AMERICAN MUSEUM

Noviltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2775, pp. 1-28, 1 fig., tables 1, 2 January 27, 1984

Results of the Archbold Expeditions. No. 1 12. The of the Huon Peninsula, Papua

S. B. MC DOWELL'

ABSTRACT The snakes known from the Huon Peninsula form are used for discriminating the species. A (from the longitude ofLae eastward) are listed and misidentification of Typhlops inornatus as Ram- discussed, mainly on the basis of specimens col- photyphlops flaviventer is corrected. The lected by the Seventh Archbold Expedition but fauna, like the frog and lizard faunas previously using other collections as well. The of discussed by Zweifel, is most easily exr iA;, as the genus in the Australian region is the result of dispersal to a Pliocene island that discussed and the following species are recognized: became joined (probably in the Pleistocene) to the D. punctulatus (including D. lineolatus); D. calli- New Guinea mainland, with a lowland fauna oc- gastra, D. salomonis, D. lorentzi, D. papuensis cupying this zone of juncture. Little, if any, en- (these four have usually been regarded as conspe- demicity is indicated for the Huon snake fauna cific); and D. gastrostictus (including D. meeki); and there is no special resemblance to the snake hemipenial morphology, dentition, and braincase fauna of nearby New Britain and Umboi.

INTRODUCTION This represents a long-delayed report on field observations of Huon Peninsula snakes, the snakes collected by the Seventh Archbold and for sharing his knowledge ofNew Guinea Expedition to the Huon Peninsula with par- herpetology; to Dr. Charles Myers (AMNH) ticular attention to the region of Mt. Rawlin- for comments on the manuscript; to the late son. Zwiefel (1980) has already reported on Mr. Hobart Van Deusen for his field obser- the frogs and lizards from there. vations and geographic information; to Dr. Karl F. Koopman, Department of Mammals ACKNOWLEDGMENTS (AMNH) for discussions of bat distributions I am indebted to Dr. Richard G. Zweifel in the New Guinea region; to Dr. Guy Mus- ofthe American Museum ofNatural History ser, Department of Mammals (AMNH) for (AMNH) for use of specimens, for use of his identification of murid stomach contents; to

' Professor of Zoology, Department of Zoology and Physiology, Rutgers University, Newark, New Jersey 07102.

Copyright © American Museum of Natural History 1984 ISSN 0003-0082 / Price $2.15 2 AMERICAN MUSEUM NOVITATES NO. 2775

Dr. Ernest E. Williams for use of specimens that of the eastern end of the Finisterre-Sa- in the Harvard Museum ofComparative Zo- ruwaket elevation (the exclusion of the west- ology (MCZ); to Dr. George Zug for use of ern end of this elevation has the practical specimens in the National Museum of Nat- advantage of concealing a colossal ignorance ural History, Smithsonian Institution of the fauna of a part of (USNM); to Dr. Alan Ziegler for use of spec- that has been insufficiently collected). "Lae" imens in the Bernice P. Bishop Museum in the literature has had various meanings, (BPBM); to Miss Alice Grandison for use of including mountains to the west or south of specimens in the British Museum (Natural Lae and separated from the Huon Peninsula History) (BM); to Dr. Gunther Peters of the by the Valley, and the in- Zoologisches Museum an der Humboldt- clusion of Lae as part of the Huon Peninsula Universitat zu Berlin (ZBM) for use of the introduces some problems of ambiguous lo- Sattelberg specimen of Typhlops inornatus; cality references. and to Dr. Marinus Hoogmoed for use of The localities of the Huon Peninsula, as material in the Reijksmuseum van Natuur- here defined, are given in Van Deusen (1977) lijke History, Leiden (RMNH). and Zweifel (1980). These papers also give a summary ofprevious work on the Huon Pen- THE GEOGRAPHIC SETTING insula fauna, to which should be added two "Huon Peninsula" is here used in the same small collections: Potter collected some spec- sense as used by Zweifel (1980), to include imens, now in the British Museum (Natural the portion of mainland New Guinea pro- History) from "Huon Gulf" which may have jecting east of the longitude of Lae, between come from the northern shore of the Gulf the Vitiaz Strait and the Huon Gulf. Lae is (i.e., the Huon Peninsula); M. C. Kurtz col- included in the "Huon Peninsula," even lected some specimens, now in the American though much of the fauna of Lae is from the Museum of Natural History, from Lae and Markham Valley; Saidor is not included in from Gusiko. the "Huon Peninsula," since Saidor is west Unless otherwise noted, specimen num- of Lae, even though Saidor is on the north bers given below are American Museum of coast of the Finisterre-Saruwaket region and Natural History numbers (AMNH). might be expected to have a fauna similar to

SYSTEMATIC ACCOUNTS TYPHLOPIDAE ly studded with glands, centrally as well as Typhlops inornatus Boulenger along scale margins; no distinct rectal cae- ZMB 24345 Sattelberg cum. This re-identification removes Ram- Vogt (1911) and Sternfeld (1913) listed photyphlopsflaviventer from the list of Huon from on the Peninsula records and that species would have "Typhlopsflaviventer" Sattelberg, a range from Ternate and Mo- basis of this specimen, which is a female Halmahera, similar in most rotai, Batjan, Batanta, Salawati, the Vogel- Typhlops inornatus, respects kop and Onin Peninsula of western New to the Bulolo specimen described and figured Guinea, Manus, New Britain and Duke of by McDowell (1974). ZMB 24345 has: scales York Is., Nissan Atoll, the Solomons, and 22-22-22, with 359 + 15 along the back be- Vuna Pi in tween rostral and terminal caudal spine; nasal Fiji. cleft almost complete, from very near rostral Sternfeld to supralabial II, horizontal in its anterodor- Typhlops depressiceps sal portion; supralabial II overlapping lower 66739 Lae end of preocular; supralabial III overlapping This specimen has already been used in the lower end of a subocular cut off from the description given by McDowell (1974); since ocular; preocular undivided, larger than ocu- then, it has been discovered that this species lar; nasals separated by rostral-prefrontal ranges to the Trobriand (Kiriwina) Islands, contact; nasals, preoculars, and rostral dense- where Heatwole (1975) recorded it (I have 1984 MC DOWELL: HUON PENINSULA SNAKES 3

seen a Trobriand specimen and agree with and Nissan Atoll) make up a superspecies and this identification); it is not unlikely that this it is interesting that one of the two Finsch- seemingly rare species has a fairly extensive hafen specimens (95532, juvenile female) distribution in New Guinea. The Lae speci- shows faint traces of light and dark rings on men was discovered about a foot under sod the body, an approach to L. boa. This feature and was, unfortunately, cut in half by the cannot be attributed entirely to juvenility, spade; attempts to recover the rear half were since Brongersma (1953) did not find cross- unsuccessful. bands even in late embryos from Fak-Fak. However, no trace of dark crossbars can be Ramphotyphlops erycinus (Werner) found in the other Finschhafen specimen (95533, adult male). This latter specimen was 66737, 95513 Lae found dead on the road, but 95532 was found In addition to these Lae records, Loveridge crawling on the tunnel ceiling about 200 ft. (1948) has recorded this species from Finsch- inside Seborgisung Cave and had a ball of hafen. The species has a fairly extensive dis- hair in the intestine. tribution in northern New Guinea, extending westward to the Mamberamo River in Irian Python amethistinus (Schneider) Jaya. 95530 Mt. Rawlinson at south ridge above Ramphotyphlops braminus (Daudin) Gang Creek, about 5200 ft.; 95531 Masba Creek, about 2000 ft.; 95141-42 Lae 66738, 66740, 95130, 95512, 126580-85 Lae The first ofthese two specimens represents It is certain that this species is now estab- the highest altitude record for the species, lished at Lae; whether or not it occurred there which elsewhere in its extensive distribution before World War II is uncertain. As an all- through the Moluccas, New Guinea and ad- female species, presumably able to propagate jacent islands, and Queensland is not known itself in a new locality from a single individ- from above 4800 ft. (AMNH 82327, Mt. Vale, ual (see McDowell, 1974), and a commensal western New Britain) and most records are of man, this form is readily introduced by from between sea level and 1000 m. The total man and has a very extensive distribution; length of 95530 was 3.47 m. (body length just where it is autochthonous is unknown. 2.94 m.); 95531 had a total length of 121 in. (3.07 m.) (collectors' measurements; both BOIDAE specimens are skinned out, except for head and tail; both are females, without note of Liasis papuanus Peters and Doria any stomach contents). Zweifel (field notes) 66761 Lae records that 95530 had been in a hole 90 ft. This species is found almost throughout above ground level in the trunk of a tree of mainland New Guinea, as well as on Mysol, the genus Syzgium. Biak, and Fergusson Islands. According to M. Another species of Python, P. boeleni, has C. Kurtz's field notes, the Lae specimen was been recorded from "Lae" by Worrell (1958; taken in the morning in a field of Kunai and as Liasis taronga). No other specimens have cane grass. been taken from Lae, although the species is known from the Eastern Highlands west of Liasis albertisii Peters and Doria Lae and from Wau, southwest of Lae, and as noted by Brongersma (1969), this "Lae" rec- 95532 Finschhafen at about 500 ft.; 95533 ord probably refers to the mountains behind same, but sea level. Also, from Lae and vi- than to Lae itself. cinity: 66746, 66756, 95531, 95535, 103869, Lae, rather 107148 Chondropython viridis (Schlegel) Widespread throughout New Guinea (also Salawati, Biak, Normanby, and Mussau) at 95131-33 Lae; 95521 Butibum R., 7 mi. n from sea level to 4000 ft. and one ofthe more of Lae; 95522-23 Kabwum, about 4500 ft.; common species. McDowell (1975) suggests 95524-25 Pindiu, 3000 ft.; 95526 Masba that this species and L. boa (Bismarck Ar- Creek; 95527 Numbut; 95528 Kotkin, Mt. chipelago other than Mussau; also Umboi Is. Rawlinson; 95529 "Huon Peninsula." Also 4 AMERICAN MUSEUM NOVITATES NO. 2775 recorded from Sattelberg by Lonnberg (1900), behaving as one species or as two in the Lae Vogt (1911), and Sternfeld (1913) region, and it is hoped that those with ready Both 95527 and 95528 had hair in the access to Candoia carinata in the Lae region stomach; the stomach was empty in the oth- will settle the question. ers, aside from nematode parasites found in It may be noted that Umboi specimens (see all except 95526. All show the northern color McDowell, 1979) are short-tails, but differ pattern for their species, with the white scales from Huon Peninsula specimens in color, scattered randomly rather than organized into having one pair oflongitudinal stripes or none a vertebral stripe (as it is in specimens from on the neck, rather than the two pairs of lon- southern New Guinea). gitudinal stripes seen in Huon Peninsula This is a widespread and fairly common specimens. New Britain specimens are long- New Guinea species (also Mysol, Salawati, tails. the Arus, and Geelvink Bay islands) but is unknown from Umboi or the Bismarck Ar- Candoia aspera (Gunther) chipelago. 95515 Masba Creek, 2000 ft.; 66741-44, 95137, 95514 Lae; BPBM 3727, 5507-08 Candoia carinata (Schneider) Busu R., nw of Lae, about 100 ft. Also re- 95516 near Pependangu, about 3200 ft.; corded from Sattelberg by Vogt (1911), and 95517 near Pindiu, about 3000 ft.; 95519- from Finschhafen by Loveridge (1948) 20 Finschhafen; 66747, 85665, 95138-30, The Masba Creek specimen had an empty 95518, 103635 Lae stomach, but BPBM 5507 (Busu R.) regur- As discussed by McDowell (1979), most gitated a partly digested frog and lizard when populations of this wide-ranging (from Su- chloroformed (F. J. Radovsky's notes on la- lawesi to the Santa Cruz Is.) species fall clear- bel) and 66741 (Lae) contained skink frag- ly into two classes: long-tails, with all indi- ments and a beetle, the latter probably sec- viduals having more subcaudals than ondarily ingested. This species is found predicted by the regression line describing the throughout New Guinea (except the Oriomo closely related species C. bibroni (expected Plateau and the higher elevations of the cen- subcaudals = 1.13 + 0.238 ventrals); and tral mountainous spine) and the Bismarck short-tails, with all individuals having fewer Archipelago, Manus Group, Geelvink Bay is- subcaudals than predicted by the regression lands, Waigoe, Batanta, Salawati, and Mysol. line for C. bibroni. These two classes, defined by subcaudal count relative to ventral count, fit well-but not exactly-with the occur- BOULENGER rence of certain color patterns, frequency of GENUS DENDRELAPHIS pelvic spurs in females, length of separation All New Guinea-Solomons-Australian between furcation of the hemipenis and of members ofthis genus (except for the Sunda- sulcus spermaticus, and shape of the post- Malayan D. pictus, known also from Mysol) orbital bone. seem closely related; they have but 13 scale The four specimens from the eastern Huon rows, lack longitudinal black stripes on most Peninsula (95516-17, 95519-20) are typical or all of their body, and have a medium or short-tails, not much different from speci- short hemipenis (not extending past the fif- mens from the eastern Papuan Peninsula and teenth subcaudal). Previous taxonomy of d'Entrecasteaux Is. Lae, on the other hand, Australasian Dendrelaphis has not taken is anomalous and of the six Lae specimens hemipenial structure into account, and when seen by me, three (66747, 95138-39) are typ- the hemipenis is considered, "D. calligastra" ical long-tails (similar to Irian Jaya, north appears to represent at least three quite dis- coast of Papua New Guinea, and Bismarck tinct species (D. lorentzi, D. papuensis, and Archipelago specimens) but the other three D. calligastra) with the last of these probably (85665, 95518, 103635) are typical short-tails. closely related to, but distinct from, the Sol- The sample is too small to permit conclusions omon Island form, D. salomonis, that has as to whether short-tails and long-tails are usually been considered conspecific with it. 1 984 MC DOWELL: HUON PENINSULA SNAKES 5

Since my re-interpretation of Dendrelaphis Group I, but is better defined by the diagonal considerably affects my identification ofHuon fold of the calyculate zone. Peninsula material, it is necessary to give a Represented on the New Guinea mainland brief summary of my conclusions. by three species, D. gastrostictus (Boulenger), On the basis of hemipenial structure, Aus- D. punctulatus (Gray), and D. calligastra tralasian Dendrelaphis fall into three groups: (Gunther), all occurring sympatrically in some the D. lorentzi Group, the D. punctulatus parts of their ranges and also occurring sym- Group, and the D. papuensis Group. The D. patrically with Group I and Group III; D. punctulatus Group contains four species, but punctulatus and D. calligastra also occur in the other two groups are monotypic. These eastern Australia. In addition, a fourth species groups and species differ as follows: (?), D. salomonis (Gunther) occurs allopat- I. Dendrelaphis lorentzi Group. The organ rically to the others throughout the Solo- is moderately long (ending opposite subcau- mons, from Bougainville to the Santa Cruz dal 12 to 15), with the major retractor muscle Is., and seems about equally similar to the attaching almost, but not quite, at its tip, so three New Guinea species; it may be con- that there is a short uninverted terminal awn specific with one of the New Guinea species, formed by the narrowed extreme tip of the but I am unable to guess which one, and rec- organ lying distal to the attachment of the ognize it as distinct in part out ofexpediency major retractor; the distal part of the organ (Misima I. Dendrelaphis are peculiar in being has numerous closely packed calyces, but each melanistic, but similar in dentition, hemi- calyx has numerous small spines along its penis, and counts to Solomon Is. specimens border, so that the general texture ofthe distal and are here referred to D. salomonis). end of the organ is spinose, with inconspic- IIA. Dendrelaphis gastrostictus (Boulen- uous longitudinal folds between the bases of ger) [including D. meeki (Boulenger) and D. the spinules; proximal to this spinose-calyc- nouhuysi (van Lidth de Jeude)]. Specially ulate region there is a region of numerous characterized by the short and wide brain- spines not mounted on calyces, but each spine case, an index of which is the shape of the much less than a subcaudal in length; this part of the supraoccipital bone exposed be- spinose zone is distinctly, but not conspicu- tween the jaw adductor muscles, covering its ously, set offfrom the spinose-calyculate zone. lateral edges, and anterior to the axial mus- Only one New Guinea species, D. lorentzi cles that cover the rear portion of the bone; (van Lidth de Jeude), but the Lesser Sunda the exposed supraoccipital is much broader D. inornatus Boulenger has a similar hemi- than long (easily ascertained by slitting the penis (although having 15 scale rows). D. lo- skin just behind the parietal scutes). rentzi is known to occur sympatrically with Maxillary teeth very numerous, 30-41, last New Guinea members of Group II. 3-5 longest (this count diagnostically greater II. Dendrelaphis punctulatus Group. The than for D. lorentzi, D. calligastra, D. pa- organ is short (not past subcaudal 10) and has puensis, but with slight overlap with D. sa- a subterminal attachment of the major re- lomonis, considerable overlap with D. punc- tractor muscles, so that (as in Group I) there tulatus). is a narrow and uninverted terminal awn, Ventrals few, 160-180 (this count diag- which may be short (D. gastrostictus, D. sa- nostically lower than for D. punctulatus, lomonis) or several caudals long (D. punc- slightly overlapping lowest variations of D. tulatus, at least some D. calligastra); distally, salomonis and D. papuensis, broadly over- the organ has large and longitudinally drawn- lapping D. calligastra, not distinguishing from out calyces that are almost or quite without D. lorentzi). spinules, so that the general texture of the Hemipenis short (ending opposite subcau- inverted distal end of the organ is one of dal 6-9) with short (usually less than two cau- closely packed longitudinal soft folds; the dals long) terminal awn that bears small spi- proximal edge of this calyculate zone forms nules and a continuation of the sulcus. a diagonal free fold partially hiding the more A blackish stripe on temporal region de- proximal spinose zone; this proximal spinose fining pale lip from dark crown (this stripe zone has numerous small spines, much as in may or may not continue forward onto snout). 6 AMERICAN MUSEUM NOVITATES NO. 2775

Known from New Guinea mainland, includ- Maxillary teeth longest posteriorly, 20 to ing the elevated interior, from Manokwari, 33 in number (slightly overlapping D. gas- and from Normanby Is. and Fergusson Is. trostictus counts and broadly overlapping IIB. D. calligastra (Gunther). Specially counts for D. calligastra, D. /orentzi, D. sa- characterized among Group II forms by hav- lomonis, and D. papuensis; however, lower ing maxillary teeth beneath prefrontal-max- counts for D. punctulatus come from Aus- illary articulation at least as long as-usually tralian material, outside the range of other longer than-the posterior maxillary teeth species of the genus with the possible excep- (seen also in D. papuensis) and by low num- tion ofD. calligastra, and in New Guinea and ber of maxillary teeth, 19-25 (diagnostically the Bismarck Archipelago, maxillary tooth lower than D. gastrostictus, overlapped by D. counts for D. punctulatus are 25-33, diag- salomonis and D. punctulatus, not distin- nostically higher than for D. lorentzi and D. guishing from D. lorentzi and D. papuensis.). calligastra and all but eastern Louisiade Is. Ventral count moderate, 169-199 (over- specimens of D. papuensis; there is broad lapping counts for all other Australasian overlap with D. salomonis in this character). species). Ventrals numerous, 185-219 (diagnosti- Exposed supraoccipital longer than wide. cally higher than for D. gastrostictus and, Terminal awn of hemipenis long to very probably, for D. lorentzi; overlap ofthis count long (at least two subcaudals) without con- with that for D. calligastra, D. papuensis, and tinuation ofthe sulcus, smooth or with a few D. salomonis is too extensive to allow iden- minute spinules; calyculate region proximal tification ofan individual specimen, although to the awn with few or no spinules; hemipenis in all regions where D. punctulatus and D. of moderate length (6-1 1, usually 8 caudals). calligastra occur together and sample is ad- A blackish stripe on temporal region iso- equate, the mean ventral count for D. punc- lating the pale upper lip from the dark crown, tulatus is higher than that for D. calligastra); continued forward on snout to the rostral. highest ventral counts for D. punctulatus- With an extensive distribution from the that is, those most different from counts for Moluccas to the Bismarck Archipelago, per- D. calligastra- come from Australian ma- haps including Normanby Is. but not the oth- terial, where maxillary count is least efficient er islands of Milne Bay Province, and to in separating D. punctulatus from D. calli- Queensland; but except on the eastern edge gastra. ofthe Eastern Highlands, not penetrating the Hemipenis as in D. calligastra, with sim- elevated interior of New Guinea, although ilarly long awn, but sulcus extending along extensively distributed around the rim ofthe awn nearly or quite to the tip; inverted organ island. ending opposite subcaudal 6-10, modally IIC. D. punctulatus (Gray) [including D. subcaudal 8. lineolatus (C. Dumeril); D. macrops (Gun- Palau Is., Mysol, Waigoe, Aru Is., New ther); D. elegans (Ogilby)]. Specially charac- Guinea, d'Entrecasteaux Archipelago, Bis- terized by large size (all Australasian Den- marck Archipelago, Torres Strait Is. and east- drelaphis with a snout-vent length of a meter ern Australia. or more belong to this species) and differs IID. D. salomonis (Gunther). With a con- from all other Australasian Dendrelaphis ex- fusing combination of the characters of D. cept some specimens of D. salomonis in: (1) punctulatus and D. calligastra (allopatric to lacking a distinct black band along the tem- both and quite possibly conspecific with one poral region isolating the dark crown from or the other). The types (BM 1946.1.5.97 and the pale upper lip, the dark crown usually 1946.1.6. 11, "Salomon Is.") have a low max- fading into the pale upper lip or extending illary tooth count (21 in each, left maxilla) raggedly onto the supralabials; and (2) having and a sharply defined dark temporal streak numerous large scale pits (slightly larger than on the dorsal ends of the supralabials, as in those of body scales) on the loreal, ocular, D. calligastra, combined with numerous large anterior temporal, and anterior supralabial sensory pits on the preocular, loreal, supra- regions. labials, and temporals and a high ventral Exposed supraoccipital longer than broad. count (190, 191, respectively) as in D. punc- 1984 MC DOWELL: HUON PENINSULA SNAKES 7

M A ~~~BC FIG. 1. Left hemipenis (inverted, opened by medial incision) of: A. Dendrelaphis lorentzi (MCZ 54329. Papua New Guinea: Western Province: Aramia R.), showing Group I organ; B. D. salomonis (MCZ 72053, Papua New Guinea; Bougainville Is.: Kunua), showing Group II organ; C. D. papuensis (AMNH 76648, Papua New Guinea: Sudest [=Tagula] Is.), showing Group III organ. 8 AMERICAN MUSEUM NOVITATES NO. 2775 tulatus. However, most specimens have the competition between the species, but in the reverse combination: a high maxillary count Solomons, where only one species is present (25-31 teeth) as in D. punctulatus, combined and this competition is absent, random vari- with a rather low ventral count (173-189 for ation is permitted in characters that are under Buka, Bougainville, Fauro, Rubiana [=New strong selective restraint in New Guinea. Georgia], Kolombangara, Choiseul, Ysabel, Quite possibly, protein electrophoretic study Malaita, and Guadalcanal, but 190-195 for of New Guinea and Solomon Dendrelaphis Vella Lavella and Gizo, and 195 for single would resolve the problem and might well Bio Is. specimen seen) suggesting D. calli- yield important insights into the general gastra. problem of species formation. All the Solo- Posterior maxillary teeth always longest. mon Is., from Buka and Bougainville to the Exposed supraoccipital longer than wide Santa Cruz Group. (but only slightly so in the type BM In addition, the melanistic Dendrelaphis of 1946.1.5.97, in which the exposed supraoc- Misima Is. (76691, 76693 [males], 76692, cipital is broader than usual for either D. cal- 76694 [females] examined) seems to fit best ligastra or D. punctulatus, thus making an with this species, having a similar hemipenis, approach to D. gastrostictus). maxillary teeth 28-29 and longest posterior- Hemipenis most like that of D. gastrostic- ly, ventrals 178-179 (males) or 189-191 (fe- tus, with short terminal awn (less than two males). subcaudals long, usually less than one) that III. Dendrelaphis papuensis Group. The bears a well-defined continuation of the sul- organ is of moderate length (ending at sub- cus spermaticus, but organ usually longer caudal 8-13; opposite subcaudal 8-12 in five (ending opposite sixth to tenth, usually eighth Woodlark Is. males), with a strictly terminal to tenth, subcaudal). attachment of the retractor longus, so that Although a blackish temporal streak may there is no suggestion of a terminal awn; dis- be present, even extending to the rostral (it tally, the organ has numerous shallow calyces does so in BPBM 3437, but not in BPBM with many small spines on their borders; 3705, both female, February 1964, Pepele, proximally there are one to five transverse Kolombangara Is.), most often there is no whorls of large spines, some of which are at concentration ofpigment along the temporal- least equal to one subcaudal in length. Spec- supralabial juncture but the dark pigment of imens from the New Guinea mainland, Fer- the crown stops abruptly and evenly on the gusson, Trobriand, Sudest, and Rossel Is. have dorsal ends of the supralabials as if defined a distinct transverse fold between the calyc- by a temporal streak. ulate and spinose zones, the obvious hom- Except for the types, conspicuous sensory ologue of the fold between the (smooth) pits absent from side ofhead in material seen calyces and small proximal spines of Group by me. II Dendrelaphis, but no such fold is evident Several hypotheses, between which I can- in Woodlark Is. specimens or in AMNH not choose at present, might explain the Sol- 42400 (Whitney South Sea Expedition, omon Islands Dendrelaphis: (1) they are a "Bougainville"; but very extensive collecting relict of a stage when the characters that now on Bougainville by Fred Parker failed to con- distinguish reproductively isolated New firm the presence of D. papuensis on that is- Guinea species were individually variable land). within populations; (2) the Solomons Den- Probably one species, Dendrelaphis pa- drelaphis is of hybrid origin, derived from puensis Boulenger. multiple invasions, perhaps before the pres- Dendrelaphis papuensis occurs on the east- ent reproductive isolation of New Guinea em Papuan peninsula, extending at least as Dendrelaphis had been achieved; (3) the far westward as Konedobu and Kila Kila, characters that mark off the species of New Central Province (MCZ specimens) and (fide Guinea Dendrelaphis (but are not in them- Fred Parker) probably into Western Prov- selves the basis for reproductive isolation or ince, and at least as far northward as Garaina, ecological distinctiveness) are maintained in Morobe Province (AMNH specimens), as well New Guinea by selective pressure arising from as Popondetta and Sangara, Northern Prov- 1 984 MC DOWELL: HUON PENINSULA SNAKES 9 ince (BPBM 3802, 3814), the Cape Vogel re- 178-193, overlapping the observed range of gion of Milne Bay Province mainland D. calligastra from the same region (173- (AMNH specimens) and Fife Bay, Milne Bay 186) but the difference between the means mainland (BM 1946.1.6.7, a decapitated (186.44 for 16 D. papuensis and 181.31 for male, cotype of Dendrelaphis schlenckeri 13 D. calligastra) is significant at the 1 per- Ogilby; another cotype ofD. schlenckeri from cent level (F1 27 = 12.47); Fergusson Is., Fife Bay, BM 1946.1.6.9, is a male D. cal- Trobriand Is., Sudest Is., and Woodlark Is. ligastra). specimens of D. papuensis approximate the On the eastern Papuan peninsula, D. pa- range for eastern Papuan peninsula speci- puensis is sympatric with D. punctulatus, D. mens, but for 11 specimens from Rossel Is. gastrostictus, and D. calligastra of Group II, the observed range is 183-203 (two lowest but in this region of overlap, D. papuensis is counts 183, 184 are two females, therefore characterized by the color of the interstitial not certainly identifiable to this species and skin of the neck (exposed by the inflation of differing from remainder of series in having the neck in members of this genus and prob- one scale pit, rather than two, and the lowest ably analogous to the dewlap display ofAno- maxillary tooth counts, 22 and 23, of the se- lis lizards); D. papuensis has black interstitial ries; omitting these two specimens, the ob- skin with a longitudinal white stripe between served range in ventrals for Rossel Is. is 189- the first and second scale rows, but the Group 203). A color pattern feature seen in many- II species show either alternating vertical but not all-D. papuensis but unknown in the white and dark bars on the interstitial skin Group II species is the presence of a pale or a spangled pattern of uniformly dark in- vertebral stripe. terstitial skin with a concealed white spot at Of the six Australasian species of Dendre- the base of each scale. laphis recognized here, all but D. papuensis Outside this region ofsympatry, interstitial and D. salomonis are known from the Huon skin pattern is not diagnostic. For example, Peninsula. BM 1946.1.6.57, male from the type series ofD. papuensis, from the Trobriand Is. (where Dendrelaphis lorentzi this is probably the only species ofthe genus) (van Lidth de Jeude) has black interstitial skin on the neck with a USNM 119505 Gusiko concealed white spot at the base ofeach scale; The above specimen, previously reported this spangled pattern is found in some D. by Loveridge (1948) as Ahaetulla calligaster calligastra from northern and northeastern schlenkeri, is a male with the hemipenis ex- New Guinea. Conversely, north of the range tending to subcaudal 13, with longitudinal of D. papuensis, D. calligastra may show a rows of small spines (each about one-sixth of longitudinal white stripe between the first and a subcaudal long), the rows of spines nearly second rows, usually in conjunction with al- to the tip and well distal to the rightward ternating black and white bars on the inter- angulation of the sulcus at subcaudal 9; an stitial skin at level of rows 5 and 6 (seen in apical awn, about three subcaudals long and some Wewak, Jayapura and Manokwari with numerous tiny spinules; no crossfold on specimens of D. calligastra and also in a Lae organ, but lips of sulcus raised as a pair of D. punctulatus, AMNH 95143). prominent folds. The 23 (left) maxillary teeth In the region of sympatry with Group II slightly (and equally) enlarged beneath the species, as well as in the Trobriand Is., D. prefrontal and at the rear of the series. Scales papuensis has a maxillary dentition similar 13-13-1 1; ventrals 181; anal divided; sub- to that of D. calligastra, with the teeth be- caudals 134 + spine. Nasal scute entire above neath the prefrontal articulation longest and nostril, a narrow strip of cornification ex- tooth count 19-23; but for 11 Rossel Is. spec- tending between nostril and internasal; su- imens the count is 22-27; four Sudest Is. pralabials 9 (fifth and sixth entering eye); one specimens have 26 maxillary teeth; and five preocular; two postoculars; temporals 2+2+2; Woodlark Is. specimens have 26-27 teeth. infralabials 10 (first six touching genials). The observed range in ventrals for D. pa- Color (as preserved) light bronze-tan above puensis from the eastern Papuan peninsula is with pale bluish gray crossbands, this pattern 10 AMERICAN MUSEUM NOVITATES NO. 2775 extending throughout the body; dorsal sur- Guinea D. lorentzi have eight supralabials face ofhead with conspicuous black spotting; except for the Tabubil specimen with nine on a longitudinal black stripe on side of neck the left side; the occurrence of nine suprala- extending along dorsal ends ofpostorbital su- bials bilaterally in the Gusiko specimen sug- pralabials as a postocular streak, represented gests a difference in frequency for this count on snout by row of black spots reaching ros- between southern New Guinea and the Huon tral; interstitial skin ofneck with bold vertical Peninsula. The Gusiko specimen is also the black and pale bars extending to the vertebral only Dendrelaphis I have seen with the entire region. Nuchal line of attachment of axial supraoccipital covered by the axial muscu- muscles at anterior edge of supraoccipital, so lature. that there is no exposed supraoccipital bone. Considering the disjunction in distribu- Except for this Huon Peninsula specimen, tion, these differences are not surprising. I do the species is known only from southern New not believe these differences are more than Guinea, at least as far west as Sabang on the intra-specific, but I wish to call attention to Lorentz River, Irian Jaya (type locality) and them because the Gusiko isolate of D. lo- as far east as the Aramia River, Western rentzi seems to be the nearest approach to an Province, Papua New Guinea (MCZ speci- endemic species among the Huon Peninsula mens), extending at least as far inland in snakes. Western Province as Tabubil, 2000 ft. (MCZ 135006). The Huon Peninsula specimen thus Dendrelaphis gastrostictus (Boulenger) represents a notable disjunction of distribu- tion (the only such case among Huon Pen- 95569 Kabwum, about 4500 ft.; 95570 Mas- insula snakes). At least the specimen "resem- ba Creek, 2100 ft.; 66669 Gusiko bling var. D. in Boulenger's 'Catalogue of The Kabwum specimen is a juvenile with Snakes'" recorded from Simbang (near 37 left maxillary teeth, 164 ventrals; the Mas- Finschhafen) by L6nnberg (1900) as Dendro- ba Creek specimen is a male with 36 left max- phis calligaster may be this species. illary teeth, 170 ventrals; the Gusiko speci- Although the structure of the hemipenis, men is a juvenile male with 34 left maxillary the black spotting on the top ofthe head (like teeth, 170 ventrals. In general, this species is calligraphic penciling) and the continuous characterized by having the highest maxillary cornification ofthe nasal scute above the nos- counts and lowest ventral counts (except for tril all indicate that this specimen is referable southern New Guinea D. lorentzi) and these to D. lorentzi, the Gusiko specimen differs in characters seem to be accentuated in the Huon several particulars from southern New Guinea Peninsula specimens. The interstitial skin of specimens. The apical awn of the hemipenis the neck is black in the Masba Creek and is longer (only 1-2 subcaudals long in south- Gusiko specimens. ern New Guinea specimens; five males, in- Except for the extension of D. calligastra cluding type of Dendrophis lorentzi, exam- into the Eastern Highlands, D. gastrostictus ined). The ventral count of the Gusiko seems to be the only species of the genus to specimen is higher than for southern New extend into the elevated interior ofthe island, Guinea specimens, which show a range of but it is not confined to high altitudes and 156-173 (both sexes; for males, 156-170). extends to the lowlands of Western Province Even when all southern New Guinea D. /o- (e.g., 59897, Lake Daviumbo, and MCZ rentzi are grouped together without distin- 54226-27, Aramia River). Nonetheless, there guishing between sexes or localities (which, are no specimens in the American Museum by ignoring possible sex dimorphism and lo- of Natural History from Lae or the lower cal differentiation, may overestimate vari- Markham River Valley, in spite of consid- ance within the sample and, thus, overesti- erable collecting there, and the species may mate the standard error of the mean), the be excluded from low elevations in north- difference between ventral count of southern eastern New Guinea, apart from the Gusiko New Guinea (N = 12) and Gusiko (N = 1) is record (on the east coast of the Huon Pen- significant at the 1 percent level (F, ,, = insula). 11.44). Additionally, all southern New The Huon Peninsula specimens differ from 1 984 MC DOWELL: HUON PENINSULA SNAKES I1I

11 Eastern Highlands-Chimbu-Western yellowish. Inside of mouth red. Bluish cast Highlands specimens and from a Mt. Missim to some dorsal scales"; for 66751 (adult fe- specimen (MCZ 44170) in having dark in- male, Lae): "Inside of mouth pink. Under- terstitial skin on the neck, without vertical side anteriorly white, becomes rust brown white bars, thus resembling two Normanby gradually to tip of tail, increasing in intensity Is. specimens. The Huon Peninsula speci- toward tail. Supralabials whitish." During his mens have significantly (at 5% level) more stay at Lae, Kurtz did observe a Dendrelaphis maxillary teeth (34-37, mean = 35.667) than in a banana tree, but did not take the speci- do the 11 Eastern Highlands-Chimbu-West- men, so that its specific identity is uncertain. em Highlands specimens (30-36, mean= His other field notes on Lae D. calligastra do 32.636). The Mt. Missim specimen has 30 not suggest the species is particularly arboreal left maxillary teeth. The Huon Peninsula at Lae: for 66749, "collected swimming in population thus seems sufficiently differen- rain pool from which tadpoles were taken"; tiated to suggest it has been isolated for some for 66751, "marshy field, midday, in grass"; considerable time. for 66755 (an adult male), on the ground in the campsite. This last specimen contains a Dendrelaphis calligastra (Gunther) skink tail in its stomach and BPBM 5136 () contains a skink and two skink USNM 119190, 119504 Gusiko; BPBM 3922 eggs (probably secondarily ingested with the Finschhafen; BPBM 5136 Busu River near skink); Stickel's field tag indicates USNM Lae; 66749, 66751, 66755 Lae; 95566 10 mi. 119504 (adult female, Gusiko) was caught w of Lae eating a frog. Although this is a common species at Lae and has been taken on the eastern coast of the Huon Peninsula, the failure of the Sev- Dendrelaphis punctulatus (Gray) enth Archbold Expedition to collect it in the 95568 Finschhafen; USNM 711482 Gusiko; Mt. Rawlinson region suggests this species is 85673, 95143 Lae; 95567 10 mi. w of Lae confined to the coastal lowlands. I find no These specimens, all females, are typical significant differences (at 5% level) in max- D. punctulatus with conspicuous large scale illary count (22-23 for Gusiko and Finsch- pits clustered on the loreal, postoculars, an- hafen, 23-24 for Lae region) or ventral count terior temporals, and dorsal ends of the su- (173-179 for Gusiko and Finschhafen; 173- pralabials; they lack a temporal streak; they 189 for Lae region but 173-178 omitting also show a characteristic of D. punctulatus specimen from west of Lae), but there may that is probably diagnostic, at least on New be a difference between the Lae region gnd Guinea: the internasal enters the dorsal rim Finschhafen-Gusiko region in the color pat- of the nostril. In 95567, the interstitial skin tern of the interstitial skin. In both regions, shows a pattern ofalternating gray and white the interstitial skin of the neck shows alter- vertical bars (in the preserved specimen), with nating vertical black and white bars, but for the uniformly dark cornified scales standing Lae region specimens, the color of the cor- out as dark spots against this pale pattern; in nified scales is not in register with the pattern 85673, the pattern ofthe interstitial neck skin of the interstitial skin, so that white bases of is similar, except that the white bars arejoined scales contrast with dark interstitial bars (Busu together by a longitudinal white stripe be- River specimen) or dark edging on the scales tween the first and second scale rows; in stands out against the white interstitial bars; 95143, there is a similar white stripe between in the Finschhafen-Gusiko specimens, the the first and second rows, but the gray bars concealed bases of the scales agree with the fuse dorsal to it, to isolate white interstitial color of the interstitial skin, so that the in- spots at the level of the fifth and sixth rows terstitial pattern of vertical bars is not dis- from the white interstitial stripe. In the adult rupted by any coarse stippling provided by Finschhafen specimen, 95568 (the juvenile the cornified scales. Gusiko specimen, USNM 711482, seems Kurtz's field notes give the color in life for similar), the interstitial skin shows black and 66749 (juvenile female, Lae) as "abdomen, white vertical bars, with the cornified scales 12 AMERICAN MUSEUM NOVITATES NO. 2775 in the black bars uniformly dark but the cor- doubtful) BM 1922.11.24.40 (Potter Field No. nified scales in the white bars bearing a white 47) "Huon Gulf" basal spot (but with the dark dorsal edges of Form with 15 scale rows: (pertinence doubt- these scales standing out as blackish spots ful) BM 1922.11.24.41 (Potter collector) against the white interstitial background). "Huon Gulf." I present these notes on interstitial neck McDowell (1972) has given reasons to be- pattern in the hope it will stimulate collectors lieve this species contains both populations to record the color in life of this region. As with 15 scale rows, mainly from northern noted in the outline oftaxonomy ofthe genus, New Guinea, and populations with 17 scale this interstitial pattern is known to be exhib- rows, mainly from southern New Guinea. The ited by neck inflation among species of Den- species is characterized by: (1) having two drelaphis and probably has a function anal- well-developed scale pits on each scale, in- ogous to the dewlap ofAnolis lizards. My own cluding those of the base of the tail (some observations, based on preserved specimens Manus specimens of S. modestus are similar and thus confined to melanin distribution, in this respect, but other Stegonotus either suggest that the interstitial neck pattern dis- lack scale pits entirely or have a few scattered tinguishes the species within a small geo- scales with a tiny pit); (2) pointed and ante- graphic region, as usual for Anolis, but shows riorly directed choanal process of palatine considerable geographic variation within the (diagnostic); (3) a juvenile head pattern species, so that the rules change from place (masked by a general darkening of the head to place as to how the species differ in this in adults, but usually still distinguishable display pattern-again, as in Anolis. In this through close inspection) involving a dark case, the pattern ofthe neck ofD. punctulatus orbital blotch, dark posterior temporal blotch, from the east coast of the Huon Peninsula a dark bar across the top ofthe head between seems similar to that of Lae D. calligastra; the orbits, and dark parietal blotches thatjoin however, around Lae, D. punctulatus has a the interorbital bar. The Lae and Munim pale interstitial pattern against which the dark Waters specimen and one of Potter's "Huon scales stand out as conspicuous spots as the Gulf' specimens have 17 scale rows and are most conspicuous element of the pattern, generally similar to southern New Guinea whereas on the east coast of the Huon Pen- specimens (such as those of Garaina in Mo- insula, D. calligastra shows a concordance of robe Province, and of Northern, Central, scale and interstitial skin color to make the Chimbu, and Milne Bay Provinces) with a inflated neck pattern one ofvertical black and large number of palatopterygoid teeth (39 in white barring without dark spots on the white 95155, 43 in 103672andBM 1922. 11.24.40). bars. I have no information about any dif- This last-mentioned specimen agrees with ferences in movements of the inflated neck specimens from farther south in having a dark by live Dendrelaphis or any precise obser- spot on the lateral end ofeach ventral, as well vations on the degree of neck-inflation of as in each subcaudal being brown anteriorly which the various species are capable. (this specimen may have come from the Huon As with D. calligastra, D. punctulatus ap- Gulf region well south of Lae); in 103672 pears to be confined to low coastal localities (adult male), many of the ventrals, particu- on the Huon Peninsula and shows little dif- larly anteriorly, lack a lateral brown spot and ferentiation between Lae and the extreme thus approach the northern (15 scale rows) eastern end ofthe peninsula. Ofthe four Huon form that has immaculate ventrals; in 95155 Peninsula Dendrelaphis, D. lorentzi and D. (juvenile male) there is a further approach to gastrostictus would appear to be "old inhab- more northern specimens in that the ventrals itants" and D. calligastra and D. punctulatus are immaculate and the subcaudals have seem to be "recent invaders." brown pigment confined to their anterome- dial edges (forming a narrow median zigzag subcaudal stripe). These specimens would Stegonotus diehli Lindholm suggest that the Lae region is part of an in- Form with 17 scale rows: 95155 Lae; 103672 tergrade zone between the southern and Munim Waters (12 mi. w ofLae); (pertinence northern forms of the species. 1 984 MC DOWELL: HUON PENINSULA SNAKES 13

However, BM 1926.5.31.16, labeled as other, as well as a significant (at 1% level) sex "Huon Gulf," Potter collector, is a typical dimorphism for both populations, with males northern specimen, indistinguishable from expected to average 4.72 more ventrals than Astrolabe Bay (Madang Province) material, do females for the same locality. For Pindiu with but 28 palatopterygoid teeth, 15 scale and Mt. Rawlinson, ventrals average 193.00 rows, immaculate ventrals, and a median dark for three males, 189.50 for two females; for zigzag on the pale gray subcaudals. If this the Lae region, ventrals average 181.50 for specimen really came from the same locality four males, 176.17 for six females. In addi- as Potter's "Huon Gulf' specimen with 17 tion, all nine Lae region specimens for which scale rows, then it would indicate that the 15 the head scutes are countable show (bilater- and 17-scaled forms are probably good ally) only one anterior temporal meeting the species, occurring sympatrically without ev- postoculars. Ofthe five Pindiu and Mt. Raw- ident introgression. But there are two other linson specimens, only the left side is count- possibilities. able in 95573 and shows two anterior tem- One possibility is that the 15-scaled spec- porals meeting the postoculars; of the other imen came from the northern coast of the four, 95574 shows two temporals meeting the Huon Gulf-that is, the Huon Peninsula. Al- postoculars bilaterally and the remaining though neither the Seventh Archbold Expe- three show two temporals meeting the post- dition nor any other collector known to me oculars on the left side and one (the more has taken this species on the peninsula east dorsal) anterior temporal meeting the post- of Lae, the typical northern form is known ocular on the right side. There is a slight, but from Saidor (MCZ 49491), near the same significant (at the 5% level) difference in num- mountain range that forms the central ele- ber of maxillary teeth: for the Lae region, vation of the Huon Peninsula; it would not observed range 15-16, mean = 15.78, s = be implausible for any snake occurring at Sai- 0.441, s,, = 0.147 (N = 9); for Pindiu and Mt. dor to extend eastward onto the Huon Pen- Rawlinson, observed range 16-17, mean = insula (as here, somewhat arbitrarily, de- 16.40, s = 0.548, s, = 0.245 (N = 5). There fined). is a similarly slight, but significant (at the 5% Another possibility is that, somehow, data level), difference in the number of dentary on Potter's specimens got transposed. The teeth: for the Lae region, observed range 18- 17-scaled "Huon Gulf' specimen bears Pot- 21, mean = 18.78, s = 0.667, s, = 0.222 ter's field number 47. But the snake bear- (N = 9) for Pindiu and Mt. Rawlinson, ob- ing Potter's field number 48 is a 15-scaled served range 19-21, mean = 19.80, s = 0.837, Stegonotus diehli from the " River del- s, = 0.374 (N = 5). There is no significant dif- ta" (BM 1926.5.31.16) suggesting that Potter ference in palatine (observed range 14-16, went to the Astrolabe Bay coast from the mean = 15.28, s = 0.726, s, = 0.194, N = 14) Huon Gulfregion just after collecting the 17- or pterygoid (observed range 23-31, mean = scaled S. diehli. The 15-scaled specimen may, 26.14, s = 2.070, s. = 0.553, N = 14) tooth then, have been collected on the Astrolabe counts, in head scute counts, or in coloration. Bay coast and somehow been mingled with I can find no differences between two Kalolo the Huon Gulf collection. specimens and the Lae region series. McDowell (1972) suggested that this species Stegonotus parvus (A. B. Mayer) may have originated as an eastern variation 95571-74 Pindiu, about 3000 ft.; 95576 Mt. of S. modestus on the Huon Peninsula, at a Rawlinson; BPBM 5439, BPBM 5441 Ka- time when the Huon Peninsula formed an lolo; 95577 Butibum River near Lae; 66750, island. Stegonotus modestus is not known 66752, 85721-23, 95154, BM 1922.11.24. from east ofAstrolabe Bay on the New Guinea 38, BM 1967.595 Lae; 95156 12 mi. w, 4 mi. mainland, but S. parvus extends westward n of Lae into Madang, East and West Provinces There is a significant (at 1% level) differ- and northern Irian Jaya to be sympatric with ence in ventral count between specimens from S. modestus. Where sympatric with S. mo- Pindiu and Mt. Rawlinson on the one hand destus, S. parvus has eight supralabial scutes, and specimens from the Lae region on the a ventral count below 190, and 17-19 max- 14 AMERICAN MUSEUM NOVITATES NO. 2775 illary teeth, thus contrasting strongly with S. then, that the Pindiu-Mt. Rawlinson region modestus from the same region, which there (five S. parvus, one S. cucullatus), in spite of has seven supralabials, 200 or more ventrals, its elevation, is more like the Markham Val- and 13 or 14 maxillary teeth. East ofthe range ley than like the northeastern highlands in of S. modestus, in the Markham Valley, Ka- the relative abundance of the species of Ste- lolo, around Lae, and in the Wau region, S. gonotus. parvus has seven (occasionally eight, but rare- All the Huon Peninsula specimens of S. ly bilaterally) supralabials, 173-187 ventrals, parvus have the usual coloration for this and 14-17 (modally, 16) maxillary teeth; far- species (and S. modestus); dark brown ver- ther east, in the central Huon Peninsula there tebrally, fading through a warm brown dor- are seven supralabials and 16 or 17 maxillary solaterally to a warm tan ventrolaterally (the teeth, but ventral count rises to 187-196 ventrolateral scales distinctly paler on their (overlapping the ventral count for Moluccan edges than on their centers), with clear whit- specimens of S. modestus). Although Mc- ish ventral surface, including the lower side Dowell (1972) states that the hemipenis of S. of the tail, and upper lip. Most stomachs ex- parvus is similar to that of S. diehli and S. amined from the Huon Peninsula series were modestus, there seems to be a hemipenial dif- empty, but 95154 (Lae) contained a skink ference separating S. modestus from other and two squamatan eggs; 95156 (west ofLae) Stegonotus (except for S. heterurus): the dis- contained two squamatan eggs; 95573 (Pin- tal one or two transverse tiers ofcalyces have diu) also contained squamatan eggs but 95574 smooth edges and are readily identified as contained a few scincid scales and an un- calyces; in the others, the distal calyces have identifiable material. spinulose rims and the entire texture of the organ appears spinose, with only a slight bas- al webbing of the more distal spines indicat- Stegonotus cucullatus ing the existence of a calyculate zone. (Dumeril, Bibron, and Dumeril) Stegonotus parvus seems to be the most 95575 Pindiu, 3000 ft.; 95157 13 mi. nw of common species of its genus on the Huon Lae; BM 1922.11.24.39 "Munum, 17 mi. Peninsula, lower Markham Valley and south- from mouth of Markham River" [=Munim ward to the Wau-Bulolo region (of 11 BPBM Waters?] specimens from the Wau region, 10 are S. Of the localities listed above, only Pindiu parvus and one is S. cucullatus). Farther south, conforms strictly to the definition of "Huon at Garaina, the species must be rare if not Peninsula" used here; the specimens from absent (17 AMNH Stegonotus from Garaina northwest of Lae and from "Munum" are comprise 16 S. cucullatus and one S. diehhl). both males, with ventrals 194 and 191, re- In the Astrolabe Bay region (24 specimens), spectively; these are unusually low counts for S. parvus (six) seems more abundant than S. this species, but matched by 85715, a female modestus (three), but less common than S. from Gurakor (farther up the Markham Val- diehli (15) and this relative abundance prob- ley) with 191, and by BPBM 4160, a juvenile ably holds for the northern edge of New male from Wau, also with 191 ventrals. The Guinea from Toem, Irian Jaya, to Wewak (of Pindiu specimen is an adult male with 200 27 Stegonotus from this region, 13 are S. ventrals, also low for this species but in fair diehli, six are S. parvus, and four each are S. agreement with 85713-14, two males from modestus and S. cucullatus). In the highlands Umi River, upper Markham Valley, with 198 of northeastern New Guinea other than the and 197 ventrals, respectively. Although these Huon Peninsula-that is, Eastern Highlands, counts are low for S. cucullatus, they are Western Highlands, and Chimbu Provinces higher than those for S. parvus from the cor- and Morobe Province west of the Ramu- responding geographic region and it is a rule Markham Valley-the most common Ste- to which I do not know any exception that gonotus is S. cucullatus (19 of 26 specimens), where S. cucullatus and S. parvus occur to- with S. parvus (six) and S. diehli (one) con- gether, the former has a higher ventral count, spicuously less abundant. It would appear, although the geographic variation in each 1 984 MC DOWELL: HUON PENINSULA SNAKES 15

species produces an overlap in ventral counts by a dog. The stomach contains the thor- for the two species when locality is not con- oughly digested remains of a murid rodent. sidered. The color ofthese specimens is as for other Morobe Province (and also Kokoda and GENUS STYPORHYNCHUS PETERS Eastern Highlands) specimens seen by me: Malnate (1975) has given a preliminary pa- cool dark smoky gray with distinct dark edg- per referring the Moluccan and New Guinea ing to the scales dorsal to the second row, the (and some Philippine) species of "Amphies- subcaudals with distinct dark edging, the ven- ma" to a separate genus Styporhynchus, char- trals with dark smudges above the angula- acterized, in part, by presence of scale pits on tion, the supralabials light gray with darker the subcaudals. Malnate and Underwood are gray borders. As with other Morobe speci- now engaged in a revision of this difficult mens, the darker edging of the dorsal scales genus and the following discussion of the is more conspicuous after long preservation Huon Peninsula forms is quite tentative; more in alcohol (the dark edging is now quite ev- definitive statements about New Guinea Sty- ident when the Pindiu specimen is immersed porhynchus must await Malnate and Under- in alcohol, but it was not at all evident when wood's revision. the specimen was unpacked more than 15 The Huon Peninsula Styporhynchus be- years previously). long to a complex within that genus contain- The sagittal crest on the parietal ofthe Pin- ing S. montanus, S. multiscutellatus, and S. diu specimen is equal to the sagittal crest on mairi; the last two are usually considered the supraoccipital, as usual for adult S. cu- conspecific, but S. mairi appears to co-exist cullatus; in S. parvus the sagittal crest is con- with S. multiscutellatus in the Oriomo Pla- fined to the supraoccipital and the parietal teau (essentially, Merauke to re- has a flat shield-shaped dorsal exposure be- gion) and Port Moresby region ofNew Guinea tween the jaw adductor muscles. The Pindiu without indications of introgression. Al- S. cucullatus is also a considerably larger though S. mairi frequently has two preocu- snake (snout-vent 1200 mm.) than S. parvus lars in Australia (where it is the only species (95571, Pindiu, the largest specimen of S. ofthe genus), New Guinea S. mairi are char- parvus known to me, has a snout-vent length acterized by a single preocular. It may be of 800 mm.); the Pindiu S. cucullatus would added that there is no evidence to indicate be only a "medium size" specimen for this that S. mairi is more closely related to S. species, which, along with Boiga irregularis multiscutellatus than it is to S. montanus. and Dendrelaphis punctulatus, comprises the Over most of New Guinea, except the el- "large colubrid" fauna of New Guinea. The evated interior, there are two members ofthe tooth counts (left side) for the Pindiu S. cu- S. mairi complex: a form with a lower ventral cullatus are: maxillary 14 + 3; palatine 17; count and preferring lower elevations (S. pterygoid 26; dentary 19 (gradual transitions multiscutellatus); and a form with higher ven- in length for dentary teeth and pre-diastemal tral count, preferring higher elevations and maxillary teeth); these counts are all within also occurring in the elevated interior of the the observed range of Eastern Highlands and island (S. montanus). Discriminating be- western Morobe Province specimens. The tween these species is not always easy, be- supralabial count of nine (fourth and fifth cause both show geographic variation and the entering eye) ofthe Pindiu specimen is modal critical value of ventral count distinguishing for Eastern Highlands-western Morobe. the two taxa varies from place to place; fur- Nothing, indeed, about the Pindiu S. cucul- thermore, the identification characteristics latus suggests even an endemic race and the additional to ventral count also change from species may well be a recent immigrant to the place to place (e.g., the feebleness of keeling Huon Peninsula, most probably from the on the first scale row that distinguishes S. eastern highlands region by way of the upper montanus in Irian Jaya is useless as a crite- Markham Valley. rion in Papua New Guinea). The following The Pindiu specimen was caught in grass is my own attempt to determine the best cri- 16 AMERICAN MUSEUM NOVITATES NO. 2775 teria for identifying material from the general The two species differ in subcaudal count, region of the Huon Peninsula. but the difference is not uniform throughout To the west ofthe Ramu-Markham Valley, the range from Aiyura to the Wau region. in the Goroka region of Eastern Highlands Neither species shows significant (at the 5% Province, there appears to be but one species, level) sexual dimorphism, but in both species S. montanus, represented in the 22 speci- the geographic variation is significant (at the mens available to me; ventrals show a sig- 5% level in the case of S. montanus, at the nificant (5% level) sex dimorphism and are: 1% level in the case of S. multiscutellatus) for males, observed range 160-171, mean = and the geographic trend is opposite in the 164.571, s = 3.345, s, = 0.894, N = 14; for two species; in S. montanus the subcaudal females, observed range 156-165, mean= count is highest in the Aiyura-Kainantu re- 160.623, s = 3.114, s, = 1.101, N = 8; sub- gion (where not significantly different, at the caudals show no significant sex dimorphism, 5% level, from the Goroka region sample) with observed range 75-89, mean = 81.526, and declines in the Kratke Mts., to decline s = 4.575, s, = 1.050, N = 19; left maxillary still further in the Wau-Bulolo region; but S. teeth show an observed range of 31-36, multiscutellatus shows highest counts in the mean = 34.143, s = 1.711, s, = 0.373, N = Wau-Bulolo region, with a decline in the 21. There is no significant correlation be- Kratke Mts. and lowest counts in Aiyura- tween subcaudals and ventrals, either within Kainantu region, where the mean for S. mul- each sex or when data for the sexes are com- tiscutellatus is below that for S. montanus, bined. although S. multiscutellatus has conspicu- When 30 specimens from farther east, on ously higher subcaudal counts than does S. the western-southern shoulder ofthe Ramu- montanus in the Wau-Bulolo region. Table Markham Valley (more precisely, 12 from 1 gives the parameter estimates. the Aiyura-Kainantu region, eight from the No significant (at the 5% level) correlation Kratke Mts., and 10 from the Bulolo-Wau between ventral and subcaudal count is de- region) are considered, it is evident that this monstrable in S. multiscutellatus in a sample sample (and each of the three component of nine from Aiyura-Kainantu, the Kratke subsamples) shows a marked bimodality in Mts., and the Bulolo-Wau region; but for 14 ventral count, with the two clusters of each S. montanus from these regions, r = -0.5985 subsample separated by a gap ofnine or more (significant at the 5% level), yielding the ventrals for which there are no observed regression equation: estimated subcaudals = counts. The clusters with higher ventral count 262.9 - 1.123 ventrals. That the slope ofthe in each subsample are not significantly (at the regression line approximates -1.0 suggests 5% level) different from one another or from that total somites (about equal to ventrals + the Goroka sample and are evidently Sty- subcaudals) approximate a constant and that porhynchus montanus; when ventral counts decrease in the number of subcaudals leads for Goroka, Aiyura-Kainantu, Kratke Mts., to a corresponding increase in the number of and Bulolo-Wau are combined, parameter ventrals (but note that while the number of estimates are: for males, observed range 160- subcaudals for Bulolo-Wau S. montanus is 171, mean = 164.769, s = 3.037, s. = 0.596, significantly lower than the mean for Goroka, N = 26; for females, observed range 156-165, there is no significant difference in ventral mean = 161.462, s = 2.847, s, = 0.789, N = count; perhaps this can be ascribed to the 13. rather high variance for ventrals in the Go- The clusters with lower ventral count would roka sample). The above regression line does seem to represent S. multiscutellatus. There not differ significantly from: estimated sub- is no significant (at 5% level) geographic dif- caudals = 242.643 - 1.000 ventrals (based ference or sexual difference in ventral counts on the mean = 242.643 for sum of ventrals for these clusters and the parameter estimates and subcaudals in this sample). In the Wau- for the combined samples are: observed range Bulolo region, the total of ventrals and sub- 145-151, mean = 148.923, s = 1.977, s= caudals for S. multiscutellatus is not diag- 0.548, N = 13. nostically different from that for S. monta- 1984 MC DOWELL: HUON PENINSULA SNAKES 17

TABLE 1 Subcaudal counts for Styporhynchus montanus and Styporhynchus multiscutellatus West and South of the Markham Valley Styporhynchus montanus Styporhynchus multiscutellatus Aiyura-Kainantu (Sample of six not distin- region guishable from 19 Goroka region specimens; combined estimates given) Observed range 75-89 Observed range 75-80 Mean = 81.440 Mean = 76.667 s = 4.263; s, = 0.853; N = 25 s = 2.887; s, = 1.667; N = 3 Kratke Mts. Observed range 72-82 Observed range 84-86 Mean = 77.000 Mean = 85.000 s = 4.041; s, = 2.021; N = 4 s = 1.414; s, = 1.000; N = 2 Bulolo-Wau Observed range 67-80 Observed range 88-96 region Mean = 74.000 Mean = 91.800 s= 5.477; s, = 2.739; N = 4 s= 2.950; s, = 1.319; N = 5 nus, but the two species differ conspicuously behind the angle ofthe jaw (a remnant ofthe in subcaudal count; farther to the northwest, posterior border of the collar). In S. mon- in the Aiyura-Kainantu region, subcaudal tanus no such marks appear, and the post- count is no longer diagnostic, but total of ocular stripe, ifnot altogether obscured, fades ventrals and subcaudals probably does dis- out posteriorly on the last supralabial. This tinguish the species (observed range 221-230 character is not completely clearcut, how- for three S. multiscutellatus, 239-249 for six ever, because S. montanus frequently has a S. montanus). checkerboard pattern of black spots and the There is no significant difference between more anterior of such spots may lie in the S. montanus and S. multiscutellatus in the general position of the posterior collar-bor- Aiyura-Wau region in the mean for left max- ders of S. multiscutellatus. illary teeth (33.353 for 17 S. montanus; In the Markham Valley and Huon Penin- 33.545 for 11 S. multiscutellatus), but there sula, ventral counts are bimodal, but the is a significant (at the 1% level) difference in higher-count cluster shows lower values than variance (for S. montanus, s = 0.931, ob- seen in the higher count cluster for the west- served range 32-35; for S. multiscutellatus, ern and southern shoulders of the Markham s = 2.162, observed range 31-37). In the re- Valley. The Huon Peninsula higher-count gion under discussion, the two species are cluster (it does not appear to occur in the most easily distinguished in thejuvenile stage, Markham Valley) has an observed range of which has a pale collar, bordered anteriorly 153-160, with mean= 156.250, s= 1.765, and posteriorly with black, in S. multiscu- s, = 0.509, N = 12. In most respects, it does tellatus but is without any collar in S. mon- not seem to differ from Styporhynchus mon- tanus. Even before sexual maturity, this col- tanus from west of the Markham Valley, but lar darkens to the same brown tint as the back has a reduced somite number, reflected in and crown in S. multiscutellatus, but the dark both the reduced ventral count and in a re- marks that formed anterior and posterior duced subcaudal count (observed range 70- borders for the collar remain, and usually can 78, mean = 74.375, s = 2.825, s, = 0.999, be made out in fully adult specimens, at least N = 8) and, consequently, with a conspicu- when they are wet, showing as a posterior ously lower sum of ventrals plus subcaudals expansion of the black postorbital stripe on (observed range 227-233, mean = 229.875, the temporal region (a remnant of the ante- s = 2.642, s, = 0.934, N = 8; for the Aiyura- rior border of the collar) and as a black mark Kainantu region, Kratke Mts., and Bulolo- 18 AMERICAN MUSEUM NOVITATES NO. 2775

Wau region, observed range is 236-249, multiscutellatus, there is no evidence of re- mean = 242.643, s = 4.03 1, s, = 1.077, N = productive isolation and I know of one spec- 14). Because ofthis reduction, ventral counts imen that appears to be intermediate: BPBM for Huon Peninsula S. montanus fall mainly 3398, an adult female from Tewep, in the in the region of the gap between the lower Sarawaket Range at 1350 m., shows distinct, ventral count cluster (S. multiscutellatus) and but inconspicuous, black spots that appear to higher ventral count cluster (S. montanus) for be the remnants of the borders of a pale ju- samples from west and south ofthe Markham venile collar and has 81 subcaudals. Valley. Nonetheless, the ventral count cluster Judging from a sample of 12 S. montanus for S. montanus is well defined for the Huon and 10 S. multiscutellatus from the Huon Peninsula-Markham Valley sample, because Peninsula (including the Lae region), the zone the lower-count cluster is also displaced to- of 150-152 ventrals lies above mean + 2s for wards lower values (143-149). S. multiscutellatus and below mean - 2s for This lower-count cluster, which seems to S. montanus. In addition, all but one of 12 correspond to the S. multiscutellatus cluster S. montanus from this region have a single of the Aiyura-Kratke-Wau sample, contains anterior temporal, whereas all 10 S. multi- two quite different forms in the Huon Pen- scutellatus have two anterior temporals, at insula-Markham Valley sample. One ofthem, least on one side of the head. In the Wau- from low elevations- around Lae and also Gu- Bulolo region, this difference in anterior tem- siko on the eastern coast of the Huon Pen- porals appears also to be diagnostic, but the insula, has a high subcaudal count (86-90, Goroka sample, apparently representing only mean = 87.250, s = 1.893, s, = 0.946, N = S. montanus, is variable in anterior temporal 4) and a distinct pale collar in the juvenile number and the character will not correctly that leaves traces of its black borders in the identify specimens from the Aiyura-Kai- adult; these snakes do not appear distinguish- nantu region. It should be noted that the di- able from S. multiscutellatus from the Bu- agnostic gap (150-152) in ventral count for lolo-Wau region and probably represent a the Huon Peninsula nearly corresponds to the recent invasion ofthe lower Markham Valley peak of the distribution for S. multiscutel- and coast of the Huon Peninsula from the latus from west and south of the Markham south. Valley. For the western and southern shoul- The other S. multiscutellatus, represented der of the Markham Valley, the diagnostic by specimens from the Rawlinson Range, at gap in ventral count (between mean + 2s for elevations of 1340 to 1580 m., has far fewer S. multiscutellatus and mean - 2s for female subcaudals (67-73, mean = 69.333, s = S. montanus) is 154-155 ventrals. 3.215, s, = 1.856, N = 3) and I can see no trace ofa former collar on the four specimens available. As noted above, there is a decrease Styporhynchus montanus in subcaudal count going from the Wau re- (van Lidth de Jeude) gion northwestward to Aiyura. The Rawlin- 95535, 95539-43, 95545 Gang Creek, Mt. son Range S. multiscutellatus has still lower Rawlinson, 4400-5200 ft.; 95546 Gumun; subcaudal counts than does the Aiyura-Kai- BPBM [Field No.] NG-28337 Kabwum, 7500 nantu sample, but seems to be a continuation ft.; BPBM [Field No.] NG-52763 Kilolo, 6900 of this trend. I interpret these Rawlinson ft.; 95006 Gevak; BPBM 3734 Sarawaket Range specimens as representing an earlier Range, 1920 m. [probably in general region invasion of the Huon Peninsula, probably of Tewep] from the west, now showing some differen- Ofthese specimens, 95541, 95546, 93006, tiation in the form ofreduced subcaudal count and BPBM 3734 contained microhylid frogs and also reduced ventral count (143-146, and 95540 contained a papery material per- mean = 144.500, s = 1.291, s, = 0.645, N = haps representing much digested frog skin; 4), and in obliteration of the collar-at least, BPBM 3734 also contained two beetle larvae. in the adult. Since microhylids are most likely to be en- Although there are conspicuous differences countered on the ground, this supports Zwei- between Lae region and Rawlinson Range S. fel's (personal commun.) impression that 1 984 MC DOWELL: HUON PENINSULA SNAKES 19

Styporhynchus is not particularly aquatic on tail the spots coalesce to form a vertebral line the Huon Peninsula. but this is not present on the body; the skin Zweifel's notes on the color in life of95 536 between the scales shows as a rather greenish are: "dark olive dorsal color, no pattern, skin yellow when the body is stretched; the under- between scales yellowish; dorsal color ex- surfaces are a dark cloudy gray." For 95537 tends onto tips ofventrals, ventrals otherwise (Gang Creek, female), "Rich brown dorsal yellowish; iris yellow-brown." To judge from color, no pattern except thin black stripe on preserved material, this absence of dorsal tail, skin between scales yellowish; pink tint pattern would hold true for another Gang to labials and chin, fading gradually to tan Creek specimen (95545) and the Gumun under tail; ventrals speckled gray; dorsal col- specimen, but the remaining specimens show or extends onto ventrals." For 95538 (Gang more or less conspicuous small black spots, Creek, male), "Dark gray-brown dorsally with separated from one another but in an alter- darker spots less than one scale in area, dark nating checkerboard arrangement. All the S. central line on tail; skin between scales gray- montanus from the Rawlinson Range show ish; ventral surfaces gray with a slight pinkish a cream-colored (in preservative) oral border tint; chin and lower labials white, upper la- for the entire supralabial series, in contrast bials lighter brown than rest of head." The to the S. multiscutellatus from the same re- Gang Creek specimens (95537-38) were col- gion, which have the last two supralabials lected by local residents, who found them brownish gray, like the temporal region; how- during preparation of gardens; the Tumnang ever, this difference will not hold outside the specimen (95150) was found among leaves Rawlinson Range and BPBM 3734, a S. beside a trail in heavy forest. montanus from the Sarawaket Range, has considerable gray suffusion of the last two Boiga irregularis (Merrem) supralabials, while BPBM 3398, a S. multi- scutellatus from Tewep, has the entire upper BPBM 5438, BPBM 5444-45, BPBM 5447 lip cream with dark borders to the suprala- Kalolo, 750 m.; 66671-72 Gusiko; 66992, bials. 95565, BPBM 3920, BPBM 3931 Finsch- hafen; 95548-53 Pindiu 2900-3000 ft.; 95554 Masba Creek, 2500 ft.; 95555-58 Numbut, Styporhynchus multiscutellatus 4500 ft.; 95559,95561 Maran, 4000 ft.; 95560 (Brongersma) Zangaren, 4000 ft.; 95151 Tumnang; BPBM 66670 Gusiko; 95537-38, 95544 Gang Creek, 3278, BPBM 3658 Singaua River (6040'S, Mt. Rawlinson, 5000-5200 ft.; 95150 Tum- 147009'E, 50 m.); 66745, 77648, 66762-64, nang, 4400 ft.; BPBM 3398 Tewep, 1350 m.; 85731-34, 95153, 95562-63 Lae; 95152 66753, 103640 Lae; 85712, 95547 Oomsis Oomsis; 95574 10 mi. w of Lae Creek, 500 ft., 17 mi. sw of Lae Also recorded from Simbang by L6nnberg The specimens from Tewep and Oomsis (1900) and from "Lialun" [=Sialum] by Vogt Creek are from slightly west ofthe Huon Pen- (191 1). insula as defined for this report, but are listed This is probably the most abundant snake because they were used in analysis ofthe Huon in the lowlands of New Guinea and the Sev- sample. The Gusiko specimen is a juvenile enth Archbold Expedition collection would male, with conspicuous black neck marks indicate that it is the commonest snake in the representing the borders of a former collar, Rawlinson Range up to 1220 m.; at higher and with 149 ventrals and 90 subcaudals. It elevations, Styporhynchus montanus seems is quite unlike the Tumnang and Gang Creek to be the most abundant snake. The habitat specimens (subcaudals 67-73, no trace ofcol- of Boiga irregularis seems varied: it was tak- lar) and similar to Lae region specimens. en within the town limits of Lae (95562), in Zweifel made notes on the color in life of a tree (95554), crawling on open ground several Rawlinson Range specimens of S. (66764), on the roofofa grass house (95560), multiscutellatus: 95150 (Tumnang, female), in kunai grass (BPBM 3278), in primary for- "The ground color is a dark greenish gray est (BPBM 3658). Of specimens with recog- overlain with a pattern ofblack spots; on the nizable stomach contents, rats or rat hair is 20 AMERICAN MUSEUM NOVITATES NO. 2775 the most frequent element (66762, 95549- Echimypera sp." and the resident people told 51, 95556, 95560), with 95556 containing an him that bandicoots were the principal food adult Rattus exulans (G. Musser determi- of Acanthophis. nation) but 95560 and 66762 containing Kurtz's field notes further state that 66757- nestling murids; 66763 contains a much-di- 58 were taken together in kunai grass in a gested skink and fin spines of a fish; 85731 low, moist spot; 66759-60 were taken in the contains a gecko tail and 95553 contains bird U.S. Army campsite area at Lae, described eggs. by Kurtz as "swampy grassland, flat terrain." Dividing the Huon Peninsula sample into Acanthophis antarcticus shows consider- four subsamples (Kalolo, Finschhafen-Gu- able geographic variation in New Guinea in siko, Rawlinson Range-Pindiu-Tumnang, ventral count, in color pattern, and in wheth- and Lae region), no significant (at 5% level) er the temporolabial scute (=supralabial 6 of difference in ventral count or subcaudal count a series ofseven supralabials) is excluded from is apparent either between geographic regions the mouth as a "lower temporal" by contact or between sexes (parameters are: ventrals of supralabial 5 with supralabial 7 (giving a 240-255, mean = 246.81, s = 3.429, s, = supralabial count of six) (see van Lidth de 0.572, N = 36; subcaudals 104-120, mean = Jeude, 1911, for discussion of this variation 110.13, s = 3.882, s, = 0.792, N = 24). in Irian Jaya Acanthophis). Lae region Acan- thophis have the temporolabial excluded from FAMILY HYDROPHIIDAE the mouth as a temporal (66759 has the tem- I follow Smith, Smith, and Sawin (1977) porolabial excluded from the mouth by hor- in referring all Australasian proteroglyphs ex- izontal division, so that there are seven su- cept Laticauda and Parapistocalamus to the pralabials, but the others have six Hydrophiidae, which thus includes all the supralabials, with the original seventh in con- "palatine draggers" of McDowell (1970). Al- tact with the fifth); in this respect, as in all though a "typical" sea snake is different in others noted by me, Lae specimens agree with many anatomical features from a "typical" those from the Markham Valley, Kratke terrestrial Australasian proteroglyph, transi- Mountains, and Bulolo region. Ventral count tional genera exist; but it is at present im- for specimens from the Markham Valley (in- possible to ally the terrestrial Australasian cluding Lae), Kratkes, and Bulolo region are: proteroglyphs to any known Asiatic, African, males 119-122, mean= 120.33, s= 1.033, s, = 0.422, N = 6; females 119-126, mean = or American genus or even generic group. 122.33, s = 2.733, s, = 1.116, N = 6. In all, each infralabial and supralabial (and the tem- Acanthophis antarcticus (Shaw) porolabial) bears a large black blotch and all 66757-60 Lae; 85740-41 "few miles west of the scales of the first row (and, on the neck, Lae" the second row) have black centers and the In addition to these Lae records, Acantho- pale ground color ofthe ventrals in much less phis is known from Gusiko (Loveridge, 1948). extensive than the black ventral spotting. I do not know of any records from the ele- Specimens from Eastern Highlands Province vated interior of the Huon Peninsula. (exclusive of the Kratke Mountains) have Ofthe Lae specimens examined, 66757 and much lower ventral counts (109 in single male 66760 contain hair in the stomach, similar seen; for females, 111-116, mean = 112.60, to Rattus hair; 66758 contains digested frag- s = 1.949, s, = 0.872, N = 5) and have the ments of Rattus exulans and 66759 contains temporolabial broadly entering the edge of a Rattus exulans skull (G. Musser determi- the mouth; the single male seen (Univ. Col- nation). Kurtz's field notes state that he re- orado 51625, Kamaliki, 8 mi. e of Goroka, moved juvenile bandicoots of the genus 5000 ft.) has black spotting on all the labial Echimypera from 66757-59; probably Kurtz scutes and scales of the first row, but 98494 misidentified these much-digested and frag- (Nivi) and 101146-49 (Dunantine River, mentary stomach contents in the field, since 5000 ft.) have black spots restricted to the 66757 was taken "near nest of bandicoot- anterior infralabials and to the sixth [=tem- 1 984 MC DOWELL: HUON PENINSULA SNAKES 21 porolabial] and seventh supralabials, with ciation has occurred, but changes are so abrupt pale ground color predominating over dark where strongly differentiated forms approach spots on the ventrals and the first scale row one another geographically that some sort of with few or no black spots. reproductive isolation is suggested, but with- The geographic pattern in New Guinea is out the ecological differentiation that would not a simple one. Baiyer River (Western permit two species to occur together without Highlands Province) specimens are similar direct competition (that is, the pattern sug- to Markham Valley specimens, as are Sepik gests biological species constituting a super- lowlands specimens. But specimens from species). Storr assigned the Western Austra- Central and Western Provinces are most like lian death adders to three distinct species: Eastern Highlands specimens, as are most Acanthophis pyrrhus Boulenger (a wholly from Irian Jaya (except for those from near Australian form), A. antarcticus (Shaw) (at former Hollandia and,fide van Lidth de Jeude least strongly analogous to the "highland" [1911], from Merauke). The nature of the New Guinea form), and A. praelongus Ram- temporolabial (whether a supralabial or a say (at least strongly analogous to the "low- temporal) is variable within some popula- land" New Guinea form). Although the anal- tions (e.g., Kubuna, Central Province; Wip- ogy with New Guinea is suggestive and it is im, Western Province; Bernhard Camp, tempting to identify the Huon Peninsula re- Idenburg River, Irian Jaya) and when the en- gion death adders as A. praelongus, the anal- tire series ofNew Guinea Acanthophis avail- ogy is not exact. For example, in Western able to me (20 males, 42 females) is plotted Australia, A. praelongus shows a greater ten- for ventral count it does not fall into two dency than does A. antarcticus to raise the discrete clusters; nor is there a complete cor- supraocular scute into a horn; but in New relation of color pattern with ventral count Guinea, it is the antarcticus-like form that or with condition ofthe temporolabial. There usually has a supraocular horn and the su- does not seem to be any real evidence that praocular is flat in the praelongus-like form. more than a single species is involved, but For the present, it seems best to call New the geographic variation within this species Guinea death adders A. antarcticus but to does not form a simple cline or easily mapped note a real problem as to how many species pattern. Perhaps certain geographic varia- exist in New Guinea and which ones (if any) tions are better adapted than others to higher are represented in Australia. altitudes and most lowland localities in east- ern New Guinea, including Lae, have been occupied by a form that is closely similar to Micropechis ikaheka (Lesson) eastern Queensland Acanthophis, but higher BPBM 5548 Kalolo, 750 m.; 95585 Pindiu, elevations have been occupied by a form with 2800 ft.; 95586 Kotkin, Mt. Rawlinson; reduced ventral count, reduced black pig- 95587 Butibum R., 7 mi. n of Lae; 66754, mentation, and with the temporolabial en- 95588 Lae; 85746 3 mi. nw of Lae; 85757 tering the mouth. This latter form has also Oomis Creek, w of Lae succeeded in entering lowland localities in Also recorded from Sattelberg by Vogt Western Province, as well as in parts of Irian (1911), and from Gusiko by Loveridge (1948). Jaya, producing a situation unknown to me Of the specimens examined by me, 95588 for any other snake known from both New contains an intact Sphenomorphus jobiensis Guinea and Australia: Oriomo Plateau (R. G. Zweifel determination) and 85746 and (southern Western Province) specimens are 95585 each contain a scincid tail. Although less similar to Queensland specimens than 85746 was taken in a clearing in rain forest are those from northeastern New Guinea. and 85757 was taken in secondary rain forest, Storr (1981) has revised the Acanthophis 95588 was taken in tall grass within the town of Western Australia, where the problem limits of Lae and 95586 was taken in a gar- seems closely comparable to that in New den. Guinea: sympatry without introgression can- The Pindiu, Mt. Rawlinson, and Lae re- not be demonstrated anywhere to prove spe- gion specimens show the pattern usual for 22 AMERICAN MUSEUM NOVITATES NO. 2775 this species in Morobe Province: head dark Aspidomorphus lineaticollis (Werner) above and laterally, with the tan anterior neck BPBM 5446 Kalolo, 750 m.; 95584 Masba showing large black spots anteriorly, but these Creek; 95583 Busu River, 12 mi. n of Lae. spots breaking up more posteriorly into scat- This species has frequently been confused tered dark scales and the body essentially pat- with A. muelleri (see McDowell, 1967) and ternless tan one-third of the way back from probably Vogt's (1911) record of"Pseudelaps the head; posterior body with 12 to 24 black- muelleri" from Sattelberg was based on A. edged russet crossbands, these bands becom- lineaticollis. This is probably the only species ing darker posteriorly and the tail with eight of its genus in the Huon Peninsula region. to 12 black rings. The Kalolo specimen shows tail a pattern more like that of Sepik Provinces The stomach of 95583 contains a skink specimens: every scale with tan edge and dark and that of95584 contains the head and other center anteriorly, but slight variations in the fragments of a Sphenomorphus (R. G. Zwei- width of the pale edging from scale to scale fel determination). give a faint suggestion ofspots on the anterior neck, becoming a speckling of scattered dark [MARINE HYDROPHIIDAE] scales against a tan ground in the midbody I have not found any records of sea snakes region; posteriorly, the scales mostly dark, from the coast ofthe Huon Peninsula; almost with a hint of about 20 crossbands (very certainly, this reflects the failure of herpetol- poorly set off from the intervening spaces) ogists to collect in the sea, rather than the and the tail with 10 black rings. absence of sea snakes from the Huon coast. The Pindiu specimen is unusual in having four (rather than three) solid maxillary teeth BIOGEOGRAPHY on each side. The geological history of the Huon Pen- insula and the faunal relationships of its liz- Toxicocalamus (Apistocalamus) loriae ards and frogs have been summarized by (Boulenger) Zweifel (1980). Analysis of the snake fauna BM 1946.1.17.57 [cotype of Pseudapistoca- of the Huon Peninsula gives results similar lamus nymani L[nnberg] probably Sattel- to those of Zweifel's analysis of the lizards. berg; MCZ 76627-28 Sattelberg; 95579-80 Table 2 lists the 24 snakes known for the Masba Creek 2300-2500 ft.; 95578, 95582 peninsula, dividing the records into those for Pindiu, 2600 ft.; 95581 Mt. Rawlinson, 4400 the low coastal region at the eastern end, those ft. for the low region around Lae, and those from Also recorded from "Lialun" [=Sialum] by the higher elevations of the interior of the Vogt (1911). Huon Peninsula. Except for 95578, with a plant rootlet and This list is almost a list of the common unidentifiable matter, the stomach is empty snakes of New Guinea. There is a striking in my specimens. McDowell (1969) has pre- lack of endemism at the specific level and sented the evidence for considering the Huon only one case, Dendrelaphis lorentzi, repre- Peninsula population ("Pseudapistocalamus sents a disjunction in distribution of consid- nymani") conspecific with Toxicocalamus erable magnitude. loriae, but it should be noted that the Huon Some of the species occurring in the "ele- Peninsula population is distinguishable by a vated interior" of the Huon Peninsula show lower mean number ofventrals, although the some incipient differentiation: Stegonotus observed range overlaps that of other eastern parvus from the interior of the peninsula dif- New Guinea populations. Ventrals of the fers from other populations of the species, Huon Peninsula sample are: males, 178-186, including the Lae population, in having a mean = 181.75, s = 4.35, s, = 2.17, N = 4; higher ventral count; Styporhynchus mon- females, 190-201, mean = 194.75, s = 5.62, tanus and S. multiscutellatus from the inte- s, = 3.25, N = 4. The largest specimen in the rior of the peninsula both differ from the Huon sample (95579) has a snout-vent length nearest populations oftheir respective species of 393 mm., tail 81 mm. in lower ventral count (interior Huon Pen- 1 984 MC DOWELL: HUON PENINSULA SNAKES 23

TABLE 2 Snakes Recorded from the Huon Peninsulaa Eastern Elevated Lowlands Lae Interior TYPHLOPIDAE Typhlops inornatus Boulenger - X Typhlops depressiceps Sternfeld X - Ramphotyphlops erycinus (Werner) X X - Ramphotyphlops braminus (Daudin) X - BOIDAE Liasis papuanus Peters and Doria X - Liasis albertisii Peters and Doria X X - Python amethistinus (Schneider) X X Chondropython viridis (Schlegel) X X Candoia carinata (Schneider) X X X Candoia aspera (Gunther) X X X COLUBRIDAE Dendrelaphis lorentzi (van Lidth de Jeude) X Dendrelaphis gastrostictus Boulenger X Dendrelaphis calligastra (Gunther) X X Dendrelaphis punctulatus (Gray) X X Stegonotus diehli Lindholm - X Stegonotus parvus (A. B. Meyer) - X X Stegonotus cucullatus (Dumeril, Bibron, and Dumeril) - X X Styporhynchus montanus (van Lidth de Jeude) - - X Styporhynchus multiscutellatus (Brongersma) X X X Boiga irregularis (Merrem) X X X HYDROPHIIDAE Acanthophis antarcticus (Shaw) X X Micropechis ikaheka (Lesson) X X X Toxicocalamus loriae (Boulenger) X _ X Aspidomorphis lineaticollis (Werner) - X X a Authority for records given in the Systematic Accounts. "Eastern Lowlands" refers to coastal records from the eastern end of the peninsula, mainly Gusiko and Finschhafen; "Lae" refers to the low elevations at or near Lae; "Elevated Interior" includes records from the Rawlinson range, Sattelberg, Kalolo, and various other localities at over 1000 ft.

insula S. multiscutellatus appears to differ Britain; see McDowell, 1975); perhaps the further from adjacent populations in lacking absence of P. boeleni, which is a high altitude the juvenile pale collar); Toxicocalamus lor- species in eastern New Guinea and (at least) iae from the interior of the Huon Peninsula the Wissel Lakes region ofIrian Jaya, permits has an unusually low ventral count; Dendre- P. amethistinus to occupy the montane re- laphis gastrostictus from the interior of the gion of the Huon Peninsula, but it is not so Huon Peninsula appears to be more like easy to explain why there are no records for d'Entrecasteaux Archipelago specimens than the low coastal region ofthe Huon Peninsula, like Eastern Highlands forms in coloration of except at Lae, since P. amethistinus is one of the interstitial skin of the neck. One species, the commonest lowland snakes elsewhere in Python amethistinus, known from 5200 ft. New Guinea. on Mt. Rawlinson, is a lowland form over' Although there are no species of snake en- most of its range in New Guinea and Aus- demic to the Huon Peninsula (revision of tralia (but there are montane records for New Styporhynchus by Malnate and Underwood 24 AMERICAN MUSEUM NOVITATES NO. 2775

may change this), study of the list of "ele- Archipelago-or even the fauna of Rooke or vated interior" forms shows that this list con- Umboi Island, lying between New Britain and tains two main components: (1) the ubiqui- the Huon Peninsula. Zweifel (1980) found tous forms (Candoia carinata, C. aspera, few special resemblances between New Brit- Boiga irregularis) that can be expected in any ain and the Huon Peninsula in the frog and New Guinea region collection of snakes; and lizard faunas, and I cannot find any snake (2) characteristic endemic (or nearly endem- species at all showing a Huon Peninsula-Bis- ic) snakes of New Guinea (Typhlops inor- marck Archipelago distribution pattern. The natus, Chondropython viridis, Stegonotus differences between my findings with snakes parvus, Dendrelaphis gastrostictus, Stypo- and Zweifel's findings with lizards and frogs rhynchus montanus, S. multiscutellatus, Mi- probably represents a difference in sample cropechis ikaheka, Toxicocalamus loriae, size: Zweifel listed 90 species of lizards and Aspidomorphus lineaticollis). I suspect that frogs for the Huon Peninsula (as here de- this assemblage represents the "old" New fined), whereas I list only 24 snakes. Thus Guinea snake fauna, probably the early in- Zweifel could note that although there were vaders ofthe Huon Peninsula when it was an no shared highland forms offrogs and lizards, island. As noted in the systematic accounts two lowland species ofLitoria (abundant and ofStyporhynchus multiscutellatus and Stego- widespread in the New Guinea region) and notus parvus, the elevated interior form is perhaps a Rana (taxonomy uncertain be- differentiated (but not, it would appear, at cause of sibling species) were shared, along the species level) from the lowland form of with many lowland species oflizards (mostly the Lae region; in these cases, we are perhaps widespread) but no highland forms. With the dealing with "old invaders" ofthe Huon Pen- snakes, there is remarkably little resemblance insula, when it was an island, that have come even when ubiquitous lowland forms are in- into contact with conspecific mainland pop- cluded. Python amethistinus, Candoia as- ulations when the Huon Peninsula became pera, C. carinata, Dendrelaphis calligastra, joined to northeastern New Guinea by the D. punctulata, and Boiga irregularis are emergence of the Markham Valley lowlands. shared (on New Britain and the Huon Pen- Python amethistinus might also belong to this insula, P. amethistinus is a highland form, "old snake fauna" of the Huon Peninsula. but elsewhere, over a broad range, it is a low- The "new snake fauna" would probably land species). Even this list may be too long, include Typhlops depressiceps, Ramphotyph- for the Candoia carinata of the Huon Pen- lops erycinus (and R. braminus, ifthat species insula is a "short-tail" (sense of McDowell, is not too recent to warrant consideration), 1979), except in the Lae region, where both Liasis papuanus and L. albertisii, Dendrela- "long-tails" and "short-tails" occur, but New phis calligastra and D. punctulatus, and Britain C. carinata are "long-tails." At pres- Acanthophis antarcticus. These species ap- ent, evidence is insufficient to prove that pear not to have penetrated the interior but "long-tails" and "short-tails" are reproduc- are confined to the most newly formed land, tively isolated and for general taxonomic pur- along the coastal lowlands. One apparently poses, the burden of proof is on those who coastal form, Dendrelaphis lorentzi, is so far would assert a reproductive isolation exists; isolated from other populations ofits species, but for the zoogeographic argument, it must and so much differentiated, that it seems like- be remembered that the "long-tails" of New ly that it is a member of the "old fauna" Britain may not be conspecific with the rather than the "new fauna." It is impossible "short-tails" of the Huon Peninsula. to guess whether Boiga irregularis, Candoia More striking are the differences between carinata, and C. aspera are very successful the Huon Peninsula and New Britain that members of the "new fauna" that have suc- would not be expected by any hypothesis of ceeded in penetrating the interior or are "old either over-water dispersal or former land fauna" elements. connection. Thus, the Aspidomorphus ofNew The snake fauna of the Huon Peninsula is Britain (there, the only terrestrial hydrophiid) strikingly unlike the fauna of the Bismarck is A. muelleri, a widespread species occurring 1984 MC DOWELL: HUON PENINSULA SNAKES 25 in the Moluccas as well as over most of New but with the attenuate form of S. hypomelas Guinea except for the Huon Peninsula, where and probably the basis for Sternfeld's [ 19 13] the genus is represented by A. lineaticollis; erroneous record of the latter from New this latter is also the only species ofthe genus Guinea). on the islands (other than the Bismarck Ar- The snake fauna of Umboi Island might chipelago) east of New Guinea and occurs be expected to be a mixture of New Britain sympatrically with A. muelleri on the Papuan and Huon Peninsula elements, favoring the peninsula. Perhaps the resources ofthe Huon latter (because of the larger species pool); it Peninsula will not support two species ofAs- is not; it is almost purely a New Britain fauna, pidomorphus and A. lineaticollis has exclud- but with a "short-tail" Candoia carinata (as ed (or exterminated, through competition) A. for the Iluon Peninsula) rather than a "long- muelleri, but I see no easy explanation for the tail" (as for New Britain). Combining my own presence on New Britain of the species of identifications of Umboi material with He- Aspidomorphus missing from the nearest diger's (1934), the snakes known from Um- mainland and islands. It may be noted that boi Island are: Ramphotyphlops subocularis the major pattern Zweifel found among New (New Britain also), Liasis boa (New Britain Britain frogs, special affinity with those ofthe also), Python amethistinus (species common Solomon Islands, does not appear among the to both New Britain and the Huon Peninsula, snakes except for one typhlopid (Rampho- but Umboi population highly peculiar in head typhlops subocularis), also known from the pattern and representing endemic geographic tip ofthe Papuan peninsula (MCZ R- 145955, variation), Candoia carinata (more like Huon Alotau, Milne Bay Province); among terres- Peninsula than like New Britain in being trial proteroglyphs, there is no suggestion of "short-tail"), C. aspera (common to both New this pattern, since the Solomons genera (Pa- Britain and Huon Peninsula), Styporhynchus rapistocalamus, Salomonelaps, Loveridge- hypomelas (New Britain also), Boiga irreg- laps) do not occur outside the Solomons and ularis (common to both New Britain and Aspidomorphus does not extend to the Sol- Huon Peninsula). omons. Although Zweifel's observations on the Liasis albertisii ofthe Huon Peninsula (and frogs and lizards and my own on snakes are most of New Guinea) is represented on New in close agreement so far as the lack of sim- Britain (and Umboi and New Ireland) by the ilarity between the Huon Peninsula and New closely related, but quite distinct, L. boa; Britain are concerned, Koopman (1979 and however, L. albertisii, rather than L. boa, personal commun.) found no such striking represents this superspecies complex on dissimilarity in the distribution of mammals Mussau I., the part of the Bismarck Archi- (mostly bats) and his (1979) analysis of the pelago farthest from the New Guinea main- small islands northeast ofNew Guinea shows land. Stegonotus is represented in the Bis- a similarity between Umboi and mainland marck Archipelago by two species: S. New Guinea in mammalian fauna that is not heterurus (perhaps only a geographic varia- apparent among snakes. tion of S. modestus of the Moluccas, Manus, However, the similarity between New Brit- and northern New Guinea, but not recorded ain and New Guinea in the mammals makes from the Huon Peninsula); and an unnamed it difficult to assign the Umboi species to one endemic with dark upper lip, perhaps related or the other source; nearly all the Umboi to S. parvus ofthe Huon Peninsula and other species are common to both sources. Fur- regions of New Guinea. The Styporhynchus thermore, Koopman (personal commun.) of New Britain and Umboi is S. hypomelas finds that although there is no striking dis- (and, if it is distinct, S. dahli), an endemic similarity between New Britain and the Huon species (or endemic complex of two closely Peninsula fauna, this represents a general related species), sharply distinguished from similarity between New Guinea and New Huon Peninsula Styporhynchus but perhaps Britain and there is no indication ofa special related to an unnamed form from more west- resemblance between the Huon Peninsula and ern New Guinea (a 15-scale-rowed species, the Bismarck Archipelago fauna. From the 26 AMERICAN MUSEUM NOVITATES NO. 2775 literature and the (not yet completely iden- turnover and accept nineteenth-century rec- tified) Seventh Archbold Expedition collec- ords of snakes as members of the present tions, Koopman finds 53 species ofmammals fauna unless the record seems to have been on the Huon Peninsula (17 Chiroptera, 36 erroneous at the time it was made. I believe- non-flying); of these, none is endemic to the but have no evidence to prove-that this Huon Peninsula and none is found on the method gives a better estimate of the true Huon Peninsula and Bismarck Archipelago fauna than would acceptance of only those and not elsewhere in New Guinea; 20 (13 records made in the last decade as the true flying, seven non-flying) species are found on present fauna, but undoubtedly some turn- Huon and the Bismarck Archipelago and also over must exist; in point of fact, the snake in other New Guinea localities; 33 (four flying, fauna of Umboi Island as given by Hediger 29 non-flying) are found on the Huon Pen- (1934) is entirely different from the fauna of insula and other New Guinea mainland lo- Umboi seen by me in the Bernice P. Bishop calities but are unknown from the Bismarck Museum and American Museum collections Archipelago. It is worth noting that although made after World War II, and I cannot from bats make up only one-third ofthe mammals my own data disprove a total turnover ofthe recorded from the Huon Peninsula, they make snake fauna of Umboi around the time of up two-thirds of the species shared with the World War II, even though I doubt any such Bismarck Archipelago. thing happened. By Koopman's (personal commun.) listing Zweifel (1980) reviewed the geology ofthe of mammal species for the New Guinea-Sol- Huon Peninsula, which indicates a Pliocene omons region, for non-flying mammals the origin as an island (in the biologist's sense of pattern is one of progressive loss of species "island," an area capable ofsupporting a ter- (or species groups) reading from west (New restrial fauna and flora and isolated by a water Guinea) to east (the Solomons) through the barrier). Zweifel documents this relatively re- Bismarck Archipelago. This is also the most cent insular origin ofthe Huon Peninsula with common pattern for bats, but among Ptero- distribution patterns of highland lizards and podidae and Emballonuridae there are a few frogs showing disjunct distributions, in the cases of progressive loss of species reading interior of the Huon Peninsula and, most from east to west or of shared Bismarck Ar- often, on the central spine of New Guinea, chipelago-Solomons endemism: the Tribe but sometimes on the isolated mountain Notopterini (also in Oceania and New Cal- ranges of the north coast or the islands east edonia) of the Pteropodidae, Pteropus ad- of New Guinea. The distributions of snakes miralitatum, Nyctimene vizeaccia, and Em- fit fairly well with this pattern based on liz- ballonura dianae. These bats would fit the ards and frogs, but there are some differences, pattern of frogs (but not lizards) noted by largely exaggerations of the peculiarities not- Zweifel (1980), ofspecial affinity between the ed by Zweifel. Zweifel noted that "a curious Bismarck Archipelago and Solomons; but aspect of the (frog and lizard) fauna is the even among Chiroptera, this is not the pre- scarcity of endemic species" but could cite dominant pattern. three poorly known frogs; I cannot cite any For Umboi, it is impossible to guess positively known endemic snake, but the most whether the differences between the mam- conspicuous disjunction, Dendrelaphis lo- malian pattern (mostly a flying mammal pat- rentzi, may represent a distinct species, since tern) and the snake pattern result from: (1) the Huon specimen is distinguishable by differences in means of dispersal; or (2) from hemipenial details and ventral count from sampling error in collection (more precisely, specimens from the southern coast of New a different bias between herpetological sam- Guinea (and it must be remembered that pling and bat sampling in which species of Malnate and Underwood's revision of Sty- the true fauna will be missed); or (3) from porhynchus may add an endemic). The ex- differences in rate offaunal turnover. Because pectation would be that such endemism as of the obvious impossibility of sampling the there is should be among forms in the inte- entire faunal diversity in any single expedi- rior, presumably the forms that have been tion, I-and others-assume zero faunal isolated for the longest time. But neither the 1984 MC DOWELL: HUON PENINSULA SNAKES 27 frogs nor the Dendrelaphis support this-the the Huon Peninsula are converging, rather very limited "endemism" in the herpeto- than diverging. fauna is from coastal localities. Zweifel noted the absence ofhigh montane LITERATURE CITED frogs of more central New Guinea from the high interior of the Huon Peninsula, even Brongersma, Leo D. though a distinct highland fauna ofHuon can 1953. Notes on New Guinean and am- be distinguished. This "highland fauna" of phibians, II. Proc. K. Nederl. Akad. Wetensch., ser. C, vol. 56, pp. 317-325, Huon is made up of medium-elevation or p1. i. even lowland forms of more central New 1969. Note on Liasis boeleni Brongersma. Guinea. This is particularly evident in the Ibid., ser. C, vol. 72, pp. 124-128, pls. snakes, as well. The "highland fauna" ofHuon 1-11. contains Python amethistinus, rather than P. Hamilton, Warren boeleni, and Toxicocalamus loriae, rather 1979. Tectonics ofthe Indonesian region. U.S. than T. stanleyanus. Geol. Surv. Profess. Papers, no. 1078 All of this is in keeping with the Huon (U.S. Govt. Printing Off. Stock no. 024- Peninsula herpetofauna being a young fauna, 001 -03207-4), pp. i-ix, 1-345, sep. map. Heatwole, Harold derived from the most likely New Guinea 1975. Biogeography ofreptiles on some ofthe species to colonize an island-the coastal islands and cays of eastern Papua-New forms. Guinea. Atoll Res. Bull., no. 180, pp. So far as resemblance of the New Britain 1-32. fauna to that of the Huon Peninsula is con- Hediger, Heini cerned, the pattern seems to depend on the 1934. Beitrag zur Herpetologie und Zoogeo- taxonomic group considered. The surprising graphie Neu Britanniens und einiger lack ofresemblance seen in the herpetofauna umliegender Gebiete. Zool. Jahrb. is not conspicuous in the bats. The special (Syst.), vol. 65, nos. 5-6, pp. 389-582. resemblance between the frog fauna of New Koopman, Karl F. 1979. Zoogeography of mammals from is- Britain and that of the Solomons is not ap- lands off the northeastern coast of New parent in the snakes or the lizards. This is Guinea. Amer. Mus. Novitates, no. consistent with settlement by dispersal across 2690, pp. 1-17. water barriers, but with the differences in ma- Lidth de Jeude, T. W. van jor means of dispersal between taxonomic 1911. Reptilien (Schlangen) [von der Neu- groups resulting in somewhat different pat- Guinea-Expedition in den Jahren 1904, terns. Aside from the Typhlopidae (which I 1907 und 1909]. Nova Guinea, vol. 9, would discount for zoogeographic discussion Zool., pp. 265-287, pl. viii. because the difficulty of collection makes the L6nnberg, Einar true ranges of the species very uncertain), 1900. Reptiles and batrachians collected in German New Guinea by the late Dr. snakes must be among the last groups to col- Erik Nyman. Ann. Mag. Nat. Hist., ser. onize an island, because they are high-order 7, vol. 6, pp. 574-582. predators and their survival depends on the Loveridge, Arthur existence of prey fauna. Because most New 1948. New Guinean reptiles and amphibians Guinea snakes feed on lizards or frogs (but in the Museum ofComparative Zo6logy no lizard or frog is primarily a snake-eater), and United States National Museum. the colonization by snakes might be expected Bull. Mus. Comp. Zool., vol. 101, no. to lag behind the colonization by lizards and 2, pp. 305-430. frogs. McDowell, Samuel B. It may be noted that Hamilton's (1979) 1967. Aspidomorphus, a genus ofNew Guinea snakes of the Family Elapidae, with analysis of the plate tectonics of this region, notes on related genera. Jour. Zool. unavailable to Zweifel at the time he was (London), vol. 151, pp. 497-543. preparing his report, do not change his geo- 1969. Toxicocalamus, a New Guinea genus of logical assumptions and do not give any sup- snakes ofthe Family Elapidae. Ibid., vol. port to any previous connection of the Huon 159, pp. 443-511. Peninsula with New Britain; New Britain and 1970. On the status and relationships of the 28 AMERICAN MUSEUM NOVITATES NO. 2775

Solomon Island elapid snakes. Ibid., vol. Sternfeld, Richard 161, pp. 145-190. 1913. Beitrage zur Schlangenfauna Neugui- 1972. The species of Stegonotus (Serpentes, neas und der benachbarten Inselgrup- Colubridae) in Papua New Guinea. Zool. pen. Sitzber. Gesell. Naturf. Freunde Meded., vol. 47, pp. 6-26. Berlin, 1913, pp. 384-389. 1974. A catalogue ofthe snakes ofNew Guinea Storr, Glenn M. and the Solomons, with special refer- 1981. The genus Acanthophis (Serpentes: ence to those in the Bernice P. Bishop Elapidae) in Western Australia. Rec. Museum, Part I. Scolecophidia. Jour. West. Aust. Mus., vol.9, no.2, pp. 203- Herpetol., vol. 8, pp. 1-57. 210. 1975. A catalogue ofthe snakes ofNew Guinea Van Deusen, Hobart M. and the Solomons, with special refer- 1978. Results of the Archbold Expeditions. ence to those in the Bernice P. Bishop No. 101. Summary of the Seventh Museum. Part II. Anilioidea and Py- Archbold Expedition to New Guinea thoninae. Ibid., vol. 9, pp. 1-80. (1964). Amer. Mus. Novitates, no. 2660, 1979. A catalogue ofthe snakes ofNew Guinea pp. 1-21. and the Solomons, with special refer- Vogt, Theodor ence to those in the Bernice P. Bishop 1911. Reptilien und Amphibien aus Kaiser- Museum. Part III. Boinae and Acro- Wilhelmsland. Sitzber. Gesell. Naturf. chordoidea (Reptilia, Serpentes). Ibid, Freunde Berlin, 1911, pp. 420-432. vol. 13, pp. 1-92. Worrell, Eric Malnate, Edmond V. 1958. A new Papuan python. Proc. Roy. Zool. 1975. Grant No. 1150 .... The classification Soc. New S. Wales, 1956-57, pp. 26- and relationships of Australasian wa- 27. tersnakes. Amer. Philos. Soc. Year Book Zweifel, Richard G. 1974, pp. 357-358. 1980. Results of the Archbold Expeditions. Smith, Hobart M., Rozella B. Smith, and H. Lewis No. 103. Frogs and lizards from the Sawin Huon Peninsula, Papua New Guinea. 1977. A summary of snake classification Bull. Amer. Mus. Nat. Hist., vol. 165, (Reptilia, Serpentes). Jour. Herpetol., no. 5, pp. 387-434. vol. 11, pp. 115-121.