A New Species of New Guinea Worm-eating Snake, Genus Toxicocalamus (Serpentes: Elapidae), from the Star Mountains of Western Province, Papua New Guinea, with a Revised Dichotomous

A NEW SPECIES OF NEW GUINEA WORM-EATING SNAKE, GENUSTOXICOCALAMUS (SERPENTES: ELAPIDAE), FROM THE STARMOUNTAINS OF WESTERN PROVINCE, PAPUA NEW GUINEA, WITHA REVISED DICHOTOMOUS KEY TO THE GENUS

MARK O’SHEA,1,4 FRED PARKER,2 AND HINRICH KAISER3

CONTENTS

Abstract ------- -- - - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 241Introduction -------- - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 242Materials and Methods ------- - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 242Toxicocalamus ernstmayri New Species ------- - - 243

Holotype -------- - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 243Etymology ------- - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 244Diagnosis ------ - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 244Comparisons ------- - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 248Description of the Holotype -------- -- - - - - - - - - - - - - - - - - 249Coloration in Life -------- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 251Coloration in Preservative ------- - - - - - - -- - - - - - - - - - - - - - - - - 251Tail Tip Morphology ------- - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 251Osteology ------- - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 252Natural History ------- - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 252

Discussion -------- - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 255Key to the Genus Toxicocalamus - - - - -- - - - - - - - - - - - - - - - - 259Acknowledgments ------- - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 260Appendix 1. Specimens Examined -------- - - - - - - - - - - - - 260Literature Cited -------- - - - - - - -- - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - 263

ABSTRACT. We describe a new species of New Guineavermivorous snake (Toxicocalamus) from a singlespecimen collected at Wangbin in the Star Mountains,

Western Province, Papua New Guinea. The newspecies is the largest known member of the genusand can be differentiated from all other Toxicocalamusby a combination of the following characters: large size(total length of the holotype 1,200 mm), dorsal headscutes in the typical ‘‘colubrid-elapid dorsal nine-scutearrangement’’; separate, single preocular and pairedpostoculars; single anterior temporal and single orpaired posterior temporals; six supralabials, with thirdand fourth supralabial contacting the orbit; dorsalscales in 15–15–15 rows; 203 ventral scales, 29subcaudal scales; and a divided anal plate. Its erstwhilestatus, misidentified as Micropechis ikaheka in thecollection of the Museum of Comparative Zoology,demonstrates the need for detailed examination ofexisting collections and is indicative of hidden diversityyet to be identified, not only in the field but also on theshelves of museum collections. We also providea revised key to the genus Toxicocalamus.

ABSTRAKSI. Kami mendeskripsikan spesies baru ularpemakan cacing (Toxicocalamus) di Papua New Guineadari koleksi spesimen di Wangbin, Pegunungan Bin-tang, Propinsi bagian Barat, Papua Nugini. Spesiesbaru ini merupakan anggota genus dan dapat dibeda-kan dari semua ular pemakan cacing tanah (Toxicoca-lamus) lainnya dengan kombinasi dari berbagai ciri-cirisebagai berikut: berukuran besar (panjang total holo-type 1200 mm), sisik di kepala bagian atas bercirikan‘‘colubrid-elapid sembilan sisik beraturan’’; terpisah,preocular tunggal and sepasang postocular; anteriortemporal tunggal dan satu atau sepasang posteriortemporal; enam supralabial, dengan supralabial ketigadan keempat menghubungi orbit; sisik di bagianpunggung berbaris 15–15–15; sisik di bagian perutberjumlah 203, sisik subcaudal berjumlah 29; danpiringan anal dibagi menjadi dua bagian. Statussebelumnya, salah diidentifikasi sebagai Micropechisikaheka di koleksi Museum Comparative Zoology. Inimenunjukkan perlunya penyelidikan secara rinci ter-hadap hasil koleksi yang ada sekarang. Hal ini jugamerupakan indikasi adanya keragaman tersembunyi

1 West Midland Safari Park, Bewdley, Worcester-shire DY12 1LF, United Kingdom; and AustralianVenom Research Unit, University of Melbourne,Victoria 3010, Australia. Author for correspondence([email protected]).

2 P.O. Box 5623, Townsville, Queensland 4810,Australia.

3 Department of Biology, Victor Valley College,18422 Bear Valley Road, Victorville, California 92395;and Department of Vertebrate Zoology, NationalMuseum of Natural History, Smithsonian Institution,Washington, DC 20013.

4 Present address, Faculty of Science and Engineer-ing, University of Wolverhampton, Wulfruna Street,Wolverhampton, West Midlands WV1 1LY, UnitedKingdom.

Bull. Mus. Comp. Zool., 161(6): 241–264, July, 2015 241

yang belum teridentifikasi, tidak hanya di lapangan tapijuga di rak-rak museum koleksi. Kami juga memberi-kan kunci revisi tentang spesies Toxicocalamus.

Key words: Elapidae, Toxicocalamus, New species,Papua New Guinea, Description, Taxonomy, Vermivory

INTRODUCTION

Toxicocalamus is a genus of enigmaticvermivorous elapid snakes with a primarilyfossorial lifestyle. Although part of the verydiverse Australopapuan terrestrial elapidradiation, Toxicocalamus is one of only sixgenera not represented on the Australiancontinent. It is endemic to the island of NewGuinea, several small offshore islands, andthe major island archipelagos to the south-east of the Papuan Peninsula (Fig. 1). Thesmall offshore islands all lie along the northcoast of Papua New Guinea (PNG) andinclude Seleo Island (Sandaun Province),Walis and Tarawai Islands (East SepikProvince), and Karkar Island (Madang Prov-ince). The southeastern archipelagos arepart of Milne Bay Province and includethe d’Entrecasteaux Archipelago (Goode-nough, Fergusson, and Normanby Islands),Woodlark Island, and the Louisiade Archi-pelago (Misima, Sudest, and Rossel Islands).Toxicocalamus is seemingly absent from theBismarck, Admiralty, and Solomons Archipel-agos, the great seasonally flooded Trans-Flyregion of southern New Guinea, the TorresStrait Islands, the Schouten Islands, and theVogelkop and associated Raja Ampat Archi-pelago of West New Guinea (Indonesia).

First described by Boulenger (1896) andlater revised by McDowell (1967, 1969), thegenus Toxicocalamus currently comprises11 species and a single subspecies, including(in order of description): T. longissimusBoulenger, 1896; T. loriae (Boulenger,1898); T. stanleyanus (Boulenger, 1903);T. preussi (Sternfeld, 1913); T. buergersi(Sternfeld, 1913); T. grandis (Boulenger,1914); T. p. angusticinctus (Bogert &Matalas, 1945); T. spilolepidotus McDowell,1969; T. holopelturus McDowell, 1969;T. misimae McDowell, 1969; T. mintoniKraus, 2009; and T. pachysomus Kraus,2009. Many of these taxa are poorly

represented in natural history collections,and we have so far located 475 specimens in28 museums worldwide. Of these, 66.3%(315 specimens) are identified as T. loriae,but considering the great variation in habitus,patterning, and scale count we have observedamong these specimens, we believe this taxonto represent a species complex. The nextbest-represented species are T. preussi andT. stanleyanus, with 55 and 37 specimens,respectively. Two island taxa, T. holopelturusand T. longissimus, are known from 18 and16 specimens, respectively, whereas ananomalous ‘‘hybrid’’ population from Garaina(Morobe Province, PNG; McDowell, 1969) isknown from 15 specimens. The remainingseven species are represented by eight orfewer specimens each, with three species,excluding the one we here describe, onlyknown from their holotypes. The largestknown species so far is T. grandis, at 960mm snout–vent length (SVL) for the singleknown specimen (BMNH 1946.1.18.34).That species was described from the SetakwaRiver (southern Papua Province, West NewGuinea, Indonesia), where it was collected in1912 during the Wollaston Expedition of1912–13 (O’Shea, 2013).

We here describe a new species, thelargest of the genus, that we discoveredamong a series of Micropechis ikaheka inthe collection of the Museum of Compara-tive Zoology, where it had languished for 45years since its collection by one of us (FP).Its description is yet another recent exampleof the unrecognized diversity of naturalhistory collections (see Kathriner et al.,2014), and it showcases their value tostudents of biological diversity.

MATERIALS AND METHODS

Characters used for evaluating and com-paring specimens were taken from 344museum specimens (Appendix 1). Abbrevia-tions for measurements and scale countsused in the description include snout–ventlength (SVL), tail length (TL), total length(TTL), number of subcaudal scales (SC),and number of infralabial scales (IL). Scales

242 Bulletin of the Museum of Comparative Zoology, Vol. 161, No. 6

were counted as suggested by McDowell(1969). Sex was determined by examinationof gonads, presence of everted hemipenes, orpresence of the retractor penis muscle.Length measurements were taken by run-ning a nonelastic string from the tip ofthe snout along the ventral medial axis ofthe body, under consideration of the pointsabout measurement accuracy raised byNatusch and Shine (2012). Measurementsof tail tips were taken using Mitutoyo digitalcalipers to the nearest 0.1 mm. X-rays ofthe holotype were taken using a KevexPXS5-724EA emitter and a Varian PanScan4030R receiver (40-kV target tube voltage) atthe Museum Support Center of the U.S.National Museum of Natural History, Smith-sonian Institution, Suitland, Maryland. Theholotype of T. grandis was X-rayed usinga Machlett X-ray tube in a Solus–Schallemitter (output settings: 55 kV, 15 mA) at theBMNH. Cranial characters were measuredon digital X-ray images in pixels using thesoftware AnalyzingDigitialImages (Museumof Science, Boston, Massachusetts). A list of

skull measurements (expanded from Dwyerand Kaiser, 1997) is presented in Table 1.Global positioning system coordinates weredetermined using individual museumrecords and published species accountswhere available; those unavailable wereobtained from Google Earth (WGS 84) tothe nearest minute. Museum acronyms aretaken from Sabaj Perez (2014), with theaddition of UPSZ, now the preferred acro-nym for the Museum of Evolution containingthe Uppsala University natural history col-lection (Mejlon, personal communication).

TOXICOCALAMUS ERNSTMAYRINEW SPECIES

Star Mountains Worm-Eating Snake

Figures 2–3, 5A, B, 6A, C

Holotype. MCZ R-145946, an adult femalefrom Wangbin village (5u14926.720S,141u15931.920E), elevation 1,468 m (4,800 ft),near the Ok Tedi River, in the Star Mountainsof the North Fly District, Western Province,

Figure 1. Satellite map showing the island of New Guinea and its surrounding islands and archipelagos. Localities for all knownmuseum specimens of Toxicocalamus are marked by white dots. The red dot indicates the locality for the holotype ofT. ernstmayri new species at Wangbin, Western Province, Papua New Guinea, whereas the yellow dot indicates the approximatetype locality of T. grandis, ‘‘launch camp’’ on the Setakwa River of Papua Province, Indonesian New Guinea. The yellow linemarks the international border between Papua New Guinea and Papua Province, Indonesia.

NEW TOXICOCALAMUS FROM PAPUA NEW GUINEA N O’Shea et al. 243

PNG, (Fig. 2), killed by a villager andcollected by FP on 23 December 1969.Originally identified and accessioned asMicropechis ikaheka Lesson, 1830.

Etymology. The species name ernstmayriis a patronym honoring the German-Amer-ican ornithologist, systematist, and evolu-tionary thinker Ernst Mayr (1904–2005).After leaving Germany in the early 1930sMayr took up a curatorial position at theAmerican Museum of Natural History inNew York. While there he wrote his de-finitive book Systematics and the Origin ofSpecies from the Viewpoint of a Zoologist(Mayr, 1942), which solved one of the bigquestions of Darwinian evolution, that ofhow one species evolved into many via themechanism of natural selection. In 1953Mayr moved to Harvard University inCambridge, Massachusetts, where he wasDirector of the MCZ from 1961–1970. Heretired in 1975 as Alexander Agassiz Pro-fessor of Zoology, Emeritus (Bradt, 2005).

There are several connections linkingErnst Mayr to this new species of Toxico-calamus, which make him, and this snake,the ideal candidates for a patronym. First,Mayr himself visited New Guinea, andduring the late 1920s he spent over 2 yearsconducting fieldwork in an area now part ofPNG, as a member of a joint Rothschild–AMNH expedition focusing on birds of

paradise (Aves, Passeriformes, Paradisaei-dae), during which he collected many newbird and orchid species. Second, the holo-type of T. ernstmayri has been housed inthe MCZ collection, mislabeled asM. ikaheka, after having arrived and beenaccessioned in June 1975, the month andyear that Mayr retired. Third, the trueidentity of this specimen was recognizedby one of us (MOS) during a visit to theMCZ in May 2014, undertaken with thefinancial support of an Ernst Mayr TravelGrant from Harvard University, awarded toenable examination of the Toxicocalamusholdings at the MCZ and the AMNH, thetwo U.S. institutions where Mayr worked.Finally, 2015, the publication year of thisdescription, marks the decennial of Mayr’spassing at age 100, and naming a NewGuinea snake after him seems a suitabletribute.

Diagnosis. Toxicocalamus ernstmayri isthe largest known species in the genusToxicocalamus and the only one with anSVL in excess of 1.0 m. It can bedistinguished from all other known Toxico-calamus by the following combination ofcharacters: large size (SVL of the holotype1,100 mm), dorsum of head exhibitingthe typical ‘‘colubrid–elapid dorsal nine-scute arrangement’’ (sensu O’Shea, 2005:12) that comprises paired internasals, paired

TABLE 1. Cranial characters measured on the holotypes of Toxicocalamus ernstmayri new species (MCZ R-145946) and T. grandis (BMNH

1946.1.18.34). Measurements were taken on the left side of paired structures and constitute maximum measurements. All measurements were

subsequently converted into ratios appropriate for comparisons (see Table 3).

Character Abbreviation Description

Skull length SL skull length from the anteriormost part of the nasal to the occipital condyleBraincase width BW broadest extent of the braincaseFrontals, total width FWPF taken at the prefrontal processFrontals, total width FWSO taken at the supraorbital ridgeParietal width ParW greatest widthSupratemporal length StL measured in a straight lineMaxilla length MxL measured in a straight lineAtlas width AtW width of the first cervical vertebra, measured across the transverse

processesAxis width AxW width of the second cervical vertebra, measured across the transverse

processesDentary–articular length D–A length of the lower jaw from the anterior tip of the dentary to the posterior

end of the articular


Figure 2. Satellite close-up maps of the region surrounding the Wangbin type locality, indicated by a red dot, of Toxicocalamusernstmayri new species, showing (A) its position on the southern versant of the Star Mountains and proximity to the Indonesianborder in relation to the Ok Tedi River and Fly River drainages, and (B) close-up view to illustrate the specific position of thelocality along the Ok Tedi River.


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al


prefrontals, a frontal between the supra-oculars, and paired parietals (Fig. 4A9), sixsupralabials, with third–fourth contactingthe orbit (Fig. 4C9, D9); dorsal scales in 15–15–15 rows; 203 ventrals, 29 subcaudals,mostly paired, a few single; divided analplate, short tail terminating with spinous,cone-shaped terminal scale (Fig. 4E9).

Comparisons. The genus Toxicocalamuscan be distinguished from all other NewGuinea elapids, including M. ikaheka butexcepting Pseudonaja textilis (Dumerilet al., 1854), by the absence of a temporo-labial scale between the fifth and sixthsupralabials (see Discussion). From P.textilis it can be differentiated by itsrelatively small eyes (diameter two-thirdsthe distance from the lower edge of theorbit to the lower edge of the upper lip) andunremarkable supraoculars, whereas thehighly active and visually alert P. textilispossesses large eyes (diameter 1.5 times thedistance from the lower edge of the orbit tothe lower edge of the upper lip), understrongly pronounced, shelved supraocularscales. Toxicocalamus ernstmayri can fur-ther be distinguished from M. ikaheka(characters in parentheses) by its short tail,equivalent to 8.3% TTL (11–14% TTL), itsventral scale count of 202 (174–191), and itslow subcaudal count of 29 (36–49). The

holotype of T. ernstmayri demonstratesa complete lack of the strongly bandedpatterning diagnostic of Papua New GuineanM. i. fasciatus (Fischer, 1884). The pattern-ing of T. ernstmayri is similar to that of thealmost patternless ‘‘yellow phase’’ M. i.ikaheka from West Papua Province, Indonesiaexemplified by the holotype of M. i. ikaheka(MNHN 7669). During the development ofanother project (O’Shea, unpublished data)we examined 132 specimens of M. ikahekafrom across the taxon’s entire range, confirm-ing the subspecific pattern differences; 15vouchers are included in Appendix 1 asexamples for these patterns.

We here compare T. ernstmayri with all12 congeners (11 species and one subspe-cies), with relevant characteristics for thesespecies given in parentheses. A more expan-sive listing of comparative characters isprovided in Table 2. Toxicocalamus ernst-mayri exhibits the regular colubrid-elapidnine-scute dorsal arrangement (Fig. 4A9)that distinguishes it from T. mintoni (fused,head-wide, frontal-supraocular scute), T.buergersi, T. preussi preussi, and T. p.angusticinctus (fused prefrontal-internasal-preocular scutes), and T. stanleyanus, T.longissimus, T. misimae, and T. mintoni (allwith fused prefrontal-preocular scutes). Thepresence of six supralabials distinguishes

Figure 3. Holotype of Toxicocalamus ernstmayri new species (MCZ R-145946) in (A) dorsal and (B) ventral view. Scale 5 5 cm.


T. ernstmayri from T. buergersi (four), andT. p. preussi, T. p. angusticinctus, and T.stanleyanus (all with five). Contact betweenthe preocular and nasal scutes separatesT. ernstmayri from T. pachysomus (inter-nasal and preocular in contact, excludingpreocular contact with nasal). Toxicocalamusernstmayri has a dorsal scale count of 15–15–15 that distinguishes it from T. p. preussi,T. p. angusticinctus (both 13–13–13), andT. longissimus (17–17–17), whereas thedivided anal plate separates it fromT. buergersi, T. preussi, T. stanleyanus, andsome Oro Province T. loriae sensu lato (analplates entire), and the mostly paired sub-caudals separate it from T. holopelturus (allsubcaudals single).

The ventral scale count of T. ernstmayri at203 is considerably lower than that of femaleT. buergersi (313–319), T. holopelturus (246–256), T. longissimus (273–304), T. preussi(291–359), and T. stanleyanus (230–281).The subcaudal scale count of 29 is signifi-cantly lower than those of female T. holo-pelturus (37–41) and T. spilolepidotus (33).

Toxicocalamus ernstmayri is longer thanany known Toxicocalamus species, with anSVL of 1,100 mm, compared with the nextlongest species: T. grandis (SVL 960 mm),T. longissimus (802 mm), T. spilolepidotus(770 mm), and T. holopelturus (760 mm).Males of all other Toxicocalamus species haveshorter SVLs. Toxicocalamus ernstmayri hasa relatively short tail (TL/TTL 5 8.3%),although females of other Toxicocalamusspecies may possess shorter tails: T. p. preussi(3.9–6.1%), T. p. angusticinctus (3.5–8.4%),T. buergersi (4.1–6.3%), T. longissimus(5.3–7.7%), T. grandis (7.7%). Its relative taillength falls within the ranges of T. loriae sensulato1 (6.5–15.6%), T. holopelturus (7.9–9.2%),

and T. stanleyanus (5.1–9.7%), and below thatof T. spilolepidotus (9.4–10.7%). Only malesare known for T. misimae, T. mintoni, andT. pachysomus, and the latter two species areknown only from holotypes with incompletetails.

The general habitus is that of a stoutsnake with a relatively broad head, distinctfrom the neck, and a short tail thatterminates abruptly. This build contrastswith most other Toxicocalamus, exceptT. grandis, and possibly T. pachysomus,which can be distinguished by head shape(distinctly pointed with small eyes), andmost particularly with the ultraslender,elongated ‘‘bootlace’’ species, T. buergersiand T. preussi.

Description of the Holotype. An adultfemale (SVL 1,100 mm, TL 100 mm, TTL1,200 mm), with 15–15–15 dorsal scalerows, imbricate, all smooth, 203 ventrals,a divided anal plate, 29 subcaudals, SC 1–6paired, SC 7–14 single, and SC 15–29paired. The head measures 21.8 mm long,from front of rostral to posterior of parietalsuture, and 19.4 mm wide, across thebroadest part of the head in line with thesuture between the fifth and sixth suprala-bials. It exhibits the normal colubrid-elapidnine-scute dorsal arrangement of pairedinternasals, paired prefrontals, frontal be-tween supraoculars, and paired parietals(Fig. 4A9). Rostral broad and high, almosttriangular and easily visible from above.Internasals small in comparison with pre-frontals (Fig. 4A9). Both internasals andprefrontals in broad contact. Frontal hexa-gonal, verging on triangular, longer thanwide because of a backward projection intothe parietals, only slightly larger than theprefrontals. Supraoculars relatively smalland not projecting, shelf-like, above theeyes. Parietals angular, at least twice as longas broad. Six supralabials, the third andfourth in contact with the orbit (Fig. 4C9,D9), the sixth being the largest. Nasal largeand divided by a suture on either side ofa relatively large naris. Loreal absent, asin all elapids; subocular absent, as in allterrestrial Papuan elapid genera except

1Based on morphological evidence (unpubl. data)and a discordant geographic distribution of availablespecimens, we believe that T. loriae, as currentlydefined, is a species complex. This is a key reason whythe range of relative tail length values shown here isconsiderably larger than for any other species in thegenus. We here refer to specimens of uncertain speciesaffinity, but currently treated as belonging to T. loriae,as T. loriae sensu lato.


Figure 4. Detailed views of the head and tail of Toxicocalamus ernstmayri new species (MCZ R-145946), presented for clarity asboth photographic and line-drawing illustrations. (A, A9) Dorsal view of the head, showing classic colubrid-elapid nine-scutearrangement (shaded), comprising paired internasals (IN), prefrontals (PF), supraoculars (SO), and parietals (P), and a singlefrontal (F). Also illustrated is the rostral (R). (B, B9) Ventral view, showing the scale arrangement of six infralabials (IL1–IL6), pairedanterior genials (AG) and posterior genials (PG), the first intergenial (IG), the mental (M), and the first ventral scute (V1). (C, C9)Left lateral view, including a separate, single preocular (PR), paired postoculars (PO), a single anterior temporal (AT), and a single


Acanthophis; temporolabial absent, in con-trast to all Papuan elapid genera exceptToxicocalamus and Pseudonaja. Nasal inbroad contact with a single elongate pre-ocular, precluding contact between thesecond supralabial and the prefrontal. Apair of small postoculars on either side, incontact with a single elongate anteriortemporal that extends backward to at leasttwice its width, to contact the fifth and sixthsupralabials, the parietal, and the posteriortemporals. Posterior temporals exist as a pair(upper small, lower large) on the right side,but as a single elongate scale on the leftside. Mental small and triangular, six infra-labials with IL 1–3 contacting pregenials(Fig. 4B9). Pregenials 1.5 times as long aswide, in broad contact along mental groove,posterior genials 1.5 times as long as wide,separated by an intergenial approximately0.5 times the size of pre- and postgenials,the intergenial forming the first in a row offive gular scales before the first ventralscute.

Scales of body large, imbricate, smooth,and distinctive, arranged in 15 rowsthroughout without posterior reduction,and without enlargement of either the firstor the midvertebral row of scales; dorsalscales of tail large and distinctive. Ventralscales broad, anal plate divided almostequally by a near-longitudinal junction, withthe left half overlapping the right half,subcaudals mostly paired, except SC 7–14,which are single; tail terminating in a large,slightly upturned conical excrescence end-ing in a slightly damaged terminal spine(Fig. 4E9).

Coloration in Life. The following de-scription is modified from Parker (1982),where this specimen was listed as part of theM. ikaheka account.

Head blackish above, lips bright yellow.Dorsum and flanks bright yellow with theanterior corner of each scale gray to black,

the vertebral row with mostly black, thenprogressively less dark pigment on each scalerow leading toward the ventrals, and with theamount of black on each scale increasingtoward the tail. Venter bright yellow.

Coloration in Preservative (45 Years Post-collection). Distinctive coloration with a yel-low-brown background color. The head isbrown above and this pigment extendslaterally to cover the rostral, preoculars,postoculars, most of the temporals, and theupper thirds of the 3rd–6th supralabials; onthe remainder of the supralabials and theinfralabials the dark pigment is confined tothe sutures, which appear highlighted. Thenasals and first two supralabials are virtuallyunmarked (Fig. 4A–D). On the yellow-brown body the darker brown pigmentmanifests as a large, dark anterior-centralspot on every dorsal scale, the posteriorportions of the scale remaining unmarkedand pale, the whole presenting an irregularreticulated appearance (Fig. 3A). Laterally,the dark pigment weakens progressively andbecomes largely confined to the anterioredge of each scale suture, although thedorsal scales of the tail are strongly markedthroughout. Ventral scales are immaculateyellow-brown, without dark markings orsuturing, but central sutures of the pairedsubcaudals are edged with brown, as a finemid-ventral zigzag (Fig. 3B). The terminalconical excrescence is immaculate yellow,in contrast to the other adjacent dorsallyvisible scales of the tail (Fig. 4E).

Tail Tip Morphology. Comparisons of thedistal portion of the tails of T. ernstmayri(Fig. 5A, B) and T. grandis (Fig. 5C, D)show that there is not only a difference inthe structure of the keratinized terminalexcrescence, but also in the dimensionsof the tail. Measurements of tail width atintervals of two to five times the length ofthe terminal excrescence (TEL) shows thatthe tail of T. ernstmayri widens more

rposterior temporal (PT). Six supralabials (SL1–SL6) are present, with SL3 and SL4 in contact with the orbit. The nasal scale (N) iselongate, measuring approximately twice as long as wide. (D, D9) Right lateral view, differing from the left view only in thepresence of two posterior temporals. (E, E9) Lateral view of the tail, showing the robust, pointed terminal scale. The scale is 3 cmfor the head views (A–D) and 5 cm for the image of the tail (E).


rapidly from the end than that of T. grandis.Whereas in T. ernstmayri, tail width dou-bles after 5 TEL, the tail of T. grandis onlyadds 70% in width at that length interval.

Osteology. We examined X-ray images ofthe skull and tail of T. ernstmayri (Fig. 6A, C)to determine the presence of diagnosticfeatures and to compare these with those ofT. grandis (Fig. 6B, D), the next largestspecies in the genus and the most similar ingeneral appearance. The skull of T. ernstmayri(Fig. 6A) appears overall slightly more robustthat of T. grandis (Fig. 6B). Easily observeddifferences include the number of maxillaryteeth (four in T. grandis and five in T.ernstmayri), the shape and dimensions of thesupratemporal, the extent of the transverseprocesses of the atlas–axis complex (axisbroader than atlas in T. ernstmayri, the reversein T. grandis), and the extent of ossifiedornamentations on the terminal vertebra.

Aside from the qualitative difference, wewere also able to contrast some characteristicsquantitatively (Table 3). In addition to the16.9% difference in the transverse dimen-

sions of the atlas–axis complex, there alsoexists a similarly large difference in thedimension of the frontal bone (15.9% inFWSO/FWPF). Smaller differences can beseen in a variety of cranial elements whencompared with skull length (e.g., MxL—10.8%, BW—9.7%, StL—9.7%, ParW—9.1%). We recognize that none of thesedifferences can be supported statisticallybecause of the availability of only singlespecimens, and they only serve to distinguishthe morphology of these bones descriptively.

Natural History. Little is known of thenatural history of any members of the genusToxicocalamus (O’Shea, 1996). They arebelieved to be fossorial, semifossorial, orterrestrial in habit, diurnal in activity, andoviparous in reproductive strategy, withclutch sizes of one to eight on record (Shineand Keogh, 1996).

Before this report, the only specimens forwhich gut content data had been reportedwere members of T. loriae sensu lato(McDowell, 1969; Shine and Keogh, 1996;O’Shea, personal observation). Numerous

Figure 5. Tails of Toxicocalamus ernstmayri new species (A, B) and T. grandis (C, D). Tails are shown in ventral (A, C) andlateral (B, D) views. The tail tip of T. ernstmayri curves downward in lateral view (B) and is slightly asymmetrical in ventral view (A),with evidence of minor breakage at the tip. In contrast, the tail tip of T. grandis is cone-shaped in ventral view (C) and does notcurve as in T. ernstmayri.


Figure 6. X-rays of the skulls (A, B) and tails (C, D) of Toxicocalamus ernstmayri new species (A, C) and T. grandis (B, D; BMNH1946.1.18.34). Key features for comparison include the shape and length of the supratemporal bone (St) and the atlas–axiscomplex (A–A). The terminal vertebra of T. ernstmayri (C) shows a significant amount of ossified ornamentation that is all butabsent in the tail of T. grandis (D).


earthworms or earthworm fragments werereported from the guts of T. loriae speci-mens from the Waghi Valley, Simbu Prov-ince, including five specimens from Kondiuwith earthworm fragments (AMNHR-75336–38, 75341, 75347), and a Kundiawaspecimen (MCZ R-111785) with a large,complete earthworm protruding from itsmouth and extending to its posterior gut.Elsewhere, a specimen of T. loriae sensulato from Aguan, Milne Bay Province(BPBM 10967), and another from Mara-tambu in the Adelbert Mountains, MadangProvince (AMNH R-82332) also containedcomplete earthworms, whereas the pres-ence of soil and plant fibers in the guts offour additional Kondiu specimens (AMNHR-75340, 75343, 75345, 75348) was consid-ered evidence of fully digested earthworms(McDowell, 1969).

Given the relative large size and lack ofstomach contents for some Toxicocalamusspecies (e.g., T. spilolepidotus with TTL 850mm, T. grandis with TTL 940 mm), it maybe difficult to believe that they haveadopted vermivorous lifestyles. In referenceto T. spilolepidotus, McDowell (1969)wrote, ‘‘The only known specimen of T.spilolepidotus2 is about the length andgirth of an average mature female NorthAmerican watersnake (Natrix sipedon) and Ifind it hard to believe that a snake this large

could have the burrowing and earthworm-eating habits of T. loriae.’’ And in referenceto T. grandis, he wrote, ‘‘As in the case ofT. spilolepidotus, it seems unlikely that solarge a snake feeds primarily on earth-worms, but its food preferences are quiteunknown.’’ It is therefore interesting thatthe holotype of T. ernstmayri, now thespecies in the genus with the largest bodysize, contains a large earthworm (see below)and no other gut contents other thana parasitic nematode, demonstrating thateven the larger members of the genusappear to feed on oligochaete worms.

McDowell (1969) also reported findinga small snail in the gut of one Waghi Valleyspecimen (AMNH R-75341), whereas a sar-cophagid fly larva was retrieved from thegut of a specimen (AMNH R-59067) fromMafulu, Owen Stanley Range, CentralProvince (Bogert and Matalas, 1945). Stern-feld (1913) reported several peculiar ‘‘ma-denartige Tiere’’ [maggot-like animals] oflength 4 cm, which he denoted as ‘‘Fliegen-maden?’’ [fly larvae?], in the gut of a spec-imen from Sattelberg, Huon Peninsula,Morobe Province. We believe it is mostlikely that the sarcophagid fly larvae repre-sent postmortem artefacts, resulting froma delay between euthanasia and preserva-tion of historical specimens in the field, andthat the ‘‘small land snail’’ found byMcDowell (1969) may have been ingestedinadvertently. On the basis of the unusualsize of Sternfeld’s ‘‘maggots,’’ we consider itpossible that these were instead megascole-cid earthworm cocoons. Subsequently wehave also learned that a captive specimen ofT. longissimus was induced to eat earth-worms, while rejecting all other potentialprey offered (Paiva, personal communica-tion), and the holotype of T. pachysomus(TTL 715 mm) also contained an earthwormin its gut (Kraus, personal communication),although this fact was not reported in theoriginal description (Kraus, 2009).

The Australasian region is inhabited bythe terrestrial megadrile oligochaete familyMegascolecidae, which contains the world’slargest earthworm species. Of 115 earth-

TABLE 3. Comparisons between cranial characteristics of Tox-

icocalamus ernstmayri new species (MCZ R-145946) and T. grandis

(BMNH 1946.1.18.34) using ratios of measurements taken from

digital X-ray images. Abbreviations are explained in Table 1.

Ratio T. ernstmayri T. grandis Difference (%)

ParW/SL 0.22 0.24 9.1FWSO/FWPF 0.63 0.73 15.9FWPF/SL 0.21 0.21 0.0MxL/SL 0.37 0.33 10.8StL/SL 0.31 0.34 9.7

2Two female specimens of T. spilolepidotus (AMNHR-85745, PNGM 21332) are now known, from Purosa,near Okapa, and the Yaiya Valley in the KratkeMountains, respectively, both localities in EasternHighlands Province, PNG, at elevation approximately1500–1740 m.


worm species recorded from New Guinea,109 belong to the Megascolecidae (Blake-more, 2006). The holotype contained a sin-gle large megascolecid earthworm (Fig. 7),likely of the genus Pheretima Kinberg, 1867(Blakemore, in litteris), now accessioned inthe MCZ annelid collection (MCZ 95752).The worm measured approximately 500 mmwhen removed upon examination of thefreshly killed snake (Parker, 1982), and it isnow 436 mm long after decades in pre-servative.

The holotype was also parasitized bya relatively large (total length 55 mm)nematode worm (Fig. 8), which has notyet been further identified.

All members of the genus Toxicocalamusare venomous. Toxicocalamus buergersipossesses extremely elongate venom glandsthat extend deep into the body cavity toextend posteriorly to almost midbody(McDowell, 1969; O’Shea, personal obser-vation); McDowell (1969) remarked, ‘‘Thevenom gland of T. grandis is fully as large asthat of the average New Guinea death adder(Acanthophis).’’ Recent investigations intothe venom composition of T. longissimus byCalvete et al. (2012) have demonstrated thatthis venom comprises primarily (92%)three-finger toxins (3FTx), combined with

smaller quantities of type-I phospholipaseA2 (PLA2) and snake venom metalloprotei-nase (SVMP). 3FTxs are key componentsfound in the neurotoxins, cardiotoxins, andplatelet inhibitors of some of the world’smost highly venomous snakes (e.g., Kini andDoley, 2010) and they are the maincomponents in the venoms of elapids(Utkin, 2013). PLA2 toxins are also commonin the neurotoxins of elapid venoms thatcause neuromuscular paralysis, whereasSVMPs cause hemorrhaging. Nothing isknown about the venom composition ortoxicity of T. ernstmayri, but given its largesize, live specimens should be treated withrespect and handled with caution.

DISCUSSION

The type locality, Wangbin (5u14926.720S,141u15931.920E), lies in the Star Mountains,which are located in the North Fly Districtof Western Province, PNG, where theprovince borders Sandaun (formerly WestSepik) Province to the north, and PapuaProvince, Indonesia, to the west (Fig. 2A).The Ok Tedi River (Fig. 2) is a northerntributary of the 1,050-km-long Fly River,the second longest river in New Guinea,which flows south and southeast into theGulf of Papua. The North Fly District isa mountainous, heavily forested region,except for the ecologically disturbed anddeforested immediate vicinity of the Ok

Figure 8. Unidentified nematode parasite (total length 5 55mm) found loose in the body cavity of Toxicocalamusernstmayri new species. We consider it likely that the parasitewas dislodged when the earthworm was removed from thesnake, but remained undetected inside the specimen.

Figure 7. Two views of a megascolecid earthworm (likely ofthe genus Pheretima Kinberg, 1867; total length 436 mm)found in the stomach of Toxicocalamus ernstmayri newspecies. Scale 5 5 cm.


Tedi Mine, an open-pit gold and coppermine with significant economic potentiallocated at the site of the former Mt. Fubilan(see Jackson, 1982; Doucette 2000), 10 kmNW of Tabubil on the Ok Tedi River(Fig. 2B). Olsobip (Fig. 9G) in the StarMountains is believed to be one of thewettest places on Earth, with an annualrainfall of up to 10 m (McKinnon et al.,2008:206). Wangbin is a village located closeto the northern shore of Lake Wangbin(Fig. 9C), approximately 5 km NE ofTabubil and 13 km ESE of the Ok TediMine (Fig. 2B). In 1969, the year theholotype of T. ernstmayri was collectedand before the development of the Ok TediMine and Tabubil (Fig. 9H), Wangbin wasa small village (Fig. 9A, B) surrounded byextensive primary mid-montane forest withsome garden areas (Fig. 9D, E). At thetime, gardens were established in partiallycleared areas, with smaller trees and un-dergrowth being removed but with largertrees left standing (Fig. 9E). The presenceof large trees may have reduced soil erosionin gardens established on slopes, an impor-tant factor affecting the environment in anarea experiencing heavy rainfall.

The holotype of T. ernstmayri wascollected by one of us (FP) during hisemploy as a kiap3 (patrol officer) while onpatrol in the North Fly District (Fig. 9F).Other elapids recorded from the Ok TediRiver and Star Mountains region of WesternProvince include Acanthophis laevis Ma-cleay, 1878, Aspidomorphus muelleri Schle-gel, 1837, M. ikaheka, T. p. angusticinctus,and T. stanleyanus. A specimen of T. loriaesensu lato was collected at Busilmin across

the provincial border in Sandaun (WestSepik) Province, but still in the StarMountains (Fig. 2A).

Australopapuan elapids primarily prey up-on vertebrates, so Toxicocalamus is perhapsunique among this elapid species assemblagein its diet of earthworms. It appears to be anespecially esoteric dietary choice for a snakethat achieves an SVL of over 1.0 m. Althoughmany Toxicocalamus exhibit a morphologythat suggests a fossorial or semifossorialexistence in keeping with this dietary prefer-ence, T. ernstmayri is morphologically similarin appearance to M. ikaheka, with which it wasinitially confused, suggesting a more active,terrestrial lifestyle.

The loss of the temporolabial scale inToxicocalamus, and the subsequent reduc-tion in the number of supralabials, isprobably a derived condition within theElapidae. Greer (1997: fig. 7.22) provideda series of head drawings that illustrate thistransition, beginning with the primitivecondition of seven supralabials as in Bun-garus candidus (Linneaus, 1758). An up-ward movement of the penultimate supra-labial, becoming a temporolabial scale as if‘‘squeezed’’ by the supralabials on eitherside, as seen in Pseudechis australis (Gray,1842), is hypothesized to precede conditionsinvolving the fusion of the temporolabialwith the anterior temporal, a characteristicof Vermicella annulata (Gray, 1841), or withthe last, now sixth, supralabial, such as inPseudonaja affinis Gunther, 1872, leading tothe eventual reduction to five supralabials,as seen in Simoselaps bimaculatus (Dumerilet al., 1854). All members of the genusToxicocalamus lack a temporolabial scaleand mirror the condition in P. affinis, withthe exception of T. preussi and T. stanleya-nus, which possess five supralabials andmirror the condition in N. bimaculatus, andT. buergersi, which exhibits a further re-duction to four supralabials. Toxicocalamuspreussi, T. stanleyanus, and T. mintonidemonstrate fusion of the preocular andprefrontal scutes; T. buergersi exhibitsfusion of the preocular, prefrontal, andinternasal scutes; whereas T. mintoni also

3Kiap is a pidgin word, derived from the Germanword Kapitan, which was applied to Australiangovernment patrol officers in the pre-independenceera. They covered considerable distances to deal withland disputes, and to apprehend murderers andcannibals to take before the courts. Fred Parker, co-author on this paper, was a kiap in PNG from 1960–73.His interest in herpetology caused him to makeextensive collections wherever he patrolled. He de-posited many specimens in MCZ, AMNH, and CAS,including the holotype of T. ernstmayri.


exhibits fusion of the frontal scute with thesupraoculars on either side, resulting ina single head-wide scute. Bogert and Porter(1966) suggested that fusion of head scutesmight be associated with a fossorial exis-tence, whereby a reduction in the numberof sutures between head scutes reduces thedegree of friction for the burrowing snake.In obligatorily fossorial species fusion ofscutes may also mirror underlying skullbone development, resulting in an overallstrengthening of the skull. These further-derived characters are not present in T.ernstmayri, which belongs to an advancedelapid genus but might be consideredrelatively basal within that genus (Metzgeret al., 2010; Pyron et al., 2013). Unfortu-nately, the age of the specimens available tous for this comprehensive study precludetheir inclusion in molecular data sets tofurther elucidate the position of individualspecies within Toxicocalamus. As it stands,however, even the few studies available toinclude Toxicocalamus in a molecular frame-work (e.g., Keogh, 1998; Scanlon and Lee,2004; Sanders et al., 2008; Metzger et al., 2010;Pyron et al., 2013) show that the position of thegenus within the Elapidae, and even itsmonophyly, is still inconclusive.

On the basis of overall cranial morphol-ogy, the skulls of both T. ernstmayri and T.grandis are fairly kinetic with a relativelysolid, elongated core. This morphology isperhaps ideally suited to a terrestrial orsemifossorial lifestyle, and to capture earth-worms with the necessary jaw pressure tosubdue them. In the absence of X-rays forthe other species of Toxicocalamus, there isunfortunately very limited utility in our dataset for broader comparisons. McDowell(1969) did include some skull characteristicsin generic and subgeneric definitions, butabsent a more comprehensive data set anda description of assessment methods, we areunable to expand on our osteologicalcomparisons at this time.

The ornamentation on the terminal verte-bra was an unexpected feature we discoveredwhen trying to determine whether the tail

was truncated or entire. Given the muchlesser degree of ornamentation in T. grandis,we wish to point out the potential diagnosticvalue in the morphology of the terminalcaudal vertebra. External aspects of tailtermination have been used traditionally asstand-out features to define unusual taxa,including some uropeltids with keratinizedtail plates such as Uropeltis melanogaster(Gray, 1858), or the ‘‘spider tail’’ of Pseudo-cerastes urarachnoides Bostanchi et al., 2006;these do not reflect the internal constructionof the tail terminus. X-rays allow a detailedstructural analysis of the terminal vertebra,and perhaps this is a useful, heretoforeoverlooked source of taxonomic information.

Ornamentation of the terminal vertebrabrought into our focus the detailed externalmorphology of Toxicocalamus tails. In thisgenus, the tail tip consists not of scales as inmost other snakes of this size, but ofa terminal excrescence in the form ofa keratinized spine or nub. The shape anddimensions of this structure appear to beuseful as an additional character for distin-guishing species, in addition to providinga tool for comparative measurements of theexpansion of the tail thickness forward fromits end. The morphometric approach to thischaracter allowed us to quantify the state-ment that T. ernstmayri has a tail thatremains relatively thick all the way towardits end, whereas by comparison T. grandishas a more slender tail. Cursory inspectionof the tail of a specimen of T. loriae sensulato (AM R9351, the holotype of Apistho-calamus lamingtoni Kinghorn, 1928) showsa very thin tail, in which the width remainsvery narrow, at only 50% greater than at thebeginning of the sharply pointed excres-cence. The tails of T. buergersi (AMR120360), T. misimae (AM R7614), and T.longissimus (AM R124904) are essentiallynubs. As we continue our investigations intothe taxonomy of the genus Toxicocalamus,we intend to pay close attention to thedimensions of tails to determine whetherthey prove useful in field identification.


Figure 9. Photographs of Wangbin and Star Mountains environs, all taken on 24 December 1969, the day after the holotype ofToxicocalamus ernstmayri new species was collected, unless otherwise stated. We are presenting these images not only to illus-trate the natural environment of T. ernstmayri, but also to provide a historical and social context for potential human-snakeinteractions. (A) The main path through Wangbin Village, which in 1969 was merely a hamlet. (B) Close-up of one of Wangbin’sresidences, showing details of access, construction, and wood storage. (C) Wangbin Lake, a small catchment surrounded by pristinerainforest. (D) The Wangbin guesthouse built by villagers in a clearing away from the main village in otherwise pristine rainforest.


KEY TO THE GENUS TOXICOCALAMUS

The following dichotomous key is a revision ofthe keys provided by McDowell (1969), O’Shea(1996), and Kraus (2009), expanded to incorpo-rate T. ernstmayri as well as some othercharacters that came to light during the exami-nation of a more comprehensive list of specimensthan used by previous authors.

1a Preocular distinct from prefrontal;always six supralabials .................. 2

1b Preocular fused with prefrontal;six or fewer supralabials .............. 7

2a Subcaudals entire ..... T. holopelturus2b Subcaudals all or mostly divided ... 33a Distinctive patterning, consisting of

a yellow central spot on everychocolate scale on head and body,except on the vertebral row; ventralscales yellow with chocolate brownsuturing ................. T. spilolepidotus

3b Patterning not as in 3a ............... 44a Habitus generally slender; size

,700 mm TTL; patterning, ifpresent, includes yellow suprala-bials and frequently a yellowcrescent-shaped marking on thenape that fades dorsally, but inspecimens where it meets mid-dorsally there may also be palecross-bars on the parietals andprefrontals ........ T. loriae sensu lato

4b Habitus robust; size .700 mmTTL; patterning not as above ..... 5

5a Internasal and preocular in con-tact, separating nasal from pre-frontal; head tapering to marked-ly pointed snout ...... T. pachysomus

5b Internasal and preocular not incontact, separated by contactbetween nasal and prefrontal;

head rounded, not tapering topointed snout ............................ 6

6a Postoculars single; point contactonly between preocular and nasal;sixth supralabial widely separatedfrom upper posterior temporal bylarge anterior temporal; pattern-ing irregular, comprising scatteredyellow spots on a brown dor-sum ........................................... T. grandis

6b Postoculars paired; broad contactbetween preocular and nasal;large sixth supralabial only nar-rowly separated from upper pos-terior temporal by narrow anteri-or temporal (posterior temporalsfused on left side in the holo-type); regular patterning compris-ing brown anterior and yellowposterior to every dorsal scale,presenting a reticulate pat-tern ............. T. ernstmayri sp. nov.

7a Anal plate divided; six suprala-bials .......................................... 8

7b Anal plate entire; fewer than sixsupralabials ............................... 10

8a 17 scale rows at midbody ...........T. longissimus

8b 15 scale rows at midbody ........... 99a Supraoculars fused with fron-

tal ................................ T. mintoni9b Supraoculars distinct from fron-

tal ................................ T. misimae10a Internasal fused with prefrontal-

preocular; anterior temporal ab-sent, allowing contact betweenfinal supralabial and parietal ..... 11

10b Internasal separate and distinctfrom prefrontal-preocular; anteri-or temporal present, preventingcontact between sixth supralabialand parietal ............. T. stanleyanus

r(E) Taro garden established in partially cleared forest with large trees left standing, located along the track fromTungenabip to Wangbin (22 December 1969); figures in right foreground for scale. (F) Kiap patrol taking a rest alonga river, having left Wangbin for Migalsimbip. Photo by Patrol Officer Mike Bell (FP is standing near the middle of thephoto, fourth from left). (G) Crossing a cane bridge en route to Olsobip (26 December 1969). (H) Tabubil ‘‘base camp’’on 8 June 1972. Tabubil is now the largest town in the North Fly District, with a 2005 population of ca. 13,800. Allphotographs by FP, unless otherwise credited.


11a 13 scale rows at midbody; fivesupralabials; postocular pre-sent ............................... T. preussi

11b 15 scale rows at midbody; foursupralabials; postocular fusedwith supraocular ......... T. buergersi

ACKNOWLEDGMENTS

The authors thank Jose Rosado and JoeMartinez for their generous assistance whileworking at the MCZ. We are very grateful toPatrick Campbell (BMNH) for providing uswith the first set of X-rays for T. grandis,Adam Baldinger (MCZ), for locating andphotographing the megascolecid earthwormpredated by T. ernstmayri, Nicholas Grif-fiths and Pumehana Imada (both BPBM)for extricating and photographing the earth-worm in the holotype of T. pachysomus,Allen Allison (BPBM) for granting permis-sion to dissect the specimen, and RobertBlakemore (College of Natural Science,Hanyang University, Seoul, South Korea)for identifying the earthworms found in gutsof Toxicocalamus specimens. MOS alsoacknowledges with thanks the hospitality ofVan Wallach and Roxane Coombs duringhis stay in Cambridge, and Christine, Lucas,Max, and Isabel Kaiser during his stay in theWashington, DC area. Finally, thanks to thefollowing collection managers and museumcurators, in addition to those alreadynamed, for permitting access to theircollections or arranging loans: Ross Sadlier(AM), David Dickey and David Kizirian(AMNH), Pumehana Imada (BPBM), AlanResetar (FMNH), Ivan Ineich (MNHN),Irvan Sidik (MZB), Heinz Grillitsch(NMW), Bulisa Iova (PNGM), EstherDondorp (RMNH), Carolyn Kovach(SAMA), Emily Braker (UCM), Paulus Keip(UPNG), Hans Mejlon (UPSZ), JeremyJacobs (USNM), and Frank Tillack (ZMB).Our thanks also go to colleagues who haveassisted with the documenting, scale count-ing, and measuring of specimens: LukasHartmann, Julious Jacobs, Christine Kaiser,Max Kieckbusch, Keliopas Krey, SvenMecke, and Øyvind Syrrist. We are also

grateful to Venancio Lopes Carvalho forpreparing the Bahasa Indonesian abstract.Finally, the authors thank the five (!)anonymous reviewers whose extremelyhelpful and constructive comments un-doubtedly improved the quality of thismanuscript. Funding for the visit by MOSto the MCZ, AMNH, and USNM wasprovided by an Ernst Mayr Travel Grantfrom Harvard University.

APPENDIX 1. SPECIMENS EXAMINED

Micropechis ikaheka (N 5 15).Indonesia: West Papua Province: Dorey (5Manokwari), E Vogelkop Peninsula, elev. 70 m[00u529S, 134u059E], MNHN 7669 (holotype).Manokwari, E Vogelkop Peninsula, elev. 70 m[02u559S, 132u179E], RMNH.RENA 5585,47926. Andai, E Vogelkop Peninsula, elev. 35m [00u559S, 134u009E], BMNH 1878.2.11.48A–B. Aitinjo, W Vogelkop Peninsula, elev. 208 m[01u259S, 132u039E], RMNH.RENA 45779. FakFak Regency, Bomberai Peninsula, elev. 380–520 m [02u559S, 132u179E], BMNH1909.4.30.12. Misool Is., Raja Ampat Islands,elev. 200 m [01u559S, 130u059E], RMNH.RENA5585, 47926 (ex 5585). Papua Province: MimikaRiver, south coast, elev. 30 m [04u349S,136u409E], BMNH 1913.10.31.220. Biak Island,Schouten Islands, Cenderawasih Bay, elev. 100m [00u589S, 135u579E], RMNH.RENA 45799.Hollandia (5 Jayapura) northeast coast, elev.175 m [02u329S, 140u429E], AMNH 62480.Papua New Guinea: Sandaun Province: Aitape,elev. 40 m [03u089S, 142u209E], MCZ 48624.Madang Province: Kaviak Plantation, KarkarIsland, elev. 30 m [04u359S, 145u559E],NHMUK 2013.283. Oro Province: Itokama, elev.800 m [09u129S, 148u159E], BPBM 36148.

Toxicocalamus buergersi (N 5 4). PapuaNew Guinea: East Sepik Province: Wewak,elev. 200 m [03u409S, 143u059E], AMNH R-75001, 75236. ‘‘Deutsch Neuguinea’’ [GermanNew Guinea], ZMB 25232 (Holotype). Maprik[03u399S, 143u029E], AM R120360.

Toxicocalamus ernstmayri (N 5 1). PapuaNew Guinea: Western Province: Wangbin, elev.1468 m [5u149S, 141u169E], MCZ R-145946(holotype).

Toxicocalamus grandis (N 5 1). Indonesia:Papua Province: Launch camp, Setakwa River,elev. 25 m [04u459S, 137u209E], BMNH1946.1.16.34 (holotype).


Toxicocalamus holopelturus (N 5 18). PapuaNew Guinea: Milne Bay Province: Mt. Rossel,Rossel Is., Louisiade Archipelago, elev. 205–700m [11u209S, 154u139E], AMNH R-76660 (holo-type); BPBM 20823–33; PNGM 25194. Cheme,Rossel Is., Louisiade Archipelago, elev. 55 m[11u199S, 154u149E], BPBM 20835–36. Mt.Yuvu, Rossel Is., Louisiade Archipelago, elev.450 m [11u229S, 154u069E], MCZ R-156548.Gobubob, Rossel Is., Louisiade Archipelago,elev. 336 m [11u219S, 154u099E], PNGM 25193.

Toxicocalamus longissimus (N 5 11). PapuaNew Guinea: Milne Bay Province: Woodlark Is.[09u139S, 152u569E], BMNH 1946.1.18.92–93(holotype, paratype). Guasopa, Woodlark Is.,elev. 12 m [09u139S, 152u569E], AM R124904,BPBM 17885–87. Kulumandau, Woodlark Is.,elev. 53 m [09u059S, 152u439E], AMNH R-76619, 76629–30, BPBM 42185. Suloga Har-bour, Woodlark Is., elev. 80 m [09u129S,152u449E], BPBM 17888. Waimunon Bay, Woo-dlark Is., elev. 72 m [09u049S, 152u509E], BPBM39702. Dikoias, Woodlark Is., elev. 73 m[09u029S, 152u459E], BPBM 42183–84. ‘‘Fergus-son Is.’’ (possibly in error) BMNH 1904.11.1.60.

Toxicocalamus loriae (N 5 212). Papua NewGuinea: Central Province: Dinawa, Owen Stan-ley Range, elev. 1,220 m [08u359S, 145u559E],BMNH 1946.1.14.53 (holotype of Apistocalamuspratti). Mafulu, Owen Stanley Range, elev. 1,270m [08u309S, 147u049E], AMNH R-59067;BMNH 1935.5.10.174. Fane to Bellavista road,Owen Stanley Range, elev. 1,065 m [08u329S,147u059E], PNGM 23158. Tapini, Owen StanleyRange, elev. 950 m [08u219S, 146u599E], USNM195619. Owers Corner, Owen Stanley Range,elev. 620 m [09u219S, 147u299E], MCZ R-150803. Siruohu, Mt. Obree, elev. 1,550 m[09u269S, 140u009E], BPBM 19603–04. Laronu,elev. 900 m [09u269S, 147u599E], BPBM 19502,19505-06. Milne Bay Province: Bunisi, Mt.Simpson, elev. 1,490 m [10u019S 149u369E],BPBM 17989, 18164, 18166. Siyomu, Mt.Simpson, elev. 1,300 m [10u019S, 149u359E],BPBM 17987–88, 18165, 30638. Agaun, Mt.Dayman, elev. 1,014 m [09u559S, 149u209E],PNGM 24649A–B; UPNG 4840. Dumae Creek,W. Agaun, Mt. Dayman, elev. 1,200 m [09u539S,149u199E], BPBM 10966–67. Bonenau, elev.1,340 m [09u539S, 149u249E], UPNG 7105,7107–08. Mai-iu River, elev. 428 m [09u419S,149u169E], UPNG 3526. Basima, Fergusson Is.,d’Entrecasteaux Archipelago, elev. 10 m[09u289S, 150u509E], BPBM 16544. Oya Waka,

Fergusson Is., d’Entrecasteaux Archipelago,elev. 990 m [09u279S, 150u339E], BPBM 16545.Oro Province: Umwate, Itokama, elev. 943 m[09u169S, 148u169E], BPBM 43028–29, 43032.Akupe camp, Itokama, elev. 723 m [09u179S,148u169E], BPBM 43027. Mt. Trafalgar, CapeNelson, elev. 187 m [09u139S, 149u099E], BPBM39813. Isurava, Kokoda Track, elev. 1,260 m[08u599S, 147u449E], BPBM 44892. Eora Creek,Kokoda Track, elev. 1,800 m [00u029S,147u449E], BPBM 44892. Kokoda Track [n/a]PNGM 22762. ‘Popondetta’ (dubious), elev. 106m [08u469S, 148u149E], AMNH R-111810, MCZR-141009. Mt. Lamington [08u569S, 148u109E],AM R9351 (holotype of Apisthocalamus laming-toni). Morobe Province: Garaina, elev. 770 m[07u539S, 147u089E], MCZ R-152432. Saurere,W. Garaina, elev. 1,440 m [07u559S, 147u059E],PNGM 22767. Saiko, elev. 1,870 m [07u579S,147u039E], BPBM 41381, 41390. Amu Creek,elev. 1,660 m [07u559S, 147u029E], BPBM41391. Wau and environs, elev. 1,220–2,000 m[07u209S, 146u439E], BPBM 17173, 17417,17451–52, 18217, 23669, PNGM 24716. KaloloCreek, elev. 965 m [07u189S, 146u439E], AMNHR-1422887, BPBM 5440, 5442. Hewieni, KaribaRiver, elev. 1,050 m [07u259S, 146u259E], BPBM17423. Aseki, Watut Valley, elev. 1,640 m[07u209S, 146u109E], BPBM 6497. Kwaplalim,W. Menyamya, Tauri River, elev. 1,500 m[07u119S, 145u589E], UCM 51552–53, NMW37670. Zenag, Markham Valley, elev. 940 m[06u579S, 146u379E], AMNH R-85744. Sattel-berg, Huon Peninsula, elev. 800 m [06u299S,147u479E], BMNH 1946.1.17.57 (cotype ofPseudapistocalamus nymani), MCZ R-76627–28, UPS 2387 (cotype of P. nymani). MasbaCreek, Huon Peninsula, elev. 760 m [06u299S,147u319E], AMNH R-95579-80. Mt. Rawlinson,Huon Peninsula, elev. 1340 m [06u319S,147u169E], AMNH R-95581. Pindui and envi-rons, Huon Peninsula, elev. 790 m [06u179S,147u189E], AMNH R-95578, 95582. Tuwop,Sarawaget Range, Huon Peninsula, elev. 1,350m [06u219S, 146u559E], BPBM 3397, 3399.Lialun, Huon Peninsula, elev. 116 m [06u069S,147u369E], ZMB 24343–44, 78770–71. MadangProvince: Astrolabe Bay, elev. 116 m [05u229S,145u399E], NMW 27383.1–2. Wanang, elev. 120m [05u159S, 145u169E], BPBM 31257. KalneRiver, Bismarck Range, elev. 1,200 m [05u319S,144u499E], UPNG 8695. Kaironk, SchraderValley, elev. 1,200 m [05u149S, 144u299E],UPNG 963–67, 3353, 5012. Matatambu, Adel-bert Range, elev. 1,140 m [04u369S, 145u549E],


UPNG 8695. E. Highlands Province: Wonenara,Yaiya Valley, elev. 1,650 m [06u489S, 145u539E],UPNG 1213–18. Agakamatasa, elev. 1,720 m[06u439S, 145u379E], MCZ R-121545. Nivi Un-ggai, elev. 2,030 m [06u139S, 145u189E], MCZ R-84142, 116791–92. Lufa, Mt. Michael, elev.1,120 m [06u209S, 145u159E], MCZ R-121546.Simbu Province: Igindi, Suai Mts., elev. 1,630 m[06u119S, 144u589E], AMNH R-98134. Kebil,Wahgi Valley, elev. 1,944 m [06u109S, 145u019E],MCZ R-85050. Kundiawa, Wahgi Valley, elev.1,585 m [06u019S, 144u589E], AMNH R-98495–98; MCZ R-83218, 84026, 84143–44, 111764–90,115586, 116774–88, 123883–86, 140818–19,145923; PNGM 864, 882, 24585–86; SAMAR9526; USNM 166280. Mintima, Wahgi Valley,elev. 1,830 m [05u589S, 144u559E], MCZ R-116789–90. Poral Ras, Wahgi Valley, elev. 1,770m [05u599S, 144u559E], MCZ R-121547–48.Kondiu, Wahgi Valley, elev. 1,600 m [05u589S,144u529E], AMNH R-75336–57. Kup, WahgiValley, elev. 1,500 m [05u579S, 144u489E],AMNH R-72780–81. Jiwaka Province: Minj,Wahgi Valley, elev. 1,560 m [05u529S,144u409E], MCZ R-141849. Banz, Wahgi Valley,elev. 1,650 m [05u479S, 144u379E], AMNH R-85743, 88060. W. Highlands Province: Dobel,Mt. Hagen, elev. 1,705 m [05u519S, 144u149E],PNGM 22160; UPNG 3992. S. HighlandsProvince: Mendi, elev. 1,750 m [06u089S,143u399E], MCZ R-121543–44. Halalinja, WagaRiver, elev. 2,140 m [06u099S, 143u229E],BMNH 1976.92. Nipa, elev. 2,070 m [06u069S,143u159E], UPNG 5811. Sandaun Province:Busilmin, Star Mts., elev. 1,880 m [04u559S,141u129E], SAMA R6275. ‘‘Neuguinea’’ NMW27382.1–2. ‘‘Deutsch Neuguinea’’ [German NewGuinea], ZMB 23533. Indonesia: W. PapuaProvince: Fak Fak Peninsula, elev. 518 m[02u579S, 132u359E], BMNH 1946.1.18.24–26;MCZ R-76634 (holotype and paratypes ofApistocalamus loennbergii). ‘‘New Guinea’’ (nofurther locality data), MZB 361.

Toxicocalamus mintoni (N 5 1). Papua NewGuinea: Milne Bay Province: Mt. Riu, SudestIs., Louisiade Archipelago, elev. 410 m [11u299S,153u259E], BPBM 20822 (holotype).

Toxicocalamus misimae (N 5 3). Papua NewGuinea: Central Province: Mekeo region[09u009S, 146u509E], AM R7614. Milne BayProvince: Mararoa, Mt. Sisa, Misima Is., Loui-siade Archipelago, elev. 350 m [10u399S,152u489E], AMNH R-76684 (holotype). Bagi-

lina, Misima Is., Louisiade Archipelago, elev. 128m [10u399S, 152u489E], BPBM 17231.

Toxicocalamus pachysomus (N 5 1). PapuaNew Guinea: Milne Bay Province: Gadowalai,Cloudy Mts., elev. 715 m [10u299S, 150u149E],BPBM 15771 (holotype).

Toxicocalamus preussi (N 5 42). Papua NewGuinea: Simbu Province: Crater Mt., elev. 2,325m [06u359S, 145u059E], BPBM 17449. Haia,elev. 908 m [06u439S, 145u059E], USNM562943–44. Gulf Province: Wabo, elev. 400 m[06u549S, 145u039E], USNM 562942. Koni,Purari River, elev. 76 m [07u159S, 145u209E],MCZ R-102176. Middletown, Kikori River, elev.63 m [07u169S, 144u099E], MCZ R-59090. NWKikori, Kikori River, elev. 37 m [07u109S,144u059E], MCZ R-150804. Western Province:S. Palmer Junction, Strickland River, elev. 65 m[05u549S, 141u329E], AMNH R-57511-12 (holo-type and paratype of Ultrocalamus preussiangusticinctus). Matkomrae, Ok Tedi River,elev. 82 m [05u499S, 141u099E], MCZ R-121551, 141008. Ningerum, Ok Tedi River, elev.86 m [05u419S, 141u089E], MCZ R-140989,141001; USNM 217500. Tabubil, Ok Tedi River,elev. 828 m [05u159S, 141u129E], UPNG 5665.Olsobip, Fly River, elev. 612 m [05u239S,141u319E], PNGM 24584; UPNG 1613. San-daun Province: Berlinhafen, Seleo Is., elev. 7 m[03u089S, 142u299E], ZMB 23948 (holotypeUltrocalamus preussi). Mt. Sapau, TorricelliRange, elev. 1,300 m [03u229S, 142u319E],BPBM 23456. Njao, Tjano River, elev. 450 m[02u489S, 141u009E], ZMA 17733. Indonesia:Papua Province: Arso, Tami River, elev. 53 m[02u569S, 140u479E], ZMA 17734B. Hollandia(5 Jayapura), elev. 100 m [02u5329S, 140u429E],FMNH 43030; ZMA 17735. Upper SermowaiRiver, elev. 95 m [02u299S, 139u509E], UIMNH19199; ZMA 17736A–C. Etik, Biri River, elev.140 m [02u179S, 139u089E], MZB 5090; ZMA17734A. SW Bernard Camp, Idenberg River,elev. 800–850 m [03u309S, 139u099E], AMNH R-62469–72. Prauwenbivak, Idenberg River, elev.40 m [02u259S, 139u109E], MZB 354, 5089, ZMA17731. Albatrosbivak, Mamberamo River, elev.40 m [02u099S, 137u529E], MZB 356, 5054–56.Pionierbivak, Mamberamo River, elev. 10 m[02u049S, 137u509E], ZMA 17732. MamberamoRiver, elev. 20 m [01u559S, 137u509E], MZB 353.

Toxicocalamus spilolepidotus (N 5 2). PapuaNew Guinea: E. Highlands Province: Purosa,Okapa, elev. 1,740 m [06u409S, 145u339E],AMNH R-85745 (holotype). Yaiya Valley, Kratke


Mts., elev. 1,500 m [07u009S, 145u499E], PNGM21332.

Toxicocalamus stanleyanus (N 5 31). PapuaNew Guinea: Central Province: Dinawa, OwenStanley Range, elev. 840 m [08u359S, 146u559E],BMNH 1946.1.7.55 (holotype), 1904.3.17.13.Madew, St. Joseph River, Owen Stanley Range,elev. 1,200 m [08u369S, 146u559E], BMNH1908.10.14.10–12. Mafulu, Owen Stanley Range,elev. 1,270 m [08u309S, 147u049E], AMNH R-59063; BMNH 1935.5.10.171–173. Gulf Prov-ince: Teduku, Eloa River, elev. 178 m [07u439S,146u299E], USNM 562945. Middletown, KikoriRiver, elev. 63 m [07u169S, 144u099E], MCZ R-59059–60; USNM 217499. Unknown locality,UPNG 8737A–B. Simbu Province: Camp II, PioRiver, elev. 300 m [06u449S, 144u529E], MCZ R-121549. Haia, elev. 640 m [06u439S, 145u059E],USNM 562941. Western Province: Matkomrae,Ok Tedi River, elev. 82 m [05u499S, 141u099E],MCZ R-121550. Ningerum, Ok Tedi River, elev.86 m [05u419S, 141u089E], MCZ R-140988;PNGM 22159. Madang Province: Hinihon toReinduk, Adelbert Range, elev. 1,000 m[04u409S, 145u229E], BPBM 5711. E. SepikProvince: Arin, elev. 560 m [03u339S,143u209E], UPNG 7171. Sandaun Province:Mt. Nibo, S. Aitape, elev. 208 m [03u239S,142u099E], AMNH R-100047. Parkop, Mt.Sapau, Torricelli Range, elev. 526 m [03u259S,142u319E], BPBM 223455. Indonesia: PapuaProvince: Ingembit, Ok Walimkan River, elev.117 m [05u389S, 141u009E], MZB 1464. Cano-bivak, Eilanden River, elev. 37 m [05u149S,139u409E], ZMA 17682. Beaufort River camp,Baliem Valley, elev. 64 m [03u279S, 139u109E],RMNH.RENA 4988, 47753–54. Siewa, elev. 80m [03u039S, 136u229E], MZB 2784. W. PapuaProvince: Fak Fak Regency, Bomberai Peninsu-la, elev. 380–520 m [02u579S, 132u359E], ZMA17683.

Toxicocalamus sp. (N 5 17). Papua NewGuinea: Morobe Province: Garaina, elev. 770 m[07u539S, 147u089E], AMNH R-95624, 101100–03, 103681, 104084–85, 107203–05; MCZ R-152428–31. Oro Province: Itokama 820 m[09u129S, 148u149E], BPBM 36185. W. High-lands Province: Jimi River, elev. 960 m [n/a],UPNG 962.

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A New Species of New Guinea Worm-eating Snake, Genus Toxicocalamus (Serpentes: Elapidae), from the Star Mountains of Western Province, Papua New Guinea, with a Revised Dichotomous

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