Sphenophryne brevicrus

ZWEIFEL, RICHARD G., 2000, Partition Of The Australopapuan Microhylid Frog Genus Sphenophryne With Descriptions Of New Species, Bulletin of the American Museum of Natural History 2000 (253), pp. 1-130 : 84-92

Sphenophryne brevicrus : Loveridge, 1948: 422.

HOLOTYPE: AMNH A66047 , collected by Hobart M. Van Deusen on the Sixth Archbold Expedition, June 21, 1959, at Piunde- Aunde Lakes , Mt. Wilhelm, 3570 m, Simbu Province, Papua New Guinea.

PARATYPES (all from Papua New Guinea): Simbu Prov.: AMNH A66040 –66046 , A66048–66054 , A92799 –92801 (C&S), A92802, A135269, A135271–135310, A135313–135333, collected by the Sixth Archbold Expedition at the type locality, June 13–28, 1959 ; AMNH A78909 , collect- ed by L. K. Wade at the type locality, June 5, 1966 ; AMS R68908 – R68924 , collected on Mt. Wilhelm by J. Hope ; BPBM 5302 , collected by G. A. Samuelson, June 12, 1967, and 13430–13033, collected by R. C. A. Rice, Sept. 22, 1968, at the type locality ; MCZ A64308 , 64310–64336 , 64338 , collected by Fred Parker, June 10, 1967, and MCZ A111896–111900 , 111967–111977 , collected by Fred Parker at the type locality ; RMNH 16765 , 16766 , 16769 , 16671–16678 , collected by M.M.J. van Balgooy, May 5, and June 4, 1965, at the type locality ; BPBM 1049 , collected by J. L. Gressitt, June 3, 1955, at Denglagu, Mt. Wilhelm , 2500 m ; BPBM 2898 , 2899 , collected by J. Sedlacek, July 3, 1963, on Mt. Wilhelm , 3800 m ; BPBM 5353 , collected by J. L. Gressitt, Aug. 2, 1969, on Mt. Wilhelm , 4000 m ; MCZ A25920–25929 + 3 untagged specimens, collected by P. J. Darlington in 1944 on Mt. Wilhelm , 3050–3660 m ; MCZ A64296 , collected by Fred Parker, June 10, 1967, on Mt. Wilhelm , 2900 –3050 m ; RMNH 16763 , 16768 , collected by M.M.J. van Balgooy, June 15 and Apr. 21, 1965, at Kombugomambuno, 3300 m, Mt. Wilhelm ; RMNH 16762 , collected by M.M.J. van Balgooy, June 14, 1965, south of Lake Aunde, 3900 m, Mt. Wilhelm ; RMNH 16770 , collected by M.M.J. van Balgooy, Apr. 26, 1965, northeast of Lake Aunde , 3600 m ; MCZ A111932 , 111933 , collected by Fred Parker, Nov. 13, 1965, on the southwest slope of Mt. Kerigomna, 2900 m. Eastern Highlands Prov.: AMNH A76580 –76581 , A76587– 76589 , A137304 – A137307 (137306 C&S), MCZ A53083–53095 , 53097 , A111934 – 111966, and SAMA R6582 , R16629 , collected by Fred Parker, July 25, 1964, and Oct. 11, 1965, on Mt. Otto, 3540 m. Western Highlands Prov.: AMNH A65276–65280 , A65302 , collected on the Spalding-Peterson Expedition, Aug. 6, 1959, on Mt. Pollam , Wahgi Dividing Range , 2740 and 2490 m .

ETYMOLOGY: The specific name derives from the Greek stenos (narrow) and dactylos (digit), and is used as an adjective.

DIAGNOSIS: Oxydactyla stenodactyla differs from other Oxydactyla except alpestris in that the toe tips are narrow, not widened or flattened, and lack terminal grooves. The larger eye size of alpestris (mean EY/SVL> 0.33) will alone properly segregate most alpestris from stenodactyla (mean EY/SVL <0.30), and combining the greater leg length of alpestris along with eye size (fig. 46) affords nearly complete separation.

DESCRIPTION OF HOLOTYPE: Adult female (gravid) with the following measurements and proportions: SVL 30.4, HW 10.6, TL 9.2, EY 2.9, EN 1.6, IN 2.5, HD 6.2, FT 10.2, TY 1.6; HW/SVL 0.349, TL/SVL 0.303, EY/SVL 0.095, EN/SVL 0.053, IN/ SVL 0.082, EN/IN 0.640, HD/SVL 0.204, FT/SVL 0.336.

Head narrower than rotund, short-legged body. Snout rounded as seen from above and laterally, scarcely projecting; nostrils easily visible from above, about equidistant from eye and tip of snout; loreal region a flat, gentle slope, canthus rostralis rounded, barely distinguished. Eyes relatively small, corneal outline just visible from beneath if eyes fully protruded, eyelid about 75% of interorbital span. Tympanum small, well separated from eye, tympanic annulus barely visible. Relative lengths of fingers 3> 4> 2> 1, first more than half length of second; tips round- ed, not disclike, no terminal grooves; subarticular and metacarpal elevations scarcely evident. Toes not webbed, relative lengths 4> 3> 5> 2> 1, first less than half length of second, tips rounded, not disclike, no terminal grooves; subarticular and inner metatarsal elevations scarcely evident. A postocular skin fold curving downward behind tympanum, becoming indistinct at foreleg insertion; skin otherwise smooth dorsally and ventrally.

All dorsal surfaces are mottled in two shades of brown, with small areas of paler ground color occasionally showing through. The ventral surfaces are pale yellow-brown with a few indistinct marks on the lower lip and scattered dark spots on the abdomen. The posterior of the thigh is largely pale, with a few dark spots.

VARIATION IN TYPE SERIES: Body proportions of adult frogs in three samples are in table 10 and regression data for the two principal samples in table 11. The largest male stenodactyla measures 30.8 mm and the largest female 31.9 mm. Judged from the presence vocal slits, males mature at about 22– 23 mm. Females 25–26 mm and larger are mature; a large (SVL 30 mm) specimen from Mt. Otto contained 23 ova about 3 mm in diameter.

Variation in the color pattern of this species has individual, ontogenetic, and geographic components. Adult frogs from Mt. Wilhelm are quite variable, but generally show modifications of the theme described for the holotype —mottling of two shades of brown on a paler ground color. At the darkest extreme, only small, vermiform spots of the pale ground color may show, or at the other extreme large areas of the back may be occupied by pale ground color alone. Dark mottling may be fine or coarse. Some individuals have moderately well-developed lumbar ocelli and a pale postorbital-dorsolateral streak (fig. 44). The ventral surfaces range from immaculate to heavily spotted, with no obvious anterior-posterior differentiation (fig. 45). Small frogs tend to show a dark postorbital streak that broadens and extends along the flank, sharply defined from the paler and largely unmarked dorsum. Traces of the dark lateral area persist in some adults.

In contrast to frogs from Mt. Wilhelm, most from the summit of Mt. Otto are quite uniform in dorsal color pattern, being unmarked light brown (fig. 43). Juveniles, like those on Mt. Wilhelm, may have a dark postocular streak that expands to cover the flank, and this pattern persists faintly in some adult frogs. Dark or light markings in the flank region may hint at the contrasting patterns seen in Mt. Wilhelm frogs, but they rarely approach this degree of patterning even in the least developed pattern. Ventral coloration is variable: A rather uniform brown color with no macroscopic pattern is the mode, but this may be broken up by pale splotches of various sizes, or the venter may be pale with coarse, irregular dark marks.

The sample of one juvenile and five adult frogs from the Wahgi Dividing Range presents a third facies. The dorsal pattern in these resembles the juvenile pattern at the other localities. The dorsum from snout to cloaca is largely unmarked, pale tan, or has faint, indistinct darkening. Two specimens show an indistinct pale vertebral hairline, and all show dark marks in the lumbar region. A dark streak passes from the tip of the snout along the canthus rostralis and the edge of the eyelid, and from there toward the groin, usually becoming ill defined posteriorly. The facial region and the area below the streak posterior to the tympanum are more or less darkened, almost as dark as the streak in some instances and almost as pale as the back in others. The ventral pattern shows the sort of variation described for the Mt. Wilhelm frogs. Two adult specimens from Mt. Kerigomna resemble those from the Sepik-Wahgi Divide, but differ in having a pale inner edge to the canthal-dorsolateral streak.

The foregoing descriptions all refer to preserved specimens. Notes by M.M.J. van Balgooy on RMNH specimens from Mt. Wilhelm stated: ‘‘Orange with black markings darker above than below,’’ and ‘‘dark grey above with orange markings, below light orange with grey spots.’’ Color transparencies of a frog from Mt. Otto (fig. 43) show a deep gray dorsal color, with markings of a similar shade on a dull orange ventral background.

ILLUSTRATIONS: Ventral pattern, fig. 45; 3rd finger terminal phalanx, fig. 71A; tip of 3rd finger, fig. 49; premaxilla, fig. 63A; hyoid, fig. 69A; sacral region, fig. 72A; vomer, fig. 65A; skull, fig. 67A; hand and foot, fig. 53A; jaw musculature, figs. 74, 78A.

CALL: I have no recording of the call, which Loveridge (1948: 422, citing information from P. J. Darlington) described as ‘‘woodeny croaking.’’ M.M.J. van Balgooy (collector’s notes accompanying RMNH specimens) stated: ‘‘Its sound, a soft chirping ‘krrr, krrr’ not unlike that of a cricket, can be heard all day.’’ Hobart M. Van Deusen, who collected this species on Mt. Wilhelm on the Sixth Archbold Expedition, wrote in the field catalog: ‘‘Call note a single short ‘enh,’ rapidly repeated (40 or 50 in 30 seconds).’’

COMPARISONS WITH OTHER SPECIES: Only O. alpestris could be confused with this species (see Diagnosis).

HABITAT AND HABITS: The principal habitat of this species is alpine grassland, although the elevations at which some specimens were taken suggest that it may also occur in sub-alpine forests. Brass (1964) described in detail the cold, wet habitat at the type locality. Wade and McVean (1969: 31) stated that these frogs form ‘‘an extensive network of burrows in the tussock grassland.’’ Notes by R. Rice accompanying a BPBM specimen stated: ‘‘Very sluggish species. Sit in shallow holes in soil and duff of tussocks and clumps of vegetation.’’ In notes accompanying RMNH specimens, M.M.J. van Balgooy characterized stenodactyla as ‘‘a very common species in the alpine region where it hides away in hollows in the soil, under shrubs and grass-tussocks from 3300–4000 m.’’ P. J. Darlington found one, together with 14 eggs, in moss under a tussock (Loveridge, 1948). The foregoing notes all refer to Mt. Wilhelm.

DISTRIBUTION: This species is known from elevations of 2490 to 4000 m in the central highlands of Papua New Guinea in Western Highlands, Simbu and Eastern Highlands Provinces (fig. 41). Populations in the four major areas of distribution—the Sepik Wahgi Divide, Mt. Wilhelm, Mt. Otto, and Mt. Kerigomna—probably are disjunct from one another. See Holotype and Paratypes for localities and specimens examined.

REMARKS: It is evident from the description of variation that frogs from at least the three principal population samples (excluding Mt. Kerigomna, with only two specimens available) are quite distinctive in color pattern. A large majority could be assigned correctly geographically on the basis of color pattern alone. One could argue not unreasonably for separate specific (or at least subspecific) status for each population. I regard the morphological similarity (other than color pattern) of the three populations as compelling evidence for conspecific status, although I would welcome the opportunity to test this hypothesis against information from advertisement calls. Naming the known populations as subspecies would, in my opinion, serve no particularly useful service, because knowledge of the variation and distribution of the species most likely is far from complete.

P. J. Darlington, who climbed Mt. Wilhelm during World War II, was the first to collect this species. Loveridge (1948) referred the specimens to Sphenophryne brevicrus , a name that as used in the past included four species by my present reckoning.

Genus Sphenophryne Peters and Doria Cornufer Tschudi, 1838: 28 , 71, part; see Zweifel

(1967b) for disposition of this name.

Sphenophryne Peters and Doria, 1878: 430 . Type species (by monotypy),

Sphenophryne cornuta Peters and Doria, 1878:

430.

DIAGNOSIS: A genus of genyophrynine microhylid frogs (sensu Zweifel, 1971 and Burton, 1986) with the following combination of morphological characters: clavicles long and slender, reaching from scapula almost to midline of pectoral girdle; frontoparietal region of skull short and wide, mean ratio of braincase to skull length, 0.56; diapophyses of sacral vertebra broadly expand- ed, mean ratio of anterior-posterior distance to lateral span, 0.47 (table 14); first finger short, less than half length of second; tips of fingers (except 1st) and toes broadened into flattened discs with terminal grooves, disc on third finger broader than that on fourth toe; terminal phalanges with the tip broadly Tshaped, span of tip of phalanx of third finger averages 2X width of base of phalanx.

CONTENT: The genus is monotypic.

MORPHOLOGY: See species account.

DISTRIBUTION: See species account.

REMARKS: Superficially, Sphenophryne may be distinguished from other genyophrynine genera by the presence of a small dermal appendage on each eyelid. Proportions of the skull and sacral vertebra set it apart from the other genera with the primitive pectoral girdle, as do the short first finger and the relative sizes of finger and toe discs.

Sphenophryne cornuta Peters and Doria Figures 31A , 47

Cornufer unicolor Tschudi, 1838: 28 , 71, part (based on two specimens, only one of which is S. cornuta ; see Zweifel, 1967b).

Sphenophryne cornuta Peters and Doria, 1878: 430 (type locality, ‘‘presso il fiume Wa Samson nella Nuovo Guinea settentrionale’’ [‘‘near the Wa Samson River in northern New Guinea ’’; this river is on the northwestern edge of the Vogelkop Peninsula , Irian Jaya, Indonesia, with its mouth about 20 km east-northeast of Sorong ]; holotype, MSNG 29479 , collected by O. Beccari in 1875).

Chaperina ceratophthalmus van Kampen, 1909: 43 (type localities, ‘‘ Noord-Fluss bei Geitenkamp’ ’ [ Geitenkamp , Lorentz River ] and ‘‘ Resi-Gipfel’ ’ [Resi Peak], Irian Jaya, Indonesia; syntypes, ZMA 5777 [Geitenkamp], collected by H. A. Lorentz, July 2, 1907, and ZMA 5778 , 5779 [Resi Peak], collected by Lorentz on August 20 and September 9, 1907).

TYPE MATERIAL: The holotype is a small individual ( SVL 21.8 mm), presumably immature (not sexed), in ‘‘Discrete condizioni’’ (fairly good condition: Capocaccia, 1957: 220). In all pertinent characters it agrees with the following composite description.

DIAGNOSIS: The presence of a small but distinct, pointed tubercle on each eyelid distinguishes this species from all other Papuan microhylids. (The large-mouthed, squat Asterophrys turpicola has elongate warts on the eyelids, but not a single slender tubercle.) Equally distinctive are the combination of flat, vertical loreal region and flat upper surface of the snout, and the large finger discs, that of the third finger being broader than that of the fourth toe.

MORPHOLOGY: Males and females mature at about 29 mm SVL. Among 106 specimens measured, the largest male measures 37.4 mm and the largest individual, probably a female, 41.4 mm. The largest specimen whose sex was determined, a female, measured 40.0 mm. Head and body relatively slender, HW/ SVL about 0.35 in adults. Snout subacutely pointed as seen from above; nostrils lateral, much closer to tip of snout than to eye; loreal region vertical and flat, distinctly set off from flat upper surface of snout. Internarial distance relatively broad (mean IN/SVL, 0.12), eye–naris distance usually slightly less than eye diameter, rarely a little greater. Eyes moderate, laterally oriented, EY/SVL about 0.11 in adults, eyelid with a small but distinct pointed tubercle near its free margin. Tympanum distinct, horizontal diameter about 63% that of the eye, no sexual dimorphism. Hind legs relatively long, TL/SVL average about 0.49 in adults. Relative lengths of fingers 3> 4> 2> 1, first quite short; terminal disc of first finger scarcely if at all expanded, without a distinct groove, discs of other fingers well developed with terminal grooves, disc of third finger averages 2.3X width of penultimate phalanx (fig. 52). Relative lengths of toes 4> 3> 5> 2> 1, all with grooved terminal discs, that of fourth toe about 1.7X width of penultimate phalanx and about 82 % of width of third finger disc. Fingers and toes with low, rounded subarticular areas; no palmar or plantar elevations evident. Toes free or with sparse basal webbing (fig. 52). Body generally smooth above and below, but with rows of widely spaced tubercles on dorsal and ventral surfaces; similar tubercles on dorsal surfaces of hind limbs; usually a tubercle on heel and others along outer edge of tarsus.

COLOR AND PATTERN: Preserved specimens are brown or gray-brown above, virtually patternless or with rather indistinct markings. Variations include a darker central figure (fig. 47), a pale vertebral hairline, and faint lighter spots. Tubercles are typically light-tipped. The dorsal surface of the thigh usually has a thin, longitudinal light line, and light lines of the same width diagonally cross the anterodorsal surface of the thigh and the dorsal surfaces of the lower leg segments in many individuals. The posterior of the thigh is slightly darker than the dorsal surface with tiny light spots. The ventral surfaces are almost completely pale in some specimens, but in others there is heavy dark mottling, especially on the chin and chest.

I recorded the following colors in life for the specimen illustrated (fig. 31A, evidently it had paled slightly when photographed): a dark reddish brown middorsal band not sharply defined from lighter reddish brown ground color of head and rest of dorsal surfaces; anterior and posterior surfaces of thighs grayish brown with numerous minute yellowish white flecks; chin and chest bright- er reddish brown than dorsum, with gray-brown mottling; paired tubercles on chest and anterior part of abdomen tipped with white, a similar row of tiny white tubercles margined with black along lateral surface of body; iris grayish gold. A frog from Western Province had a golden line along the canthus rostralis reaching to the spike on the eyelid and a pale vertebral hairline in addition to the dorsal line on the thigh; chin and chest gray with splotches of orange; iris grayish gold with a red streak before and behind the pupil. Another specimen from the same region was largely gray beneath with a white midventral hairline and no red streak in the grayish gold iris.

Menzies (1976, pl. 12d) illustrated in color a specimen of much grayer aspect than those described, and (p. 61) mentioned one specimen whose ‘‘ventral side was bright red all over.’’ Brongersma and Venema (1962: 104) stated ‘‘it has a brown back and a rusty-red belly.’’

VARIATION IN SIZE AND PROPORTIONS: Park- er (1934: 154) gave the maximum length as 41 mm. The largest specimen I measured is 41.5 mm SVL (ZMA, uncataloged from Heuvelbivak, Lorentz River, Irian Jaya, probably female but not sexed). A female syntype of Chaperina ceratophthalmus (ZMA 5779) measures 40.0 mm. Males reach at least 37.4 mm (ZMA 5777, syntype of C. ceratophthalmus ; RMNH 16641, one of several with this number), but few males attain 36 mm whereas many females do. I measured and sexed 61 specimens (RMNH) from Mabilibol, Sibil Valley, Star Mountains, Irian Jaya. Males mature at about 28 mm and females at only a slightly larger size: The smallest males with vocal slits are that size; females of about 27–28 mm have apparently immature ova, whereas enlarging ova are present at 29 mm SVL.

Variation in proportions among adults in the large sample from the Sibil Valley region of West Irian is set forth in table 12, and regression statistics are presented in table 13. I detect no geographic trends in size or proportions in this widespread species. Measurements of individuals in scattered, smaller samples fall within the ranges set by the Sibil specimens.

ILLUSTRATIONS: 3rd finger terminal phalanx, fig. 71M; premaxilla, fig. 64D; sacral region, fig. 73; vomer, fig. 65P; skull, fig. 66; hand and foot, fig. 52; mandibular musculature, fig. 75B.

CALL: Menzies (1976: 61) described the call as ‘‘a long rattling chuckle lasting about one second and so loud that it can be heard from several hundred metres distance.’’ I heard this call near Tabubil but was unable to record it. However, a recording by Ian Redmond made available through the courtesy of Mr. Redmond and the British Library of Wildlife Sounds is available for analysis (fig. 76B). The calls (N = 4 from one individual, BMNH 1980.673) are 1.9–3.1 sec in length (mean 2.4) and consist of 42–67 notes (mean 52.2), each note comprised of 2 or 3 (rarely 4) discrete pulses. There is a dominant frequency at 3000 Hz and a slightly lesser peak at 1200 Hz. The note repetition rate is 21.6–23.0 notes per second (mean 22.1), with no obvious change in rate over the course of a call. Temperature was not noted.

In addition to the call described above, the species also has another that is possibly territorial in function. This call includes one to three brief (0.03–0.05 sec) pulsed notes with several emphasized harmonics. Single- and double-note calls are in the majority; triplenote calls are infrequent. Single or initial notes are more rapidly pulsed than second or third notes and include a more nearly complete set of harmonics (fig. 76A). In one recorded instance, a frog called 18 times in the space of 3 min 49 sec (average, once every 12.7 sec). Many of these calls (10 of 19) appeared to elicit responses from, or were in response to, similar calls from an individual nearby.

COMPARISONS WITH OTHER SPECIES: Sphenophryne cornuta is that rare creature, a Papuan microhylid that readily can be identified to genus and species without resorting to dissection or call analysis. No other species shares its combination of a small dermal appendage on the eyelid, fingers with enlarged discs larger than those on the toes, and vertical loreal region. Scansorial species of Cophixalus and Oreophryne may be similar in size and general morphology, but none has the eyelid appendage.

HABITAT AND HABITS: My field experience with this species is limited to two areas—the Adelbert Mountains in coastal Madang Province, and the vicinity of Tabubil, Western Province. At an elevation of 670 m in the former region, I found individuals at night at heights of less than 2 m on vegetation beside a stream in rainforest regrowth. They shared this habit and habitat with two other microhylids of similar size, Cophixalus biroi (Méhelÿ) and C. cheesmanae Parker. These two were calling, whereas I heard no calls from cornuta . Near Tabubil, frogs gave possibly territorial calls (see above) from perches 1–2 m up in shrubs, saplings, or tangled vegetation not close to any running water.

Menzies (1976: 60 –61) remarked that ‘‘calling males seem to be well spaced out for I have never found, or heard, two close together,’’ and added that cornuta ‘‘is probably as much terrestrial as scansorial although males, at least, do climb about on low vegetation.’’ I find nothing else in the literature on the ecology of this species.

DISTRIBUTION: On the north coast of New Guinea, Sphenophryne cornuta has been found at low to moderate elevations from near the tip of the Vogelkop Peninsula eastward to the Adelbert Mountains north of Madang. Records on the south coast extend from the Bomberi Peninsula in Irian Jaya eastward to slightly east of Port Moresby (fig. 48). Spotty but fairly intensive collecting on the Huon Peninsula and eastward on the north coast has produced no cornuta . Elevations of collection range from virtually sea level, as at Kerema and Katau, to 2600 m in the Hellwig Mountains of Irian Jaya, though most localities lie between about 400 and 1250 m. There are no insular records.

LOCALITY RECORDS AND SPECIMENS EXAMINED: IRIAN JAYA: Bomberi ( BPBM 5290 ) ; Camp VI, Utakwa River ( BMNH 1913 . 11.1.139) ; foot of Charles Lewis Mtns. ( BMNH 1897.3.23.5) ; Geitenkamp, Lorentz River ( ZMA 5777 , syntype of Chaperina ceratophthalmus ) ; Hellwig Mtns., 2600 m ( ZMA, unnumbered) ; Heuvelbivak, 800 m, Lorentz River ( ZMA, unnumbered) ; Mt. Kohari , between Modder-lust and Kasawari ( MCZ A7611 ) ; Launch Camp, Setekwa Riv- er ( BMNH 1913.11 .1.136–138) ; near the Wa Samson River , about 20 km ENE Sorong ( MSNG 29479 , holotype) ; Peramelesbivak, Lorentz River ( ZMA unnumbered) ; Resi Peak, Lorentz River ( ZMA 5778 , 5779 , syntypes of Chaperina ceratophthalmus ) ; Sabang, Lorentz River ( ZMA, unnumbered) ;

(triangle) on the Aru Islands.

Sibil Valley, 1250 m (AMNH A84503; BPBM 1045–1047, 3167, 3171, 3173, 3175); Mabilibol (RMNH 16607, 16609 [4], 16613 [2], 16636 [2], 16637 [2], 16638 [20], 16641 [8], 16642, 16644 [12], 16647 [2], 16654 [2], 16659; Sibil Valley; Sibil (RMNH 16663 [3]). PAPUA NEW GUINEA: Central Prov.: Vikaiku, Angabunga (St. Joseph) River (MSNG 29929); Bellavista, via Fane, 1460 m (PNGM, James Menzies, personal commun.; not examined by author); Musgrave River, 18 km E Sogeri, 460 m (MCZ A88020). Gulf Prov.: Gihiteri Village, Omati River (Schultz-Westrum 350 [ZSM]); Kerema (UPNG 3666–3668, 3670, 3671); Weiana, 8 km S, 1 km E Soliabedo, 460 m (MCZ A64105, 64106); Camp III, Nimi River, 13.5 km S, 1 km E Soliabedo, 430 m (MCZ A64111–64114); Uraru, Purari River 90 m (MCZ A64108, 64115). Madang Prov.: Wanuma, Adelbert Mtns., 670 m (AMNH A83048–83052); Wanuma, 1000 m (BPBM 1203). Simbu Prov: Bomai area (AMNH A76586), Bomai, Tive Plateau, 1070 m (MCZ A59696–59701); Karimui, 1100 m (AMNH A77101; MCZ A64104, 64109, 68338–63341); Camp II, Pio River, 6.5 km S, 1 km E Soliabedo, 300 m (MCZ A64110). Western Prov.: Derongo, 400 m (AMNH A82286; MCZ A80006, 81231–81234); Megalsimbip (UPNG 6731–6733); Imigabip, 1280 m (MCZ A80995–80997); Menemsorae (MCZ A80005); Bolangon, 1280 m (MCZ A80998); Yongtau No. 1, 240 m (MCZ A80993, 80994); Haidauwogam, 240 m, 27 km S, 5 km E Tabubil (AMNH A130542–130544); Finalbin, 840 m, 5 km N, 2 km W Tabubil (AMNH A130545, A130546, A130548, A130549, UPNG 8275); Katau (MSNG 29480); junction of Strickland and Rentoul rivers (BMNH 1980.673); junction of Strickland and Tomu rivers (BMNH 1980.674); Kiunga (MCZ A80003, 80004; USNM 203856); Emeti, Bamu River (MCZ A87269, 87270). West Sepik Prov.: Upper course of the Reinjamur, Torricelli Mtns., 650–700 m (south of Paup; Wandolleck, 1911: 4); Mt. Somoro, Torricelli Mtns., 730 –1420 m (AMNH A78145– 78149); Mt. Nibo, Torricelli Mtns., 700– 1550 m (AMNH A78150–78155, A92803– 92804 (C&S), A135334–135369 (the last C&S), A135334 –135339; Lumi, 530 m (AMNH A78156, A78157).

Figures

Fig. 31. Examples of Austrochaperina, Liophryne, and Sphenophryne. A. S. cornuta, AMNH A130544, adult. B. A. brevipes, AMNH A130534, SVL 21.9 mm. C. A. guttata, QM J67256, SVL 43.2 mm. D. A. macrorhyncha, MZB 3564, SVL 36.8 mm. E. A. rivularis, holotype, AMNH A130551, SVL 49.0 mm. F. A. palmipes, AMNH A130550, adult. G. L. similis, AMNH A130564, SVL 40.0 mm. H. A. derongo, MZB 3563, SVL 36.7 mm. Photos C, D, and H by S. Richards.

Fig. 47. Sphenophryne cornuta, AMNH A83051, SVL 38.5 mm.