AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2723, pp. 1-14, figs. 1-10 February 26, 1982 A New Montane Microhylid from Papua New Guinea, and Comments on the Status of the Genus

RICHARD G. ZWEIFEL1 AND ALLEN ALLISON2

ABSTRACT Discovery of an undescribed, ground-dwelling these are ones common to many microhylid species of the largely scansorial genus species of similar ground-dwelling habits and do led to the re-evaluation of the monotypic genus not necessarily indicate closeness of relationship. Aphantophryne to see if this genus, presently Evaluation of previously unstudied features of the synonymized within Cophixalus, should be re- anatomy of C. pansus does not support revival of vived to accommodate its terrestrial type species, Aphantophryne either as a monotypic genus or to C. pansus, and the new form. The two species include the new species. The latter is described share some presumably derived features, but as Cophixalus sphagnicola, new species.

INTRODUCTION In 1969 Zweifel collected four individuals tion of Allison's ecological studies by pro- of a small, ground-dwelling and apparently viding a name for the undescribed species; unnamed microhylid atop Mount Kaindi, and to investigate whether there is sufficient Morobe Province, Papua New Guinea. The justification for reviving the genus Aphan- habitus of these was sufficiently like tophryne Fry " 1916" [1917] to accommodate that of montane that he set both its type species (presently assigned to them aside for later consideration in the Cophixalus) and the new frog described course of a proposed revision of that genus. herein. Later, Allison conducted an extensive eco- logical study of this species. He submitted METHODS specimens to Mr. James Menzies, who de- Standard measurements were made and termined that the species has the skeletal utilized in generating ratios used to compare characteristics of the genus Cophixalus rath- proportions: body length, from snout to vent er than of Sphenophryne. The present paper (SV); length of tibia, from heel to outer side has two purposes: to facilitate the comple- of flexed knee (TL); distance from anterior

I Curator, Department of Herpetology, American Museum of Natural History. 2 Assistant Director, Wau Ecology Institute, Wau, Papua New Guinea.

Copyright © American Museum of Natural History 1982 ISSN 0003-0082. / Price $2.28 2 AMERICAN MUSEUM NOVITATES NO. 2723 edge (corner) of eye opening to center of ex- (1934). Zweifel recognized that this squat, ternal naris (EN); internarial distance, be- short-legged terrestrial frog with rounded toe tween centers of external nares (IN); head tips differed from typical Cophixalus, which width, at jaw articulation (HW); diameter of are scansorial frogs with expanded digital orbit, measured between anterior and pos- tips, but he preferred not to revive a mono- terior edges of eye opening (Eye). Measure- typic genus on these grounds alone, so he ments, expressed in millimeters, were made referred pansa to the genus Cophixalus and, with vernier calipers or ocular micrometer. hence, Aphantophryne to the synonymy of Statistics are given in the format: mean + Cophixalus, where it has remained. The dis- standard error of mean (range). covery of a new terrestrial, montane species somewhat resembling C. pansus calls for ACKNOWLEDGMENTS comparison between the two and with other This research utilized specimens deposited Cophixalus to determine if revival of Aphan- in the American Museum of Natural History, tophryne may be warranted. New York (AMNH), the Bernice P. Bishop Cophixalus was one of five genera of Museum, Honolulu (BPBM), and the Aus- sphenophrynine microhylids recognized by tralian Museum, Sydney (AM). We are Parker (1934), who diagnosed them on the grateful to Dr. Alan A. Ziegler (BPBM) and basis of characteristics of the pectoral girdle. Dr. Harold Cogger (AM) for their coopera- The five formed a graded series distinguished tion. Zweifel's participation in the R/V Alpha by the reduction and loss of ventral elements Helix Expedition in 1969, supported by the of the girdle: Sphenophryne (girdle com- National Science Foundation, contributed plete); (clavicle reduced); Mi- importantly to the study. Part of the research crobatrachus (clavicle supposedly lost); Co- was carried out while Zweifel was a Visiting phixalus (clavicle and procoracoid lost); Curator in the Australian Museum. Aphantophryne (clavicle, procoracoid and, supposedly, sternal plate lost). Several STATUS OF Aphantophryne changes in the generic level have Fry ("1916" [1917]) described the genus taken place since Parker's revision. Zweifel and species Aphantophryne pansa on the (1956) referred Aphantophryne to the syn- basis of six specimens taken at an elevation onymy of Cophixalus, and later (1971) re- of 3719 m. (12,200 ft.) in the Owen Stanley defined the Sphenophryninae, transferring Mountains of British New Guinea (now Pa- the genus Genyophryne to it. Tyler (1978) pua New Guinea). The principal feature disposed of Microbatrachus as probably thought to be diagnostic of the new genus based on a juvenile Sphenophryne. Menzies was the lack of a sternal plate. Parker, in his and Tyler (1977) removed the genera Co- revision of the (1934), recog- piula and Choerophryne from the synonymy nized Aphantophryne as a monotypic genus, of Cophixalus. relying on Fry's description in the absence of specimens to examine. Loveridge (1948) FEATURES OF Cophixalus pansus referred Aphantophryne to the synonymy of PERTINENT TO GENERIC STATUS Asterophrys after examining (but not dis- GENERAL MORPHOLOGY: The features of secting) one of Fry's paratypes, and without morphology that impart to C. pansus an ap- reference to the nature of the pectoral girdle. pearance rather different from other Cophix- With relatively fresh material available, alus are ones common to ground-dwelling Zweifel (1956) determined that the type microhylids of several genera: short hind species does possess a sternal plate and oth- legs, narrow toe tips, broad head, small eyes. erwise has the skeletal characteristics of the Leg length of pansus is the shortest relative genus Cophixalus3 as diagnosed by Parker to body length of any Cophixalus. No other 3See Menzies, Tyler, and Zweifel (1980) for the use Cophixalus has digits with no trace of disc of this generic name. development and no trace of terminal 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 3

A B FIG. 1. Left hands of Cophixalus pansus (A) and C. sphagnicola (B) in palmar aspect. Scale lines marked in mm. grooves (fig. 1A). The terminal phalanges bear what we assume to be vestiges of the T- shaped extensions of scansorial species (fig. 2D). Relative head width and relative eye size are very nearly at the extremes for the genus in pansus. VOMERINE BONES: The shape of the vo- mers may be diagnostic of microhylid genera (Zweifel, 1972; Menzies and Tyler, 1977), but that of pansus does not differ signifi- A B cantly from that of more typical Cophixalus (fig. 3D). SACRAL VERTEBRA: The sacral vertebra of pansus has the diapophyses relatively little expanded. In this respect it differs slightly if at all from scansorial species of Cophixalus (fig. 4). PECTORAL GIRDLE: The sternal plate of pansus is weakly developed compared with A a other Cophixalus. Specimens of other C D species cleared and stained with Alizarin Red often show calcified tissue in the basal of the but no calcification is FIG. 2. Terminal phalanges of third fingers to part sternum, show relative development of T-shaped process- present in a specimen of pansus. In other es. A. Cophixalus verrucosus, a scansorial spe- respects the girdle is like other Cophixalus. cies. B. sp., a genus of ground-dwelling Fry's ("1916" [1917]) impression of the dis- species, formerly referred to Cophixalus. C. Co- tinctness of the pectoral musculature of pan- phixalus sphagnicola. D. C. pansus. Scale lines sus was accurate within the limited context marked in 0.25 mm. increments. 4 AMERICAN MUSEUM NOVITATES NO. 2723

A

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C D

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B FIG. 3. Vomers of right side in ventral aspect. A. Cophixalus parkeri, a scansorial species. B. Copiula sp. C. Cophixalus sphagnicola. D. C. pansus. Stippled areas represent cartilage asso- ciated with vomers and sphenethmoids. Scale lines marked in 0.5 mm. increments. of material available to him, but he had no other Cophixalus for comparison. C THROAT MUSCULATURE: Variation in throat musculature is assuming significance FIG. 4. Sacral vertebrae in dorsal aspect. A. in assessing relationships among frogs (Ty- Cophixalus riparius, a scansorial species. B. C. ler, 1971, 1972, 1979; Emerson, 1976). The sphagnicola. C. C. pansus. Scale lines marked in superficial mandibular musculature of C. mm. pansus (fig. 5) is basically similar to that of other microhylids (Tyler, 1979, fig. 4:8; sus than in several other Cophixalus dis- Emerson, 1976, fig. 5): well-developed inter- sected. In these latter species the muscle hyoideus, intermandibularis (with a single may be represented only by fibers distally supplementary slip) and submentalis (inter- along the mandible, hidden by the interman- mandibularis anterior) muscles. dibularis. If the intermandibularis is loos- In pansus there is a fairly sturdy muscle ened medially and reflected, the muscle in (imand d., in fig. 5) lying dorsal to the inter- question reflects with it and may not be rec- mandibularis, between the main body of the ognized. intermandibularis and the lateral geniohyoi- What presumably is the homologous mus- deus. It originates on the mandible and in- cle is well developed in Copiula, reaching serts along with the intermandibularis in the essentially to the median raphe, but is com- region of the median raphe; part of the mus- pletely hidden by the intermandibularis. cle is visible without reflecting the interman- Emerson (1976, fig. 4) illustrates and notes dibularis or its accessory slip. Investigation an "unusual condition" of the intermandib- of other Cophixalus and the related genus ularis in Kalophrynus pleurostigma, a micro- Copiula shows that this muscle is not unique hylid thought not to be closely related to any to pansus, though better developed in pan- Papuan species. Here, the intermandibularis 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 5

(intermandibularis posterior of Emerson) is divided into smaller anterior and larger pos- - wsubmen. terior parts, with a space between. Such a condition could be a precursor to the division genhy.I1. seen in Cophixalus and Copiula. In any imand. d. event, the close anatomical relationship of genhy.m. the intermandibularis (in the strict sense) and the muscle in question suggests that pending suppl. more comprehensive and detailed study the latter be referred to as a dorsal element of ----~ imand. the intermandibularis. FEATURES OF THE NEW SPECIES PERTINENT TO GENERIC STATUS interhy. GENERAL MORPHOLOGY: Because of its relatively short hind legs and lack of expand- ed digital discs, this species more closely re- sembles pansus than it does the more typical scansorial species of Cophixalus. Head width, however, is close to average for the FIG. 5. Superficial throat musculature of Go- genus, and the eyes are relatively large. Ter- phixalus pansus, right side only. Abbreviations: minal processes of the phalanges are only genhy 1., geniohyoideus lateralis; genhy in., ge- slightly broader than those of pansus (fig. niohyoideus medialis; imand., Intermandibularis; 2C), and although the tips of the digits are imand. d., dorsal element of intermandibularis; not expanded, their shape is disclike, and submen., submentalis; suppl., supplementary slip weak terminal grooves are present (fig. iB). of intermandibularis. Scale line marked in mm. VOMERINE BONES: The shape of the vo- mer is much like that of pansus and typical Cophixalus (fig. 3). an arboreal or at least climbing way of life: SACRAL VERTEBRA: The expansion of the complete absence of toe discs and terminal diapophyses is similar to that of pansus and grooves; relatively small eyes; short legs. of scansorial Cophixalus-neither greatly The relatively broad head is also seen in expanded nor narrowed (fig. 4). many terrestrial Papuan microhylids, though PECTORAL GIRDLE: In a double-stained it is not so clearly an adaptation to terrestrial the sternal shows no cal- life. specimen, region Aphantophryne can be diagnosed on the cification and little sign of cartilage. Hence, basis of the unique derived conditions of there is a resemblance to the condition in rounded digital tips without discs or terminal pansus. grooves; the head, eye, and limb proportions THROAT MUSCULATURE: We cannot iden- are supporting but less distinctive character- tify a dorsal element of the intermandibularis istics. The distinction is blurred, however, in the new species, but since it can be diffi- when the new species is considered. Its dig- cult to see even in larger species of Cophix- ital tips are not at all expanded, but are alus, the apparent absence in this small form somewhat flattened and disclike, and bear should not be attributed significance. weak terminal grooves. The condition is in- termediate between typical expanded discs DISCUSSION of scansorial species and the state in pansus. From the foregoing, it is evident that Co- An earlier decision that Aphantophryne phixalus pansus differs from more typical did not warrant recognition as a monotypic scansorial Cophixalus chiefly in ways asso- genus (Zweifel, 1956) considered the foot ciated with a ground-dwelling as opposed to structure, pectoral girdle and general habitus 6 AMERICAN MUSEUM NOVITATES NO. 2723

FIG. 6. Cophixalus in dorsal aspect. A. C. pansus, AMNH 58123, x2.2. B. C. sphagnicola ho- lotype, AMNH 83077, x2.9.

of pansus. The examination of additional in- ternal characteristics in the present investi- Cophixalus sphagnicola, new species gation reveals nothing there unique to the Figure 6B species, and the discovery of a new species HOLOTYPE: AMNH 83077 (field no. RZ that is in some ways intermediate between 9162), adult female, collected by R. G. Zwei- pansus and more typical Cophixalus further fel at about 2300 m., near the summit of diminishes its distinctiveness. Cophixalus Mount Kaindi, 5 km. west of Wau, Morobe pansus represents the extreme of adaptation Province, Papua New Guinea, on September to terrestrial existence seen in the Cophix- 7, 1969. alus evolutionary line, but to segregate it in PARATYPES: AMNH 83076, 83078, 83079, a monotypic genus would serve little pur- AMR 31242, same locality as holotype; BPBM pose. To group it with the new species in a 5636, 6354, 6357, 6362, Bulldog Road, 2400 genus apart from Cophixalus would be in- m., 8 km. SSW Wau; BPBM 6347, 6348, advisable. Not only would such a genus be Bulldog Road, 2775-2800 m., 13 km. SSE much less effectively diagnosed than would Wau; BPBM 6293, 6294, 6296-6299, Mt. the monotypic genus, but such an associa- Kumbak, 2800 m., Bulldog Road, 15 km. tion would imply a closeness of relationship SSW Wau; BPBM 8311, Bulldog Road, that present information cannot support. The 2500-2700 m.; BPBM 6250, 6251, 6262, Mt. presumed derived characters that link the Amungwiwa, 3000 m., 20 km. SW Wau; two species-leg length and toe structure- BPBM 8308-8310, Mt. Missim, 2400 m., 10 are ones notorious for parallel development km. NE Wau. All localities are in Morobe in Papuan microhylids. Similar morphology Province, Papua New Guinea. is seen in several montane, ground-dwelling DEFINITION AND DIAGNOSIS: Cophixalus species of other genera. sphagnicola is a small species, males reach- 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 7 ing about 18.5 mm. and females about 20.6 6.5; Eye, 2.3; EN, 1.1; IN, 1.7; eyelid, 1.4; mm. in length from snout to vent. The hind interorbital span, 2.0. legs are short, first finger less than one-half VARIATION IN THE TYPE SERIES: Females the length of the second, and tips of digits range up to 20.6 mm. snout to vent length unexpanded but disclike and with terminal and are gravid at as small as 16.3 mm. The grooves. The only other species of Cophix- largest male measures 18.5 mm., and one of alus that lacks expanded terminal discs on 15.8 mm. has vocal slits so presumably is both fingers and toes is C. pansus. Larger mature. Selected proportions vary as follows size (females to 28 mm., males to 22 mm. (N = 17): TL/SV, 0.384 + 0.005 (0.35-0.42); SV), longer first finger and absence of ter- HW/SV, 0.376 + 0.005 (0.35-0.42); IN/SV, minal grooves on digits distinguish C. pan- 0.106 + 0.001 (0.097-0.113); EN/IN, 0.630 + sus from C. sphagnicola. 0.009 (0.55-0.71); EN/SV, 0.066 + 0.001 DESCRIPTION OF HOLOTYPE: Body slightly (0.059-0.073); Eye/SV, 0.131 + 0.002 (0.119- wider than moderately broad head (HW/SV, 0.148). 0.398); snout shorter than eye, viewed from Color patterns are variable, although the above appears bluntly pointed, nearly trun- dark face mask effect is present to a degree cate; loreal region flat and almost vertical; in all specimens. Most have an elongate dark nares broadly spaced (IN/SV, 0.104; EN/IN, postorbital-supratympanic mark that is 0.647); eyes relatively large (Eye/SV, 0.141), scarcely seen in the holotype. The upper sur- interorbital space about 1.4 times width of face of the snout may be dark, as in the ho- upper eyelid; tympanum indistinct, located lotype, or may be pale, sharply set off along about one-half its diameter behind posterior a transverse midorbital line. A more or less corner of eye. Fingers unwebbed, relative well-developed dorsolateral dark streak lengths 3>4>2>1, first finger less than one- often is present, and the middorsal area thus half length of second; tips of fingers 2-4 disc- defined may be distinctly paler than the lat- like with terminal grooves, but not broader eral area below the marking. One specimen than penultimate phalanges; first finger blunt- has a narrow light midvertebral line irregu- ly pointed with only a faint terminal groove; larly edged with darker pigment, and another no distinct subdigital or palmar tubercles- has a dark X-shaped mark in the scapular only low, rounded elevations. Toes un- region. Ventral surfaces range from dark webbed, relative lengths 4>3>5>2>1, first brown with tiny light flecks to pale with a toe less than one-half length of second, tips broken network of melanophores. The av- of toes disclike but not broader than penul- erage condition of pale spots enclosed within timate phalanges; a low, rounded inner meta- a weak darker reticulum is seen in the ho- tarsal tubercle, other subdigital and plantar lotype. In life, the dorsal ground color is elevations scarcely evident. Hind legs rela- grayish brown to reddish brown, and the tively short (TL/SV, 0.380). Head, body and ventral surfaces are gray with lighter gray front limbs smooth dorsally and ventrally; a flecks. few small warts present on sides ofbody pos- The skeleton was examined in one cleared teriorly and on upper surfaces of hind limbs. and stained specimen, a female 15 mm. Teeth lacking. A median palatal tubercle fol- snout-vent length. The coracoids are the lowed by a low denticulate ridge. only ossified ventral elements of the pectoral Eyelids, snout, and interoribtal area dark girdle. There are no procoracoids or clavicles, brown; side of head-snout to tympanum- and the sternum is scarcely discernible. Some dark brown; body and limbs lighter brown skeletal elements are illustrated in figures 2, 3, dorsally with indistinct small, darker mark- and 4. ings; a dark brown triangular patch below CALL: The call is a train of high-pitched cloaca. All ventral surfaces, including limbs, peeps. Individual notes uttered by two frogs with small pale spots enclosed in a brown (fig. 7) averaged 0.32 and 0.26 sec. in length, reticulum. with little variation. Midpoint frequencies of MEASUREMENTS: SV, 16.3; TL, 6.2; HW, the longer calls ranged between 3250 and 8 AMERICAN MUSEUM N;OVITATES NO. 2723

4.

O N I .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 2.2

6-

4-

2-

0 9 9 9 p 9 a ff . .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 TIME IN SECONDS

FIG. 7. Audiospectrograms of calls of Cophixalus sphagnicola, 45 Hz. filter. Recorded September 12, 1980, by A. Allison on Bulldog Road, Morobe Province, Papua New Guinea. Lower figure shows part of call of A. Allison no. 7194, calling 8 cm. within a mossy bank at a temperature of 12.3°C. Upper figure shows part of call of a second individual (not captured) calling nearby and recorded at the same time as the first frog. Differences seen in frequency, note repetition rate and note length could result from either or both of temperature and body size differences, though a substantial temperature difference under these circumstances is unlikely.

3450 Hz., whereas those of the shorter calls afternoon, especially if there is rain. On very were between 3750 and 3950 Hz. The num- rainy days calling may continue at a low level ber of notes in a train varied from 15 to 27 throughout the day. This species rarely calls in a sample of 150 calls counted on the sum- at night. On Bulldog Road and adjacent areas mit of Mt. Kaindi and on Bulldog Road (2850 of the Ekuti Dividing Range the pattern is m.), approximately 10 km. from Mt. Kaindi. similar except that calling activity, after During peak activity, individuals may call peaking at first light, generally continues every 30 to 60 sec. throughout the day except during infrequent Calling activity on Mt. Kaindi begins sunny periods. Substratum temperatures shortly before dawn, peaks at first light and while the frogs are calling range from 11 to then drops off, typically ceasing by midday 150C. (fig. 8). Some calling generally resumes in the ECOLOGICAL NOTES: The holotype and 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 9

three AMNH paratypes were found beneath 50- chunks of wood and sticks lying beside a trail through heavily mossed forest on Mt. Kain- di, where this species commonly occurs in surface litter beneath clumps of Gleichenia 40- ferns or Rubus vines along the road to the (n microwave repeater station. See Gressitt and -J Nadkarni (1978) for a general description of 4 30- u the area and its vegetation. 0 Cophixalus a: sphagnicola reaches its greatest abundance- w m more than five individuals per square me- 2 20- ter-in second-growth habitat at the edge of z heavily mossed forest. On Bulldog Road, built and abandoned during World War II and now heavily overgrown, it occurs most 10- frequently in clumps of Sphagnum and other mosses along road banks or under mats of Vaccinium amblyandrum covering the for- 06:00 oe:oo l0:00 12:00 14:00 mer road surface. At high elevations on the TIME OF DAY Ekuti Divide (e.g., Mt. Amungwiwa) it oc- FIG. 8. Number of calls of Cophixalus sphag- curs in alpine grassland, inhabiting Des- nicola tabulated during 15 minute periods along champsia-Danthonia grass tussocks. It is no- a 10 m. section of road on the summit of Mt. where common in the forest interior and is Kaindi, 2362 m., Morobe Province, Papua New virtually absent from deeply shaded mossy Guinea on February 9, 1981. About 15 male frogs forest. were resident in the study area. In the lower part of its elevational range (2300-2600 m.) C. sphagnicola is sympatric with Barygenysflavigularis Zweifel, a slight- 2700 m.). Cophixalus sphagnicola and C. ly larger diurnal terrestrial microhylid (Zwei- pansus are similar ecologically. They occur fel, 1972) that inhabits the interior of the for- in much the same habitat and, judged by est but also occurs in small numbers at the their calling activity, both are active mainly forest edge. At least five species of Cophix- in the morning and less so during the rest of alus, all nocturnally active, occur sympatri- the day. They are the only diurnal terrestrial cally with sphagnicola on the summit of Mt. frogs on the study site. The only other ter- Kaindi: C. cryptotympanum Zweifel, C. restrial species at this elevation is Spheno- kaindiensis Zweifel, C. parkeri Loveridge, phryne rhododactyla (Boulenger), a large C. riparius Zweifel and C. variegatus (van microhylid that roams the forest floor at Kampen). The last four are scansorial night. species; three of them commonly call from Cophixalus sphagnicola breeds through- shrubs or low trees, whereas riparius may out the year on Bulldog Road. Up to 20 eggs be found on vertical road banks as well as per clutch are laid in a shallow depression high in the taller trees. Cophixalus crypto- beneath the surface litter and are attended tympanum may call from bushes, but is most by the male until they hatch in about three often found on the ground beside seeps or months. Males occasionally call while with along roadside ditches. Its main activity is the eggs. As is presumably the case with all nocturnal though it may call during late after- New Guinea microhylids, development is noon. direct; there is no aquatic tadpole stage. At the Bulldog Road study site, about 2850 COMPARISONS WITH OTHER SPECIES: Be- m., C. sphagnicola is sympatric with C. cause of its lack of expanded digital discs, pansus, C. riparius, C. variegatus and pos- Cophixalus sphagnicola is more likely to be sibly C. parkeri (known from a maximum of confused with species of other genera than 10 AMERICAN MUSEUM NOVITATES NO. 2723 with other Cophixalus. With of such descriptive of the frog's habitat on the mossy small size, one must clear and stain repre- forest floor. sentative specimens to be sure of the perti- DISTRIBUTION: Cophixalus sphagnicola is nent skeletal characteristics. The absence of known only from mountains near Wau, Mo- clavicles and of procoracoids coupled with robe Province, Papua New Guinea, at ele- the eleutherognathine condition of the upper vations from about 2300 to 3000 m. (7500- jaw (premaxillae not meeting on the midline), 9800 ft.). Evidently the species is distributed will distinguish sphagnicola from small in- among disjunct populations (fig. 10). Local- dividuals of Sphenophryne and Phrynoman- ities and specimens examined are listed un- tis with similar toe structure. Barygenys has der Paratypes, above. vertical ridges on the snout. Although the new species has been com- Cophixalus pansus (Fry) pared with C. pansus in determining generic Figure 6A status, it does not necessarily follow that the Aphantophryne pansa Fry, " 1916" (1917), p. 772 two species are each other's closest rela- (type locality, "Mount Scratchley, on the Owen tives. As has been indicated, their common Stanley Range, British New Guinea . .. 12,200 and presumably derived characteristics may ft. [3719 mr.]"4; holotype, AM R5238, collected represent parallel adaptations to terrestrial by A. Giulianetti in September or October existence. 1896).5 Cophixalus sphagnicola shares some char- Asterophrys pansa pansa: Loveridge, 1948, p. acteristics with a sibling pair of scansorial 417. species, C. shellyi Zweifel and C. pipilans Cophixalus pansus: Zweifel, 1956, p. 41. Zweifel (Zweifel, 1980). These two are small DIAGNOSIS: Cophixalus pansus differs frogs, maximum length about 20-22 mm. from all microhylids of the New Guinea re- snout to vent, with the first finger greatly re- gion in the following combination of char- duced and with a dark face mask. Fingers, acters: upper jaw eleutherognathine, clav- except the first, bear terminal discs, but icles and procoracoids lacking, tips of fingers these are relatively small and are smaller and toes rounded without terminal grooves than those on the toes; the reverse is true in and with no trace of discs. typical scansorial Cophixalus. The internar- In general appearance, C. pansus resem- ial span in shellyi is relatively the greatest in bles a ground-dwelling species of Phryno- the genus, with sphagnicola and pipilans mantis or Sphenophryne more than it does being closest to it. The call in all three its scansorial congeners. The eleutherogna- species is a series of high-pitched peeps. thine upper jaw (vs. symphygnathine) distin- Reduction of the first finger must be con- guishes pansus from any Phrynomantis, and sidered a derived character, and small size, the presence of clavicles differentiates reduced terminal digital expansions, and Sphenophryne. broad internarial span may also be derived. DESCRIPTION AND VARIATION: Cophixal- No case can be made for regarding the pig- us pansus is a rather squat, relatively broad- mentation or call either primitive or derived. headed frog with short hind legs (the shortest There is no way of determining whether the relative to body length in its genus) and small shared derived characters of the three eyes. Females reach a maximum length from species reflect common ancestry, but pend- snout to vent of about 28 mm., males 22 mm. ing additional evidence, it is reasonable to consider C. sphagnicola to be a montane ter- restrial derivative of the shellyi-pipilans line I Mt. Scratchley lies on the border between the Cen- tral and Northern provinces of Papua New Guinea; it is of Cophixalus. uncertain in which province the holotype was collected. ETYMOLOGY: The specific epithet derives 5Fry chose and illustrated a holotype from among the from the Latinized Greek sphagnus, moss, six specimens available to him but did not specify a combined with the Latin suffix -cola (com- catalog number. However, AM R5238 is segregated and mon gender), inhabiting, to form an adjective bears a label reading "type." 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 11

8-

IN 4

0 . 6 0 .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS

FIG. 9. Audiospectrogram of call of Cophixalus pansus, 300 Hz. filter. Allen Allison no. 7195, recorded September 12, 1980 on Bulldog Road, Morobe Province, Papua New Guinea, calling at a temperature of 12.30C. from 5 cm. below the surface of mossy substratum.

The snout is rounded, verging on bluntly end of the abdomen. There is, however, pointed in slightly desiccated specimens, much variation in both dorsal and ventral canthus rounded, loreal region sloping and color patterns. Occasional individuals are shallowly concave to virtually flat, nostrils dark above and below except for the under- relatively close together. The tympanum is sides of the fingers and toes. The vertebral indistinct and small, less than one-half the stripe may be lacking, or the dorsum may be diameter of the eye. Most individuals show pale, or marked with darker pigment in vary- a prominent postorbital-supratympanic fold ing degrees. The ventral surfaces likewise of skin. The dorsal body surface ranges from exhibit a range from completely unmarked to smooth to rather warty, possibly in part a uniformly dark, though heavily marked frogs reflection of the modes of killing and pres- are a small minority. ervation. Relative lengths of fingers are Variation in the several proportions cal- 3>4>2>1, with the first finger well devel- culated shows no significant sexual dimor- oped, one-half or more the length of the phism, and there are no significant differ- adpressed second finger. Toes are ences between northern (Morobe Province) 4>3>5>2>1, the first being less than one- and southern (Central and Northern prov- half the length of the second. Tips of fingers inces) samples. Statistics for the pooled sam- (fig. 1) and toes are bluntly pointed with no ples (N = 38) are: TL/SV, 0.313 + 0.002 trace of terminal grooves. There are no sub- (0.29-0.34); HW/SV, 0.397 + 0.002 (0.36- articular or palmar tubercles, and only very 0.43); IN/SV, 0.083 + 0.001 (0.072-0.094); low, rounded inner and outer metatarsal el- EN/IN, 0.919 + 0.010 (0.68-1.00); EN/SV, evations. Some features of the skeleton and 0.067 ± 0.001 (0.060-0.075); Eye/SV, musculature have been discussed and illus- 0.106 + 0.001 (0.095-0.116). trated above. CALL: The call is a train of clicks with en- A typical Cophixalus pansus in preserva- ergy widely distributed between about 1700 tive is dark purplish brown above with a nar- and 3700 Hz. but with peaks indicated row pale vertebral line and a pale curved around 2000 and 3000 Hz. (fig. 9). Eight calls postorbital streak, and is largely pale yellow of one individual recorded at 12.3°C. aver- beneath except for dark areas on the chest, aged 1.64 + 0.04 sec. in length (range 1.44- at the base of each arm and at the posterior 1.72). Mean clicks per call were 34.6 + 0.8 12 AMERICAN MUSEUM NOVITATES NO. 2723

WAU

e

POPONDETTA 0

0 10 20 1. MORESBY

FIG. 10. Distribution of Cophixalus sphagnicola (solid and half-filled circles) and C. pansus (open and half-filled circles) in Papua New Guinea. Area above 2400 m. is shaded. 1982 ZWEIFEL AND ALLISON: MICROHYLID FROG 13

(32-39) and mean clicks per sec. 20.7 + 0.7 6253-6259, 5513); Mt. Kumbak, Bulldog (18.1-24.7). Ten calls of a second individual Road, about 13-15 km. south-southwest of recorded at 12.9°C. have the following sta- Wau, about 2775-2800 m. (BPBM 6276, tistics: length, 2.17 + 0.05 (1.87-2.32); clicks 6281-6292, 6345, 6346, 6473, 6476, 6477, per call, 52.9 ± 0.8 (47-57); clicks per sec., 6479-6481); Bulldog Road, 2700 m., 35 km. 24.1 ± 0.4 (22.9-27.3). The interval between south of Wau (BPBM 8312, 8313). Northern calls averaged 1 min. 25 sec. (0:47-2:13, N = Province or Central Province: Mt. Scratch- 7) in the first individual and 1 min. 33 sec. ley, 12,200 ft. (3619 m.) (AM R5238, holo- (0:44-2:50, N = 11). These calls were re- type). Northern Province: Mt. Albert Ed- corded from the northern disjunct population ward, southwest slope, 3680-3811 m. along Bulldog Road. Recordings of calls (AMNH 58103-58105, 58120-58137). Cen- from the southern population sound the tral Province: Wharton Range, 2100-2850 m. same, but were not analyzed in detail. (BPBM 5300, 5301); Murray Pass, Wharton ECOLOGICAL NOTES: Cophixalus pansus Range, 2800-3000 m. (AMNH 53397-53401, is ecologically similar to C. sphagnicola (see 58106-58119, 58138-58141, 81133-81138; above). On Bulldog Road it tends to be BPBM 5292-5299; AM R20839, R20948- slightly more common than sphagnicola and 20951, R21000, R21211-21215, R23464- at higher elevations in the Wau area (e.g., 23508, R24304). grass tussocks on Mt. Amungwiwa-ca. 3200 m.) it is the dominant species. Breeding occurs throughout the year and females may produce up to 30 but generally LITERATURE CITED only 10 to 12 eggs per clutch. The eggs are attended by the male and development is di- Emerson, Sharon B. 1976. A preliminary report on the superficial rect. throat musculature of the Microhylidae Cophixalus pansus attains a much larger and its possible role in tongue action. adult size than does C. sphagnicola. The Copeia, no. 3, pp. 546-551, figs. 1-5. ecological consequences of this difference Fry, Dene B. are under study by Allison. Preliminary in- 1916 [1917]. Description of Aphantophryne, formation suggests that the two species differ a new batrachian genus from New significantly in the kinds and size range of Guinea; with comparative notes on the prey. pectoral musculature. Proc. Linn. Soc. DISTRIBUTION: Cophixalus pansus is New South Wales, vol. 41, part 4, pp. found at high elevations in the Owen Stanley 770-785, pls. 54-55. Gressitt, J. L., and Nalini Nadkarni Mountains and, in an evidently disjunct pop- 1978. Guide to Mt Kaindi: background to ulation, in mountains south of Wau (fig. 10). montane New Guinea ecology. Wau Apparently it is absent from similar eleva- Ecology Inst. Handbook no. 5, pp. tions in the mountains of the Kuper Range vi + 135, pls. 1-4, figs. 1-48, tables 1- (e.g., Mt. Missim, 3200 m.) which form the 12. north side of the Wau Valley. Elevations of Loveridge, Arthur localities with adequate data range from 1948. New Guinean reptiles and about 2800 to 3800 m. A distance of about in the Museum of Comparative Zoology 130 km. separates the northernmost of the and the United States National Mu- Owen localities from the of seum. Bull. Mus. Comp. Zool., vol. Stanley closest 101, no. 2, pp. 305-430. the northern collection sites, but the inter- Menzies, J. I., and Michael J. Tyler vening area has been little studied, and pan- 1977. The systematics and adaptations of sus may occur in much of it. some Papuan microhylid frogs which SPECIMENS EXAMINED: Morobe Prov- live underground. Jour. Zool. (London), ince: Mt. Amungwiwa, about 3000 m., about vol. 183, pp. 431-464, figs. 1-14, pls. 1- 20 km. southwest of Wau (BPBM 6246-6248, 3, tables 1-2. 14 AMERICAN MUSEUM NOVITATES NO. 2723

Menzies, J. I., Michael J. Tyler, and Richard G. figs. 4:1-4:10. In William E. Duellman Zweifel (ed.), The South American herpeto- 1980. Cophixalus Boettger, 1892 (Amphibia, fauna: its origin, evolution, and dispers- Salientia): Proposed designation of type al. Mono. no. 7, Mus. Nat. Hist. Univ. species under the plenary powers. Z. N. Kansas. (S.) 2298. Bull. Zool. Nomen. vol. 36, Zweifel, Richard G. no. 4, pp. 231-235. 1956. Results ofthe Archbold Expeditions. No. Parker, H. W. 72. Microhylid frogs from New Guinea, 1934. A monograph of the frogs of the family with descriptions of new species. Amer. Microhylidae. London, British Museum Mus. Novitates, no. 1766, pp. 1-49, (Natural History), viii + 208 pp., figs. figs. 1-9, tables 1-4. 1-67, 2 maps. 1971. Results of the Archbold Expeditions. Tyler, Michael J. No. 96. Relationships and distribution 1971. The phylogenetic significance of vocal of Genyophryne thomsoni, a microhylid sac structure in hylid frogs. Univ. Kan- frog of New Guinea. Ibid., no. 2469, pp. sas Publ. Mus. Nat. Hist., vol. 19, no. 1-13, figs. 1-6. 4, pp. 319-360, figs. 1-10, tables 1-3. 1972. Results of the Archbold Expeditions. 1972. Superficial mandibular musculature, vo- No. 97. A revision of the frogs of the cal sacs and the phylogeny of Australo- subfamily , family Mi- Papuan leptodactylid frogs. Rec. S. crohylidae. Bull. Amer. Mus. Nat. Aust. Mus., vol. 16, no. 9, pp. 1-20, Hist., vol. 148, art. 3, pp. 411-456, figs. figs. 1-5. 1-79, tables 1-9. 1978. The systematic status of the Aru Islands 1980. Results of the Archbold Expeditions. microhylid frog, genus Microbatrachus No. 103. Frogs and lizards from the Roux. Jour. Nat. Hist., vol. 12, pp. 457- Huon Peninsula, Papua New Guinea. 460. Ibid., vol. 165, art. 5, pp. 387-434, figs. 1979. Herpetofaunal relationships of South 1-14, 1 table. America with Australia, pp. 73-106,