Research Article |
Corresponding author: Stephen J. Richards ( steve.richards@adelaide.edu.au ) Academic editor: Uwe Fritz
© 2022 Stephen J. Richards, Paul M. Oliver.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Richards SJ, Oliver PM (2022) A new species of insular treefrog in the Litoria thesaurensis species group from the Nakanai Mountains, New Britain, Papua New Guinea. Vertebrate Zoology 72: 1067-1076. https://doi.org/10.3897/vz.72.e91422
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The Islands of East Melanesia have a unique and highly endemic frog fauna derived entirely from overseas colonisation events. Within East Melanesia New Britain is a notable centre of frog diversity and endemism, with at least 15 endemic species, mostly in the ceratobatrachid genus Cornufer. Here we describe the first endemic pelodryadid treefrog from New Britain. The new species is a member of the Litoria thesaurensis species group but can be distinguished from near relatives by aspects of body size, webbing extent, bone pigmentation and male advertisement call. The two known specimens of the new species were collected in Hill Forest on karst basement in the Nakanai Mountains in East New Britain. The new species provides new evidence of diversification of insular PelodrydidaePelodrydidae, and reinforces New Britain, and especially the predominantly karst Nakanai mountains, as a hotspot of frog diversity in East Melanesia. In light of high rates of forest loss and conversion New Britain is also a region of significant conservation concern.
East Melanesia, forest loss, island endemism, karst endemism, overwater dispersal, Pelodryadidae, taxonomy
The islands of East Melanesia (also sometimes in part referred to as Northern Melanesia) extend from Manus in the west through New Ireland, New Britain and the Solomon Islands in the east, and under some interpretations through to Vanuatu and Fiji (
The frog fauna of East Melanesia shows a particularly marked differentiation from that of nearby mainland New Guinea, being dominated by a single radiation of direct-developing species in the family Ceratobatrachidae that has diversified greatly in ecology and associated body form (
The most diverse and widespread lineage of Litoria Tschudi, 1838 in East Melanesia comprises two species in the L. thesaurensis species group. This is a group of moderately-sized treefrogs with variable colouration and extent of finger webbing (
The new species described herein and its putative relatives are assigned to the genus Litoria based on having a horizontal pupil following
The type material of the new species was fixed in 10% formalin and subsequently stored in 70% ethanol and lodged at the South Australian Museum. Comparative material was examined in the collections of the Natural History Museum, London (
Measurements (to the nearest 0.1 mm) were taken with dial calipers and a stereomicroscope fitted with an ocular micrometer. Morphological measurements largely follow
Advertisement calls were recorded with a Marantz PMD-661 Solid-state Recorder and Sennheiser ME66 microphone at distances of 0.5–2 m. Ambient temperatures were noted during recordings. Calls were analysed using Avisoft-SASLab Pro (v4.34, available from Avisoft Bioacoustics: http://www.avisoft.com/) following procedures and terminology recommended by
A species of Litoria that can be distinguished from all other taxa by the following unique combination of characters: moderately small size (male SVL 38.7–41.7 mm); finger webbing moderate, not extending beyond base of second phalanx on any fingers; toe webbing moderate, extending to base of second phalanx on both sides of toe 4; finger discs moderately expanded (3FD/SVL 0.059–0.062); toe discs moderately expanded (4TD/SVL 0.052–0.058); bones of limbs green in life; lower forelimbs and hindlimbs with low crenulated dermal ridges along lateral edges; male throat and neck smooth or ridged, but lacking numerous obvious round bumps in life; venter plain off-white in life; dorsal colouration mottled pale grey or brown with scattered very small dark-brown flecks; head lacking dark lateral band or mask; iris reddish-brown with prominent sky blue outer rim; and call very long (> 13 s), consisting of more than 90 notes repeated at a rate of 6.7 notes/s.
Habitus moderately robust (Fig.
Fingers moderately long, relative lengths 3>4>2>1, terminal discs moderately wide (3FD/SVL 0.062), with circum-marginal grooves (Fig.
Toes moderately short, relative lengths 4>5>3>2>1, terminal discs moderately wide (4TD/SVL 0.058) with circum-marginal grooves (Fig.
In life at night dorsal and upper-lateral surfaces of torso varying shades of mottled light to medium brown (Fig.
In preservative dorsal and upper lateral surfaces of head and torso mottled with varying shades of light brown and light grey, anterior portion of head distinctly greyer than remainder of dorsal surfaces, scattered small dark-brown flecks on dorsal surfaces of torso clearly apparent, with smaller number of small white flecks. Exposed surfaces of limbs show similar base colouration to dorsum, but also have areas of off-white mottling and lack dark-brown flecks. Vent bordered dorsally by dark brown blotch and then irregular white band. Dorsal surfaces of digits buff with varying amounts of fine brown stippling, stippling especially dense on outer digits, but tending to very sparse on inner digits. Ventral surfaces buff, largely unpatterned, except for small areas of dense stippling close to lower jaw, and along extremities of forelimbs and hindlimbs.
SVL 41.7; TL 22.1; HW 15.3; HL 14.8; EYE 5.3; TYM 2.9; IN 3.3; EN 4.2; 3FD 2.6; 3FP 1.4; 4TD 2.4; 4TP 1.8.
The sole paratype (
Details of species in the Litoria thesaurensis Group in life. A Litoria insularis sp. nov. paratype
We recorded a single complete call produced by the holotype at an air temperature of 22.0°C. It was recorded from approximately 2 m distance and the call of this species is very quiet so detailed analysis of some features was not possible due to reduced resolution and high background noise. The call is a long (13.9 s) series of 94 short, distinctly pulsed notes repeated rapidly (6.7 notes/s) at relatively uniform inter-note intervals of 0.106–0.144 s (mean = 0.120, SD = 0.012, n = 25). Twenty-five notes measured in detail are 0.027–0.050 s long (mean = 0.034, SD = 0.005) and contain 3–5 pulses (3 pulses = 4 notes, 4 pulses = 17 notes, 5 pulses = 4 notes). Energy in the call is distributed broadly between ~0.5 and 3 kHz and dominant frequency is between 1260 and 2156 Hz in five randomly selected notes. The call produced by the holotype is illustrated and compared with two calls of a sympatric L. thesaurensis in Fig.
Waveform (top) and spectrogram (bottom) of (A) an entire call of Litoria insularis sp. nov.; B a three second section from the middle of this call illustrating the distinctly pulsed nature of the notes; C two consecutive calls of L. thesaurensis recorded in sympatry with the new species, showing strikingly different call structure. The strong signal at ~2.2 s on A) is a call of the New Britain endemic microhylid frog Oreophryne brachypus.
Litoria insularis sp. nov. was found in sympatry with L. thesaurensis and is most likely to be confused with that species, but differs in having wider finger discs (3FD/SVL 0.059–0.062 versus 0.037–0.052) and toe discs (4TD/SVL 0.052–0.058 versus 0.038–0.047); dorsum with scattered small dark-brown flecks (versus base colouration and pattern highly variable, but usually not with small scattered dark-brown flecks) (Fig.
It has been suggested that Litoria thesaurensis as currently construed may be a complex of several species (
The new species differs from other species in the L. thesaurensis species group as follows; from L. flavescens Kraus & Allison, 2004 by its smaller size (male SVL 38.7–41.7 mm versus 44.5–45.6), by having green limb bones (versus white), and by having a light grey and/or brown dorsum (versus yellow); from L. lutea by its smaller maximum size (adult male maximum 42 mm versus 62 mm), and dorsum with scattered small dark-brown flecks (versus dark brown flecks absent (Fig.
Only two other species of Litoria occur in East Melanesia. Litoria insularis sp. nov. can be readily distinguished from these as follows: from L. infrafrenata (Günther, 1867) by its much smaller size (male SVL 41.2 mm versus up to at least 135 mm), mottled brown or grey dorsal colouration (versus typically plain green or brown), and by lacking a prominent white labial strip (versus present); and from L. lododesma Menzies, Richards & Tyler, 2008 by its much larger size (max male SVL 41.2 mm versus 23.8 mm), dorsum brown or grey with scattered brown flecks (versus typically green without pattern), and the absence of white lateral stripe on the head (versus usually present).
Known only from the type locality in the Nakanai Mountains in the eastern portion of the island of New Britain (Fig.
The type series was collected in relatively undisturbed Hill Forest (Fig.
Litoria insularis is known only from a single locality despite moderately intense survey effort elsewhere in New Britain, suggesting that it may have a localised distribution. The type locality is immediately adjacent to a road that has been constructed into the interior of the island from Pomio on the coast and this will undoubtedly improve access to, and increase pressure on, the surrounding forest. Rainforests across New Britain are also under threat from rapid expansion of Oil Palm plantations so, given the combination of very limited knowledge about the new species’ distribution, ecology and habitat requirements, and potential threats, we recommend that it be listed as Data Deficient under the IUCN Redlist Criteria.
The name insularis alludes to both the new species’ likely endemic status on the island of New Britain, and the potential that it is further restricted to the Nakanai Mountains, which are themselves a ‘sky island’.
Litoria insularis sp. nov. is currently known only from a single site in upper Hill Forest, and we consider that it is probably endemic to New Britain, and potentially to the Nakanai Mountains. Although numerous endemic ceratobatrachid frogs have been described from New Britain, Litoria insularis sp. nov. provides the first evidence of a pelodryadid treefrog endemic to this island. This new species brings the number of endemic frogs on New Britain to 16 species, the highest number of any island in East Melanesia (Bougainville [including adjacent Buka]) is closest with seven endemic species) (
The new species also serves to emphasise the important role that island colonisation and diversification has played in the evolution of the Litoria thesaurensis species group. While not species rich, this radiation now includes two species that are only known from islands that have never had connections with the much more species-rich frog fauna of nearby New Guinea (the other being L. lutea from Bougainville and the Solomon Islands). Acoustic evidence further suggests that the diversity of insular endemic taxa in this clade remains underestimated (
Fieldwork in Papua New Guinea by SJR was approved by the PNG National Research Institute, the PNG Department of Environment and Conservation (now Conservation and Environment Protection Authority) and the East New Britain Provincial Government and would not have been possible without the generous support of Florence Pasparea of East New Britain’s Environment and Conservation Management Committee and Alois Magogo of Pomio Local Level Government. We are particularly grateful to the communities of Irena, Marmar and Muro villages for their hospitality and assistance. Rose Singadan and Paulus Kei provided assistance at the University of PNG and we are also grateful to C. Kovach, D. Capone and M. Hutchinson (South Australian Museum), Barry Clarke (
Material examined
Litoria lutea (n = 11):
Litoria thesaurensis (n = 40):