https://doi.org/10.24199/j.mmv.1972.33.08 7 February 1972

AMPHIBIANS AND REPTILES FROM SIMBAI, BISMARCK-,

By David S. Woodruff The Biological Laboratories, Harvard University, Cambridge, Massachusetts, 02138

Abstract

Frogs: Litoria angiana, L. arfakiana, L. micromembrana, Nyctimystes disrupter, N. foricula, N. kubori; skinks: Sphenomorphus flavipes, Emoia pallidiceps, Scincclla stanleyana, S. prehensicauda; and the snake Natrix montana are reported from Simbai (altitude 1770 m) in the Bismarck-Schrader mountains of New Guinea. Taxonomic problems associated with N. disrupta and the lygosomines are discussed but not resolved. N. disrupta is added to the list of montane, riparian anurans known to be host to endoparasitic leeches. Marked sexual dichromatism in S. prehensicauda is described and illustrated. The Simbai anuran fauna appears to be similar to that of the adjacent Kaironk Valley and to that of the Wahgi-Sepik Divide 40 km further S. Finally, Rappaport's study of the regulation of the consumption of small animals by natives, and Bulmer and Tyler's comparison of modern and native classifications of frogs in this area are discussed.

Introduction 2540-3810 mm. Originally the bases of the ranges were covered with lowland rainforest, This contribution to the herpetology of New the flanks of the ranges up to about 1825 m Guinea concerns a small collection of six with lower montane rainforest, and the top of species of hylid frogs and five species of reptiles the ridge with montane or cloud forest. Today constituting the first collection reported from the valleys of the Simbai and Kaironk arc the Simbai Valley. populated by Karam and Maring horticultura- Simbai Patrol Post is located about 113 km lists and much of the lower montane forest has W. of Madang on the NE. side of the Bismarck- been cleared or disturbed (PI. 8, fig. 2). Schrader mountain range (Fig. 1). The settle- Except in the main valleys where settlement ment lies in the upper valley of the Simbai occurs at higher altitudes, only remnants of River at an altitude of about 1740 in. The virgin forest occur below about 1 600 m. Above Schrader Range lies to the N. and W. of Sim- that level the forests arc still relatively undis- bai, the Bismarck Range to the S. and E. The turbed along much of the Bismarck-Schrader SW. side of the ranges are drained at this Range (PL 8, fig. 3). At Simbai, however, the point by the Kaironk River. The Simbai- grasslands and cultivated areas are continuous Kaironk divide is about 2000 m above sea from the Simbai to the Kaironk, so separating level, while in this region the ridges of the and effectively isolating the forests of the ranges vary from 2225 m to 2625 m. The Schrader Range from those of the Bismarck lowlands on either side of the ranges are less and other ranges in the central highlands. than 600 m above sea level. Although the Bismarck-Schrader Ranges are The climate and ecology of the neighbour- still relatively remote they have been visited ing Kaironk Valley have been described by by several biologists and anthropologists. Three Bulmer and Tyler (1968). Conditions near parties collected amphibians and reptiles in the summit of Mt. Kominjim just north of the Simbai-Kaironk area prior to the author's Simbai have been described by Gilliard and visit. Undescribed collections made by H. G. Le Croy (1968), and Rappaport (1967) pro- Cogger in the upper Kaironk Valley and by vides a detailed account of environmental con- Gilliard on Mt. Kominjim are ditions in the Bismarck Range a few miles the late E. T in the collections of the Australian Museum S. of Simbai. The climate of this region is of Natural History subtropical; annual rainfall varies between and the American Museum

57 58 DAVID S. WOODRUFF

M4°F

5°S

BAIYER

MT. HAGEN

Fig. 1 Area of the central — highlands of New Guinea discussed in the text showing the rela- tionships of the main mountain ranges. Inset shows the location of the larger map.

respectively. R. N. H. Bulmer collected nearly Species Represented 500 frogs in connection with a detailed study Anura of the ecology of the Karam in the upper Litoria angiana Kaironk. This collection, which contains at (Boulenger) least 19 species, has been reported by Bulmer Specimens: Two adult males (D 14472 and Tyler (1968). The present collection was 14478). made at the Anglican Mission, Simbai, on 1 Discussion: These specimens agree with Jan., 1966. All specimens were collected dur- Tyler's (1968) description of this morpho- ing the day in nearby native gardens and along logically variable species. Both specimens have the banks of the swiftly flowing Simbai River, a continuous fold on the posterior surface of at an altitude of about 1770 m. The specimens the forearms and nuptial pads a little larger have been deposited in the National Museum but of the same shape of those illustrated by of Victoria; their registration numbers are Tyler (1963, Fig. 3B). This species is now indicated below. known throughout the mountains of New AMPHIBIANS AND REPTILES FROM SIMBAI 59

Guinea, occurring near streams at altitudes of on Mt. Victoria in the 1225-2750 m. (Boulenger 1897) and is widespread in the The Australian hylid frogs previously placed highlands between 1525 and 2275 m (Zweifel in the genus Hyla are here referred to the 1958, Tyler 1963b). One of the five syntypes genus Litoria Tchudi on the authority of of N. papua in the British Museum has subse- Tyler (1971). quently been referred to N. semipalmata Litoria arfakiana Peters and Doria. (Parker 1936). More recently Tyler (1963b) has suggested that the type series may still not Specimens: Three adult males (D14473, be conspecific. The validity 14480-1). of N. disrupta may therefore depend, in part, on the selection of Discussion: These males agree with those a lectotype from among the four remaining described by Tyler (1968). The long unweb- syntypes in the British Museum and a single bed fingers, tubercles on the upper eyelid, heel, syntype in the Museum of Comparative Zoo- and vent (absent from the tarsi), and the logy. As Boulenger's original description of prominent pointed snout, appear to character- N. papua includes an illustration of one indi- ize this species. In life two of the males were vidual (1897, PL 1, fig. 5) the specimen brown dorsally, the other was olive green. represented should become the lectotype. Until L. arfakiana has been collected throughout the such a lectotype is designated N. disrupta must mountains of New Guinea at altitudes of 775- be regarded as a species of doubtful validity 1825 m. possibly synonymous, in part, with N. papua. Litoria micromembrana (Tyler). When captured this frog exuded a slimy secretion Specimens: 13 adults (D 14462-71, 14475, with a sickly odour. A large endo- 14477, 14479). parasitic leech was found in the dorsal lymph sac. Although such parasitism has not been Discussion: These specimens agree closely reported previously in this species, these un- with Tyler's (1968) description of this species. usual infestations are now known to occur in All of the specimens were collected in vegeta- most of the common riparian anura in the tion along the side of the river. This species : Rana grisea, Litoria is now known from a number of localities in angiana, L. arfakiana, L. micromembrana, N. the highlands of central New Guinea at alti- kubori, and N. papua (Mann and Tyler 1963, tudes of 1225-2450 m. Tyler, Parker and Bulmer 1966, Tyler 1968). Nyctimystes disrupta Tyler. Nyctimystes foricula Tyler. Specimen: One adult male (D 1421 8). Discussion: Tyler (1963c) distinguished Specimens: One adult male and one juvenile this species from the morphologically similar (D 14030-1).

N. papua on the basis of differences in pal- Discussion : The specimens agree closely with pebral venation, the degree of webbing on the Tyler's (1963c) original description, being fourth finger, and the ratio of the eye-nostril green above and yellow below in life. The to internarial distance. While the above speci- palpebral venation differs from that of the men agrees closely with the description of sympatric N. kubori in that the veins are

Tyler's holotype it is intermediate between broad, parallel, and oblique. This species has these two taxa in all three diagnostic characters. been reported previously only from the Kai- Specimens collected by H. G. Cogger (pers. ronk Valley. comm. 1966) were also morphologically inter- Nyctimystes mediate between N. disrupta and N. papua as kubori Zweifel. presently defined. Specimens: Four adult males (D 142 14-7). N. disrupta was previously known from the Discussion: This species was described by Kaironk Valley and from near Baiyer River, Zweifel (1958) and subsequently by Tyler about 50 km to the SW. (Tyler 1963c). N. (1963b, c). The present series conform closely papua was described from specimens collected to the males described by Tyler (1963b). This E .

60 DAVID S. WOODRUFF

species is now known from a number of locali- due to the presence of a pigment in the blood ties in the central highlands of New Guinea at plasma, which gives the muscles, bones, and altitudes between 1525 and 1975 m. other tissues a striking green colour. They Reptiua: Squamata report that females are similarly coloured internally but do not appear green Scincella stanleyana (Boulenger). externally because of the presence of brown pigment in Specimens: Seven adults and four juveniles the scales. from Simbai (D 140 18-28). An additional This species is now known from several individual was collected in montane forest at localities in the central highlands about of New 2130 m on the trail between Simbai and Guinea. Kundan (D 14029). Discussion: This small, ground dwelling Sphenomorphus flavipes (Parker). skink was common beneath rotting vegetation Specimen: One adult (D 14005). in the gardens around Simbai. Adults have Discussion: This handsome arboreal skink snout-vent lengths of 48-59 mm. All specimens has a body length of have a 90 mm and a total length dark brown lateral line separating the of 180 mm. Dorsal surfaces copper coloured back from are dark brown the cream under- with cream spots while the belly is a fine surface. In immature individuals there is also mosaic of cream, a yellow, and brown. The tail broken, white, dorso-latcral line extending is marked with 6-7 from the upper longitudinal brown stripes. eyelid onto the flanks. This The specimen was collected 4 m above the species is widespread in the mountains of New ground in a small Casuarina tree. Guinea (Loveridgc 1948, and undescribed Greer and Raizes (1969) note that collections in the Museum of Comparative the predominant ground colour of Zoology ) both sexes of this species is brown, and that there are Scincella three prehensicauda (Loveridge). distinct colour morphs: crossbanded, striped, Specimens: Six females and plain. (D14012-7) and The present specimen belongs in four males (D14008-11). the last group. Loveridge (1948, p. 354) was Discussion: This therefore small arboreal skink with mistaken in describing the variation striking white in eyelids is unusual in possessing his three specimens (referred to Lygosoma marked sexual (Leiolopisma) dichromatism (PI. 8, fig. 1). flavipes) as a case of sexual In life, females are brown dorsally with darker dichromatism. brown crossbars and As in small paler patches, Scincella prehensicauda this species particularly on the also sides. There are conspicu- possesses a green blood pigment (Greer ous dark stripes on and Raizes the sides of the neck. This 1969). According to these authors colouration is little this changed on preservation. phenomenon is known in only one other In contrast the males, amniote: which are uniformly Leiolopisma virens anolis, an arbo- green in life, are real skink a pale golden brown colour from New Guinea and the Solomon in alcohol. The females Is. The are also slightly larger apparent uniqueness of the green blood than the males, as pigment, indicated by the following coupled with certain morphological measurements of individuals features and with undamaged similarity of distribution patterns tails (data in mm). indicate that these three species should probab- ly be placed in a No. Total-length Tail length single genus (Greer, in prepa- Females 4 135-151 70-76 ration). Males 3 J21-I25 6.V68 The Karam natives are well aware of this Emoia pallidiceps De Vis. dimorphism, pointing out to the author in Specimens: One adult and one pidgin Frmlish Jjuvenile that: 'dispela man bilong dis- (D14006-7). pela mari' and vice versa. Discussion: Loveridge (1948) treated Greer and Raizes (1969) have shown how this species as a race of E. baudini. Brown the green colour of the (1953) males of this species is in a subsequent revision of the baudini com- AMPHIBIANS AND REPTILES FROM SIMBAI 61

plex resurrected E. pallidiceps as a distinct to the fact that the collection was made in species. The specimens from Simbai agree native gardens near the river, rather than in closely with Brown's description. The adult the lower montane or montane forest. At this measures 53 mm from snout to vent and is time, therefore, the data suggest that the anu- in 128 mm total length. It is dark brown ran fauna of these two areas on either side of dorsally with black sides and there is a white the mountain range are probably very similar. lateral line running from the ear to the shoul- Although herpetological exploration of the der, continuing as a series of spots to the hind New Guinea highlands is still in its infancy the limb. anurans of the Wahgi-Sepik Divide, about 40 km south of Simbai, have Natrix montana Jeude. been the subject of a number of recent papers by Tyler (1963a, Specimen: One juvenile (D 14004). b, c, 1968) and Zweifel (1956a, b, 1958, Discussion: The specimen has a total length 1962). It can be seen in Fig. 1 that the Bis- of 190 mm, a tail length of 43 mm, divided marck-Schrader Range and the Wahgi-Sepik anal and subcaudal scales, and 15 rows of Divide converge near Mt. Wilhelm in the E. scales at the mid-body. It was collected beneath The forested ridges of these ranges, which are a log on the river bank. still largely unsurveyed, vary in altitude be- tween 2100-3100 m with summits rising up Discussion 4510 m. As the lower montane and mon- Anurans of the Bismarck-Schrader Range tane forests appear to be continuous along Collecting over a period of several years, these mountain ranges, it might be expected Buhner and Tyler (1968) were able to report that the fauna of the Bismarck-Schrader Range 19 species of anurans from the upper Kaironk near Simbai would be similar in species com- Valley and the adjacent forests of the Schrader position to that of the Wahgi-Sepik Divide Range. Ten species of hylids, eight microhy- near Banz or Nondugl. lids, and one ranid occur in that area: The comparison of the faunal lists of these two areas shows that in fact the two are re- Litoria angiana, L. arfakiana, L. bulmeri, markably similar. Sixteen of the 18 described L. micromembrana, L. moclica, Nyctimystes species collected in the upper Kaironk Valley disrupta, N. joricula, N. kubori, N. narinosa, are known from elsewhere in the central high- N. sp. indet, Sphenophryne brevicrus, Cophi- lands. All but one of these 16 (Cophixalus xalus parkeri, C. riparius, C. shellyi, C. variegatus) have been collected on the Wahgi- variegatus, Asterophrys wilhelmana, Xenoba- Sepik Divide. Only two taxa (Litoria bulmeri trachus rostratus, Barygenys sp., and Rana and Nyctimystes joricula) have not yet been grisea. reported from outside the Kaironk-Simbai area. On the other hand, there are five This is essentially a montane fauna: most of species (L. becki, L. darlingtoni, iris, these species are not known from below 1200 L. Metopostira ocellata, and C. darlingtoni) known from m and all are restricted to altitudes of more the Wahgi-Sepik Divide than 600 m. Generally speaking the hylids are which so far have not been found on the Bismarck-Schrader the dominant group in the cleared areas along Range. More detailed comparison of the valley floor while the microhylids are re- the two faunas will have to await the availability of stricted to the forests. data deriv- ed from additional field work, and On the basis of present knowledge, the the clarifi- cation of a number of adjacent Simbai Valley contains the six hylids distributional and taxonomic problems. reported above and Litoria bulmeri which Gilliard collected at 2530 m on Mt. Kominjim (Tyler 1968). All species reported from the Anurans and Man in the Bismarck- Simbai watershed are represented in the larger Schrader Range Kaironk collection. The absence of microhy- Finally attention is drawn to the work of two lids in the Simbai collection is probably due anthropologists who have studied the relations 62 DAVID S. WOODRUFF

between the natives of this area and the non- Acknowledgements domestic fauna. Rappaport (1967) studied the I am indebted to Dr. H. G. Cogger for his role of ritual in the ecology of the Tsembaga assistance in the comparison of the Simbai group of the Maring people who live about specimens with collections in the Australian 10 km south of Simbai. Rappaport (1967) Museum, to Professor R. N. H. Bulmer and and Bulmer (pers. comm. 1966) have drawn Dr. A. E. Greer for useful discussions, to W. attention to the fact that frogs, lizards, and S. Kimpton for assistance in the field, to Turid other small animals constitute a small but Holldobler for the preparation of the photo- important part of the diet of the Maring and graphs, and to Michael J. Tyler for his help Karam people. On several occasions the author in checking my hylid identifications, reading, saw children carrying home frogs wrapped in and helpfully criticizing the manuscript. I am leaves or secured by the legs. Although chil- especially indebted to Father John Cottier and dren in this area do not exhibit symptoms of his wife for their generous hospitality at the severe protein deficiency, Rappaport presented Anglican Mission, Simbai. Grateful acknow- evidence suggesting that their protein intake ledgement is also made to the Trustees of the may be inadequate. It is very interesting, there- National Museum of Victoria for a contribution fore, that in the Tsembaga he found that towards the cost of the author's field work in taboos operated to make most of the subsidiary New Guinea. sources of animal protein available to women and children but precluded them from adoles- References cents and adult males. He concluded that such Boulenger, G. A., 1897. Descriptions of new lizards ritualistic regulation maximizes the value of and frogs from Mount Victoria, Owen Stanley Range, New Guinea, collected by Mr. A. S. such foods for the population as a whole. Anthony. Ann. Mag. Nat. Hist. 6(19): 6-13. It is partly because of the dietary importance Brown, W. C, 1953. Results of the Archbold of small animals like frogs to the natives of Expeditions. No. 69. A review of New Guinea lizards allied to Emoia baudini and Emoia this area that their knowledge of the natural physicae (Scincidae). Amer. Mus. Novit. 1627: history of such animals is so remarkable. Bul- 1-25. Bulmer, R. N. H., and M. J. Tyler, 1968. Karam mer, in his field study of the Karam in the classification of frogs. /. Polynesian Soc. 77: upper Kaironk Valley, recorded the native 333-385. names of all specimens he received. He was Gilliard, E. T., and M. Le Croy. 1968. Birds of the Schrader Mountain region, New Guinea. subsequently able to show that the Karam Results of the American Museum of Natural classification of frogs is very similar to that History Expedition to New Guinea in 1964. adopted by trained herpetologists (Bulmer and Amer. Mus. Novit. 2343: 1-41. Greer, A. E., and G. Raizes, 1969. Green blood Tyler 1968). The exceptions to this generali- pigment in lizards. Science 166: 392-393. zation are particularly interesting as they in- Loveridge, A., 1948. New Guinean reptiles and amphibians volve Litoria angiana and Nyctimystes dismpta, in the Museum of Comparative Zoology and the United States National Museum. taxa which have troubled museum workers. Bull. Mus. Comp. Zool. 101: 305-430. In the case of L. angiana the Karam, like the Mann, K. H., and M. J. Tyler, 1963. Leeches as endoparasites of frogs. Nature 197: 1224-1225. museum taxonomists, have attempted to de- Parker, H. W., 1936. A collection of reptiles and scribe the variation of this highly variable amphibians from the mountains of British New species by reference to a series of taxa. In Guinea. Ann. Mag. Nat. Hist. 10(17): 66-93. Rappaport, R. A.. 1967. Pigs for the Ancestors. the second case Bulmer and Tyler note that Ritual in the Ecology of a New Guinea People. the Karam recognize two speciemes as being New Haven, Yale, xx + 311 pp. Tyler, M. J., 1963a. taxonomic equivalent to the taxon N. disrupta. It may A study of amphi- bians and reptiles of the Central Highlands of well be that future studies will reveal the New Guinea, with notes on their ecology and presence of sibling species presently confused biology. 1. Anura: Microhylidae. Trans. R. Soc. S. Aust. 86: 11-29. under this name. Linguistic studies of the type , 1963b. A taxonomic study of amphibians carried out by Bulmer guide may biologists to and reptiles of the Central Highlands of New a number of hitherto unrecognised species com- Guinea, with notes on their ecology and biology. 2. Anura: plexes. Ranidae and Hvlidae. Ibid. 86: 105- 130. AMPHIBIANS AND REPTILES FROM SIMBAI 63

, 1963c. An account of collections of frogs , 1962. Results of the Archbold Expeditions. from Central New Guinea. Rec. Aust. Mus 26- No. 83. Frogs of the microhylid genus Cophixa- 113-130. lus from the mountains of New Guinea. Ibid. , 1968. Papuan hylid frogs of the genus 2087: 1-26. Hyla. Zool. Verhand. 96: 1-203. , 1971. The phylogenetic significance of vocal sac structure in hylid frogs. Univ. Kansas Pubis. Mus. nat. Hist. 19: 319-360. Explanation of Plate 8 , F. Parker, and R. N. H. Bulmer, 1966. Observations on endoparasitic leeches infesting frogs in New Guinea. Rec. S. Aust. Mus. 15: Fig. 1 Sexual dichromatism 356-359. — in Scincella prehensi- cauda. The upper two specimens are females, Zweifel, R. G., 1956a. Results of the Archbold the lower pair are males. Expeditions. No. 72. Microhylid frogs of New Fig. 2—Simbai Patrol Post (altitude 1740 m) and Guinea, with descriptions of new species. Amer 600 m airstrip near the headwaters of the Mus. Novit. 1766: 1-49. Simbai River. Note that the forest has been , 1956b. Notes on microhylid frogs, genus cleared from the valley floor and lower Cophixalus, from New Guinea. Ibid. 1785: 1-8. ridges. , 1958. Results of the Archbold Expeditions. Fig. 3 —Montane or cloud forest 20-30 m high on No. 78. Frogs of the Papuan hylid genus the trail between Simbai and Kundan at an Nyctimystes. Ibid. 1896: 1-51. altitude of about 2100 m. MEM. NAT. MUS. VICT. 33 PLATE 8