Papuahyla Donnellan, Mahony & Richards, 2025

Donnellan, Stephen C., Mahony, Michael J., Esquerré, Damien, Brennan, Ian G., Price, Luke C., Lemmon, Alan, Lemmon, Emily Moriarty, Günther, Rainer, Monis, Paul, Bertozzi, Terry, Keogh, J. Scott, Shea, Glenn M. & Richards, Stephen J., 2025, Phylogenomics informs a generic revision of the Australo-Papuan treefrogs (Anura: Pelodryadidae), Zoological Journal of the Linnean Society 204 : -

Papuahyla Donnellan, Mahony & Richards , gen. nov.

( Fig. 26)

ZooBank LSID: urn:lsid:zoobank.org:act:D17E72ED-A24F-49F0-B9FE-45A99516009D Type species: Litoria chloristona Menzies, Richards & Tyler, 2008 .

Content: Nine species— Papuahyla albolabris ( Wandolleck, 1911) comb. nov., Papuahyla bibonius * ( Kraus and Allison, 2004a) comb. nov., Papuahyla chloristona * ( Menzies, Richards & Tyler, 2008) comb. nov., Papuahyla contrastens ( Tyler, 1968a) comb. nov., Papuahyla eurynastes ( Menzies, Richards & Tyler, 2008) comb. nov., Papuahyla lodesdema * ( Menzies, Richards & Tyler, 2008) comb. nov., Papuahyla louisiadensis * ( Tyler, 1968a) comb. nov., Papuahyla mystax ( Van Kampen, 1906) comb. nov., Papuahyla rubrops * ( Kraus and Allison, 2004b) comb. nov.

Diagnosis: Papuahyla can be diagnosed from the other members of the Drymomantis Sub-clade as follows: from Amnihyla by a Type 1 vs. Type 3 tadpole oral disc, Type 1 vs. Type 6 or 7 overall tadpole morphology; and by a right triangular vs. fusiform, oval, rectangular, spike, or teardrop call envelope shape. It can be diagnosed further from A. amnicola by unornamented vs. prominent tubercles on the hindlimb. It can be diagnosed from Exedrobatrachus by unornamented vs. tubercules on hindlimb, by a right triangular vs. fusiform call envelope shape, and 13 sites in the mitochondrial ND4 alignment ( Table 3); from Exochohyla by absence of the rostral spike, unornamented vs. tubercles or crenulations on hindlimb, small vs. large egg size, a Type 1 vs. Type 1A tadpole oral disc, and Type 1 vs. Type 6 overall tadpole morphology; from Hyalotos by a pigmented vs. transparent tympanum, unornamented vs. crenulations on hindlimb, small vs. medium or large eggs, and by a right triangular vs. teardrop call envelope shape; from Ischnohyla by a Type 1 vs. Type 3 tadpole oral disc, Type 1 vs. Type 2A overall tadpole morphology, and by a right triangular vs. left triangular, left teardrop, or oval call envelope shape; from Kallistobatrachus by unornamented vs. tubercles or crenulations on hindlimb, a Type 1 vs. Type 1B tadpole oral disc, Type 1 vs. Type 7 overall tadpole morphology, and by a right triangular vs. fusiform, spike, or teardrop call envelope shape; from Lathrana by unornamented vs. tubercules on hindlimb, small vs. medium eggs, and by a right triangular vs. oval call envelope shape, and a note rate change across the call vs. none; from Nasutibatrachus and Teretistes by the absence of a rostral spike; further from Teretistes by full or reduced vs. minimal toe webbing, small vs. large eggs, and a Type 1 vs. Type 3 tadpole oral disc; from Viridihyla by ossified vs. cartilaginous intercalary structures, small vs. large eggs, and by a right triangular vs. fusiform, left teardrop, or rectangular call envelope shape. Papuahyla can be diagnosed by a right triangular call envelope shape vs. a spike-fusiform in Carichyla and fusiform-spike in Drymomantis , respectively ( Table 2), and from Carichyla and Drymomantis by 13 and 16 sites, respectively, in the mitochondrial ND4 alignment ( Table 3). Refer to Tables 1, 2, and 3. Diagnosis of Papuahyla from all other genera in the Drymomantis Sub-clade is supported by 94 sites distributed across eight AHE loci (Supporting Information, AHE loci diagnostic sites).

Distribution and ecology: New Guinea and surrounding islands, Bismarck and Admiralty Archipelagos, and the Maluku and East Nusa Tenggara provinces of Indonesia. Arboreal frogs that are found in open permanent or seasonal grassy, reedy, or sago swamps in natural or altered habitats, usually not in closed forests ( Menzies 2006, Anstis 2017), with most species in lowlands, except for D. contrastens which is a New Guinean highland species.

Etymology: Papua is a non-Latin non-Greek place name, a noun in apposition to a second noun ( Hyla ), without any need to modify either word. See etymology for Amnihyla above for the derivation of Hyla .

Remarks: Papuahyla is the equivalent to part of the Litoria bicolor Group and to the Litoria albolabris Group of Tyler and Davies (1978).

In the absence of molecular genetic data and clear evidence of affinity from morphology for Hyla mystax Van Kampen, 1906 and Hyla albolabris Wandolleck, 1911 , we placed these two taxa conservatively in Papuahyla .

For Hyla mystax, Menzies (2006) discussed its relationships with the Litoria bicolor assemblage. Tyler (1968a) compared it with other green in life New Guinean species within or near to the size range of H. mystax which served to distinguish it from these species without providing direction for its affinities. Tyler and Davies (1978) placed it, without discussion, in the Litoria bicolor Group. Richards and Donnellan (2023) demonstrated that H. mystax is morphologically similar to Kallistobatrachus beryllinus .

Hyla albolabris Wandolleck, 1911 is a small green species with spotted ventral surfaces and a SVL of 20–22mm ( Menzies 2006). Most of the original specimens are lost and the remaining two are in such poor condition being extremely dehydrated, distorted, and brittle that comparison with other species is not achievable ( Menzies 2006). Tyler (1968a) discussed the original series in detail and concluded it may have been composite as there appears to have been at least two size classes represented. Tyler and Davies (1978) placed it in their Litoria albolabris Group.

Two species, Hyla contrastens and Litoria eurynastes , that lack genetic data presently are conservatively included in Papuahyla because they were considered to be closely related to L. bicolor in their original descriptions ( Tyler 1968a, Menzies et al. 2008) and these associations have not been refuted subsequently, and because they are more geographically allied with Papuahyla being remote from Drymomantis that is exclusively Australian and do not share distribution with Carichyla which in New Guinea only occurs in the Trans-Fly region. Papuahyla eurynastes is likely to be a species complex ( Menzies et al. 2008).