often white- Magnumoculus gen. nov. diverged 13 MYA from its tipped) tubercles. nearest living relatives, being Badiohyla gen. nov.. Species within Badiohyla gen. nov. are separated from Badiohyla gen. nov. diverged 13 MYA from its nearest the other five genera (Nyctimystes, Nigreosoculus gen. living relatives being Magnumoculus gen. nov.. nov., Occultatahyla gen. nov., Magnumoculus gen. nov. In turn both Magnumoculus gen. nov. and Badiohyla gen. and Albogibba gen. nov.) by having a unique “reticulated” nov. as a pair diverged from their nearest living relatives palpebral venation, half-webbed fingers and a size in 22 MYA being Nyctimystes and other genera. males with SVL to maximum of 50 mm to 100 mm Albogibba gen. nov. diverged 16.7 MYA from its nearest depending on the species. They are further defined as living relatives in other genera. follows: The snout is relatively short, blunt and high (E-N/ The subgenus Ratiobrunneis subgen. nov. diverged 10 IN 0.94), with distinct canthus rostralis and oblique loreal MYA from its nearest living relatives in the nominate region. The vomerine teeth are in two patches between subgenus. the posterior edges of the choanae. Distribution: The two species in the subgenus The palpebral venation forms a heavy reticulum. The Ratiobrunneis subgen. nov. are known from West Sepik,

Chimbu, Southern Highlands and Western Provinces Both A. ingens sp. nov. and A. humeralis comprising the (Papua New Guinea) as well as on the Utakwa River, entirety of the nominate subgenus Albogibba gen. nov. Eipomek (Papua, Indonesia). are separated from the two species in the other Etymology: In Latin “Ratiobrunneis” means brown subgenus Ratiobrunneis subgen. nov. by the following: pattern, which is exactly the character state of the dorsal Being a large species (males to 100 mm. in snout to vent surface of these frogs. length); of a uniform dorsal coloration (green in life, Content: Badiohyla (Ratiobrunneis) zweifeli (Tyler, 1967) purple in alcohol-preserved specimens); palpebral (type species); B. (Ratiobrunneis) granti (Boulenger, venation is a reticulum without obvious orientation and in 1914). the nominate subgenus of this genus (these two ALBOGIBBA (ALBOGIBBA) INGENS SP. NOV. species), the male bears a spine-like process on the anterior surface of the proximal part of the humerus LSIDurn:lsid:zoobank.org:act:DFF9FB1D-9B0B-4AB3- (upper arm). B787-188F6417AB0A These characters also separate A. ingens sp. nov. and A. Holotype: A preserved specimen at the American humeralis from all other tree frogs in Australia and New Museum of Natural History, Manhatten, New York, USA, Guinea. reptile collection, specimen number AMNH 49691 Ratiobrunneis subgen. nov. are similar in most respects collected from 6 kilometers southwest of the Bernhard to A. ingens sp. nov. and A. humeralis but are separated Camp, 1200 meters, Idenburg River region, Irian Jaya, as follows: Male SVL of up to 80 mm; not including a Indonesia, Latitude -3.48 S., Longitude 139.2 E. spine-like process on the anterior surface of the proximal This facility allows access to its holdings. part of the humerus (upper arm) in males and with a Paratypes: Two preserved specimens at the American unique dorsal pattern of black vermiculations on the body Museum of Natural History, Manhatten, New York, USA, and limbs, with unmarked greyish lower flanks. reptile collection, specimen numbers AMNH 49695 and Photos of a male A. ingens sp. nov. in life from near the AMNH 49697 collected from 4 kilometers southwest of type locality can be found at: the Bernhard Camp, 1200 meters, Idenburg River region, https://www.naturepl.com/stock-photo-tree-frog-nature- Irian Jaya, Indonesia, Latitude -3.48 S., Longitude 139.2 image01361627.html E. and Diagnosis: Albogibba ingens sp. nov. has until now been treated as a north-west population of A. humeralis https://timlaman.photoshelter.com/image/ (Boulenger, 1912), known in most texts as Nyctimystes I0000WOJyFmCZ70g humeralis (e.g. Zweifel 1958 or Menzies 2006). A photo of a female A. humeralis in life can be found in Both would key out as A. humeralis in both Zweifel (1958) Menzies (1976) on plate 8 at top left and Menzies (2006) or Menzies (2006). at photo 90 (same image). A. ingens sp. nov. is readily separated from A. humeralis Distribution: A. ingens sp. nov. is distributed on the by the following suite of characters: Iris is reddish-brown; northern side of the central cordillera of New Guinea west white tubercles on the flanks are broad and blunt in of the Huon Peninsula in Papua New guinea, at least as profile and sometimes yellowish instead of white; flanks far west as the Mamberamo River in Irian Jaya. A. and venter are white or yellow; upper lip has a thick humeralis occurs south of the central cordillera, including yellow or white bar of consistent thickness running from in Central Province, Papua New guinea. tip of snout along top of mouth; fingers yellow or white Etymology: “Ingens” is Latin for “huge”, or “big”. As the and toes orangeish. Dorsum usually a brilliant emerald adults of this species are very large, the species name is green. Dorsum and legs smooth and covered with appropriate. numerous tiny evenly spaced granules. Forearms have OCCULTATAHYLA GEN. NOV. numerous tubercles. LSIDurn:lsid:zoobank.org:act:1CDD42D2-B7BC-4F75- By contrast A. humeralis has a brown iris; white tubercles ABF0-F57A8383CFC2 on flanks are not broad or flattened, but well defined and Type species: Nyctimystes semipalmata Parker, 1936. more-or-less conical in shape or pointed. There is no Diagnosis: Species of Nyctimystes, Occultatahyla gen. thick yellow bar of consistent thickness running from tip nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., of snout along top of mouth, but instead either simply the Badiohyla gen. nov. and Albogibba gen. nov. are normal greenish colour of the upper surface, or separated from all other Australasian (Australian and sometimes a very thin, narrow white edge (hairline). Toes New Guinea) Tree Frogs (Pelodryadidae) by the following are greenish and pads light green or sometimes suite of characters: yellowish in males. Venter is whitish. Dorsum varies in Large dark eyes, which have vertically elliptical pupils colour, but is invariably green, sometimes with indistinct and a lower eyelid with a characteristic reticulum or mottling and only sometimes covered with numerous tiny palpebral venation of fine pigmented lines. evenly spaced granules. There is usually a distinct, crenulated skin fold along the Forearms not with numerous tubercles. hind edge of the forearm and the foot, the heel often Both A. ingens sp. nov. and A. humeralis sometimes having a small flap of skin. have scattered yellow spots or rings on the dorsum and Males are usually considerably smaller than females. sometimes a red, brown or violet flush on the flanks. Nyctimystes Stejneger, 1916 as defined in this paper, are Males usually have a yellowish throat, versus cream or separated from the other five genera (Occultatahyla gen. white in females.

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 118 Australasian Journal of Herpetology nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., relatively few horizontal cross-connections; pale-tan to Badiohyla gen. nov. and Albogibba gen. nov.) by having near black iris; rear of thighs barred/mottled with brown, sparse, broken vertically-oriented palpebral venation caramel, or blue-gray. across the nictitating membrane and in having adult Species within Magnumoculus gen. nov. are separated males without vocal slits. from the other five genera (Nyctimystes, Nigreosoculus Frogs in the subgenus Magnummanibus subgen. nov. are gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. separated from those in the nominate subgenus and Albogibba gen. nov.) by the following suite of Nyctimystes and the subgenus Asperohyla subgen. nov. characters: The head is broad (HL/HW <1.0), the snout by their possession of very long, prominent heel lappets. high and blunt, the canthus rostralis distinct, broadly Frogs in the subgenus Asperohyla subgen. nov. are spaced nostrils (EN/IN ratio <1), and the loreal region separated from the other two subgenera by having a nearly vertical. A strong, slightly curved, supratympanic completely hidden tympanum and a dorsal skin fold begins at the posterior corner of the eye and roughened by tiny, conical asperities rather than by disappears above the insertion of the forelimb. The variable sized and more rounded irregularities as seen in tympanum is small but distinct. The hands have little some other New Guinea tree frog species; iris is brown in webbing; the outer fingers are about one-quarter webbed. life. The vomerine teeth are in two patches between the Species within Occultatahyla gen. nov. are separated internal nares, nearly on a line connecting the posterior from the other five genera (Nyctimystes, Nigreosoculus margins of the nares. The skin is minutely granular gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. above, coarser beneath. There is no tubercle on the heel. and Albogibba gen. nov.) by large size (adult females 50 A nearly straight, very slightly crenulated fold of skin is to 84 mm. in snout to vent length); internarial distance present on the outer surface of the forearm. A moderate distinctly smaller than distance from eye to naris (E-N/IN body size (40-60 mm S-V), predominantly brown averages 1.34 (range 1.2-1.5)); the head is broad, colouration, an eyelid venation composed of a network flattish, with depressed snout, closely spaced nares, and with numerous horizontal connections with relatively few oblique loreal region. Palpebral venation is in oblique vertical interconnections. There is decoration on the lines (more vertical than horizontal and with few forearm and tarsus in the form of rows of (often white- horizontal interconnections), with a few horizontal tipped) tubercles. interconnections. A dermal fold passes from the posterior Species within Badiohyla gen. nov. are separated from corner of the eye over the upper edge of the tympanum the other five genera (Nyctimystes, Nigreosoculus gen. and down to the insertion of the forelimb, sometimes nov., Occultatahyla gen. nov., Magnumoculus gen. nov. becoming obscure when leaving the tympanum. The and Albogibba gen. nov.) by having a unique “reticulated” outer fingers are approximately half-webbed, the third palpebral venation, half-webbed fingers and a size in and fifth toes webbed to the disc. A light-coloured dermal males with SVL to maximum of 50 mm to 100 mm ridge or series of tubercles extends from the elbow to the depending on the species. They are further defined as disc of the fourth finger. A similar but less distinct ridge or follows: The snout is relatively short, blunt and high (E-N/ line of tubercles occupies the outer edge of the tarsus IN 0.94), with distinct canthus rostralis and oblique loreal and fifth toe. A very small heel tubercle is present. The region. The vomerine teeth are in two patches between skin of the dorsum is smooth to very slightly granular, that the posterior edges of the choanae. of the venter is always coarsely granulate. Slit-like, vocal The palpebral venation forms a heavy reticulum. The sac openings are usually present in the floor of the tympanum is distinct and is separated from the eye by mouth. A small patch of very fine horny tubercles is found approximately the diameter of the tympanum. A fold of on the first finger of males. The iris is dark or black in skin passes from the posterior corner of the eye, over the colour. upper edge of the tympanum and down, becoming Exceptional to the preceding is the subgenus Webpede indistinct above the insertion of the forelimb. The skin of subgen. nov. which conforms to the above genus the dorsum is minutely roughened, that of the venter diagnosis, save for the following differences: The heel is coarsely granular. A row of tubercles is present along the without a tubercle, or in rare cases has a very small one. outer edge of the forearm and there is a similar row on The skin is smooth above, granular below. A weak, wavy the tarsus. There is only a slight suggestion of a heel fold is present on the outer surface of the forearm. The tubercle. The outer fingers are approximately half- head is broader than long; the canthus rostralis is not webbed, the toes, except the fourth toe, are webbed to distinct; the loreal region is oblique; Unique to this the disc. subgenus is that the palpebral venation is distinct, but is The body and head are light brown to purple dorsally, reduced to individual pigment spots and thin, meandering either with irregular dark brown blotches or markings lines, although the lines are still mainly oriented largely in absent. The legs have a similar ground color, with or a vertical direction, with few horizontal interconnections. without irregular darker spots and bands present on the Species within Nigreosoculus gen. nov. are separated tibia. from the other five genera (Nyctimystes, Occultatahyla Species within Albogibba gen. nov. are separated from gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. the other five genera (Nyctimystes, Nigreosoculus gen. and Albogibba gen. nov.) by the following suite of nov., Occultatahyla gen. nov., Magnumoculus gen. nov. characters: Vocal slits present; a very small heel tubercle; and Badiohyla gen. nov.) by one or other of the following basal webbing on the hand; exposed tympanum; vertical unique suites of characters: lines of palpebral reticulum oriented obliquely and with 1/ Being a large species (males to 100 mm. in snout to

vent length); of a uniform dorsal coloration (green in life, skin fold along the hind edge of the forearm and the foot, purple in alcohol-preserved specimens); palpebral the heel often having a small flap of skin. Males are venation is a reticulum without obvious orientation and in usually considerably smaller than females. the nominate subgenus of this genus, the male bears a Species within Occultatahyla gen. nov. are separated spine-like process on the anterior surface of the proximal from the other five genera (Nyctimystes, Nigreosoculus part of the humerus (upper arm) (nominate subgenus gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. Albogibba subgen. nov.), or: and Albogibba gen. nov.) by large size (adult females 50 2/ As above, but male SVL of up to 80 mm; not including to 84 mm. in snout to vent length); internarial distance a spine-like process on the anterior surface of the distinctly smaller than distance from eye to naris (E-N/IN proximal part of the humerus (upper arm) in males and averages 1.34 (range 1.2-1.5)); the head is broad, with a unique dorsal pattern of black vermiculations on flattish, with depressed snout, closely spaced nares, and the body and limbs, with unmarked greyish lower flanks oblique loreal region. Palpebral venation is in oblique (Ratiobrunneis subgen. nov.). lines (more vertical than horizontal and with few According to Duellman et al. (2016), each of the six horizontal interconnections), with a few horizontal preceding genera had the following divergences from one interconnections. A dermal fold passes from the posterior another. corner of the eye over the upper edge of the tympanum Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its and down to the insertion of the forelimb, sometimes nearest living relatives in other genera. becoming obscure when leaving the tympanum. The The subgenus Magnummanibus subgen. nov. diverged outer fingers are approximately half-webbed, the third 11.5 MYA from its nearest living relatives in the nominate and fifth toes webbed to the disc. A light-coloured dermal subgenus. ridge or series of tubercles extends from the elbow to the disc of the fourth finger. A similar but less distinct ridge or Occultatahyla gen. nov. diverged 16.7 MYA from its line of tubercles occupies the outer edge of the tarsus nearest living relatives in other genera. and fifth toe. A very small heel tubercle is present. The Nigreosoculus gen. nov. diverged 14.8 MYA from its skin of the dorsum is smooth to very slightly granular, that nearest living relatives in other genera. of the venter is always coarsely granulate. Slit-like, vocal Magnumoculus gen. nov. diverged 13 MYA from its sac openings are usually present in the floor of the nearest living relatives, being Badiohyla gen. nov.. mouth. A small patch of very fine horny tubercles is found Badiohyla gen. nov. diverged 13 MYA from its nearest on the first finger of males. The iris is dark or black in living relatives being Magnumoculus gen. nov.. colour. In turn both Magnumoculus gen. nov. and Badiohyla gen. Exceptional to the preceding is the subgenus Webpede nov. as a pair diverged from their nearest living relatives subgen. nov. which conforms to the above genus 22 MYA being Nyctimystes and other genera. diagnosis, save for the following differences: The heel is Albogibba gen. nov. diverged 16.7 MYA from its nearest without a tubercle, or in rare cases has a very small one. living relatives in other genera. The skin is smooth above, granular below. A weak, wavy The subgenus Ratiobrunneis subgen. nov. diverged 10 fold is present on the outer surface of the forearm. The MYA from its nearest living relatives in the nominate head is broader than long; the canthus rostralis is not subgenus. distinct; the loreal region is oblique; Unique to this Distribution: New Guinea and immediately adjacent subgenus is that the palpebral venation is distinct, but is islands on the continental shelf. reduced to individual pigment spots and thin, meandering lines, although the lines are still mainly oriented largely in Etymology: In Latin, “Occultata” means “hidden”, and a vertical direction, with few horizontal interconnections. with “Hyla” meaning “tree frog”, the genus name in effect Fingers with much webbing, the outer finger webbed means hidden tree frog. This reflects the excellent almost to disc. camoflague in the wild state of these species. According to Duellman et al. (2016), Occultatahyla gen. Content: Occultatahyla semipalmatus (Parker, 1936) nov. diverged 16.7 MYA from its nearest living relatives in (type species); O. avocalis (Zweifel, 1958); O. daymani other genera. (Zweifel, 1958); O. fluviatilis (Zweifel, 1958); O. foricula (Tyler, 1963); O. perimetri (Zweifel, 1958); O. rueppelli Nyctimystes Stejneger, 1916 as defined in this paper, are (Boettger, 1895). separated from the other five genera (Occultatahyla gen. nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., WEBPEDE SUBGEN. NOV. Badiohyla gen. nov. and Albogibba gen. nov.) by having LSIDurn:lsid:zoobank.org:act:96A567B5-A1A4-44DF- sparse, broken vertically-oriented palpebral venation 8336-6B97080036D8 across the nictitating membrane and in having adult Type species: Hyla rueppelli Boettger, 1895. males without vocal slits. Diagnosis: Species of Nyctimystes, Occultatahyla gen. Frogs in the subgenus Magnummanibus subgen. nov. are nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., separated from those in the nominate subgenus Badiohyla gen. nov. and Albogibba gen. nov. are Nyctimystes and the subgenus Asperohyla subgen. nov. separated from all other Australasian (Australian and by their possession of very long, prominent heel lappets. New Guinea) Tree Frogs (Pelodryadidae) by the following Frogs in the subgenus Asperohyla subgen. nov. are suite of characters: Large dark eyes, which have separated from the other two subgenera by having a vertically elliptical pupils and a lower eyelid with a completely hidden tympanum and a dorsal skin characteristic reticulum or palpebral venation of fine roughened by tiny, conical asperities rather than by pigmented lines. There is usually a distinct, crenulated Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 120 Australasian Journal of Herpetology variable sized and more rounded irregularities as seen in interconnections. A dermal fold passes from the posterior some other New Guinea tree frog species; iris is brown in corner of the eye over the upper edge of the tympanum life. and down to the insertion of the forelimb, sometimes The other four genera including species previously becoming obscure when leaving the tympanum. The treated as being within Nyctimystes are dealt with outer fingers are approximately half-webbed, the third elsewhere in this paper. and fifth toes webbed to the disc. A light-coloured dermal Distribution: The single species constituting this ridge or series of tubercles extends from the elbow to the subgenus is known only from Halmahera and Morotai disc of the fourth finger. A similar but less distinct ridge or Islands in the Moluccas, Indonesia. line of tubercles occupies the outer edge of the tarsus Etymology: “Webpede” is derived from the Latin words and fifth toe. A very small heel tubercle is present. The “web” meaning “web” in English and the Latin “Pede” skin of the dorsum is smooth to very slightly granular, that meaning “paw”, or in the context of a frog, the hand or of the venter is always coarsely granulate. Slit-like, vocal front foot. sac openings are usually present in the floor of the mouth. A small patch of very fine horny tubercles is found Content: Occultatahyla (Webpede) rueppelli (Boettger, on the first finger of males. The iris is dark or black in 1895) (monotypic). colour. NIGREOSOCULUS GEN. NOV. Exceptional to the preceding is the subgenus LSIDurn:lsid:zoobank.org:act:8FB6CBD7-F9BC- Planusrususpes subgen. nov. which conforms to the 4C0C-8D2A-071067F436C2 above genus diagnosis, save for the following Type species: Nyctimystes cheesmani Tyler, 1964. differences: The heel is without a tubercle, or in rare Diagnosis: Species of Nyctimystes, Occultatahyla gen. cases has a very small one. The skin is smooth above, nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., granular below. A weak, wavy fold is present on the outer Badiohyla gen. nov. and Albogibba gen. nov. are surface of the forearm. The head is broader than long; separated from all other Australasian (Australian and the canthus rostralis is not distinct; the loreal region is New Guinea) Tree Frogs (Pelodryadidae) by the following oblique; Unique to this subgenus is that the palpebral suite of characters: venation is distinct, but is reduced to individual pigment Large dark eyes, which have vertically elliptical pupils spots and thin, meandering lines, although the lines are and a lower eyelid with a characteristic reticulum or still mainly oriented largely in a vertical direction, with few palpebral venation of fine pigmented lines. There is horizontal interconnections. usually a distinct, crenulated skin fold along the hind Species within Nigreosoculus gen. nov. are separated edge of the forearm and the foot, the heel often having a from the other five genera (Nyctimystes, Occultatahyla small flap of skin. Males are usually considerably smaller gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. than females. and Albogibba gen. nov.) by the following suite of Nyctimystes Stejneger, 1916 as defined in this paper, are characters: Vocal slits present; a very small heel tubercle; separated from the other five genera (Occultatahyla gen. basal webbing on the hand; exposed tympanum; vertical nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., lines of palpebral reticulum oriented obliquely and with Badiohyla gen. nov. and Albogibba gen. nov.) by having relatively few horizontal cross-connections; pale-tan to sparse, broken vertically-oriented palpebral venation near black iris; rear of thighs barred/mottled with brown, across the nictitating membrane and in having adult caramel, or blue-gray. males without vocal slits. Species within Magnumoculus gen. nov. are separated Frogs in the subgenus Magnummanibus subgen. nov. are from the other five genera (Nyctimystes, Nigreosoculus separated from those in the nominate subgenus gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. Nyctimystes and the subgenus Asperohyla subgen. nov. and Albogibba gen. nov.) by the following suite of by their possession of very long, prominent heel lappets. characters: The head is broad (HL/HW <1.0), the snout Frogs in the subgenus Asperohyla subgen. nov. are high and blunt, the canthus rostralis distinct, broadly separated from the other two subgenera by having a spaced nostrils (EN/IN ratio <1), and the loreal region completely hidden tympanum and a dorsal skin nearly vertical. A strong, slightly curved, supratympanic roughened by tiny, conical asperities rather than by fold begins at the posterior corner of the eye and variable sized and more rounded irregularities as seen in disappears above the insertion of the forelimb. The some other New Guinea tree frog species; iris is brown in tympanum is small but distinct. The hands have little life. webbing; the outer fingers are about one-quarter webbed. Species within Occultatahyla gen. nov. are separated The vomerine teeth are in two patches between the from the other five genera (Nyctimystes, Nigreosoculus internal nares, nearly on a line connecting the posterior gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. margins of the nares. The skin is minutely granular and Albogibba gen. nov.) by large size (adult females 50 above, coarser beneath. There is no tubercle on the heel. to 84 mm. in snout to vent length); internarial distance A nearly straight, very slightly crenulated fold of skin is distinctly smaller than distance from eye to naris (E-N/IN present on the outer surface of the forearm. A moderate averages 1.34 (range 1.2-1.5)); the head is broad, body size (40-60 mm S-V), predominantly brown flattish, with depressed snout, closely spaced nares, and colouration, an eyelid venation composed of a network oblique loreal region. Palpebral venation is in oblique with numerous horizontal connections with relatively few lines (more vertical than horizontal and with few vertical interconnections. There is decoration on the horizontal interconnections), with a few horizontal forearm and tarsus in the form of rows of (often white- tipped) tubercles.

Species within Badiohyla gen. nov. are separated from Badiohyla gen. nov. diverged 13 MYA from its nearest the other five genera (Nyctimystes, Nigreosoculus gen. living relatives being Magnumoculus gen. nov.. nov., Occultatahyla gen. nov., Magnumoculus gen. nov. In turn both Magnumoculus gen. nov. and Badiohyla gen. and Albogibba gen. nov.) by having a unique “reticulated” nov. as a pair diverged from their nearest living relatives palpebral venation, half-webbed fingers and a size in 22 MYA being Nyctimystes and other genera. males with SVL to maximum of 50 mm to 100 mm Albogibba gen. nov. diverged 16.7 MYA from its nearest depending on the species. They are further defined as living relatives in other genera. follows: The snout is relatively short, blunt and high (E-N/ The subgenus Ratiobrunneis subgen. nov. diverged 10 IN 0.94), with distinct canthus rostralis and oblique loreal MYA from its nearest living relatives in the nominate region. The vomerine teeth are in two patches between subgenus. the posterior edges of the choanae. Distribution: New Guinea, including offshore islands on The palpebral venation forms a heavy reticulum. The the continental shelf. tympanum is distinct and is separated from the eye by Etymology: In Latin Nigreosoculus means black eye, in approximately the diameter of the tympanum. A fold of reflection of the near black iris in some species. skin passes from the posterior corner of the eye, over the upper edge of the tympanum and down, becoming Content: Nigreosoculus cheesmani (Tyler, 1964) (type indistinct above the insertion of the forelimb. The skin of species); N. bivocalis (Kraus, 2012); N. calcaratus the dorsum is minutely roughened, that of the venter (Menzies, 2014); N. eucavatus (Menzies, 2014); N. coarsely granular. A row of tubercles is present along the intercastellus (Kraus, 2012); N. kuduki (Richards, 2007); outer edge of the forearm and there is a similar row on N. latratus (Menzies, 2014); N. montana (Peters and the tarsus. There is only a slight suggestion of a heel Doria, 1878); N. obsoletus (Lönnberg, 1900); N. tubercle. The outer fingers are approximately half- persimilis (Zweifel, 1958); N. traunae (Menzies, 2014). webbed, the toes, except the fourth toe, are webbed to BADIOHYLINA SUBTRIBE NOV. the disc. BADIOHYLA GEN. NOV. The body and head are light brown to purple dorsally, LSIDurn:lsid:zoobank.org:act:D0D7D60F-CD8F-454D- either with irregular dark brown blotches or markings 9EBB-357AEF6BF5CD absent. The legs have a similar ground color, with or Type species: Nyctimystes kubori Zweifel, 1958. without irregular darker spots and bands present on the Diagnosis: Species of Nyctimystes, Occultatahyla gen. tibia. nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., Species within Albogibba gen. nov. are separated from Badiohyla gen. nov. and Albogibba gen. nov. are the other five genera (Nyctimystes, Nigreosoculus gen. separated from all other Australasian (Australian and nov., Occultatahyla gen. nov., Magnumoculus gen. nov. New Guinea) Tree Frogs (Pelodryadidae) by the following and Badiohyla gen. nov.) by one or other of the following suite of characters: unique suites of characters: Large dark eyes, which have vertically elliptical pupils 1/ Being a large species (males to 100 mm. in snout to and a lower eyelid with a characteristic reticulum or vent length); of a uniform dorsal coloration (green in life, palpebral venation of fine pigmented lines. purple in alcohol-preserved specimens); palpebral There is usually a distinct, crenulated skin fold along the venation is a reticulum without obvious orientation and in hind edge of the forearm and the foot, the heel often the nominate subgenus of this genus, the male bears a having a small flap of skin. Males are usually spine-like process on the anterior surface of the proximal considerably smaller than females. part of the humerus (upper arm) (nominate subgenus Nyctimystes Stejneger, 1916 as defined in this paper, are Albogibba subgen. nov.), or: separated from the other five genera (Occultatahyla gen. 2/ As above, but male SVL of up to 80 mm; not including nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., a spine-like process on the anterior surface of the Badiohyla gen. nov. and Albogibba gen. nov.) by having proximal part of the humerus (upper arm) in males and sparse, broken vertically-oriented palpebral venation with a unique dorsal pattern of black vermiculations on across the nictitating membrane and in having adult the body and limbs, with unmarked greyish lower flanks males without vocal slits. (Ratiobrunneis subgen. nov.). Frogs in the subgenus Magnummanibus subgen. nov. are According to Duellman et al. (2016), each of the six separated from those in the nominate subgenus preceding genera had the following divergences from one Nyctimystes and the subgenus Asperohyla subgen. nov. another. by their possession of very long, prominent heel lappets. Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its Frogs in the subgenus Asperohyla subgen. nov. are nearest living relatives in other genera. separated from the other two subgenera by having a The subgenus Magnummanibus subgen. nov. diverged completely hidden tympanum and a dorsal skin 11.5 MYA from its nearest living relatives in the nominate roughened by tiny, conical asperities rather than by subgenus. variable sized and more rounded irregularities as seen in Occultatahyla gen. nov. diverged 16.7 MYA from its some other New Guinea tree frog species; iris is brown in nearest living relatives in other genera. life. Nigreosoculus gen. nov. diverged 14.8 MYA from its Species within Occultatahyla gen. nov. are separated nearest living relatives in other genera. from the other five genera (Nyctimystes, Nigreosoculus Magnumoculus gen. nov. diverged 13 MYA from its gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. nearest living relatives, being Badiohyla gen. nov.. and Albogibba gen. nov.) by large size (adult females 50

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 122 Australasian Journal of Herpetology to 84 mm. in snout to vent length); internarial distance present on the outer surface of the forearm. A moderate distinctly smaller than distance from eye to naris (E-N/IN body size (40-60 mm S-V), predominantly brown averages 1.34 (range 1.2-1.5)); the head is broad, colouration, an eyelid venation composed of a network flattish, with depressed snout, closely spaced nares, and with numerous horizontal connections with relatively few oblique loreal region. Palpebral venation is in oblique vertical interconnections. There is decoration on the lines (more vertical than horizontal and with few forearm and tarsus in the form of rows of (often white- horizontal interconnections), with a few horizontal tipped) tubercles. interconnections. A dermal fold passes from the posterior Species within Badiohyla gen. nov. are separated from corner of the eye over the upper edge of the tympanum the other five genera (Nyctimystes, Nigreosoculus gen. and down to the insertion of the forelimb, sometimes nov., Occultatahyla gen. nov., Magnumoculus gen. nov. becoming obscure when leaving the tympanum. The and Albogibba gen. nov.) by having a unique “reticulated” outer fingers are approximately half-webbed, the third palpebral venation, half-webbed fingers and a size in and fifth toes webbed to the disc. A light-coloured dermal males with SVL to maximum of 50 mm to 100 mm ridge or series of tubercles extends from the elbow to the depending on the species. They are further defined as disc of the fourth finger. A similar but less distinct ridge or follows: The snout is relatively short, blunt and high (E-N/ line of tubercles occupies the outer edge of the tarsus IN 0.94), with distinct canthus rostralis and oblique loreal and fifth toe. A very small heel tubercle is present. The region. The vomerine teeth are in two patches between skin of the dorsum is smooth to very slightly granular, that the posterior edges of the choanae. of the venter is always coarsely granulate. Slit-like, vocal The palpebral venation forms a heavy reticulum. The sac openings are usually present in the floor of the tympanum is distinct and is separated from the eye by mouth. A small patch of very fine horny tubercles is found approximately the diameter of the tympanum. A fold of on the first finger of males. The iris is dark or black in skin passes from the posterior corner of the eye, over the colour. upper edge of the tympanum and down, becoming Exceptional to the preceding is the subgenus indistinct above the insertion of the forelimb. The skin of Planusrususpes subgen. nov. which conforms to the the dorsum is minutely roughened, that of the venter above genus diagnosis, save for the following coarsely granular. A row of tubercles is present along the differences: The heel is without a tubercle, or in rare outer edge of the forearm and there is a similar row on cases has a very small one. The skin is smooth above, the tarsus. There is only a slight suggestion of a heel granular below. A weak, wavy fold is present on the outer tubercle. The outer fingers are approximately half- surface of the forearm. The head is broader than long; webbed, the toes, except the fourth toe, are webbed to the canthus rostralis is not distinct; the loreal region is the disc. The body and head are light brown to purple oblique; Unique to this subgenus is that the palpebral dorsally, either with irregular dark brown blotches or venation is distinct, but is reduced to individual pigment markings absent. The legs have a similar ground color, spots and thin, meandering lines, although the lines are with or without irregular darker spots and bands present still mainly oriented largely in a vertical direction, with few on the tibia. horizontal interconnections. Species within Albogibba gen. nov. are separated from Species within Nigreosoculus gen. nov. are separated the other five genera (Nyctimystes, Nigreosoculus gen. from the other five genera (Nyctimystes, Occultatahyla nov., Occultatahyla gen. nov., Magnumoculus gen. nov. gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. and Badiohyla gen. nov.) by one or other of the following and Albogibba gen. nov.) by the following suite of unique suites of characters: characters: Vocal slits present; a very small heel tubercle; 1/ Being a large species (males to 100 mm. in snout to basal webbing on the hand; exposed tympanum; vertical vent length); of a uniform dorsal coloration (green in life, lines of palpebral reticulum oriented obliquely and with purple in alcohol-preserved specimens); palpebral relatively few horizontal cross-connections; pale-tan to venation is a reticulum without obvious orientation and in near black iris; rear of thighs barred/mottled with brown, the nominate subgenus of this genus, the male bears a caramel, or blue-gray. spine-like process on the anterior surface of the proximal Species within Magnumoculus gen. nov. are separated part of the humerus (upper arm) (nominate subgenus from the other five genera (Nyctimystes, Nigreosoculus Albogibba subgen. nov.), or: gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. 2/ As above, but male SVL of up to 80 mm; not including and Albogibba gen. nov.) by the following suite of a spine-like process on the anterior surface of the characters: The head is broad (HL/HW <1.0), the snout proximal part of the humerus (upper arm) in males and high and blunt, the canthus rostralis distinct, broadly with a unique dorsal pattern of black vermiculations on spaced nostrils (EN/IN ratio <1), and the loreal region the body and limbs, with unmarked greyish lower flanks nearly vertical. A strong, slightly curved, supratympanic (Ratiobrunneis subgen. nov.). fold begins at the posterior corner of the eye and According to Duellman et al. (2016), each of the six disappears above the insertion of the forelimb. The preceding genera had the following divergences from one tympanum is small but distinct. The hands have little another. webbing; the outer fingers are about one-quarter webbed. Nyctimystes Stejneger, 1916 diverged 14.8 MYA from its The vomerine teeth are in two patches between the nearest living relatives in other genera. internal nares, nearly on a line connecting the posterior The subgenus Magnummanibus subgen. nov. diverged margins of the nares. The skin is minutely granular 11.5 MYA from its nearest living relatives in the nominate above, coarser beneath. There is no tubercle on the heel. subgenus. A nearly straight, very slightly crenulated fold of skin is

Occultatahyla gen. nov. diverged 16.7 MYA from its gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. nearest living relatives in other genera. and Albogibba gen. nov.) by large size (adult females 50 Nigreosoculus gen. nov. diverged 14.8 MYA from its to 84 mm. in snout to vent length); internarial distance nearest living relatives in other genera. distinctly smaller than distance from eye to naris (E-N/IN Magnumoculus gen. nov. diverged 13 MYA from its averages 1.34 (range 1.2-1.5)); the head is broad, nearest living relatives, being Badiohyla gen. nov.. flattish, with depressed snout, closely spaced nares, and Badiohyla gen. nov. diverged 13 MYA from its nearest oblique loreal region. Palpebral venation is in oblique living relatives being Magnumoculus gen. nov.. lines (more vertical than horizontal and with few horizontal interconnections), with a few horizontal In turn both Magnumoculus gen. nov. and Badiohyla gen. interconnections. A dermal fold passes from the posterior nov. as a pair diverged from their nearest living relatives corner of the eye over the upper edge of the tympanum 22 MYA being Nyctimystes and other genera. and down to the insertion of the forelimb, sometimes Albogibba gen. nov. diverged 16.7 MYA from its nearest becoming obscure when leaving the tympanum. The living relatives in other genera. outer fingers are approximately half-webbed, the third The subgenus Ratiobrunneis subgen. nov. diverged 10 and fifth toes webbed to the disc. A light-coloured dermal MYA from its nearest living relatives in the nominate ridge or series of tubercles extends from the elbow to the subgenus. disc of the fourth finger. A similar but less distinct ridge or Distribution: Confined to the highlands of the central line of tubercles occupies the outer edge of the tarsus cordillera of New Guinea. and fifth toe. A very small heel tubercle is present. The Etymology: In Latin “Badio” means the colour beige and skin of the dorsum is smooth to very slightly granular, that “Hyla” is a tree frog. Many specimens in this tree frog of the venter is always coarsely granulate. Slit-like, vocal genus are in fact beige coloured, meaning that the genus sac openings are usually present in the floor of the name is entirely descriptive. mouth. A small patch of very fine horny tubercles is found Content: Badiohyla kubori (Zweifel, 1958) (type species); on the first finger of males. The iris is dark or black in B. gularis (Parker, 1936). colour. MAGNUMOCULUS GEN. NOV. Exceptional to the preceding is the subgenus LSIDurn:lsid:zoobank.org:act:7CC692B4-52AB-48BC- Planusrususpes subgen. nov. which conforms to the 84C2-4BF322A74060 above genus diagnosis, save for the following differences: The heel is without a tubercle, or in rare Type species: Nyctimystes narinosa Zweifel, 1958. cases has a very small one. The skin is smooth above, Diagnosis: Species of Nyctimystes, Occultatahyla gen. granular below. A weak, wavy fold is present on the outer nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., surface of the forearm. The head is broader than long; Badiohyla gen. nov. and Albogibba gen. nov. are the canthus rostralis is not distinct; the loreal region is separated from all other Australasian (Australian and oblique; Unique to this subgenus is that the palpebral New Guinea) Tree Frogs (Pelodryadidae) by the following venation is distinct, but is reduced to individual pigment suite of characters: spots and thin, meandering lines, although the lines are Large dark eyes, which have vertically elliptical pupils still mainly oriented largely in a vertical direction, with few and a lower eyelid with a characteristic reticulum or horizontal interconnections. palpebral venation of fine pigmented lines. There is Species within Nigreosoculus gen. nov. are separated usually a distinct, crenulated skin fold along the hind from the other five genera (Nyctimystes, Occultatahyla edge of the forearm and the foot, the heel often having a gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. small flap of skin. Males are usually considerably smaller and Albogibba gen. nov.) by the following suite of than females. characters: Vocal slits present; a very small heel tubercle; Nyctimystes Stejneger, 1916 as defined in this paper, are basal webbing on the hand; exposed tympanum; vertical separated from the other five genera (Occultatahyla gen. lines of palpebral reticulum oriented obliquely and with nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., relatively few horizontal cross-connections; pale-tan to Badiohyla gen. nov. and Albogibba gen. nov.) by having near black iris; rear of thighs barred/mottled with brown, sparse, broken vertically-oriented palpebral venation caramel, or blue-gray. across the nictitating membrane and in having adult Species within Magnumoculus gen. nov. are separated males without vocal slits. from the other five genera (Nyctimystes, Nigreosoculus Frogs in the subgenus Magnummanibus subgen. nov. are gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. separated from those in the nominate subgenus and Albogibba gen. nov.) by the following suite of Nyctimystes and the subgenus Asperohyla subgen. nov. characters: The head is broad (HL/HW <1.0), the snout by their possession of very long, prominent heel lappets. high and blunt, the canthus rostralis distinct, broadly Frogs in the subgenus Asperohyla subgen. nov. are spaced nostrils (EN/IN ratio <1), and the loreal region separated from the other two subgenera by having a nearly vertical. A strong, slightly curved, supratympanic completely hidden tympanum and a dorsal skin fold begins at the posterior corner of the eye and roughened by tiny, conical asperities rather than by disappears above the insertion of the forelimb. The variable sized and more rounded irregularities as seen in tympanum is small but distinct. The hands have little some other New Guinea tree frog species; iris is brown in webbing; the outer fingers are about one-quarter webbed. life. The vomerine teeth are in two patches between the Species within Occultatahyla gen. nov. are separated internal nares, nearly on a line connecting the posterior from the other five genera (Nyctimystes, Nigreosoculus margins of the nares. The skin is minutely granular Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 124 Australasian Journal of Herpetology above, coarser beneath. There is no tubercle on the heel. of characters: Colour in life in normal conditions is usually A nearly straight, very slightly crenulated fold of skin is a unform emerald green above, but occasionally may be present on the outer surface of the forearm. A moderate fawn, green, purple-green or blue. Hind side of thighs body size (40-60 mm S-V), predominantly brown more or less unform and without black and yellow colouration, an eyelid venation composed of a network marbling or spots. Hind edge of forearms are smooth or with numerous horizontal connections with relatively few with at most a few low discontinuous tubercles; hind edge vertical interconnections. There is decoration on the of foot is smooth. Fingers with conspicuous webbing that forearm and tarsus in the form of rows of (often white- reaches at least as far as the punultimate phalanx of the tipped) tubercles. fourth finger; Vomerine teeth present. Distribution: New Guinea, including offshore islands on Frogs in the the nominate subgenus Shireenhoserhylea the continental shelf. gen. nov. are readily separated from those in the Etymology: In Latin “Magnumoculus” means big eyes, in subgenus Emeraldhyla subgen. nov. by having an iris, or reflection of the big eyes in all species. iris immediately above and below the pupil that is a Content: Magnumoculus narinosus (Zweifel, 1958) (type brilliant red or bright orange in colour and whitish species); N. ocreptus (Menzies, 2014); N. myolae underneath, versus an iris that is yellow or dull orange in (Menzies, 2014); N. cryptochrysos (Kraus, 2012). colour and strongly yellow to orange underneath in PELODRYANINI TRIBE NOV. Emeraldhyla subgen. nov.. PELODRYAS GÜNTHER, 1858. The taxon S. luteiventris (Ogilby, 1907), originally described as “Hyla luteiventris” with a type locality of Type species: Rana caerulea White, 1790. Brisbane, Queensland, is herein resurrected from the Diagnosis: Species in the genus Pelodryas Günther, synonymy of S. gracilenta (Peters, 1869) with a type 1858 are readily separated from all other Australasian locality of Mackay, North Queensland. Tree Frogs (Pelodryadidae) by the following unique suite Differences between the two nominate taxa are explained of characters: Colour in life is usually a near unform in Anstis (2013) on pages 209-211, where she treats emerald-green in life, but varies from dark purplish-green them as a single species but also notes “appears to to fawn or at times even blue, often with scattered white comprise more than one species awaiting further or yellow spots or dots; there is no obvious white stripe description”. running along the edge of the lower jaw; the hind side of thighs lacks black or yellow marbling, either being plain Hence S. luteiventris (Ogilby, 1907) appears in the list of or sometimes pinkish or orangeish; hind edge of forearm species within this genus. is smooth, or with at most a few, low, discontinuous The species S. megaviridis sp. nov. from Eungella, near tubercles; the skin on top is smooth or slightly leathery; Mackay in north Queensland, formally described herein, venter is white and coarsely granular; hind edge of foot is has until now been treated as a northern population of smooth; fingers have conspicuous webbing, but reaching “Hyla chloris Boulenger, 1893” from the Richmond River, no further than the base of the penultimate phalanx of the in New South Wales. fourth finger; toes are about 2/3 webbed; finger and toe However in many respects this taxon is intermediate discs are large and obvious; vomerine teeth present and between the two forms now known as S. chloris prominent, between and behind the choanae; there is (Boulenger, 1893) and S. xanthomera (Davies, obvious thick, supratympanic gland and the tympanum is McDonald, and Adams, 1986), which have a 4 MYA large and obvious. divergence from one another according to Duellman et al. Duellman et al. (2016) found that the four named and (2016). currently recognized species within this genus diverged Because the Eungella population is also reproductively from their nearest living relatives, 24.7 MYA, these being isolated from both other populations, as well as being species within the genus Shireenhoserhylea gen. nov.. clearly divergent morphologically, it is herein formally Distribution: Most parts of continental Australia, except named as a new species, S. megaviridis sp. nov.. the coldest regions as well as parts of New Guinea. The most closely related species to this genus Because the most common species P. caerulea (White, (Shireenhoserhylea gen. nov.) are the morphologically 1790) is often commensal of human habitation and a divergent species within the genus Pelodryas Günther, popular pet, the distribution has expanded in Australia 1858, type species being Rana caerulea White, 1790. and New Guinea in modern times. According to Duellman et al. (2016) the two species Content: Pelodryas caerulea (White, 1790) (type groups diverged 24.7 MYA, making genus level division a species); P. cavernicola (Tyler and Davies, 1979); P. common-sense decision. splendida (Tyler, Davies and Martin, 1977); P. gilleni The Litoria kumae Menzies and Tyler, 2004 species (Spencer, 1896). group, being the New Guinea members of the genus SHIREENHOSERHYLINA SUBTRIBE NOV. Shireenhoserhylea gen. nov., as well as S. bella SHIREENHOSERHYLEA GEN. NOV. (McDonald, Rowley, Richards and Frankham, 2016) from Cape York, Australia had according to Duellman et al. LSIDurn:lsid:zoobank.org:act:A50DB4CA-5049-4818- (2016) diverged from the Australian Hyla chloris 9BF2-98ECA036D81C Boulenger, 1893 group 16.6 MYA. Type species: Hyla chloris Boulenger, 1893. Therefore the mainly New Guinea group is herein placed Diagnosis: Frogs in the genus Shireenhoserhylea gen. in a new subgenus Emeraldhyla subgen. nov.. nov. are readily separated from all other Australasian Distribution: Wetter parts of the east coast of Australia, Tree Frogs (Pelodryadidae) by the following unique suite excluding colder parts of the far south, New Guinea and Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 125

offshore islands. The nominate subgenus Shireenhoserhylea subgen. nov. is restricted to the East Shireenhoserhylea chloris Coast of Australia, south of the northern wet tropics in Queensland. The subgenus Emeraldhyla subgen. nov. occurs in New Guinea and offshore islands as well as the far north of Cape York, Queensland. Etymology: Named in honour of Shireen Hoser, of Park Orchards, Victoria, Australia, who also happens to be my wife, in recognition of decades of work supporting wildlife conservation globally, including successfully defending herpetology against taxonomic vandalism by members of the Wolfgang Wüster gang of thieves in the 1990’s. This was done via a successful submission to the ICZN in 1998. That submission, (Hoser 1998), successfully stopped Robert Sprackland illegally renaming the species Ourimbah, NSW originally described as Odatria keithhornei Wells and Wellington, 1985 the obscenely coined name “Varanus teriae” being that of Sprackland’s wife. The ICZN correctly struck out the illegally coined Shireenhoserhylea gen. nov., as well as S. bella patronym of Sprackland, this being the second ruling by (McDonald, Rowley, Richards and Frankham, 2016) from the ICZN in favour of the Wells and Wellington papers, Cape York, Australia had according to Duellman et al. which was not so much a defence of Wells and (2016) diverged from the Australian Hyla chloris Wellington (1985), but rather a defence of the Boulenger, 1893 group 16.6 MYA. Therefore the mainly International Code of Zoological Nomenclature and the New Guinea group is herein placed in this new subgenus rules that are supposed to bind all scientists. Emeraldhyla subgen. nov.. Content: Shireenhoserhylea chloris (Boulenger, 1893) Distribution: Species within Emeraldhyla subgen. nov. (type species); S. aruensis (Horst, 1883); S. auae are found in New Guinea, including offshore islands as (Menzies and Tyler, 2004); S. bella (McDonald, Rowley, well as the northern parts of Cape York, Australia. Richards and Frankham, 2016); S. callista (Kraus, 2013); Etymology: Emeraldhyla literally means Emerald green S. elkeae (Günther and Richards, 2000); S. eschata tree frog, which is exactly what these species are! (Kraus and Allison, 2009); S. gracilenta (Peters, 1869); S. kumae (Menzies and Tyler, 2004); S. luteiventris (Ogilby, Content: Shireenhoserhylea (Emeraldhyla) kumae 1907); S. megaviridis sp. nov.; S. robinsonae (Oliver, (Menzies and Tyler, 2004); (type species); S. Stuart-Fox and Richards 2008); S. xanthomera (Davies, (Emeraldhyla) aruensis (Horst, 1883); S. (Emeraldhyla) McDonald, and Adams, 1986). auae (Menzies and Tyler, 2004); S. (Emeraldhyla) bella (McDonald, Rowley, Richards and Frankham, 2016); S. EMERALDHYLA SUBGEN. NOV. (Emeraldhyla) callista (Kraus, 2013); S. (Emeraldhyla) LSIDurn:lsid:zoobank.org:act:F0C25610-5D1F-40E6- elkeae (Günther and Richards, 2000); S. (Emeraldhyla) B0CC-E343EC59991E eschata (Kraus and Allison, 2009); S. (Emeraldhyla) Type species: Litoria kumae Menzies and Tyler, 2004. robinsonae (Oliver, Stuart-Fox and Richards 2008). Diagnosis: Frogs in the genus Shireenhoserhylea gen. SHIREENHOSERHYLEA (SHIREENHOSERHYLEA) nov. are readily separated from all other Australasian MEGAVIRIDIS SP. NOV. Tree Frogs (Pelodryadidae) by the following unique suite LSIDurn:lsid:zoobank.org:act:3541C57B-6D88-4D1A- of characters: AC3E-4E95ABF9E6AD Colour in life in normal conditions is usually a unform Holotype: A preserved specimen at the Queensland emerald green above, but occasionally may be fawn, Museum, Brisbane, Queensland, Australia, specimen green, purple-green or blue. Hind side of thighs more or number J35941, collected from Broken River, Eungella less unform and without black and yellow marbling or National Park, (near Mackay), Queensland, Australia, spots. Hind edge of forearms are smooth or with at most Latitude -21.175 S., Longitude 148.5083 E. a few low discontinuous tubercles; hind edge of foot is This government-owned facility allows access to its smooth. Fingers with conspicuous webbing that reaches holdings. at least as far as the punultimate phalanx of the fourth finger; Vomerine teeth present. Paratypes: Ten preserved specimens at the Queensland Museum, Brisbane, Queensland, Australia, specimen Frogs in the the nominate subgenus Shireenhoserhylea numbers J35906, J35912, J35965, J35970, J35979, gen. nov. are readily separated from those in the J35980, J36002, J36007, J36016 and J36025 all subgenus Emeraldhyla subgen. nov. by having an iris, or collected from within 10 km of the type locality above. iris immediately above and below the pupil that is a brilliant red or bright orange in colour and whitish Diagnosis: Shireenhoserhylea megaviridis sp. nov. has underneath, versus an iris that is yellow or dull orange in until now been treated as a population of S. chloris colour and strongly yellow to orange underneath in (Boulenger, 1893) known in most texts as “Litoria chloris”, Emeraldhyla subgen. nov.. including in Anstis (2013) and Cogger (2014). A more northerly (wet tropics in Queensland) population The Litoria kumae Menzies and Tyler, 2004 species previously treated as S. chloris was formally named in group, being the New Guinea members of the genus

1986 and is herein known as S. xanthomera (Davies, colour and strongly yellow to orange underneath in McDonald, and Adams, 1986). Emeraldhyla subgen. nov.. According to Duellman et al. (2016), the two preceding Frogs in the genus Shireenhoserhylea gen. nov. are species diverged from one another 4 MYA. readily separated from all other Australasian Tree Frogs In many respects S. megaviridis sp. nov. is intermediate (Pelodryadidae) by the following unique suite of between the two forms, but due to divergence between characters: Colour in life in normal conditions is usually a each, as well as geographic disjunction, it is appropriate unform emerald green above, but occasionally may be that this population be treated as a full species as fawn, green, purple-green or blue. Hind side of thighs opposed to subspecies of one or other. more or less unform and without black and yellow S. megaviridis sp. nov. would key as S. chloris (as “Litoria marbling or spots. Hind edge of forearms are smooth or chloris”) in Anstis (2013) or Cogger (2014). with at most a few low discontinuous tubercles; hind edge The morphologically similar and closely related S. of foot is smooth. Fingers with conspicuous webbing that xanthomera is separated from both S. megaviridis sp. reaches at least as far as the punultimate phalanx of the nov. and S. chloris by having bright orange-yellow sides fourth finger; Vomerine teeth present. and upper arms, hands and feet as well as inner surfaces Photos of L. chloris in life can be found in Hoser (1989) of thighs being bright orange, with whitish or yellow on page 36, top two images, Vanderduys (2012) at page undersides. 30, both images, Cogger (2014) on p. 156 at bottom right This is opposed to light yellow sides and upper arms, and Anstis (2013) at page 168 both images. hands and feet as well as inner surfaces of thighs being Photos of S. megaviridis sp. nov. in life can be found in dark purplish-red or brown often with a purple surface in Anstis (2013) at page 170 (metamorphosed young), two S. chloris. top left images, and adults in life can be seen online at: S. megaviridis sp. nov. is separated from both S. https://lyrebird-gallery.smugmug.com/Frogs/Litoria- xanthomera and S. chloris by having brilliant blue on the chloris/i-zHZKnxq inner surfaces of the thighs. and: The tadpoles of S. xanthomera are separated from both https://www.flickr.com/photos/huntermccall/31855340567/ S. megaviridis sp. nov. and S. chloris by having a and: narrowly arched jaw sheath, versus not so narrow in the https://www.flickr.com/photos/jaricornelis/38736419370/ other two species. and: S. megaviridis sp. nov. is separated from S. chloris by the https://www.flickr.com/photos/reptileshots/24091549126/ expansion of flash colour under the lower left leg to and: occupy the entire lower surface, versus not so in S. https://www.flickr.com/photos/reptileshots/31194264913/ chloris. and: The upper arm of S. megaviridis sp. nov. is usually yellow, versus usually greenish-yellow in S. chloris. https://www.flickr.com/photos/144043627@N08/ In dorsal colouration, pre-metamorphosing tadpoles of S. 40530915481/ megaviridis sp. nov. are like those of S. xanthomera and and: not S. chloris. In S. chloris the anterior of the tadpole is https://www.flickr.com/photos/piazzi1969/5772240176/ greyish with peppering and any markings are indistinct. Photos of S. xanthomera in life can be found in There are no obvious spots or blotches, with two spots on Vanderduys (2012) on page 78 in both images, Cogger the snout anterior to the eyes being indistinct. Only the (2014) on page 196 at top and Anstis (2013) on page 339 anterior fringe is pale. in all photos. By contrast the relevant tadpoles of both S. xanthomera Distribution: Known with certainty only from the vicinity and S. megaviridis sp. nov. have a generally pale anterior of Eungella National Park in north-east Queensland and end and snout, with two distinct large greyish brown dots generally south of the area known as the Burdekin Gap, on the snout anterior to the eyes. Anteriorly, the relevant north Queensland and north of the St. Lawrence Gap, S. xanthomera are whitish, versus yellowish-brown in S. east Queensland, both being major lowland megaviridis sp. nov.. biogeographical barriers in the Queensland region. The other two species within the subgenus However based on specimens in State Museums, the Shireenhoserhylea gen. nov., namely S. gracilenta potential limits of this taxon are likely to be Caithu State (Peters, 1869) and S. luteiventris (Ogilby, 1907) are both Forest, Latitude -19.58 S., Longitude 147.43 E. in the separated from the three preceding species by the north and Bruce Highway at the junction of St. Lawrence, following characters: The green colour of the forearms Latitude -22.3667 S., Longitude 149.45 E in the south stops abruptly with a sharp edge at the elbow. The skin and of course confined to the east by the Pacific Ocean covering the tympanum is finely granular. The rim of the and in the west of the coastal zone by drier lowland lower jaw, especially near the tip of the snout is narrowly habitats. edged with white. Etymology: The word “mega” usually means lots and Frogs in the the nominate subgenus Shireenhoserhylea “viridis” means green and as the relevant species is very gen. nov. are readily separated from those in the green, the species name S. megaviridis sp. nov. is wholly subgenus Emeraldhyla subgen. nov. by having an iris, or appropriate. iris immediately above and below the pupil that is a brilliant red or bright orange in colour and whitish underneath, versus an iris that is yellow or dull orange in

SUMMAVIRIDIS GEN. NOV. Adult females are about 37 mm in body length (modified LSIDurn:lsid:zoobank.org:act:6187EF08-0B88-40F9- from Tyler, 1978). 837F-C0C06F50525F Specimens of the morphologically similar Llewellynura Type species: Hyla (Litoria) vagabunda Peters and Wells and Wellington, 1985 are readily separated by their Doria, 1878. much smaller adult size of less than 25 mm body length. Diagnosis: The genus Summaviridis gen. nov. is The species within the morphologically similar genus monotypic for a West Papuan species, S. vagabunda that Rotundaura gen. nov. are separated from this genus does not appear to have close affinities to any other (Summaviridis gen. nov.) by the tympanum being fully species or genus. Tyler (1968) placed it in a group on its exposed and round, versus the upper surface being cut own, although Menzies (2006) associated it with his so- at the rear by a skin fold forming a straight line and a called “Litoria gracielenta complex”. However S. significantly blunter snout in Rotundaura gen. nov.. vagabunda has unwebbed hands, versus heavily webbed Distribution: The type species for Summaviridis gen. in the other species, which in effect scuttles any close nov. is only known from the two types, a male and a association at the genus level. female, being from Vogelkop Peninsula, Papua (New Summaviridis gen. nov. are readily separated from all Guinea) (female) and Seram (= Ceram) Island, Molucca other Australasian (Australian and New Guinea) Tree Islands, Indonesia. Frogs (Pelodryadidae) by the following unique suite of Etymology: The name Summaviridis in Latin means characters: A broad head, short limbs, unwebbed fingers, intense or deep green in reflection of the colour of the a prominent fold of skin across the chest and (in life) a dorsal surface of these frogs in life. dark green dorsal colouration. Content: Summaviridis vagabunda (Peters and Doria, In further detail the Summaviridis gen. nov. is separated 1878) (monotypic). from all other Australasian (Australian and New Guinea) PUSTULATARANINI TRIBE NOV. Tree Frogs (Pelodryadidae) as follows: The head is PUSTULATARANA GEN. NOV. broader than long (HL/HW 0.951), its length slightly less LSIDurn:lsid:zoobank.org:act:E2F68230-6BBB-4518- than one-third of the snout to vent length (HL/S-V 0.310). A9C2-C8432FE558DE The snout is rounded when viewed from above and in Type species: Litoria longirostris Tyler and Davies, 1977. profile. The nostrils are more lateral than superior, their distance from the end of the snout slightly less than that Diagnosis: The type species in this genus Litoria from the eye. The distance between the eye and the naris longirostris Tyler and Davies, 1977 is phylogenetically is greater than the internarial span (E-N/IN 1.154). The and morphologically closest to species within canthus rostralis is straight and poorly defined. The eye Llewellynura Wells and Wellington, 1985. However is large, its diameter greater than the distance separating Duellman et al. (2016) found a divergence of 18 MYA it from the nostril. Three-quarters of the tympanum is between this and other living species within Llewellynura. visible, the superior rim of the tympanic annulus is hidden Duellman et al. (2016) also found a divergence of 20.9 beneath the supra-tympanic fold. The tympanal diameter MYA between “Litoria longirostris Tyler and Davies, 1977” is equivalent to two-thirds of the eye diameter. The and species within the genus Mahoneybatrachus Wells vomerine teeth are in two broadly oval series situated and Wellington, 1985. between the choanae. The tongue is broadly cordiform With such deep divergences between the three groups with a deeply indented posterior border. The fingers are and obvious morphological divergence, the most sensible very long and slender with narrow lateral fringes; in classification system (taxonomy) calls for them to be decreasing order of length 3>4>2>1; unwebbed. The placed in three separate groups, or genera. Hence the terminal discs and sub-articular tubercles are large and erection of the genus Pustulatarana gen. nov. herein. prominent. The hind limbs are short and slender with a Because this genus is monotypic for “Litoria longirostris TL/S-V ratio of 0.513. Toes in decreasing order of length Tyler and Davies, 1977”, the diagnosis for the species is 4>5>3 >2>1. The webbing between the outer and fourth the same as for the genus. The keys in Cogger (2014) toe reaches half-way up the penultimate phalanx on the and Anstis (2013) for this species taxon, in effect also fifth toe, and the sub-articular tubercle at the base of the diagnose this genus. penultimate phalanx on the fourth. The skin on the dorsal The single known living species within the genus surfaces is smooth with a few minute flattened tubercles. Pustulatarana gen. nov. is readily separated from all The throat and chest are slightly tubercular; the other Australasian Tree Frogs (Pelodryadidae) by the abdomen, sides of the body and lower surface of the following suite of characters: A smallish sized tree frog thighs are extremely granular. There is a conspicuous being 27 mm in body length. Brown to greenish brown or curved supra-tympanic fold, and a prominent skin fold even yellow above, with obscure and irregular darker across the chest. The dorsal surfaces are in life dark flecks and blotches. Creamy-white below, finely peppered green and in preservative dark blue. There is a faint trace with dark blackish-brown or shading of other lighter of a narrow white line above a broader brown line on the colour on the throat, with heavy concentrations of canthus rostralis, upper eyelid and supra-tympanic fold. stippling on chin, chest and flanks of some specimens, The backs of the thighs are pale brown and marked with and ...... Continued in AJH Issue 46... small irregularly-shaped, pale yellow spots. The backs of Hoser, R. T. 2020. For the first time ever! An overdue the tarsus and forearm are pale brown. The ventral review and reclassification of Australasian Tree Frogs surfaces are pale yellow with leaden blue patches at the (Amphibia: Anura: Pelodryadidae), including formal descriptions of 12 tribes, 11 subtribes, 34 genera, 26 angles of the jaws, and small faint brown spots on the subgenera, 62 species and 12 subspecies new to science. remainder of the mandibular border and on the throat. Australasian Journal of Herpetology 44-46:1-192. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved AustralasianAustralasian JournalJournal ofof HerpetologyHerpetology

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

Hoser, R. T. 2020. For the first time ever! An overdue review and reclassification of Australasian Tree Frogs (Amphibia: Anura: Pelodryadidae), including formal descriptions of 12 tribes, 11 subtribes, 34 genera, 26 subgenera, 62 species and 12 subspecies new to science. Australasian Journal of Herpetology 44-46:1-192. ISSUE 46, PUBLISHED 5 JUNE 2020 Hoser, R. T. 2020. For the first time ever! An overdue review and reclassification of Australasian Tree Frogs (Amphibia: Anura: Pelodryadidae), including formal descriptions of 12 tribes, 11 subtribes, 34 genera, 26 130 Australasiansubgenera, 62 species Journal and 12 subspecies of Herpetologynew to science. Australasian Journal of Herpetology 44-46:1-192. ... Continued from AJH Issue 45 ... zone of apparently unsuitable habitat of significant geological antiquity and are therefore reproductively Underside of thighs have irregular darker patches and isolated and therefore evolving in separate directions. hind isde of thigh has irregular fine creamish coloured They are also morphologically divergent, warranting stripes. Skin is leathery and with numerous scattered identification of the unnamed population at least to tubercles which may or not be arranged in well-defined subspecies level as done herein. longitudinal rows, including sometimes some of medium to large size and a prominent one on the eyelid. Belly is The zone dividing known populations of each species is smooth except for some granular skin on the lower belly only about 30 km in a straight line. and thighs. Vomerine teeth present, but weakly P. longirostris tozerensis subsp. nov. is separated from P. developed and between the choanae. Fingers lack longirostris longirostris by having a dorsum covered with webbing but have large oval discs. Toes poorly to numerous small pointed tubercles and some small folds moderately webbed. A minute rounded outer metatarsal of skin not obviously arranged in a longitudinal manner, tubercle and a moderate-zied oval inner metatarsal versus small pointed and larger tubercles including large tubercle. Tympanum distinct. rounded ones on the dorsum, some of which are clearly Pustulatarana gen. nov. are separated from the genus arranged in a longitudinal manner down the dorsum. Llewellynura Wells and Wellington, 1985 by the large White on the upper lip, under the eye of P. longirostris oval discs on the forelimbs and larger body size (27 mm tozerensis subsp. nov. extends beyond the eye towards vs 20 mm). Pustulatarana gen. nov. are separated from the snout, but does not do so in P. longirostris the genus Mahoneybatrachus Wells and Wellington, longirostris. 1985, by having reduced toe webbing, versus well The single known living species within the genus developed webbing on the feet. Pustulatarana gen. nov., P. longirostris is readily Living specimens of Pustulatarana longirostris (Tyler and separated from all other Australasian Tree Frogs Davies, 1977) are depicted in Vanderduys (2012) on (Pelodryadidae) by the following suite of characters: A page 49 at bottom (2 images), as well as in Anstis (2013) smallish sized tree frog being 27 mm in body length. on pages 234-237, including tadpoles and in Cogger Brown to greenish brown or even yellow above, with (2014) on page 172 at top. obscure and irregular darker flecks and blotches. Creamy-white below, finely peppered with dark blackish- Distribution: Known only from McIllwraith Range, Mount brown or shading of other lighter colour on the throat, Tozer and William Thomson Ranges on Cape York with heavy concentrations of stippling on chin, chest and Peninsula in Far North Queensland, Australia. flanks of some specimens. Underside of thighs have Etymology: In Latin Pustulata means warty and rana irregular darker patches and hind isde of thigh has means frog, so the name literally means warty frog, which irregular fine creamish coloured stripes. Skin is leathery sums up the morphology of the species in the genus. and with numerous scattered tubercles which may or not Content: Pustulatarana longirostris (Tyler and Davies, be arranged in well-defined longitudinal rows, including 1977). sometimes some of medium to large size and a PUSTULATARANA LONGIROSTRIS TOZERENSIS prominent one on the eyelid. Belly is smooth except for SUBSP. NOV. some granular skin on the lower belly and thighs. LSIDurn:lsid:zoobank.org:act:6FA68D51-7733-43A7- Vomerine teeth present, but weakly developed and A381-129BE1723EAD between the choanae. Fingers lack webbing but have Holotype: A preserved specimen at the Queensland large oval discs. Toes poorly to moderately webbed. A Museum, Brisbane, Queensland, Australia, specimen minute rounded outer metatarsal tubercle and a number J92480 collected from the south side of Mount moderate-zied oval inner metatarsal tubercle. Tympanum Tozer, far north Queensland, Australia, Latitude -13.1 S., distinct. Longitude 143.2 E. Pustulatarana gen. nov. are separated from the genus This facility gives access to its holdings. Llewellynura Wells and Wellington, 1985 by the large Paratypes: Five preserved specimens at the Queensland oval discs on the forelimbs and larger body size (27 mm Museum, Brisbane, Queensland, Australia, specimen vs 20 mm). numbers J92481-J92485 all collected from the south side Pustulatarana gen. nov. are separated from the genus of Mount Tozer, far north Queensland, Australia, Latitude Mahoneybatrachus Wells and Wellington, 1985, by -13.1 S., Longitude 143.2 E. having reduced toe webbing, versus well developed Diagnosis: Pustulata longirostris tozerensis subsp. nov. webbing on the feet. has until now been treated as a northern outlier Distribution: P. longirostris tozerensis subsp. nov. is population of nominate P. longirostris (Tyler and Davies, found in the vicinity of rainforest streams running from 1977), with a type locality of Rocky River (Latitude 13.46 just south of Mount Tozer to include the Sir William S., Longitude 142.23 E), McIlwraith Range, Cape York Thompson Range, north of Lockhart River, far north Peninsula, Queensland, Australia and found generally in Queensland, Australia. the rainforested tributaries of the Rocky River system. Etymology: The subspecies name reflects the locality By contrast P. longirostris tozerensis subsp. nov. is found that most specimens of this taxon have been found to in the rainforest streams running from the south side of date, namely the environs of Mount Tozer. Mount Tozer and including the Sir William Thompson LLEWELLYNURA WELLS AND WELLINGTON, 1985 Range, north of Lockhart River, far north Queensland, Type species: Hyla dorsalis microbelos Cogger, 1966. Australia. The two populations are separated by a dry Diagnosis: The type species Hyla dorsalis microbelos

Cogger, 1966 is now recognized as a full species MICROLITORIA SUBGEN. NOV. separate to the morphologically similar New Guinea LSIDurn:lsid:zoobank.org:act:6DA3701D-5A02-4C6C- taxon originally described as Litoria dorsalis Macleay, B19A-8670E0ECD0BA 1878. The putative taxon Llewellynura microbelos Type species: Litoria dorsalis Macleay, 1878. (Cogger, 1966), with a type locality of Cairns in Diagnosis: Species within the genus Llewellynura Wells Queensland, Australia has long been recognized as a so- and Wellington, 1985 are readily separated from all other called composite species (e.g. Anstis 2013). Two new Australasian Tree Frogs (Pelodryadidae) by the following forms previously included in L. microbelos are formally suite of characters: they are tiny in size, being about 20 named within this paper. mm in body length as adults. Colour is variegated dark Species within the genus Llewellynura Wells and and brown dorsally; a row of widely spaced dorsolateral Wellington, 1985 are readily separated from all other tubercles and ridges may be on each side, or Australasian Tree Frogs (Pelodryadidae) by the following alternatively just a few randomly scattered small suite of characters: they are tiny in size, being about 20 tubercles on either side of upper flank; no pectoral fold; mm in body length as adults. Colour is variegated dark several conspicuous tubercles above each eye; tiny, and brown dorsally; a row of widely spaced dorsolateral slender and agile and with a sharply pointed snout. The tubercles and ridges may be on each side, or dorsum may be grey, brown, yellow or red, often flecked alternatively just a few randomly scattered small or mottled with darker colour. There is usually a broad tubercles on either side of upper flank; no pectoral fold; dark band from behind the eye to the groin and an even several conspicuous tubercles above each eye; tiny, darker stripe along the snout, through the eye and over slender and agile and with a sharply pointed snout. The the base of the arm to the flank. Snout and upper dorsum may be grey, brown, yellow or red, often flecked surfaces of the limbs are peppered with dark brown. or mottled with darker colour. There is usually a broad There is a conspicuous dark bar along the front edge of dark band from behind the eye to the groin and an even the arm. darker stripe along the snout, through the eye and over Ventral surface is white except for a fine peppering of the base of the arm to the flank. Snout and upper brown on the throat, chest and limbs. Skin is smooth surfaces of the limbs are peppered with dark brown. dorsally, with at least some small tubercles or warts or There is a conspicuous dark bar along the front edge of skin folds and usually several small subercles over each the arm. Ventral surface is white except for a fine eye. Throat skin is smooth, but belly is granular. Finger peppering of brown on the throat, chest and limbs. Skin is and toe discs are moderate in size but distinct. Fingers smooth dorsally, with at least some small tubercles or lack webbing and toes are less than half webbed. The warts or skin folds and usually several small subercles disc and nearly two phlanges are free on the outer side of over each eye. Throat skin is smooth, but belly is the fourth toe. The inner metatarsal tubercle is prominent granular. Finger and toe discs are moderate in size but and there is no outer one. Tympanum ranges from small distinct. Fingers lack webbing and toes are less than half to indistinct. Second finger longer than first. webbed. The disc and nearly two phlanges are free on The Australian species within Llewellynura being of the the outer side of the fourth toe. The inner metatarsal nominate subgenus Llewellynura are separated from the tubercle is prominent and there is no outer one. New Guinea species herein placed in the subgenus Tympanum ranges from small to indistinct. Second finger Microlitoria subgen. nov. by lacking vomerine teeth. longer than first. These are present in species of Microlitoria subgen. nov.. The Australian species within Llewellynura being of the According to Duellman et al. (2016), the two subgenera nominate subgenus Llewellynura are separated from the diverged from one another 14.8 MYA. New Guinea species herein placed in the subgenus Microlitoria subgen. nov. by lacking vomerine teeth. Distribution: Species within the subgenus Microlitoria subgen. nov. are known from various locations in New Distribution: The three species within the nominate Guinea. subgenus Llewellynura are found in the tropical parts of northern Australia from three main areas being: The three species within the nominate subgenus Llewellynura are found in the tropical parts of northern 1/ The north-west Kimberley Division of Western Australia from three main areas being: Australia, centred around the Mitchell Plateau (L. fukker sp. nov.), and: 1/ The north-west Kimberley Division of Western Australia, centred around the Mitchell Plateau (L. fukker 2/ The top end of the Northern Territory, from Litchfield sp. nov.), and: National Park in the west and extending east to include Groote Eylandt, and also including the Tiwi Islands (L. 2/ The top end of the Northern Territory, from Litchfield yehbwudda sp. nov.), and: National Park in the west and extending east to include Groote Eylandt, and also including the Tiwi Islands (L. 3/ Cape York Queensland, extending as far south as yehbwudda sp. nov.), and: Townsville (L. microbelos). Records south of there are almost certainly specimens inadvertently moved in recent 3/ Cape York Queensland, extending as far south as times by humans. Townsville (L. microbelos). Records south of there are almost certainly specimens inadvertently moved in recent Species within the subgenus Microlitoria subgen. nov. are times by humans. known from various locations in New Guinea. Etymology: The species in the subgenus have until now Content: Llewellynura microbelos (Cogger, 1966) (type been treated by most publishing authors as being within species); L. dorsalis (Macleay, 1878); L. fukker sp. nov.; the genus “Litoria Tschudi, 1838”. Due to them being L. jeudii (Werner, 1901); L. timida (Tyler and Parker,

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. 1972); L. yehbwudda sp. nov.. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 132 Australasian Journal of Herpetology among the tinyest species in the putative genus, they are nostril and is well defined at this point. herein called “Micro” as in small and hence in full L. yehbwudda sp. nov. has a yellow-grey iris (rarely seen “Microlitoria”. with a slight orange tinge) and a prominent well-defined Content: Llewellynura (Microlitoria) dorsalis (Macleay, row of skin folds and tubercles running in a line on either 1878); L. (Microlitoria) jeudii (Werner, 1901); L. side of the back of the dorsum and a second similar well- (Microlitoria) timida (Tyler and Parker, 1972). defined row of smaller tubercles running along each side LLEWELLYNURA (LLEWELLYNURA) FUKKER SP. of the mid flank. The white labial marking under the eye NOV. does not extend as far foreward as the nostril. LSIDurn:lsid:zoobank.org:act:DBE25DB2-10D7-4380- Anstis (2013) on page 247, gives detail of differences A5CF-40F842506D43 between the tadpoles and metamorps of Queensland L. Holotype: A preserved specimen at the Western microbelos, and Northern Territory L. yehbwudda sp. nov. Australian Museum, Perth, Western Australia, Australia, (which are treated by her provisionally as a separate specimen number R56470, collected 20 km south east of population of L. microbelos). Mitchell Plateau Camp, Latitude -14.95 S., Longitude Significantly the images of two metamorphs (one of each 125.95 E. species) on page 247 bottom of Anstis (2013) readily This government-owned facility allows access to its shows the diagnostic differences also in the adults of holdings. both species as outlined herein. Paratypes: Thirteen preserved specimens at the Photos of L. microbelos in life can be seen in Vanderduys Western Australian Museum, Perth, Western Australia, (2012) on page 54 in two images, Cogger (2014) on page Australia, specimen numbers R43337, R43361- R43370, 174 bottom left and Anstis (2013) on page 245 at top left R56470 and R62203 all collected from Mitchell Plateau, and second from top on right. Western Australia, Australia, Latitude -14.8667 S., Photos of fukker sp. nov. in life can be seen in Tyler and Longitude125.8333 E. Davies (1986) in the colour plates at number 27 and in Diagnosis: The three species within the nominate Tyler et al. (1994) on plate 32 at bottom. subgenus Llewellynura Wells and Wellington, 1985 have Photos of L. yehbwudda sp. nov. in life can be seen in until 2019 been treated as a single species, namely L. Anstis (2013) on page 245 at top right. microbelos (Cogger, 1966). However this putative taxon Species within the genus Llewellynura Wells and in fact consists of three separate and widely allopatric Wellington, 1985 are readily separated from all other species. Australasian Tree Frogs (Pelodryadidae) by the following This has been known for many years and Anstis (2013) suite of characters: they are tiny in size, being about 20 stated a view that more than one species was within mm in body length as adults. Colour is variegated dark putative L. microbelos as described in her book. and brown dorsally; a row of widely spaced dorsolateral As a trio, they are found in the tropical parts of northern tubercles and ridges may be on each side, or Australia from three main areas being: alternatively just a few randomly scattered small 1/ The north-west Kimberley Division of Western tubercles on either side of upper flank; no pectoral fold; Australia, centred around the Mitchell Plateau (L. fukker several conspicuous tubercles above each eye; tiny, sp. nov.), and: slender and agile and with a sharply pointed snout. The dorsum may be grey, brown, yellow or red, often flecked 2/ The top end of the Northern Territory, from Litchfield or mottled with darker colour. There is usually a broad National Park in the west and extending east to include dark band from behind the eye to the groin and an even Groote Eylandt and also including the Tiwi Islands (L. darker stripe along the snout, through the eye and over yehbwudda sp. nov.), and: the base of the arm to the flank. Snout and upper 3/ Cape York Queensland, extending as far south as surfaces of the limbs are peppered with dark brown. Townsville (L. microbelos). Records south of there are There is a conspicuous dark bar along the front edge of almost certainly specimens inadvertently moved in recent the arm. Ventral surface is white except for a fine times by humans, such as when being translocated in peppering of brown on the throat, chest and limbs. Skin is boxes of fruit via trucks and trains. smooth dorsally, with at least some small tubercles or All three species would key out as L. microbelos in either warts or skin folds and usually several small subercles Anstis (2013) or Cogger (2014). over each eye. Throat skin is smooth, but belly is The three species can however be readily separated from granular. Finger and toe discs are moderate in size but one another as follows: distinct. Fingers lack webbing and toes are less than half L. microbelos with a type locality of Cairns in far north webbed. The disc and nearly two phlanges are free on Queensland, and confined to the Cape York area, has an the outer side of the fourth toe. The inner metatarsal orange iris and tubercles on the flanks are mainly, but not tubercle is prominent and there is no outer one. entirely small, blunt and widely scattered and not in any Tympanum ranges from small to indistinct. Second finger well-defined longitudinal rows. The white labial marking longer than first. under the eye does extend foreward beyond the nostril, The three Australian species within Llewellynura as but is ill-defined at this point. outlined in the description above, being of the nominate L. fukker sp. nov. from the north-west Kimberley in subgenus Llewellynura are separated from the New Western Australia has an orange iris and very few widely Guinea species herein placed in the subgenus scattered tubercles on the flanks. The white labial Microlitoria subgen. nov. by lacking vomerine teeth. marking under the eye extends foreward beyond the These are present in species of Microlitoria subgen. nov..

According to Duellman et al. (2016), the two subgenera The three species can however be readily separated from diverged from one another 14.8 MYA. one another as follows: Distribution: L. fukker sp. nov. appears to be restricted L. microbelos with a type locality of Cairns in far north to the north-west Kimberley Division of Western Queensland, and confined to the Cape York area, has an Australia, centred around the Mitchell Plateau, Western orange iris and tubercles on the flanks are mainly, but not Australia. entirely small, blunt and widely scattered and not in any Etymology: The call is this species is a series of very well-defined longitudinal rows. The white labial marking high-pitched rapidly repeated whirring notes at the rate of under the eye does extend foreward beyond the nostril, about 3 per second. These are produced in bouts lasting but is ill-defined at this point. about 6-10 seconds and increase in speed towards the L. fukker sp. nov. from the north-west Kimberley in end of the series. The call is like that of a small cicada Western Australia has an orange iris and very few widely insect. Frogs call in large choruses including dozens or scattered tubercles on the flanks. The white labial even hundreds of males and individual males will call marking under the eye extends foreward beyond the between notes of nearby males, making the general nostril and is well defined at this point. sound appear as a loud ear pearcing din. L. yehbwudda sp. nov. has a yellow-grey iris (rarely seen The local Aboriginals in the area are called the with a slight orange tinge) and a prominent well-defined Wunambal people. As they try to sleep adjacent to the row of skin folds and tubercles running in a line on either swamps these frogs start calling from, they will wake up side of the back of the dorsum and a second similar well- and yell “Fukker” and hence the species name. defined row of smaller tubercles running along each side LLEWELLYNURA (LLEWELLYNURA) YEHBWUDDA of the mid flank. The white labial marking under the eye SP. NOV. does not extend as far foreward as the nostril. LSIDurn:lsid:zoobank.org:act:616665B3-A265-4CA7- Anstis (2013) on page 247, gives detail of differences A606-196085938E47 between the tadpoles and metamorps of Queensland L. Holotype: A preserved specimen at the Museum and Art microbelos, and Northern Territory L. yehbwudda sp. nov. Gallery of the Northern Territory, Darwin, Northern (which are treated by her provisionally as a separate Territory, Australia, specimen number R22599 collected population of L. microbelos). from Howard Springs, an outer Darwin suburb, Darwin, Significantly the images of two metamorphs (one of each Australia, Latitude -12.45 S., Longitude 131.05 E. species) on page 247 bottom of Anstis (2013) readily This government-owned facility allows access to its shows the diagnostic differences also in the adults of holdings. both species as outlined herein. Paratypes: Eleven preserved specimens at the Museum Photos of L. microbelos in life can be seen in Vanderduys and Art Gallery of the Northern Territory, Darwin, Northern (2012) on page 54 in two images, Cogger (2014) on page Territory, Australia, specimen numbers R00129-R00135, 174 bottom left and Anstis (2013) on page 245 at top left R12063, R22600, R27707 and R27714 all collected from and second from top on right. Howard Springs, an outer Darwin suburb, Darwin, Photos of fukker sp. nov. in life can be seen in Tyler and Australia, Latitude -12.45 S., Longitude 131.05 E. Davies (1986) in the colour plates at number 27 and in Diagnosis: The three species within the nominate Tyler et al. (1994) on plate 32 at bottom. subgenus Llewellynura Wells and Wellington, 1985 have Photos of L. yehbwudda sp. nov. in life can be seen in until 2019 been treated as a single species, namely L. Anstis (2013) on page 245 at top right. microbelos (Cogger, 1966). However this putative taxon Species within the genus Llewellynura Wells and in fact consists of three separate and widely allopatric Wellington, 1985 are readily separated from all other species. Australasian Tree Frogs (Pelodryadidae) by the following This has been known for many years and Anstis (2013) suite of characters: they are tiny in size, being about 20 stated a view that more than one species was within mm in body length as adults. Colour is variegated dark putative L. microbelos as described in her book. and brown dorsally; a row of widely spaced dorsolateral As a trio, they are found in the tropical parts of northern tubercles and ridges may be on each side, or Australia from three main areas being: alternatively just a few randomly scattered small 1/ The north-west Kimberley Division of Western tubercles on either side of upper flank; no pectoral fold; Australia, centred around the Mitchell Plateau (L. fukker several conspicuous tubercles above each eye; tiny, sp. nov.), and: slender and agile and with a sharply pointed snout. The dorsum may be grey, brown, yellow or red, often flecked 2/ The top end of the Northern Territory, from Litchfield or mottled with darker colour. There is usually a broad National Park in the west and extending east to include dark band from behind the eye to the groin and an even Groote Eylandt and also including the Tiwi Islands (L. darker stripe along the snout, through the eye and over yehbwudda sp. nov.), and: the base of the arm to the flank. Snout and upper 3/ Cape York Queensland, extending as far south as surfaces of the limbs are peppered with dark brown. Townsville (L. microbelos). Records south of there are There is a conspicuous dark bar along the front edge of almost certainly specimens inadvertently moved in recent the arm. Ventral surface is white except for a fine times by humans, such as when being translocated in peppering of brown on the throat, chest and limbs. Skin is boxes of fruit via trucks and trains. smooth dorsally, with at least some small tubercles or All three species would key out as L. microbelos in either warts or skin folds and usually several small subercles Anstis (2013) or Cogger (2014). over each eye. Throat skin is smooth, but belly is

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 134 Australasian Journal of Herpetology granular. Finger and toe discs are moderate in size but diagnosis) comprised just one species as in the type distinct. Fingers lack webbing and toes are less than half spcies. A morphologically similar species M. aurifera webbed. The disc and nearly two phlanges are free on (Anstis, Tyler, Roberts, Price and Doughty, 2010) is the outer side of the fourth toe. The inner metatarsal clearly within this genus, as outlined by the authors in tubercle is prominent and there is no outer one. their description. Tympanum ranges from small to indistinct. Second finger Anstis (2013) stated of M. meiriana “may include two longer than first. species”, but gave no indication as to which populations The three Australian species within Llewellynura as they referred to. outlined in the description above, being of the nominate However in line with other biogeographical splits in subgenus Llewellynura are separated from the New northern Australia, the obvious presumption wold be she Guinea species herein placed in the subgenus was referring to the morphologically distinct Kimberley Microlitoria subgen. nov. by lacking vomerine teeth. (West Australia) specimens as being the putative new These are present in species of Microlitoria subgen. nov.. species. According to Duellman et al. (2016), the two subgenera Investigations showed a split within Kimberley animals diverged from one another 14.8 MYA. into two distinctive groups as well as another distinctive Distribution: L. yehbwudda sp. nov. occurs in the top form south of the Gulf of Carpentaria. Yet another end of the Northern Territory, from Litchfield National Park distinctive and geographically isolated population occurs in the west, extending east to include Groote Eylandt and in the Victoria River area of the north-west Northern also including the Tiwi Islands near Darwin. Territory. All four are formally named within this paper as Etymology: These frogs are common on the outskirts of new species on the basis of morphological differences Darwin in the Northern Territory, including the type locality and separation by well-known biogeographical barriers of of Howard Springs. In the 1980’s surviving native known antiquity. Australian Aboriginals, in this region known as the Hence the genus Mahonabatrachus now includes six Larrakia people, were eking out a miserable existence species. after the British Empire stole their land and exterminated Distribution: Rock escarpment country of the top end of most of the population. Bored women and childen would the Northern Territory and nearby Western Australia, scour the vegetation of the local swamp at Howard including the Kimberley District of Western Australia and Springs looking for the small frogs to cook on the the southern edge of the Gulf of Carpentaria. campfire, which they then ate whole. Content: Mahonabatrachus meiriana (Tyler, 1969) (type When a Larrakia child would grab a frog he’d scream to species); M. aurifera (Anstis, Tyler, Roberts, Price and his mates “yehbwudda” and hence the species name for Doughty, 2010); M. chriswilliamsi sp. nov.; M. this taxon. marionanstisae sp. nov.; M. pailsae sp. nov.; M. roypailsi MAHONABATRACHUS WELLS AND WELLINGTON, sp. nov.. 1985 MAHONABATRACHUS CHRISWILLIAMSI SP. NOV. Type species: Hyla meiriana Tyler, 1969. LSIDurn:lsid:zoobank.org:act:D5398E68-EDB4-4F32- Diagnosis: Known as the “Rock Hole Frogs”, living frogs 9EC8-86369DDFEEC3 in the genus Mahonabatrachus Wells and Wellington, Holotype: A preserved specimen at the South Australian 1985 are all readily separated from all other Australasian Museum, Adelaide, South Australia, Australia, specimen Tree Frogs (Pelodryadidae) by the following unique suite number R15638 collected from the Admiralty Gulf area, of characters: In colour they are usually irregularly North-west Kimberley Division, Western Australia, mottled above with metallic fawn, brown or reddish brown Australia. Latitude -14.50 S., Longitude 125.83 E. and dark brown, the small low tubercles on the back This government-owned facility allows access to its sometimes tending to be light-centered and/or dark holdings. edged. Sometimes these frogs are dominantly dark Paratype: A preserved specimen at the Western brown but with a gold-orange dorsolateral stripe from Australian Museum, Perth, Western Australia, Australia, snout to eye and continuing above the eye above the specimen number R44319, collected from Crystal Creek tympanum along the body almost to the groin. Limbs of at Crystal Head, from the Admiralty Gulf area in the all specimens are always coloured with irregular dark North-west Kimberley Division of Western Australia, brown cross bands and all frogs have barred or spotted Australia, Latitude -14.5167 S., Longitude 125.7833 E. lips to some degree and lower surfaces are whitish. Diagnosis: Mahonabatrachus chriswilliamsi sp. nov., M. The skin is leathery to finely granular above with marionanstisae sp. nov., M. pailsae sp. nov. and M. numerous low, rounded or sometimes slightly pointed roypailsi sp. nov. have all been treated as populations of tubercles, that are fairly evenly spaced. No pectoral fold. a widespread putative taxon M. meiriana (Tyler, 1969), Moderate sized but conspicuous diss on fingers and toes. originally described as “Hyla meriana”, with a type locality Fingers free and toes with moderately developed of 98 miles north of Mainoru, Northern Territory (NT), webbing, including reaching the disc of the fifth toe. Australia. Adults average 20 mm in length. The five morphologically similar and obviously closely According to Duellman et al. (2016), the species in this related species can be readily separated on the basis of genus diverged from their nearest living relatives 20.9 differences in morphology and colour. MYA. Nominate M. meiriana is herein confined to the Until recently this genus as defined by Wells and Arnhemland Escarpment of the Northern Territory and Wellington (1985) and as defined herein (with a modified immediately adjacent outliers in a region bound by the Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 135

type locality in Arnhemland (98 miles north of Mainoru, with spikey edge of same colour as pigment below (either Northern Territory, Australia), west to Litchfield National reddish-brown or grey). There are two well-defined white Park, NT. labial bars beneath the eye and at least one on the snout M. pailsae sp. nov. is the taxon in this species group anterior to this (on each side). found in the region generally bound by the Daly River in Hind limbs have semi-distinct grey and brown bands and the north and West Baines/Victoria River in the south- the forelimbs are of grey background colour with a strong west, including escarpment country within this zone. light brown overlay. The front limbs also have a small M. marionanstisae sp. nov. is found generally around the number of well defined irregular shaped, dark brown Ord River region of the Kimberley District of Western spots or blotches. The iris is brilliant red/orange on the Australia including immediately adjacent parts of North- upper half and light grey below, with demarcation west Northern Territory, including areas bounded by the between both being well defined by a black midline. West Baines/Victoria River in the east and Durack River In M. meiriana and the other species except for M. in the West. pailsae sp. nov., the interdigital webbing reaches the M. chriswilliamsi sp. nov. is confined to the north-west base of the terminal discs of all toes except the fourth Kimberley division from Durack River in the east and where it extends as far as the subarticular tubercle at the generally north of the Mitchell Plateau. Specimens in the base of the penultimate phalanx and is united to the disc southern Kimberley are tentatively assigned to this by a narrow lateral fringe. species. M. pailsae sp. nov. is readily separated from the other M. roypailsi sp. nov. is restricted to low escarpments four species by the following suite of characters: This south of the Gulf of Carpentaria in the Northern Territory. species has more extensive webbing of the feet than all All five species would be identified as M. meiriana by the other species, in this taxon being unique among the five diagnosis in Tyler (1969) or that of Cogger (2014). in reaching mid-way up the penultimate phalanx of the All five species and a sixth species in this genus M. fourth toe. aurifera (Anstis, Tyler, Roberts, Price and Doughty, 2010) In colour the dorsum is generally a dark brown, are separated from all other Australasian Tree Frogs accentuated by the dark grey (rather than light grey) (Pelodryadidae) by the following unique suite of interspaces and blotches on the dorsum. The dorsal characters: They are an extremely small species with a surface of the thigh is similar to the colour of the head maximum snout to vent length of 22.5 mm, characterised and back (the light markings on the posterior face do not by an extremely high E-N/IN ratio (1.286-1.600), short extend upon it). The ventral surfaces are much more and unwebbed fingers with prominent, transversely oval heavily and extensively marked; the throat is usually a discs and extensively webbed toes (the webbing reaching uniform dark brown and only the base of the discs of all toes except the fourth); infrequently stippled with brown and the ventral surface of vomerine teeth and outer metatarsal tubercles are the thighs is suffused with brown in most specimens.The present. In life there are striking post-femoral markings; iris is a dull orange colour above and grey with an orange the snout is evenly rounded and not particularly tinge below and there is either no border between upper prominent or projecting. and lower parts of the iris, or if present it is greyish, not I can report for the first time that M. aurifera (Anstis, Tyler, black and peppered and indistinct. Dark labial bars or Roberts, Price and Doughty, 2010) is most simply markings are prominent anterior to the eye. separated from the other five species by having no more Limbs are generally light brown with indistinct darker than five rounded tubercles in a row between front and markings. hind leg on the flank (counted in a line and ignoring stray M. marionanstisae sp. nov. is readily separated from the tubercles above or below the line), versus six or more in other four species by the following suite of characters: A the other five species and the relevant tubercles are generally faded and indistinct colouration of the dorsum, largeish, rounded and blunt in form in M. aurifera, versus which is a combination of indistinct dark brown and light smaller and tending towards being more pointed in the brown markings (versus for example reddish-brown and other five species. It also readily separated from the other light grey in M. meiriana). The demarkation between five species by having an iris that is orange above and upper iris and lower iris by way of black border is either dull orange below with distinctive strips/patches of black indistinct or absent. The upper eye is a dull orange-brown pigment on each of the north-south-east-west axis. and extends well below half-way with only the very lower M. aurifera is further separated from the other five part of the eye either dark grey or greyish in colour, species (all treated by the relevant authors as M. although in many specimens the entire eye is a dull meiriana) by the suite of characters outlined in the orange-brown. Labial bars of any colour (dark or light) original description of Anstis et al. (2010), including a are generally indistinct. Forelimbs are yellowish-brown slightly more pointed snout and tadpoles having a unique and generally unmarked, except for some scattered black, gold and red pigment pattern as well as a patches of darker peppering. Hindlimbs are light brown continuous papillary border around the oral disc (neither with heavy dark peppering giving them a marbled of which are seen in the other five species). appearance. M. meiriana is readily separated from the other four The hind legs are relatively shorter in this species than in species by the following suite of characters: The dorsum the other four. In this species there is a TL/S-V range of is generally dark reddish-brown with light grey blotches 0.476-0.556 with a mean of 0.515, versus TL/S-V range and marks occupying 15 to 40 percent of the dorsum. of 0.541-0.640 and the mean 0.554 in the other four There is an average of 11 moderately enlarged tubercles species (derived from Tyler, 1969).

M. chriswilliamsi sp. nov. is readily separated from the Australia, Australia, generally west of the Durack River other four species by the following suite of characters: A and north of the Mitchell Plateau. Populations from the dorsum that is dark brown in colour with semi-distinct south-west Kimberley are tentatively referred to this black spots and blotches of irregular shape, but generally taxon. scallered across the dorsum and limbs. On the limbs the Etymology: Named in honour of Chris Williams, of New black markings tend to form bars. Many of the tubercles South Wales, Australia, a former president of the on the body are white tipped. Australian Herpetological Society in Sydney, New South In common with M. marionanstisae sp. nov. the pale light Wales, ex employee of Tarango Zoo in Sydney (poor brown patch encompassing the upper heal of the upper thing) and John Weigel’s Reptile Park business (same) in back foot is well defined and prominent. recognition of his many contributions to herpetology in The upper iris is bright red, lower iris is grey and the Australia in often difficult circumstances. black line demarcating the two halves of the eye is well MAHONABATRACHUS MARIONANSTISAE SP. NOV. defined. LSIDurn:lsid:zoobank.org:act:C0D5E065-F72F-45BD- In this species at the top of the iris is a large patch of 96A8-1DEC01B4EA86 dark pigment in the form of black peppering, forming a Holotype: A preserved specimen at the South Australian reasonably large blotch. In all other species there is Museum, Adelaide, South Australia, Australia, specimen either a tiny black dot or nothing at all at the same place number R14513 collected in the Lake Argyle area, in the eye. Kimberley Division of Western Australia, Australia, M. chriswilliamsi sp. nov. is also unusual in that the Latitude -16.22 S., Longitude 128.90 E. This government- tubercles on the upper surface of the back leg are large, owned facility allows access to its holdings. pointed and white-tipped, numbering 6-9. Labial markings Paratypes: Four preserved specimens at the Western are distinct, but the white is either in the form of small Australian Museum, Perth, Western Australia, Australia, spots or alternatively as very narrow bars. Between these specimen numbers R162530-33 from Kununurra, white bits are areas of brown, the brown being the Kimberley Division of Western Australia, Australia, dominant colour of the labials and snout. Latitude 15.6049 S., Longitude 128.7655 E. The upper surface of the head is brown with black spots Diagnosis: Mahonabatrachus chriswilliamsi sp. nov., M. and blotches (versus for example reddish-brown and light marionanstisae sp. nov., M. pailsae sp. nov. and M. grey in M. meiriana). roypailsi sp. nov. have all been treated as populations of M. roypailsi sp. nov. is similar in most respects to M. a widespread putative taxon M. meiriana (Tyler, 1969), meiriana as described above, but the upper iris is dull originally described as “Hyla meriana”, with a type locality orange in colour, labial markings and those on the limbs of 98 miles north of Mainoru, Northern Territory (NT), are indistinct; the anterior of the snout is brownish as Australia. opposed to purplish in colour and the dorsum has beige The five morphologically similar and obviously closely as opposed to light grey interspaces. related species can be readily separated on the basis of M. meiriana in life is depicted on page 173 of Cogger differences in morphology and colour. (2014) and Anstis (2013) on page 242 at top right and Nominate M. meiriana is herein confined to the right middle and online at: Arnhemland Escarpment of the Northern Territory and https://www.flickr.com/photos/stephenmahony/ immediately adjacent outliers in a region bound by the 36225998011/ type locality in Arnhemland (98 miles north of Mainoru, and: Northern Territory, Australia), west to Litchfield National https://www.flickr.com/photos/mattsummerville/ Park, NT. 40940670621/ M. pailsae sp. nov. is the taxon in this species group and: found in the region generally bound by the Daly River in https://www.flickr.com/photos/ryanfrancis/26315692613/ the north and West Baines/Victoria River in the south- and: west, including escarpment country within this zone. https://www.flickr.com/photos/euprepiosaur/7240116716/ M. marionanstisae sp. nov. is found generally around the M. chriswilliamsi sp. nov. in life is depicted online at: Ord River region of the Kimberley District of Western https://www.flickr.com/photos/54876436@N08/ Australia including immediately adjacent parts of North- 19647295261/ west Northern Territory, including areas bounded by the M. marionanstisae sp. nov. is depicted on page 242 of West Baines/Victoria River in the east and Durack River Anstis (2013) at top left and online at: in the West. https://www.flickr.com/photos/23031163@N03/ M. chriswilliamsi sp. nov. is confined to the north-west 8519050483/ Kimberley division from Durack River in the east and and: generally north of the Mitchell Plateau. Specimens in the https://www.flickr.com/photos/14807473@N08/ southern Kimberley are tentatively assigned to this 49397917677/ species. M. pailsae sp. nov. in life is depicted online at: M. roypailsi sp. nov. is restricted to low escarpments https://www.flickr.com/photos/68921296@N06/ south of the Gulf of Carpentaria in the Northern Territory. 14697052000/ All five species would be identified as M. meiriana by the Distribution: Mahonabatrachus chriswilliamsi sp. nov. is diagnosis in Tyler (1969) or that of Cogger (2014). found in the north Kimberley division of Western All five species and a sixth species in this genus M.

aurifera (Anstis, Tyler, Roberts, Price and Doughty, 2010) In colour the dorsum is generally a dark brown, are separated from all other Australasian Tree Frogs accentuated by the dark grey (rather than light grey) (Pelodryadidae) by the following unique suite of interspaces and blotches on the dorsum. The dorsal characters: They are an extremely small species with a surface of the thigh is similar to the colour of the head maximum snout to vent length of 22.5 mm, characterised and back (the light markings on the posterior face do not by an extremely high E-N/IN ratio (1.286-1.600), short extend upon it). The ventral surfaces are much more and unwebbed fingers with prominent, transversely oval heavily and extensively marked; the throat is usually a discs, and extensively webbed toes (the webbing uniform dark brown and only reaching the base of the discs of all toes except the infrequently stippled with brown and the ventral surface of fourth); vomerine teeth and outer metatarsal tubercles the thighs is suffused with brown in most specimens.The are present. In life there are striking post-femoral iris is a dull orange colour above and grey with an orange markings; the snout is evenly rounded and not tinge below and there is either no border between upper particularly prominent or projecting. and lower parts of the iris, or if present it is greyish, not I can report for the first time that M. aurifera (Anstis, Tyler, black and peppered and indistinct. Dark labial bars or Roberts, Price and Doughty, 2010) is most simply markings are prominent anterior to the eye. separated from the other five species by having no more Limbs are generally light brown with indistinct darker than five rounded tubercles in a row between front and markings. hind leg on the flank (counted in a line and ignoring stray M. marionanstisae sp. nov. is readily separated from the tubercles above or below the line), versus six or more in other four species by the following suite of characters: A the other five species and the relevant tubercles are generally faded and indistinct colouration of the dorsum, largeish, rounded and blunt in form in M. aurifera, versus which is a combination of indistinct dark brown and light smaller and tending towards being more pointed in the brown markings (versus for example reddish-brown and other five species. It also readily separated from the other light grey in M. meiriana). The demarkation between five species by having an iris that is orange above and upper iris and lower iris by way of black border is either dull orange below with distinctive strips/patches of black indistinct or absent. The upper eye is a dull orange-brown pigment on each of the north-south-east-west axis. and extends well below half-way with only the very lower M. aurifera is further separated from the other five part of the eye either dark grey or greyish in colour, species (all treated by the relevant authors as M. although in many specimens the entire eye is a dull meiriana) by the suite of characters outlined in the orange-brown. Labial bars of any colour (dark or light) original description of Anstis et al. (2010), including a are generally indistinct. Forelimbs are yellowish-brown slightly more pointed snout and tadpoles having a unique and generally unmarked, except for some scattered black, gold and red pigment pattern as well as a patches of darker peppering. Hindlimbs are light brown continuous papillary border around the oral disc (neither with heavy dark peppering giving them a marbled of which are seen in the other five species). appearance. M. meiriana is readily separated from the other four The hind legs are relatively shorter in this species than in species by the following suite of characters: The dorsum the other four. In this species there is a TL/S-V range of is generally dark reddish-brown with light grey blotches 0.476-0.556 with a mean of 0.515, versus TL/S-V range and marks occupying 15 to 40 percent of the dorsum. of 0.541-0.640 and the mean 0.554 in the other four There is an average of 11 moderately enlarged tubercles species (derived from Tyler, 1969). with spikey edge of same colour as pigment below (either M. chriswilliamsi sp. nov. is readily separated from the reddish-brown or grey). There are two well-defined white other four species by the following suite of characters: A labial bars beneath the eye and at least one on the snout dorsum that is dark brown in colour with semi-distinct anterior to this (on each side). black spots and blotches of irregular shape, but generally Hind limbs have semi-distinct grey and brown bands and scallered across the dorsum and limbs. On the limbs the the forelimbs are of grey background with a strong light black markings tend to form bars. Many of the tubercles brown overlay. The front limbs also have a small number on the body are white tipped. of well defined irregular shaped, dark brown spots or In common with M. marionanstisae sp. nov. the pale light blotches. The iris is brilliant red/orange on the upper half brown patch encompassing the upper heal of the upper and light grey below, with demarcation between both back foot is well defined and prominent. being well defined by a black midline. The upper iris is bright red, lower iris is grey and the In M. meiriana and the other species except for M. black line demarcating the two halves of the eye is well pailsae sp. nov., the interdigital webbing reaches the defined. base of the terminal discs of all toes except the fourth In this species at the top of the iris is a large patch of where it extends as far as the subarticular tubercle at the dark pigment in the form of black peppering, forming a base of the penultimate phalanx and is united to the disc reasonably large blotch. In all other species there is by a narrow lateral fringe. either a tiny black dot or nothing at all at the same place M. pailsae sp. nov. is readily separated from the other in the eye. four species by the following suite of characters: This M. chriswilliamsi sp. nov. is also unusual in that the species has more extensive webbing of the feet than all tubercles on the upper surface of the back leg are large, other species, in this taxon being unique among the five pointed and white-tipped, numbering 6-9. Labial markings in reaching mid-way up the penultimate phalanx of the are distinct, but the white is either in the form of small fourth toe. spots or alternatively as very narrow bars. Between these

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 138 Australasian Journal of Herpetology white bits are areas of brown, the brown being the 26, also collected from the type locality (above). dominant colour of the labials and snout. Diagnosis: Mahonabatrachus chriswilliamsi sp. nov., M. The upper surface of the head is brown with black spots marionanstisae sp. nov., M. pailsae sp. nov. and M. and blotches (versus for example reddish-brown and light roypailsi sp. nov. have all been treated as populations of grey in M. meiriana). a widespread putative taxon M. meiriana (Tyler, 1969), M. roypailsi sp. nov. is similar in most respects to M. originally described as “Hyla meriana”, with a type locality meiriana as described above, but the upper iris is dull of 98 miles north of Mainoru, Northern Territory (NT), orange in colour, labial markings and those on the limbs Australia. are indistinct; the anterior of the snout is brownish as The five morphologically similar and obviously closely opposed to purplish in colour and the dorsum has beige related species can be readily separated on the basis of as opposed to light grey interspaces. differences in morphology and colour. M. meiriana in life is depicted on page 173 of Cogger Nominate M. meiriana is herein confined to the (2014) and Anstis (2013) on page 242 at top right and Arnhemland Escarpment of the Northern Territory and right middle and online at: immediately adjacent outliers in a region bound by the https://www.flickr.com/photos/stephenmahony/ type locality in Arnhemland (98 miles north of Mainoru, 36225998011/ Northern Territory, Australia), west to Litchfield National and: Park, NT. https://www.flickr.com/photos/mattsummerville/ M. pailsae sp. nov. is the taxon in this species group 40940670621/ found in the region generally bound by the Daly River in and: the north and West Baines/Victoria River in the south- https://www.flickr.com/photos/ryanfrancis/26315692613/ west, including escarpment country within this zone. and: M. marionanstisae sp. nov. is found generally around the https://www.flickr.com/photos/euprepiosaur/7240116716/ Ord River region of the Kimberley District of Western M. chriswilliamsi sp. nov. in life is depicted online at: Australia including immediately adjacent parts of North- https://www.flickr.com/photos/54876436@N08/ west Northern Territory, including areas bounded by the 19647295261/ West Baines/Victoria River in the east and Durack River M. marionanstisae sp. nov. is depicted on page 242 of in the West. Anstis (2013) at top left and online at: M. chriswilliamsi sp. nov. is confined to the north-west https://www.flickr.com/photos/23031163@N03/ Kimberley division from Durack River in the east and 8519050483/ generally north of the Mitchell Plateau. Specimens in the and: southern Kimberley are tentatively assigned to this https://www.flickr.com/photos/14807473@N08/ species. 49397917677/ M. roypailsi sp. nov. is restricted to low escarpments M. pailsae sp. nov. in life is depicted online at: south of the Gulf of Carpentaria in the Northern Territory. https://www.flickr.com/photos/68921296@N06/ All five species would be identified as M. meiriana by the 14697052000/ diagnosis in Tyler (1969) or that of Cogger (2014). Distribution: M. marionanstisae sp. nov. is found All five species and a sixth species in this genus M. generally around the Ord River region of the Kimberley aurifera (Anstis, Tyler, Roberts, Price and Doughty, 2010) District of Western Australia including immediately are separated from all other Australasian Tree Frogs adjacent parts of North-west Northern Territory. It is found (Pelodryadidae) by the following unique suite of in the area bounded by the West Baines/Victoria River in characters: They are an extremely small species with a the east and Durack River in the West. maximum snout to Etymology: Named in honour of Marion Anstis of New vent length of 22.5 mm, characterised by an extremely South Wales, Australia, author of numerous books and high E-N/IN ratio (1.286-1.600), short and unwebbed papers on frogs and a past president of the Sydney- fingers with prominent, transversely oval discs and based Australian Herpetological Society, in recognition of extensively webbed toes (the webbing reaching the base her wide contributions to herpetology over more than 40 of the discs of all toes except the fourth); vomerine teeth years. and outer metatarsal tubercles are present. In life there MAHONABATRACHUS PAILSAE SP. NOV. are striking post-femoral markings; the snout is evenly rounded and not particularly prominent or projecting. LSIDurn:lsid:zoobank.org:act:28C5D12E-F360-4C85- 961E-ACF32EE5882E I can report for the first time that M. aurifera (Anstis, Tyler, Roberts, Price and Doughty, 2010) is most simply Holotype: A preserved specimen in the National separated from the other five species by having no more Museum of Victoria, Melbourne, Victoria, Australia, than five rounded tubercles in a row between front and specimen number D10811, collected at Japser Gorge, hind leg on the flank (counted in a line and ignoring stray Northern Territory, Australia, Latitude -16.03 S., tubercles above or below the line), versus six or more in Longitude 130.68 E. the other five species and the relevant tubercles are This government-owned facility allows access to its largeish, rounded and blunt in form in M. aurifera, versus holdings. smaller and tending towards being more pointed in the Paratypes: 16 preserved specimens at the National other five species. It also readily separated from the other Museum of Victoria, Melbourne, Victoria, Australia, five species by having an iris that is orange above and specimen numbers D10773-74, D10812-16 and D10818- dull orange below with distinctive strips/patches of black Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 139

pigment on each of the north-south-east-west axis. indistinct or absent. The upper eye is a dull orange-brown M. aurifera is further separated from the other five and extends well below half-way with only the very lower species (all treated by the relevant authors as M. part of the eye either dark grey or greyish in colour, meiriana) by the suite of characters outlined in the although in many specimens the entire eye is a dull original description of Anstis et al. (2010), including a orange-brown. Labial bars of any colour (dark or light) slightly more pointed snout and tadpoles having a unique are generally indistinct. Forelimbs are yellowish-brown black, gold and red pigment pattern as well as a and generally unmarked, except for some scattered continuous papillary border around the oral disc (neither patches of darker peppering. Hindlimbs are light brown of which are seen in the other five species). with heavy dark peppering giving them a marbled M. meiriana is readily separated from the other four appearance. species by the following suite of characters: The dorsum The hind legs are relatively shorter in this species than in is generally dark reddish-brown with light grey blotches the other four. In this species there is a TL/S-V range of and marks occupying 15 to 40 percent of the dorsum. 0.476-0.556 with a mean of 0.515, versus TL/S-V range There is an average of 11 moderately enlarged tubercles of 0.541-0.640 and the mean 0.554 in the other four with spikey edge of same colour as pigment below (either species (derived from Tyler, 1969). reddish-brown or grey). There are two well-defined white M. chriswilliamsi sp. nov. is readily separated from the labial bars beneath the eye and at least one on the snout other four species by the following suite of characters: A anterior to this (on each side). dorsum that is dark brown in colour with semi-distinct Hind limbs have semi-distinct grey and brown bands and black spots and blotches of irregular shape, but generally the forelimbs are of grey background with a strong light scallered across the dorsum and limbs. On the limbs the brown overlay. The front limbs also have a small number black markings tend to form bars. Many of the tubercles of well defined irregular shaped, dark brown spots or on the body are white tipped. blotches. The iris is brilliant red/orange on the upper half In common with M. marionanstisae sp. nov. the pale light and light grey below, with demarcation between both brown patch encompassing the upper heal of the upper being well defined by a black midline. back foot is well defined and prominent. In M. meiriana and the other species except for M. The upper iris is bright red, lower iris is grey and the pailsae sp. nov., the interdigital webbing reaches the black line demarcating the two halves of the eye is well base of the terminal discs of all toes except the fourth defined. where it extends as far as the subarticular tubercle at the In this species at the top of the iris is a large patch of base of the penultimate phalanx and is united to the disc dark pigment in the form of black peppering, forming a by a narrow lateral fringe. reasonably large blotch. In all other species there is M. pailsae sp. nov. is readily separated from the other either a tiny black dot or nothing at all at the same place four species by the following suite of characters: This in the eye. species has more extensive webbing of the feet than all M. chriswilliamsi sp. nov. is also unusual in that the other species, in this taxon being unique among the five tubercles on the upper surface of the back leg are large, in reaching mid-way up the penultimate phalanx of the pointed and white-tipped, numbering 6-9. Labial markings fourth toe. are distinct, but the white is either in the form of small In colour the dorsum is generally a dark brown, spots or alternatively as very narrow bars. Between these accentuated by the dark grey (rather than light grey) white bits are areas of brown, the brown being the interspaces and blotches on the dorsum. The dorsal dominant colour of the labials and snout. surface of the thigh is similar to the colour of the head The upper surface of the head is brown with black spots and back (the light markings on the posterior face do not and blotches (versus for example reddish-brown and light extend upon it). The ventral surfaces are much more grey in M. meiriana). heavily and extensively marked; the throat is usually a M. roypailsi sp. nov. is similar in most respects to M. uniform dark brown and only meiriana as described above, but the upper iris is dull infrequently stippled with brown, and the ventral surface orange in colour, labial markings and those on the limbs of the thighs is suffused with brown in most are indistinct; the anterior of the snout is brownish as specimens.The iris is a dull orange colour above and opposed to purplish in colour and the dorsum has beige grey with an orange tinge below and there is either no as opposed to light grey interspaces. border between upper and lower parts of the iris, or if M. meiriana in life is depicted on page 173 of Cogger present it is greyish, not black and peppered and (2014) and Anstis (2013) on page 242 at top right and indistinct. Dark labial bars or markings are prominent right middle and online at: anterior to the eye. https://www.flickr.com/photos/stephenmahony/ Limbs are generally light brown with indistinct darker 36225998011/ markings. and: M. marionanstisae sp. nov. is readily separated from the https://www.flickr.com/photos/mattsummerville/ other four species by the following suite of characters: A 40940670621/ generally faded and indistinct colouration of the dorsum, and: which is a combination of indistinct dark brown and light https://www.flickr.com/photos/ryanfrancis/26315692613/ brown markings (versus for example reddish-brown and and: light grey in M. meiriana). The demarkation between https://www.flickr.com/photos/euprepiosaur/7240116716/ upper iris and lower iris by way of black border is either M. chriswilliamsi sp. nov. in life is depicted online at: Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 140 Australasian Journal of Herpetology https://www.flickr.com/photos/54876436@N08/ in the West. 19647295261/ M. chriswilliamsi sp. nov. is confined to the north-west M. marionanstisae sp. nov. is depicted on page 242 of Kimberley division from Durack River in the east and Anstis (2013) at top left and online at: generally north of the Mitchell Plateau. Specimens in the https://www.flickr.com/photos/23031163@N03/ southern Kimberley are tentatively assigned to this 8519050483/ species. and: M. roypailsi sp. nov. is restricted to low escarpments https://www.flickr.com/photos/14807473@N08/ south of the Gulf of Carpentaria in the Northern Territory. 49397917677/ All five species would be identified as M. meiriana by the M. pailsae sp. nov. in life is depicted online at: diagnosis in Tyler (1969) or that of Cogger (2014). https://www.flickr.com/photos/68921296@N06/ All five species and a sixth species in this genus M. 14697052000/ aurifera (Anstis, Tyler, Roberts, Price and Doughty, 2010) Distribution: M. pailsae sp. nov. is found in the region are separated from all other Australasian Tree Frogs generally bound by the Daly River in the north and West (Pelodryadidae) by the following unique suite of Baines/Victoria River in the south-west, including characters: They are an extremely small species with a escarpment country within this zone. maximum snout to vent length of 22.5 mm, characterised Etymology: Named in honour of Lyn Pails, wife of Roy by an extremely high E-N/IN ratio (1.286-1.600), short Pails, a well-known herpetologist from Ballarat, Victoria, and unwebbed fingers with prominent, transversely oval Australia, who along with her husband has made a discs and extensively webbed toes (the webbing reaching valuable contribution to herpetology spanning many the base of the discs of all toes except the fourth); decades. vomerine teeth and outer metatarsal tubercles are present. In life there are striking post-femoral markings; MAHONABATRACHUS ROYPAILSI SP. NOV. the snout is evenly rounded and not particularly LSIDurn:lsid:zoobank.org:act:AAF1E11E-E5B2-49E0- prominent or projecting. AE67-592D37998585 I can report for the first time that M. aurifera (Anstis, Tyler, Holotype: A preserved specimen at the Australian Roberts, Price and Doughty, 2010) is most simply Museum, Sydney, New South Wales, Australia, specimen separated from the other five species by having no more number R.55405 collected at Glyde River, being about 10 than five rounded tubercles in a row between front and km east of the Mcarthur River Camp, in the Northern hind leg on the flank (counted in a line and ignoring stray Territory, Australia, Latitude -16.433 S., Longitude tubercles above or below the line), versus six or more in 136.150 E. the other five species and the relevant tubercles are This government-owned facility allows access to its largeish, rounded and blunt in form in M. aurifera, versus holdings. smaller and tending towards being more pointed in the Paratypes: Six preserved specimens at the Australian other five species. It also readily separated from the other Museum, Sydney, New South Wales, Australia, specimen five species by having an iris that is orange above and numbers R.55414, R.55377, R.55387, R.55396, dull orange below with distinctive strips/patches of black R.55406 and R.55429 all from the type locality (above). pigment on each of the north-south-east-west axis. Diagnosis: Mahonabatrachus chriswilliamsi sp. nov., M. M. aurifera is further separated from the other five marionanstisae sp. nov., M. pailsae sp. nov. and M. species (all treated by the relevant authors as M. roypailsi sp. nov. have all been treated as populations of meiriana) by the suite of characters outlined in the a widespread putative taxon M. meiriana (Tyler, 1969), original description of Anstis et al. (2010), including a originally described as “Hyla meriana”, with a type locality slightly more pointed snout and tadpoles having a unique of 98 miles north of Mainoru, Northern Territory (NT), black, gold and red pigment pattern as well as a Australia. continuous papillary border around the oral disc (neither The five morphologically similar and obviously closely of which are seen in the other five species). related species can be readily separated on the basis of M. meiriana is readily separated from the other four differences in morphology and colour. species by the following suite of characters: The dorsum Nominate M. meiriana is herein confined to the is generally dark reddish-brown with light grey blotches Arnhemland Escarpment of the Northern Territory and and marks occupying 15 to 40 percent of the dorsum. immediately adjacent outliers in a region bound by the There is an average of 11 moderately enlarged tubercles type locality in Arnhemland (98 miles north of Mainoru, with spikey edge of same colour as pigment below (either Northern Territory, Australia), west to Litchfield National reddish-brown or grey). There are two well-defined white Park, NT. labial bars beneath the eye and at least one on the snout M. pailsae sp. nov. is the taxon in this species group anterior to this (on each side). found in the region generally bound by the Daly River in Hind limbs have semi-distinct grey and brown bands and the north and West Baines/Victoria River in the south- the forelimbs are of grey background with a strong light west, including escarpment country within this zone. brown overlay. The front limbs also have a small number M. marionanstisae sp. nov. is found generally around the of well defined irregular shaped, dark brown spots or Ord River region of the Kimberley District of Western blotches. The iris is brilliant red/orange on the upper half Australia including immediately adjacent parts of North- and light grey below, with demarcation between both west Northern Territory, including areas bounded by the being well defined by a black midline. West Baines/Victoria River in the east and Durack River In M. meiriana and the other species except for M.

pailsae sp. nov., the interdigital webbing reaches the defined. base of the terminal discs of all toes except the fourth In this species at the top of the iris is a large patch of where it extends as far as the subarticular tubercle at the dark pigment in the form of black peppering, forming a base of the penultimate phalanx and is united to the disc reasonably large blotch. In all other species there is by a narrow lateral fringe. either a tiny black dot or nothing at all at the same place M. pailsae sp. nov. is readily separated from the other in the eye. four species by the following suite of characters: This M. chriswilliamsi sp. nov. is also unusual in that the species has more extensive webbing of the feet than all tubercles on the upper surface of the back leg are large, other species, in this taxon being unique among the five pointed and white-tipped, numbering 6-9. Labial markings in reaching mid-way up the penultimate phalanx of the are distinct, but the white is either in the form of small fourth toe. spots or alternatively as very narrow bars. Between these In colour the dorsum is generally a dark brown, white bits are areas of brown, the brown being the accentuated by the dark grey (rather than light grey) dominant colour of the labials and snout. interspaces and blotches on the dorsum. The dorsal The upper surface of the head is brown with black spots surface of the thigh is similar to the colour of the head and blotches (versus for example reddish-brown and light and back (the light markings on the posterior face do not grey in M. meiriana). extend upon it). The ventral surfaces are much more M. roypailsi sp. nov. is similar in most respects to M. heavily and extensively marked; the throat is usually a meiriana as described above, but the upper iris is dull uniform dark brown and only orange in colour, labial markings and those on the limbs infrequently stippled with brown and the ventral surface of are indistinct; the anterior of the snout is brownish as the thighs is suffused with brown in most specimens.The opposed to purplish in colour and the dorsum has beige iris is a dull orange colour above and grey with an orange as opposed to light grey interspaces. tinge below and there is either no border between upper M. meiriana in life is depicted on page 173 of Cogger and lower parts of the iris, or if present it is greyish, not (2014) and Anstis (2013) on page 242 at top right and black and peppered and indistinct. Dark labial bars or right middle and online at: markings are prominent anterior to the eye. https://www.flickr.com/photos/stephenmahony/ Limbs are generally light brown with indistinct darker 36225998011/ markings. and: M. marionanstisae sp. nov. is readily separated from the https://www.flickr.com/photos/mattsummerville/ other four species by the following suite of characters: A 40940670621/ generally faded and indistinct colouration of the dorsum, and: which is a combination of indistinct dark brown and light https://www.flickr.com/photos/ryanfrancis/26315692613/ brown markings (versus for example reddish-brown and and: light grey in M. meiriana). The demarkation between https://www.flickr.com/photos/euprepiosaur/7240116716/ upper iris and lower iris by way of black border is either M. chriswilliamsi sp. nov. in life is depicted online at: indistinct or absent. The upper eye is a dull orange-brown https://www.flickr.com/photos/54876436@N08/ and extends well below half-way with only the very lower 19647295261/ part of the eye either dark grey or greyish in colour, M. marionanstisae sp. nov. is depicted on page 242 of although in many specimens the entire eye is a dull Anstis (2013) at top left and online at: orange-brown. Labial bars of any colour (dark or light) https://www.flickr.com/photos/23031163@N03/ are generally indistinct. Forelimbs are yellowish-brown 8519050483/ and generally unmarked, except for some scattered patches of darker peppering. Hindlimbs are light brown and: with heavy dark peppering giving them a marbled https://www.flickr.com/photos/14807473@N08/ appearance. 49397917677/ The hind legs are relatively shorter in this species than in M. pailsae sp. nov. in life is depicted online at: the other four. In this species there is a TL/S-V range of https://www.flickr.com/photos/68921296@N06/ 0.476-0.556 with a mean of 0.515, versus TL/S-V range 14697052000/ of 0.541-0.640 and the mean 0.554 in the other four Distribution: M. roypailsi sp. nov. is restricted to low species (derived from Tyler, 1969). escarpments south of the Gulf of Carpentaria in the M. chriswilliamsi sp. nov. is readily separated from the Northern Territory. other four species by the following suite of characters: A Etymology: M. roypailsi sp. nov. is named in honour of dorsum that is dark brown in colour with semi-distinct Roy Pails, a well-known herpetologist from Ballarat, black spots and blotches of irregular shape, but generally Victoria, Australia in recognition for his many services to scallered across the dorsum and limbs. On the limbs the herpetology spanning many decades. He has been a black markings tend to form bars. Many of the tubercles world leader in breeding numerous species of reptile over on the body are white tipped. many decades. His website at: http:// In common with M. marionanstisae sp. nov. the pale light www.pailsforscales.com.au/about/ describes himself as brown patch encompassing the upper heal of the upper follows: back foot is well defined and prominent. “There comes a time in your life when you meet a bloke The upper iris is bright red, lower iris is grey and the and you’re convinced he’s lost his marbles. The man black line demarcating the two halves of the eye is well behind Pails For Scales!”

This is how Roy comes across to many people. He is Tympanum distinct. thoroughly obsessed with reptiles and literally lives and Duellman et al. (2016) found that the three species in this breathes them every waking hour of his life. The man is a genus grouping diverged from their nearest living living legend! relatives 20.8 MYA, which alone strongly supports the His current enterprise “Pails for scales conservation” is contention that these species should be placed in a fighting an immensely difficult campaign for wildlife separate genus. Morphological divergence of the said conservation. Competing for funds against government- taxa further vindicates this contention. owned zoos and predatory property enterprises like the Distribution: The three species are confined to Terri Irwin business means that on the ground and escarpments of Arnhemland, Northern Territory, Australia “hands on” wildlife conservation enterprises like those of Salmocularana personata (Tyler, Davies and Martin, Pails are being literally starved of funds. 1978); the east Kimberley District of Western Australia S. This is happening in 2020, while the government-owned staccato (Doughty and Anstis, 2007) and the north-west and government backed fake conservation business Kimberley District of Western Australia S. saxacola sp. enterprises masquerading as charities, squander cash in nov.. the form of donations from misguided donors and tax- Etymology: In Latin Salmocularana is an abbreviation of payer funded hand outs in the name of wildlife the words “Salmon eyed frog” in reflection of the usual conservation and literally do nothing significantly useful to colour of the upper iris in these species. save threatened species. Content: Salmocularana personata (Tyler, Davies and SALMOCULARANINA SUBTRIBE NOV. Martin, 1978) (type species); S. saxacola sp. nov.; S. SALMOCULARANA GEN. NOV. staccato (Doughty and Anstis, 2007). LSIDurn:lsid:zoobank.org:act:44A0AE6A-A67A-48D8- SALMOCULARANA SAXACOLA SP. NOV. 9286-CDF304C066A4 LSIDurn:lsid:zoobank.org:act:27E63C72-487A-4563- Type species: Litoria personata Tyler, Davies and Martin, 8FA9-96790A7F1193 1978. Holotype: A preserved adult female specimen at the Diagnosis: The three species in the genus Western Australian Museum, Perth, Western Australia, Salmocularana gen. nov. are small species that breed in Australia, specimen number R167737 collected at Little escarpment rock hills in Arnhemland, Northern Territory Mertens Falls, Mitchell Plateau, West Kimberley District, and also the Kimberley Ranges of north-west Australia. Western Australia, Australia, Latitude -14.8222 S., The three known living species within the genus Longitude 125.7108 E. This government-owned facility Salmocularana gen. nov. (one formally described within allows access to its holdings. this paper) are readily separated from all other Paratypes: Three preserved specimens at the Western Australasian Tree Frogs (Pelodryadidae) by the following Australian Museum, Perth, Western Australia, Australia, unique suite of characters: specimen numbers R167738, R167739 and R167740 Pale grey-brown, fawn, reddish, orange or pink above, collected from Little Mertens Falls, Mitchell Plateau, West with dorsal markings ranging from nothing more than Kimberley District, Western Australia, Australia, Latitude - slight peppering at the anterior part of the dorsum as 14.8222 S., Longitude 125.7108 E. seen in Salmocularana personata (Tyler, Davies and Diagnosis: Until now, S. saxacola sp. nov. would have Martin, 1978), indistinct marbling and blotches as seen in been diagnosed and treated as a population of S. S. staccato (Doughty and Anstis, 2007) or an intense staccato (Doughty and Anstis, 2007) as defined by Anstis randomised configuration of dark brown pigment on a (2013) or Cogger (2014), under the name “Litoria light beige to grey background in S. saxacola sp. nov.; staccato”. specimens have a distinct or semi-distinct dark brownish The two geographically disjunct species and the closely or purple head stripe from just in front of the nostril, related and morphologically similar species S. personata through the eye and ear to upper flank just behind the (Tyler, Davies and Martin, 1978) are however readily forelimbs. Lips are whitish to some extent, ranging from a separated as follows: By colour they are separated as distinct yellow bar, to merely light and peppered brown. follows: All may be pale grey-brown, fawn, reddish, Ventral surfaces whitish. Skin may be smooth, with orange or pink above. extremely tiny scattered tubercles above as in S. Each species has dorsal markings as follows: personata, or with a very limited number of scattered 1/ Nothing more than slight peppering at the anterior part small tubercles above the arm and on the upper flank of the dorsum as seen in S. personata. and nearby dorsum in S. staccato to scattered medium sized tubercles above the arm and on the upper flank 2/ Indistinct marbling and blotches as seen in S. staccato. and nearby dorsum in S. saxacola sp. nov.. Underneath 3/ An intense randomised configuration of dark brown the skin is coarsely granular, except on the throat, where pigment on a light beige to grey background in S. it is smooth. Vomerine teeth are in two short clumps saxacola sp. nov.. between the choanae. Finger and toe discs are The three species are further separated by one or other moderate, wider than the digits. Fingers lack webbing, of the following dorsal skin configurations: toes being about half webbed, including not reaching the 1/ Skin may be smooth, with extremely tiny scattered disc of the fifth toe but extending no more than half way tubercles above as in S. personata. along the penultimate phalanx. Second finger is longer 2/ Skin is smooth with a very limited number of scattered than the first. There is a small oval inner metatarsal small tubercles above the arm and on the upper flank tubercle and a smaller outer metatarsal tubercle. and nearby dorsum in S. staccato.

3/ Skin is smooth with scattered medium sized tubercles or purple head stripe from just in front of the nostril, above the arm and on the upper flank and nearby through the eye and ear to upper flank just behind the dorsum in S. saxacola sp. nov.. forelimbs. Lips are whitish to some extent, ranging from a S. personata has a well-defined, thick dark brown stripe distinct yellow bar, to merely light and peppered brown. running from snout, through eye, the entire tympanum Ventral surfaces whitish. Skin may be smooth, with and along the entire flank, although it tends to fade extremely tiny scattered tubercles above as in S. posteriorly to become a greyish purple colour. personata, or with a very limited number of scattered By contrast the same stripe is indistinct in S. staccato small tubercles above the arm and on the upper flank and never goes midway along the flank. and nearby dorsum in S. staccato to scattered medium In S. saxacola sp. nov. the same stripe is broken along sized tubercles above the arm and on the upper flank the entire length, but the contrast between dark and light and nearby dorsum in S. saxacola sp. nov.. Underneath is sharp and the blotches of the line do run along the the skin is coarsely granular, except on the throat, where entire length of the flank of the body. The dark colour is it is smooth. Vomerine teeth are in two short clumps usually purple to brown and the light of the dorsum is between the choanae. Finger and toe discs are usually beige to light grey. There is a well-defined thin moderate, wider than the digits. Fingers lack webbing, dark crescent running along the upper ridge of the toes being about half webbed, including not reaching the tympanum in this species that is not seen in either other disc of the fifth toe but extending no more than half way species. along the penultimate phalanx. Second finger is longer than the first. There is a small oval inner metatarsal Fingers and toes of S. personata and S. staccato are tubercle and a smaller outer metatarsal tubercle. whitish and without obvious markings, at most having Tympanum distinct. indistinct tinges in parts, versus whitish with obvious and well-defined purplish spots, bars, flecks or blotches. Duellman et al. (2016) found that the three species in this genus grouping diverged from their nearest living The three species can also be differentiated by their relatives 20.8 MYA, which alone strongly supports the tadpole stages. S. personata has large pre- contention that these species should be placed in a metamorphosis tadpoles that are blackish grey in dorsal separate genus. Morphological divergence of the said colour with two distinct yellow lines running down the taxa further vindicates this contention. dorsum on either side of the medial line and continuing down the upper sides of each side of most of the tail. By Distribution: S. saxacola sp. nov. is known only from the contrast the tadpoles of the other two species at the escarpment country of the north-west Kimberley Division same stage of development are yellowish brown in of Western Australia. More specifically the known range colour. of this species is bound by Spring Creek, Latitude - 15.1997 S., Longitude 126.0881 E. in the east, Katers In tadpoles of S. staccato the two distinct yellow lines Island, Latitude - 4.4666 S., Longitude 125.5333 E. running down the dorsum on either side of the medial line in the north and Harding Range, Latitude -16.3231 S., are effectively absent. Longitude 124.7589 E. in the south. The ocean to the In S. saxacola sp. nov. the two distinct yellow lines west otherwise constrains the species. running down the dorsum on either side of the medial line Etymology: The species name derives from the rock- are present on the body, but not on the tail. They are dwelling habits of these frogs, being confined to nowhere near as brilliant in colour or contrast as those extremely rocky escarpment country. seen in S. personata. GENUS LITORIA TSCHUDI, 1838 Anstis (2013) provides comparative images of all of S. personata, S. saxacola sp. nov. and S. staccato, Type species: Litoria freycineti Tschudi, 1838. including images of tadpoles and metamorphosed frogs Diagnosis: The genus Litoria, with type species L. of each species, all of which clearly show the diagnostic freycineti Tschudi, 1838 is herein restricted to the type differences between each species detailed herein. species and the closely related L. latopalmata Günther, In terms of S. saxacola sp. nov., Anstis (2013) has 1867, which as a pair diverged from their nearest images of this species in life on pages 315 (top right), common ancestor 13.3 MYA according to Duellman et al. 316, tadpoles at top and two images below on right, and (2016). page 317 at bottom left (young frog). This divergence and morphological differences are exactly why the species previously included in Litoria The three known living species within the genus have been assigned to other genera. Pustulatarana gen. nov. (one formally described within this paper) are readily separated from all other Litoria as defined herein, are readily separated from all Australasian Tree Frogs (Pelodryadidae) by the following other Australasian Tree frogs (Pelodryadidae) by the unique suite of characters: following suite of characters: Species are a frog that is fawn to dark above, being either immaculate or with Pale grey-brown, fawn, reddish, orange or pink above, markings on the back, with or without a warty exterior, the with dorsal markings ranging from nothing more than warts if present being small and flattish, the markings if slight peppering at the anterior part of the dorsum as present typically being a series of darker blotches or seen in Salmocularana personata (Tyler, Davies and variegations in either an irregular or regular pattern. Martin, 1978), indistinct marbling and blotches as seen in There is a broad dark, canthal stripe, almost completely S. staccato (Doughty and Anstis, 2007) or an intense interrupted in front of the eye and continues behind the randomised configuration of dark brown pigment on a eye to effectively overwrite the tympanum and extend to light beige to grey background in S. saxacola sp. nov.; the flanks where it invariably breaks up into a series of specimens have a distinct or semi-distinct dark brownish Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 144 Australasian Journal of Herpetology black spots or blotches extending ro the groin. There is a partially barred. The groin and hind side of thighs are pale glandular stripe from below the eye to the base of pale yellow, spotted and variegated with dark brown. the forelimb. Limbs are variegated darker but usually with Venter is whitish. Dorsal surface is smooth and leathery, an irregular pattern of spots, blotches or bars. Lower jaw throat is smooth and the venter granular. Vomerine teeth is variegated with yellow and dark brown, often forming a are between the choanae. Finger discs are small and reticulum, often with a barred appearance. The venter is barely distinct, while toe discs are of similar size. Fingers whitish. Ventral surface and flanks are granular. There is are unwebbed and toes extensive webbing between no dorsolateral skin fold. Finger and toe discs are small them, with the webbing reaching the disc of the fifth toe. and project slightly but noticeably, beyond the lateral There is a small inner and tiny outer metatarsal tubercle. edges of the penultimate phlanges. Fingers are free, Tympanum is distinct and second finger is slightly shorter without webbing. Toes have well developed webbing, the than the first. Body length averages 75 mm (subgenus webbing not reaching the disc of the fifth toe and Ferelitoria subgen. nov.). extending no more than half way along the penultimate Distribution: The subgenus Paralitoria subgen. nov. is phalanx. There is a small inner and minute outer found on the eastern third of Cape York Peninsula, Torres metatarsal tubercle. Vomerine teeth are in two clusters Strait islands and southern New Guinea. Subgenus between the choanae. The tympanum is distinct and Ferelitoria subgen. nov. is found in the dry tropics of adults average 40-45 mm snout to rear. northern Australia, being the top end of the Northern Distribution: Most of the eastern third of Australia. Territory and Western Australia. Content: L. freycineti Tschudi, 1838 (type species); L. Etymology: Para in Latin means “not quite” and hence latopalmata Günther, 1867. the name “Paralitoria” meaning not quite Litoria. PARALITORIA GEN. NOV. Content: Paralitoria nigrofrenata (Günther, 1867) (type LSIDurn:lsid:zoobank.org:act:65126912-F61A-4380- species); P. spaldingi (Hosmer, 1964); P. watjulumensis 9CD2-05592C940390 (Copland, 1957). Type species: Hyla nigrofrenata Günther, 1867. FERELITORIA SUBGEN. NOV. Diagnosis: The genus Paralitoria gen. nov. diverged LSIDurn:lsid:zoobank.org:act:45B26663-ECD5-46D7- from its nearest common ancestor, being that of the B907-49C08FF6C5B8 genera Litoria and Quasilitoria gen. nov. being the most Type species: Hyla latopalmata watjulumensis Copland, closely related genera, some 15.5 MYA according to 1957. Duellman et al. (2016). Diagnosis: The genus Paralitoria gen. nov. diverged Paralitoria gen. nov. is separated from all other from its nearest common ancestor, being that also of the Australasian tree frogs (Pelodryadidae) by the following genera Litoria and Quasilitoria gen. nov. some 15.5 MYA suite of characters, being one or other of the following: according to Duellman et al. (2016). 1/ Fawn, grey or brown above, being immaculate; a Paralitoria gen. nov. is separated from all other conspicuous black stripe from the snout, through the Australasian tree frogs (Pelodryadidae) by the following nostril to the eye, where it is almost entirely broken by a suite of characters, being one or other of the following: pale vertical bar in front of the eye as a band continuing 1/ Fawn, grey or brown above, with limited flecks or other below the tympanum and extending back to the forearm markings, often as a peppering; a conspicuous narrow with minimal loss of width along the length, where after a black stripe from the snout, through the nostril to the eye, break, it continues obliquely along the mid flank. There is continuing through the tympanum (effectively over-writing a pale glandular stripe from below the eye to the forearm. it) and extending past the forearm with minimal loss of There is a conspicuous dark brown stripe along the front width along the length to the anterior flank. There is a edge of the tibial region. Lips are not spotted or barred. pale glandular stripe from below the eye to the forearm. The groin and hind side of thighs are pale yellow, spotted There is a conspicuous dark brown stripe along the front and variegated with dark brown. Venter is whitish. Dorsal edge of the tibial region. The lower lips are faintly or surface is smooth and leathery, throat is smooth and the partially barred. The groin and hind side of thighs are venter granular. Vomerine teeth are between the pale yellow, spotted and variegated with dark brown. choanae. Finger discs are small and distinct, while toe Venter is whitish. Dorsal surface is smooth and leathery, discs are smaller and indistinct. Fingers are unwebbed throat is smooth and the venter granular. Vomerine teeth and toes have moderate webbing between them. There is are between the choanae. Finger discs are small and a small inner and tiny outer metatarsal tubercle. barely distinct, while toe discs are of similar size. Fingers Tympanum is distinct and second finger is slightly shorter are unwebbed and toes extensive webbing between than the first. Body length averages 50 mm (subgenus them, with the webbing reaching the disc of the fifth toe. Paralitoria subgen. nov.), or: There is a small inner and tiny outer metatarsal tubercle. 2/ Fawn, grey or brown above, with limited flecks or other Tympanum is distinct and second finger is slightly shorter markings, often as a peppering; a conspicuous narrow than the first. Body length averages 75 mm (subgenus black stripe from the snout, through the nostril to the eye, Ferelitoria subgen. nov.), or: continuing through the tympanum (effectively over-writing 2/ Fawn, grey or brown above, being immaculate; a it) and extending past the forearm with minimal loss of conspicuous black stripe from the snout, through the width along the length to the anterior flank. There is a nostril to the eye, where it is almost entirely broken by a pale glandular stripe from below the eye to the forearm. pale vertical bar in front of the eye as a band continuing There is a conspicuous dark brown stripe along the front below the tympanum and extending back to the forearm edge of the tibial region. The lower lips are faintly or with minimal loss of width along the length, where after a

break, it continues obliquely along the mid flank. There is 1/ Back is without darker markings or mottling, at most a pale glandular stripe from below the eye to the forearm. there being a slight peppering on the upper surfaces; There is a conspicuous dark brown stripe along the front anterior black head stripe when present is interrupted at edge of the tibial region. Lips are not spotted or barred. least in part by a slight vertical bar in front of the eye; The groin and hind side of thighs are pale yellow, spotted posterior dark head stripe is as wide as and overwriting and variegated with dark brown. Venter is whitish. Dorsal the tympanum or not quite so and if not, then the lower surface is smooth and leathery, throat is smooth and the part of the tympanum is not blackened and snout is only venter granular. Vomerine teeth are between the moderately pointed, (Q. axillaris, Q. coplandi, Q. inermis, choanae. Finger discs are small and distinct, while toe Q. pallida, Q. tornieri) (subgenus Quasilitoria subgen. discs are smaller and indistinct. Fingers are unwebbed nov.), or alternatively: and toes have moderate webbing between them. There is 2/ With darker markings and/or stripes on the dorsal a small inner and tiny outer metatarsal tubercle. surface, with some longitudinal folds on the back, a Tympanum is distinct and second finger is slightly shorter strongly pointed snout and a well defined yellow bar in than the first. Body length averages 50 mm (subgenus front of the eye, breaking the black line from snout past Paralitoria subgen. nov.). eye, the yellow bar being blocked at the top by a small Distribution: Subgenus Ferelitoria subgen. nov. is found and well-defined area of black. in the dry tropics of northern Australia, being the top end Forefingers light yellowish or white, peppering on the light of the Northern Territory and Western Australia. surfaces of the snout. Tympanum has a distinctive pale Etymology: Fere in Latin means “Like”, which is accurate rim (Q. nasuta, Q. peninsulae) (subgenus as these frogs are “like” Litoria frogs. Vultusamolitoria subgen. nov.). Content: Paralitoria (Ferelitoria) watjulumensis (Copland, Distribution: Northern third of Australia, extending south 1957) (type species); P. ( Ferelitoria) spaldingi (Hosmer, along the east coast of Australia, to the New South 1964). Wales North coast. PARALITORIA SUBGEN. NOV. Etymology: “Quasi” in Latin means “like” which sums up Type species: Hyla nigrofrenata Günther, 1867. the species in the genus, as they are “like” Litoria. Diagnosis: As identified within the genus diagnosis Content: Quasilitoria inermis (Peters, 1867) (type within Paralitoria gen. nov. within this paper. species); Q. pallida (Davies, Martin and Watson 1983); Distribution: The subgenus Paralitoria subgen. nov. is Q. coplandi (Tyler, 1968); Q. mickpughi sp. nov.; Q. found on the eastern third of Cape York Peninsula, Torres mippughae sp. nov.; Q. nasuta (Gray, 1842); Q. Strait islands and southern New Guinea. peninsulae (De Vis, 1884); Q. tornieri (Neiden, 1923); Q. Content: Paralitoria nigrofrenata (Günther, 1867) axillaris (Doughty, 2011). (monotypic). VULTUSAMOLITORIA SUBGEN. NOV. Etymology: As for the genus. LSIDurn:lsid:zoobank.org:act:26361CC4-C5D6-428C- QUASILITORIA GEN. NOV. A4B6-7BCC8F897A37 LSIDurn:lsid:zoobank.org:act:535D6B63-119D-414C- Type species: Pelodytes nasutus Gray, 1842. 98F7-F83196290D9D Diagnosis: Frogs in the subgenus Vultusamolitoria Type species: Chiroleptes inermis Peters, 1867. subgen. nov. are separated from frogs in the nominate subgenus of Quasilitoria subgen. nov. by the following Diagnosis: The genus Quasilitoria gen. nov. as defined suite of characters: having darker markings and/or stripes herein, diverged from its nearest common ancestor with on the dorsal surface, with some longitudinal folds on the genus Litoria Tschudi, 1838, (as defined in this paper), back, a strongly pointed snout and a well defined yellow being the next most closely related genus, some 13.3 bar in front of the eye, breaking the black line from snout MYA according to Duellman et al. (2016). past eye, the yellow bar being blocked at the top by a Species of Quasilitoria gen. nov. are separated from all small and well-defined area of black. Forefingers light other Australasian tree frogs (Pelodryadidae) by the yellowish or white, peppering on the light surfaces of the following suite of characters: snout. Tympanum has a distinctive pale rim (Q. nasuta, The dorsum is smooth or with some scattered low round Q. peninsulae) (subgenus Vultusamolitoria subgen. nov.). tubercles; front toe discs are no wider than the In turn frogs of the nominate subgenus are separated penultimate phalanx and toe discs only slightly wider, but from Vultusamolitoria subgen. nov. by the following suite otherwise also small. of characters: Back is without darker markings or Front edge of thigh has either a continous black stripe or mottling, at most there being a slight peppering on the alternatively an arrangement of broken black spots or upper surfaces; anterior black head stripe when present blotches. is interrupted at least in part by a slight vertical bar in Lower surfaces white, throat with or without some front of the eye; posterior dark head stripe is as wide as mottling, smooth on the throat and chest and slightly and overwriting the tympanum or not quite so and if not, granular on the belly. then the lower part of the tympanum is not blackened, Dorslolateral folds are either absent or very indistinct. and snout is only moderately pointed, (Q. axillaris, Q. Inner metatarsal tubercle is small and outer one is tiny. coplandi, Q. inermis, Q. pallida, Q. tornieri) (subgenus Prominent vomerine teeth. Quasilitoria subgen. nov.). Fingers unwebbed, while toes are half to three quarters Common to both subgenera and when combined with the webbed. Furthermore, one or other of the following relevant characters above, species in this genus character suites: (Quasilitoria gen. nov.) are separated from all other Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 146 Australasian Journal of Herpetology

Australasian Tree Frogs (Pelodryadidae) by the following from the type form and would are almost certainly worthy additional characters: of recognition as full species, but in the absence of The dorsum is smooth or with some scattered low round molecular data for each form, I have conservatively tubercles; front toe discs are no wider than the named each as subspecies. penultimate phalanx and toe discs only slightly wider, but All three forms would identify as Quasilitoria inermis, otherwise also small. Front edge of thigh has either a identified as “Litoria inermis”, using the key in Cogger continous black stripe or alternatively an arrangement of (2014) as a basis to separate them from all other species broken black spots or blotches. Lower surfaces white, of Australasian tree frog (Pelodryadidae). throat with or without some mottling, smooth on the throat All Q. inermis are further separated from morphologically and chest and slightly granular on the belly. Dorslolateral similar species within Pelodryadidae where they occur by folds are either absent or very indistinct. Inner metatarsal having noticeable warts or tubercles on the dorsum, tubercle is small and outer one is tiny. Prominent poorly defined lateral headstripe and finely reticulated vomerine teeth. Fingers unwebbed, while toes are half to thigh markings. three quarters webbed. Q. inermis dunphyi subsp. nov. restricted to the According to Duellman et al. (2016) this subgenus Kimberley District of Western Australia is readily diverged from the other 9.3 MYA. separated from both other subspecies by having a Distribution: Northern third of Australia, extending south generally dark purplish-brown dorsum, versus grey brown along the east coast of Australia, to the New South in Q. inermis davidtribei subsp. nov. and yellow-brown in Wales North coast. Q. inermis inermis. Etymology: “Vultusamo” in Latin means “looks like” Q. inermis dunphyi subsp. nov. is further separated from which sums up the species in the genus, as they “look both other subspecies by having a limited number of like” Litoria. relatively small warts or tubercles on the dorsum and Content: Quasilitoria (Vultusamolitoria) nasuta (Gray, upper flanks, these being roundish in shape, versus 1842) (type species); Q. (Vultusamolitoria) peninsulae numerous well-defined warts or tubercles, often squarish (De Vis, 1884). or rectangular in shape in both Q. inermis inermis and Q. QUASILITORIA SUBGEN. NOV. inermis davidtribei subsp. nov.. Type species: Chiroleptes inermis Peters, 1867. Adult male Q. inermis dunphyi subsp. nov. have strong Diagnosis: As identified within the genus diagnosis lightening of the upper lip, versus not so in Q. inermis within Paralitoria gen. nov. within this paper. davidtribei subsp. nov. and slight in Q. inermis inermis. Distribution: Northern tropics of Australia, mainly in the Q. inermis davidtribei subsp. nov. from the top end of the Northern Territory and Western Australia. Northern Territory, is separated from both Q. inermis inermis and Q. inermis dunphyi subsp. nov. by having a Etymology: As for the genus. limited amount of black pigment intruding on the upper Content: Quasilitoria (Quasilitoria) inermis (Peters, 1867) tympanum, versus not so in the other two subspecies (type species); Q. (Quasilitoria) pallida (Davies, Martin (none intrudes). and Watson 1983); Q. (Quasilitoria) coplandi (Tyler, Q. inermis davidtribei subsp. nov. has significant dark 1968); Q. mickpughi sp. nov.; Q. mippughae sp. nov.; Q. barring or patches on the dorsum of the hind limbs, as (Quasilitoria) tornieri (Neiden, 1923); Q. (Quasilitoria) well as patches of dark pigment bordering the upper lip, axillaris (Doughty, 2011). which is not seen in the other two subspecies. QUASILITORIA INERMIS DAVIDTRIBEI SUBSP. NOV. Q. inermis inermis in life is depicted in Anstis (2013) on LSIDurn:lsid:zoobank.org:act:FC0A19BD-2B20-405E- page 212 at top right and page 214 at top. A449-D56B6A0C1C68 Q. inermis davidtribei subsp. nov. in life is depicted in Holotype: A preserved specimen at the South Australian Cogger (2014) on page 167 at bottom. Museum, Adelaide, South Australia, Australia, specimen Q. inermis dunphyi subsp. nov. is depicted in Anstis number R23285, collected from the Jabiru Airstrip, (2013) on page 212 at top left. Northern Territory, Australia, Latitude -12.6667 S., Distribution: The subspecies Q. inermis davidtribei Longitude 132.8333 E. subsp. nov. appears to be confined to the top end of the This government-owned facility allows access to its Northern Territory, Australia. holdings. Etymology: The subspecies Q. inermis davidtribei subsp. Paratypes: Preserved specimens at the South Australian nov. is named in honour of David Tribe, program director Museum, Adelaide, South Australia, Australia, specimen at the New South Wales Gould League, a well-known numbers R23286-300, R23343-52, R23312-25, R23352 wildlife conservation organisation. and R23284 all collected from the Jabiru Airstrip, In the 1990’s this organisation had its income stream Northern Territory, Australia, Latitude -12.6667 S., virtually annihilated by the predatory “donation stealing” Longitude 132.8333 E. practices of the Steve Irwin business. Diagnosis: The subspecies Quasilitoria (Quasilitoria) Irwin took the business model of marketing his property inermis davidtribei subsp. nov. and Quasilitoria empire to conservation-minded people to a low never (Quasilitoria) inermis dunphyi subsp. nov. have both until previously seen in Australian history. The net result was now been treated as regional populations of the species that dozens of long-established grass-roots wildlife Quasilitoria (Quasilitoria) inermis (Peters, 1867), with a conservation organisations in Australia were effectively type locality of Rockhampton in coastal Queensland, starved of funds and forced to shut down, with Australia. Both relevant subspecies are highly divergent

enormously negative consequences for wildlife tympanum, versus not so in the other two subspecies. conservation in Australia. Q. inermis davidtribei subsp. nov. has significant dark The Irwin business also militarized State Wildlife barring or patches on the dorsum of the hind limbs, as departments and police forces to raid wildlife well as patches of dark pigment bordering the upper lip, conservation organisations they saw as rivals to their own which is not seen in the other two subspecies. business. Q. inermis inermis in life is depicted in Anstis (2013) on QUASILITORIA INERMIS DUNPHYI SUBSP. NOV. page 212 at top right and page 214 at top. LSIDurn:lsid:zoobank.org:act:A67D73DE-9699-46AA- Q. inermis davidtribei subsp. nov. in life is depicted in ACE5-654618116D3F Cogger (2014) on page 167 at bottom. Holotype: A preserved specimen at the Western Q. inermis dunphyi subsp. nov. is depicted in Anstis Australian Museum, Perth, Western Australia, Australia, (2013) on page 212 at top left. specimen number R81887 collected from Kununurra, Distribution: The subspecies Q. inermis dunphyi subsp. Western Australia, Australia, Latitude -15.7667 S., nov. appears to be confined to the Kimberley District of Longitude 128.7333 S. North-western Australia. This government-owned facility allows access to its Etymology: The subspecies is named in honour of Milo holdings. Dunphy, 1928-1996, the son of Myles Dunphy, in Paratypes: Four preserved specimens at the Western recognition for his political activism to preserve Australian Museum, Perth, Western Australia, Australia, wilderness areas in New South Wales, Australia. specimen numbers R81884, R81885, R81886, R81888, QUASILITORIA (QUASILITORIA) MICKPUGHI SP. NOV. collected from Kununurra, Western Australia, Australia, LSIDurn:lsid:zoobank.org:act:961A988F-9086-4AF2- Latitude -15.7667 S., Longitude 128.7333 S. 9530-F1989D44D356 Diagnosis: The subspecies Quasilitoria (Quasilitoria) Holotype: A preserved specimen at the South Australian inermis dunphyi subsp. nov. and Quasilitoria Museum, Adelaide, South Australia, Australia, specimen (Quasilitoria) inermis davidtribei subsp. nov. have both number R.43642 collected at Jabiluka, Northern Territory, until now been treated as regional populations of the Australia, Latitude -12.52 S., Longitude 132.88 E. species Quasilitoria (Quasilitoria) inermis (Peters, 1867), This government-owned facility allows access to its with a type locality of Rockhampton in coastal holdings. Queensland, Australia. Both relevant subspecies are Paratypes: 1/ A preserved specimen at the Australian highly divergent from the type form and would are almost Museum, Sydney, New South Wales, Australia, specimen certainly worthy of recognition as full species, but in the number R.88556 collected at Jabiluka, Northern Territory, absence of molecular data for each form, I have Australia, Latitude -12.52 S., Longitude 132.88 E. conservatively named each as subspecies. 2/ A preserved specimen at the Northern Territory Art All three forms would identify as Quasilitoria inermis, Gallery and Museum, specimen number R.02503 identified as “Litoria inermis”, using the key in Cogger collected from 6.5 km south west of Oenpelli, Northern (2014) as a basis to separate them from all other species Territory, Australia, Latitude -12.367 S., Longitude of Australasian tree frog (Pelodryadidae). 133.033 E. All Q. inermis are further separated from morphologically Diagnosis: The two species Quasilitoria mickpughi sp. similar species within Pelodryadidae where they occur by nov. and Quasilitoria mippughae sp. nov. were both until having noticeable warts or tubercles on the dorsum, now treated as populations of Q. coplandi (Tyler, 1968), poorly defined lateral headstripe and finely reticulated although at the time Tyler described Q. coplandi he did thigh markings. not inspect any specimens of either of these newly Q. inermis dunphyi subsp. nov. restricted to the named putative species. Kimberley District of Western Australia is readily His relevant sample of frogs was confined to south-west separated from both other subspecies by having a Northern Territory and the nearby Kimberley Division of generally dark purplish-brown dorsum, versus grey brown Western Australia animals (Tyler 1969). in Q. inermis davidtribei subsp. nov. and yellow-brown in All three species would be identified and diagnosed as Q. Q. inermis inermis. coplandi in both Cogger (2014) and Anstis (2013). Q. inermis dunphyi subsp. nov. is further separated from All three species are separated from all other similar both other subspecies by having a limited number of species of tree frog (Pelodryadidae) in northern Australia relatively small warts or tubercles on the dorsum and by the following suite of characters: Webbing reaches the upper flanks, these being roundish in shape, versus base of the terminal disc on the fifth toe. numerous well-defined warts or tubercles, often squarish The finger and toe discs are distinctly dilated and or rectangular in shape in both Q. inermis inermis and Q. approximately twice the width of the penultimate inermis davidtribei subsp. nov.. phlanges. Lateral head stripe is absent. Snount-vent Adult male Q. inermis dunphyi subsp. nov. have strong length is under 45 mm. lightening of the upper lip, versus not so in Q. inermis All of Quasilitoria mickpughi sp. nov., Q. mippughae sp. davidtribei subsp. nov. and slight in Q. inermis inermis. nov. and Q. coplandi can be readily separated from one Q. inermis davidtribei subsp. nov. from the top end of the another on the basis of colouration. Northern Territory, is separated from both Q. inermis Q. coplandi (Tyler, 1968) with a type locality of Inverway inermis and Q. inermis dunphyi subsp. nov. by having a Station, Northern Territory, Australia, part of the Victoria limited amount of black pigment intruding on the upper River district, in the far north-west of the State, near the

Western Australian border, and including all populations QUASILITORIA (QUASILITORIA) MIPPUGHAE SP. from this general area and west into the Kimberley NOV. District of Western Australia, is a light fawn to beige LSIDurn:lsid:zoobank.org:act:5AB69DE5-107A-46E1- colour on top without obvious or distinct markings. There 8A55-3C57580F1089 are blotches, flecks or peppering on the back, but all are Holotype: A preserved specimen at the Museum and Art faded and generally merge with the overall dorsal Gallery of the Northern Territory, Darwin, Northern colouration. Territory, Australia, specimen number R25754 collected By contrast Q. mickpughi sp. nov. from the top end of the at Chinaman’s Waterhole, 35 km east of Mount Isa, Northern Territory, with a centre of distribution on the Queensland, Australia, Latitude -20.725 S., Longitude Arnhemland Escarpment and including areas south-east 139.779 E. of there including the southern shores of the Gulf of This government-owned facility allows access to its Carpentaria, has a well-defined dorsal colouration holdings. consisting of a light yellow to pinkish orange-background Paratypes: Preserved specimens at the Museum and Art overlaid with numerous dark purple blotches and marks Gallery of the Northern Territory, Darwin, Northern occupying about half the surface area, giving the frog a Territory, Australia, specimen numbers R25755, R25756, distinctly darker colour. The contrast between dark and R21713 and R21714 and specimens at the South light remains intense on the snout, versus only indistinct Australian Museum, Adelaide, South Australia, Australia, markings on Q. coplandi. specimen numbers R54356 and R54358, all collected at The species Q. mippughae sp. nov. with a centre of Chinaman’s Waterhole, 35 km east of Mount Isa, distribution on the Selwyn Ranges of North-west Queensland, Australia, Latitude -20.725 S., Longitude Queensland and nearby hilly aras, extending as far north 139.779 E. as about Lawn Hill (still in Queensland), is characterised Diagnosis: The two species Quasilitoria mippughae sp. by having a yellow to beige coloured dorsum and well nov. and Quasilitoria mickpughi sp. nov. were both until scattered irregular, but well defined blackish spots and now treated as populations of Q. coplandi (Tyler, 1968), blotches, with the region anterior to the eyes having although at the time Tyler described Q. coplandi he did either no darker pigment or only a very limited amount (in not inspect any specimens of either of these newly contrast to both other species). named putative species. His relevant sample of frogs All three species are generally light in a well-defined zone was confined to south-west Northern Territory and between the eyes, but have darker pigment both anterior Kimberley Division of Western Australia animals (Tyler and posterior to this, except in the case of Q. mippughae 1969). sp. nov. where there is generally no darker pigment anterior, or very little. All three species would be identified and diagnosed as Q. coplandi in both Cogger (2014) and Anstis (2013). Anstis (2013) reports on differences in the jaw structure All three species are separated from all other similar of tadpoles within the species herein defined as Q. species of tree frog (Pelodryadidae) in northern Australia mickpughi sp. nov. and Q. coplandi. by the following suite of characters: Webbing reaches the Q. coplandi in life is depicted in Anstis (2013) on page base of the terminal disc on the fifth toe. The finger and 175 at top right. toe discs are distinctly dilated and approximately twice Q. mickpughi sp. nov. in life is depicted in Anstis (2013) the width of the penultimate phlanges. Lateral head stripe on page 177 at top right and as a tadpole below that. is absent. Snount-vent length is under 45 mm. The specimen depicted on page 159 of Cogger (2014) All of Quasilitoria mickpughi sp. nov., Q. mippughae sp. top right, is also referred to this taxon. nov. and Q. coplandi can be readily separated from one Q. mippughae sp. nov. in life is depicted on page 32 of another on the basis of colouration. Vanderduys (2012) in both photos. Q. coplandi (Tyler, 1968) with a type locality of Inverway Numerous photos of all three species can be found Station, Northern Territory, Australia, part of the Victoria online at: River district, in the far north-west of the State, near the http://www.flickr.com Western Australian border, and including all populations when doing a search for “Litoria coplandi”. from this general area and west into the Kimberley Distribution: Q. mickpughi sp. nov. is believed to be District of Western Australia, is a light fawn to beige restricted to the Arnhemland region of the top-end of the colour on top without obvious or distinct markings. There Northern Territory, although specimens from the east of are blotches, flecks or peppering on the back, but all are the Northern Territory, including those south of the Gulf of faded and generally merge with the overall dorsal Carpentaria are also tentatively referred to this taxon on colouration. the basis of obvious morphological similarity. By contrast Q. mickpughi sp. nov. from the top end of the Etymology: Named in honour of Mick Pugh of Geelong, Northern Territory, with a centre of distribution on the Victoria, Australia, a former president of the Victorian Arnhemland Escarpment and including areas south-east Association of Amateur Herpetologists Incorporated, in of there including the southern shores of the Gulf of recognition of his many decades of service to Carpentaria, has a well-defined dorsal colouration herpetology and wildlife conservation in general. consisting of a light yellow to pinkish orange-background Mick Pugh was hounded out of herpetology by a corrupt overlaid with numerous dark purple blotches and marks and dysfunctional State wildlife department that saw his occupying about half the surface area, giving the frog a expertise and reputation as a “threat” to their own lesser distinctly darker colour. The contrast between dark and “experts” employed at their business “Zoos Victoria”. light remains intense on the snout, versus only indistinct

markings on Q. coplandi. Museum, Perth, Western Australia, Australia, specimen The species Q. mippughae sp. nov. with a centre of number R115931, collected from about 14 km north distribution on the Selwyn Ranges of North-west north-west of the junction of the Calder River and Queensland and nearby hilly aras, extending as far north Bachsten Creek, West Kimberley District, Western as about Lawn Hill (still in Queensland), is characterised Australia, Australia, Latitude-15.9833 S., Longitude by having a yellow to beige coloured dorsum and well 125.3167 E. scattered irregular, but well defined blackish spots and Diagnosis: Until now the subspecies Quasilitoria tornieri blotches, with the region anterior to the eyes having serventyi subsp. nov. has been treated as nominate either no darker pigment or only a very limited amount (in Quasilitoria tornieri tornieri (Nieden, 1923) as defined and contrast to both other species). separated from other relevant species by Cogger (2014). All three species are generally light in a well-defined zone Both subspecies are separated from all other congeners between the eyes, but have darker pigment both anterior by the fact the dorsum is usually smooth, the lateral head and posterior to this, except in the case of Q. mippughae stripe is well defined, particularly before the eye, thigh sp. nov. where there is generally no darker pigment markings are strongly reticulated and the stripe along the anterior, or very little. edge of tarsus is uninterrupted. Anstis (2013) reports on differences in the jaw structure However Q. tornieri serventyi subsp. nov. is readily of tadpoles within the species herein defined as Q. separated from that taxon (type locality of Port Essington, mickpughi sp. nov. and Q. coplandi. NT), by the following suite of characters: Q. coplandi in life is depicted in Anstis (2013) on page Breeding adult males have a slight yellowish tinge on the 175 at top right. whitish lips and groin, versus strong deep yellow in Q. Q. mickpughi sp. nov. in life is depicted in Anstis (2013) tornieri tornieri. In both sexes of adult Q. tornieri serventyi on page 177 at top right and as a tadpole below that. subsp. nov. the upper hind limb are strongly peppered The specimen depicted on page 159 of Cogger (2014) with grey or with faint darker markings or blotches, versus top right, is also referred to this taxon. not so in Q. tornieri tornieri. Q. mippughae sp. nov. in life is depicted on page 32 of The (hind) toe pads of Q. tornieri tornieri are slightly Vanderduys (2012) in both photos. larger than the lateral edges of the penultimate phlanges, Numerous photos of all three species can be found versus not so in Q. tornieri serventyi subsp. nov.. online at: The tadpoles of both subspecies also differ. http://www.flickr.com Metamorphosing tadpoles of Q. tornieri tornieri have a when doing a search for “Litoria coplandi”. large amount of blackish pigment in blotches on the outer Distribution: Q. mippughae sp. nov. is believed to be edges and inner areas of the fins, versus only small restricted to the Selywyn and nearby ranges in north-west amounts on the edges and inner areas in Q. tornieri Queensland, being a distribution centred on the town of serventyi subsp. nov.. Mount Isa, Queelsland, Australia. Images of both Q. tornieri serventyi subsp. nov. and Q. Etymology: Named in honour of Mip Pugh of Geelong, tornieri tornieri in life and their tadpoles can be seen on Victoria, Australia, in recognition of her services to pages 321-324 of Anstis 2013. herpetology in Australia, especially with regards to the Distribution: Q. tornieri serventyi subsp. nov. is found in captive breeding of Australian agamid lizards and the Kimberley District of Western Australia, extending to educating many others in terms of her expertise the Daly River region of the Northern Territory. Q. tornieri developed over many years. tornieri is found throughout the rest of the range of the species, this being the top end of the Northern Territory QUASILITORIA TORNIERI SERVENTYI SUBSP. NOV. and the Gulf of Carpentaria into far north-west LSIDurn:lsid:zoobank.org:act:361FEC86-D280-46F6- Queensland. B781-77F012216C06 Etymology: The subspecies is named in honour of Holotype: A preserved specimen at the Western Vincent Serventy, born 6 January 1916, and died 8 Australian Museum, Perth, Western Australia, Australia, September 2007, who authored many popular books specimen number R50671 collected from Drysdale River about Australian wildlife and who was regarded as an National Park, Kimberley District, Western Australia, avid conservationist in his active years. He lived in Australia, Latitude -14.73 S., Longitude 126.93 E. Western Australia and New South Wales. This facility allows access to its holdings. SAGUNURINI TRIBE NOV. Paratypes: 1/ A preserved specimen at the Western SAGANURA WELLS AND WELLINGTON, 1985 Australian Museum, Perth, Western Australia, Australia, Type species: Hyla burrowsae Scott, 1942. specimen number R50670 collected from Drysdale River National Park, Kimberley District, Western Australia, Diagnosis: Living frogs in the genus Saganura Wells Australia, Latitude -14.73 S., Longitude 126.93 E. and Wellington, 1985 are all readily separated from all other Australasian Tree Frogs (Pelodryadidae) by the 2/ A preserved male specimen at the Western Australian following unique suite of characters: It is a uniform light Museum, Perth, Western Australia, Australia, specimen green above, commonly with scattered light fawn spots, number R99065 collected from 6 km north of Lake or dark brown with irregular bright green patches and Gilbert, near Beverley Springs, West Kimberley District, light brown flecks. Western Australia, Australia (in leaf litter around rocks), Latitude -16.5042 S., Longitude 125.275 E. There is a narrow, black canthal streak, widening behind the eye to form a dark broad band extending almost to 3/ A preserved specimen at the Western Australian the groin; this dark band often broken up by groups of Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 150 Australasian Journal of Herpetology irregular white blotches and/or spots. There is an from relatively few specimens. obscure, narrow light green zone along the Etymology: The frogs in this genus are absolutely supratympanic ridge. There is no pale line along the spectacular. They are big, bold and bright and posterior edge of the upper jaw. Venter is pinkish-white, herpetologists who set eyes on them invariably exclaim throat darker with a greyish tinge. Groin and hind side of “Wow!”. Hence the simple and easy to remember genus thighs is a uniform pale brown. Dorsal surface smooth or name “Wowrana”. with scattered small tubercles. Chin smooth and other Content: Wowrana dux (Richards and Oliver, 2006) (type lower surfaces are granular. Vomerine teeth are between species); W. graminea (Boulenger, 1905); W. hunti or behind the choanae, with hind edge of vomerine teeth (Richards, Oliver, Dahl and Tjaturadi. 2006); W. always further back than the choanae. Prectoral fold pallidofemora (Kraus, 2018); W. pterodactyla (Oliver, present. Finger and toe discs large. Fingers with Richards and Donnellan, 2018); W. nullicedens (Kraus, conspicuous basal webbing, being nearly a third webbed, 2018); W. sauroni (Richards and Oliver, 2006). toes are about three quarters webbed. There is a large PARAWOWRANA SUBGEN. NOV. inner and small outer metatarsal tubercle. Tympanum is LSIDurn:lsid:zoobank.org:act:D130450F-D9D5-471B- distinct. Second finger is much longer than first; when B7A1-212F2774E134 pressed together, the tip of the first finger reaches no further than the base of the disc of the second finger. Type species: Litoria hunti Richards, Oliver, Dahl, and Heel of adpressed hindlimb reaches to the eye or Tjaturadi, 2006. beyond. Adults attain about 55 mm in total length Diagnosis: Frogs in the genus Wowrana gen. nov. in the (adapted and modified from Cogger 2014). nominate subgenus are readily separated from all other According to Duellman et al. (2016), the species in this Australasian (Australian and New Guinea) Tree Frogs genus diverged from their nearest living relatives 27.6 (Pelodryadidae) by the following unique suite of MYA. characters: Distribution: South-west Tasmania, including highlands Large size (adult males over 60 mm); uniform bright and coast. green dorsal colouration in life (blue in preservative). Fully webbed hands. No prominent and enlarged parotoid Content: Sagunura burrowsae (Scott, 1942) (monotypic). glands, no white labial stripe, SVL not over 85 mm and a WOWRANAINI TRIBE NOV. call consisting of a single, relatively short (0.206-0.379 WOWRANA GEN. NOV. second) grunt with an unusual clumped pattern of pulses. LSIDurn:lsid:zoobank.org:act:9F436614-24BC-49DE- The single species within the subgenus Parawowrana 8AFB-FDFB8D8B5671 subgen. nov. is defined as above, except that it instead is Type species: Litoria dux Richards and Oliver, 2006. slightly smaller than the preceding species (adult males Diagnosis: Frogs in the genus Wowrana gen. nov. in the 50-55 mm), has a white labial stripe (in common with the nominate subgenus are readily separated from all other much larger species in Sandyrana Wells and Wellington, Australasian (Australian and New Guinea) Tree Frogs 1985, that grow to in excess of 100 mm in body length) (Pelodryadidae) by the following unique suite of and has a characteristically longer call than those in the characters: nominate subgenus (0.69-0.9 second) (Richards et al. Large size (adult males over 60 mm); uniform bright 2006). green dorsal colouration in life (blue in preservative). Duellman et al. (2016) found that the species in the Fully webbed hands. No prominent and enlarged parotoid genus Wowrana gen. nov. diverged from their nearest glands, no white labial stripe, SVL not over 85 mm and a living relatives 26.5 MYA, being species within Sandyrana call consisting of a single, relatively short (0.206-0.379 Wells and Wellington, 1985. The type form of second) grunt with an unusual clumped pattern of pulses. Nyctimystes Stejneger, 1916, namely Nyctimantis papua The single species within the subgenus Parawowrana Boulenger, 1897 diverged from this genus 36.1 MYA, subgen. nov. is defined as above, except that it instead is making genus level assignment of Wowrana gen. nov. an slightly smaller than the preceding species (adult males obvious choice. 50-55 mm), has a white labial stripe (in common with the Distribution: The sole species currently placed within much larger species in Sandyrana Wells and Wellington, the subgenus Parawowrana subgen. nov. is known only 1985, that grow to in excess of 100 mm in body length) from the type locality (Utai, Sanduan Province, northern and has a characteristically longer call than those in the Papua New Guinea). nominate subgenus (0.69-0.9 second) (Richards et al. Etymology: “Para” in Latin, means “not quite” or “almost” 2006). and as this frog is not quite the same as nominate Duellman et al. (2016) found that the species in the Wowrana gen. nov. the subgenus name makes sense. genus Wowrana gen. nov. diverged from their nearest Content: Wowrana (Parawowrana) hunti (Richards, living relatives 26.5 MYA, being species within Sandyrana Oliver, Dahl and Tjaturadi. 2006) (monotypic). Wells and Wellington, 1985. The type form of SANDYRANINA SUBTRIBE NOV. Nyctimystes Stejneger, 1916, namely Nyctimantis papua SANDYRANA WELLS AND WELLINGTON, 1985 Boulenger, 1897 diverged from this genus 36.1 MYA, Type species: Hyla infrafrenata Günther, 1867. making genus level assignment of Wowrana gen. nov. an obvious choice. Diagnosis: Frogs in the genus Sandyrana Wells and Wellington, 1985 are readily separated from all other Distribution: All species within Wowrana gen. nov. are Australasian Tree Frogs (Pelodryadidae) by the following confined to forested locations on the island of New unique suite of characters: Large frogs, which as adults Guinea and in general most species are currently known Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 151

can grow to in excess of 100 mm. Dorsal colour of adults flanks greyish-white, often with brown under the throat. in life is usually uniform green above in normal Axilla and groin are flesh coloured. Hind side of thighs is circumstances, but may otherwise range from fawn, mottled with pale and dark brown. Skin is smooth to through green, purplish or even blue. Fingers with leathery above, and granular on the venter. Vomerine conspicuous webbing, reaching at least as far as the teeth are present and between the choanae. There is no base of the penultimate phalanx of the fourth finger. Hind pectoral fold. edge of forearm is smooth, or with at most a few low, There is no enlarged tubercle or crenulated ridge along discontinuous tubercles. Hind edge of feet is smooth. the hind edge of the forearm; Finger and toe discs are Hind side of thighs is more or less uniform, without black large. Fingers lack webbing and toes are nearly and yellow marbling, spots or blotches. Webbing completely webbed, with webbing reaching the base of reaching no further than the base of the penultimate the penultimate phalanx of the fourth toe. phalanx of the fourth finger. A distinctive white or pink There is a prominent inner metatarsal tubercle and no stripe along the edge of the lower jaw, extending back to outer one. Heel of adpressed hind limb goes well beyond the level of the forelimb and not in a configuration of the eye. Tympanum is large and obvious, with a well- blobs, spots or random markings otherwise tending to be developed supratympanic fold. Above this, there is a in a linear manner. Vomerine teeth present. semidistinct beige coloured stripe, extending to form Duellman et al. (2016) found that the species within the broken blotches along a line running along the margin of genus Sandyrana diverged from their nearest living the upper flank. This is not however in the form of a relatives 26.5 MYA. distinct, well marked whitish stripe. Distribution: Throughout New Guinea and west as far as There is also no pale line along the posterior edge of the Halmahera Island as well as New Britain, New Ireland, upper jaw, although this region of the head is usually a Aru Islands and north-east Queensland, Australia. light bluey, purplish grey in colour. Second finger is much Content: Sandyrana infrafrenata (Günther, 1867) (type longer than the first, the tip of the first finger goes no species); S. militarius (Ramsay, 1878); S. multicolor further than the base of the disc of the second finger (Günther, 2004); S. purpureolatus (Oliver, Richards, when they are pressed togeather. Tjaturadi, and Iskandar, 2007); S. sanguinolenta (Van Males get to 80 mm in body length and females 100 mm. Kampen, 1909). The genus Euscelis Fitzinger, 1843 is separated from all other Australasian Tree Frogs (Pelodryadidae) by the TRIBE AND SUBTRIBE DESCRIPTIONS following unique suite of characters: Colouration is a pale fawn to dark brown above, usually immaculate, or with ADELYNHOSERHYLEINI TRIBE NOV. darker markings, ranging from flecks to blotches and LSIDurn:lsid:zoobank.org:act:D4FD8FB3-7C85-423F- including one that forms a transverse bar between the eyes; the tympanum has a pale rim; dorsal surface 8ACB-3017A21AAE90 generally smooth or sometimes leathery, with one Type genus: Adelynhoserhylea gen. nov. species having slight warts; webbing may reach the disc Diagnosis: The tribe Adelynhoserhyleini tribe nov. is best of the fifth toe, but usually not, and generally extending defined and diagnosed by defining the four component no more than halfway along the penultimate phalanx; genera separately. discs on fingers and toes are small and inconspicuous Species of tree frogs within the genus Adelynhoserhylea and barely wider than digits; fingers unwebbed; second gen. nov. are readily separated from all other finger slightly longer than first; anterior head stripe is Australasian Tree Frogs (Pelodryadidae) by the following present, usually narrow but always continuous, but unique suite of characters: Vomerine teeth present. sometimes ill-defined, not interrupted by a vertical bar in Fingers with conspicuous webbing, reaching at least as front of the eye; posterior head stripe is narrow, no more far as the base of the penultimate phalanx of the fourth than half as wide as and not enclosing the tympanum; finger. A conspicuous serrated ridge along the forearm there is a moderate inner metatarsal tubercle and a small and another along the hind edge of the foot. outer metatarsal tubercle is present; vomerine teeth The morphologically similar genus Jackyhoserhylea gen. present; groin is yellow and heavily blotched with black. nov. with species confined to the New Guinea subregion Whitish ventrally with granular skin. No dorsolateral skin is readily separated from Adelynhoserhylea gen. nov. as fold. defined above, by having dermal fringes on limbs poorly Distribution: Most species are confined to New Guinea defined, fingers only part-webbed and they do not not and Cape York, Australia, but one genus Euscelis have a well-developed crenulated fold along the outer Fitzinger, 1843 occurs in the wetter parts of the east edge of the fore and hind-limbs, this always being either coast of south-east Australia, extending, from Victoria, reduced or absent. through New South Wales and south east Queensland The monotypic species compring the entirety of the and then to the wet tropics of north-east Queensland. genus Leucodigirana gen. nov. is readily separated from Content: Adelynhoserhylea gen. nov. (type genus); all other Australasian Tree Frogs (Pelodryadidae) by the Euscelis Fitzinger, 1843; Jackyhoserhylea gen. nov.; following unique suite of characters: Brown to golden Leucodigirana gen. nov.. brown across most of the dorsum, the body and limbs with numerous scattered dark or light brown spots and blotches, among which are usually smaller pale brown or cream spots and blotches. Venter is white and lower

LEUCODIGIRANINA SUBTRIBE NOV. Distribution: All species are confined to Australia or LSIDurn:lsid:zoobank.org:act:41D58607-4C02-45E2- potentially southern New Guinea with the genus 897E-9D87851B9004 occurring in the wetter parts of the east coast of south- Type genus: Leucodigirana gen. nov. east Australia, extending, from Victoria, through New Diagnosis: The subtribe Leucodigiranina subtribe nov. is South Wales and south east Queensland and then to the best defined and diagnosed by defining the two wet tropics of north-east Queensland. component genera separately. Content: Leucodigirana gen. nov. (type genus); Euscelis The monotypic species compring the entirety of the Fitzinger, 1843. genus Leucodigirana gen. nov. is readily separated from COGGERDONIANI TRIBE NOV. all other Australasian Tree Frogs (Pelodryadidae) by the LSIDurn:lsid:zoobank.org:act:342B883B-E1C6-4D69- following unique suite of characters: Brown to golden 8019-3096C62A9FEE brown across most of the dorsum, the body and limbs Type genus: Coggerdonia Wells and Wellington, 1985. with numerous scattered dark or light brown spots and Diagnosis: The tribe Coggerdoniaini tribe nov. is blotches, among which are usually smaller pale brown or monotypic for the genus Coggerdonia Wells and cream spots and blotches. Venter is white and lower Wellington, 1985. Living frogs in the genus Coggerdonia flanks greyish-white, often with brown under the throat. Wells and Wellington, 1985 are all readily separated from Axilla and groin are flesh coloured. Hind side of thighs is all other Australasian Tree Frogs (Pelodryadidae) by the mottled with pale and dark brown. Skin is smooth to following unique suite of characters: A slender frog being leathery above, and granular on the venter. Vomerine light brown, fawn or light green above, with dark brown teeth are present and between the choanae. There is no patches or flecks arranged in distinct longitudinal lines. pectoral fold. There is no enlarged tubercle or crenulated There is a dark brown to black stripe running from the tip ridge along the hind edge of the forearm; Finger and toe of the snout below the canthus to the eye, somewhat discs are large. Fingers lack webbing and toes are nearly triangular in shape as it widens towards the eye and then completely webbed, with webbing reaching the base of remaining broad as it extends across the tympanum and the penultimate phalanx of the fourth toe. There is a beyond along the flank to the rear of the body. prominent inner metatarsal tubercle and no outer one. This dark coloured stripe is bordered on the lower edge Heel of adpressed hind limb goes well beyond the eye. with a well-defined line of white, also running from the Tympanum is large and obvious, with a well-developed upper lip. Sometimes this stripe will break up into a supratympanic fold. Above this, there is a semidistinct series of adjacent spots. The hind parts of the thighs are beige coloured stripe, extending to form broken blotches dark brown with orange or reddish spots. Belly is whitish along a line running along the margin of the upper flank. to light brown. Skin is smooth above and coarsely This is not however in the form of a distinct, well marked granular below, except under the throat, which is also whitish stripe. There is also no pale line along the smooth. Vomerine teeth are prominent between and posterior edge of the upper jaw, although this region of behind the choanae. There is a distinct pectoral fold. the head is usually a light bluey, purplish grey in colour. Finger and toe discs are small and not much wider than Second finger is much longer than the first, the tip of the the digits. Fingers have basal webbing only and toes are first finger goes no further than the base of the disc of the about three quarters webbed. Inner metatarsal tubercle is second finger when they are pressed togeather. Males large and there is no outer one. Tympanum is large and get to 80 mm in body length and females 100 mm. distinct and the second finger is longer than the first. The genus Euscelis Fitzinger, 1843 is separated from all Adult size 50 60 mm (derived from Cogger, 2014). other Australasian Tree Frogs (Pelodryadidae) by the According to Duellman et al. (2016), the single living following unique suite of characters: Colouration is a pale member of this genus diverged from its nearest living fawn to dark brown above, usually immaculate, or with relatives 30.8 MYA, supporting tribe level classification for darker markings, ranging from flecks to blotches and this (currently) monotypic genus. including one that forms a transverse bar between the Distribution: Wetter parts of south-western Western eyes; the tympanum has a pale rim; dorsal surface Australia, Australia. generally smooth or sometimes leathery, with one Content: Coggerdonia Wells and Wellington, 1985 species having slight warts; webbing may reach the disc (monotypic). of the fifth toe, but usually not, and generally extending CYCLORANININI TRIBE NOV. no more than halfway along the penultimate phalanx; discs on fingers and toes are small and inconspicuous LSIDurn:lsid:zoobank.org:act:2A17B806-9DFA-4BC0- and barely wider than digits; fingers unwebbed; second B4BA-7E2231EC4756 finger slightly longer than first; anterior head stripe is Type genus: Cyclorana Steindachner, 1867. present, usually narrow but always continuous, but Diagnosis: The tribe Cycloraninini tribe nov. is best sometimes ill-defined, not interrupted by a vertical bar in defined by separately diagnosing each of the eight front of the eye; posterior head stripe is narrow, no more component genera. than half as wide as and not enclosing the tympanum; Species within the genera Cyclorana, Mitrolysis and there is a moderate inner metatarsal tubercle and a small Neophractops are all readily separated from all other outer metatarsal tubercle is present; vomerine teeth Australasian Tree Frogs (Pelodryadidae) by the following present; groin is yellow and heavily blotched with black. unique suite of characters: Whistish ventrally with granular skin. No dorsolateral skin They are heavily built, often rotund, frogs; Inner fold. metatarsal tubercle is shovel-shaped; no fronto-parietal

foramen in adults; vomerine teeth are present and largely All species within Ranoidea, Chirodryas and Crottyanura between the choanae; tongue large and oval-shaped; gen. nov. as defined herein, while morphologically similar, Pupil horizontal; tympanum distinct in all but one species are sufficiently divergent from one another to warrant group (Invisibiliaauris subgen. nov.); phlanges are simple beling treated as separate genera. and tips with tiny discs or none at all; first finger opposed All species within the genera Ranoidea, Chirodryas and to remainder and toes are webbed. Crottyanura gen. nov. are readily separated from all other The genus Cyclorana is readily separated from the Australasian Tree Frogs (Pelodryadidae) by the following genera Mitrolysis and Neophractops by having a marked, suite of characters: straight dorsolateral skin fold and a very stout build; toes Vomerine teeth are present; fingers are free or webbed one third webbed; hind side of thighs lacking white spots. only at the base; first finger is longer than, equal to or The genus Neophractops is readily separated from the only slightly shorter than the second finger; no outer genera Mitrolysis and Cyclorana by having no definite metatarsal tubercle. straight dorsolateral skin fold and toes that are three The genera Ranoidea and Chirodryas are separated from quarters webbed. the genus Crottyanura gen. nov. by having flanks that are The genus Mitrolysis is separated from the genera strongly granular below a distinct, glandular dorso-lateral Cyclorana and Neophractops by one or other of: skin fold with densely packed, large, rounded, usually 1/ Having no definite straight dorsolateral skin fold and pale coloured granules, contrasting with the smooth toes that are less than half webbed, or: tubercular skin on the back above the skin fold. 2/ Having a marked, straight dorsolateral skin fold and a In contrast Crottyanura gen. nov. has flanks that are very slender build; toes half webbed; hind side of thighs smooth or with a few scattered granules, not contrasting has numerous white spots. with the skin on the back. The subgenus Paramitrolysis subgen. nov. with the type Ranoidea species are readily separated from species species of Cyclorana verrucosa Tyler and Martin, 1977 is within the genus Chirodryas by having either a smooth readily separated from the nominate subgenus (of dorsum with few if any rounded warts or tubercles, or if Mitrolysis) by having the following unique combination of present in any number or size, are arranged in regular characters: A blunt snout with nostril distinctly nearer to longitudinal rows near the vertebral line. the tip than to the eye; a blackish stripe on the side of the By contrast Chirodryas species have large warts on the head from the snout through the eye and distinct back that are irregularly scattered and not configured in tympanum to the forelimb; a dorsum with numerous skin regular longitudinal rows. folds or large tubercles, with either: 1/ Many being white- The two subgenera within Ranoidea are separated as tipped, highlighted by being surrounded by dark blackish follows: Subgenus Ranoidea has a generally smooth pigment as well as a dorsal colouration dominated by dorsum with at most a few scattered, low tubercles. By brilliant dark lime green or large patches of brilliant dark contrast subgenus Sandgroperanura subgen. nov. has lime green, or 2/ A dorsum with numerous skin folds or obvious flat warts on the back that are arranged in large tubercles not marked in any way and a dorsal regular longitudinal rows near the vertebral line. colouration of beige, overlain with scattered and faded The build of Crottyanura gen. nov. while solid, is slender light olive green patches or blotches in irregular fashion. when compared to both Ranoidea and Chirodryas. The The subgenus Invisibiliaauris subgen. nov. with the type species in Crottyanura gen. nov. are further diagnosed by species Cyclorana cryptotis Tyler and Martin, 1977 is being olive green with brownish tinge above and a readily separated from the other two subgenera by distinctive light mid-vertebral line (in C dahlii but not often having a hidden ear, being covered by skin, in stark in C. crottyi sp. nov.); having the hind side of the thighs contrast to the other subgenera which have an obvious mottled, marbled or spotted with white (in C dahlii but not and exposed tympanum. in C. crotty sp. nov.); a finely granular dorsum; smooth The genus Crottyanura gen. nov. consists of two species, white-coloured venter; no dorsolateral skin fold; fully namely C. dahlii (Boulenger, 1896), with a type locality of webbed toes; no outer metatarsal tubercle and inner Daly River, Northern Territory and found in the general metatarsal tubercle is not shovel-shaped. region of the type locality, being the western half of the All species within the genera Ranoidea, Chirodryas and top end of the Northern Territory and into immediately Crottyanura gen. nov. are readily separated from all other adjacent north-west Western Australia, as well as the Australasian Tree Frogs (Pelodryadidae) by the following newly named species Crottyanurua crottyi sp. nov. from suite of characters: the eastern side of the Gulf of Carpentaria and the Vomerine teeth are present; fingers are free or webbed western side of Cape York in Queensland. only at the base; first finger is longer than, equal to or Both species are readily separated from other only slightly shorter than the second finger; no outer Australasian Tree Frogs (Pelodryadidae) by the following metatarsal tubercle. diagnosis. The genera Ranoidea and Chirodryas are separated from Ranoidea Tschudi, 1838 as defined herein includes only the genus Crottyanura gen. nov. by having flanks that are the Bell Frog group of species, excluding the species strongly granular below a distinct, glandular dorso-lateral associated with the Chirodryas raniformis Keferstein, skin fold with densely packed, large, rounded, usually 1867 species group, herein placed in the genus pale coloured granules, contrasting with the smooth Chirodryas and the tropical species Chiroleptes dahlii tubercular skin on the back above the skin fold. In Boulenger, 1896 herein placed in the new genus contrast Crottyanura gen. nov. has flanks that are smooth Crottyanura gen. nov.. or with a few scattered granules, not contrasting with the Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 154 Australasian Journal of Herpetology skin on the back. a fine, continuous reticulum over the back, head and Ranoidea species are readily separated from species limbs. The flanks have a bluish metal sheen. White within the genus Chirodryas by having either a smooth below, commonly with brown on the throat; axilla and dorsum with few if any rounded warts or tubercles, or if groin flesh coloured; hind side of thighs are dark brown. present in any number or size, are arranged in regular Skin leathery, finely granular, or with numerous small longitudinal rows near the vertebral line. scattered warts. Granular below (M. nannotis (Andersson, By contrast Chirodryas species have large warts on the 1916), M. cottoni sp. nov. M. lorica (Davies and back that are irregularly scattered and not configured in McDonald, 1979)) (nominate subgenus), or: regular longitudinal rows. 2/ Vomerine teeth conspicuous in two rows on a line with The two subgenera within Ranoidea are separated as the hind edge of the choanae; with or without pectoral follows: Subgenus Ranoidea has a generally smooth fold; fingers with conspicuous webbing, reaching at least dorsum with at most a few scattered, low tubercles. as far as the base of the penultimate phalanx of the By contrast subgenus Sandgroperanura subgen. nov. fourth finger; hind edge of forearm is smooth or with at has obvious flat warts on the back that are arranged in most a few, low, discontinuous tubercles; hind edge of regular longitudinal rows near the vertebral line. foot is smooth; hind side of thighs more or less uniform and without black and yellow markings or marbling; Gedyerana gen. nov. is readily separated from all other colour brown to blackish above, never green, with or Australasian Tree Frogs (Pelodryadidae) by being the without a broad, darker vertebral patch and other only Australasian Tree Frog species having a lower contrasting markings; a dark dorsal patch, if present, is eyelid with a characteristic reticulatum or palpebral inconspicuous and commencing from a line joining the venation of fine, pigmented lines and a horizontally centre of each eye; tympanum covered by skin (adult size elliptical pupil. is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. All other species within this region with a lower eyelid nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola with a characteristic reticulatum or palpebral venation of (Liem, 1974)) (subgenus Amnisrana subgen. nov.). fine, pigmented lines have a vertically elliptical pupil, Distribution: Most parts of continental Australia including including species within Nyctimystes Stejneger, 1916 the tropics and the arid zone. sensu lato, as defined by other authors including Cogger (2014) at page 197 under the heading “genus Content: Cyclorana Steindachner, 1867 (type genus); Nyctimystes Stejneger, 1916”. Chirodryas Keferstein, 1867; Crottyanura gen. nov.; Gedyerana gen. nov.; Mitrolysis Cope, 1889; Mosleyia Both species within the genus Gedyerana gen. nov. Wells and Wellington, 1985; Neophractops Wells and would be diagnosed as either “Litoria dayi” in Anstis Wellington, 1985; Ranoidea Tschudi, 1838. (2013) or “Nyctimystes dayi” in Cogger (2014). RANOIDEINA SUBTRIBE NOV. These frogs have large protruding eyes, with a dark iris, a broad head and a slender body. LSIDurn:lsid:zoobank.org:act:2CEAF103-5F6F-4B55- 993B-7026C0498838 Males get to about 45 mm in body length, but females are considerably larger and get to about 60 mm in length. Type genus: Ranoidea Tschudi, 1838. Their tadpoles have large sucker mouths. Diagnosis: The subtribe Ranoideina subtribe nov. Dorsal colour ranges from grey, brown, yellow, orange or (comprising the two genera Ranoidea Tschudi, 1838 red, with or without mottling, spots or flecks and with or (type genus) and Chirodryas Keferstein, 1867) is best without discrete white or cream blotches or ocelli on the defined by disgnosing and defining the two component upper surfaces. genera. Ranoidea Tschudi, 1838 as defined herein includes only the Bell Frog group of species, excluding Snout is moderately rounded or acuminate (tending to be the species associated with the Chirodryas raniformis pointed) and the genus occurs in rainforest stream Keferstein, 1867 species group, herein placed in the habitats. genus Chirodryas and the tropical species Chiroleptes The genus Mosleyia is readily separated from all other dahlii Boulenger, 1896 herein placed in the new genus Australasian Tree Frogs (Pelodryadidae) by one or other Crottyanura gen. nov.. of the two unique suites of characters: All species within Ranoidea, Chirodryas and Crottyanura 1/ Vomerine teeth prominent and largely between the gen. nov. as defined herein, while morphologically similar, choanae. No pectoral fold; A series of small enlarged are sufficiently divergent from one another to warrant tubercles along the hind edge of the forearm forming a beling treated as separate genera. Crottyanura gen. nov. low but distinct crenulated ridge; fingers with strong basal is morphologically convergent, but is in fact placed in the webbing; toes nearly fully webbed; a prominent inner nominate subtribe of Cycloranina subtribe nov. (formally metatarsal tubercle and a small outer one. Heel of defined in this paper by elimination of other subtribes) adpressed hind limb reaches to eye or beyond; first finger instead of the subtribe Ranoideina subtribe nov.. much smaller and shorter than the second and when All species within the genera Ranoidea, Chirodryas and pressed together the tip of the first finger reaches no Crottyanura gen. nov. are readily separated from all other further than the base of the disc of the second finger; Australasian Tree Frogs (Pelodryadidae) by the following tympanum indistinct; no pale stripe along the suite of characters: supratympanic ridge or along posterior edge of upper jaw; adult snout-vent length 65 mm; dorsal pattern is Vomerine teeth are present; fingers are free or webbed mostly grey or dull green, almost blackish and consists of only at the base; first finger is longer than, equal to or a series of relatively narrow dark lines and spots forming only slightly shorter than the second finger; no outer metatarsal tubercle.

The genera Ranoidea and Chirodryas are separated from These frogs have large protruding eyes, with a dark iris, a the genus Crottyanura gen. nov. by having flanks that are broad head and a slender body. Males get to about 45 strongly granular below a distinct, glandular dorso-lateral mm in body length, but females are considerably larger skin fold with densely packed, large, rounded, usually and get to about 60 mm in length. Their tadpoles have pale coloured granules, contrasting with the smooth large sucker mouths. Dorsal colour ranges from grey, tubercular skin on the back above the skin fold. brown, yellow, orange or red, with or without mottling, In contrast Crottyanura gen. nov. has flanks that are spots or flecks and with or without discrete white or smooth or with a few scattered granules, not contrasting cream blotches or ocelli on the upper surfaces. Snout is with the skin on the back. moderately rounded or acuminate (tending to be pointed) Ranoidea species are readily separated from species and the genus occurs in rainforest stream habitats. within the genus Chirodryas by having either a smooth The genus Mosleyia is readily separated from all other dorsum with few if any rounded warts or tubercles, or if Australasian Tree Frogs (Pelodryadidae) by one or other present in any number or size, are arranged in regular of the two unique suites of characters: longitudinal rows near the vertebral line. 1/ Vomerine teeth prominent and largely between the By contrast Chirodryas species have large warts on the choanae. No pectoral fold; A series of small enlarged back that are irregularly scattered and not configured in tubercles along the hind edge of the forearm forming a regular longitudinal rows. low but distinct crenulated ridge; fingers with strong basal The two subgenera within Ranoidea are separated as webbing; toes nearly fully webbed; a prominent inner follows: Subgenus Ranoidea has a generally smooth metatarsal tubercle and a small outer one. Heel of dorsum with at most a few scattered, low tubercles. By adpressed hind limb reaches to eye or beyond; first finger contrast subgenus Sandgroperanura subgen. nov. has much smaller and shorter than the second and when obvious flat warts on the back that are arranged in pressed together the tip of the first finger reaches no regular longitudinal rows near the vertebral line. further than the base of the disc of the second finger; The build of Crottyanura gen. nov. while solid, is slender tympanum indistinct; no pale stripe along the when compared to both Ranoidea and Chirodryas. The supratympanic ridge or along posterior edge of upper species in Crottyanura gen. nov. are further diagnosed by jaw; adult snout-vent length 65 mm; dorsal pattern is being olive green with brownish tinge above and a mostly grey or dull green, almost blackish and consists of distinctive light mid-vertebral line (in C dahlii but not often a series of relatively narrow dark lines and spots forming in C. crottyi sp. nov.); having the hind side of the thighs a fine, continuous reticulum over the back, head and mottled, marbled or spotted with white (in C dahlii but not limbs. The flanks have a bluish metal sheen. White in C. crotty sp. nov.); a finely granular dorsum; smooth below, commonly with brown on the throat; axilla and white-coloured venter; no dorsolateral skin fold; fully groin flesh coloured; hind side of thighs are dark brown. webbed toes; no outer metatarsal tubercle and inner Skin leathery, finely granular, or with numerous small metatarsal tubercle is not shovel-shaped. scattered warts. Granular below (M. nannotis (Andersson, 1916), M. cottoni sp. nov. M. lorica (Davies and Distribution: Wetter parts of south-east and south-west McDonald, 1979)) (nominate subgenus), or: Australia. 2/ Vomerine teeth conspicuous in two rows on a line with Content: Ranoidea Tschudi, 1838 (type genus); the hind edge of the choanae; with or without pectoral Chirodryas Keferstein, 1867. fold; fingers with conspicuous webbing, reaching at least GEDYERANINA SUBTRIBE NOV. as far as the base of the penultimate phalanx of the LSIDurn:lsid:zoobank.org:act:032114B1-C75A-4721- fourth finger; hind edge of forearm is smooth or with at B3D0-A56EF6704859 most a few, low, discontinuous tubercles; hind edge of Type genus: Gedyerana gen. nov. foot is smooth; hind side of thighs more or less uniform Diagnosis: The subtribe Gedyeranina subtribe nov. is and without black and yellow markings or marbling; best defined by disgnosing and defining the two colour brown to blackish above, never green, with or component genera. without a broad, darker vertebral patch and other Gedyerana gen. nov. is readily separated from all other contrasting markings; a dark dorsal patch, if present, is Australasian Tree Frogs (Pelodryadidae) by being the inconspicuous and commencing from a line joining the only Australasian Tree Frog species having a lower centre of each eye; tympanum covered by skin (adult size eyelid with a characteristic reticulatum or palpebral is 35-50 mm body length) (M. michaelsmythi sp. nov.; M. venation of fine, pigmented lines and a horizontally nyakalensis (Liem, 1974); M. pilloti sp. nov.; M. rheocola elliptical pupil. (Liem, 1974)) (subgenus Amnisrana subgen. nov.). All other species within this region with a lower eyelid Distribution: North-east Australia. with a characteristic reticulatum or palpebral venation of Content: Gedyerana gen. nov. (type genus); Mosleyia fine, pigmented lines have a vertically elliptical pupil, Wells and Wellington, 1985. including species within Nyctimystes Stejneger, 1916 DARANINANURINI TRIBE NOV. sensu lato, as defined by other authors including Cogger LSIDurn:lsid:zoobank.org:act:E6C1197B-B538-443D- (2014) at page 197 under the heading “genus 996F-9AA74B74A1BE Nyctimystes Stejneger, 1916”. Type genus: Daraninanura gen. nov. Both species within the genus Gedyerana gen. nov. Diagnosis: The tribe Daraninanurini tribe nov. is best would be diagnosed as either “Litoria dayi” in Anstis defined by diagnosing and defining the single genus. (2013) or “Nyctimystes dayi” in Cogger (2014). The genus Daraninanura gen. nov. monotypic for the type

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 156 Australasian Journal of Herpetology species D. brevipalmata (Tyler, Martin and Watson, 1972) Skin below is granular. Small but prominent vomerine is readily separated from all other Australasian Tree teeth are located mostly behind the choanae. A pectoral Frogs (Pelodryadidae) by the following unique suite of fold is indistinct. Finger and toes discs are moderate, characters: Rich brown to chocolate brown above, being only a little wider than the digits. Fingers with occasionally with scattered small black flecks. There is a distinct basal webbing and the toes are fully webbed. wide canthal stripe running from snout to eye, continuing There is a small but prominent inner metatarsal tubercle, past the eye as a wide black band, almost over-writing no outer one. Tympanum is indistinct. Second finger is the standard (for frogs) sized tympanum, continuing to larger than the first; adult size to 45 mm in lenth. the flank and sometimes bordered above with white or The tadpole is free-swimming, elongated and flattened, yellow. The upper lip has a narrow white or yellow stripe, and reaches a total length of 40 mm prior to narrowly edged below with brown, which continues as a metamorphosis. The body is dark brown to black above, glandular stripe from the angle of the mouth to the base with fine silver chromatophores extending onto the flanks. of the forearm. The lower flanks are yellowish with Darker spots may be present on the dorsal surface, while scattered black spots, flecks or peppering. Groin is green the ventral surface is darkly pigmented. The tail fin and or blue green. There are no red or orange spots on the muscle are covered with fine melanophores. The tail is hind side of the thighs. There is a dark stripe along the moderately thick and has a rounded tip. The eyes are front edge of the hindlimb. Venter is white to light yellow. dorso-lateral, and the mouth is ventral. Top of iris is silver to gold in colour. Skin is smooth to The oral disc is large relative to closely-related species, slightly leathery above and coarsely granular below. and the oral papillae have a wide anterior gap. There are Snout is rounded in shape. Vomerine teeth are prominent two rows of anterior labial teeth and three posterior rows between the choanae. There is no pectoral fold. Finger (Hero et al. 1995; Anstis 2013). and toe discs are of medium size, fingers are unwebbed In terms of morphologically similar and potentially and toes about one third webbed. There is a prominent sympatric species the warty back of Fiacumminganurea inner metatarsal tubercle and an indistinct small outer gen. nov. distinguishes species in this genus from the tubercle. The second finger is longer than the first, the morphologically similar species Dryopsophus nudidigita first finger being so short that when pressed together with (Copland, 1962) and its lack of a distinct tympanum the second, it reaches no further than the base of the distinguishes it from Dryopsophus citropa (Dümeril and disc of the second. Bibron, 1841). Duellman et al. (2016) found the type and only species in Frogs in the genus Dryopsophus Fitzinger, 1843 as the genus Daraninanura gen. nov. to have diverged from defined herein, are readily separated from all other its nearest living relative 35.1 MYA, necessitating the Australasian Tree Frogs (Pelodryadidae) by the following transfer of the species D. brevipalmata (Tyler, Martin and unique suite of characters: Watson, 1972) to a new genus and also a monotypic Vomerine teeth present and the hind edge is behind the tribe. choanae; fingers are free or only webbed at the base; the Photos of Daraninanura brevipalmata (Tyler, Martin and first finger is much shorter and smaller than the second Watson, 1972), can be found in Cogger (2014) on page and when pressed together the tip of the first finger 153 (two images), Vanderduys (2012) on page 27, Eipper reaches no further than the base of the disc of the and Rowland (2018) on page 119 at top and Anstis second finger; the heel of the adpressed hindlimb (2013) on pages 156 (right side), 157 and 158. reaches to the eye or beyond; and one or other of the Distribution: Known only from wet sclerophyll forests of following three suites of characters: the north coast of New South Wales, north from about 1/ There is a narrow light line along at least the posterior Gosford and into the wetter parts of south-east edge of the upper jaw, from below the eye to the Queensland, Australia. glandular region behind the angle of the mouth; the Content: Daraninanura gen. nov. (monotypic). tympanum is distinct; the tympanic annulus is clearly FIACUMMINGANURINI TRIBE NOV. visible (subgenus Dryopsophus), or: LSIDurn:lsid:zoobank.org:act:846EAAC5-5DCD-4A61- 2/ There is a narrow light line along at least the posterior 969E-AA8CC9E2C3A8 edge of the upper jaw, from below the eye to the Type genus: Fiacumminganura gen. nov. glandular region behind the angle of the mouth; the Diagnosis: The tribe Fiacumminganurini tribe nov. is tympanum is indistinct; the tympanic annulus is indistinct, best defined by disgnosing the two component genera. (subgenus Leucolatera), or: The three morphologically similar species within 3/ There is no light line along the posterior edge of the Fiacumminganurea gen. nov. are readily separated from upper jaw; there is a narrow light green, white or gold all other Australasian Tree Frogs (Pelodryadidae) by the stripe, sometimes with indefinite edges, but always following unique suite of characters: bordered below with dark brown or black along the supra- A colouration that is grey to green above, with irregular tympanic ridge; fingers have barely a trace of webbing darker mottling, marbling or flecks, the latter often and the white or gold supratympanic streak is sharp forming irregular cross-bands on the limbs. Ventral edged (subgenus Ausverdarana subgen. nov.) surface white or yellow, or white becoming yellow towards Distribution: Wetter parts of the east coast of Australia, the rear. Lower and concealed surface of limbs are mainly near the coast. yellowish. Content: Fiacumminganurea gen. nov. (type genus); Skin is somewhat leathery, with a few tiny whitish Dryopsophus Fitzinger, 1843. tubercles on the back becoming numerous on the sides. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 157

DRYOPSOPHINA SUBTRIBE NOV. scattered small pointed tubercles on an otherwise LSIDurn:lsid:zoobank.org:act:A85B6140-C767-49C9- smooth body in Kumanjayiwalkerus gen. nov.. 9543-A5BD1A806823 Furthermore species within Pengilleyia invariably have Type genus: Fiacumminganura gen. nov. green spots, flecks or blotches on the back versus none Diagnosis: The subtribe Dryopsophina subtribe nov. is in Kumanjayiwalkerus gen. nov.. best defined by disgnosing the the component generus. Kumanjayiwalkerus gen. nov.. has a strongly contrasting Frogs in the genus Dryopsophus Fitzinger, 1843 as reddish-brown upper iris, with grey below, versus either defined herein, are readily separated from all other weakly contrasting reddish-brown upper iris or the iris Australasian Tree Frogs (Pelodryadidae) by the following being grey all over in Pengilleyia. unique suite of characters: The genera Colleeneremia Wells and Wellington, 1985 Vomerine teeth present and the hind edge is behind the and Audaxura gen. nov. are all readily separated from all choanae; fingers are free or only webbed at the base; the other Australasian Tree Frogs (Pelodryadidae) by the first finger is much shorter and smaller than the second following unique suite of characters: Grey, brown or fawn and when pressed together the tip of the first finger above, usually with a broad, darker vertebral band, reaches no further than the base of the disc of the bounded on either side by a lighter brown zone. There is second finger; the heel of the adpressed hindlimb a blackish stripe along the side of the head, continuing reaches to the eye or beyond; and one or other of the behind the eye, over the base of the forelimb and along following three suites of characters: the side of the body almost to the groin. Dorsal surface of 1/ There is a narrow light line along at least the posterior the body and limbs is flecked with dark brown or black edge of the upper jaw, from below the eye to the and sometimes small to medium sized patches of darker glandular region behind the angle of the mouth; the pigment. Hind side of thighs brown with fine white spots. tympanum is distinct; the tympanic annulus is clearly Groin is usually lemon-yellow. Ventral surface white, visible (subgenus Dryopsophus), or: cream or yellowish. 2/ There is a narrow light line along at least the posterior Dorsal surface usually smooth or with numerous tiny edge of the upper jaw, from below the eye to the granules above and coarsely granular below. Vomerine glandular region behind the angle of the mouth; the teeth almost entirely behind the choanae. Pectoral fold tympanum is indistinct; the tympanic annulus is indistinct, prominent. Finger and toe discs large. Toes about two (subgenus Leucolatera), or: thirds webbed. Inner metatarsal tubercle moderate and elongated, outer tubercle is small and rounded. 3/ There is no light line along the posterior edge of the Tympanum is large, rounded and distinct. Second finger upper jaw; there is a narrow light green, white or gold longer than first. Average adult size 35 mm in length. stripe, sometimes with indefinite edges, but always bordered below with dark brown or black along the supra- Species within the genus Colleeneremia are separated tympanic ridge; fingers have barely a trace of webbing from the morphologically similar species within the genus and the white or gold supratympanic streak is sharp Audaxura gen. nov., their closest living relatives as edged (subgenus Ausverdarana subgen. nov.). follows: Colleeneremia species always exhibits a very broad, dark stripe on the side of the head and body. This Distribution: Wetter parts of the east coast of Australia, is not found in species within Audaxura gen. nov.. mainly near the coast. Furthermore Audaxura gen. nov. have pale regular or Content: Dryopsophus Fitzinger, 1843 (monotypic). irregular stripes or patches on the dorsal surface of the KUMANJAYIWALKERINI TRIBE NOV. body not exhibited in the same configuration or form in LSIDurn:lsid:zoobank.org:act:F7FA4EDF-1AC8-47B8- any Colleeneremia species. Colleeneremia have shorter B0BA-0B54DE669D22 limbs than Audaxura gen. nov.. The TL/S-V ratios Type genus: Kumanjayiwalkerus gen. nov. provides a means of distinguishing the genera, this being Diagnosis: The tribe Kumanjayiwalkerini tribe nov. is 0.335-0.432 for Colleeneremia and 0.477-0.520 for best diagnosed by way of defining each of the six Audaxura gen. nov.. component genera. Within Colleeneremia the two subgenera are separated The genera Pengilleyia Wells and Wellington, 1985 as as follows: defined within this paper, and Kumanjayiwalkerus gen. 1/ Fingers with only rudimentary webbing, being the nov. are as a pair, both readily separated from all other subgenus Colleeneremia, or: Australasian Tree Frogs (Pelodryadidae) by the following 2/ With conspicuous webbing on the fingers, reaching at unique suite of characters: least as far as the base of the penultimate phalanx of the Vomerine teeth present; fingers with conspicuous fourth finger being the subgenus Balatusrana subgen. webbing reaching at least as far as the base of the nov.. penultimate phalanx of the fourth finger; hind edge of Brevicrusyla gen. nov. includes two species from the forearm is smooth, or with at most a few low, Indonesian side of New Guinea that are morphologically discontinuous tubercles; hind edge of foot is smooth; similar to species within the genera Colleeneremia Wells hind side of thighs with contrasting black and yellow bars and Wellington, 1985 and Audaxura gen. nov. as detailed or marbling, at least dorsally. above, but are readily separated from them in that adults The genus Pengilleyia Wells and Wellington, 1985 is of Brevicrusyla gen. nov. while having a smooth dorsum readily separated from the genus Kumanjayiwalkerus on the body like the other two genera, instead has gen. nov. by having a back that is either very warty or tubercles on the head, that is not seen in the other two moderately warty, versus virtually smooth or with well genera.

Tadpoles of Brevicrusyla gen. nov. have a long muscular is not found in species within Audaxura gen. nov.. tail, with narrow dorsal and ventral fins, versus a Furthermore Audaxura gen. nov. have pale regular or relatively short tail, with broad fins in species within irregular stripes or patches on the dorsal surface of the Audaxura gen. nov. and Colleeneremia. body not exhibited in the same configuration or form in The nine known living species within the genus any Colleeneremia species. Colleeneremia have shorter Rawlinsonia Wells and Wellington, 1985 (one formally limbs than Audaxura gen. nov.. The TL/S-V ratios described within this paper) are readily separated from all provides a means of distinguishing the genera, this being other Australasian Tree Frogs (Pelodryadidae) by the 0.335-0.432 for Colleeneremia and 0.477-0.520 for following unique suite of characters: smallish frogs Audaxura gen. nov.. usually about 35 mm in body length. Fingers are free or Within Colleeneremia the two subgenera are separated only webbed at the base. First finger is much shorter and as follows: smaller than the second, when pressed together the tip of 1/ Fingers with only rudimentary webbing, being the the first finger reaches no further than the base of the subgenus Colleeneremia, or: disc of the second finger. Hind edge of vomerine teeth 2/ With conspicuous webbing on the fingers, reaching at are between the choanae. Usually one or a pair of least as far as the base of the penultimate phalanx of the semidistinct mid dorsal patches, darker than the ground fourth finger being the subgenus Balatusrana subgen. colour, extending from a line joining the centre of the nov.. eyes; white stripe below eye, if present does not extend Brevicrusyla gen. nov. includes two species from the beyond the anterior edge of the base of the forelimb. Indonesian side of New Guinea that are morphologically Distribution: Most parts of continental Australia and New similar to species within the genera Colleeneremia Wells Guinea. and Wellington, 1985 and Audaxura gen. nov. as detailed Content: Kumanjayiwalkerus gen. nov. (type genus); above, but are readily separated from them in that adults Pengilleyia Wells and Wellington, 1985; Audaxura gen. of Brevicrusyla gen. nov. while having a smooth dorsum nov.; Brevicrusyla gen. nov.; Colleeneremia Wells and on the body like the other two genera, instead has Wellington, 1985; Rawlinsonia Wells and Wellington, tubercles on the head, that is not seen in the other two 1985. genera. Tadpoles of Brevicrusyla gen. nov. have a long AUDAXURINA SUBTRIBE NOV. muscular tail, with narrow dorsal and ventral fins, versus LSIDurn:lsid:zoobank.org:act:CF7BDF2F-FFF5-4B08- a relatively short tail, with broad fins in species within AAB9-DE506607B437 Audaxura gen. nov. and Colleeneremia. Type genus: Audaxura gen. nov. Duellman et al. (2016) found that the genera Diagnosis: The subtribe Audaxurina subtribe nov. Colleeneremia and Audaxura gen. nov. diverged from comprises three genera, being Audaxura gen. nov., one another 17 MYA, with a similar divergence indicated Brevicrusyla gen. nov. and Colleeneremia Wells and by Pyron and Weins (2011). Wellington, 1985. Distribution: Species in this tribe are found in most parts All species in this subtribe are readily separated from all of Australia and New Guinea. other Australasian Tree Frogs (Pelodryadidae) by the Content: Audaxura gen. nov. (type species); Brevicrusyla following unique suite of characters: gen. nov.; Colleeneremia Wells and Wellington, 1985. Grey, brown or fawn above, usually with a broad, darker RAWLINSONINA SUBTRIBE NOV. vertebral band, bounded on either side by a lighter brown LSIDurn:lsid:zoobank.org:act:FD2468BD-A21E-4F61- zone. There is a blackish stripe along the side of the BBA8-A97170185AB3 head, continuing behind the eye, over the base of the Type genus: Rawlinsonia Wells and Wellington, 1985. forelimb and along the side of the body almost to the Diagnosis: Rawlinsonina subtribe nov. is monotypic for groin. Dorsal surface of the body and limbs is flecked the genus Rawlinsonia Wells and Wellington, 1985 and with dark brown or black and sometimes small to medium so the genus diagnosis is the same as that for the tribe. sized patches of darker pigment. Hind side of thighs The nine known living species within the genus brown with fine white spots. Rawlinsonia Wells and Wellington, 1985 (one formally Groin is usually lemon-yellow. Ventral surface white, described within this paper) are readily separated from all cream or yellowish. Dorsal surface usually smooth or with other Australasian Tree Frogs (Pelodryadidae) by the numerous tiny granules above and coarsely granular following unique suite of characters: smallish frogs below. Vomerine teeth almost entirely behind the usually about 35 mm in body length. Fingers are free or choanae. Pectoral fold prominent. Finger and toe discs only webbed at the base. large. First finger is much shorter and smaller than the second, Toes about two thirds webbed. Inner metatarsal tubercle when pressed together the tip of the first finger reaches moderate and elongated, outer tubercle is small and no further than the base of the disc of the second finger. rounded. Tympanum is large, rounded and distinct. Hind edge of vomerine teeth are between the choanae. Second finger longer than first. Average adult size 35 mm Usually one or a pair of semidistinct mid dorsal patches, in length. darker than the ground colour, extending from a line Species within the genus Colleeneremia are separated joining the centre of the eyes; white stripe below eye, if from the morphologically similar species within the genus present does not extend beyond the anterior edge of the Audaxura gen. nov., their closest living relatives as base of the forelimb. follows: Colleeneremia species always exhibits a very Duellman et al. (2016) found that the living species in the broad, dark stripe on the side of the head and body. This Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 159

genus Rawlinsonia diverged from their nearest living unwebbed or slightly webbed, long fingers, large finger relatives 23.2 MYA, giving support to the erection of this discs and fully webbed toes. Intercalary structures are genus by Wells and Wellington, 1985 and this is before broad or elongate and ossified. Straight canthus rostralis. one considers the significant morphological divergence of The hyoid plate lacks alary processes. The eggs are the species group. large and unpigmented, or alternatively: Distribution: Wetter parts of south-east Australia, 2/ Medium to large species wlth long and variously ranging from South Australia, through Victoria and webbed fingers, long hindlimbs, pigmented or Tasmania, along the New South Wales coast to south- unpigmented bones and very highly variable dorsal east Queensland, with outlier populations at Eungella, coloration. The intercalary stiuctures are small and west of Mackay and the Atherton Tableland and adjacent cartilaginous. The hyoid plate bears pedunculate alary mountains in the southern wet tropics of North processes. The ova are small and pigmented. Queensland. Frogs in the subgenus Alliuma subgen. nov. are Content: Rawlinsonia Wells and Wellington, 1985 separated from all other Australasian (Australian and (monotypic). New Guinea) Tree Frogs (Pelodryadidae) by the following MAXINEHOSERRANINI TRIBE NOV. suite of characters: They are medium-sized montane tree LSIDurn:lsid:zoobank.org:act:876587D4-4B6A-4CA2- frogs with spiniform tubercles on the hindlimbs, but no 86B6-44504B5B98C7 spiniform tubercles on the body, a green and brown Type genus: Maxinehoserranae gen. nov. blotched dorsum, and yellow colouration on the hidden surfaces of the thighs.The ventral surface is variegated Diagnosis: The tribe Maxinehoserranini tribe nov. is with dark pigments. Moderate to extensive finger extremely speciose and contains 13 morphologically webbing. Vocal slits present in males; strongly curved divergent, but phylogenetically close genera. canthus rostralis. Therefore the most sensible way to define and diagnose this tribe is by way of doing so by defining and Frogs in the subgenus Naveosrana subgen. nov. are diagnosing each genus as a group. separated from all other Australasian (Australian and New Guinea) Tree Frogs (Pelodryadidae) by the following Species within Maxinehoserranae gen. nov. are suite of characters: A small (under 50 mm snout-vent in separated from all other Australasian (Australian and both sexes) grey or brown species of frog with broad half- New Guinea) Tree Frogs (Pelodryadidae) by the following webbed fingers bearing large discs and having a strongly suite of characters: curved canthus rostralis. The intercalary structures are No vomerine teeth (Australian species) or in two small elongate and cartilaginous. The hyoid plate lacks alary patches between the choanae (New Guinea species); processes. Ova are small and unpigmented. Further dorsal colour is usually green, or occasionally fawn or a diagnostic characters of this subgenus are as follows: mixture of green and bronze; dorsal surface has a broad The dorsum varies from grey to dark brown with pale vertebral band of bronze bordered on either side by markings including light pigment over the site of the green; in terms of flecks or blotches, there are at most a cutaneous blood vessels. The snout is gently rounded few dark flecks on the dorsal surface; dorsal surface is and the canthus rostralis sharply curved. The fingers smooth; at least one strong pectoral fold; no tubercles have broad discs and are approximately half webbed. above the eye; brown head streak present; tympanum The toes are webbed to the base of the discs. Tubercles brown; internarial distance/eye-naris distance ratio is less occur commonly on the upper eyelids, below the anus, than 1.0. on the back of the thighs and on the heels. Frogs within the subgenus Vegrandihyla subgen. nov. are The cranial elements are reduced. The nasals are small separated from the nominate subgenus and very widely separated medially. They do not Maxinehoserranae subgen. nov. by having concealed articulate with the sphenethmoid, which appears to be surfaces of the legs being bright red in colour (in life), lobulated anteriorly in retaining a double condition, and versus blue-black, brown, yellow to orange in colour in all does not extend between the nasals. The frontoparietal other species. fontanelle is large and ovoid. The squamosal has a short Vegrandihyla subgen. nov. are further separated from zygomatic ramus and slightly longer otic ramus. The New Guinea species within Maxinehoserranae subgen. quadratojugal is not developed. The pars facialis is nov. by their non-overlapping HL/HW ratios, 1.027-1.189 shallow and the short posterior process does not versus 1.243-1.254. In terms of the Australian species in articulate with the maxillary process of the nasal. The Maxinehoserranae subgen. nov. the magnitude of the alary processes of the premaxillaries are well developed, difference is less, being a mean HL/HW for Vegrandihyla bifurcated at their extremities, widely separated medially subgen. nov. 1.04, versus 1.10 in the Australian species and perpendicular to the pars dentalis. The palatine in Maxinehoserranae subgen. nov.. processes are well developed and do not articulate with The described species in the genus Angularanta gen. each other medially. The prevomers are reduced. The nov. are most easily separated from all other Australasian sacral diapophyses are broadly expanded and the ilia (Australian and New Guinea) Tree Frogs (Pelodryadidae) extend one third along their length. No flange is present by separation of each of the relevant subgenera. on the third metacarpal and the intercalary structures are Frogs in the nominate subgenus Angularanta subgen. long and cartilaginous. Alary processes of the hyoid plate nov. are separated from all other Australasian (Australian are lacking. The adductor mandibulae externus and New Guinea) Tree Frogs (Pelodryadidae) by one of superficialis is absent. the two following suites of characters: Frogs within the subgenus Scelerisqueanura subgen. 1/ Small to medium sized, stream-breeding species with Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 160 Australasian Journal of Herpetology nov. are separated from all other Australasian (Australian thirds that from the eye. The distance from the eye and and New Guinea) Tree Frogs (Pelodryadidae) by the the naris is less than the internarial span. The canthus following suite of characters: Small (adult males 25.8- rostralis is well defined and strongly rounded. The eyes 30.3 mm) frogs. Dorsum is chocolate brown, with or are large and prominent the eye diameter being greater without paler patches. Short, narrow fringed, half-webbed than the eye to naris distance. fingers and webbed toes. The finger lengths are as The tympanum is covered with skin, only the inferior half follows 3>4>2>1. The webbing between the third and of the annulus is visible. The tympanum diameter fourth fingers extends to a point slightly below the paired approximates the equivalent of two fifths of the eye subarticular tubercles at the base of the penultimate diameter, separated from the eye by a distance greater phalanx on the fourth toe. Broadly spaced nares (E-N/IN than its own diameter. 0.657-0.758). No vomerine teeth. The head is slightly The tongue shape is somewhat distorted and roughly longer than broad (HL/HW 1.031), its length equivalent to cordiform in shape with a very slightly indented posterior more than one-third of the snout to vent length (HL/S-V margin. 0.356). The snout is not prominent; abrupt and truncate The two species Angularanta impura and A. oxyeei sp. when viewed from above and very slightly rounded in nov. constitute the entirety of the subgenus Raucus profile. The nostrils are lateral, their distance from the subgen. nov. and are separated from all other species end of the snout slightly less than that from the eye. The within Angularanta gen. nov. and all other Australasian distance between the eye and the naris is less than the (Australian and New Guinea) Tree Frogs (Pelodryadidae) internarial span (E-N/IN 0.694). The canthus rostralis is by the following suite of characters: unpigmented bones; well defined and very slightly curved. The eye is large single subarticular tubercle on fourth finger; reduced toe and conspicuous, its diameter greater than the eye to webbing; reduced hand webbing (versus moderate to naris distance. The tympanum is covered with skin and extensive in one or both in all species of Angularanta very small, its diameter equivalent to one-third of the eye gen. nov.), narrow lateral fringes on fingers; dark brown diameter, separated from the eye by a distance greater to blackish chin in males; dark brown to reddish brown than its own diameter. The tongue is broadly cordiform dorsum; dark brown canthal stripe; upper-lip may be with weakly indented posterior border. white, with a very thin white line bordered by black or The terminal discs are prominent. Long and slender brown; concealed surfaces of thighs are brown with hindlimbs with a TL/S-V ratio of 0.584. yellow spots; yellow to orange iris, sometimes with a Toe lengths 4>5=3>2>1. The webbing of all toes except green upper margin; ventrum white; slightly pointed snout for the fourth reaches the base of the discs, while on the when viewed from above, side on or below; a raucous fourth toes the webbing reaches the subarticular tubercle call; IN/SV 0.072-0.08, TY/SV 0.067-0.075, HW/SV 0.34- at the base of the penultimate phalanx and continues to 0.3, HL/SV 0.34-0.3, EN/IN 1.1-1.2. the disc via a narrow fringe. The dorsal and lateral The five described species in the genus Bellarana gen. surfaces of the body are finely pitted and striated. There nov. are readily separated from all other Australasian is an inconspicuous supratympanic fold. The throat and (Australian and New Guinea) Tree Frogs (Pelodryadidae) chest are smooth. Abdomen and nearby halves of the by the following suite of characters: The head is high and ventral surface of the thighs are coarsely granular. There usually broader than long (average HL/HW 0.878-1.012), is a small pigmented nuptial pad at the base of the first its length usually less than one-third of the snout to vent finger. Vocal sac openings are exceptionally long, length (average HL/S-V 0.300-0.347). The snout is not extending from the base of the tongue to the angles of prominent; when viewed from above it is evenly rounded the jaw. or obtusely angular; in profile it is rounded or acutely Ventrally the frogs are a pale creamish colour, stippled angular. The nostrils are more lateral than superior, their with dark brown on the throat. distance from the end of the snout less than that from the Frogs within the subgenus Longuscrusanura subgen. eye. The distance between the eye and the naris is either nov. are separated from all other Australasian (Australian more or less than the internarial span (average E-N/IN and New Guinea) Tree Frogs (Pelodryadidae) by the 0.509-0.884). The canthus rostralis may be curved, following suite of characters: Small size with male slightly defined or straight. The eye is moderate to large, maximum length 23 mm, female 28 mm. Fingers are its diameter greater than the distance separating it from slightly fringed and slightly less than half webbed and of the nostril. The tympanum is visible, its diameter moderate length. The toes are fully webbed and the equivalent to one-third to one-quarter of the eye hindlimbs are particularly long, with a ratio of tibia length diameter. The vomerine teeth are in two short oblique to snout-vent length of 0.59-0.68. series situated between the choanae. The tongue is The dorsum is greyish (in preservative) and marked with cordiform wih a slightly indented posterior border. pale, cryptic markings resembling lichens. The fingers are moderately long and are equipped with The snout is short and very slightly pointed. narrow lateral fringes; in decreasing order of length Ovarian eggs are unpigmented. 3>4>2>1; the webbing is not extensive, not reaching the Vomerine teeth are present in some individuals and sub-articular tubercle at the base of the penultimate absent in others. phalanx on the fourth finger. The terminal discs are moderate. The snout is prominent, slightly projecting and obtusely pointed when viewed from above and slightly projecting The hind limbs are long with a moderate to high TL/S-V in profile. The nostrils are more lateral than superior, their ratio (0.524-0.689). Toes in decreasing order of length distance from the end of the snout approximately two 4>3>or= 5>2>1. The toes are webbed to the discs with

the exception of the fourth which is webbed to the sub- in dorsal view and truncate in lateral view. Rostral spike articular tubercle at the base of the penultimate phalanx. in males extending from tip of upper jaw; mean spike The skin on the dorsal surfaces is smooth or at most length varies among populations and species from slightly tubercular with very small and widely scattered different localities. Canthus rostralis moderately well tubercles distributed evenly across the upper body and defined, slightly curved; loreal region concave. Nares upper surfaces of the limbs. The throat is smooth and the closer to tip of snout (excluding spike) than to eye, chest, abdomen and lower femora granular. There is a oriented laterally. Eyes moderately large, protruding in prominent, curved supra-tympanic fold extending from both dorsal and lateral views; pupil horizontal. Upper jaw the posterior corner of the eye to the shoulder. On the protruding marginally beyond lower jaw. Tympanum small posterior surface of the forearm are tubercles which are with distinct to indistinct annulus, bordered dorsally by a widely separated from one another, in juxtaposition in a fleshy supratympanic fold extending to the superior edge distinct row, or replaced by a continuous fold. There is a of the insertion of the upper arm. Choanae small and poorly defined dermal ridge on the posterior surface of circular, situated close to anterior and lateral edge of the heel and tarsus. palate; no vomerine teeth visible; tongue The colouration is very highly variable, including at times, fleshy and ovoid and usually with a slightly indented strong sexual dimorphism in many locations. posterior edge. Vocal slits present in males. Dorsal skin Fluvirana gen. nov. are a genus of stream-dwelling frogs tuberculate; ventral skin finely to coarsely granular on only known from the north-west New Guinea (Indonesia) throat, abdomen and tibia; remaining ventral surfaces of separated from all other similar species in New Guinea limbs smooth; additional tubercles also present in (including those sympatric in northern New Guinea) by clusters around the vent, and to a variable extent on having a stout build, only slightly pointed snout from sides of ankles and upper forearms. above or below, also being slightly truncate; distinctive Fingers with relative lengths 3>4>2>1; fleshy, whitish to finger webbing on otherwise short, thick fingers, an translucent webbing between all digits, forming a narrow absence of distinctive markings on the dorsum; brownish basal strip between 1 and 2, extending to disc on distal or grey dorsum; limited markings on upper limbs, and edge of 2 and proximal edge of 4 and to penultimate usually in the form of scattered flecks or indistinct bars or phalanx on both sides on 3. Terminal finger discs spots; white to whitish underparts; dorsal skin that is expanded or narrower (depending on species) and either smooth or only moderately granular or tuberculate; usually wider than toe discs and with distinct circum- short thick limbs which are greyish in colour and spotted marginal grooves. Nuptial pads dark brown, roughly tear- with white; relatively dull concealed areas of limbs; shaped with point of tear oriented in posteroventral moderate tympanic fold that covers the top section of the direction. Indistinct unpigmented bifid subarticular otherwise exposed tympanum. tubercles usually visible at the base of penultimate The species in the genus Hopviridi gen. nov. are readily phalanx on all fingers, further indistinct unpigmented separated from all other Australasian (Australian and subarticular tubercles in series on finger four, indistinct New Guinea) Tree Frogs (Pelodryadidae) by the following proximal metacarpal tubercles at base of finger one, and suite of characters: Small, montane species. They are small distal metacarpal tubercles at base of three and uniformly green, sometimes lacking flash markings on the four. undersides of limbs or alternatively pink with yellow spots Toes moderately long, relative lengths 4>3>5>2>1. All in H. leucova (Tyler, 1968) or otherwise bright orange in digits with extensive fleshy, opaque webbing, basal H. chloronota (Boulenger, 1911). Short, slightly fringed, between 1 and 2, extending to anterior end of half-webbed fingers, fully webbed toes and eggs penultimate phalanx on distal edge of 2 and 3 and unpigmented, 2 mm in diameter ova. Maximum size of proximal edge of five, to halfway along penultimate females is 30 mm snout-vent. The snout is short with a phalanx on proximal edge of three and four, and base of low internarial span. Vomerine teeth are absent. penultimate tubercle on distal edge of four. Terminal discs The genus Incertanura gen. nov. are separated from all slightly expanded to not really expanded and otherwise, other Australasian (Australian and New Guinea) Tree narrower than finger discs and with distinct circum- Frogs (Pelodryadidae) by the following suite of marginal grooves. Indistinct unpigmented subarticular characters: A long (up to at least 2.8 mm) erectile rostral tubercles on penultimate phalanx of all toes, single on 1- spike in males that is circular in cross section; relatively 3, bifid on 4 and 5. No other tubercles clearly apparent. small size (SVL up to 30.5 mm); slender build (HW/SVL Small, indistinct unpigmented metatarsal tubercle at base 0.22-0.26); tibia approximately half length of the body of 1. (TL/SVL 0.049-0.053); moderately large eyes (EYE/SVL In preservative, ground colour of all dorsal surfaces is 0.10-0.12); moderately small tympanum (TYM/SVL 0.42- medium brown, with extensive blueish green and darker 0.53); small, rounded and often green tubercles brown flecking, maculations, vermiculations and/or extending across the mid-dorsum in life; brown to blotches across all dorsal surfaces, blueish pattern yellowish-brown dorsal colouration with, or without, elements sometimes coalesce into three indistinct distinct green transverse bands; and orange colouration transverse blotches, overall darker pigmentation in concealed areas of the thighs and axilla. elements tend to be densest on body, and less The five species within Incertanura gen. nov. are further concentrated and more finely reticulated towards distal defined as follows: Body very slender and elongate, tibia extremities of limbs. approximately half length of body; head wider than body Rostral spike is usually light brown, with extensive darker in dorsal profile, clearly distinct from neck. Snout rounded brown flecking at base, tending towards unpatterned at

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 162 Australasian Journal of Herpetology tip. White or yellow patches often, but not always, chest are smooth except for a few flattened tubercles. present on either side of head, extending from The abdomen and lower femora are coarsely granular. posterolateral edge of eye, below tympanum and around There is a short row of tubercles on the outer surface of axilla. Venter predominantly buff, internal organs the fourth finger. A more conspicuous dermal ridge sometimes visible, throat sometimes offwhite, and/or with extends along the outer surface of the tarsus and fifth two clusters of dark brown maculations laterally, tubercles toe. There are numerous tubercles below the anus and around vent and sometimes those along outer edge of two extremely prominent femoral tubercles. The supra- limbs, off-white. tympanic fold is inconspicuous. Appearance in life is as follows: Dorsal base colouration Predominantly green in life and may be marked with gold is light to dark brown, with variable amounts of green and and black. The intercalary structures are elongate and darker brown spotting, mottling or banding on the body, ossified. The hyoid plate lacks alary processes. The ova limbs and head. Dorsal and lateral tubercles on torso and are small and pigmented (see below). limbs often, but not always, green and contrasting The dorsum is green, stippled with black and against the brown base colouration. Head brown, often occasionally marked quite extensively with gold. The with extensive green pigmentation, usually including a fingers are short and slender, with about one third ring webbing; the toes are almost fully webbed. The snout is around the eyes and more variably a green transverse slightly rounded in profile. band between the eyes. Dense clusters of dark-brown to The cranial elements are poorly developed. The nasals almost black maculations extend to a variable degree are widely separated medially and do not articulate with across the limbs, lateral portions of torso and in patches the sphenethmoid. The sphenethmoid remains double in across the back. Off-white patches sometimes present the adult. The frontoparietal foramen is large and ovoid. below tympanum and on exposed surfaces of lower The squamosals have short zygomatic rami and very hindlimbs. Iris pattern complex; base colouration is much longer otic rami. The pars facialis of the maxillary is usually light brownish, with extensive darker brown shallow and the posterior process does not make contact vermiculations; rim of pupil orange. Hidden regions of with the maxillary process of the nasal. The alary axilla and groin orange (adapted from Oliver et al. processes of the premaxillary are broad, widely 2019a). separated medially, and perpendicular to the pars The genus Inlustanura gen. nov. are separated from all dentalis. The palatine processes are well developed and other Australasian (Australian and New Guinea) Tree do not articulate with each other medially. The prevomers Frogs (Pelodryadidae) by the following suite of are very much reduced. characters: The known species are characterised by fully A phlange is present on the distal surface of the 3rd webbed outer fingers, a large rounded head with metacarpal. The sacral diapophyses are broadly dilated, inconspicuous to large eyes (depending on the species), and the intercalary structures are elongate and ossified. and distinctive well developed dermal folds on the There are no alary processes on the hyoid plate. The posterior surface of the forearm, anus, tibia, tarsus and adductor mandibulae externus superficialis is absent. fifth toe. Although in at least one species the The ova are small and pigmented (brown animal pole) development of the fold on the tarsus may be reduced. and laid in stagnant marshes. The mean ovidiameters These frogs have a distinctive dorsal colouration of green are 1.2-1.7 mm. Tadpoles have moderately developed intersperced with yellow spots in life (in preservative a fins and the labial teeth comprise 2 upper and 3 lower distinctive colouration in which blue and violet are the rows (Tyler 1963; Menzies 1972). The chromosome predominant pigments), with yellow spots absent in at number is 26. least some specimens of one species. These frogs are Moechaeanura gen. nov. is further separated from a moderately sized species. Females attain a maximum number of morphologically similar New Guinea species snout to vent length of 35-48 mm and males 39-42 mm. within the genus Variabilanura gen. nov. by its very Species within Moechaeanura gen. nov. are different reproductive mode. Moechaeanura gen. nov. morphologically similar to those species in the genus species have small brown pigmented eggs (average ova Variabilanura gen. nov., with some differences between size of 1.2-1.7 mm) that are deposited in water, while the two detailed in this description. species in the genus Variabilanura gen. nov. attach a Moechaeanura gen. nov. are separated from all other mass of large green eggs with an average ova size of 2.5 Australasian (Australian and New Guinea) Tree Frogs mm to vegetation overhanging slow moving streams. (Pelodryadidae) by the following suite of characters: They In Variabilanura gen. nov. the ova are laid in groups of are small to medium species (male maximum snout-vent from 4 to 37 (mean 14) on the leaves of trees length 34 mm, female maximum snout-vent length 38 mm overhanging water, and around the stems of vegetation at with short, three webbed fingers and almost fully webbed the edge of the water. The ova are surrounded by a very toes. large mass of clear albumen. A period of approximately The webbing of the fourth toe extends to a point midway fourteen days is spent within the spawn clump, and the between the disc and the sub-articular tubercle at the tadpole emerges possessing internal gills and capable of base of the penultimate phalanx, (versus reaches the coordinated movements (Tyler 1978). sub-articular tubercle at the base of the penultimate Frogs within the subgenus Aspercutis subgen. nov. phalanx on the fourth finger in the genus Variabilanura (within Moechaeanura gen. nov.) conform to the above gen. nov.). The skin on the dorsal surfaces of the head, diagnosis for the genus Moechaeanura gen. nov., except body and limbs is minutely roughened. The throat and for their more extensive finger webbing, highly

tuberculate dorsum (versus smooth to slightly tuberculate 4>5=3>2>1. On the fifth toe, the webbing extends two- in the nominate subgenus), distinctively crenulated ridges thirds up the penultimate phalanx of the fifth and to the following the outer surfaces of the lower limbs, silvery- base of the penultimate phalanx on the fourth. gold and heavily veined iris and fully truncate snout The skin on the dorsal surface of the head, body and (versus normal to slightly truncate in the nominate limbs is either smooth with numerous small, spaced subgenus) which as a full diagnosis separates them from apart, scattered tubercles all over, ranging down to all other New Guinea frogs. smooth all over but for a few small and only slightly Frogs within the subgenus Telaater subgen. nov. are developed tubercles on the scapular region. separated from all other New Guinea tree frogs by having The throat, abdomen and lower surface of the thighs are full black webbing on hands and feet, the venter with very weakly granular. There is a row of small tubercles extensive areas of black, white and yellow and a extending posteriorly from the angle of the jaws and a transparent periphery on the tympanic membrane patch of similar sized tubercles beneath the anus. combined with a small adult size of less than 30 mm. the Of the tubercles at the angles of the jaws those anterior placement of these species within a subgenus within a to the tympanum are green, grey or white, whilst those greater Moechaeanura gen. nov. is tentative and posterior to it are usually white. elevation to full genus may be required as further The scapular tubercles are usually green, and the lateral evidence is obtained. surfaces of the body between the axilla and the groin are The species in the genus Ornatanura gen. nov. are one or other of being 1/ Liberally spotted with white, 2/ readily separated from all other Australasian (Australian Grey marks or spots on a cream background, 3/ Back and New Guinea) Tree Frogs (Pelodryadidae) by the marks in the form of large spots or blotches, circled by following suites of characters: These species are white and on an otherwise light lime green background. characterised by their small size (males 23.4-30.0 mm, The ventral surfaces are creamish with less dense females 27.4-35.4 mm), moderate to relatively long limbs stippling than appears on the dorsum and although (average TL/S-V 0.522-0.604) and broadly spaced nares dense stippling occurs in some specimens. (average E-N/IN 0.611-0.818). The colouration is highly The greatest density of groups of chromatophores on the variable. ventral surface appear on the throat and particularly The dorsal surface of the head, body and limbs may towards the labial margins, with more on the upper than range from light sandy grey, or brown to dark grey and is lower margin and greatest intensity between eye and ear, darkened by the presence of very dense, minute, dark although the exact configuration varies between brown to black stippling. The anterior portion of the head specimens and species. bordered laterally by the canthus rostralis, and posteriorly Males vary in size from 23.4 mm to 30.0 mm and females to a line between the anterior portions of the upper from 27.4-35.4mm. The head length is consistently eyelids is an immaculate pale green, although in some longer than broad (average HL/HW 1.028-1.155) and the species this may be reduced to be brown or grey with head length varies from one-third to considerably more some lime green spots or markings. than one-third of the snout to vent length, the complete The head is longer than broad (average HL/HW 1.132), HL/S-V range its length equivalent to more than one-third of the snout being 0.333-0.404. The E-N/IN range is 0.611-0.818 and to vent length (average HL/S-V 0.350). The snout is the eye diameter is consistently smaller than the evenly rounded when viewed from above and projects internarial span. The average TL/S-V range is 0.522- slightly in profile. The nostrils are more lateral than 0.604. superior, their distance from the tip of the snout less than that from the eye. The diploid chromosome number is 26. The distance between the eye and the naris is less than The morphologically similar species Bellarana the internarial span (average E-N/IN 0.727). The canthus micromembrana (Tyler, 1963) from Madang, is most rostralis is well defined and distinctly curved. readily separated from the species within Ornatanura gen. nov. by its distinctive orange upper iris, versus The eye is prominent and bulges above the head, its yellowish in Ornatanura gen. nov. species. diameter greater than the eye to naris distance and less than the internarial span. The superior one-quarter to Species within the genus Nasuscuspis gen. nov. are one-third of the tympanum is hidden beneath the supra- separated from all other Australasian (Australian and tympanic fold. The diameter of the tympanum is New Guinea) Tree Frogs (Pelodryadidae) by the following equivalent to less than one third of the eye diameter to suite of characters: They are montane species, growing slightly more than one-third of the eye diameter. The to a maximum of 50 mm snout vent length in females and vomerine teeth are in two small oblique series between males to 45 mm. The dorsum is a mixture of greys, the choanae. The tongue is almost circular and lacks a greenish greys and ochres. The fingers and toes are posterior indentation. extensively webbed. There are highly developed dermal appendages including crenulated ridges on the posterior The fingers are long and lack lateral fringes; in surfaces of the radius and tarsus, prominent dermal folds decreasing order of length 3>4>2>1. There is a small above the vent; a row of very large tubercles on the vestige of webbing between the third and fourth fingers. undersurface of the mandible, and an elongate rostral, The terminal discs are not prominent, with the degree of dermal spike always in males and females although in expansion varying slightly between species. some reduced form in females of some species. The hind limbs are relatively long with an average TL/S-V Rotundaura gen. nov. is separated from all other ratio of 0.558. Toes in decreasing order of length

Australasian (Australian and New Guinea) Tree Frogs The genus Variabilanura gen. nov. are separated from all (Pelodryadidae) by the following suite of characters: A other Australasian (Australian and New Guinea) Tree very high E-N/IN ratio (average 1.435), short limbs with a Frogs (Pelodryadidae) by the following suite of low TL/S-V ratio (average 0.482), wholly unwebbed characters: They are small montane species (males 24.3- fingers and incompletely webbed toes characterise this 34.1 mm; females 30.1-43 mm). In life they are green, species. usually with yellow spots or markings (in preservative the The head is long and flattened and longer than broad dorsal surface is blue, usually with darker and (average HL/HW 1.073), its length more than one-third of occasionally lighter markings). There is usually a deep the snout to vent length (average HL/S-V 0.355). The violet, orange, yellow or red patch in the groin and snout is rather prominent, gently rounded when viewed similarly coloured markings occur on the axilla, posterior from above and strongly rounded in profile. The nostrils surface of the thighs, tibia and tarsus. The head is as are more superior than lateral, their distance from the broad as long or longer than broad (HL/HW 0.910-1.192), end of the snout about one-half that from the eye, and its length less than or greater than one-third of the snout separated from each other by a distance which is to vent length (HL/S-V 0.328-0.384). The shape of the equivalent to approximately two-thirds of the eye to naris snout is highly variable, being from prominent to distance (average E-N/IN 1.435). The canthus rostralis is inconspicuous and strongly or only slightly rounded when straight and inconspicuous and the loreal region oblique. viewed from above and in profile. The nostrils are lateral The eye is small, its diameter less than the eye to naris in specimens with prominent snouts and more lateral distance. The tympanum is visible, its diameter than superior in specimens with inconspicuous snouts. equivalent to two-thirds of the eye diameter and The nares are very much nearer the eye than the tip of separated from the eye by a distance equivalent to the snout in the latter individuals and almost equidistant approximately one-half its own diameter. The vomerine in the former. The distance between the eye and the naris teeth are in two obliquely oval series in juxtaposition on is less than the internarial span (E-N/IN 0.667-0.965). the midline between the small, obliquely oval choanae. The canthus rostralis is curved and slightly or well The tongue is small, triangular and very feebly indented defined. The eye is large and prominent, its diameter on its posterior border. The fingers are long, slender and greater than the distance separating it from the nostril. unwebbed. The terminal discs are large and the sub- The tympanum is covered with skin, its diameter articular tubercles prominent. The fingers in decreasing equivalent to from one-third to slightly less than one-half order of length 3>4>2>1. The hind limbs are short with a the eye diameter. Vomerine teeth are present in the TL/S-V ratio of 0.482. The toes are incompletely webbed, majority of specimens, and confined to small circular the webbing reaching the base of the penultimate series on slight elevations between the choanae. The phalanx on the fourth toe and half-way up the penultimate tongue is broadly cordiform with a slightly indented phalanx of the fifth. Toes in decreasing order of length posterior margin. The fingers are short and equipped with 4>5=3>2>1. The skin on the dorsal surfaces of the head very slight lateral fringes; in decreasing order of length and body is smooth but for a few flattened tubercles on 3>4>2>1. The webbing reaches the sub-articular tubercle the back. The throat is smooth and the chest, abdomen at the base of the penultimate phalanx on the fourth and back of the thighs are coarsely granular. There is an finger. The terminal discs are prominent. The hind limbs inconspicuous supratympanic fold barely obscuring any are variable with a TL/S-V ratio of 0.485-0.587. Toes in of the tympanum and a pronounced skin fold across the decreasing order of length 4>5>3>2>1. The webbing chest. There is neither a tarsal ridge nor a dermal between the toes reaches the base of the disc of all toes, appendage on the heel. Dimensions: of gravid female except the fourth where it reaches the tubercle at the holotype are snout-vent 33.2 mm; tibia length 15.8 mm; base of the penultimate phalanx. The skin on the dorsal head length 11.8 mm; head width 11.0 mm; eye to naris surface of the head, body and limbs is minutely distance 3.3 mm; internarial span 2.3 mm; eye diameter roughened. The throat and chest are slightly granular. 3.1 mm; tympanum diameter 2.0 mm. The dorsal There are a row of tubercles on the posterior surface of surfaces of the head and body and limbs are a very pale the forearm, and conspicuous tubercles below the anus. brown with indistinct darker and lighter marbling. Marbling Femoral tubercles are usually present. The supra- is present and most conspicuous on the dorsal surface of tympanic fold is inconspicuous. In preservative and in life, the thigh and tibia. The ventral surfaces of the body and dull orange patches may extend on to the dorsum in the limbs are dull cream with an irregular brown patch on the axillary region. throat (modified from Tyler 1978). The lateral surfaces of the body are frequently heavily Specimens of the morphologically similar Llewellynura pigmented with black or deep yellow (in life, being violet Wells and Wellington, 1985 are readily separated by their in preservative), marked with large white patches. There much smaller adult size of less than 25 mm body length. is invariably a yellow, orange, red or violet patch in the Species of Rotundaura gen. nov. are separated from the groin (in life) and the same colouration may be present in genus Summaviridis gen. nov. (and in turn all other the axilla and on the posterior surface of the tibia and Australasian (Australian and New Guinea) Tree Frogs tarsus. There is a white patch beneath the eye extending (Pelodryadidae)), by the tympanum being fully exposed to the angle of the jaws or on the lateral surfaces of the and round, versus the upper surface being cut at the rear body. The ventral surface of the body is cream to white by a well-defined and prominent skin fold forming a (white in subgenus Sudesanura subgen. nov.), with or straight line and a significantly blunter snout in without sparse blue stippling on the throat near the angle Rotundaura gen. nov. versus pointed, when both are of the jaws. The lower surface of the hind limbs is viewed in profile side on. creamish yellow or dull yellow. These species are often

beautifully pigmented in life being marked with cream, base of the disc of the second finger. orange, yellow, green, violet and black (Tyler, 1962). Distribution: New Guinea and tropical parts of northern The ovum is pale green in colour and has a diameter of Australia. approximately 2.5 mm. The ova are laid in groups of from Content: Maxinehoserranae gen. nov. (type genus); 4 to 37 (mean 14) on the leaves of trees overhanging Angularanta gen. nov.; Bellarana gen. nov.; Drymomantis water, and around the stems of vegetation at the edge of Peters, 1882; Fluvirana gen. nov.; Hopviridi gen. nov.; the water. The ova are surrounded by a very large mass Incertanura gen. nov.; Inlustanura gen. nov.; of clear albumen. A period of approximately fourteen Moechaeanura gen. nov.; Ornatanura gen. nov.; days is spent within the spawn clump and the tadpole Nasuscuspis gen. nov.; Rotundaura gen. nov.; emerges possessing internal gills, being capable of Variabilanura gen. nov.. coordinated movements. DRYMONTANTINA SUBTRIBE NOV. The mouth is anterior in position, and surrounded by a LSIDurn:lsid:zoobank.org:act:14F658D7-CE72-464A- band of papillae on the inferior and lateral borders. There B24A-C8371BB6A549 are two upper and three lower rows of labial teeth. Type genus: Drymomantis Peters, 1882. Species in the subgenus Sudesanura subgen. nov. (type Diagnosis: The subtribe Drymomantina subtribe nov. is species “Litoria havina Menzies, 1993”) are separated monotypic for the genus Drymomantis Peters, 1882 and from those in the nominate subgenus by having a red therefore the subtribe diagnosis is the same as for the patch in the groin, versus a deep violet, orange or yellow genus. patch in the groin in the nominate subgenus. Frogs in the subtribe Drymomantina subtribe nov. thereby Species within Sudesanura subgen. nov. are further including genus Drymomantis Peters, 1882 are all readily defined as follows: Small (SV max. 30.05 mm); head separated from all other Australasian Tree Frogs narrow (average HL/HW 1.08); Head always longer than (Pelodryadidae) by the following unique suite of wide (average HL/HW>1.0) except in females, which lack characters: the rostral spike, where HL=HW; canthus rounded, These are small elongated and agile frogs averaging concave, lores oblique, nostrils more or less lateral, about 25 mm in body length with moderately pointed widely placed (average EN/IN 0.63 in males and EN/IN snouts, varying somewhat between species; brown head 0.54 in females); snout with a prominent pointed rostral streak present or absent; dorsal surface is uniform green spike; vomerine teeth absent; eye large (average EY/SV or fawn, sometimes with darker flecks and with at most a 0.11); tympanum visible, upper margin covered by skin paler vertebral zone; there is not a broad vertebral band fold. Fore limb with indistinct row of raised tubercles of bronze bordered on either side with green; skin is down outer side; hind limb without heel lappets or other smooth above; finely granular on the throat and coarsely dermal appendages; fingers half-webbed, toes fully granular below; at least one strong pectoral fold, no webbed; subarticular tubercles poorly developed. Legs dorsolateral fold and no tubercles above each eye; always long (average TL/SV>0.54). Dorsum usually fingers webbed at the base and toes moderately webbed. immaculate, bright pale green, yellow or fawn brown A moderate oval inner and small rounded outer reduced to a very narrow band on the thighs and ceasing metatarsal tubercle; vomerine teeth absent, or if present, at ankle and wrist, leaving hands and feet virtually the hind edge of vomerine teeth are between the colourless; concealed thighs and axillae bright cherry- choanae; first finger is much smaller than the second red; white band on upper lip, snout to axilla; raised when pressed together, the top of the first finger reaching tubercles on fore limb white; ventral surfaces pure white. no further than the base of the disc of the second finger. Frogs in the genus Drymomantis Peters, 1882 this single According to Duellman et al. (2016), the members of this genus being the sole member of the subtribe genus diverged from their nearest living relatives 21.5 Drymomantina subtribe nov. (being the only other MYA confirming tribe-level recognition as being subtribe in the tribe) are all readily separated from all appropriate. other Australasian Tree Frogs (Pelodryadidae) by the following unique suite of characters: These are small Content: Drymomantis Peters, 1882 (monotypic). elongated and agile frogs averaging about 25 mm in NYCTIMYSTINI TRIBE NOV. body length with moderately pointed snouts, varying LSIDurn:lsid:zoobank.org:act:D093F148-149A-4BB1- somewhat between species; brown head streak present 973B-E0C3B72AF48B or absent; dorsal surface is uniform green or fawn, Type genus: Nyctimystes Stejneger, 1916. sometimes with darker flecks and with at most a paler Diagnosis: The tribe Nyctimystini tribe nov. includes a vertebral zone; there is not a broad vertebral band of total of six genera, until recently placed by many authors bronze bordered on either side with green; skin is smooth within an expanded version of the putative genus above; finely granular on the throat and coarsely granular Nyctimystes Stejneger, 1916, sensu Cogger (2014), below; at least one strong pectoral fold, no dorsolateral excluding Australian species previously treated as being fold and no tubercles above each eye; fingers webbed at within Nyctimystes but having a horizontally elliptical the base and toes moderately webbed. A moderate oval pupil. inner and small rounded outer metatarsal tubercle; However the diagnosis of Nyctimystes has been vomerine teeth absent, or if present, the hind edge of expanded (e.g. Duellman et al. 2016) and now shrunk vomerine teeth are between the choanae; first finger is significantly in this (2020) paper. much smaller than the second when pressed together, All genera within this tribe, namely Nyctimystes, the top of the first finger reaching no further than the Occultatahyla gen. nov., Nigreosoculus gen. nov.,

Magnumoculus gen. nov., Badiohyla gen. nov. and The skin is smooth above, granular below. A weak, wavy Albogibba gen. nov. are separated from all other fold is present on the outer surface of the forearm. The Australasian (Australian and New Guinea) Tree Frogs head is broader than long; the canthus rostralis is not (Pelodryadidae) by the following suite of characters: distinct; the loreal region is oblique; Unique to this Large dark eyes, which have vertically elliptical pupils subgenus is that the palpebral venation is distinct, but is and a lower eyelid with a characteristic reticulum or reduced to individual pigment spots and thin, meandering palpebral venation of fine pigmented lines. There is lines, although the lines are still mainly oriented largely in usually a distinct, crenulated skin fold along the hind a vertical direction, with few horizontal interconnections. edge of the forearm and the foot, the heel often having a Species within Nigreosoculus gen. nov. are separated small flap of skin. Males are usually considerably smaller from the other five genera (Nyctimystes, Occultatahyla than females. gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. In further detail, the relevant genera are separated from and Albogibba gen. nov.) by the following suite of one another as follows: characters: Vocal slits present; a very small heel tubercle; Nyctimystes Stejneger, 1916 are separated from the basal webbing on the hand; exposed tympanum; vertical other five genera (Occultatahyla gen. nov., Nigreosoculus lines of palpebral reticulum oriented obliquely and with gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. relatively few horizontal cross-connections; pale-tan to and Albogibba gen. nov.) by having sparse, broken near black iris; rear of thighs barred/mottled with brown, vertically-oriented palpebral venation across the caramel, or blue-gray. nictitating membrane and in having adult males without Species within Magnumoculus gen. nov. are separated vocal slits. from the other five genera (Nyctimystes, Nigreosoculus Frogs in the subgenus Magnummanibus subgen. nov. are gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. separated from those in the nominate subgenus and Albogibba gen. nov.) by the following suite of Nyctimystes and the subgenus Asperohyla subgen. nov. characters: The head is broad (HL/HW <1.0), the snout by their possession of very long, prominent heel lappets. high and blunt, the canthus rostralis distinct, broadly Frogs in the subgenus Asperohyla subgen. nov. are spaced nostrils (EN/IN ratio <1), and the loreal region separated from the other two subgenera by having a nearly vertical. A strong, slightly curved, supratympanic completely hidden tympanum and a dorsal skin fold begins at the posterior corner of the eye and roughened by tiny, conical asperities rather than by disappears above the insertion of the forelimb. The variable sized and more rounded irregularities as seen in tympanum is small but distinct. The hands have little some other New Guinea tree frog species; iris is brown in webbing; the outer fingers are about one-quarter webbed. life. The vomerine teeth are in two patches between the Species within Occultatahyla gen. nov. are separated internal nares, nearly on a line connecting the posterior from the other five genera (Nyctimystes, Nigreosoculus margins of the nares. The skin is minutely granular gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. above, coarser beneath. There is no tubercle on the heel. and Albogibba gen. nov.) by large size (adult females 50 A nearly straight, very slightly crenulated fold of skin is to 84 mm. in snout to vent length); internarial distance present on the outer surface of the forearm. A moderate distinctly smaller than distance from eye to naris (E-N/IN body size (40-60 mm S-V), predominantly brown averages 1.34 (range 1.2-1.5)); the head is broad, colouration, an eyelid venation composed of a network flattish, with depressed snout, closely spaced nares, and with numerous horizontal connections with relatively few oblique loreal region. Palpebral venation is in oblique vertical interconnections. There is decoration on the lines (more vertical than horizontal and with few forearm and tarsus in the form of rows of (often white- horizontal interconnections), with a few horizontal tipped) tubercles. interconnections. A dermal fold passes from the posterior Species within Badiohyla gen. nov. are separated from corner of the eye over the upper edge of the tympanum the other five genera (Nyctimystes, Nigreosoculus gen. and down to the insertion of the forelimb, sometimes nov., Occultatahyla gen. nov., Magnumoculus gen. nov. becoming obscure when leaving the tympanum. The and Albogibba gen. nov.) by having a unique “reticulated” outer fingers are approximately half-webbed, the third palpebral venation, half-webbed fingers and a size in and fifth toes webbed to the disc. A light-coloured dermal males with SVL to maximum of 50 mm to 100 mm ridge or series of tubercles extends from the elbow to the depending on the species. They are further defined as disc of the fourth finger. A similar but less distinct ridge or follows: The snout is relatively short, blunt and high (E-N/ line of tubercles occupies the outer edge of the tarsus IN 0.94), with distinct canthus rostralis and oblique loreal and fifth toe. A very small heel tubercle is present. The region. The vomerine teeth are in two patches between skin of the dorsum is smooth to very slightly granular, that the posterior edges of the choanae. of the venter is always coarsely granulate. Slit-like, vocal The palpebral venation forms a heavy reticulum. The sac openings are usually present in the floor of the tympanum is distinct and is separated from the eye by mouth. A small patch of very fine horny tubercles is found approximately the diameter of the tympanum. A fold of on the first finger of males. The iris is dark or black in skin passes from the posterior corner of the eye, over the colour. upper edge of the tympanum and down, becoming Exceptional to the preceding is the subgenus Webpede indistinct above the insertion of the forelimb. The skin of subgen. nov. which conforms to the above genus the dorsum is minutely roughened, that of the venter diagnosis, save for the following differences: The heel is coarsely granular. A row of tubercles is present along the without a tubercle, or in rare cases has a very small one. outer edge of the forearm and there is a similar row on the tarsus. There is only a slight suggestion of a heel Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 167

tubercle. The outer fingers are approximately half- the posterior edges of the choanae. webbed, the toes, except the fourth toe, are webbed to The palpebral venation forms a heavy reticulum. The the disc. tympanum is distinct and is separated from the eye by The body and head are light brown to purple dorsally, approximately the diameter of the tympanum. A fold of either with irregular dark brown blotches or markings skin passes from the posterior corner of the eye, over the absent. The legs have a similar ground color, with or upper edge of the tympanum and down, becoming without irregular darker spots and bands present on the indistinct above the insertion of the forelimb. The skin of tibia. the dorsum is minutely roughened, that of the venter Species within Albogibba gen. nov. are separated from coarsely granular. A row of tubercles is present along the the other five genera (Nyctimystes, Nigreosoculus gen. outer edge of the forearm and there is a similar row on nov., Occultatahyla gen. nov., Magnumoculus gen. nov. the tarsus. There is only a slight suggestion of a heel and Badiohyla gen. nov.) by one or other of the following tubercle. The outer fingers are approximately half- unique suites of characters: webbed, the toes, except the fourth toe, are webbed to 1/ Being a large species (males to 100 mm. in snout to the disc. The body and head are light brown to purple vent length); of a uniform dorsal coloration (green in life, dorsally, either with irregular dark brown blotches or purple in alcohol-preserved specimens); palpebral markings absent. The legs have a similar ground color, venation is a reticulum without obvious orientation and in with or without irregular darker spots and bands present the nominate subgenus of this genus, the male bears a on the tibia. spine-like process on the anterior surface of the proximal Species within Magnumoculus gen. nov. are separated part of the humerus (upper arm) (nominate subgenus from the other five genera (Nyctimystes, Nigreosoculus Albogibba subgen. nov.), or: gen. nov., Occultatahyla gen. nov., Badiohyla gen. nov. 2/ As above, but male SVL of up to 80 mm; not including and Albogibba gen. nov.) by the following suite of a spine-like process on the anterior surface of the characters: The head is broad (HL/HW <1.0), the snout proximal part of the humerus (upper arm) in males and high and blunt, the canthus rostralis distinct, broadly with a unique dorsal pattern of black vermiculations on spaced nostrils (EN/IN ratio <1), and the loreal region the body and limbs, with unmarked greyish lower flanks nearly vertical. A strong, slightly curved, supratympanic (Ratiobrunneis subgen. nov.). fold begins at the posterior corner of the eye and Distribution: New Guinea including nearby offshore disappears above the insertion of the forelimb. The islands on the continental shelf. tympanum is small but distinct. The hands have little Content: Nyctimystes Stejneger, 1916 (type genus); webbing; the outer fingers are about one-quarter webbed. Albogibba gen. nov.; Occultatahyla gen. nov.; The vomerine teeth are in two patches between the Nigreosoculus gen. nov.; Badiohyla gen. nov.; internal nares, nearly on a line connecting the posterior Magnumoculus gen. nov.. margins of the nares. The skin is minutely granular above, coarser beneath. There is no tubercle on the heel. BADIOHYLINA SUBTRIBE NOV. A nearly straight, very slightly crenulated fold of skin is LSIDurn:lsid:zoobank.org:act:97DD93A0-93D2-43A1- present on the outer surface of the forearm. A moderate 9776-EFF15C005132 body size (40-60 mm S-V), predominantly brown Type genus: Badiohyla gen. nov. colouration, an eyelid venation composed of a network Diagnosis: This tribe includes two genera, namely with numerous horizontal connections with relatively few Badiohyla gen. nov. and Magnumoculus gen. nov.. vertical interconnections. There is decoration on the Species of Nyctimystes, Occultatahyla gen. nov., forearm and tarsus in the form of rows of (often white- Nigreosoculus gen. nov., Magnumoculus gen. nov., tipped) tubercles. Badiohyla gen. nov. and Albogibba gen. nov. are In terms of the nominate subtribe Nyctimystina subtribe separated from all other Australasian (Australian and nov. the four component genera are defined as stated New Guinea) Tree Frogs (Pelodryadidae) by the following below. suite of characters: Nyctimystes Stejneger, 1916 as defined in this paper, are Large dark eyes, which have vertically elliptical pupils separated from the other five genera (Occultatahyla gen. and a lower eyelid with a characteristic reticulum or nov., Nigreosoculus gen. nov., Magnumoculus gen. nov., palpebral venation of fine pigmented lines. There is Badiohyla gen. nov. and Albogibba gen. nov.) by having usually a distinct, crenulated skin fold along the hind sparse, broken vertically-oriented palpebral venation edge of the forearm and the foot, the heel often having a across the nictitating membrane and in having adult small flap of skin. Males are usually considerably smaller males without vocal slits. than females. Frogs in the subgenus Magnummanibus subgen. nov. are Species within Badiohyla gen. nov. are separated from separated from those in the nominate subgenus the other five genera (Nyctimystes, Nigreosoculus gen. Nyctimystes and the subgenus Asperohyla subgen. nov. nov., Occultatahyla gen. nov., Magnumoculus gen. nov. by their possession of very long, prominent heel lappets. and Albogibba gen. nov.) by having a unique “reticulated” Frogs in the subgenus Asperohyla subgen. nov. are palpebral venation, half-webbed fingers and a size in separated from the other two subgenera by having a males with SVL to maximum of 50 mm to 100 mm completely hidden tympanum and a dorsal skin depending on the species. They are further defined as roughened by tiny, conical asperities rather than by follows: The snout is relatively short, blunt and high (E-N/ variable sized and more rounded irregularities as seen in IN 0.94), with distinct canthus rostralis and oblique loreal some other New Guinea tree frog species; iris is brown in region. The vomerine teeth are in two patches between Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 168 Australasian Journal of Herpetology life. Albogibba subgen. nov.), or: Species within Occultatahyla gen. nov. are separated 2/ As above, but male SVL of up to 80 mm; not including from the other five genera (Nyctimystes, Nigreosoculus a spine-like process on the anterior surface of the gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. proximal part of the humerus (upper arm) in males and and Albogibba gen. nov.) by large size (adult females 50 with a unique dorsal pattern of black vermiculations on to 84 mm. in snout to vent length); internarial distance the body and limbs, with unmarked greyish lower flanks distinctly smaller than distance from eye to naris (E-N/IN (Ratiobrunneis subgen. nov.). averages 1.34 (range 1.2-1.5)); the head is broad, Distribution: New Guinea, including offshore islands on flattish, with depressed snout, closely spaced nares, and the continental shelf. oblique loreal region. Palpebral venation is in oblique Content: Badiohyla gen. nov. (type species); lines (more vertical than horizontal and with few Magnumoculus gen. nov.. horizontal interconnections), with a few horizontal PELODRYANINI TRIBE NOV. interconnections. A dermal fold passes from the posterior LSIDurn:lsid:zoobank.org:act:35805C8F-3448-4565- corner of the eye over the upper edge of the tympanum 995C-D9BFA707C3CD and down to the insertion of the forelimb, sometimes becoming obscure when leaving the tympanum. The Type genus: Pelodryas Günther, 1858. outer fingers are approximately half-webbed, the third Diagnosis: The tribe Pelodryanini tribe nov. is best and fifth toes webbed to the disc. A light-coloured dermal diagnosed by way of defining each of the three ridge or series of tubercles extends from the elbow to the component genera. disc of the fourth finger. A similar but less distinct ridge or Species in the genus Pelodryas Günther, 1858 are line of tubercles occupies the outer edge of the tarsus readily separated from all other Australasian Tree Frogs and fifth toe. A very small heel tubercle is present. The (Pelodryadidae) by the following unique suite of skin of the dorsum is smooth to very slightly granular, that characters: of the venter is always coarsely granulate. Slit-like, vocal Colour in life is usually a near unform emerald-green in sac openings are usually present in the floor of the life, but varies from dark purplish-green to fawn or at mouth. A small patch of very fine horny tubercles is found times even blue, often with scattered white or yellow on the first finger of males. The iris is dark or black in spots or dots; there is no obvious white stripe running colour. along the edge of the lower jaw; the hind side of thighs Exceptional to the preceding is the subgenus lacks black or yellow marbling, either being plain or Planusrususpes subgen. nov. which conforms to the sometimes pinkish or orangeish; hind edge of forearm is above genus diagnosis, save for the following smooth, or with at most a few, low, discontinuous differences: The heel is without a tubercle, or in rare tubercles; the skin on top is smooth or slightly leathery; cases has a very small one. The skin is smooth above, venter is white and coarsely granular; hind edge of foot is granular below. A weak, wavy fold is present on the outer smooth; fingers have conspicuous webbing, but reaching surface of the forearm. The head is broader than long; no further than the base of the penultimate phalanx of the the canthus rostralis is not distinct; the loreal region is fourth finger; toes are about 2/3 webbed; finger and toe oblique; Unique to this subgenus is that the palpebral discs are large and obvious; vomerine teeth present and venation is distinct, but is reduced to individual pigment prominent, between and behind the choanae; there is spots and thin, meandering lines, although the lines are obvious thick, supratympanic gland and the tympanum is still mainly oriented largely in a vertical direction, with few large and obvious. horizontal interconnections. Frogs in the genus Shireenhoserhylea gen. nov. are Species within Nigreosoculus gen. nov. are separated readily separated from all other Australasian Tree Frogs from the other five genera (Nyctimystes, Occultatahyla (Pelodryadidae) by the following unique suite of gen. nov., Magnumoculus gen. nov., Badiohyla gen. nov. characters: Colour in life in normal conditions is usually a and Albogibba gen. nov.) by the following suite of unform emerald green above, but occasionally may be characters: Vocal slits present; a very small heel tubercle; fawn, green, purple-green or blue. Hind side of thighs basal webbing on the hand; exposed tympanum; vertical more or less unform and without black and yellow lines of palpebral reticulum oriented obliquely and with marbling or spots. Hind edge of forearms are smooth or relatively few horizontal cross-connections; pale-tan to with at most a few low discontinuous tubercles; hind edge near black iris; rear of thighs barred/mottled with brown, of foot is smooth. Fingers with conspicuous webbing that caramel, or blue-gray. reaches at least as far as the punultimate phalanx of the Species within Albogibba gen. nov. are separated from fourth finger; Vomerine teeth present. the other five genera (Nyctimystes, Nigreosoculus gen. Frogs in the the nominate subgenus Shireenhoserhylea nov., Occultatahyla gen. nov., Magnumoculus gen. nov. gen. nov. are readily separated from those in the and Badiohyla gen. nov.) by one or other of the following subgenus Emeraldhyla subgen. nov. by having an iris, or unique suites of characters: iris immediately above and below the pupil that is a 1/ Being a large species (males to 100 mm. in snout to brilliant red or bright orange in colour and whitish vent length); of a uniform dorsal coloration (green in life, underneath, versus an iris that is yellow or dull orange in purple in alcohol-preserved specimens); palpebral colour and strongly yellow to orange underneath in venation is a reticulum without obvious orientation and in Emeraldhyla subgen. nov.. the nominate subgenus of this genus, the male bears a The genus Summaviridis gen. nov. is monotypic for a spine-like process on the anterior surface of the proximal West Papuan species, S. vagabunda that does not part of the humerus (upper arm) (nominate subgenus appear to have close affinities to any other species or Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 169

genus. Tyler (1968) placed it in a group on its own, The species within the morphologically similar genus although Menzies (2006) associated it with his so-called Rotundaura gen. nov. are separated from this genus “Litoria gracielenta complex”. However S. vagabunda has (Summaviridis gen. nov.) by the tympanum being fully unwebbed hands, versus heavily webbed in the other exposed and round, versus the upper surface being cut species, which in effect scuttles any close association at at the rear by a skin fold forming a straight line and a the genus level. significantly blunter snout in Rotundaura gen. nov.. Summaviridis gen. nov. is therefore only tentatively Distribution: Most parts of continental Australia, except placed in the tribe Pelodryanini tribe nov. and subtribe the coldest regions as well as most parts of New Guinea. Shireenhoserhylina subtribe nov.. Content: Pelodryas Günther, 1858 (type genus); Summaviridis gen. nov. are readily separated from all Shireenhoserhylea gen. nov.; Summaviridis gen. nov.. other Australasian (Australian and New Guinea) Tree SHIREENHOSERHYLINA SUBTRIBE NOV. Frogs (Pelodryadidae) by the following unique suite of LSIDurn:lsid:zoobank.org:act:5315C7D8-DADE-4350- characters: A broad head, short limbs, unwebbed fingers, BB70-8F144939DD39 a prominent fold of skin across the chest and (in life) a Type genus: Shireenhoserhylea gen. nov. dark green dorsal colouration. Diagnosis: The subtribe Shireenhoserhylina subtribe In further detail the Summaviridis gen. nov. is separated nov. is best diagnosed by way of defining each of the two from all other Australasian (Australian and New Guinea) component genera. Tree Frogs (Pelodryadidae) as follows: The head is Frogs in the genus Shireenhoserhylea gen. nov. are broader than long (HL/HW 0.951), its length slightly less readily separated from all other Australasian Tree Frogs than one-third of the snout to vent length (HL/S-V 0.310). (Pelodryadidae) by the following unique suite of The snout is rounded when viewed from above and in characters: profile. The nostrils are more lateral than superior, their distance from the end of the snout slightly less than that Colour in life in normal conditions is usually a unform from the eye. The distance between the eye and the naris emerald green above, but occasionally may be fawn, is greater than the internarial span (E-N/IN 1.154). The green, purple-green or blue. Hind side of thighs more or canthus rostralis is straight and poorly defined. The eye less unform and without black and yellow marbling or is large, its diameter greater than the distance separating spots. Hind edge of forearms are smooth or with at most it from the nostril. Three-quarters of the tympanum is a few low discontinuous tubercles; hind edge of foot is visible, the superior rim of the tympanic annulus is hidden smooth. Fingers with conspicuous webbing that reaches beneath the supra-tympanic fold. The tympanal diameter at least as far as the punultimate phalanx of the fourth is equivalent to two-thirds of the eye diameter. The finger; Vomerine teeth present. vomerine teeth are in two broadly oval series situated Frogs in the the nominate subgenus Shireenhoserhylea between the choanae. The tongue is broadly cordiform gen. nov. are readily separated from those in the with a deeply indented posterior border. The fingers are subgenus Emeraldhyla subgen. nov. by having an iris, or very long and slender with narrow lateral fringes; in iris immediately above and below the pupil that is a decreasing order of length 3>4>2>1; unwebbed. The brilliant red or bright orange in colour and whitish terminal discs and sub-articular tubercles are large and underneath, versus an iris that is yellow or dull orange in prominent. The hind limbs are short and slender with a colour and strongly yellow to orange underneath in TL/S-V ratio of 0.513. Toes in decreasing order of length Emeraldhyla subgen. nov.. 4>5>3 >2>1. The webbing between the outer and fourth The genus Summaviridis gen. nov. is monotypic for a toe reaches half-way up the penultimate phalanx on the West Papuan species, S. vagabunda that does not fifth toe, and the sub-articular tubercle at the base of the appear to have close affinities to any other species or penultimate phalanx on the fourth. The skin on the dorsal genus. Tyler (1968) placed it in a group on its own, surfaces is smooth with a few minute flattened tubercles. although Menzies (2006) associated it with his so-called The throat and chest are slightly tubercular; the “Litoria gracielenta complex”. However S. vagabunda has abdomen, sides of the body and lower surface of the unwebbed hands, versus heavily webbed in the other thighs are extremely granular. There is a conspicuous species, which in effect scuttles any close association at curved supra-tympanic fold, and a prominent skin fold the genus level. across the chest. The dorsal surfaces are in life dark Summaviridis gen. nov. are readily separated from all green and in preservative dark blue. There is a faint trace other Australasian (Australian and New Guinea) Tree of a narrow white line above a broader brown line on the Frogs (Pelodryadidae) by the following unique suite of canthus rostralis, upper eyelid and supra-tympanic fold. characters: A broad head, short limbs, unwebbed fingers, The backs of the thighs are pale brown and marked with a prominent fold of skin across the chest and (in life) a small irregularly-shaped, pale yellow spots. The backs of dark green dorsal colouration. the tarsus and forearm are pale brown. The ventral In further detail the Summaviridis gen. nov. is separated surfaces are pale yellow with leaden blue patches at the from all other Australasian (Australian and New Guinea) angles of the jaws, and small faint brown spots on the Tree Frogs (Pelodryadidae) as follows: The head is remainder of the mandibular border and on the throat. broader than long (HL/HW 0.951), its length slightly less Adult females are about 37 mm in body length (modified than one-third of the snout to vent length (HL/S-V 0.310). from Tyler, 1978). The snout is rounded when viewed from above and in Specimens of the morphologically similar Llewellynura profile. The nostrils are more lateral than superior, their Wells and Wellington, 1985 are readily separated by their distance from the end of the snout slightly less than that much smaller adult size of less than 25 mm body length. from the eye. The distance between the eye and the naris Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 170 Australasian Journal of Herpetology is greater than the internarial span (E-N/IN 1.154). The PUSTULATARANINI TRIBE NOV. canthus rostralis is straight and poorly defined. The eye LSIDurn:lsid:zoobank.org:act:C57E50A3-84EC-4921- is large, its diameter greater than the distance separating A71A-A0F1B28D4305 it from the nostril. Three-quarters of the tympanum is Type genus: Pustulatarana gen. nov. visible, the superior rim of the tympanic annulus is hidden Diagnosis: The tribe Pustulataranini tribe nov. is best beneath the supra-tympanic fold. The tympanal diameter diagnosed by way of defining each of the seven is equivalent to two-thirds of the eye diameter. The component genera. vomerine The single known living species within the genus teeth are in two broadly oval series situated between the Pustulatarana gen. nov. is readily separated from all choanae. The tongue is broadly cordiform with a deeply other Australasian Tree Frogs (Pelodryadidae) by the indented posterior border. The fingers are very long and following suite of characters: slender with narrow lateral fringes; in decreasing order of A smallish sized tree frog being 27 mm in body length. length 3>4>2>1; unwebbed. The terminal discs and sub- Brown to greenish brown or even yellow above, with articular tubercles are large and prominent. The hind obscure and irregular darker flecks and blotches. limbs are short and slender with a TL/S-V ratio of 0.513. Creamy-white below, finely peppered with dark blackish- Toes in decreasing order of length 4>5>3 >2>1. The brown or shading of other lighter colour on the throat, webbing between the outer and fourth toe reaches half- with heavy concentrations of stippling on chin, chest and way up the penultimate phalanx on the fifth toe, and the flanks of some specimens. Underside of thighs have sub-articular tubercle at the base of the penultimate irregular darker patches and hind isde of thigh has phalanx on the fourth. The skin on the dorsal surfaces is irregular fine creamish coloured stripes. Skin is leathery smooth with a few minute flattened tubercles. The throat and with numerous scattered tubercles which may or not and chest are slightly tubercular; the abdomen, sides of be arranged in well-defined longitudinal rows, including the body and lower surface of the thighs are extremely sometimes some of medium to large size and a granular. There is a conspicuous curved supra-tympanic prominent one on the eyelid. Belly is smooth except for fold, and a prominent skin fold across the chest. The some granular skin on the lower belly and thighs. dorsal surfaces are in life dark green and in preservative Vomerine teeth present, but weakly developed and dark blue. There is a faint trace of a narrow white line between the choanae. Fingers lack webbing but have above a broader brown line on the canthus rostralis, large oval discs. Toes poorly to moderately webbed. A upper eyelid and supra-tympanic fold. The backs of the minute rounded outer metatarsal tubercle and a thighs are pale brown and marked with small irregularly- moderate-zied oval inner metatarsal tubercle. Tympanum shaped, pale yellow spots. The backs of the tarsus and distinct. forearm are pale brown. The ventral surfaces are pale yellow with leaden blue patches at the angles of the jaws, Pustulatarana gen. nov. are separated from the genus and small faint brown spots on the remainder of the Llewellynura Wells and Wellington, 1985 by the large mandibular border and on the throat. Adult females are oval discs on the forelimbs and larger body size (27 mm about 37 mm in body length (modified from Tyler, 1978). vs 20 mm). Pustulatarana gen. nov. are from the genus Mahoneybatrachus Wells and Wellington, 1985, by Specimens of the morphologically similar Llewellynura having reduced toe webbing, versus well developed Wells and Wellington, 1985 are readily separated by their webbing on the feet. much smaller adult size of less than 25 mm body length. Species within the genus Llewellynura Wells and The species within the morphologically similar genus Wellington, 1985 are readily separated from all other Rotundaura gen. nov. are separated from this genus Australasian Tree Frogs (Pelodryadidae) by the following (Summaviridis gen. nov.) by the tympanum being fully suite of characters: they are tiny in size, being about 20 exposed and round, versus the upper surface being cut mm in body length as adults. Colour is variegated dark at the rear by a skin fold forming a straight line and a and brown dorsally; a row of widely spaced dorsolateral significantly blunter snout in Rotundaura gen. nov.. tubercles and ridges may be on each side, or Distribution: Wetter parts of the east coast of Australia, alternatively just a few randomly scattered small excluding colder parts of the far south, New Guinea and tubercles on either side of upper flank; no pectoral fold; offshore islands. The nominate subgenus several conspicuous tubercles above each eye; tiny, Shireenhoserhylea subgen. nov. is restricted to the East slender and agile and with a sharply pointed snout. The Coast of Australia, south of the northern wet tropics in dorsum may be grey, brown, yellow or red, often flecked Queensland. or mottled with darker colour. There is usually a broad The subgenus Emeraldhyla subgen. nov. occurs in New dark band from behind the eye to the groin and an even Guinea and offshore islands as well as the far north of darker stripe along the snout, through the eye and over Cape York, Queensland. the base of the arm to the flank. Snout and upper The type species for the monotypic genus Summaviridis surfaces of the limbs are peppered with dark brown. gen. nov. is only known from the two types, a male and a There is a conspicuous dark bar along the front edge of female, being from Vogelkop Peninsula, Papua (New the arm. Ventral surface is white except for a fine Guinea) (female) and Seram (= Ceram) Island, Molucca peppering of brown on the throat, chest and limbs. Skin is Islands, Indonesia. smooth dorsally, with at least some small tubercles or Content: Shireenhoserhylea gen. nov. (type species); warts or skin folds and usually several small subercles Summaviridis gen. nov.. over each eye. Throat skin is smooth, but belly is granular. Finger and toe discs are moderate in size but

distinct. Fingers lack webbing and toes are less than half choanae. Finger and toe discs are moderate, wider than webbed. The disc and nearly two phlanges are free on the digits. Fingers lack webbing, toes being about half the outer side of the fourth toe. The inner metatarsal webbed, including not reaching the disc of the fifth toe tubercle is prominent and there is no outer one. but extending no more than half way along the Tympanum ranges from small to indistinct. Second finger penultimate phalanx. Second finger is longer than the longer than first. first. There is a small oval inner metatarsal tubercle and a The Australian species within Llewellynura being of the smaller outer metatarsal tubercle. Tympanum distinct. nominate subgenus Llewellynura are separated from the The genus Litoria, with type species L. freycineti Tschudi, New Guinea species herein placed in the subgenus 1838 is herein restricted to the type species and the Microlitoria subgen. nov. by lacking vomerine teeth. closely related L. latopalmata Günther, 1867, which as a Known as the “Rock Hole Frogs”, living frogs in the genus pair diverged from their nearest common ancestor 13.3 Mahonabatrachus Wells and Wellington, 1985 are all MYA according to Duellman et al. (2016). readily separated from all other Australasian Tree Frogs This divergence and morphological differences are (Pelodryadidae) by the following unique suite of exactly why the species previously included in Litoria characters: In colour they are usually irregularly mottled have been assigned to other genera. above with metallic fawn, brown or reddish brown and Litoria as defined herein, are readily separated from all dark brown, the small low tubercles on the back other Australasian Tree frogs (Pelodryadidae) by the sometimes tending to be light-centered and/or dark following suite of characters: Species are a frog that is edged. Sometimes these frogs are dominantly dark fawn to dark above, being either immaculate or with brown but with a gold-orange dorsolateral stripe from markings on the back, with or without a warty exterior, the snout to eye and continuing above the eye above the warts if present being small and flattish, the markings if tympanum along the body almost to the groin. Limbs of present typically being a series of darker blotches or all specimens are always coloured with irregular dark variegations in either an irregular or regular pattern. brown cross bands and all frogs have barred or spotted There is a broad dark, canthal stripe, almost completely lips to some degree and lower surfaces are whitish. interrupted in front of the eye and continues behind the The skin is leathery to finely granular above with eye to effectively overwrite the tympanum and extend to numerous low, rounded or sometimes slightly pointed the flanks where it invariably breaks up into a series of tubercles, that are fairly evenly spaced. No pectoral fold. black spots or blotches extending ro the groin. There is a Moderate sized but conspicuous diss on fingers and toes. pale glandular stripe from below the eye to the base of Fingers free and toes with moderately developed the forelimb. Limbs are variegated darker but usually with webbing, including reaching the disc of the fifth toe. an irregular pattern of spots, blotches or bars. Lower jaw Adults average 20 mm in length. is variegated with yellow and dark brown, often forming a The three species in the genus Salmocularana gen. nov. reticulum, often with a barred appearance. The venter is are small species that breed in escarpment rock hills in whitish. Ventral surface and flanks are granular. There is Arnhemland, Northern Territory and also the Kimberley no dorsolateral skin fold. Finger and toe discs are small Ranges of north-west Australia. and project slightly but noticeably, beyond the lateral The three known living species within the genus edges of the penultimate phlanges. Fingers are free, Salmocularana gen. nov. (one formally described within without webbing. Toes have well developed webbing, the this paper) are readily separated from all other webbing not reaching the disc of the fifth toe and Australasian Tree Frogs (Pelodryadidae) by the following extending no more than half way along the penultimate unique suite of characters: Pale grey-brown, fawn, phalanx. There is a small inner and minute outer reddish, orange or pink above, with dorsal markings metatarsal tubercle. Vomerine teeth are in two clusters ranging from nothing more than slight peppering at the between the choanae. The tympanum is distinct and anterior part of the dorsum as seen in Salmocularana adults average 40-45 mm snout to rear. personata (Tyler, Davies and Martin, 1978), indistinct The genus Paralitoria gen. nov. diverged from its nearest marbling and blotches as seen in S. staccato (Doughty common ancestor, being that of the genera Litoria and and Anstis, 2007) or an intense randomised configuration Quasilitoria gen. nov. being the most closely related of dark brown pigment on a light beige to grey genera, some 15.5 MYA according to Duellman et al. background in S. saxacola sp. nov.; specimens have a (2016). distinct or semi-distinct dark brownish or purple head Paralitoria gen. nov. is separated from all other stripe from just in front of the nostril, through the eye and Australasian tree frogs (Pelodryadidae) by the following ear to upper flank just behind the forelimbs. Lips are suite of characters, being one or other of the following: whitish to some extent, ranging from a distinct yellow bar, 1/ Fawn, grey or brown above, being immaculate; a to merely light and peppered brown. Ventral surfaces conspicuous black stripe from the snout, through the whitish. Skin may be smooth, with extremely tiny nostril to the eye, where it is almost entirely broken by a scattered tubercles above as in S. personata, or with a pale vertical bar in front of the eye as a band continuing very limited number of scattered small tubercles above below the tympanum and extending back to the forearm the arm and on the upper flank and nearby dorsum in S. with minimal loss of width along the length, where after a staccato to scattered medium sized tubercles above the break, it continues obliquely along the mid flank. There is arm and on the upper flank and nearby dorsum in S. a pale glandular stripe from below the eye to the forearm. saxacola sp. nov.. Underneath the skin is coarsely There is a conspicuous dark brown stripe along the front granular, except on the throat, where it is smooth. edge of the tibial region. Lips are not spotted or barred. Vomerine teeth are in two short clumps between the The groin and hind side of thighs are pale yellow, spotted Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 172 Australasian Journal of Herpetology and variegated with dark brown. Venter is whitish. Dorsal eye, the yellow bar being blocked at the top by a small surface is smooth and leathery, throat is smooth and the and well-defined area of black. Forefingers light yellowish venter granular. Vomerine teeth are between the or white, peppering on the light surfaces of the snout. choanae. Finger discs are small and distinct, while toe Tympanum has a distinctive pale rim (Q. nasuta, Q. discs are smaller and indistinct. Fingers are unwebbed peninsulae) (subgenus Vultusamolitoria subgen. nov.). and toes have moderate webbing between them. There is Distribution: Most parts of Australia and New Guinea, a small inner and tiny outer metatarsal tubercle. although the majority of species are in northern Australia. Tympanum is distinct and second finger is slightly shorter Content: Pustulatarana gen. nov. (type species); than the first. Body length averages 50 mm (subgenus Llewellynura Wells and Wellington, 1985; Paralitoria subgen. nov.), or: Mahonabatrachus Wells and Wellington, 1985; 2/ Fawn, grey or brown above, with limited flecks or other Salmocularana gen. nov.; Litoria Tschudi, 1838; markings, often as a peppering; a conspicuous narrow Paralitoria gen. nov.; Quasilitoria gen. nov.. black stripe from the snout, through the nostril to the eye, SALMOCULARANINA SUBTRIBE NOV. continuing through the tympanum (effectively over-writing LSIDurn:lsid:zoobank.org:act:F925B9F0-D4E1-4663- it) and extending past the forearm with minimal loss of A618-8FC4120624B0 width along the length to the anterior flank. There is a Type genus: Salmocularana gen. nov. pale glandular stripe from below the eye to the forearm. There is a conspicuous dark brown stripe along the front Diagnosis: The subtribe Salmocularanina subtribe nov. edge of the tibial region. The lower lips are faintly or is best diagnosed by way of defining each of the four partially barred. The groin and hind side of thighs are component genera. pale yellow, spotted and variegated with dark brown. The three species in the genus Salmocularana gen. nov. Venter is whitish. Dorsal surface is smooth and leathery, are small species that breed in escarpment rock hills in throat is smooth and the venter granular. Vomerine teeth Arnhemland, Northern Territory and also the Kimberley are between the choanae. Finger discs are small and Ranges of north-west Australia. barely distinct, while toe discs are of similar size. Fingers The three known living species within the genus are unwebbed and toes extensive webbing between Salmocularana gen. nov. (one formally described within them, with the webbing reaching the disc of the fifth toe. this paper) are readily separated from all other There is a small inner and tiny outer metatarsal tubercle. Australasian Tree Frogs (Pelodryadidae) by the following Tympanum is distinct and second finger is slightly shorter unique suite of characters: than the first. Body length averages 75 mm (subgenus Pale grey-brown, fawn, reddish, orange or pink above, Ferelitoria subgen. nov.). with dorsal markings ranging from nothing more than Species of Quasilitoria gen. nov. are separated from all slight peppering at the anterior part of the dorsum as other Australasian tree frogs (Pelodryadidae) by the seen in Salmocularana personata (Tyler, Davies and following suite of characters: Martin, 1978), indistinct marbling and blotches as seen in The dorsum is smooth or with some scattered low round S. staccato (Doughty and Anstis, 2007) or an intense tubercles; front toe discs are no wider than the randomised configuration of dark brown pigment on a penultimate phalanx and toe discs only slightly wider, but light beige to grey background in S. saxacola sp. nov.; otherwise also small. Front edge of thigh has either a specimens have a distinct or semi-distinct dark brownish continous black stripe or alternatively an arrangement of or purple head stripe from just in front of the nostril, broken black spots or blotches. Lower surfaces white, through the eye and ear to upper flank just behind the throat with or without some mottling, smooth on the throat forelimbs. Lips are whitish to some extent, ranging from a and chest and slightly granular on the belly. Dorslolateral distinct yellow bar, to merely light and peppered brown. folds are either absent or very indistinct. Inner metatarsal Ventral surfaces whitish. Skin may be smooth, with tubercle is small and outer one is tiny. Prominent extremely tiny scattered tubercles above as in S. vomerine teeth. Fingers unwebbed, while toes are half to personata, or with a very limited number of scattered three quarters webbed. small tubercles above the arm and on the upper flank Furthermore, one or other of the following character and nearby dorsum in S. staccato to scattered medium suites: sized tubercles above the arm and on the upper flank and nearby dorsum in S. saxacola sp. nov.. Underneath 1/ Back is without darker markings or mottling, at most the skin is coarsely granular, except on the throat, where there being a slight peppering on the upper surfaces; it is smooth. Vomerine teeth are in two short clumps anterior black head stripe when present is interrupted at between the choanae. Finger and toe discs are least in part by a slight vertical bar in front of the eye; moderate, wider than the digits. Fingers lack webbing, posterior dark head stripe is as wide as and overwriting toes being about half webbed, including not reaching the the tympanum or not quite so and if not, then the lower disc of the fifth toe but extending no more than half way part of the tympanum is not blackened, and snout is only along the penultimate phalanx. Second finger is longer moderately pointed, (Q. axillaris, Q. coplandi, Q. inermis, than the first. There is a small oval inner metatarsal Q. pallida, Q. tornieri) (subgenus Quasilitoria subgen. tubercle and a smaller outer metatarsal tubercle. nov.), or alternatively: Tympanum distinct. 2/ With darker markings and/or stripes on the dorsal The genus Litoria, with type species L. freycineti Tschudi, surface, with some longitudinal folds on the back, a 1838 is herein restricted to the type species and the strongly pointed snout and a well defined yellow bar in closely related L. latopalmata Günther, 1867, which as a front of the eye, breaking the black line from snout past pair diverged from their nearest common ancestor 13.3

MYA according to Duellman et al. (2016). markings, often as a peppering; a conspicuous narrow This divergence and morphological differences are black stripe from the snout, through the nostril to the eye, exactly why the species previously included in Litoria continuing through the tympanum (effectively over-writing have been assigned to other genera. it) and extending past the forearm with minimal loss of Litoria as defined herein, are readily separated from all width along the length to the anterior flank. There is a other Australasian Tree frogs (Pelodryadidae) by the pale glandular stripe from below the eye to the forearm. following suite of characters: Species are a frog that is There is a conspicuous dark brown stripe along the front fawn to dark above, being either immaculate or with edge of the tibial region. The lower lips are faintly or markings on the back, with or without a warty exterior, the partially barred. The groin and hind side of thighs are warts if present being small and flattish, the markings if pale yellow, spotted and variegated with dark brown. present typically being a series of darker blotches or Venter is whitish. Dorsal surface is smooth and leathery, variegations in either an irregular or regular pattern. throat is smooth and the venter granular. Vomerine teeth There is a broad dark, canthal stripe, almost completely are between the choanae. Finger discs are small and interrupted in front of the eye and continues behind the barely distinct, while toe discs are of similar size. Fingers eye to effectively overwrite the tympanum and extend to are unwebbed and toes extensive webbing between the flanks where it invariably breaks up into a series of them, with the webbing reaching the disc of the fifth toe. black spots or blotches extending ro the groin. There is a There is a small inner and tiny outer metatarsal tubercle. pale glandular stripe from below the eye to the base of Tympanum is distinct and second finger is slightly shorter the forelimb. Limbs are variegated darker but usually with than the first. Body length averages 75 mm (subgenus an irregular pattern of spots, blotches or bars. Lower jaw Ferelitoria subgen. nov.). is variegated with yellow and dark brown, often forming a Species of Quasilitoria gen. nov. are separated from all reticulum, often with a barred appearance. The venter is other Australasian tree frogs (Pelodryadidae) by the whitish. Ventral surface and flanks are granular. There is following suite of characters: no dorsolateral skin fold. Finger and toe discs are small The dorsum is smooth or with some scattered low round and project slightly but noticeably, beyond the lateral tubercles; front toe discs are no wider than the edges of the penultimate phlanges. Fingers are free, penultimate phalanx and toe discs only slightly wider, but without webbing. Toes have well developed webbing, the otherwise also small. Front edge of thigh has either a webbing not reaching the disc of the fifth toe and continous black stripe or alternatively an arrangement of extending no more than half way along the penultimate broken black spots or blotches. Lower surfaces white, phalanx. There is a small inner and minute outer throat with or without some mottling, smooth on the throat metatarsal tubercle. Vomerine teeth are in two clusters and chest and slightly granular on the belly. Dorslolateral between the choanae. The tympanum is distinct and folds are either absent or very indistinct. Inner metatarsal adults average 40-45 mm snout to rear. tubercle is small and outer one is tiny. Prominent The genus Paralitoria gen. nov. diverged from its nearest vomerine teeth. Fingers unwebbed, while toes are half to common ancestor, being that of the genera Litoria and three quarters webbed. Quasilitoria gen. nov. being the most closely related Furthermore, one or other of the following character genera, some 15.5 MYA according to Duellman et al. suites: (2016). 1/ Back is without darker markings or mottling, at most Paralitoria gen. nov. is separated from all other there being a slight peppering on the upper surfaces; Australasian tree frogs (Pelodryadidae) by the following anterior black head stripe when present is interrupted at suite of characters, being one or other of the following: least in part by a slight vertical bar in front of the eye; 1/ Fawn, grey or brown above, being immaculate; a posterior dark head stripe is as wide as and overwriting conspicuous black stripe from the snout, through the the tympanum or not quite so and if not, then the lower nostril to the eye, where it is almost entirely broken by a part of the tympanum is not blackened, and snout is only pale vertical bar in front of the eye as a band continuing moderately pointed, (Q. axillaris, Q. coplandi, Q. inermis, below the tympanum and extending back to the forearm Q. pallida, Q. tornieri) (subgenus Quasilitoria subgen. with minimal loss of width along the length, where after a nov.), or alternatively: break, it continues obliquely along the mid flank. There is 2/ With darker markings and/or stripes on the dorsal a pale glandular stripe from below the eye to the forearm. surface, with some longitudinal folds on the back, a There is a conspicuous dark brown stripe along the front strongly pointed snout and a well defined yellow bar in edge of the tibial region. Lips are not spotted or barred. front of the eye, breaking the black line from snout past The groin and hind side of thighs are pale yellow, spotted eye, the yellow bar being blocked at the top by a small and variegated with dark brown. Venter is whitish. Dorsal and well-defined area of black. Forefingers light yellowish surface is smooth and leathery, throat is smooth and the or white, peppering on the light surfaces of the snout. venter granular. Vomerine teeth are between the Tympanum has a distinctive pale rim (Q. nasuta, Q. choanae. Finger discs are small and distinct, while toe peninsulae) (subgenus Vultusamolitoria subgen. nov.). discs are smaller and indistinct. Fingers are unwebbed Distribution: Most parts of Australia and parts of and toes have moderate webbing between them. There is southern New Guinea, although the majority of species a small inner and tiny outer metatarsal tubercle. are in northern Australia. Tympanum is distinct and second finger is slightly shorter Content: Salmocularana gen. nov. (type species); Litoria than the first. Body length averages 50 mm (subgenus Tschudi, 1838; Paralitoria gen. nov.; Quasilitoria gen. Paralitoria subgen. nov.), or: nov.. 2/ Fawn, grey or brown above, with limited flecks or other Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 174 Australasian Journal of Herpetology

SAGUNURINI TRIBE NOV. a single, relatively short (0.206-0.379 s) grunt with an LSIDurn:lsid:zoobank.org:act:55665C91-6E14-4F47- unusual clumped pattern of pulses. The single species AE34-E3E87D4FAC05 within the subgenus Parawowrana subgen. nov. is Type genus: Saganura Wells and Wellington, 1985. defined as above, except that it instead is slightly smaller Diagnosis: The tribe Saganurini tribe nov. is monotypic than the preceding species (adult males 50-55 mm), has for the type genus and therefore the tribe diagnosis is the a white labial stripe (in common with the much larger same as for the genus Saganura Wells and Wellington, species in Sandyrana Wells and Wellington, 1985, that 1985. grow to in excess of 100 mm in body length) and has a characteristically longer call than those in the nominate Living frogs in the genus Saganura Wells and Wellington, subgenus (0.69-0.9 second) (Richards et al. 2006). 1985 are all readily separated from all other Australasian Tree Frogs (Pelodryadidae) by the following unique suite Duellman et al. (2016) found that the species in the of characters: genus Wowrana gen. nov. diverged from their nearest living relatives 26.5 MYA, being species within Sandyrana It is a uniform light green above, commonly with scattered Wells and Wellington, 1985. The type form of light fawn spots, or dark brown with irregular bright green Nyctimystes Stejneger, 1916, namely Nyctimantis papua patches and light brown flecks. There is a narrow, black Boulenger, 1897 diverged from this genus 36.1 MYA, canthal streak, widening behind the eye to form a dark making genus level assignment of Wowrana gen. nov. an broad band extending almost to the groin; this dark band obvious choice. often broken up by groups of irregular white blotches and/or spots. There is an obscure, narrow light green Frogs in the genus Sandyrana Wells and Wellington, zone along the supratympanic ridge. There is no pale line 1985 are readily separated from all other Australasian along the posterior edge of the upper jaw. Venter is Tree Frogs (Pelodryadidae) by the following unique suite pinkish-white, throat darker with a greyish tinge. Groin of characters: Large frogs, which as adults can grow to in and hind side of thighs is a uniform pale brown. Dorsal excess of 100 mm. Dorsal colour of adults in life is surface smooth or with scattered small tubercles. Chin usually uniform green above in normal circumstances, smooth and other lower surfaces are granular. Vomerine but may otherwise range from fawn, through green, teeth are between or behind the choanae, with hind edge purplish or even blue. Fingers with conspicuous webbing, of vomerine teeth always further back than the choanae. reaching at least as far as the base of the penultimate Prectoral fold present. Finger and toe discs large. Fingers phalanx of the fourth finger. Hind edge of forearm is with conspicuous basal webbing, being nearly a third smooth, or with at most a few low, discontinuous webbed, toes are about three quarters webbed. There is tubercles. Hind edge of feet is smooth. Hind side of a large inner and small outer metatarsal tubercle. thighs is more or less uniform, without black and yellow Tympanum is distinct. Second finger is much longer than marbling, spots or blotches. first; when pressed together, the tip of the first finger Webbing reaching no further than the base of the reaches no further than the base of the disc of the penultimate phalanx of the fourth finger. A distinctive second finger. Heel of adpressed hindlimb reaches to the white or pink stripe along the edge of the lower jaw, eye or beyond. Adults attain about 55 mm in total length extending back to the level of the forelimb and not in a (adapted and modified from Cogger 2014). configuration of blobs, spots or random markings According to Duellman et al. (2016), the species in this otherwise tending to be in a linear manner. Vomerine genus diverged from their nearest living relatives 27.6 teeth present. MYA being more than sufficient justification for erecting Distribution: All species within Wowrana gen. nov. are this new tribe. confined to forested locations on the island of New Distribution: South-west Tasmania, including highlands Guinea and in general most species are currently known and coast. from relatively few specimens. Content: Sagunura burrowsae (Scott, 1942) (monotypic). Species within the genus Sandyrana Wells and Wellington, 1985 are found throughout New Guinea and WOWRANAINI TRIBE NOV. west as far as Halmahera Island as well as New Britain, LSIDurn:lsid:zoobank.org:act:5756C747-7B83-47E0- New Ireland, Aru Islands and north-east Queensland, 95CE-F3AB9E9C3341 Australia. Type genus: Wowrana gen. nov. Content: Wowrana gen. nov. (type genus); Sandyrana Diagnosis: The tribe Wowranaini tribe nov. includes two Wells and Wellington, 1985. genera. It is best defined by diagnosing each, noting that SANDYRANINA SUBTRIBE NOV. each genus also occupies one subtribe each. LSIDurn:lsid:zoobank.org:act:4B9B0AAD-FDD5-4EE7- Frogs in the genus Wowrana gen. nov. in the nominate AC3B-DFE5893944C8 subgenus are readily separated from all other Type genus: Sandyrana Wells and Wellington, 1985. Australasian (Australian and New Guinea) Tree Frogs (Pelodryadidae) by the following unique suite of Diagnosis: The subtribe Sandyranina subtribe nov. is characters: monotypic for the type genus and therefore the tribe diagnosis is the same as for the genus Sandyrana Wells Large size (adult males over 60 mm); uniform bright and Wellington, 1985. green dorsal colouration in life (blue in preservative). Fully webbed hands. Frogs in the genus Sandyrana Wells and Wellington, 1985 are readily separated from all other Australasian No prominent and enlarged parotoid glands, no white Tree Frogs (Pelodryadidae) by the following unique suite labial stripe, SVL not over 85 mm and a call consisting of of characters: Large frogs, which as adults can grow to in Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 175

excess of 100 mm. Dorsal colour of adults in life is Litoria lesueuri (Dumeril and Bibron, 1841) (Anura: usually uniform green above in normal circumstances, Hylidae) and comments on population declines of L. but may otherwise range from fawn, through green, booroolongensis. Transactions of the Royal Society of purplish or even blue. Fingers with conspicuous webbing, South Australia 122: 33-44. reaching at least as far as the base of the penultimate Anstis, M., Tyler, M. J., Roberts, J. D., Price, L. C. and phalanx of the fourth finger. Hind edge of forearm is Doughty, P. 2010. A new species of Litoria (Anura: smooth, or with at most a few low, discontinuous Hylidae) with a highly distinctive tadpole from the north- tubercles. Hind edge of feet is smooth. Hind side of western Kimberley region of Western Australia. Zootaxa thighs is more or less uniform, without black and yellow (online PRINO)2550:39-57. marbling, spots or blotches. Webbing reaching no further Anstis, M., Rowley, J. J. L. and Altig. R. I. 2016a. than the base of the penultimate phalanx of the fourth Morphological clarifications of Australian hylid and finger. A distinctive white or pink stripe along the edge of limnodynastid tadpoles. Zootaxa (online PRINO) the lower jaw, extending back to the level of the forelimb 4126:146-150. and not in a configuration of blobs, spots or random Anstis, M., Price, L. C., Dale Roberts, J., Catalano, S. R., markings otherwise tending to be in a linear manner. Hines, H. B., Doughty, P. and Donnellan, S.C. 2016b. Vomerine teeth present. Revision of the water-holding frogs, Cyclorana Duellman et al. (2016) found that the species within the platycephala (Anura: Hylidae), from arid Australia, genus Sandyrana diverged from their nearest living including a description of a new species. Zootaxa (Online relatives 26.5 MYA making subtribe level classification for PRINO), 4126(4):451-479. the group wholly appropriate. Barbour, T. 1908. Some new reptiles and amphibians. Distribution: Throughout New Guinea and west as far as Bulletin of the Museum of Comparative Zoology. Harvard, Halmahera Island as well as New Britain, New Ireland, Cambridge, Massachusetts 51:315-325. Aru Islands and north-east Queensland, Australia. Barbour, T. 1912. A contribution to the zoogeography of Content: Sandyrana Wells and Wellington, 1985. the East Indian Islands. Memoirs of the Museum of SUMMARY Comparative Zoology. Harvard, Cambridge, For the first time ever, the Australian and New Guinean Massachusetts 44:1-203. frog family Pelodryadidae have a robust classification at Barbour, T. 1921. Reptiles and amphibians from the all levels. The classification of the group has effectively British Solomon Islands. Proceedings of the New been brought up to date. England Zoölogical Club. Cambridge, Massachusetts Well known but hitherto unnamed species have been 7:91-112. formally named, which is an important first step for some Barker, J. and Grigg, G. C. 1977. A field guide to which are already facing extinction. Australian Frogs. Rigby, Australia:229 pp. The tribe and genus-level classification of the family is Barker, J., Grigg, G. C. and Tyler, M. J. 1995. A Field also based on simple logic and common sense, which Guide to Australian Frogs. Edition 2. Surrey Beatty and also happen to be core principles of science. When the Sons, Chipping Norton, Australia. smoke, mirrors and lies of the Wolfgang Wüster gang of Bell, R. C., MacKenzie, J. B., Hickerson, M., Chavarría, thieves clears after the death of the key members of the K. L., Cunningham, M. J., Williams, S. R. and Moritz, C. gang, there is little doubt that the classification within this 2012. Comparative multi-locus phylogeography confirms paper will be derided as both logical and self evident. multiple vicariance events in co-distributed rainforest One hopes that for the sake of the frogs themselves and frogs. Proceedings of the Royal Society of London. their ultimate long-term survival, as outlined by Hoser Series B, Biological Sciences 279:991-999. (2019a, 2019b), that the classification system within this Bell, R. C., Webster, G. N. and Whiting, M. J. 2017. paper is in fact adopted sooner, rather than later. Breeding biology and the evolution of dynamic sexual REFERENCES CITED dichromatism in frogs. Journal of Evolutionary Biology Ahl, E. 1929. Beschreibung eines neuen Laubfrosches 30(12):2104-2115. der Gattung Hyla von Java. Zoologischer Anzeiger Bevelander, J. 2014. Persistence of the Eastern Dwarf 85:269-271. Tree Frog (Litoria fallax) in Melbourne: habitat Ahl, E. 1935. Beschreibung eines neuen Laubfrosches characteristics and climatic niche. MSC Thesis, aus Südaustralien. Zoologischer Anzeiger 109:252-253. University of Melbourne, Australia. Andersson, L. G. 1916. Results of Dr. E. Mjöberg’s Boettger, O. 1895. Liste der Reptilien und Batrachier der Swedish scientific expeditions to Australia 1910-1913. I. Insel Halmaheira nach den Sammlungen Prof. Dr. W. 9. Batrachians from Queensland. Kongliga Svenska Kükenthal’s. Zoologischer Anzeiger 18:129-138. Vetenskaps-Akademiens Handlingar 52:1-20. Boettger, O. 1900. Die Reptilien und Batrachier. Anstis, M. 2013. Tadpoles and frogs of Australia. Reed / Abhandlungen der Senckenbergischen Naturforschenden New Holland, Sydney, Australia:829 pp. Gesellschaft, Frankfurt am Main 25:325-400. Anstis, M. and Tyler, M. J. 2005. Breeding biology of Böhme, W. 2014. Herpetology in Bonn. Mertensiella. Litoria microbelos (Cogger) (Anura: Hylidae). Deutsche Gesellschaft für Herpetologie und Transactions of the Royal Society of South Austalia Terrarienkunde 21:i–vi+1-262. 129:43-48. Böhme, W. and Bischoff, W. 1984. Die Anstis, M., Alford, R. and Gillespie, G. 1998. Breeding Wirbeltiersammlungen des Museums Alexander Koenig. biology of Litoria booroolongensis (Moore, 1961), and III. Amphibien und Reptilien. Bonner Zoologische

Monographien 19:151-213. Natural History, Series 8, 16:402-404. Bolt, A. 2020. George Floyd and the US race riots: Be Brongersma, L. D. 1953. Notes on New Guinean reptiles Warned! Herald-Sun, 31 May, posted online at: and amphibians. III. Proceedings of the Koninklijke https://www.heraldsun.com.au/blogs/andrew-bolt/george- Nederlandse Akademie van Wetenschappen. Series C, floyd-and-the-us-race-riots-be-warned/news-story/ Biological and Medical Sciences 56:572-583. f42b7d239067f528b0ebbc0c4480fff9 Brown, W. C. 1952. The amphibians of the Solomon (last downloaded 31 May 2020) Islands. Bull. Mus. Cornp. Zool. Harvard.:107:1-64. Boulenger, G. A. 1882. Catalogue of the Batrachia Bungard, M., Ward, M. and Griffin, M. 2020. George Gradientia and Batrachia Apoda in the collection of the Floyd protests LIVE updates: White House riots intensify British Museum. Trustees of the British Museum, UK:127 as violence ensues across the US, curfews imposed on pp. and 9 pl. multiple cities. Sydney Morning Herald, 1 June, posted Boulenger, G. A. 1883. Description of new species of online at: reptiles and batrachians in the British Museum. Annals https://www.smh.com.au/world/north-america/george- and Magazine of Natural History, Series 5, 12:161-167. floyd-riots-live-updates-minneapolis-national-guard- Boulenger, G. A. 1887a. An account of the batrachians deployed-as-police-officer-charged-with-murder- obtained in Burma by M. L. Fea of the Genoa Civic 20200531-p54y6p.html Museum. Annali del Museo Civico di Storia Naturale di (last downloaded on 1 June 2020) Genova. Series 2, 5:418-424. Burns, E. L. 2004. Phylogeography, Population History Boulenger, G. A. 1887b. Second contribution to the and Conservation Genetics of the Endangered Green herpetology of the Solomon Islands. Proceedings of the and Golden Bell Frog (Litoria aurea). PhD Thesis, Zoological Society of London 1887:333-338. University of New South Wales, Australia:140 pp. Boulenger, G. A. 1892. Description of a new tree-frog Burns, E. and Crayn, D. M. 2006. Phylogenetics and from New South Wales. Proceedings of the Linnean evolution of bell frogs (Litoria aurea species-group, Society of New South Wales, Series 2, 7:403. Anura: Hylidae) based on mitochondrial ND4 sequences. Boulenger, G. A. 1896. Descriptions of a new snake and Molecular Phylogenetics and Evolution 39:573-579. a new frog from north Australia. Proc. Zool. Soc. Lond. Burt, C. E, and Burt. M. D. 1932. Herpetological results of 1895:867. the Whitney South Sea Expedition. VI. Pacific island Boulenger, G. A. 1897a. Descriptions of new lizards and amphibians and reptiles in the collection of the American frogs from Mount Victoria, Owen Stanley Range, New Museum of Natural History. Bulletin of the American Guinea, collected by Mr. A. S. Anthony. Annals and Museum of Natural History 63:461-597. Magazine of Natural History, (6):19: 6-13. Capocaccia, L. 1957. Catalogo dei tipi di anfibi del Museo Boulenger, G. A. 1897b. A list of the reptiles and Civico di Storia Naturale di Genova. Annali del Museo batrachians collected by Mr. Alfred Everett in Lombok, Civico di Storia Naturale di Genova. Serie 3, 69:208-222. Flores, Sumba, and Savu, with descriptions of new Clyne, D. 1969. Australian Frogs. Lansdowne Press, species. Annals and Magazine of Natural History, (6):19: Melbourne, Victoria, Australia:112 pp. 503-509. Cogger, H. G. 1966. A new hylid frog from Australia. Boulenger, G. A. 1898. Fourth report on additions to the Australian Zoologist 13:220-227. batrachian collection in the Natural-History Museum. Cogger, H. G. 1979. Type specimens of reptiles and Proceedings of the Zoological Society of London amphibians in the Australian Museum. Records of the 1898:473-482. Australian Museum 32:164-210. Boulenger, G. A. 1905. Description of new tailless Cogger, H. G. 2014. Reptiles and Amphibians of Australia batrachians in the collections of the British Museum. (Seventh Edition). CSIRO Publishing, Collingwood, Annals and Magazine of Natural History 7(16-20):180- Victoria, Australia:1033 pp. 184. Cogger, H. G., Cameron, E. E. and Cogger, H. M. 1983. Boulenger, G. A. 1911. Descriptions of three new tree- Zoological Catalogue of Australia (1) Amphibia and frogs discovered by Mr. A. E. Pratt in Dutch New Guinea. Reptilia, Australian Government Publishing Service, Annals and Magazine of Natural History, Series 8, 8:55- Canberra, ACT, Australia:319 pp. 56. Condit, J. M. 1964. A list of the types of hylid frogs in the Boulenger, G. A. 1912. On some treefrogs allied to Hyla collection of the British Museum (Natural History). Journal caerulea, with remarks on noteworthy secondary sexual of the Ohio Herpetological Society 4:85-98. characters in the family Hylidae. Zoologische Jahrbücher. Cope, E. D. 1866. On the structures and distribution of Abteilung für Systematik, Geographie und Biologie der the genera of the arciferous Anura. Journal of the Tiere. Jena 15:211-218. Academy of Natural Sciences of Philadelphia. Series Boulenger, G. A. 1914. An annotated list of the 2(6):67-112. batrachians and reptiles collected by the British Cope, E. D. 1867. On the families of the raniform Anura. Ornithologists’ Union Expedition and the Wollaston Journal of the Academy of Natural Sciences of Expedition in Dutch New Guinea. Transactions of the Philadelphia. Series 2, 6:189-206. Zoological Society of London 20:247-275. Copland, S. J. 1957. Australian tree frogs of the genus Boulenger, G. A. 1915. Description of a new tree-frog of Hyla. Proceedings of the Linnean Society of New South the genus Hyla discovered by Mr. A. E. Pratt in the Arfak Wales 82:9-108. Mountains, Dutch New Guinea. Annals and Magazine of Copland, S. J. 1960. A new tree-frog (genus Hyla) from

Queensland. Proceedings of the Linnean Society of New Donnellan, S. C. and Mahony, M. J. 2004. Allozyme, South Wales 85:154-156. chromosomal and morphological variability in the Litoria Copland, S. J. 1961. A new name for Hyla pearsoni, lesueuri species group (Anura:Hylidae), including a preoccupied (Amphibia). Proceedings of the Linnean description of a new species. Australian Journal of Society of New South Wales 86:168. Zoology 52:1-28. Copland, S. J. 1963a. Hyla phyllochrous Günther Donnellan, S. C., McGuigan, K., Knowles, R., Mahony, M. (Amphibia) as an addition to the fauna of Victoria, with a and Moritz, C. 1999. Genetic evidence for species description of a new race and a note on the name of the boundaries in frogs of the Litoria citropa species-group genus. Proceedings of the Linnean Society of New South (Anura : Hylidae). Australian Journal of Zoology, 47:275- Wales 87:137-140. 293. Copland, S. J. 1963b. Size at metamorphosis of the frog Doughty, P. 2011. An emerging frog diversity hotspot in Hyla aureus raniformis (Keferstein). Proceedings of the the northwest Kimberley of Western Australia: another Linnean Society of New South Wales 88:105-107. new frog species from the high rainfall zone. Records of Court of Appeal Victoria. 2014. Hoser v Department of the Western Australian Museum 26:209-216. Sustainability and Environment [2014] VSCA 206 (5 Doughty, P. and Anstis, M. 2007. A new species of rock- September 2014). dwelling hylid frog (Anura:Hylidae) from the eastern Courtice, G. P. and Grigg, G. C. 1975. A taxonomic Kimberley region of Western Australia. Records of the revision of the Litoria aurea complex (Anura : Hylidae) in Western Australian Museum 23:241-257. South Eastern Australia. Aust. Zool. 18:149-63. Dubois, A. 1984. Miscellanea nomenclatorica Covacevich, J. 1974. The status of Hyla irrorata De Vis, batrachologica (II). Alytes. Paris 3:83-84. 1884 (Anura: Hylidae). Memoirs of the Queensland Dubois, A. 2005 Amphibia Mundi. 1.1. An ergotaxonomy Museum 17:49-53. of recent amphibians. Alytes 23:1-24. Coventry, A. J. 1970. Reptile and amphibian type Dubois, A. 2007. Genitives of species and subspecies specimens housed in the National Museum of Victoria. nomina derived from personal names should not be Memoirs of the National Museum of Victoria. Melbourne emended. Zootaxa 1550:49-68. 31:115-124. Dubois, A. 2018. The correct spelling of the nomen Czechura, G. V., Ingram, G. J. and Liem, D. S. S. 1987. Nyctimystes cheesmani Tyler, 1964 The genus Nyctimystes (Anura: Hylidae) in Australia. (Amphibia, Anura), with some comments on Latin Records of the Australian Museum 39:333-338. grammar and the Rules of the Code. Alytes, 35(1-4):75- Daan, S. and Hillenius, D. 1966. Catalogue of the type 84. specimens of amphibians and reptiles in the Zoological Dubois, A. and Frétey, T. 2016. A new nomen for a Museum, Amsterdam. Beaufortia. Amsterdam 13:117- subfamily of frogs (Amphibia, Anura). Dumerilia. Paris 144. 6:17-23. Daudin, F. -M. 1802 “An. XI”. Histoire Naturelle des Dubois, A., Bauer, A. M., Ceriaco, L. M. P., Dusoulier, F., Rainettes, des Grenouilles et des Crapauds. Quarto Fretey, T., Lobl, I., Lorvelec, O., Ohler, A., Stopiglia, R. version. Levrault, Paris. and Aescht, E. 2019. The Linz Zoocode project: a set of Daudin, F. -M. 1803 “An. XI”. Histoire Naturelle, Générale new proposals regarding the terminology, the Principles et Particulière des Reptiles; Ouvrage Faisant suit à and Rules of zoological nomenclature. First report of l’Histoire Naturelle Générale et Particulière, Composée activities (2014-2019). Bionomina (online), 17:1-111. par Leclerc de Buffon; et Rédigée par C.S. Sonnini, Duellman, W. E. 1977. Liste der rezenten Amphibien und Membre de Plusieurs Sociétés Savantes. Volume 8. F. Reptilien. Hylidae, Centrolenidae, Pseudidae. Das Dufart., Paris. Tierreich 95:1-225. Davies, M. M., Martin, A. A. and Watson, G. F. 1983. Duellman, W. E. 1993. Amphibian Species of the World: Redefinition of the Litoria latopalmata species group Additions and Corrections. Special Publication. Natural (Anura: Hylidae). Transactions of the Royal Society of History Museum, University of Kansas 21:iii+372. South Australia 107(2):87-108. Duellman, W. E. and Trueb, L. 1986 Biology of Davies, M. M., McDonald, K. R. and Adams, M. 1986. A Amphibians. McGraw-Hill, New York, USA:670 pp. new species of green tree frog (Anura: Hylidae) from Duellman, W. E., Marion, A. B. and Blair Hedges, S. Queensland, Australia. Proceedings of the Royal Society 2016. Phylogenetics, classification, and biogeography of of Victoria 98:63-71. the treefrogs (Amphibia: Anura: Arboranae). Zootaxa De la Riva, I., Bosch, J. and Márquez, R. 2004. The (online):4104:1-109. advertisement calls of two New Guinea species Duméril, A. H. A. 1853. Mémoire sur les batraciens of Litoria (Amphibia, Anura, Hylidae). Amphibia-Reptilia anoures, de la famille des hylaeformes ou rainettes, 25:173-78. comprenent la description d’un genre nouveau et de Dennis, A. J. and Cunningham, M. J. 2006. Litoria onze espèces nouvelles. Annales des Sciences richardsi sp. nov., a new treefrog (Anura: Hylidae) from Naturelles. Zoologie et Biologie Animale. Paris. New Guinea. Memoirs of the Queensland Museum 52:65- (3)19:135-179. 69. Duméril, A. M. C. and Bibron, G. 1841. Erpétologie De Vis, C. W. 1884. On new species of Hyla. Genérale ou Histoire Naturelle Complète des Reptiles. Proceedings of the Royal Society of Queensland 1:128- Volume 8. Librarie Enclyclopedique de Roret, Paris. 792 130. pp.

Duméril, A. M. C. and Duméril, A. H. A. 1851. Catalogue and Co., Ltd., London, UK. méthodique de la collection des reptiles du Muséum Gassó Miracle, M. E., van den Hoek Ostende, L. W. and d’Histoire Naturelle de Paris. Gide et Baudry/Roret, Paris, Arntzen, J. W. 2007. Type specimens of amphibians in 224 pp. the National Museum of Natural History, Leiden, The Eipper, S. 2012. A guide to … Australian frogs in Netherlands. Zootaxa (online) 1482:25-68. captivity. Reptile Publications, Burleigh, Queensland, Gillespie, G. R. 2001. The role of introduced trout in the Australia:151 pp. decline of the spotted tree frog (Litoria spenceri) in south- Eipper, S. and Rowland, P. 2018. A Naturalist’s Guide to eastern Australia. Biological Conservation 100:187-198. the Frogs of Australia. John Beaufoy Publishing, Oxford, Gillespie, G. R. 2002. Ecology of the Spotted Tree Frog UK:176 pp. (Litoria spenceri): An Investigation of Causes of Faivovitch, J., Haddad, C. F. B., Garcia, P. C. A., Frost, D. Population Decline. PhD Thesis. University of Melbourne, R., Campbell, J. A. and Wheeler, W. C. 2005. Systematic Australia. review of the frog family Hylidae, with special reference to Gillespie, G. R. 2004. Action Statement, Spotted Tree Hylinae: phylogenetic analysis and taxonomic revision. Frog, Litoria spenceri. Department of Sustainability and Bull. Amer. Mus. Nat. Hist. 294:1-240. Environment, Victoria, Australia:10 pp. Fitzinger, L. J. F. J. 1826. Neue Classification der Gillespie, G. R. 2010. Population age structure of the Reptilien nach ihren Natürlichen Verwandtschaften nebst spotted tree frog Litoria spenceri: insights einer Verwandtschafts-Tafel und einem Verzeichnisse der into population declines. Wildlife Research 37:19-26. Reptilien-Sammlung des K. K. Zoologisch Museum’s zu Gillespie, G. R. 2011. Life history variation in the spotted Wien. J. G. Heubner, Wien. tree frog Litoria spenceri (Anura: Fitzinger, L. J. F. J. 1860. Die Ausbeute der Hylidae) from South Eastern Australia. Herpetologica österreichischen Naturforscher an Säugethieren und 67:10-22. Reptilien während der Weltumsegelung Sr. Majestät Gillespie, G. R. 2012. Egg compliment variation amongst Fregatte Novara. Sitzungsberichte der Kaiserlichen temperate stream-breeding Hylid frogs in south-eastern Akademie der Wissenschaften, Mathematisch- Australia. Proceedings of the Royal Society of Victoria Naturwissenschaftliche Classe 42:383-416. 123:153-160. Fletcher, J. J. 1898. Contributions to a more exact Gillespie, G. R. and Hollis, G. J. 1996. Distribution and knowledge of the geographical distribution of Australian habitat of the Spotted Tree Frog Litoria spenceri Dubois Batrachia. No. V. Proceedings of the Linnean Society of (Anura: Hylidae), and an assessment of potential causes New South Wales, (2)12:660-684. of population declines. Wildlife Research 23:49-75. Forliti, A. and Baenen, J. 2020. Four Minneapolis officers Gillespie, G. R., Hunter, D., Berger, L. and Marantelli, G. fired after death of black man. Sydney Morning Herald, 2015. Rapid decline and extinction of a montane frog 27 May, online at: population in southern Australia follows detection of the https://www.smh.com.au/world/north-america/four- amphibian pathogen Batrachochytrium dendrobatidis. minneapolis-officers-fired-after-death-of-black-man- Animal Conservation 18:295-302. 20200527-p54wqk.html Goldman, J., Hill, L. and Stanbury, P. J. 1969. Type (last downloaded on 27 May 2020) specimens in the Macleay Museum, University of Sydney. Forcart, L. 1953. Die Amphibien und Reptilien von II. Amphibians and reptiles. Proceedings of the Linnean Sumba ihre zoogeographischen Beziehungen und Society of New South Wales 93:427-438. Revision der Unterarten von Typhlops polygrammicus. Gray, J. E. 1841. Description of some new species and Verhandlungen der Naturforschenden Gesellschaft in four new genera of reptiles from western Australia, Basel 64:356-388. discovered by John Gould, Esq.. Annals and Magazine of Frost, D. R. 2013. Amphibian Species of the World: An Natural History, 1(7): 6-91. Online Reference. Version 6.0 (30 April 2020). New York, Gray, J. E. 1842. Description of some hitherto unrecorded USA: American Museum of Natural History. species of Australian reptiles and batrachians. Zoological Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, Miscellany, Part 2:51-57. A., Haddad, C. F. B., de Sá, R. O., Channing, A., Guibé, J. 1948. Catalogue des Types d’Amphibiens du Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Muséum National d’Histoire Naturelle. Imprimerie Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nationale, Paris, France. Nussbaum, R. A., Lynch, J. D., Green, D. M. and Günther, A. 1858. Catalogue of the Batrachia Salienata in Wheeler, W. C., 2006. The amphibian tree of life. Bulletin the Collection of the British Museum, London. British of the American Museum of Natural History 297:1-370. Museum, UK:xvi+160 pp. Fry, D. B. 1912. Description of Austrochaperina, a new Günther, A. C. L. G. 1863a. On new species of genus of Engystomatidae from North Australia. Records batrachians from Australia. Ann. Mag. Nat. Hist. 3(11):26- of the Australian Museum 9:87-106. 28. Fry, D. B. 1913. A re-examination of Macleay’s New Günther, A. C. L. G. 1863b. Observations on Australian Guinea and Queensland frog types. Memoirs of the tree-frogs living in the Society’s Menageria. Proceedings Queensland Museum 2:46-50. of the Zoological Society of London 1863:249-251, 1 pl.. Fry, D. B. 1915. Herpetological notes. Proceedings of the Günther, A. C. L. G. 1864. Third contribution to our Royal Society of Queensland 27:60-95. knowledge of batrachians from Australia. Proceedings of Gadow, H. F. 1901. Amphibia and Reptiles. Macmillan the Zoological Society of London 1864:46-49. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 179

Günther, A. C. L. G. 1867. Additions to the knowledge of Hero, J. –M., Littlejohn, M. and Marantelli, G. 1991. Australian reptiles and fishes. Annals and Magazine of Frogwatch field guide to Victorian Frogs. Department of Natural History, (3)20:45-68. Conservation and Environment, Victoria, Australia:108 Günther, A. C. L. G. 1873. Description of two new species pp. of frogs from Australia. Annals and Magazine of Natural Hero, J. –M., Watson, G. F. and Gillespie, G. 1995. The History, (4)11:349-350. tadpole of Litoria spenceri Spencer (Anura: Hylidae). Günther, A. C. L. G. 1876. Descriptions of new species of Proceedings of the royal Society of Victoria 100:39-43. reptiles from Australia, collected by Hr. Dämel for the Hiaso, J. and Richards, S.J. 2006. A new species of Godeffroy Museum. Journal des Museum Godeffroy. Litoria (Anura: Hylidae) from Tagula Island, Papua New Hamburg 5:45-47. Guinea. Science in New Guinea 28:34-38. Günther, A. C. L. G. 1897. Descriptions of new species of Horst, R. 1883. On new and little-known frogs from the lizards and of a tree-frog from north-eastern Queensland. Malayan archipelago. Notes from the Leyden Museum Novitates Zoologicae. Tring 4:403-406. 5:235-244. Günther, R. 2003. Notable distribution records of New Hoser, R. T. 1989. Australian Reptiles and Frogs. Pierson Guinea frog species and genera beyond their known and Co., Sydney, NSW, Australia:238 pp. ranges (Amphibia: Anura). Faunistische Abhandlungen Hoser, R. T. 1991. Endangered Animals of Australia. Staatliches Museum fur Tierkunde Dresden: 209-216. Pierson and Co., Mosman, NSW, 240 pp. Günther, R. 2004a. Description of a new treefrog species Hoser, R. T. 1993. Smuggled: The Underground Trade in from western New Guinea showing extreme colour Australia’s Wildlife. Apollo Publishing, Moss Vale, NSW. polymorphism (Anura, Hylidae, Litoria). Mitteilungen aus 160 pp. dem Museum für Naturkunde in Berlin. Zoologische Hoser, R. T. 1995. Release into hell. Monitor:Journal of Reihe 80:251-260. the Victorian Herpetological Society, 7(2):77-88. Günther, R. 2004b. Two new treefrog species of the Hoser, R. T. 1996. Smuggled-2: Wildlife Trafficking, Crime genus Litoria (Anura: Hylidae) from the west of New and Corruption in Australia. Kotabi Publishing, Doncaster, Guinea. Zoologische Abhandlungen. Staatliches Museum Victoria, 3108, Australia:280 pp. für Tierkunde in Dresden 54:163-175. Hoser, R. T. 1998. Comments on the proposed Günther, R. 2006a. A remarkable new species of the conservation of the specific name of Varanus teriae genus Litoria (Amphibia, Anura, Hylidae) from north- Sprackland, 1991 (Reptilia, Squamata) (Case 3043; see western New Guinea. Zoosystematics and Evolution BZN 54: 100-103, 250-251; 55: 37-39). Bulletin of 82:179-190. Zoological Nomenclature 55(2):113-114. Günther, R. 2006b. Derived reproductive modes in New Hoser, R. T. 2007. Wells and Wellington - It’s time to bury Guinean anuran amphibians and description of a new the hatchet. Calodema Supplementary Paper 1:1-9. species with paternal care in the genus Callulops Hoser, R. T. 2009. Creationism and contrived science: A (Microhylidae). Journal of Zoology 268:153-170. review of recent python systematics papers and the Günther, R. 2006c. A new species of treefrog of the resolution of issues of taxonomy and nomenclature. genus Litoria (Anura, Hylidae) from Biak Island off Australasian Journal of Herpetology 2:1-34. (3 February). northwest New Guinea. Salamandra 42:117-128. Hoser, R. T. 2012a. Exposing a fraud! Afronaja Wallach, Günther, R. 2008. Two new hylid frogs (Anura: Hylidae: Wüster and Broadley 2009, is a junior synonym of Litoria) from western New Guinea. Vertebrate Zoology. Spracklandus Hoser 2009! Australasian Journal of Museum für Tierkunde, Dresden 58:103-112. Herpetology 9 (3 April 2012):1-64. Günther, R. 2014. New data on Litoria mucro (Amphibia, Hoser, R. T. 2012b. Robust taxonomy and nomenclature Anura, Hylidae) from New Guinea. Abhandlungen des based on good science escapes harsh fact-based Naturwissenschaftlichen Vereins zu Bremen 47:225-231. criticism, but remains unable to escape an attack of lies Günther, R. and Richards, S. J. 2000. A new species of and deception. Australasian Journal of Herpetology the Litoria gracilenta group from Irian Jaya (Anura: 14:37-64. Hylidae). Herpetozoa 13:27-43. Hoser, R. T. 2013a. The science of herpetology is built on Günther, R. and Richards, S. J. 2005. Three new evidence, ethics, quality publications and strict mountain stream dwelling Litoria (Amphibia: Anura: compliance with the rules of nomenclature. Australasian Hylidae) from western New Guinea. Russian Journal of Journal of Herpetology 18:2-79. Herpetology 12:195-212. Hoser, R. T. 2013b. Monitor Lizards reclassified with Hansen, N. and Crosby, K. 2016. Habitat use of the some common sense (Squamata: Sauria: Varanidae). threatened Booroolong Frog (Litoria booroolongensis) in Australasian Journal of Herpetology 21: 41-58. the Central West Catchment Management Area. Hoser, R. T. 2014. The break up of Odatria Australian Zoologist 38(2):161-170. (Kimberleyvaranus) glebopalma (Mitchell, 1955) into Häupl, M. and Tiedemann, F. 1978. Vertebrata 1. three obvious subspecies. Australasian Journal of Typenkatalog der Herpetologischen Sammlung. Kataloge Herpetology 24:16-19. der Wissenschaftlichen Sammlungen des Hoser, R. T. 2015a. Dealing with the “truth haters” ... a Naturhistorischen Museums in Wien 2:7-34. summary! Introduction to Issues 25 and 26 of Heatwole, H., De Bavay, J., Webber, P. and Webb, G. Australasian Journal of Herpetology. Including “A timeline 1995. Faunal survey of New England. IV. The frogs. of relevant key publishing and other events relevant to Memoirs of the Queensland Museum 38:229-249. Wolfgang Wüster and his gang of thieves.” and a Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 180 Australasian Journal of Herpetology

“Synonyms list”. Australasian Journal of Herpetology Australasian Journal of Herpetology 39:53-63. 25:3-13. Hoser, R. T. 2020a. From a putative new taxon to a mutt! Hoser, R. T. 2015b. The Wüster gang and their proposed Formal descriptions of three new genetically divergent “Taxon Filter”: How they are knowingly publishing false Mountain Pygmy Possums from Victoria and New South information, recklessly engaging in taxonomic vandalism Wales closely associated with Burramys parvus Broom, and directly attacking the rules and stability of zoological 1896. Australasian Journal of Herpetology 42:3-10. nomenclature. Australasian Journal of Herpetology Hoser, R. T. 2020b. Small and overlooked … six new 25:14-38. species of Pygmy Possum, Genus Cercartetus Gloger, Hoser, R. T. 2015c. Best Practices in herpetology: Hinrich 1841 sensu lato from the Australasian bioregion. Kaiser’s claims are unsubstantiated. Australasian Journal Australasian Journal of Herpetology 42:11-22. of Herpetology 25:39-52. Hoser, R. T. 2020c. A new subgenus, three new species Hoser, R. T. 2015d. PRINO (Peer reviewed in name only) and one new subspecies of Ringtailed Possums journals: When quality control in scientific publication (Marsupialia: Petauridae) from the north of Australia. fails. Australasian Journal of Herpetology 26:3-64. Australasian Journal of Herpetology 42:23-30. Hoser, R. T. 2015e. Rhodin et al. 2015, Yet more lies, Hoser, R. T. 2020d. The ability to conserve a threatened misrepresentations and falsehoods by a band of thieves species begins when they are named! New species of intent on stealing credit for the scientific works of others. Rock Wallaby (Marsupialia: Macropodidae: Petrogale) Australasian Journal of Herpetology 27:3-36. from northern Australia. Australasian Journal of Hoser, R. T, 2015f. Comments on Spracklandus Hoser, Herpetology, 42:42-49. 2009 (Reptilia, Serpentes, ELAPIDAE): request for Hoser, R. T. 2020e. An overdue refinement of the confirmation of the availability of the generic name and taxonomy of the Australian Ring Tailed Dragons, Genus for the nomenclatural validation of the journal in which it Ctenophorus Fitzinger, 1843, Subgenus Tachyon Wells was published (Case 3601; see BZN 70: 234-237; and Wellington, 1985, including the formal descriptions of comments BZN 71:30-38, 133-135). (unedited version) eight new species. Australasian Journal of Herpetology, Australasian Journal of Herpetology 27:37-42. 42:50-64. Hoser, R. T. 2015g. Hitherto overlooked species of reptile Hoskin, C. J. 2007. Description, biology and conservation from Northern Australia: A result of science, taxonomy, of a new species of Australian tree frog (Amphibia: molecular biology, systematics, history and forensic Anura: Hylidae: Litoria) and an assessment of the herpetology. Australasian Journal of Herpetology 30:21- remaining populations of Litoria genimaculata Horst, 27. 1883: systematic and conservation implications of an Hoser, R. T. 2018a. New Australian lizard taxa within the unusual speciation event. Biological Journal of the greater Egernia Gray, 1838 genus group of lizards and Linnean Society 91:549-563. the division of Egernia sensu lato into 13 separate Hosmer, W. 1964. A new frog of the genus Hyla from genera. Australasian Journal of Herpetology, 36:49-64. Northern Territory, Australia. American Museum Hoser, R. T. 2018b. Six new species of Dwarf Goanna, Novitates 2182:1-7. Worrellisaurus Wells and Wellington, 1984 from Hunter, D. A. 2012. National Recovery Plan for Australia. Australasian Journal of Herpetology 37:24-37. Booroolong Frog Litoria booroolongensis. Office of Hoser, R. T. 2018c. Varanus kingorum Storr, 1980, Environment and Heritage, Hurstville, NSW, Australia. Varanus minor Weigel, 1985, a damaging case of Hunter, D. A. and Smith, M. J. 2013. Multiscale habitat taxonomic vandalism by John Weigel and Worrellisaurus assessment for the endangered Booroolong Frog (Litoria bigmoreum sp. nov., a new species of small monitor Booroolongensis): Implications for threatened species lizard, from the East Kimberley division of Western management in rural landscapes of southeastern Australia. Australasian Journal of Herpetology 37:38-43. Australia. Herpetological Conservation and Biology Hoser, R. T. 2018d. A new species within the Odatria 8:122-130. glauerti (Squamata: Varanidae) species complex. Hunter, D. A., Smith, M. J., Scroggie, M. P. and Gilligan, Australasian Journal of Herpetology 37:44-45. D. 2011. Experimental examination of the potential for Hoser, R. T. 2018e. The description of a fourth three introduced fish species to prey on tadpoles of the subspecies of Odatria (Kimberleyvaranus) glebopalma endangered Booroolong frog, Litoria booroolongensis. (Mitchell, 1955) (Reptilia: Squamata: Varanidae). Journal of Herpetology 45:181-185. Australasian Journal of Herpetology 37:46-47. Ingram, G. J. and Corben, C. J. 1990. Litoria electrica: a Hoser, R. T. 2019a. 11 new species, 4 new subspecies new treefrog from western Queensland. Memoirs of the and a subgenus of Australian Dragon Lizard in the genus Queensland Museum 28:475-478. Tympanocryptis Peters, 1863, with a warning on the Ingram, G. J., Corben, C. J. and Hosmer, W. 1982. Litoria conservation status and long-term survival prospects of revelata: a new species of tree-frog from eastern some newly named taxa. Australasian Journal of Australia. Memoirs of the Queensland Museum 20:635- Herpetology 39:23-52. 637. Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Ingram, G. J., Nattrass, A. E. O. and Czechura, G. V. Kaiser et al. (2013), Jane Melville et al. (2018 and 2019): 1993. Common names for Queensland frogs. Memoirs of Australian Agamids and how rule breakers, liars, thieves, the Queensland Museum 33:221-224. taxonomic vandals and law breaking copyright infringers James, C. H. 1998. Molecular systematics and are causing reptile species to become extinct. phylogeography of the Litoria bicolor species group

(Anura: Hylidae) from Eastern Australia. PhD thesis, Kraus, F. 2013b. A new treefrog of the Litoria gracilenta University of Queensland, Brisbane, Australia. group (Hylidae) from Papua New Guinea. Proceedings of James, C. H. and Moritz, C. 2000. Intraspecific the Biological Society of Washington 126:151-160. phylogeography in the sedge frog Litoria fallax (Hylidae) Kraus, F. 2018. Taxonomy of Litoria graminea (Anura: indicates pre-Pleistocene vicariance of an open forest Hylidae), with descriptions of two closely related new species from eastern Australia. Molecular Ecology 9:349- species. Zootaxa (online PRINO):4457:264-284. 358. Kraus, F. and Allison, A. 2004a. Two new treefrogs from Johnston, G. R. and Richards, S. J. 1994. A new species Normanby Island, Papua New Guinea. Journal of of Litoria (Anura: Hylidae) from New Guinea and a Herpetology 38:197-207. redefinition of Litoria leucova (Tyler, 1968). Memoirs of Kraus, F. and A. Allison. 2004b. A new species of Litoria the Queensland Museum 37:273-279. (Anura: Hylidae) from southeastern New Guinea. Kaiser, H. 2012a. SPAM email sent out to numerous Herpetologica 60:97-103. recipients on 5 June 2012. Kraus, F. and Allison, A. 2009 New species of frogs from Kaiser, H. 2012b. Point of view. Hate article sent as Papua New Guinea. Bishop Museum Occasional Papers attachment with SPAM email sent out on 5 June 2012. 104:1-36. Kaiser, H. 2013. The Taxon Filter, a novel mechanism Krell, F. and Marshall, S. A. 2017. New Species designed to facilitate the relationship between taxonomy Described From Photographs: Yes? No? and nomenclature, vis-à-vis the utility of the Code’s Sometimes? A Fierce Debate and a New Declaration of Article 81 (the Commission’s plenary power). Bulletin of the ICZN. Insect Systematics and Diversity, 1(1):2017:3- Zoological Nomenclature 70(4) December 2013:293-302. 19. Kaiser, H. 2014a. Comments on Spracklandus Hoser, Lamb, J. 1911. Description of three new batrachians from 2009 (Reptilia, Serpentes, ELAPIDAE): request for southern Queensland. Annals of the Queensland confirmation of the availability of the generic name and Museum 10:26-28. for the nomenclatural validation of the journal in which it Laurance, W. F., McDonald, K. R. and Speare, R. 1996. was published. Bulletin of Zoological Nomenclature, Epidemic disease and the catastrophic decline of 71(1):30-35. Australian rain forest frogs. Conservation Biology 10:406- Kaiser H. 2014b. Best Practices in Herpetological 413. Taxonomy: Errata and Addenda. Herpetological Review, Lesson, R. -P. 1829. Fig. 7. Duperrey, L.-I. ed., Voyage 45(2):257-268. autour du Monde, exécuté par Ordre du Roi, sur la Kaiser, H., Crother, B. L., Kelly, C. M. R., Luiselli, L., Corvette de sa Majesté, La Coquille, pendant les années O’Shea, M., Ota, H., Passos, P., Schleip, W. D. and 1822, 1823, 1824, 1825. Atlas. Livraison 11: Paris, Arthur Wüster, W. 2013. Best practices: In the 21st Century, Bertrand. Taxonomic Decisions in Herpetology are Acceptable Only Lesson, R. -P. 1831. Zoologie. Duperrey, L.-I. ed., Voyage When supported by a body of Evidence and Published autour du Monde, exécuté par Ordre du Roi, sur la via Peer-Review. Herpetological Review 44(1):8-23. Corvette de sa Majesté, La Coquille, pendant les années Keferstein, W. M. 1867. Über einige neue oder seltene 1822, 1823, 1824, 1825. Volume 2, Part 1 Livraison 25, Batrachier aus Australien und dem tropischen Amerika. Paris, Arthur Bertrand. Nachrichten von der Georg-Augusts-Universität und der Leunis, J. 1844. Synopsis der drei Naturreiche ... Mit Königl. Gesellschaft der Wissenschaften zu Göttingen vorzüglicher Berücksichtigung der nützlichen und 18:342-361. schädlichen Naturkörper Deutschlands. Erster Theil, Keferstein, W. M. 1868. Über die Batrachier Australiens. Zoologie. Hahn’fche Hofbuchhandlung, Hannover. Archiv für Naturgeschichte. Berlin 34:251-290. Liem, D. S. S. 1974a. A review of the Litoria nannotis Kraus, F. 2007. A new species of treefrog of the genus species group, and a description of a new species of Litoria (Hylidae) from the Louisiade Islands, Papua New Litoria from northern Queensland, Australia (Anura: Guinea. Herpetologica, 63:365-374. Hylidae). Memoirs of the Queensland Museum 17:151- Kraus, F. 2009. New species of Toxicocalamus 168. (Squamata: Elapidae) from Papua New Guinea. Liem, D. S. S. 1974b. A new species of the Litoria bicolor Herpetologica 65(4):460-457. species group from southeast Queensland, Australia Kraus, F. 2010. More range extensions for Papuan (Anura: Hylidae). Memoirs of the Queensland Museum reptiles and amphibians. Herpetological Review 41:246- 17:169-174. 248. Liem, D. S. S, and Ingram, G. J. 1977. Two new species Kraus, F. 2012a. Identity of Nyctimystes cheesmani of frogs (Anura: Myobatrachidae, Pelodryadidae) from (Anura: Hylidae), with description of two new related Queensland and New South Wales. Victorian Naturalist species. Zootaxa (Online PRINO) 3493:1-26. 94:255-262. Kraus, F. 2012b. A new species of Nyctimystes (Anura: Lönnberg, E. 1900. Reptiles and batrachians collected in Hylidae) from Papua New Guinea. Herpetologica 68:541- German New Guinea by the late Dr. Erik Nyman. Annals 550. and Magazine of Natural History, (7):6:574-582. Kraus, F. 2013a. Morphological data show that Hyla dayi Loveridge, A. 1945. New tree-frogs of the genera Hyla Gunther, 1897 (Amphibia: Anura: Hylidae) should never and Nyctimystes from New Guinea. Proceedings of the have been assigned to Nyctimystes. Memoirs of the Biological Society of Washington 58:53-58. Queensland Museum 56:581-587. Loveridge, A. 1948. New Guinean reptiles and

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 182 Australasian Journal of Herpetology amphibians in the Museum of Comparative Zoology and Menzies, J. I. and Tippett, J. 1976. Chromosome United States National Museum. Bulletin of the Museum numbers of Papuan hylid frogs and the karyotype of of Comparative Zoology, Harvard 101:305-430. Litoria infrafrenata (Amphibia, Anura, Hylidae). Journal of Loveridge, A. 1950. New frogs of the genera Cyclorana Herpetology 10:167-173. and Hyla from southeastern Australia. Proceedings of the Menzies, J. I. and Tyler, M. J. 2004. Litoria gracilenta Biological Society of Washington 63:131-138. (Anura: Hylidae) and related species in New Guinea. Macleay, W. J. 1878. The batrachians of the “Chevert” Australian Journal of Zoology 52:191-214. Expedition. Proceedings of the Linnean Society of New Menzies, J. I. and Zug, G. R. 1979. Papuan tree frogs of South Wales 2:135-138. the Litoria thesaurensis group (Salientia: Hylidae). Mahony, M., Knowles, R., Foster, R. and Donnellan, S. Micronesica 15:325-333. 2001.Systematics of the Litoria citropa (Anura: Hylidae) Menzies, J. I. and Zweifel, R. G. 1974. Systematics of Complex in Northern New South Wales and Southern Litoria arfakiana of New Guinea and sibling species Queensland, Australia, With the Description of a New (Salientia, Hylidae). Am. Mus. Novit. No. 2558. Species. Records of the Australian Museum 53:37-48. Menzies, J. I. and Zweifel, R. G. 1976. Specific identity of Main, A. R. 1965. The frogs of southern Western specimens in the type series of the Papnan hylid frog Australia. Handbook No. 8., West Australian Naturalists Litoria arfakiana. Ann. Mus. Civ. Stor. Nut. ‘Giacom Doria’ Club, Perth, Australia. 81:17-21. Martin, A. A., Watson, G. F., Gartside, D. F., Littlejohn, M. Menzies, J. I., Richards, S. J. and Tyler, M. J. 2009. J. and Loftus-Hills, J. 1978. A new species of the Litoria Systematics of the Australo-Papuan tree frogs known as peronii complex (Anura: Hylidae) from eastern Australia. Litoria bicolor (Anura: Hylidae) in the Papuan Region. Proceedings of the Linnean Society of New South Wales Australian Journal of Zoology 56:257-280. 103:23-35. Merrem, B. 1820. Versuch eines Systems der Amphibien. McDonald, K. R. 1997. A new stream-dwelling Litoria Tentamen Systematis Amphibiorum. Marburg, Johann from the Melville Range, Queensland, Australia. Christian Krieger, Hesse (Germany). Memoirs of the Queensland Museum 42:307-309. Mertens, R. 1922. Verzeichnis der Typen in der McDonald, K. R., Rowley, J. J. L., Richards, S. J. and herpetologischen Sammlung des Senckenbergischen Frankham, G. J. 2016 A new species of treefrog (Litoria) Museum. Senckenbergiana Biologica 4:162-183. from Cape York Peninsula, Australia. Zootaxa (online Mertens, R. 1930. Die von Dr. F. Kopstein auf den PRINO) 4171:153-169. Molukken und einigen Benachbarten Inseln gesammelten McDowell, S. B. 1969. Toxicocalamus, a New Guinea Froschlurche. Zoologische Mededelingen. Leiden genus of snakes of the family Elapidae. Journal of 13:141-150. Zoology, London 159:443-511. Mertens, R. 1964. Die systematische Stellung des zentral Méhely, L. von. 1897. Further contributions to the australischen Laubfrosches Hyla gilleni. herpetology of New-Guinea [In Hungarian and English]. Senckenbergiana Biologica 45:15-21. Természetrajzi Füzetek. Budapest 20:398-419. Mertens, R. 1967. Die herpetologische Sektion des Menzies, J. I. 1969. A new species of tree frog (Hyla) Natur-Museums und Forschungs-Institutes Senckenberg from Papua. Transactions of the Royal Society of South in Frankfurt a. M. nebst einem Verseichnis ihrer Typen. Australia 93:165-168. Senckenbergiana Biologica 48(A):1-106. Menzies, J. I. 1972. Papuan tree frogs of the Litoria Meyer, A. B. 1874. Eine Mittheilung von Hrn. Dr. Adolf nigropunctata group. Herpetologica 28(4):291-300. Bernhard Meyer über die von ihm auf Neu-guinea und Menzies, J. I. 1976. Handbook of common New Guinea den Inseln Jobi, Mysore und Mafoor im Jahre 1873 Frogs. Wau Ecology Institute, PNG:75 pp. gesammelten Amphibien. Monatsberichte der Menzies, J. 1993. Systematics of Litoria iris (Anura, Königlichen Preussische Akademie des Wissenschaften Hylidae) and its allies in New-Guinea and a note on zu Berlin 1874:128-140. sexual dimorphism in the group. Australian Journal of Meyer, A. B. 1887. Verzeichness der von mir in den Zoology 41:225-255. Jahren 1870-1873 im ostindischen Archipel Menzies, J. I. 2006. Frogs of New Guinea and the gesammelten Reptilien und Batrachier. Abhandlungen Solomon Islands, Pensoft, Sofia. und Berichte des Zoologischen und Anthropologisch- Ethnographischen Museums zu Dresden 1886-1887 (Art. Menzies, J. I. 2014a. Notes on Nyctimystes Species 2):1-16. (Anura, Hylidae) of New Guinea: The Nyctimystes narinosus Species Group with Descriptions of Two New Meyer, E. and Agnew, L. 2013. A range extension of the Species. Transactions of the Royal Society of South water holding frog (Cyclorana platycephala) in Australia, Incorporated: incorporating the Records of the Queensland. Queensland Naturalist, 51:19-22. South Australian Museum 138(1):135-143. Minister for the Environment, Commonwealth Menzies, J. I. 2014b. Notes on Nyctimystes (Anura: Government of Australia. 2017. Threatened species Hylidae), tree frogs of New Guinea, with descriptions of scientific committee, Conservation Advice, Litoria four new species. Alytes. Paris 30:42-68. spenceri, Spotted tree frog. Commonwealth of Australia, Department of Environment:10 pp. Menzies, J. I. and Johnston, G. R. 2015. The structure of the male proboscis in the New Guinean tree frogs, Litoria Mitchell, F. J. 1948. A revision of the lacertilian genus pronimia and Litoria havina (Anura:Hylidae). Australian Tympanocryptis. Records of the South Australian Journal of Zoology 63:175-180. Museum 9:57-86.

Moore, J. A. 1961. The frogs of eastern New South Peters, W. C. H. 1867. Herpetologische Notizen. Wales. Bull. Am. Mtrs. Nat. Hist. 121:149-386. Monatsberichte der Königlichen Preussische Akademie Murray, B. R. and Hose, G. C. 2005. Life-history and des Wissenschaften zu Berlin 1867:13-37. ecological correlates of decline and extinction in the Peters, W. C. H. 1869. Über neue Saurier endemic Australian frog fauna. Austral Ecology 30:564- (Chaunolaemus multicarinatus, Tropidolepisma richardi 571. und Gymnodactylus steudneri) und Batrachier Nieden, F. 1923. Anura I. Subordo Aglossa und (Cyclorhamphus fasciatus und Hyla gracilenta). Phaneroglossa, Sectio 1. Arcifera. Das Tierreich 46: Monatsberichte der Königlichen Preussische Akademie xxxii+584. des Wissenschaften zu Berlin 1869:786-790. Noble, G. K. 1931. The Biology of the Amphibia. New Peters, W. C. H. 1871. Über einige Arten der York and London: McGraw-Hill. herpetologischen Sammlung des Berliner zoologischen Obst, F. J. 1977. Die Herpetologische Sammlung des Museums. Monatsberichte der Königlichen Preussische Staatlichen Museums für Tierkunde Dresden und ihre Akademie des Wissenschaften zu Berlin 1871:644-652. Typusexemplare. Zoologische Abhandlungen. Staatliches Peters, W. C. H. 1873a. Über zwei Gisftschlangen aus Museum für Tierkunde in Dresden 34:171-186. Afrika und über neue oder weniger bekannte Gattungen Ogilby, J. D. 1890. Report on a zoological collection from und Arten von Batrachiern. Monatsberichte der British New Guinea. I. Reptiles, batrachians, and fishes. Königlichen Preussische Akademie des Wissenschaften Records of the Australian Museum 1:89-101. zu Berlin 1873:411-418. Ogilby, J. D. 1907. A new tree frog from Brisbane. Peters, W. C. H. 1873b. Über eine neue Schildrötenart, Proceedings of the Royal Society of Queensland 20:31- Cinosternon Effeldtii und einige andere neue oder 32. weniger bekannte Amphibien. Monatsberichte der Oken, L. 1816. Lehrbuch der Naturgeschichte. Vol. 3. Königlichen Preussische Akademie des Wissenschaften Zoologie. Abtheilung 2. Atlas. C. H. Reclam, Leipzig. zu Berlin 1873:603-618. Oliver, P. M. and Richards, S. J. 2007. A new species of Peters, W. C. H. 1874. Über neue Amphibien (Gymnopis, montane stream-dwelling Litoria from Papua, Indonesia Siphonops, Polypedates, Rhacophorus, Hyla, Clyclodus, (Anura: Hylidae). Hamadryad 31(2):299-303. Euprepes, Clemmys). Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin Oliver, P. M., Richards, S. J.,Tjaturadi, B. and Iskandar, 1874: 616-624. D. T. 2007. A new large green species of Litoria (Anura: Hylidae) from western New Guinea. Zootaxa (PRINO Peters, W. C. H. 1877. Herpetologische Notizen. II. online) 1519:17-26. Bemerkungen über neue oder weniger bekannte Amphibien. Monatsberichte der Königlichen Preussische Oliver, P. M., Stuart-Fox, D. and Richards, S. J. 2008. A Akademie des Wissenschaften zu Berlin 1877:415-423. new species of treefrog (Hylidae, Litoria) from the southern lowlands of New Guinea. Current Herpetology Peters, W. C. H. 1878. Herpetologischen Notizen. 2, 27(1):35-42. Bemerkungen über neue oder weniger bekannte Amphibien. Mber. dt. Akad. Wiss. Berlin 1878:415-423. Oliver, P. M., Günther, R., Mumpuni, and Richards, S. J. 2019a. Systematics of New Guinea treefrogs (Litoria: Peters, W. C. H. 1880. Über neue oder weniger bekannte Pelodryadidae) with erectile rostral spikes: an extended Amphibien des Berliner Zoologischen Museums description of Litoria pronimia and a new species from (Leposoma dispar, Monopeltis (Practogonus) jugularis, the Foja Mountains. Zootaxa (PRINO online) 4604:335- Typhlops depressus, Leptocalamus trilineatus, Xenodon 348. punctatus, Elapomorphus erythronotus, Hylomantis fallax). Monatsberichte der Königlichen Preussische Oliver, P. M., Richards, S. J. and Donnellan, S. C. 2019b. Akademie des Wissenschaften zu Berlin 1880:217-224. Two new species of treefrog (Pelodrydidae: Litoria) from southern New Guinea elucidated by DNA barcoding. Peters, W. C. H. 1882. Über eine neue Batrachier der Zootaxa (PRINO Online) 4609(3):469-484 . Gattung Hyperolius und Limnodytes (Hylorana) aus Africa. Sitzungsberichte der Gesellschaft Parker, H. W. 1936. A collection of reptiles and Naturforschender Freunde zu Berlin 1882: 8-10. amphibians from the Mountains of British New Peters, W. C. H. and Doria, G. 1878. Catalogo dei rettili e Guinea. Annals and Magazine of Natural History 17:66- dei batraci raccolti da O. Beccari, 93. L. M. D’Albertis e A. A. Bruijn nella SottoRegione Parker, H. W. 1938. The races of the Australian frog Hyla AustroMalese. Ann. Mus. Civ. Stor. Nat. 13:323-450. aurea Lesson. Annals and Magazine of Natural History, Series 11(2):302-305. Pyron, A. and Wiens, J. J. 2011 A large-scale phylogeny of Amphibia with over 2,800 species, and a revised Parker, H. W. 1940. The Australasian frogs of the family classification of extant frogs, salamanders, and Leptodactylidae. Novitates Zoologicae. Tring 42:1-106. caecilians. Molecular Phylogenetics and Evolution Péron, F. 1807. Voyage de Decouvertes aux Terres 61:543-583. Australes, execute par ordre de sa majeste l’Empereur et Ramsay, E. P. 1878. Description of a new species of Roi, sur les Corvettes la Geographe, la Naturaliste et la Pelodryas, from New Ireland. Proceedings of the Linnean Goulette le Casuarina, pendant les annees 1800, 1801, Society of New South Wales 2:28-30. 1803 et 1804. Volume 1. Imprimerie imperial, Paris. Regan, L. 2002. Saving the Booroolong Frog. Peters, W. C. H. 1863. Fernere Mittheilungen über neue Australasian Science. Control Publications. Hawksburn, Batrachier. Monatsberichte der Königlichen Preussische Victoria, Australia 23(9):34-35. Akademie des Wissenschaften zu Berlin 1863: 445-470. Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 184 Australasian Journal of Herpetology

Rensch, B. 1936. Die Geschichte des Sundabogens. 5BD, UK (also commonly cited as “The Rules”, Eine tiergeographische Untersuchung. Berlin: Gebrüder “Zoological Rules” or “ICZN 1999”). Borntraeger. Robinson, M. 1993. A field guide to frogs of Australia. Reynolds, S. J. 2007. Some common names for top end Reed, Chatswood, New South Wales, Australia:112 pp. frogs. Northern Territory Naturalist 19:60-68. Rosauer, D., Laffan, S. W., Crisp, M. D., Donnellan, S. C. Richards, S. J. 1992. The tadpole of the Australian frog and Cook, L. G. 2009. Phylogenetic endemism: a new Litoria nyakalensis (Anura, Hylidae), and a key to the approach for identifying geographical concentrations of torrent tadpoles of northern Queensland. Alytes, 10:99- evolutionary history. Molecular Ecology 18:4061-4072. 103. Rowley, J. J. L. and Cutajar, T. P. 2018. Rediscovery of Richards, S. J. 2001. A new torrent-dwelling frog (Anura, the Booroolong Frog Litoria booroolongensis on the Hylidae, Litoria) from the mountains of Indonesian New Australian New England Tablelands after more than 40 Guinea (West Papua). Memoirs of the Queensland years. Herpetological Review. 49:620-621. Museum 47:733-739. Saunders, S. 2019. Morrison and Shorten’s Big Australia: Richards, S. J. 2002. Rokrok: An illustrated field guide to The overpopulation problem. News article published 17 frogs of the Kikori Integrated Conservation and April 2019 and downloaded from Development Project area of Papua New Guinea. WWF, https://independentaustralia.net/politics/politics-display/ Papua New Guinea:33 pp. morrison-and-shortens-big-australia-the-overpopulation- Richards, S. J. 2005. A new species of treefrog (Anura: problem,12584 Hylidae: Litoria) from the Huon Peninsula, Papua New (last downloaded on 7 February 2020). Guinea. Zootaxa (online PRINO)1052:29-39. Savage, J. M. 1986. Nomenclatural notes on the Anura Richards, S. J. 2007a. A new species of “spike-nosed” (Amphibia). Proceedings of the Biological Society of frog from northern New Guinea (Anura: Hylidae: Litoria). Washington 99:42-45. Zootaxa (online PRINO):1525 (1):51-59. Schneider, J. G. 1799. Historia Amphibiorum Naturalis et Richards, S. J. 2007b. A new species of Nyctimystes Literarariae. Fasciculus Primus. Continens Ranas, (Anura, Hylidae) from Papua New Guinea and comments Calamitas, Bufones, Salamandras et Hydros in Genera et on poorly-known members of the genus. Phyllomedusa. Species Descriptos Notisque suis Distinctos. Friederici Belo Horizonte 6:105-118. Frommanni, Jena. Richards, S. J. and Alford, R. A. 2005. Structure and Scott, E. O. G. 1942. A new Hyla from Cradle Valley, dynamics of a rainforest frog (Litoria genimaculata) Tasmania. Records of the Queen Victoria Museum. population in northern Queensland. Australian Journal of Launceston 1:5-11. Zoology 53:229-236. Shaw, G. 1802. General Zoology or Systematic Natural Richards, S. J. and Iskandar, D. T. 2001. A new tree frog History. Volume III, Part 1. Amphibia. Thomas Davison, (Anura, Hylidae, Litoria) from the mountains of Irian Jaya, London, UK. Indonesia. Alytes 18:141-152. Shea, G. M. and Kraus, F. 2007. A list of herpetological Richards, S. and Iskandar, D. T. 2006. A new species of type specimens in the collections of the Papua New torrent-dwelling frog (Hylidae, Litoria) from the mountains Guinea National Museum and Art Gallery and University of New Guinea. Current Herpetology 25:57-63. of Papua New Guinea. Zootaxa (online PRINO) 1514:37- Richards, S. J. and Johnston, G. R. 1993. A new species 60. of Nyctimystes (Anura: Hylidae) from the Star Mountains, Shea, G. M. and Sadlier, R. A. 1999. A catalogue of the Papua New Guinea. Memoirs of the Queensland non-fossil amphibian and reptiles type specimens in the Museum 33:73-76. collection of the Australian Museum; types currently, Richards, S. J. and Oliver, P. M. 2006a. Two new species previously and purportedly present. Technical Reports of of large green canopy-dwelling frogs the Australian Museum 15:1-91. (Anura: Hylidae: Litoria) from Papua New Guinea. Sonnini de Manoncourt, C. S., and Latreille, P. A. 1801 Zootaxa (PRINO online):1295:41-60. “An. X”. Histoire Naturelle des Reptiles, avec Figures Richards, S. J. and Oliver, P. M. 2006b. A new species of dissinées d’après Nature. Volume 2. Paris: Deterville. torrent-dwelling Litoria (Anura: Hylidae) from the Kikori Spencer, B. 1896. Amphibia. pp. 152-175. in Spencer, B. Integrated Conservation and Development Project area, (ed.), Report on the Work of the Horn Scientific Papua New Guinea. Salamandra 42(4):231-238. Expedition to Central Australia. Part 2. Zoology, Dulau Richards, S. J, Oliver, P. M, Dahl, C. and Tjaturadi, B. and Co., London, UK. 2006. A new species of large green treefrog (Anura: Spencer, B. 1901. Two new species of frogs from Hylidae: Litoria) from northern New Guinea. Zootaxa Victoria. Proceedings of the Royal Society of Victoria (online PRINO) 1208:57-68. 13:176-178. Richards, S. J., Oliver, P. M., Krey, K. and Tjaturadi, B. Steindachner, F. 1867. Reise der österreichischen 2009. A new species of Litoria (Amphibia: Anura: Fregatte Novara um die Erde in den Jahren 1857, 1858, Hylidae) from the foothills of the Foja Mountains, Papua 1859 unter den Bafehlen des Commodore B. von Province, Indonesia. Zootaxa (online PRINO):2277:1-13. Wüllerstorf-Urbair. Zologischer Theil. 1. Amphibien. Ride, W. D. L. (ed.) et al. (on behalf of the International Wien: K. K. Hof- und Staatsdruckerei. Zoologie 1(4):1-70. Commission on Zoological Nomenclature) 1999. Stejneger, L. 1916. New generic name for a tree-toad International code of Zoological Nomenclature. The from New Guinea. Proceedings of the Biological Society Natural History Museum - Cromwell Road, London SW7 of Washington 29:86.

Straughan, I. R. 1969. The Hyla bicolor complex (Anura, Tyler, M. J. 1985. The first collections of frogs from South Hylidae) in North Queensland. Proceedings of the Royal Australia. Mitteilungen aus dem Zoologischen Museum in Society of Queensland 80:43-54. Berlin 61:335-337. Stuart, S. N., Hoffmann, M., Chanson, J., Cox, N., Tyler, M. J. 1992. Encyclopedia of Australian Animals: Berridge, R., Ramani, P. and Young, B. (eds.). 2008. Frogs. The Australian Museum / National Photographic Threatened Amphibians of the World. International Union Index of Wildlife. Angus and Robertson, Pymble, NSW, for the Conservation of Nature, Lynx Editions, Barcelona, Australia:109 pp. Spain; Gland. Switzerland; Conservation International, Tyler, M. J. and Anstis, M. A. 1975. Taxonomy and Arlington, Virginia, USA. biology of the frogs of the Litoria citropa complex (Anura : Thomson, S. A., Littlejohn, M. J., Robinson, W. A. and Hylidae). Rec. South Aust. Mus. (Adelaide) 17:41-50. Osborne, W. S. 1996. Taxonomy of the Litoria aurea Tyler, M. J. and M. Anstis, 1983. Replacement name for complex: a re-evaluation of the Southern Tableland Litoria glandulosa Tyler and Anstis 1975 (Anura: populations of the Australian Capital Territory and New Hylidae). Transactions of the Royal Society of South South Wales. Australian Zoologist 30(2):158-169. Australia 107:129-130. Tschudi, J. J. von. 1838. Classification der Batrachier mit Tyler, M. J. and Davies, M. M. 1977. A new species of Berücksichtigung der fossilen Thiere dieser Abtheilung hylid frog from northern Australia. Copeia 1977:620-623. der Reptilien. Neuchâtel: Petitpierre. Tyler, M. J. and Davies, M. M. 1978. Species groups Tyler, M. J. 1962. New hylid frog from the Central within the Australopapuan hylid frog genus Litoria Highlands of New Guinea. Records of the South Tschudi. Australian Journal of Zoology 63 Australian Museum 14:253-257. (supplement):1-47. Tyler, M. J. 1963a. A taxonomic study of amphibians and Tyler, M. J., and Davies, M. M. 1979. A new species of reptiles of the central highlands of New Guinea with notes cave-dwelling hylid frog from Mitchell Plateau, Western on their ecology and biology II Ranidae and Hylidae. Australia. Transactions of the Royal Society of South Transactions of the Royal Society of South Australia Australia 103:149-153. 86:105-130. Tyler, M. J. and Davies, M. M. 1983. A new species of Tyler, M. J. 1963b. An account of collections of frogs from Litoria (Anura: Hylidae) from Irian Jaya, New Guinea. Central New Guinea. Records of the Australian Museum Copeia:803-808. 26:113-130. Tyler, M. J. and Davies. M. M. 1985. A new species of Tyler, M. J. 1964a. An investigation of the systematic Litoria (Anura: Hylidae) from New South Wales, Australia. position and synonymy of Hyla montana Peters and Doria Copeia 1985:145-149. (Anura, Hylidae). Zoologische Abhandlungen. Staatliches Tyler, M. J. and Davies. M. M. 1986. Frogs of the Museum für Tierkunde in Dresden 27:265-270. Northern Territory. Conservation Commission of the Tyler, M. J. 1964b. Two new species of Australian frogs of Northern Territory, Darwin, NT, Australia:77 pp. the hylid genus Nyctimystes. Transactions of the Royal Tyler, M. J., and Dobson, J. 1973. On the identity, Society of South Australia 88:111-114. authorship and fate of the type specimens of Rana Tyler, M. J. 1964c. Results of the Archbold Expedition. caerulea. Herpetologica 29:373-375. No. 85. A new hylid frog from the Eastern Highlands of Tyler, M. J. and Knight, F. 2009. Field Guide to the Frogs new Guinea. American Museum Novitates 2187:1-6. of Australia. CSIRO Publishing, Collingwood, Victoria, Tyler, M. J. 1965. Taxonomic studies of some hylid frogs Australia. of Australia and New Guinea. Proceedings of the Tyler, M. J. and Martin, A. A. 1977. Taxonomic studies of Zoological Society of London 145:91-106. the some Australian leptodactylid frogs of the genus Tyler, M. J. 1967. A new species of frog of the hylid genus Cyclorana Steindachner. Records of the South Australian Nyctimystes from the highlands of New Guinea. Museum 17:261-276. Transactions of the Royal Society of South Australia Tyler, M. J. and Parker, F. 1972. Additions to the hylid 91:191-195. frog fauna of New Guinea, with description of a new Tyler, M. J. 1968a. A taxonomic study of hylid frogs of the species, Litoria timida. Transactions of the Royal Society Hyla lesueuri complex occurring in north-western of South Australia 96:157-163. Australia. Records of the South Australian Museum Tyler, M. J. and Parker, F. 1974. New species of hylid and 15:711-727. leptodactylid frogs from southern New Guinea. Tyler, M. J. 1968b. Papuan hylid frogs of the genus Hyla. Transactions of the Royal Society of South Australia Zoologische Verhandelingen:1-203. 98:71-77. Tyler, M. J. 1968c. An additional Australian species of the Tyler, M. J., Martin, A. A. and Watson, G. F. 1972. A new hylid frog genus Nyctimystes with notes on Hyla dayi species of hylid frog from New South Wales. Proceedings Günther. Arkiv för Zoologi. Stockholm 2(20):501-504. of the Linnean Society of New South Wales 97:82-86. Tyler, M. J. 1969. A synopsis of the frogs of the genus Tyler, M. J., Davies, M. and Martin, A. A. 1977. A new Hyla of north-western Australia, with the description of a species of large, green tree frog from northern Western new species. Records of the South Australian Museum Australia. Trans. R. Soc. S. Aust. 101:133-138. 16:1-11. Tyler, M. J., Davies, M. and King, M. 1978a. The Tyler, M. J. 1971. The phylogenetic significance of vocal Australian frog Chiroleptes dahlii Boulenyer: its sac structure in hylid frogs. University of Kansas systematic position, morphology, chromosomes and Publications. Museum of Natural History 19:319-360. distribution. Trans. R. Soc. S, Aust. 102:17-23.

Tyler, M. J., Davies, M. and Martin, A. A. 1978b. A new Dresden 13:1-15. species of hylid frog from the Northern Territory. Wandolleck, 0. 1911. Dic Amphibien und Reptilien der Transactions of the Royal Society of South Australia papuanischen Ausbeute Dr Schlaginhaufens. 102:151-157. Abh. Ber. K. Zool. AnlhropoL-Mhnol. Afus. Dresden Vanderduys, E. 2012. Field guide to the Frogs of 13(6):1-15. Queensland. CSIRO, Collingwood, Victoria, Australia:192 Watson, G. F., Loftus-Hills, J. and Littlejohn, M. J. 1971. pp. The Litoria ewingi complex (Anura: Hylidae) in south- Tyler, M. J., Davies, M. M. and Martin, A. A. 1981. New eastern Australia. Aust. Journal of Zoology 19:401-416. and rediscovered species of frogs from the Derby- Watson, G. F., Littlejohn, M. J., Hero, J. -M. and Broome area of Western Australia. Records of the Robertson, P. 1991. Conservation Status, Ecology and Western Australian Museum 9:147-172. Management of the Spotted Tree Frog (Litoria spenceri). Tyler, M. J., Davies, M. and Martin, A. A. 1982. Biology, Arthur Rylah Institute Technical Report Series No. 116. morphology and distribution of the Australian fossorial Department of Conservation and Environment: frog Cyclorana cryptotis (Anura: Hylidae). Copeia, Heidelberg, Victoria, Australia. 1982:260-264. Wells, R. W. and Wellington, C. R. 1985. A classification Tyler, M. J., Davies, M. M. and Aplin, K. P. 1986. A new of the Amphibia and Reptilia of Australia. Australian stream-dwelling species of Litoria (Anura: Hylidae) from Journal of Herpetology, Supplementary Series, (1):1-61. New Guinea. Transactions of the Royal Society of South Weijola, W., Vahtera, V., Koch, A., Schmitz, A. and Kraus, Australia 110:63-67. F. 2020. Taxonomy of Micronesian Tyler, M. J., Smith, L. A. and Johnstone, R. E. 1994. monitors (Reptilia: Squamata: Varanus): endemic status Frogs of Western Australia (Revised edition). Western of new species argues for caution in pursuing eradication Australian Museum, Perth, Western Australia, plans. R. Soc. Open Sci. 7:1-28. Australia:187 pp. Werner, F. 1898. Vorläufige Mittheilung über die von Van Beurden, E. and McDonald, K. R. 1980. A new Herrn Prof. F. Dahl im Bismark-Archipel gesammelten species of Cyclorana (Anura: Hylidae) from northern Reptilien und Batrachier. Zool. Anzeiger 21:552-556. Queensland. Transactions of the Royal Society of South Werner, F. 1901. Beschreibung neuer Frösche aus Australia 104:193-195. Bolivia, Ostindien und Neu-Guinea. Zoologischer Vanderduys, E. 2012. Field guide to the frogs of Anzeiger 24:97-103. Queensland. CSIRO, Collingwood, Victoria, Australia:192 White, A. M., Whitford, R. W. and Mahony, M. J. 1994. A pp. new species of Litoria (Anura: Hylidae) from eastern Van Kampen, P. N. 1906. Amphibien. Nova Guinea. Australia. Proceedings of the Linnean Society of New Leiden 5:163-180. South Wales 114:3-10. Van Kampen, P. N. 1909. Die Amphibienfauna von Neu- White, J. 1790. Journal of a Voyage to New South Wales Guinea, nach der Ausbeute der niederlänischen Süd- with Sixty-five Plates of Non-descript Animals, Birds, Neu-Guinea Expeditionen von 1904-1905 und 1907. Lizards, Serpents, Curious Cones of Trees and Other Nova Guinea. Leiden 9:31-49. Natural Productions. J. Debrett, London, UK. Van Kampen, P. N. 1919. Die Amphibienfauna von Neu- Wichmann, A. 1912. Entdeckungsgeschichte von Neu- Guinea. Bijdragen tot de Dierkunde/ Contributions to Guinea (1885 bis 1902). Nova Guinea 2:607-608. Zoology. Amsterdam 21:51-56. Wiens, J. J., Kuczynski, C. A., Hua, X. and Moen, D. S. Van Kampen, P. N. 1923. Amphibia of the Indo-Australian 2010. An expanded phylogeny of treefrogs (Hylidae) Archipelago. Brill: Leiden. based on nuclear and mitochondrial sequence data. Van Tuijl, L. 1995. Revised catalogue of the type Molecular Phylogenetics and Evolution, 55:871-882. specimens of recent amphibians and reptiles in the Withers, P. C. 1993. Metabolic depression during “Zoölogisch Museum”, University of Amsterdam, The aestivation in the Australian frogs, Neobatrachus and Netherlands. Bulletin Zoölogisch Museum, Universiteit Cyclorana. Australian Journal of Zoology, 41:467-473. van Amsterdam 14:125-144. Withers, P. C. 1995. Cocoon formation and structure in Victorian Civil and Administrative Tribunal (VCAT). 2015. the aestivating Australian desert frogs, Neobatrachus and Hoser v Department of Environment Land Water and Cyclorana. Australian Journal of Zoology, 43:429-441. Planning (Review and Regulation) [2015] VCAT 1147 (30 Withers, P. C. 1998. Evaporative water loss and the role July 2015, judgment and transcript). of cocoon formation in Australian frogs. Australian Vogt, T. 1912. Beitrag zur Reptilien und Amphibienfauna Journal of Zoology, 46:405-418. der Südseeinseln. Sitzungsberichte der Gesellschaft Woodruff, D. S. 1972. Australian anuran chromosome Naturforschender Freunde zu Berlin 1912:1-13. numbers. Herpetological Review 4:208. Wagler, J. 1830. Natürliches System der Amphibien, mit Wüster, W. 2020. Hate post on ICZN List server posted vorangehender Classification der Säugthiere und Vogel. on 6 May 2020 at 7:44 AM UK time. Ein Beitrag zur vergleichenden Zoologie. J. G. Cotta, Zaczek, Z. 2019. Former PM Kevin Rudd says Australia München, Stuttgart and Tübingen. should drastically increase its population to 50 MILLION Wandolleck, B. 1910. Die Amphibien und Reptilien der to make sure nation can defend itself in the face of papuanischen Ausbeute Dr. Schlaginhaufens. threats from China. Daily Mail (Australia) (27 November), Abhandlungen und Berichte des Zoologischen und published online at: https://www.dailymail.co.uk/news/ Anthropologisch-Ethnographischen Museums zu article-7729689/Kevin-Rudd-says-Australia-increase- Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 187

population-50-MILLION-defend-China.html Zweifel, R. G. 1960. Results of the 1958-1959 Gilliard Zweifel, R. G. 1956. Results of the Archbold Expeditions New Britain Expedition 3. Notes on the frogs of New No. 72. Microhylid frogs from New Guinea, with Britain. Amer. Mus. Novit. 2023:1-27. descriptions of new species. Amer. Mus. Novit. 1766:1- Zweifel, R. G. 1980. Results of the Archbold Expeditions 49. No. 103. Frogs and lizards from the Huon Peninsula, Zweifel, R. G. 1958. Results of the Archbold Papua New Guinea. Bulletin of the American Museum of Expeditions No. 78. Frogs of the Papuan hylid Natural History:387-434. genus Nyctimystes. Amer. Mus. Novit. Zweifel, R. G. 1983. Two new hylid frogs from Papua 1896:1-51. New Guinea and a discussion of the Nyctimystes papua species group. American Museum Novitates:1-21. Note: PRINO is an acronym for “Peer reviewed in name only”. Such publications are in effect NOT peer reviewed and therefore material published in such journals must be treated with a high degree of scepticism. The worst offender in the Herpetology space is the online “journal” Zootaxa, which has no effective peer review although being aggressively marketed as being peer reviewed. CONFLICTS OF INTEREST None.

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

Australasian Tree Frog (Family Pelodryadidae Günther, 1858), genus and species list (incl. page numbers for descriptions).

PELODRYADIDAE GUNTHER, 1858

ADELYNHOSERHYLEINI TRIBE NOV. … 151 Paramitrolysis subgen. nov. … 21 Adelynhoserhylea gen. nov. … 10 M. verrucosa (Tyler and Martin, 1977) (type species) Adelynhoserhylea adelynhoserae sp. nov. (M. verrucosa inornata subsp. nov.) … 22 (type species) … 10 Invisibiliaauris subgen. nov. … 22 A. myola (Hoskin, 2007) M. cryptotis (Tyler and Martin, 1977) (type species) A. serrata (Andersson, 1916) M. flavoranae sp. nov. … 23 A. exophthalmia (Tyler, Davies and Aplin, 1986) M. leucodorsalinea sp. nov. … 24 Yikesanura subgen. nov. … 10 Neophractops Wells and Wellington, 1985 … 25 A. yikes sp. nov. (type species) … 11 Neophractops platycephalus (Günther, 1873) A. eucnemis Lönnberg, 1900 (type species) Jackyhoserhylea gen. nov. … 11 N. occidentalis (Anstis, Price, Dale Roberts, Catalano, Jackyhoserhylea genimaculata (Horst, 1883) Hines, Doughty and Donnellan, 2016) (type species) N. rosea sp. nov. … 25 J. ernieswilei sp. nov. … 13 Crottyanura gen. nov. … 26 J. jackyhoserae sp. nov. … 14 Crottyanura crottyi sp. nov. (type species) … 27 LEUCODIGIRANINA SUBTRIBE NOV. … 152 C. dahlii (Boulenger, 1896) Leucodigirana gen. nov. … 15 RANOIDEINA SUBTRIBE NOV. … 154 Leucodigirana andiirrmalin McDonald, 1997 Ranoidea Tschudi, 1838 … 29 (monotypic) Ranoidea aurea (Lesson, 1831) (type species) Euscelis Fitzinger, 1843 … 16 Sandgroperanura subgen. nov. … 29 Euscelis lesueurii (Dümeril and Bibron, 1841) R. cyclorhyncha (Boulenger, 1892) (type species) (type species) R. moorei (Copland, 1957) E. booroolongensis (Moore, 1961) Chirodryas Keferstein, 1867 … 30 (E. booroolongensis dorsaruber subsp. nov.) … 16 Chirodryas raniformis Keferstein, 1867 (type species) (E. booroolongensis occultatum subsp. nov.) … 17 C. castanea (Steindachner, 1867) E. jungguy (Donnellan and Mahony, 2004) C. sloppi sp. nov. … 30 E. wilcoxi (Günther, 1864) GEDYERANINA SUBTRIBE NOV. … 155 Gedyerana gen. nov. … 31 COGGERDONIANI TRIBE NOV. … 152 Gedyerana gedyei sp. nov. (type species) … 32 Coggerdonia Wells and Wellington, 1985 … 18 G. dayi (Günther, 1897) Coggerdonia adelaidensis (Gray, 1841) (type species) Mosleyia Wells and Wellington, 1985 … 33 Mosleyia nannotis (Andersson, 1916) (type species) CYCLORANININI TRIBE NOV. … 152 M. cottoni sp. nov. … 34 Cyclorana Steindachner, 1867 … 19 M. lorica (Davies and McDonald, 1979) Cyclorana novaehollandiae Steindachner, 1867 Amnisrana subgen. nov. … 34 (type species) M. rheocola (Liem, 1974) (type species) C. australis (Gray, 1842) M. michaelsmythi sp. nov. … 35 Mitrolysis Cope, 1889. … 19 M. nyakalensis (Liem, 1974) Mitrolysis alboguttata (Günther, 1867) (type species) M. piloti sp. nov. … 36 (M. alboguttata dumptrashensis subsp. nov.) … 20 M. brevipes (Peters, 1871) DARANINANURINI TRIBE NOV. … 155 M. cultripes (Parker, 1940) Daraninanura gen. nov. … 37 M. longipes (Tyler and Martin, 1977) Daraninanura brevipalmata M. maculosa (Tyler and Martin, 1977) (Tyler, Martin and Watson, 1972) (type species) M. maini (Tyler and Martin, 1977) M. manya (van Buerden and Macdonald, 1980) FIACUMMINGANURINI TRIBE NOV. … 156 M. vagitus (Tyler, Davies and Martin, 1981) Fiacumminganurea gen. nov. … 38 Fiacumminganurea fiacummingae sp. nov. (type species) … 38 F. spenceri (Dubois, 1984) F. timdalei sp. nov. … 40

DRYOPSOPHINA SUBTRIBE NOV. … 157 RAWLINSONINA SUBTRIBE NOV. … 158 Dryopsophus Fitzinger, 1843 … 42 Rawlinsonia Wells and Wellington, 1985 … 61 Dryopsophus citropa (Dümeril and Bibron, 1841) Rawlinsonia ewingi (Duméril and Bibron, 1841) (type species) (type species) (D. citropa gippslandensis subsp. nov.) … 43 R. alpina (Fry, 1915) Leucolatera subgen. nov. … 42 R. corbeni Wells and Wellington, 1985 D. subglandulosa (Tyler and Anstis, 1983) R. jervisensis (Duméril and Bibron, 1851) (type species) R. littlejohni (White, Whitford and Mahoney, 1994) D. daviesae (Mahony, Knowles, Foster and R. paraewingi Donnellan, 2001) (Wilson, Loftus-Hills and Littlejohn, 1971) Ausverdarana subgen. nov. … 42 R. revelata (Ingram, Corben and Hosmer, 1982) D. phyllochroa (Günther, 1863) (type species) R. ventrileuco sp. nov. … 61 D. barringtonensis (Copland, 1957) R. verreauxii (Duméril, 1853) D. jarrodthomsoni sp. nov. … 44 D. kroombitensis (Hoskin, Hines, Meyer, MAXINEHOSERRANINI TRIBE NOV. … 159 Clarke and Cunningham, 2013) Maxinehoserranae gen. nov. … 62 D. nudidigita (Copland, 1962) Maxinehoserranae bicolor (Gray, 1842) D. pearsoniana (Copland, 1961) (type species) D. piperata (Tyler and Davies, 1985) M. albolabris (Wandolleck, 1911) M. bibonius (Kraus and Allison, 2014) KUMANJAYIWALKERUS TRIBE NOV. … 157 M. brettbarnetti sp. nov. … 63 Kumanjayiwalkerus gen. nov. … 44 M. chloristona (Menzies, Richards and Tyler, 2008) Kumanjayiwalkerus kumanjayi sp. nov. M. eurynastes (Menzies, Richards and Tyler, 2008) (type species) … 45 M. lodesdema (Menzies, Richards and Tyler, 2008) K. rothii (De Vis, 1884) M. maxinehoserae sp. nov. … 66 Pengilleyia Wells and Wellington, 1985 … 46 M. mystax (Van Kampen, 1906) Pengilleyia tyleri (Martin, Watson, Gartside, M. piersoni sp. nov. … 67 Littlejohn and Loftus-Hills, 1979) (type species) M. viranula (Menzies, Richards and Tyler, 2008) P. amboinensis (Horst, 1883) Vegrandihyla subgen. nov. … 63 P. darlingtoni (Loveridge, 1945) M. contrastens (Tyler, 1968) P. everetti (Boulenger, 1897) Angularanta gen. nov. … 69 P. obtusirostris (Meyer, 1875) Angularanta arfakiana (Peters and Doria, 1878) P. peronii (Tschudi, 1838) A. becki (Loveridge, 1943) AUDAXURINA SUBTRIBE NOV. … 158 A. brongersmai (Loveridge, 1945) Audaxura gen. nov. … 47 A. bulmeri (Tyler, 1968) Audaxura congenita A. chydaeus sp. nov. … 73 (Peters and Doria, 1878) (type species) A. communia sp. nov. … 74 A. capitula (Tyler, 1968) A. extentacrus sp. nov. … 75 A. pygmaea (Meyer, 1875) A. flavescens (Kraus and Allison, 2004) A. quadrilineata (Tyler and Parker, 1974) A. longicrus (Boulenger, 1911) Brevicrusyla gen. nov. … 48 A. lutea (Boulenger, 1887) Brevicrusyla wisselensis (Tyler, 1968) (type species) (A. lutea leucopunctata subsp. nov.) … 79 B. umbonata (Tyler and Davies 1983) A. milneana (Loveridge, 1945) Colleeneremia Wells and Wellington, 1985 … 48 A. mukherjii sp. nov. … 76 Colleeneremia rubella, (Gray, 1842) (type species) A. oenicolen (Menzies and Zweifel, 1974) C. bogfrog sp. nov. … 50 A. pratti (Boulenger, 1911) C. chunda sp. nov. … 52 A. quaeinfernas sp. nov. … 77 C. dunnyseat sp. nov. … 54 A. solomonis (Vogt, 1912) C. electrica (Ingram and Corben, 1990) A. thesaurensis (Peters, 1877) C. watdat sp. nov. … 56 A. vulgarans sp. nov. … 78 C. wifi sp. nov. … 58 A. wollastoni (Boulenger, 1914) Balatusrana subgen. nov. … 50 Alliuma subgen. nov. … 70 C. dentata (Keferstein, 1868) (type species) A. spartacus (Richards and Oliver, 2006) (C. dentata toowoombaensis subsp. nov.) … 60 (type species) A. spinifera (Tyler, 1968) A. macki (Richards, 2001)

Longuscrusanura subgen. nov. … 71 N. chrisdahli (Richards, 2007) A. napaea (Tyler, 1968) N. hilli (Hiaso and Richards, 2006) Naveosrana subgen. nov. … 71 N. humboldtorum (Günther, 2006) A. dorsivena (Tyler, 1968) (type species) Rotundaura gen. nov. … 99 A. fuscula (Oliver and Richards, 2007) Rotundaura jeudei (Werner, 1901) (monotypic) Raucus subgen. nov. … 72 Variabilanura gen. nov. … 99 A. impura (Peters and Doria, 1878) Variabilanura iris (Tyler, 1962) (type species) A. oxyeei sp. nov. … 80 V. tomcottoni sp. nov. … 102 Scelerisqueanura subgen. nov. … 72 V. majikthise (Johnston and Richards, 1994) A. louisiadensis (Tyler, 1968) V. mareku (Günther 2008) (type species) (including subspecies) V. mucro (Menzies, 1993) (A. brunetus subsp. nov.) … 81 V. ollauro (Menzies, 1993) Bellarana gen. nov. … 82 Sudesanura subgen. nov. … 100 Bellarana angiana (Boulenger, 1915) V. havina (Menzies, 1993) B. angularis (Loveridge, 1945) DRYMONTANTINA SUBTRIBE NOV. … 165 B. megalops (Richards and Iskandar, 2006) Drymomantis Peters, 1882 … 103 B. micromembrana (Tyler, 1963) Drymomantis fallax (Peters, 1882) (type species) Fluvirana gen. nov. … 83 D. ausviridis sp. nov. … 103 Fluvirana rara (Günther and Richards, 2005) D. celantur sp. nov. … 105 (type species) D. cooloolensis (Liem, 1974) F. rivicola (Günther and Richards, 2005) D. glauerti (Copland, 1957) F. scabra (Günther and Richards, 2005) D. northdstradbrokeensis sp. nov. … 106 Hopviridi gen. nov. … 83 D. olongburensis (Liem and Ingram, 1977) Hopviridi leucova (Tyler, 1968) (type species) H. chloronota (Boulenger, 1911) NYCTIMYSTINI TRIBE NOV. … 165 Incertanura gen. nov. … 84 Nyctimystes Stejneger, 1916 … 107 Incertanura pronimia (Menzies, 1993) (type species) Nyctimystes papua (Boulenger, 1897) (type species) I. cuspis sp. nov. … 85 N. disruptus Tyler, 1963 I. fakfakensis sp. nov. … 86 N. mondoensis sp. nov. … 109 I. pinocchio (Oliver, Günther and Richards, 2019) N. oktediensis Richards and Johnston, 1993 I. vivissimia (Oliver, Richards and Donnellan, 2019) N. tyleri Zweifel, 1983 Inlustanura gen. nov. … 87 Magnummanibus subgen. nov. … 108 Inlustanura multiplica (Tyler, 1964) (type species) N. pulchra (Wandolleck, 1910) (type species) I. gasconi (Richards, Oliver, Krey and Tjaturadi, 2009) N. charlottae sp. nov. … 110 I. inluster sp. nov. … 87 N. doggettae sp. nov. … 110 Moechaeanura gen. nov. … 88 Asperohyla subgen. nov. … 109 M. nigropunctatus (Meyer, 1874) (type species) N. trachydermis Zweifel, 1983 (type species) M. albatermacula sp. nov. … 92 N. aspera sp. nov. … 111 M. biakensis (Günther, 2006) N. georgefloydi sp. nov. … 112 M. christianbergmanni (Günther, 2008) Albogibba gen. nov. … 114 M. rubrops (Kraus and Allison, 2004) Albogibba humeralis (Boulenger, 1912) (type species) M. triton sp. nov. … 91 A. ingens sp. nov. … 117 M. umarensis (Günther, 2008) Ratiobrunneis subgen. nov. … 115 M. vocivincens (Menzies, 1972) A. zweifeli (Tyler, 1967) M. wapogaensis (Richards and Iskandar, 2001) A. granti (Boulenger, 1914) Aspercutis subgen. nov. … 89 Occultatahyla gen. nov. … 117 M. verae (Günther, 2004) (type species) Occultatahyla semipalmatus (Parker, 1936) M. singadanae (Richards, 2005) (type species) Telaater subgen. nov. … 90 O. avocalis (Zweifel, 1958) M. richardsi (Dennis and Cunningham, 2006) O. daymani (Zweifel, 1958) (type species) O. fluviatilis (Zweifel, 1958) M. spica sp. nov. … 92 O. foricula (Tyler, 1963) Ornatanura gen. nov. … 93 O. perimetri (Zweifel, 1958) Ornatanura modica (Tyler, 1968) Webpede subgen. nov. … 119 O. leucopicturas sp. nov. … 94 O. rueppelli (Boettger, 1895) O. parscinereo sp. nov. … 95 Nigreosoculus gen. nov. … 120 O. parsviridis sp. nov. … 97 Nigreosoculus cheesmani (Tyler, 1964) Nasuscuspis gen. nov. … 98 N. bivocalis (Kraus, 2012) Nasuscuspis prora (Menzies, 1969) (type species) N. calcaratus (Menzies, 2014)

N. eucavatus (Menzies, 2014) Mahonabatrachus Wells and Wellington, 1985 … N. intercastellus (Kraus, 2012) 134 N. kuduki (Richards, 2007) Mahonabatrachus meiriana (Tyler, 1969) N. latratus (Menzies, 2014) (type species) N. montana (Peters and Doria, 1878) M. aurifera N. obsoletus (Lönnberg, 1900) (Anstis, Tyler, Roberts, Price and Doughty, 2010) N. persimilis (Zweifel, 1958) M. chriswilliamsi sp. nov. … 134 N. traunae (Menzies, 2014) M. marionanstisae sp. nov. … 136 BADIOHYLINA SUBTRIBE NOV. … 167 M. pailsae sp. nov. … 138 Badiohyla gen. nov. … 121 M. roypailsi sp. nov. … 140 Badiohyla kubori (Zweifel, 1958) (type species) SALMOCULARANINA SUBTRIBE NOV. … 172 B. gularis (Parker, 1936) Salmocularana gen. nov. … 142 Magnumoculus gen. nov. … 123 Salmocularana personata Magnumoculus narinosus (Zweifel, 1958) (Tyler, Davies and Martin, 1978) (type species) N. ocreptus (Menzies, 2014) S. saxacola sp. nov. … 142 N. myolae (Menzies, 2014) S. staccato (Doughty and Anstis, 2007) N. cryptochrysos (Kraus, 2012) Litoria Tschudi, 1838 … 143 Litoria freycineti Tschudi, 1838 (type species) PELODRYANINI TRIBE NOV. … 168 L. latopalmata Günther, 1867. Pelodryas Günther, 1858 … 124 Paralitoria gen. nov. … 144 Pelodryas caerulea (White, 1790) (type species) Paralitoria nigrofrenata (Günther, 1867) (type species) P. cavernicola (Tyler and Davies, 1979) Ferelitoria subgen. nov. … 144 P. splendida (Tyler, Davies and Martin, 1977) P. watjulumensis (Copland, 1957) (type species) P. gilleni (Spencer, 1896) P. spaldingi (Hosmer, 1964) SHIREENHOSERHYLINA SUBTRIBE NOV. … 169 Quasilitoria gen. nov. … 145 Shireenhoserhylea gen. nov. … 124 Quasilitoria inermis (Peters, 1867) (type species) Shireenhoserhylea chloris (Q. inermis davidtribei subsp. nov.) … 146 (Boulenger, 1893) (type species) (Q. inermis dunphyi subsp. nov.) … 147 S. gracilenta (Peters, 1869) Q. axillaris (Doughty, 2011) S. luteiventris (Ogilby, 1907) Q. coplandi (Tyler, 1968) S. megaviridis sp. nov. … 125 Q. mickpughi sp. nov. … 147 S. xanthomera (Davies, McDonald, and Adams, 1986) Q. mippughae sp. nov. … 148 Emeraldhyla subgen. nov. … 125 Q. pallida (Davies, Martin and Watson 1983) S. kumae (Menzies and Tyler, 2004) (type species) Q. tornieri (Neiden, 1923) S. aruensis (Horst, 1883) (Q. tornieri serventyi subsp. nov.) … 149 S. auae Menzies and Tyler, 2004 Vultusamolitoria subgen. nov. … 145 S. bella Q. nasuta (Gray, 1842) (type species) (McDonald, Rowley, Richards and Frankham, 2016) Q. peninsulae (De Vis, 1884) S. callista (Kraus, 2013) S. elkeae (Günther and Richards, 2000) SAGUNURINI TRIBE NOV. … 174 S. eschata (Kraus and Allison, 2009) Sagunura Wells and Wellington, 1985 … 149 S. robinsonae (Oliver, Stuart-Fox and Richards 2008) Sagunura burrowsae (Scott, 1942) (type species) Summaviridis gen. nov. … 127 Summaviridis vagabunda WOWRANAINI TRIBE NOV. … 174 (Peters and Doria, 1878) (monotypic) Wowrana gen. nov. … 150 . Wowrana dux PUSTULATARANINI TRIBE NOV. … 170 (Richards and Oliver, 2006) (type species) Pustulatarana gen. nov. … 127 W. graminea (Boulenger, 1905) Pustulatarana longirostris (Tyler and Davies, 1977) W. nullicedens (Kraus, 2018) (type species) W. pallidofemora (Kraus, 2018) (P. longirostris tozerensis subsp. nov.) … 130 W. pterodactyla Llewellynura Wells and Wellington, 1985 … 130 (Oliver, Richards and Donnellan, 2018) Llewellynura microbelos (Cogger, 1966) (type species) W. sauroni (Richards and Oliver, 2006) L. fukker sp. nov. … 132 Parawowrana subgen. nov. … 150 L. yehbwudda sp. nov. … 133 W. hunti (Richards, Oliver, Dahl and Tjaturadi. 2006) Microlitoria subgen. nov. … 131 (type species) L. dorsalis (Macleay, 1877) (type species) SANDYRANINA SUBTRIBE NOV. … 174 L. timida (Tyler and Parker, 1972) Sandyrana Wells and Wellington, 1985 … 150 Sandyrana infrafrenata Günther, 1867 (type species)

Hoser 2020 - Australasian Journal of Herpetology 44-46:1-192. S. militarius (Ramsay, 1878) Hoser, R. T. 2020. For the first time ever! An overdue review and reclassification of Australasian Tree Frogs S. multicolor (Günther, 2004) (Amphibia: Anura: Pelodryadidae), includingAvailable formal online at www.herp.net descriptions of 12 tribes, 11 subtribes, 34 genera, 26 S. purpureolatus subgenera, 62 species andCopyright- 12 subspecies new Kotabi to science. Publishing (Oliver, - All rightsRichards, reserved Tjaturadi, and Iskandar, 2007) Australasian Journal of Herpetology 44-46:1-192. S. sanguinolenta (Van Kampen, 1909) AustralasianAustralasian JournalJournal ofof HerpetologyHerpetology