Amphibia, Anura, Microhylidae)

Zootaxa 3641 (3): 271–281 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3641.3.6 http://zoobank.org/urn:lsid:zoobank.org:pub:EBED01BD-3791-4B8C-BC77-51A66FBCD198 From a dwarf to a giant: Revalidation of Callulops valvifer (Barbour, 1910), (Amphibia, Anura, Microhylidae)

RAINER GÜNTHER Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt-University, Invalidenstr. 43, 10115 Berlin, Germany

Abstract

According to the most recent revision of the subfamily Asterophryinae, the species Pomatops valvifera was considered to be a synonym of Phrynomantis (now Callulops) robusta. On the basis of recently collected material from near the type locality of Pomatops valvifera on the Bomberai Peninsula in western New Guinea, the invalidity of the genus name is confirmed but the species name is revalidated. Callulops valvifer (new combination) was hitherto unequivocally known from a single specimen of less than 30 mm snout-vent length. With a length of more than 70 mm for males and of more than 80 mm for females, this species is now among the largest of the currently known 22 species of the genus Callulops.

Key words: Pomatops valvifera; Liophryne kampeni; morphology; bioacoustics, Fakfak Mountains; western New Guinea

Introduction

The genus Pomatops, and its sole species P. valvifera, were erected by Barbour in 1910 to accommodate a microhylid frog of less than 30 mm snout-vent length that was found in the stomach of a natricine snake, Tropidonophis mairii (Gray, 1841). This snake was collected by A. E. Pratt at Fak Fak (today also spelled Fakfak) on the Bomberai Peninsula, then north-western Dutch New Guinea, today Papua Province of Indonesia. As the main feature to diagnose the new genus, Barbour (1910) referred to “a flap of skin extending for some distance anterior and posterior to the position of the eye. These flaps are sufficiently developed so that they may be laid down and thus completely cover the whole eye”. The monotypic genus Pomatops was considered valid by Van Kampen (1923) and Nieden (1926), but Parker (1934) synonymised it with Asterophrys. He correctly found that the diagnostic features given by Barbour do not apply. However, he maintained the species Asterophrys valvifera and considered Liophryne kampeni Boulenger, 1914 as a synonym of that species. Asterophrys valvifera was considered valid also by Loveridge (1948) and by Zweifel (1956). In his revision of the subfamily Asterophryinae Zweifel (1972) synonymised Asterophrys valvifera and Liophryne kampeni with Phrynomantis robusta. Dubois (1988) noted that the genus name Phrynomantis was preoccupied by a South African frog genus and he replaced it with the name Callulops Boulenger, 1888, for the at-present 22 Papuan taxa (Frost 2013, Günther et al. 2012). According to Zweifel (1972) Phrynomantis robusta, now Callulops robustus, “is widely distributed throughout New Guinea from southeastern islands to the Vogelkop Peninsula”. The opinion that all frogs determined to be Callulops robustus from the mainland of New Guinea really belong to this species was doubted, for example, by Menzies (1975, 2006), Günther (2003), Kraus & Allison (2009), and Oliver et al. (2012). Kraus (2012) recently confined the areal distribution of Callulops robustus to the type locality of Misima Island (Louisiade Archipelago, east of the eastern end of mainland New Guinea) and resurrected Mantophryne microtis Werner, 1901, as a valid species of Callulops. If this taxonomic action proves to be correct, all frogs from mainland New Guinea that were hitherto treated as Callulops robustus belong to other species. New species of Callulops from the main island of New Guinea were described, after Zweifel´s (1972) revision of the genus, by Richards et al. (1995), Kraus & Allison (2003, 2009), Oliver et al. (2012), and Günther et al. (2012). Although Oliver et al. (2012) and Kraus (2012) suggested that Pomatops valvifera could be a valid taxon, they lacked material to define this taxon.

Accepted by M. Vences: 6 Mar. 2013; published: 24 Apr. 2013 271 A detailed redescription of Pomatops valvifera in its appropriate genus Callulops will be given below on the basis of recently collected material at the type locality. This is necessary because of the very cursory and misleading original description by Barbour (1910), based moreover on only a single juvenile specimen.

Redescription and revalidation

Callulops valvifer (Barbour, 1910), new combination (the name valvifera was changed here to valvifer in order to choose the correct Latin suffix for the species name in combination with the masculine genus name Callulops).

Original name: Pomatops valvifera Barbour, 1910: 89; Type locality: “Fak Fak, Northwestern Dutch New Guinea”;

Pomatops valvifera (van Kampen 1923: 147); Pomatops valvifera (Nieden 1926: 63); Asterophrys valvifera (Parker 1934: 64); Asterophrys valvifera (Loveridge 1948: 417); Asterophrys valvifera (Zweifel 1956: 8); Phrynomantis robusta (Zweifel 1972: 494); Callulops robustus (Dubois 1988: 4); Callulops robustus (Menzies 2006: 244); Callulops robustus (Frost 2001); Callulops robustus (Kraus 2012: 257).

Material and methods

During a collection trip in the Fakfak Mountains, northwestern tip of the Bomberai Peninsula, together with native colleagues I collected two large Callulops specimens and recorded calls of this species: Field number 7877=ZMB 70532, female, collected on 8 September 2008, 700 m a.s.l., 2°51´S and 132°17É; and FN 7924=ZMB 70533, male, collected on 11 September 2008, 500 m a.s.l., 2°52´S and 132°17É. Moreover, I studied a juvenile specimen that was collected by D. Telnov on 24 September 2010 in the Fakfak Mountains, 900 m NN, 2°50´06´´S and 132°18´22´É. This specimen is stored in the collection of the Naturkundemuseum Erfurt (Germany), inventory number NME A 1925/13. The male frog was detected at night by tracking its advertisement calls and the female was also found at night with the aid of head-torches. Both frogs were anaesthetized with chlorobutanol, fixed in 2–3 % formalin in the field for three weeks and transferred to 75% ethanol for permanent storage. Samples of thigh muscle tissue were extracted from both specimens before fixation and stored in 95% ethanol to permit DNA analyses. The juvenile specimen was fixed in 80% ethanol and muscle tissue from a thigh was extracted two years after fixation. Measurements of SUL, TaL, and TL (defined below) to the nearest 0.1 mm were made with digital callipers, and all others were made with an ocular micrometer fitted to a dissecting microscope. Abbreviations are as follow:

SUL snout-urostyle length, from tip of snout to distal tip of urostyle bone (SUL and snout-vent length differ insignificantly, but SUL is more accurately measured); TL tibia length, outer distance between knee and heel (with calliper gently pressed); TaL length of tarsus, outer distance with tarsal and heel joints held at right angles; L4T length of fourth toe, from tip of toe to proximal end of inner metatarsal tubercle; L3F length of third finger (to proximal end of inner metacarpal tubercle); F1D transverse diameter of first finger disc; F3D transverse diameter of third finger disc; T1D transverse diameter of first toe disc;

272 · Zootaxa 3641 (3) © 2013 Magnolia Press GÜNTHER T4D transverse diameter of fourth toe disc; HL head length, from tip of snout to posterior margin of tympanum; HW head width, taken in the region of the tympana; END distance from anterior corner of orbital opening to centre of naris; IND internarial distance between centres of nares; ED eye diameter, from anterior to posterior corner of orbital opening; TyD horizontal diameter of tympanum; SL snout length, from an imaginary line connecting the centres of the eyes to tip of snout; EST distance from eye corner to tip of snout.

Calls were recorded with a Sony Digital Audio Tape (DAT) Walkman TCD-D 100 and a Sennheiser microphone MKE 300 and analysed with Avisoft-SASLab Pro (v4.34) software. Genomic DNA isolation and mitochondrial fragment amplification (12S and 16S rRNA) were performed according to Köhler & Günther (2008). Forward and reverse strands were aligned using CodonCode Aligner v. 3.5.7 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye. The consensus sequences were aligned using MAFFT (Katoh & Toh 2008); substitution models were obtained from jModeltest v. 0.1.1 ((Posada 2008); AIC and BIC; 12S rRNA: GTR+G, 16S rRNA: GTR+G). Phylogenetic analyses of the combined dataset (671 base pairs of the 12S rRNA gene and 488 base pairs of the 16S gene) were performed using maximum parsimony (MP) as implemented in PAUP* v. 4.0b010 (Swofford 2002), maximum likelihood (ML) using RAxML Blackbox (Stamatakis et al. 2008), and Bayesian inference (BI) using MrBayes v. 3.1.2 (Ronquist & Huelsenbeck 2003). MP parameters: heuristic search with 10 random additions (maximum number of saved trees), tree bisection and reconstruction (TBR) branch swapping, number of bootstrap replicates = 100. ML parameters: bootstrap replicates = 100; BI parameters: 1,000,000 generations, sample frequency = 20, number of chains = 4, burnin value = 35,001. Comparisons of Callulops valvifer with congeners are based on material stored in the collection of the Museum für Naturkunde, Berlin (ZMB) and on data published in the respective original descriptions or comparative studies (Zweifel 1972, Burton 1986, Menzies 2006). Diagnosis. With a SVL of 72.3 mm (male) and 81.4 mm (female) the species is among the largest species of the genus. Legs fairly short (TL/SUL 0.41–0.43), eyes medium-sized (ED/SUL 0.111–0.118). All fingers and toes with well developed circum-marginal grooves, the discs of fingers the same size or smaller than that of toes (T4D/ F3D 1.0–1.23). Internarial distance notably wider than distance eye-naris (END/IND 0.75–0.81). Dorsal surfaces of juveniles blackish with whitish spots, that of adults uniformly brownish. Advertisement call consists of a series of 4–8 very loud, harsh, croaking notes; note length (except first note, which is nearly always distinctly shorter) 192–261 milliseconds (ms), note repetition rate 1.24–1.30 notes per second (s). Dominant frequency at 1.1 kHz. Redescription (based on the three above-mentioned recently collected specimens). For measurements and ratios see Table 1. Head not distinct from body, only slightly more narrow than body. Snout truncate in dorsal view, rounded and slightly protruding in lateral view. Canthus rostralis curved and rounded; loreal region concave; nares near snout tip, directed laterally. Distance between nares greater than distance between nostril and eye (END/IND 0.75–0.81). Eye medium-sized (ED/SUL 0.111–0.118) in adult, larger (ED/SUL 0.148) in juvenile. Tympanum clearly visible with supratympanal fold extending to half way between tympanum and forearm insertion. Extremities robust, with strong muscles especially on hind legs. Inner metacarpal tubercle well expressed, all other palmar and subarticular tubercles large but indistinct; digital discs notably wider than penultimate phalanges, all with terminal grooves; no webs between fingers or toes; relative length of fingers 3>4>2>1. Inner metatarsal tubercle well developed, no outer one; subarticular tubercles on toes more distinct than those on fingers; all toe discs wider than penultimate phalanges and with terminal grooves; about same width or wider than finger discs (F3D/T4D 1.0–1.23); relative length of toes 4>3>5>2>1. Skin smooth on all dorsal and ventral surfaces. Colour of dorsal surfaces of the adult male in preservative uniformly dark brown, that of the female medium brown. Lower surface uniformly light grey-brown in the female (compare Fig. 1) and slightly darker in the male; throat and chest in both specimens darker than the remaining ventral surfaces. Female with vague lumbar ocelli. Dorsal base of all finger discs and toe discs with a light spot. Dorsal colour of the adult specimens in life uniformly brown, flanks and ventral surfaces light grey. The male in life was slightly darker than the female. Iris golden with blackish venation; a light-blue patch on anterior and posterior corner of eye in the female. From comparisons with obviously closely related adult and juvenile specimens of Callulops wondiwoiensis, recently described by Günther et al. (2012), it appears that the specimen NME A 1925/13 is a juvenile Callulops

REVALIDATION OF CALLULOPS VALVIFER Zootaxa 3641 (3) © 2013 Magnolia Press · 273 and resembles other juveniles of the C. “robustus-group”. It seems to be characteristic for species of the C. robustus-group that juveniles show another colouration than adults (see also Kraus 2012). Comparisons of the 12S and 16S genes from this juvenile specimen with those of the adults confirmed the conspecifity of all three Fakfak frogs (Fig. 5). The juvenile frog exhibits a different colouration and a different eye size compared to the adults. Most biometric characters, however, coincide with that of the adults (Table 1). The most important differences between the juvenile and the adults are in colouration. Dorsal and lateral surfaces of the juvenile are dark brown, probably blackish in life, covered with many whitish (possibly yellowish in life) spots and its ventral surfaces are brownish (probably also blackish in life), as well as with numerous whitish spots (Fig. 2). It looks very much like the holotype of Pomatops valvifera described by Barbour (photos of that specimen are available online at the MCZ herp division website). Various authors (van Kampen 1923, Parker 1934, Loveridge 1948, and Oliver et al. 2012) used this obviously juvenile colouration erroneously to distinguish the latter taxon from adult specimens of other Callulops taxa.

TABLE 1. Biometrical values of an adult male (ZMB 70539), an adult female ( ZMB 70532), and a juvenile (NME A 1925/ 13) of Callulops valvifer. All measurements in mm. Abbreviations are explained in the section “Material and methods”.

Inventory-No. ZMB 70533 ZMB 70532 NME A 1925/13 Mean Range SVL 72.3 81.4 61.0 TL 31.1 34.6 26.9 TaL 20.0 21.7 17.5 T4L 30.9 33.7 27.0 T4D 2.2 2.7 2.0 T1D 1.8 2.0 1.8 F3L 18.6 21.4 16.2 F3D 2.2 2.2 1.8 F1D 2.0 2.0 1.5 HL 22.4 25.9 17.6 HW 28.3 32.0 22.5 END 4.5 5.7 3.5 IND 6.0 7.0 4.9 ED 8.5 9.0 5.5 TyD 3.3 5.2 3.4 EST 8.2 9.5 5.2 SL 9.2 10.5 6.9 TL/SUL 0.43 0.43 0.41 0.42 0.41–0.43 TaL/SUL 0.28 0.27 0.27 0.27 0.27–0.28 T4D/SUL 0.030 0.033 0.031 0.031 0.031–0.033 T4D/T1D 1.22 1.35 1.33 1.30 1.22–1.35 F3D/SUL 0.030 0.027 0.031 0.029 0.027–0.031 F3D/F1D 1.10 1.10 1.14 1.11 1.10–1.14 T4D/F3D 1.00 1.23 1.00 1.08 1.00–1.23 HL/SUL 0.31 0.32 0.31 0.31 0.31–0.32 HW/SUL 0.39 0.39 0.38 0.39 0.38–0.39 HL/HW 0.79 0.81 0.82 0.81 0.79–0.82 END/IND 0.75 0.81 0.80 0.79 0.75–0.81 ED/SUL 0.118 0.111 0.148 0.126 0.111–0.148 TyD/ED 0.39 0.58 0.39 0.45 0.39–0.58

274 · Zootaxa 3641 (3) © 2013 Magnolia Press GÜNTHER FIGURE 1. Dorsolateral and ventral view of a female Callulops valvifer (ZMB 70532) in life. This specimen was erroneously designated as a male in a former paper by Günther (2009, Fig. 10a).

FIGURE 3. Landscape along the road from Fakfak Town to Kokas where Callulops valvifer was found.

Neither in the adult nor in the juvenile specimens, studied by myself, is the upper eyelid more strongly developed than in other Callulops frogs. This observation confirms the statement by Parker (1934) and Zweifel (1972) that the genus Pomatops, which is mainly based on that character, has no validity. Distribution and ecological notes. I heard advertisement calls of C. valvifer at 300 m to 900 m a.s.l. along the road from Fakfak Town to Kokas (Fig. 3). This region was formerly almost certainly completely covered by primary rain forest, but today primary and undisturbed forest remain only on steep slopes. Elsewhere in the Fakfak Mountains, primary forest has been selectively or partly deforested or even completely replaced by plantations. Callulops valvifer was found mainly in secondary forest where the canopy was partly closed. It inhabited humus soil interspersed with dead wood and leaves, roots, and boulders of different sizes. All four observed adult

276 · Zootaxa 3641 (3) © 2013 Magnolia Press GÜNTHER specimens (two were collected) were observed near hollows in the ground to which they retreated when people approached. The juvenile was found under a fallen log. The taxonomic affiliation of the specimens listed as A. valvifera (now Callulops valvifer) by Parker (1936) from Mount Mafulu and by Zweifel (1956) from Mount Mafulu and Hollandia (now Jayapura) is unclear. Therefore the range of that taxon for the present seems to be restricted to the Fakfak Mountains. Vocalisation. Calls are very loud and dominate all other nearby amphibian voices. Frogs called mainly in front of burrows in the first hours of darkness. Calls of one male followed one another after gaps of from 50 seconds to several minutes. Three calls from one male could be analysed more exactly. Call duration was 3.08–4.84 s and calls contained 4–6 notes (Fig. 4). Note repetition rate was 1.24–1.30 notes/s. Mean note length of 12 notes (excluding the first) was 243 ms, SD 22.5, range 192–261 ms. Duration of the first note ranged from 69–107 ms. Mean length of 12 internote intervals was 716 ms, SD 120, range 581–988 ms. Mean number of pulses/s among 15 notes was 198, SD 10.8, range 173–213. Dominant frequency was at 1.1 kHz (Fig. 4). Two calls from another male, which was too distant to allow proper call analysis, contained 6 and 8 notes, call duration was 4.63 and 6.28 s, and note repetition rate was 1.27 and 1.30 notes/s. From these data it follows that note length (except for the first note), pulses/s, and note repetition rate were fairly constant and that the last internote interval was always considerably longer than the others. Calls are neither tuned nor with frequency modulations.

FIGURE 4. Waveform (above left ), spectrogram (below left) of call notes 2–4, and power spectrum of note 2 (right), of an advertisement call of Callulops valvifer consisting of four notes, recorded on 11 September 2008, 22 °C.

Comparisons with other species. Callulops valvifer differs from the recently described species of Metamagnusia and Pseudocallulops (Günther 2009) by its more robust body, shorter legs, less expanded finger and toe tips, and discs of toes broader than those of fingers (vice versa in the latter two genera). Callulops glandulosus (Zweifel, 1972), C. sagittatus Richards, Burton, Cunningham & Dennis, 1995, C. stictogaster (Zweifel, 1972), and C. wilhelmanus (Loveridge, 1948), completely lack expanded digital tips and terminal grooves; C. valvifer has expanded digital tips, all with terminal grooves. Callulops biakensis Günther, Stelbrink & von Rintelen, 2012 has a shorter adult body length (53.6–67.8 mm versus 72.3–81.4 mm in C. valvifer), a shorter internarial distance (END/IND 0.67–0.71 versus 0.75–0.81), smaller eyes (ED/SUL 0.90–0.97 versus 0.111–0.118 in adults), and a different note repetition rate (3.06–3.48 notes/s in C. biakensis versus 1.24–1.30 notes/s in C. valvifer). Callulops boettgeri (Méhely, 1901) and C. dubius (Boettger, 1895) from the island of Halmahera, C. eremnosphax Kraus & Allison, 2009 from the Gulf Province of Papua New Guinea, C. humicola (Zweifel, 1972) from Eastern Highlands District of Papua New Guinea, C. kopsteini (Mertens, 1930) from the Sula Islands, and C. mediodiscus Oliver, Richards & Tjaturadi, 2012 from the Southern Highlands Province of Papua New Guinea, all with snout-vent lengths of under 50 mm, are much smaller than C. valvifer. Callulops comptus (Zweifel, 1972) is not longer than 60 mm (SVL), has shorter hind legs (TL/SVL 0.30–0.39 versus 0.41–0.43 ), an orange or pink postocular patch, and a boldly mottled ventral surface.

REVALIDATION OF CALLULOPS VALVIFER Zootaxa 3641 (3) © 2013 Magnolia Press · 277 Callulops doriae Boulenger, 1888 has a tuberculate dorsal surface showing numerous blackish spots, each with a central white cap. Callulops fojaensis Oliver, Richards & Tjaturadi, 2012 is smaller than C. valvifer (male less than 55 mm SVL), with shorter hind legs (TL/SVL 0.34), and has a higher note repetition rate (1.68–1.94 notes per s). Callulops fuscus (Peters, 1867) is smaller than C. valvifer (less than 60 mm SVL versus more than 70 mm), utters advertisement calls with three notes/s and seems to be restricted to Ambon, Seram, and Batanta Islands (Menzies 2006). Liophryne kampeni Boulenger, 1914 from the Mimika River in the southern Papua Province of Indonesia, which certainly belong to the genus Callulops, should no longer be treated as a synonym of C. robustus. This was underlined also by Kraus (2012). Callulops kampeni is smaller (the holotype has a SVL of 55 mm according to Parker, 1934) than C. valvifer, has broader toe tips (according to measurements by Zweifel 1972, the ratio T4D/ F3D is 1.33 in the holotype of C. kampeni, but is 1.00–1.23 in C. valvifer). Moreover, the dorsal outline of the snout of C. kampeni (see Fig. 2B in Kraus & Allison 2009) differs clearly from that of C. valvifer. It is roundish in dorsal view in C. kampeni but truncate in C. valvifer. The proper status of C. kampeni should be formulated when enough material for a sophisticated diagnosis is available. Callulops marmoratus Kraus & Allison, 2003 is smaller , has longer hind legs (TL/SVL 0.45–0.49), larger eyes , and a mottled dorsal surface. Callulops microtis (Werner, 1901) was recently resurrected from synonymy with C. robustus by Kraus (2012). According to his data and my study of the holotype (ZMB 16499), this species differs from C. valvifer mainly by its skin structure and colouration: Flanks and the dorsum (less intensive) are tubercular in the former species but smooth in the latter, and the lateral surfaces have a uniformly grey-brown or red-brown colour in C. valvifer, but are dark-grey with white flecking in C. microtis. Juvenile color pattern is also different from C. valvifer. Callulops omnistriatus Kraus & Allison, 2009 is notably smaller (8 males 55.0–59.6 mm SVL and 10 females 49.9–66.9 mm SVL) than C. valvifer. Body ratios of both are overlapping. It should be mentioned that both species have similar 12S and 16S mtDNA sequences (Fig. 5 and Table 2). More detailed studies must show whether C. omnistriatus and C. valvifer are really distinct species or whether the former is possibly a junior synonym of the latter.

C. wondiwoiensis ZMB 63874 100 98 100 C. wondiwoiensis ZMB 70316 96 97 100 C. wondiwoiensis ZMB 70315 - 52 C. wondiwoiensis ZMB 62037 64 99 - 100 C. biakensis ZMB 64107 72 100 65 C. biakensis ZMB 64108

100 C. yapenensis ZMB 62566 100 100 C. valvifer NME A1925/13 88 80 100 C. valvifer ZMB 70532 96 82 99 C. valvifer ZMB 70533

C. omnistriatus BPBM 32098

100 100 A. turpicola ZMB 62043 100

A. turpicola ZMB 64105

0.04

FIGURE 5. Bayesian phylogram of the concatenated data set (12S rRNA and 16S rRNA). The numbers on branches are maximum parsimony bootstrap values, maximum likelihood bootstrap values, and posterior probabilities of Bayesian inference (from top to bottom).

Species pair 12S 16S C. yapenensis—C. biakensis 2.3 4.5–5.0 C. yapenensis—C. wondiwoiensis 3.0–4.4 7.1–7.6 C. yapenensis—C. valvifer 3.0–3.2 6.9–7.1 C. biakensis—C. wondiwoiensis 2.1–4.3 6.2–6.9 C. biakensis—C. valvifer 2.8–3.0 7.6–7.8 C. wondiwoiensis—C. valvifer 2.8–3.9 7.1–7.6 C. yapenensis—C. omnistriatus 3.6 5.5 C. biakensis—C. omnistriatus 3.4 6.2–6.7 C. valvifer—C. omnistriatus 0.4–0.5 1.9 C. wondiwoiensis—C. omnistriatus 3.2–3.9 6.2–6.4

Callulops personatus (Zweifel, 1972) has an orange or yellow ground colour, which is clearly different from that of C. valvifer (pers. comm. F. Kraus, Febr. 2013), it has, except on the loreal region, a black head, its dorsal surfaces are not uniformly coloured as in C. valvifer, but exhibit a pattern of grey (blackish in life) flecks and blotches, and its belly is pale orange vs. light grey in C. valvifer. Callulops robustus was redefined recently by Kraus (2012). Males have a snout-vent length of from 60 to 65 mm and females of from 60 to 78 mm, and thus are somewhat smaller than C. valvifer. Moreover, C. robustus has less developed digital discs and terminal grooves and most likely occurs, according to Kraus (2012), only on Misima Island in the Louisiade Archipelago. Juveniles of C. robustus differ in colouration from that of C. valvifer. Callulops wondiwoiensis Günther, Stelbrink & von Rintelen, 2012 has a smaller body (49.5–61.5 mm SUL), shorter call notes (mean note length 147 ms versus 243 ms in C. valvifer), a shorter internote interval length (mean 371 ms versus 716 ms), and utters more notes/s (mean 2.17 in C. biakensis and 1.28 in C. valvifer). Callulops yapenensis Günther, Stelbrink & von Rintelen, 2012 is obviously smaller (the single known adult male measures 48.4 mm SUL), has larger eyes (ED/SUL 0.135 versus 0.111–0.118 in adults), mottled dorsal surfaces (versus uniform dorsal surfaces in C. valvifer), shorter call notes and internote intervals, and a clearly higher note repetition rate (2.38–2.80 versus 1.24–1.30 notes/s). Besides morphological and bioacoustic differences between Callulops valvifer, C. yapenensis, C. wondiwoiensis and C. biakensis (all localized in western New Guinea and immediately adjacent islands), there are also sequence differences in the 12S and 16S genes, which support species status for all four forms (Fig. 5).

Acknowledgements

Field work and collection of voucher specimens was permitted by representatives of Belai Besar Konservasi Sumber Daya Alam (KSDA), Sorong, Papua Province of Indonsia (PPI). Marthinus Kapisa (Biak/PPI); Andreas, Frank, and Apner Piahar (Kampung Lusiperi near Fakfak Town/PPI); and Christian Bergmann (Berlin/Germany) helped during field work. Family Piahar also permitted the collection of frogs on their property. B. Stelbrink, T. von Rintelen, and F. Köhler conducted the molecular investigations, and F. Tillack (all Museum für Naturkunde/Berlin) provided technical help. U. Scheidt (Naturkundemuseum Erfurt/Germany) lent the juvenile specimen of Callulops valvifer. F. Kraus (Fort Collins, Colorado/USA) and R. Arndt (Pomona, New Jersey/USA) improved my draft with a number of very helpful comments. To all of them I am deeply grateful.

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