Three new species of Callulops ( Anura : Microhylidae ) from western New Guinea

Various works in recent years have shown that the microhylid genus Callulops is not a monophyletic group, and that the „central species“ Callulops robustus is not, as previously assumed, distributed over the entire island of New Guinea. Further, Callulops robustus apparently does not occur in the northwestern part of the island. This species is there replaced by a number of other species, and at present no sympatric occurrence of species in the C. robustus-group are known. On the basis of our own collections in the Wondiwoi Mountains at the base of the Wandammen Peninsula as well as on the islands of Yapen and Biak (Cenderawasih Bay) in the years 1998 to 2003, we here describe three new species in the Callulops robustus-group.


Introduction
About three-quarters of the Papuan batrachofauna belongs to the family Microhylidae and all Pa puan microhylids are members of the subfamily Aste rophryinae (Frost et al. 2006, Köhler & Günther 2008).The most speciose Papuan asterophryine genus is Oreophryne, with presently about 50 species, followed by Cophixalus with 35 species.The genus Xenorhina contains 29 species, Austrochaperina 25, Callulops 18, and Albericus 15 species.The remaining 16 genera contain fewer than 10 species each (Menzies 2006, Frost 2011).The genus Callulops was revised by zweiFel (1972) under the name Phrynomantis, but this name had to be changed to Callulops for name priority reasons (Dubois 1988).The genus was considered monophyletic by burton (1986), but Köhler & Günther (2008) doubted this view.With the description of the new genera Metamagnusia and Pseudocallulops, Günther (2009) split the former Callulops into three supposedly monophyletic groups.Other recent publications on the genus Callulops are descriptions of new species (Kraus & allison 2003(Kraus & allison , 2009; oliver, richarDs & tjaturaDi 2012) and a re-diagnosis of the type species Callulops robustus connected with a resurrection of Mantophryne microtis werner, 1901 by Kraus (2012).
The senior author collected Callulops frogs in the Wondiwoi Mountains (Wandammen Peninsula), on Yapen Island, on Biak Island, and on the Fakfak Peninsula in the 1990's and in the first decade of this century.All localities are in the Papua Province of Indonesia, in the far western part of the island of New Guinea.Although all these frogs are morphologically similar to Callulops robustus (boulenGer, 1898), there remained serious doubts about their proper allocation.Because of these doubts they were preliminary classified as Callulops cf.robustus, for example by Köhler & Günther (2008).Closer examination of this material, including comparisons of their mtDNA, have shown that they belong to four different species, three of them new to science.Here these new species, from the Wondiwoi Mountains, Biak Island, and Yapen Island (Fig. 1), are formally described.

Material and methods
Most frogs were collected after locating them by their advertisement calls, at night, on the ground, and at the entrances to their burrows; others were found moving on the ground surface.Some specimens were photographed in life the next day.Tissue probes were taken from thigh muscles were taken from some specimens and stored in 75 % ethanol for later DNA sequencing.All specimens were preserved in 2 % formalin in the field and were later transferred to 75 % ethanol in the collection of the Museum für Naturkunde.Two specimens from the Wondiwoi Mountains population (ZMB 70530 and ZMB 70531) were cleared and stained as osteological preparations according to a method modified from DinGerKus & uhler (1977).
Measurements were made only on preserved specimens with a sliding calliper (< 10 mm) or with a binocular dissecting microscope fitted with an ocular micrometer (< 10 mm) to the nearest 0.1 mm, as follows: SUL snout-urostyle length from tip of snout to distal tip of urostyle bone; SUL differs only slightly from snoutvent length (SVL), but is more accurately measured.Both measures will be treated as synonymous in the following text; TL tibia length: external distance between knee and ankle; TaL length of tarsus: external distance, tarsal and ankle joints held at right angles; T4L length of fourth toe: from tip of toe to proximal end of inner metatarsal tubercle; T4D transverse diameter of disc of fourth toe; T1D transverse diameter of disc of first toe; F3L length of third finger; F3D transverse diameter of disc of third finger; F1D transverse diameter of disc of first finger; 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; EST distance from anterior corner of eye to tip of snout; SL snout length, from an imaginary line connecting the centres of the eyes to the tip of snout.
Advertisement calls were recorded under natural conditions with a Sony Digital Audio Tape (DAT) Walkman TCD-D 100 and a Sennheiser microphone MKE 300 and analysed with Avisoft-SAS Lab software.All specimens are currently stored in the Museum für Naturkunde Berlin (ZMB) and given registration numbers of this institution.Part of the material will be transferred to the Museum Zoologicum Bogoriense (MZB) at Cibinong after completion of our studies.
Comparisons of the new species with congeners are mostly based on data published in the respective original descriptions or comparative studies (zweiFel 1972, burton 1986, Menzies 2006).

Diagnosis.
A medium-sized Callulops (snout-urostyle lengths of five adult males 49.5 -57.2 mm, mean 53.4 mm, of four adult females 56.0-61.5 mm, mean 59.0 mm, and of one presumed subadult female 51.9 mm) with short legs (TL/SUL 0.38 -0.42), moderately expanded terminal discs on fingers and toes, those of toes slightly wider than those of fingers (F3D/T4D 0.75 -0.93).All finger and toe discs with terminal grooves.Head broader than long (HL/HW 0.84 -0.93), truncate in dorsal view.Eye medium-sized (ED/SUL  adult (ZMB 62568) females, and three juveniles in the type series.The SUL of the males is 49.5 -57.2 mm, mean 53.4 (SD 3.07) and that of the four adult females is 56.0 -61.5 mm, mean 58.9 mm (SD 2.32).Various body ratios of adult specimens are shown in Tab. 1. Their colouration is very similar to that of the holotype and the paratype shown in Fig. 2 and Fig. 3. Some specimens exhibited in life scattered small yellow spots in the region behind the eyes and/or anterior back, which spots disappeared in preservation fluid.Snout-urostyle lengths of three juveniles was 22.0 -28.5 mm.Juveniles differ significantly from adults.They have a clearly larger eye (ED/SUL 0.132 -0.140 vs. 0.107 -0.125) and smaller tympanum (TyD/SUL 0.045 -0.053 vs. 0.052 -0.065).Moreover, basic colour of dorsal surfaces is black with numerous white spots and irregular white flecks, and with orange or yellow patches on the central and anterior dorsum (Fig. 4a).Ventral surfaces were also blackish with nu merous white spots in life (Fig. 4b).Black colour changed to grey in preserved specimens.
Distribution and ecological notes.We collected C. wondiwoiensis at from 500 m to 700 m a.s.l. and heard them calling in primary rain forest between 300 m and 750 m a.s.l. on the eastern slopes and valleys of the Wondiwoi Mountains at the base of the Wandammen Peninsula.Most adult males, some females, and one of the juvenile specimens were found at the entrances to burrows in humus or stony loam soil.One specimen was excavated at the end of a burrow of one meter length and at a depth of 25 cm.No specimen was found under leaf litter or on vegetation.The new species was fairly common but distances between calling males were pronounced (at minimal distances of about 20 m).While adult specimens did not show an apparent preference for bodies of water, all juveniles were found proximate to a waterfall (Fig. 5).Calling specimens were heard in all months when the senior author visited the Wondiwoi Mountains, i.e. in April, May, June, July, August, and September.
The ovaries of a ripe female (ZMB 70531), collected at the end of July 1998, contained 16 eggs, with a maximum diameter of 4.0 mm.

Vocalisation.
Males called predominantly during late dusk and in the first few hours after dusk; few calls were heard during early dusk and in later night hours.Calls consisted of a series of very loud croaking notes and were emitted at long intervals, i.e. intervals between calls lasted several minutes and longer.They sounded very much like calls of the female mallard duck, Anas plathyrhynchos.No calls were noted during the day.Males called mostly from the entrance to their burrows, sometimes also from within the burrow, or from a few meters away.In some cases antiphony broad (HL/HW 0.90).Head is truncate viewed from above and rounded in profile, nostrils elevated, laterally directed, and near tip of snout.Canthus rostralis rounded, loreal region slightly concave.Eye moderately large (ED/SUL 0.107), its horizontal diameter slightly longer than distance from eye to snout tip (EST/ED 0.95), supratympanal fold strongly developed and reaching from eye to fore arm.Tympanum, except its margins, clearly visible and half the size of eye (TyD/ED 0.49).Internarial distance clearly wider than distance eye-naris (END/IND 0.75).The forelegs are moderately long and heavily built, fingers end as small discs, all with terminal grooves.Relative length of fingers 3 > 2 = 4 > 1, metacarpal tubercles and subarticular tubercles poorly developed.Hind legs fairly short and of heavy build, toe discs larger than finger discs, all discs with terminal grooves.Inner metatarsal tubercle and subarticular tubercles weakly developed, no outer metatarsal tubercle.Relative length of toes 4 > 3 > 5 > 2 > 1.No webs between fingers or toes.All dorsal and ventral surfaces smooth, except for some tubercles on posterior dorsum.Dorsal surfaces uniformly dark brown, abdomen and inferior thighs light grey-brown, chest and throat darker brown than abdomen but lighter than dorsum.Measurements (in mm) of the holotype: SUL 57.2, TL 22.7, TaL 14.9, T4L13.5, T4D 1.9, T1D 1.5, F3L13.5,F3D 1.5,F1D 1.3,HL 19.2,HW 21.4,END 3.6,IND 4.8,ED 6.1,TyD 3.0,EST 5.8,SL 8.7.Morphological variation in the type series.There are five adult males, four adult and one presumed sub-     with grooves; the new species has slightly expanded digital tips, all with terminal grooves.
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)   intends to resurrect the name of the species Pomatops valvifera from synonymy with C. robustus on the base of this material.Differences between P. valvifera and C. wondiwoiensis will be outlined in the planned paper.
Etymology.Named after the collection site of the type series, the Wondiwoi Mountains on the Wandammen Peninsula, Cenderawasih Bay, north-western New Guinea.
Callulops yapenensis sp.nov.Callulops personatus (zweiFel, 1972) has, except on the loreal region, a black head, its dorsal surfaces not uniformly coloured as in C. wondiwoiensis, but with a pattern of grey (blackish in life) flecks and blotches, and its belly is pale orange vs. light grey in the new species.
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 larger than C. wondiwoiensis.Moreover, C. robus tus occurs, according to Kraus (2012), only locality on Misima Island in the Louisiade Archi pelago.
Callulops microtis (werner, 1901) was recently resurrected from synonymy with C. robustus by Kraus (2012).According to his data and our study of the holotype (ZMB 16499), this species differs from C. wondiwoiensis 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 dorsal surfaces have a uniformly grey-brown or red-brown colour in C. wondiwoiensis, but are darkgrey with white flecking in C. microtis.
Considering the fact that Callulops robustus was restricted to Misima Island, Liophryne kampeni boulenGer, 1914 from the Mimika River region in the southern Papua Province of Indonesia, and Pomatops valvifera barbour, 1910 from the Fakfak Mountains, throat of the Vogelkop Peninsula (both now obviously belong in the genus Callulops), can no longer be treated as synonyms of Callulops robustus.Callulops kampeni has broader toe tips (according to measurements by zweiFel 1972, the ratio T4D/F3D is 1.33 in the holotype, which is the only hitherto known specimen of C. kampeni, but is 1.07 -1.29 in C. wondiwoiensis) and a smaller tympanum (TyD/SVL 0.049 vs. 0.052-0.065,and TyD/ED 0.43 vs. 0.44 -0.57).Moreover, the dorsal outline of the snout of C. kampeni (see Fig. 2B in Kraus & Allison 2009) differs clearly from that of C. wondiwoiensis, and odontoids on the vomer are weakly developed in the latter species but are strongly developed in the former (boulenGer 1914).
The senior author in 2008 collected some Callulops frogs in the Fakfak Mountains and in the near future Waira Mountain and on adjacent mountains (Fig. 10).
Vocalisation.Calling behaviour is much the same as in Callulops wondiwoiensis.The advertisement calls of two males were analysed (Fig. 11).Mean call length of seven calls was 3.73 s (SD 1.24), range 1.43 -4.52 s.Calls contained 4 -11 notes, mean 8.4 (SD 2.76) notes per call.Mean note length of 59 notes was 139 ms (SD 21.9), range 60 -163 ms.Mean internote interval length of 52 intervals was 294 ms (SD 37.5), range 247 -431 ms.The note repetition rate in seven calls varied from 2.4 -2.8, mean 2.55 notes/s.The pecially inner metatarsal tubercle more conspicuous than tubercles on fingers and hands.No webs on toes.Relative length of toes 4 > 3 > 5 > 2 > 1.
Distribution and ecological notes.The holotype was found on an eastern slope of Waira Mountain, about 7 km NEN of the village of Kontinuae (Konti), at 620 m a.s.l.This slope was covered by patches of primary and secondary rain forest.Selective logging was then in progress.The forest substrate was mainly clay, loam, and humus, and covered by some leaf litter and dry wood.Ferns and shrubs were abundant in places and sparsely distributed in others.The animal was encountered in front of a burrow and calling from time to time.A few other calling Callulops yapenensis sp.nov.were heard at between 300 and 700 m a.s.l. on   1.2 kHz.This second peak seems not to be obligatory because it was not expressed in the calls of the second specimen.Calls were recorded at an air temperature of 22 °C.
Advertisement call is much like that of C. wondi woiensis and structure of calls and notes are very similar in both species.Differences of single parameters are given in the next paragraph.
Comparisons with other species.Callulops yapen ensis sp.nov.differs from the recently described species of Metamagnusia and Pseudocallulops (Günther 2009), among others, by its more chunky body with shorter legs, less expanded finger and toe tips, and discs of toes broader than those of fingers (vice versa in the latter two genera).pulse repetition rate in 58 notes varied from 110-211, mean 168, pulses/s.The frequency spectrum extends from 0.4 to about 7 kHz with dominant frequency at 2.2 kHz (Fig. 12).There is a second frequency peak at  Callulops kampeni, which is currently treated as a synonym of C. robustus (zweiFel 1972, andFrost 2011) and the taxonomic status of which is to be redetermined, differs from C. yapenensis by its uniform brown dorsal colour, larger size, smaller eye (ED/SUL 0.115 vs. 0.135) different snout shape (see Fig. 2B in Kraus & allison 2009).
Etymology.Named after the collection site of the type series, Yapen Island, Cenderawasih Bay, north-western New Guinea.Diagnosis.With males up to 67.8 mm SUL, a largesized Callulops.Snout not completely truncate as in Callulops fuscus has a dark brown dorsum without pattern (C.yapenensis has a grey dorsum mottled with dark brown), an advertisement call with three notes per second, and seems to be restricted to Ambon, Seram, and Batanta Islands (Menzies 2006).

Callulops biakensis sp. nov.
Callulops humicola has a dark brown dorsum without pattern and a bright yellow or flesh coloured mottled ventrum and lives at higher elevations (1500 -2000 m a.s.l.).
Callulops marmoratus has longer hind legs (TL/SVL 0.45 -0.49 vs. 0.43 in C. yapenensis) and a different advertisement call (significantly longer notes, fewer notes per second, and a lower dominant frequency).
Callulops microtis with SVL of males of 51.6 -58.8 mm is larger than the new species, has a shorter hind leg (TL/SUL 0.35 -0.43 vs. 0.43 in the new species), smaller eye (ED/SUL 0.100 vs. 0.135 and ED/ EST 1.00 -1.30 vs. 1.44), less pronounced terminal grooves on fingers and toes, and its dorsal surfaces are more pustulose than those in C. yapenensis.
Callulops omnistriatus is larger (males 55 -59.6 mm SVL vs. 48.2mm in the new species), and its dorsum is uniformly grey-brown versus mottled in C. yapenensis.
Callulops personatus has, except on loreal region, a black head, and its belly is pale orange vs. light grey in the new species.
Callulops robustus males have a snout-vent length from 60 to 65 mm and females from 60 to 78 mm and with this are evidently larger than C. yapenensis.More over, this species has a uniformly coloured dorsum and is restricted to Misima Island.
The senior author in 2008 collected some Callulops frogs in the Fakfak Mountains which obviously be- from posterior corner of eye to a short distance behind tympanum.Except for this fold, a very fine middorsal fold, and a few minor tubercles on flanks, all dorsal and ventral surfaces smooth.Subarticular tubercles more developed on feet than on hands, three poorly defined metacarpal tubercles and a well expressed inner metatarsal tubercle.Relative length of fingers 3 > 4 > 2 > 1 and relative length of toes 4 > 3 > 5 > 2 > 1.All fingers and toes with well developed grooved terminal discs, those on fingers narrower than those on toes (T4D/ F3D 1.15).and toes without webbing.
All dorsal and lateral surfaces uniformly brown, some tiny tubercles on posterior flanks are darker and with a whitish tip, and tympana are noticeably lighter brown than their surroundings.Ventral surfaces show a mixture of grey and brown, the brownish fractions many other Callulops, but snout tip slightly rounded.Leg fairly long for the genus (TL/SUL 0.43 -0.47).Eye small (ED/SUL 0.90 -0.94).All fingers and toes with medium-sized discs.Dorsal surfaces with inconspicuous tubercles and uniformly dark brown colour.Advertisement call lasts about two seconds, mean note length 117 ms, mean internote interval length 217 ms, and note repetition rate 3.06 -3.48 notes per s.Description of the holotype.Head in the region of tympana only slightly more narrow than body, broader than long (HL/HW 0.85).Snout truncate with slightly rounded tip, canthus rostralis rounded and curved outward superior of the nostril, loreal region slightly sloped and slightly concave.Nostrils situated laterally almost at tip of snout, scarcely visible from above.Supratympanic fold clearly expressed and reaches Distribution and ecological notes.Callulops biak ensis is fairly common on Biak Island.We collected specimens near Biak Town on the south coast, and near the village of Korem on the north coast (Fig. 16).Moreover, we heard males calling on eastern Supiori, at various sites along the north coast between Supiori and Korem, and across the island between Korem and Biak Town.All were at elevations of from 30 m to 400 m a.s.l.All specimens were encountered in secondary forest (in partly primary forest on Supiori) where trees and bushes grew more or less One specimen was in front of a burrow, another, surprisingly, was at about one meter height on a tree trunk, and a couple showed mating behaviour on a predominant on chest and throat.For measurements of the holotype see Table 2. Iris black with minute silvery speckles and a golden inner margin.
Morphological variation in the type series.For mea surements and ratios of the type series see Table 2.Both specimens (two males) from near Biak Town have a greater SUL than male and female from Arwe.In ter est ingly, the largest specimen is a male of 67.8 mm SUL.The only female paratype (Figs. 14 and 15) is gravid, its largest eggs measure 3.8 mm in diameter.Colouration is much the same in all paratypes, except ZMB 64108 which has some very irregular whitish flecks on the dorsum.Vocalisation.As in both former new species, the advertisement call consists of a series of loud croaking notes.Fifteen calls from one male were analysed.Eleven calls belonged to a series which lasted a total of eleven minutes, i.e. this male uttered one call per minute.Shortest time between two successive calls was 40 s.Mean number of notes per call was 7.0 (SD 0.38), range 6 -8.Mean call length was 2.12 s (SD 0.15), range 1.78 -2.52 s.The first note was clearly shorter than all others, and the last note was mostly the longest in all calls.First and last internote interval were longest in all calls (Fig. 17).Typically, the pulse repetition rate was slower at the beginning and at the end of a note, and faster in the middle of a note.Mean note length of 105 notes was 117 ms (SD 26.7), range 37 -152 ms.Mean internote interval length of 90 intervals was 217 ms (SD 28.6), range 169 -320.Note repetition rate varied between 3.06 -3.48, mean 3.3, notes/s (SD 0.11) in 15 calls.Pulse repetition rate varied from 132 -235, mean 196 pulses/s in 105 notes.Frequency band reaches from 0,5 to 7 kHz, dominant frequency at 2.2 kHz (Fig. 18).Calls were recorded at an air temperature of 24 °C.fallen and rotting log.During the last, the male tried several times to crawl under its mate.It could be supposed that the male tried to stimulate its mate that way to later amplexus.Unfortunately, this mating behaviour finished unsuccessfully.Calling was more common during and after rainfall and in the first hours of evening.No calling was heard during the day.Callulops robustus is restricted to Misima Island, at the easternmost extremity of the geographic range of the genus Callulops.
Because we have only a single specimen of Cal lulops yapenensis, statistics of metrical parameters is not possible.The following metrical characters of C. yapenensis are not in the range of variation of C. bi akensis and could be confirmed as real differences if more material becomes available: T4D/SUL 0.031 in C. yapenensis vs. 0.032 -0.035 in C. biakensis; F3D/ F1D 1.27 vs. 1.17 -1.25; T4D/F3D 1.07 vs. Callulops fuscus seems to be resticted to Ambon, Seram, and Batanta Islands in the west of the Vogelkop Peninsula.
Callulops humicola has a reddish ear patch and a strongly mottled ventrum, with no such ear patch and a uniformly coloured ventrum in C. biakensis.
Callulops marmoratus has a mottled dorsum, and it is uniform in C. biakensis.
Callulops personatus has a black head and its belly is pale orange vs. grey-brown in C. biakensis.ference and other head proportions seem to be "good" diagnostic characters.
-Note repetition rate (seven calls from C. yape nen sis and 15 calls from C. biakensis were analysed) was on average 2.55 notes/s (SD 0.17), range 2.38 -2.80 notes/s in C. yapenensis, and 3.30 (SD 0.11), range 3.06 -3.48 notes/s in C. biakensis (t = 12.74, p = 0.000) (Fig. 23).Etymology.The new species is named after its type locality and, to date, the only known site of its occurrence.
Molecular evidence.Callulops omnistriatus from eastern New Guinea has a very similar morphology compared to some species from western New Guinea and was therefore included in our phylogenetic analysis.3.2 -3.9 6.2 -6.4 The phylogenetic tree (Fig. 24) shows a highly supported Callulops clade with C. cf.robustus from the Fakfak Mountains + C. omnistriatus as sister group to a clade comprising the three new species.While each of the new species is well supported (with the exception of C. yapenensis, where only one sequence was available), their relationships are poorly supported.This might possibly be explained by short intervals between divergence events in that clade.However, the considerable morphological and bioacoustic differences between these species are also reflected by genetic p-distance values ranging from 2.1 -4.4 % in 12S and 4.5 -7.8 % in 16S, respectively (Table 3).This range of genetic distances is usually considered indicative of distinct species status in frogs (e.g.Fouquet et al. 2007).

Fig. 1 .
Fig. 1.Map of the Papua Province of Indonesia with type localities of the new taxa from the Wondiwoi Mountains at the base of the Wandammen Peninsula (1), Yapen Island (2), and Biak Island (3).
n 62 (3) 2012 in adults 0.107 -0.125) and internarial distance clearly wider than distance eye-naris (END/IND 0.64 -0.88).Dorsal surfaces grey-brown to reddish-brown with diminutive white dots predominantly on flanks; ventral surfaces light grey-brown, throat and especially chin darker than belly.Iris bronze with strong blackish pigmentation.Mainly located in burrows.Advertisement call consists of a series of 6 -9 loud croaking notes and is very similar to the calls of female mallard duck, Anas plathyrhynchos linnaeus, 1758.Mean note length 147 ms, mean note repetition rate 2.18 notes/s, and dominant frequency around 1.75 kHz .Description of the holotype.The body appears chunky and is wider than the head.Head is shorter than Aligner v. 3.5.7 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye.The 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 (12S + 16S) 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 & huel sen becK 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 pa ra meters: bootstrap replicates = 100; BI parameters: 1,000,000 generations, sample frequency = 20, number of chains = 4, burnin value = 35,001.Types of Callulops microtis and C. fuscus were examined by the authors; other values were taken from the original descriptions and comparative treatments.
Fig. 24 is by B. stelbrinK, all others are by the senior author.
notes/s (n = 12).The pulse repetition rate varied from 119 to 239 per second, mean 189 pulses/s (n = 74).Frequency spectrum reaches from 0.5 to about 6 kHz, with dominant frequency at 1.75 kHz (Fig.7).Calls were recorded at air temperatures of from 21 to 23 °C.Comparisons with other species.Callulops wondi woiensis sp.nov.differs from the recently described species of Metamagnusia and Pseudocallulops (Gün ther 2009), among others, by its more chunky body, of neighbouring males occurred.Advertisement calls of three different males were analysed.The first note was nearly always the shortest, followed in length by the second and the third notes.Notes are clearly pulsed, with interpulse intervals longer in the first half of a note.First and last internote intervals are mostly longer than the other intervals (Figs.5, 6).Mean call length of 12 calls was 3.49 s, minimum 2.43 and maximum 5.47 s.Calls contained 6-9 notes, mean 7.4 notes per call.Note length of 94 notes was 147 ms on average, min.59 ms, max.214 ms.Mean internote length of 71 intervals was 371 ms, min.255 ms, max.909

Fig. 4b .
Fig. 4b.Ventral view of a juvenile specimen of Callulops won di woiensis sp.nov. in life.

Fig. 5 .
Fig. 5. Waterfall and surroundings of the mountain stream "Sungai Maja" at 550 m a.s.l., habitat of Callulops wondiwoiensis sp.nov.and more than 20 other anuran species.

Fig. 7 .
Fig. 7. Power spectrum of an advertisement call note of Cal lu lops wondiwoiensis sp.nov.

Fig. 6 .
Fig. 6.Wave form (above) and spectrogram (below) of first five notes of an advertisement call of Callulops wondiwoiensis sp.nov.consisting of seven notes.

Fig. 10 .
Fig. 10.Surroundings of the Waira Mountain north of the village of Konti, where Callulops yapenensis sp.nov.males were heard calling.
Callulops glandulosus, C. sagittatus, C. stictogaster, and C. wilhelmanus completely lack expanded digital tips with grooves, while the new species has slightly expanded digital tips, all with terminal grooves.Callulops boettgeri, C. dubius, C. eremnosphax, and C. kopsteini are clearly smaller than C. yapenensis.Callulops comptus has shorter hind legs (TL/SVL 0.30 -0.39 versus 0.43), an orange or pink postocular patch, and a boldly mottled ventral surface.Callulops doriae has a larger body (SVL up to 100 mm), and a verrucous dorsal surface showing numerous blackish spots, each with a central white cap.Callulops fojaensis has shorter hind legs (TL/SVL 0.34), longer call notes (mean 225 ms versus 139 ms in C. yapenensis), and a lower dominant frequency (1.3 kHz versus 2.2 kHz).

Fig. 12 .
Fig. 12. Power spectrum of an advertisement call note of Cal lu lops yapenensis sp.nov.

Fig. 17 .
Fig. 17.Wave form (above) and spectrogram (below) of an advertisement call of Callulops biakensis sp.nov.consisting of eight notes.

Fig. 18 .
Fig. 18.Power spectrum of an advertisement call note of Cal lulops biakensis sp.nov.

Fig. 24 .
Fig. 24.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).

Table 1 .
Body ratios of five adult males, four adult females, and one subadult female of the type series of Callulops won diwoiensis sp.nov.Abbreviations are explained in the section "Material and methods".
ms.The note repetition rate varied from between 1.65 and 2.59 notes per second, mean 2.17 Kraus & allison, 2003dened tips, terminal grooves well developed, no webs.Except inner metacarpal tubercle, all other palmar and subarticular tubercles scar cely visible.Relative length of fingers 3 > 4 > 2 > 1, first finger almost as long as second, no webs.All toes with broadened tips, that of fourth toe a little wider than that of third finger.Subarticular tubercles and es-Callulops marmoratusKraus & allison, 2003has longer hind legs (TL/SVL 0.45 -0.49) and a mottled dorsal surface.
Diagnosis.With 48.2 mm SUL, a medium-sized species of the genus.Eye large (ED/SUL 0.135), hind leg medium-sized (TL/SUL 0.43).Finger tips and toe tips dilated to small discs, all with terminal grooves.Toe tips insignificantly wider than finger tips (T4D/ F3D 1.07).Dorsal and lateral surfaces light greybrown with a dense pattern of middle-and dark-brown smaller and larger flecks which are connected with one another in many cases.Conspicuous are two irregular pale patches above insertion of fore legs, and large lumbar "ocelli" of a similar colour.Abdomen uniformly light grey, chest and throat brownish with whitish spots.Body sides with many light pustules.

Table 2 .
Biometrical values of the holotype (ZMB 64107) and three paratypes of Callulops biakensis sp.nov.ZMB 64109 is a female, all others are males.All measurements in mm.Abbreviations are explained in the section "Material and methods".
Callulops doriae has a larger body and numerous striking blackish spots on dorsal surfaces vs. no spots in C. biakensis.Callulops fojaensis has a shorter hind leg (TL/SVL (TL/SVL 0.34) and longer call notes (mean 225 ms vs. 117 ms in C. biakensis).