New Species of Hypsilurus (Squamata: Agamidae) from Papua New Guinea

New Species of Hypsilurus (Squamata: Agamidae) from Papua New Guinea FRED KRAUS 1,2 AND SHEPHERD MYERS 1 1 Bernice P. Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817 USA

Journal of Herpetology, Vol. 46, No. 3, 396 401, 2012 Copyright 2012 Society for the Study of Amphibians and Reptiles New Species of Hypsilurus (Squamata: Agamidae) from Papua New Guinea FRED KRAUS 1,2 AND SHEPHERD MYERS 1 1 Bernice P. Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817 USA ABSTRACT. We describe a new species of agamid lizard of the genus Hypsilurus from northern Papua New Guinea. The new species is characterized by its small adult size, scalational features, small nuchal and dorsal crests, and distinctive dorsal color pattern of brown and white chevrons. It is a member of the H. godeffroyi species group, but otherwise, its closest relative is not obvious. The species is currently known from two of the outlying North Coast ranges, but following the pattern seen in several other species endemic to this region, it will likely prove to occur along much of the northern lowlands of New Guinea. Agamid lizards do not present great diversity in New Guinea. Five genera are present, but four of those genera are represented by only one or two species (Allison, 1996), and most of these are shared with adjacent portions of Australia. The arboreal genus Hypsilurus Peters, 1867 presents an exception, with 16 species distributed in the Papuan region (Manthey and Denzer, 2006) and two additional species in the rain forests of Queensland (Manthey and Denzer, 2006; Shea, 2008), with the origin of Hypsilurus godeffroyi remaining undetermined (Crombie and Pregill, 1999; Bauer and Watkins-Colwell, 2001; Manthey and Denzer, 2006). All of the Papuan species are endemic to New Guinea, one or more of its adjacent islands, or the Solomon Islands. These lizards are poorly known, in part because they have long suffered from taxonomic confusion and misassignment, but this situation has improved slowly at least for taxonomic understanding. Moody (1980) recognized that these lizards are part of the Australian agamid radiation and removed them from the Asian Gonocephalus. Manthey and Denzer (2006) provided an extensive review of type specimens, rediagnosed all previously named species, described four new species and one new subspecies, synonymized one subspecies, removed another name from synonymy, and provided a key to all members of the genus. This work has improved taxonomic understanding of the genus considerably and provided a sufficiently credible revision that one of us (FK) has been able to easily identify all Hypsilurus residing in the extensive collections of the Australian Museum and Bishop Museum, as well as most of those animals collected by him in the course of continued biotic surveys in Papua New Guinea. Doing so has resulted in a significant expansion of our understanding of the geographic ranges of several of the Papuan species (Kraus, 2010). However, one species of Hypsilurus obtained during the course of recent herpetological surveys in New Guinea is unassignable to any known form, and it clearly represents a new species not included in the revision of Manthey and Denzer (2006). We describe it herein. MATERIALS AND METHODS We collected specimens under applicable national and provincial permits, euthanized them in accordance with applicable University of Hawaii animal-care-and-use procedures, fixed them in 10% buffered formalin, and transferred them to 70% ethanol for storage. We measured snout vent length (SV); tail length (TL); forelimb length ([FL] from body wall to tip of longest digit); and hindlimb length ([HL] from 2 Corresponding Author. E-mail: fkraus@hawaii.edu DOI: 10.1670/11-159 body wall to tip of longest digit) to the nearest 1 mm with a plastic ruler; and maximum head width (HW), tympanum width (TY), eye diameter (EY), and length of the longest nuchal scale ([NL] taken orthogonal to the long axis of the body) to the nearest 0.1 mm with either vernier calipers or an ocular micrometer fitted to a binocular dissecting scope, depending on specimen size; we measured mass to the nearest 0.1 g in the field with a Pesola scale. Scalational counts included numbers of mid-body scale rows, supralabials, infralabials, and scales along the upper eye margin. In several Hypsilurus there is a row of enlarged scales ventral to the infralabials that begins anteriorly along the ventral margin of the anterior infralabials but which becomes progressively separated from the infralabials as the row continues posteriorly. Manthey and Denzer (2006) referred to this row of enlarged scales as the submaxillaries and introduced a new character the maximum number of rows of small scales between the infralabials and submaxillaries. We include that same character here but instead refer to the enlarged row of scales as submandibles. We also introduce one new scalational feature here: the number of gular scale rows lying between the posteriormost of the enlarged submandibles. We determined sex by dissection and maturity by presence of eggs or enlarged ova (females) or convoluted, pigmented vasa deferentia and swollen testes (males). We based diagnostic features and comparisons to other species on data provided in Manthey and Denzer (2006) and by reference to specimens (Appendix I) housed in the Bernice P. Bishop Museum, Honolulu (BPBM), California Academy of Science (CAS), Museum of Comparative Zoology, Harvard University (MCZ), and Australian Museum, Sydney (AMS). Type specimens of the new species are deposited in the BPBM, MCZ, and Papua New Guinea National Museum (PNGNM). Locality coordinates for the new species use GPS datum AUS 66 (specimens from West Sepik Province) or WGS 84 (specimens from East Sepik Province). Hypsilurus capreolatus, sp. nov. Figures 1 3 Holotype. BPBM 23413 (field no. FK 11409), adult female, obtained by native collectors at Parkop, 3.424578S, 142.518668E, 420 m (GPS datum AUS 66), Torricelli Mts., West Sepik Province, Papua New Guinea, 13 May 2005. Paratypes (n = 28). Adult males (n = 2): Papua New Guinea: East Sepik Province: Bainyik (MCZ 152896); West Sepik Province: Torricelli Mts.: 3.2 km SSE Mt. Sapau summit, 3.393298S, 142.528268E, 550 m (BPBM 23420). Subadult males (n = 9): West Sepik Province: Torricelli Mts.: Parkop, 3.424578S, 142.518668E, 420 m (BPBM 23414, 23417 19), 3.2 km SSE Mt. Sapau summit,

NEW LIZARD FROM NEW GUINEA 397 FIG. 1. Portrait of the holotype of Hypsilurus capreolatus (BPBM 23413) in life, showing the distinctive pattern of dorsal chevrons characteristic of the species. 3.393298S, 142.528268E, 550 m (PNGNM 25172); East Sepik Province: Prince Alexander Mts.: Joromba, 3.57928338S, 143.50058E, 87 m (BPBM 35621), along Joromba River, 3.578858S, 143.500338E, 230 m (BPBM 35622), Herimuri, 3.58891668S, 143.5012338E, 67 m (BPBM 35623 24). Adult female (n = 1): East Sepik Province: Bainyik (MCZ 152897). Subadult females (n = 3): West Sepik Province: Torricelli Mts.: Parkop, 3.424578S, 142.518668E, 420 m (BPBM 23415); East Sepik Province: Prince Alexander Mts.: Nioguangia, 3.55548S, 143.5381338E, 12 m (BPBM 34751, 35631). Unsexed juveniles (n = 13): West Sepik Province: Torricelli Mts.: Parkop, 3.424578S, 142.518668E, 420 m FIG. 2. Dorsal crests of (A) Hypsilurus capreolatus (PNGNM 25172) and (B) Hypsilurus longii (CAS 94023), showing relative height differences. Anterior is to the right side; scale bars = 5 mm. FIG. 3. Views of (A) side of head and (B) throat of holotype of Hypsilurus capreolatus (BPBM 23413).

398 F. KRAUS AND S. MYERS (BPBM 23412, 23416, PNGNM 25171); East Sepik Province: Prince Alexander Mts.: Nioguangia, 3.55548S, 143.5381338E, 12 m (BPBM 35625 30, 35632 35). Diagnosis. Hypsilurus capreolatus is a small species for the genus (adult males, 97 115 mm; adult females, 100 119 mm) characterized by having the unique combination of small, homogeneous dorsal scales; a nuchal crest with small, lanceolate scales approximately one-third diameter of tympanum (range, 0.15 0.69; mean, 0.31), separated from a dorsal crest of very low, largely overlapping scales; 114 158 scale rows around mid-body (mean, 128.6); a row of enlarged submandibles extending from mental to below rictus; a maximum of three to four scales present between infralabials and submandibles; scales of gular sac small, smooth or tuberculate, juxtaposed to subimbricate, and of similar size anteriorly and posteriorly, lacking enlarged scales anteriorly or medially, and with 30 42 (mean, 36.0) rows of gular scales extending between submandibular scale rows at level of the posterior-most infralabials; weakly enlarged scales at rictus and below tympanum; HW/SV = 0.16 0.22 (mean, 0.19); TY/EY = 0.51 0.94 (mean, 0.62); with a dorsal pattern of five long, brown chevrons across the dorsum between the nuchal crest and hindlimbs, these separated by four light chevrons (Fig. 1); and throat and sides of neck spotted with black. Comparisons with Other Species. Hypsilurus capreolatus may be distinguished from H. modestus by having (vs. lacking) a dorsal crest; from H. boydii, H. dilophus, and H. spinipes by having homogeneous dorsal scales (vs. having large dorsal scales scattered among the smaller); from H. auritus, H. geelvinkianus, and H. nigrigularis in having (vs. lacking) a row of enlarged submandibular scales extending from mental to below rictus and a rounded (vs. acuminate) base of the gular pouch near the clavicular region (vs. on the chest); from H. tenuicephalus in having only a single enlarged scale (vs. many) below the tympanum, the submandibles extending to the level of the rictus (vs. ending at the level of the middle of the lower jaw), and a broader head (HW/SV > 0.15 in H. capreolatus vs. HW/SV < 0.12 in H. tenuicephalus); from H. hikidanus and H. schultzewestrumi in lacking (vs. having) a series of enlarged midventral scales on the throat; from H. binotatus and H. ornatus in having the nuchal and dorsal crests separate (vs. continuous) and the gular scales smooth or tuberculate (vs. keeled or tuberculate); and from H. macrolepis and H. papuensis in lacking (vs. having) greatly enlarged scales below the tympanum and in having smooth or tuberculate (vs. strongly keeled) gular scales. Hypsilurus capreolatus differs from H. bruijnii in its smaller adult size (SV up to 119 mm in H. capreolatus vs. up to 143 mm in H. bruijnii), in usually having four (three in 14% of 29 specimens) instead of two to three rows of scales between the infralabials and submandibles, in having (vs. lacking) weakly enlarged scales below the tympanum and rictus, and in being dorsally banded (vs. brown with small dark spots in H. bruijnii); it differs from H. schoedei in usually having four instead of two to three rows of scales between the infralabials and submandibles, in having a broader head (HW/ SV > 0.15 in H. capreolatus vs. HW/SV < 0.15 in H. schoedei), in having much smaller gular scales, and in having only four light bands on the back (vs. having either a uniformly brown or bluegreen dorsum or with five to 10 light bands). Hypsilurus capreolatus is never uniformly brown above, whereas H. schoedei usually is; H. capreolatus never has a V-shaped marking on nape, whereas H. schoedei sometimes does. Using the key provided by Manthey and Denzer (2006), H. capreolatus would seem most similar to H. magnus and H. longii. From both of these species, H. capreolatus differs in its much smaller size (adult males to 115 mm, adult females to 119 mm in H. capreolatus vs. up to 235 and 210 mm, respectively, for H. longii and 223 and 180 mm, respectively, for H. magnus). From H. longii, H. capreolatus further differs in having a lower nuchal crest (scale length approximately one-third the width of tympanum in H. capreolatus vs. one-half the width of tympanum in H. longii); a lower dorsal crest (Fig. 2); smaller gular scales (30 42 rows between enlarged submandibles at level of posterior-most infralabial in H. capreolatus vs. 20 21 in H. longii); more mid-body scale rows (114 158 in H. capreolatus vs. 94 102 in H. longii); more scales along upper margin of eye (16 24 in H. capreolatus vs. 15 17 scales in H. longii); anterior gulars approximately same (no more than twice) size as posterior gulars (vs. 5 10-fold difference in size in H. longii); and sides of neck spotted with black (vs. unspotted white or brown in H. longii). The dorsal bands of H. capreolatus are also better defined, more oblique, and extend farther ventrally than do those of H. longii. From H. magnus, H. capreolatus further differs in having a lower nuchal crest (NL/TY = 0.15 0.69; mean, 0.31 in H. capreolatus vs. NL/TY = 0.28 1.78; mean, 0.59 in H. magnus); anterior gulars approximately same (no more than twice) size as posterior gulars (vs. three- to five-fold difference in size in H. magnus); a dorsal pattern of long brown chevrons (vs. short mid-dorsal bars in H. magnus); and throat spotted with black (vs. streaked with black lines in H. magnus). In life, H. magnus is dark purple brown heavily speckled with tan and the throat is bright yellow or white streaked with dark purple brown. In contrast, H. capreolatus is pale tan with brown chevrons and speckles dorsally, laterally, and on the throat (Fig. 1). The adult female holotype of H. capreolatus had an SV of 100 mm and a mass of 29.1 g, and an adult male paratype (BPBM 23420) had an SV of 97 mm and a mass of 32.0 g; that of an adult male H. magnus (BPBM 34769) had an SV of 210 mm and a mass of 287 g, well illustrating the size difference between the species. Description of Holotype. An adult female with enlarged oviductal eggs. Rostral semilunar, bordered by first supralabials, five small scales dorsally, and two (right) and one (left) small scales dorsolaterally. Enlarged nasal perforated by external naris, bordered by eight small scales. Scales on snout and top of head keeled, imbricate. Canthus of sharply raised scales extending from snout to above eye. Upper eyelid bordered by 21 scales, lower by 22. Temporal scales slightly enlarged, keeled; one scale below tympanum enlarged to twice size of its neighbors. Supralabials 12 (right), 11 (left); infralabials nine (right), 10 (left). A row of enlarged submandibles from mental to below rictus, 13 (right), 12 (left); these separated by one (right) or two (left) rows of small scales from three (right) or two (left) scales lying between submandibles and anterior edge of tympanum that are enlarged to twice the size of adjacent scales; a maximum of four scale rows between infralabials and submandibles; 42 rows of gular scales across throat between posteriormost submandibles. Mental semilunar with scalloped margins, bordered by first infralabials, first submandibles, and two anterior gulars. Gular pouch small; anterior scales on gular pouch small, subimbricate, rhomboidal, almost pyramidal, with raised points on posterior of each scale; posterior scales on gular pouch smaller, granular; scales on chin approximately equal to or twice size of those on gular pouch (Fig. 3). Dorsal scales homogeneous, rhomboidal, imbricate, with strong keels oriented at an upward angle posteriorly, in 131 rows at mid-body; smallest laterally, enlarging dorsolaterally and ventrolaterally; with keels becoming progressively more

NEW LIZARD FROM NEW GUINEA 399 TABLE 1. Morphometric and meristic data for type series of Hypsilurus capreolatus. The categories males and females include all specimens that could be sexed, whether mature or not. Adults (n = 4) Subadults (n = 12) Juveniles (n = 13) Males (n = 11) Females (n = 5) Character Mean Range Mean Range Mean Range Mean Range Mean Range SV 108 97 119 69 55 82 48 40 55 76 55 115 86 66 119 TL 332 317 350 203 160 256 133 106 164 233 160 346 259 194 350 FL 53.3 47 60 35.2 29 42 25.4 21 30 38.4 29 55 42.6 32 60 HL 91.8 85 95 65.3 51 79 44.6 36 52 70.2 51 95 75.6 60 94 HW 17.5 16.1 18.6 12.9 11.1 14.6 9.8 8.5 11.0 13.8 11.1 18.6 14.6 12.2 17.6 EY 5.8 5.1 6.1 4.0 3.2 5.0 3.2 2.6 3.6 4.4 3.2 6.1 4.6 3.4 6.1 TY 4.0 3.4 4.7 2.6 1.9 3.3 1.9 1.6 2.2 2.8 1.9 4.4 3.2 2.1 4.7 NL 2.7 1.4 3.8 1.0 0.5 1.6 0.4 0.3 0.6 1.4 0.5 3.8 1.5 0.9 3.2 TL/SV 3.1 2.9 3.3 3.0 2.9 3.1 2.7 2.5 3.0 3.0 2.9 3.3 3.0 2.9 3.2 FL/SV 0.49 0.48 0.51 0.51 0.45 0.54 0.53 0.49 0.56 0.51 0.47 0.54 0.50 0.45 0.53 HL/SV 0.86 0.79 0.98 0.94 0.89 0.99 0.94 0.88 1.0 0.93 0.81 0.99 0.90 0.79 0.99 HW/SV 0.16 0.15 0.18 0.19 0.17 0.21 0.21 0.19 0.22 0.18 0.16 0.21 0.17 0.15 0.19 TY/EY 0.70 0.63 0.77 0.65 0.51 0.94 0.60 0.51 0.74 0.64 0.51 0.79 0.71 0.51 0.94 NL/TY 0.66 0.41 0.86 0.39 0.21 0.65 0.21 0.15 0.27 0.46 0.21 0.86 0.45 0.33 0.68 Midbody scale rows 139.3 131 158 129.4 117 142 124.6 114 146 131.6 117 158 132.4 125 140 Supralabials 10.8 10 12 11.2 9 12 11.2 10 13 11.0 9 12 11.2 10 12 Infralabials 9.0 9 10.1 9 12 9.7 8 12 9.8 9 12 9.8 9 11 Scales along upper eyelid 19.3 18 21 19.6 18 22 20.8 16 24 19.8 18 22 18.8 18 21 Scales btw submandibles and infralabials 3.5 3 4 3.8 3 4 4.0 4 3.7 3 4 3.8 3 4 Gular rows btw posterior submandibles 38.0 34 42 36.8 33 42 34.7 30 42 36.5 33 42 38.6 34 42 horizontally oriented ventrally. Ventral scales larger, rhomboidal, imbricate, with sharply pointed keels oriented posteriorly. Nuchal crest of nine to 10 relatively small lanceolate scales that are each typically separated from its neighbors by one or two small intervening scales. Nuchal crest separated from dorsal crest by a gap approximately one-third the length of nuchal crest. Dorsal crest of low, enlarged, pointed, backward-directed scales that overlap their posterior neighbors; these scales gradually lose their distinctiveness from adjacent scales approximately halfway along tail. Limbs relatively long, with the hindlimbs longer than the forelimbs (FL/SV = 0.51, HL/SV = 0.85). Base of tail laterally compressed. Brown with two indistinct darker blotches on nape, another larger blotch above the pelvic region, and five dark-bordered brown chevrons with forward-pointing apices between these blotches; these brown blotches and chevrons separated by five paler, dirty-tan chevrons. Sides lighter brown speckled with dark brown; tail with dark-bordered brown bands approximately twice the width of paler intervening ground. Venter dirty white with small dark brown spots scattered laterally. Chin and throat dirty white with scattered, small dark brown spots (Fig. 3). Tympanum light tan. Measurements. Measurements (in mm) are SV = 100, TL = 317, FL = 51, HL = 85, HW = 16.1, EY = 5.1, TY = 3.4, and NL = 1.4. Variation. Specimens range in SV from 40 to 119 mm, with most being juveniles or immature lizards. Adults of both sexes are of similar size, reaching approximately 120 mm SV, and there are no obvious sexual differences in any morphometric or meristic feature (Table 1). Animals <55 mm SV could not be reliably sexed, but immature animals between 55 and 82 mm SV could be. Some morphometric features seem to vary ontogenetically, with relative HW and limb lengths decreasing with age, whereas relative TL and relative NL increase (Table 1). It is also clear to the naked eye that the nuchal and dorsal crests of young animals are less well developed than in adults. There are suggestions of differences in meristic features between age classes, with adults seeming to have more midbody scale rows, more gulars between the submandible, and fewer scales along the upper eyelid than juveniles (Table 1), but conclusions drawn from this sample are tempered by the small number of adults available. One specimen (BPBM 35622) lacks the slightly enlarged scale below the tympanum. Variation in color pattern is slight, with the primary difference being in the relative contrast of the dorsal chevrons in preserved animals, which vary from being rather obscure to bold. In life, these pattern elements were always obvious. One specimen (PNGNM 25172) has five tan chevrons dorsally instead of the more usual four. Venters are usually unspotted dirty white or with only a few dark brown spots, these most often concentrated laterally; however, three specimens (BPBM 23415, 35623; PNGNM 25172) consisting of one immature female and two immature males varying in SV from 68 to 82 mm, have more spots, and these have a more even distribution across the venter. The dark gular spotting is absent in some of the smallest lizards but is always evident in the adults and subadults. Color pattern does not seem to differ ontogenetically or between sexes. Color in Life. Field notes for BPBM 23414 (immature male, SV = 62 mm) read: Dorsum dull gray-brown banded obscurely with darker brown. Venter brown, lighter on chin and with pale orange-yellow on throat BPBM 23415 (immature female, SV = 68 mm) had the throat lemon yellow, and the belly and undersides of the rear limbs had a pale salmon-red wash. BPBM 35621 (immature male, SV = 70 mm) also had a yellow throat. The holotype was dirty white with brown chevrons irregularly margined with black spots (Fig. 1). Ecological Notes. Animals were collected in both primary lowland rainforest and in secondary forest near villages, occurring from near sea level to 550-m elevation. Forest canopies in these areas varied from 25 to 35 m in height. Etymology. The trivial epithet is a Latin masculine adjective of capreolus meaning rafter, from which is derived the French

400 F. KRAUS AND S. MYERS FIG. 4. Map of eastern New Guinea showing known localities for Hypsilurus capreolatus. Filled circle = type locality. (and, hence, English) chevron. It refers to the diagnostic pattern of this species. Range. Distributed along the lower elevations of the Prince Alexander and Torricelli ranges and associated lowlands of the Sepik River in northern Papua New Guinea (Fig. 4). Like many other lowland species of the northern ranges, it is likely to be distributed across the northern lowlands of the entire island west of the Huon Peninsula. DISCUSSION Following the definitions provided by Manthey and Denzer (2006), H. capreolatus is clearly a member of their H. godeffroyi species group, having adult TL/SV > 3.0, homogeneous dorsal scalation, enlarged submandibular scales, and the anterior scales of the gular pouch slightly larger than the posterior scales. Within that group, its relationships are less clear. It shares with H. magnus, H. ornatus, and H. tenuicephalus the feature that the size differentiation between anterior and posterior scales of the gular pouch is rather minor, whereas the other members of the H. godeffroyi species group have a much more striking size demarcation. However, H. capreolatus differs from each of these species in several important features and is not more clearly similar to one species than another. Resolution of its relationships to these species will have to await explicit phylogenetic analysis that is currently hindered by insufficient material of several Hypsilurus species for conducting either morphological or molecular analyses. Hypsilurus capreolatus brings to 14 the number of Hypsilurus species known from the main island of New Guinea, with a few additional species occurring on some of its satellite islands (Manthey and Denzer, 2006). Several species remain known only from their type localities (e.g., H. bruijnii, H. nigrigularis, H. ornatus, H. schoedei) or have uncertain provenance (H. godeffroyi, H. tenuicephalus). It remains uncertain to what extent these represent taxa with truly restricted geographic ranges or whether they reflect a lack of collecting effort coupled with prior taxonomic confusion. One limitation of the revision of Manthey and Denzer (2006) is that even though it is taxonomically extensive, it is based primarily on older collections in European museums that generally do not have geographically extensive holdings of Papuan material. As a result, some species reported from only geographically limited areas by Manthey and Denzer (2006) have been shown to have much larger distributional ranges than initially thought (Kraus, 2010). The same may prove true for some of the species listed above, as well as for our new species. Currently, H. capreolatus is known only from two of the North Coast mountain ranges in Papua New Guinea (Prince Alexanders, Torricellis). Several other species of reptiles and amphibians newly described in recent years from the north coast of New Guinea in the far west of the island (Cophixalus balbus, Hylarana volkerjane, Hylophorbus picoides, Lechriodus platyceps, Litoria purpureolata) have been found to range far eastward to several of the North Coast ranges in Papua New Guinea (Kraus and Allison, 2006; Kraus, 2010, unpubl. data), and many other species endemic to the North Coast ranges are known to be extensively distributed across that area (cf. Papuan Herpetofauna website at http://www. bishopmuseum.org/research/pbs/papuanherps/). Indeed, few, if any, seem limited to single mountain ranges within that region, even though these mountain ranges are derived from many different terranes having offshore origins (Pigram and Davies, 1987) that could be expected to promote some degree of single-range endemism. Consequently, although H. capreolatus is known currently from only two adjacent North Coast ranges, we predict that it will prove to be distributed more widely across additional mountain ranges of the northern portion of New Guinea. Acknowledgments. We thank J. Vindum (CAS) and J. Rosado (MCZ) for loan of specimens; R. Sadlier for providing access to specimens at AMS; C. Kishinami and P. Imada for specimen processing and documentation; D. Cameron for helpful advice on Latin epithets and grammar; and I. Haguna, J. Jiramori, A. Nakung, S. Simalken, J. Slapcinsky, C. Wintawa, D. Yaripma, B. Yawi, and all the people of Parkop for providing field or logistical assistance during our expeditions. We thank the PNG National Museum and Art Gallery for providing in-country collaborative assistance and the Department of Environment and Conservation; National Research Institute; and the East Sepik and West Sepik provincial governments for permission to conduct this research. This research was supported by National Science Foundation grants DEB-0103794 and DEB-0743890. This is contribution 2012-005 to the Pacific Biological Survey. LITERATURE CITED ALLISON, A. 1996. Zoogeography of amphibians and reptiles of New Guinea and the Pacific region. In A. Keast and S. E. Miller (eds.), The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. pp.407 436. SPB Academic Publishing, Amsterdam, The Netherlands. BAUER, A. M., AND G. J. WATKINS-COLWELL. 2001. On the origin of the types of Hypsilurus godeffroyi (Reptilia: Squamata: Agamidae) and early German contributions to the herpetology of Palau. Micronesica 34: 73 84. CROMBIE, R. I., AND G. K. PREGILL. 1999. A checklist of the herpetofauna of the Palau Islands (Republic of Belau), Oceania. Herpetological Monographs 13:29 80. KRAUS, F. 2010. More range extensions for Papuan reptiles and amphibians. Herpetological Review 41:246 248. KRAUS, F., AND A. ALLISON. 2006. Range extensions for reptiles and amphibians along the northern versant of Papua New Guinea. Herpetological Review 37:364 368. MANTHEY, U., AND W. DENZER. 2006. A revision of the Melanesian- Australian angle head lizards of the genus Hypsilurus (Sauria: Agamidae: Amphibolurinae), with description of four new species and one new subspecies. Hamadryad 30:1 40.

NEW LIZARD FROM NEW GUINEA 401 MOODY, S. 1980. Phylogenetic and historical biogeographical relationships of the genera in the family Agamidae (Reptilia, Lacertilia). Unpubl. Ph.D. diss., University of Michigan, Ann Arbor. PIGRAM, C., AND H. DAVIES. 1987. Terranes and the accretion history of the New Guinea Orogen. BMR Journal of Australian Geology and Geophysics 10:193 211. SHEA, G. M. 2008. Rejection of an Australian distribution of the Solomon Islands agamid lizard Hypsilurus longii (Macleay, 1877). Hamadryad 33:127 129. Accepted: 18 March 2012. APPENDIX I Specimens Examined Hypsilurus auritus: Indonesia: Papua Province: Manokwari Division: Oransbari (BPBM 3294), Kebar Valley (BPBM 3845 46). Hypsilurus binotatus: Papua New Guinea: Western Province: Lake Murray, 6.78338S, 141.4338E (AMS R24282). Hypsilurus dilophus: Papua New Guinea: East Sepik Province: near Yefehi, 3.60528S, 143.49278E, 400 m (BPBM 34750); Northern Province: Ajoa River, 9.200108S, 149.088788E, 850 m (BPBM 37321), Ajoa River, 9.238848S, 149.135598E, 400 m (BPBM 37322 24); West Sepik Province: Trefas (BPBM 23158, 23166, 23173 74). Hypsilurus hikidanus: Indonesia: Papua Province: Paniai Division: Itoda (BPBM 3396), Enarotali (BPBM 3739 41); Papua New Guinea: Chimbu Province: Toromambuno, Mt. Wilhelm, 5.84420288S, 145.10180188E (BPBM 2693 96). Hypsilurus longii: Papua New Guinea: Bougainville Island: Buin, 6.8338S, 155.7338E (AMS R11259-61, R11419, R11454-57), Kunua (CAS 94023), Turiboiru (CAS 107815); Solomon Islands: Isabel Island: Alu Alu, 8.38S, 159.558E (AMS R137241); Kaipitu River, near Alu Alu, 8.38S, 159.558E (AMS R137243); Makira Island (AMS R127290); Makira Island: Sgsgna, (AMS R130465). Hypsilurus macrolepis: Solomon Islands: Choiseul Island: Malangona, 7.0498S, 156.7788E (BPBM 2488); Isabel Island: Alu Alu, 8.38S, 159.558E (AMS R137242); Makira Island: Sgsgna, (AMS R130464); San Cristobal Island: 10.68S, 161.758E (AMS R10407-10), Mami (BPBM 2264), Kirakira (BPBM 2531 33, 5868, 6725 26); St. Anna Island: 10.816668S, 162.458E (AMS R131114-19); Ugi Island: 10.258S, 161.758E (AMS R42686, R131112-13). Hypsilurus magnus: Papua New Guinea: East Sepik Province: Huriak, 3.58461668S, 143.5007338E, 70 m (BPBM 34769); Southern Highlands Province: confluence of Libano and Hegigio rivers, 6.39906668S, 142.97613338E, 250 m (BPBM 28232 34), Namosado, 6.258S, 142.78338E (AMS R122435-40, R122442-44, R122470), Bobole, 6.208S, 142.76668E (AMS R122441), Fogamaiyu, 6.51668S, 143.08338E (AMS R122474); West Sepik Province: Menawa River, 8.4 km N, 11.4 km E of Utai, 3.3128S, 141.6888E, 560 m (BPBM 23161, 23164); Western Highlands Province: Baiyer River, 5.558S, 144.008E (AMS R28690). Hypsilurus modestus: Papua New Guinea: East New Britain Province: Marmar, 5.51902178S, 151.50180188E, 30 m (BPBM 22174), 2.6 km NNW Marmar, 5.498958S, 151.48938E, 520 m (BPBM 22172 73, 22175 76); Morobe Province: Apele, Mt. Shungol, 6.790658S, 146.665358E, 330 m (BPBM 18664); West Sepik Province: Torricelli Mts.: Parkop, 3.424578S, 142.518668E, 420 m (BPBM 23421 29). Hypsilurus papuensis: Papua New Guinea: Central Province: Laronu, 9.443878S, 147.983838E, 880 m (BPBM 19507-08), Moroka, 9.437968S, 147.598048E, 262 m (BPBM 22177); Milne Bay Province: Alotau (BPBM 15830, 17252, 17726), Normanby Island: S end Sewa Bay, Wassasaia River, 10.05163048S, 150.97168958E, 50 m (BPBM 17251), Siyomu, 10.01449298S, 149.59699458E, 1300 m (BPBM 17253), Bunisi, 10.017138S, 149.600158E, 1420 m (BPBM 17254). Hypsilurus schultzewestrumi: Indonesia: Papua Province: Bokondini, 40 km. N. of Baliem Valley, 3.638S, 138.68E, 1400 m (BPBM 3296); Papua New Guinea: Chimbu Province: Goglme, 5.933338S, 145.033338E (AMS R29126 28); Eastern Highlands Province: Roka Estate Village on Goroka-Daulo Pass Road, 6.058S, 145.48E (AMS R64740 71); Koge, 6.108S, 145.016668E (AMS R68895); Irumbafoi, 6.233338S, 145.266668E (AMS R86887); Enga Province: Yaramanda, West Baiyer Valley, 5.616668S, 143.9166668E (AMS R14849 50); Madang Province: Fungoi, Kaironk Valley, Schrader Mts., 5.3333338S, 144.416666678E (AMS R21190 91, 23039, 24328 30); West Sepik Province: Telefomin, 5.1367758S, 141.6333338E (BPBM 3858); Western Highlands Province: Minj (BPBM 2090).