A New Species of New Guinea Worm-Eating Snake, GenusToxicocalamus(Serpentes: Elapidae), From the Star Mountains of Western Province, Papua New Guinea, With a Revised Dichotomous Key to the Genus

2015·5.7 MB·English

by  Mark O’Shea

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 KEY TO THE GENUS
MARKO’SHEA,1,4FREDPARKER,2ANDHINRICHKAISER3
CONTENTS
Abstract 241 Western Province, Papua New Guinea. The new
----------------------------------------------------------------------
Introduction 242 species is the largest known member of the genus
--------------------------------------------------------------
MaterialsandMethods 242 andcanbedifferentiatedfromallotherToxicocalamus
-----------------------------------------
ToxicocalamusernstmayriNewSpecies 243 byacombinationofthefollowingcharacters:largesize
---------
Holotype 243 (total length of the holotype 1,200 mm), dorsal head
Etymology---------------------------------------------------------------- 244 scutesinthetypical‘‘colubrid-elapiddorsalnine-scute
Diagnosis ------------------------------------------------------------- 244 arrangement’’; separate, single preocular and paired
Compariso-n--s------------------------------------------------------------ 248 postoculars; single anterior temporal and single or
Descriptionof---t-h---e---H----o--l--o--t-y--p---e------------------------------ 249 pairedposterior temporals;six supralabials, withthird
-------------------------- and fourth supralabial contacting the orbit; dorsal
ColorationinLife 251
---------------------------------------------- scales in 15–15–15 rows; 203 ventral scales, 29
ColorationinPreservative 251
------------------------------- subcaudalscales;andadividedanalplate.Itserstwhile
TailTipMorphology 251
----------------------------------------- status, misidentified as Micropechis ikaheka in the
Osteology 252
-------------------------------------------------------------- collection of the Museum of Comparative Zoology,
NaturalHistory 252
--------------------------------------------------- demonstrates the need for detailed examination of
Discussion 255
----------------------------------------------------------------- existingcollectionsandisindicativeofhiddendiversity
KeytotheGenusToxicocalamus 259
---------------------- yettobeidentified,notonlyinthefieldbutalsoonthe
Acknowledgments 260
--------------------------------------------------- shelves of museum collections. We also provide
Appendix1.SpecimensExamined 260
-------------------- arevisedkeytothegenusToxicocalamus.
LiteratureCited 263
------------------------------------------------------
ABSTRAKSI. Kami mendeskripsikan spesies baru ular
pemakancacing(Toxicocalamus)diPapuaNewGuinea
ABSTRACT. WedescribeanewspeciesofNewGuinea
dari koleksi spesimen di Wangbin, Pegunungan Bin-
vermivorous snake (Toxicocalamus) from a single
tang, Propinsi bagian Barat, Papua Nugini. Spesies
specimencollectedatWangbinintheStarMountains,
baru ini merupakan anggota genus dan dapat dibeda-
kan dari semua ular pemakan cacing tanah (Toxicoca-
1West Midland Safari Park, Bewdley, Worcester- lamus)lainnyadengankombinasidariberbagaiciri-ciri
shire DY12 1LF, United Kingdom; and Australian sebagai berikut: berukuran besar (panjang total holo-
Venom Research Unit, University of Melbourne, type 1200 mm), sisik di kepala bagian atas bercirikan
Victoria 3010, Australia. Author for correspondence ‘‘colubrid-elapid sembilan sisik beraturan’’; terpisah,
([email protected]). preocular tunggal and sepasang postocular; anterior
2P.O. Box 5623, Townsville, Queensland 4810, temporal tunggal dan satu atau sepasang posterior
Australia. temporal; enam supralabial, dengan supralabial ketiga
3Department of Biology, Victor Valley College, dan keempat menghubungi orbit; sisik di bagian
18422BearValleyRoad,Victorville,California92395; punggung berbaris 15–15–15; sisik di bagian perut
and Department of Vertebrate Zoology, National berjumlah 203, sisik subcaudal berjumlah 29; dan
Museum of Natural History, Smithsonian Institution, piringan anal dibagi menjadi dua bagian. Status
Washington,DC20013. sebelumnya, salah diidentifikasi sebagai Micropechis
4Presentaddress,FacultyofScienceandEngineer- ikaheka di koleksi Museum Comparative Zoology. Ini
ing, University of Wolverhampton, Wulfruna Street, menunjukkan perlunya penyelidikan secara rinci ter-
Wolverhampton, West Midlands WV1 1LY, United hadap hasil koleksi yang ada sekarang. Hal ini juga
Kingdom. merupakan indikasi adanya keragaman tersembunyi
Bull. Mus. Comp. Zool., 161(6): 241–264, July, 2015 241
242 Bulletin of the Museum of Comparative Zoology, Vol. 161, No. 6
yangbelumteridentifikasi,tidakhanyadilapangantapi represented in natural history collections,
juga di rak-rak museum koleksi. Kami juga memberi- andwehavesofarlocated475specimensin
kankuncirevisitentangspesiesToxicocalamus.
28 museums worldwide. Of these, 66.3%
Key words: Elapidae, Toxicocalamus, New species, (315 specimens) are identified as T. loriae,
PapuaNewGuinea,Description,Taxonomy,Vermivory butconsideringthegreatvariationinhabitus,
patterning,andscalecountwehaveobserved
INTRODUCTION
amongthesespecimens,webelievethistaxon
Toxicocalamus is a genus of enigmatic to represent a species complex. The next
vermivorous elapid snakes with a primarily best-represented species are T. preussi and
fossorial lifestyle. Although part of the very T. stanleyanus, with 55 and 37 specimens,
diverse Australopapuan terrestrial elapid respectively. Two island taxa, T. holopelturus
radiation, Toxicocalamus is one of only six and T. longissimus, are known from 18 and
genera not represented on the Australian 16 specimens, respectively, whereas an
continent.ItisendemictotheislandofNew anomalous‘‘hybrid’’populationfromGaraina
Guinea, several small offshore islands, and (MorobeProvince,PNG;McDowell,1969)is
the major island archipelagos to the south- known from 15 specimens. The remaining
east of the Papuan Peninsula (Fig. 1). The seven species are represented by eight or
small offshore islands all lie along the north fewer specimens each, with three species,
coast of Papua New Guinea (PNG) and excluding the one we here describe, only
include Seleo Island (Sandaun Province),
known from their holotypes. The largest
Walis and Tarawai Islands (East Sepik
known species so far is T. grandis, at 960
Province), and Karkar Island (Madang Prov-
mm snout–vent length (SVL) for the single
ince). The southeastern archipelagos are
known specimen (BMNH 1946.1.18.34).
part of Milne Bay Province and include
ThatspecieswasdescribedfromtheSetakwa
the d’Entrecasteaux Archipelago (Goode-
River (southern Papua Province, West New
nough, Fergusson, and Normanby Islands),
Guinea,Indonesia),whereitwascollectedin
Woodlark Island, and the Louisiade Archi-
1912 during the Wollaston Expedition of
pelago (Misima, Sudest, and Rossel Islands).
1912–13 (O’Shea, 2013).
Toxicocalamus is seemingly absent from the
We here describe a new species, the
Bismarck,Admiralty,andSolomonsArchipel-
largest of the genus, that we discovered
agos, the great seasonally flooded Trans-Fly
among a series of Micropechis ikaheka in
region of southern New Guinea, the Torres
the collection of the Museum of Compara-
Strait Islands, the Schouten Islands, and the
tiveZoology,whereithadlanguishedfor45
Vogelkop and associated Raja Ampat Archi-
years since its collection by one of us (FP).
pelagoofWestNewGuinea(Indonesia).
Itsdescriptionisyetanotherrecentexample
First described by Boulenger (1896) and
of the unrecognized diversity of natural
laterrevisedbyMcDowell(1967,1969),the
history collections (see Kathriner et al.,
genus Toxicocalamus currently comprises
2014), and it showcases their value to
11speciesandasinglesubspecies,including
students of biological diversity.
(in order of description): T. longissimus
Boulenger, 1896; T. loriae (Boulenger,
MATERIALS AND METHODS
1898); T. stanleyanus (Boulenger, 1903);
T. preussi (Sternfeld, 1913); T. buergersi Characters used for evaluating and com-
(Sternfeld, 1913); T. grandis (Boulenger, paring specimens were taken from 344
1914); T. p. angusticinctus (Bogert & museumspecimens(Appendix1).Abbrevia-
Matalas,1945);T.spilolepidotusMcDowell, tions for measurements and scale counts
1969; T. holopelturus McDowell, 1969; used in the description include snout–vent
T. misimae McDowell, 1969; T. mintoni length (SVL), tail length (TL), total length
Kraus, 2009; and T. pachysomus Kraus, (TTL), number of subcaudal scales (SC),
2009. Many of these taxa are poorly andnumberofinfralabialscales(IL).Scales
NEW TOXICOCALAMUS FROM PAPUA NEW GUINEA N O’Shea et al. 243
Figure1. SatellitemapshowingtheislandofNewGuineaanditssurroundingislandsandarchipelagos.Localitiesforallknown
museum specimens of Toxicocalamus are marked by white dots. The red dot indicates the locality for the holotype of
T.ernstmayrinewspeciesatWangbin,WesternProvince,PapuaNewGuinea,whereastheyellowdotindicatestheapproximate
typelocalityofT.grandis,‘‘launchcamp’’ontheSetakwaRiverofPapuaProvince,IndonesianNewGuinea.Theyellowline
markstheinternationalborderbetweenPapuaNewGuineaandPapuaProvince,Indonesia.
were counted as suggested by McDowell skull measurements (expanded from Dwyer
(1969). Sex was determined by examination and Kaiser, 1997) is presented in Table 1.
ofgonads,presenceofevertedhemipenes,or Global positioning system coordinates were
presence of the retractor penis muscle. determined using individual museum
Length measurements were taken by run- records and published species accounts
ning a nonelastic string from the tip of where available; those unavailable were
the snout along the ventral medial axis of obtained from Google Earth (WGS 84) to
the body, under consideration of the points the nearest minute. Museum acronyms are
about measurement accuracy raised by taken from Sabaj Pe´rez (2014), with the
Natusch and Shine (2012). Measurements addition of UPSZ, now the preferred acro-
of tail tips were taken using Mitutoyo digital nymfortheMuseumofEvolutioncontaining
calipers to the nearest 0.1 mm. X-rays of the Uppsala University natural history col-
the holotype were taken using a Kevex lection (Mejlon, personal communication).
PXS5-724EA emitter and a Varian PanScan
4030Rreceiver(40-kVtargettubevoltage)at TOXICOCALAMUS ERNSTMAYRI
the Museum Support Center of the U.S. NEW SPECIES
NationalMuseumofNaturalHistory,Smith-
Star Mountains Worm-Eating Snake
sonian Institution, Suitland, Maryland. The
holotype of T. grandis was X-rayed using Figures 2–3, 5A, B, 6A, C
a Machlett X-ray tube in a Solus–Schall
emitter(outputsettings:55kV,15mA)atthe Holotype.MCZR-145946,anadultfemale
BMNH. Cranial characters were measured from Wangbin village (5u14926.720S,
on digital X-ray images in pixels using the 141u15931.920E), elevation 1,468 m (4,800 ft),
software AnalyzingDigitialImages (Museum neartheOkTediRiver,intheStarMountains
of Science, Boston, Massachusetts). A list of of the North Fly District, Western Province,
244 Bulletin of the Museum of Comparative Zoology, Vol. 161, No. 6
TABLE1. CranialcharactersmeasuredontheholotypesofToxicocalamusernstmayrinewspecies(MCZR-145946)andT.grandis(BMNH
1946.1.18.34).Measurementsweretakenontheleftsideofpairedstructuresandconstitutemaximummeasurements.Allmeasurementswere
subsequentlyconvertedintoratiosappropriateforcomparisons(seeTable3).
Character Abbreviation Description
Skulllength SL skulllengthfromtheanteriormostpartofthenasaltotheoccipitalcondyle
Braincasewidth BW broadestextentofthebraincase
Frontals,totalwidth FW takenattheprefrontalprocess
PF
Frontals,totalwidth FW takenatthesupraorbitalridge
SO
Parietalwidth ParW greatestwidth
Supratemporallength StL measuredinastraightline
Maxillalength MxL measuredinastraightline
Atlaswidth AtW widthofthefirstcervicalvertebra,measuredacrossthetransverse
processes
Axiswidth AxW widthofthesecondcervicalvertebra,measuredacrossthetransverse
processes
Dentary–articularlength D–A lengthofthelowerjawfromtheanteriortipofthedentarytotheposterior
endofthearticular
PNG, (Fig. 2), killed by a villager and paradise (Aves, Passeriformes, Paradisaei-
collected by FP on 23 December 1969. dae), during which he collected many new
Originally identified and accessioned as bird and orchid species. Second, the holo-
Micropechis ikaheka Lesson, 1830. type of T. ernstmayri has been housed in
Etymology. The species name ernstmayri the MCZ collection, mislabeled as
is a patronym honoring the German-Amer- M. ikaheka, after having arrived and been
ican ornithologist, systematist, and evolu- accessioned in June 1975, the month and
tionary thinker Ernst Mayr (1904–2005). year that Mayr retired. Third, the true
After leaving Germany in the early 1930s identity of this specimen was recognized
Mayr took up a curatorial position at the by one of us (MOS) during a visit to the
American Museum of Natural History in MCZ in May 2014, undertaken with the
New York. While there he wrote his de- financial support of an Ernst Mayr Travel
finitive book Systematics and the Origin of Grant from Harvard University, awarded to
Species from the Viewpoint of a Zoologist enable examination of the Toxicocalamus
(Mayr, 1942), which solved one of the big holdings at the MCZ and the AMNH, the
questions of Darwinian evolution, that of two U.S. institutions where Mayr worked.
how one species evolved into many via the Finally, 2015, the publication year of this
mechanism of natural selection. In 1953 description, marks the decennial of Mayr’s
Mayr moved to Harvard University in passing at age 100, and naming a New
Cambridge, Massachusetts, where he was Guinea snake after him seems a suitable
Director of the MCZ from 1961–1970. He tribute.
retired in 1975 as Alexander Agassiz Pro- Diagnosis. Toxicocalamus ernstmayri is
fessor of Zoology, Emeritus (Bradt, 2005). the largest known species in the genus
There are several connections linking Toxicocalamus and the only one with an
Ernst Mayr to this new species of Toxico- SVL in excess of 1.0 m. It can be
calamus, which make him, and this snake, distinguished from all other known Toxico-
the ideal candidates for a patronym. First, calamus by the following combination of
Mayr himself visited New Guinea, and characters: large size (SVL of the holotype
during the late 1920s he spent over 2 years 1,100 mm), dorsum of head exhibiting
conducting fieldwork in an area now part of the typical ‘‘colubrid–elapid dorsal nine-
PNG, as a member of a joint Rothschild– scute arrangement’’ (sensu O’Shea, 2005:
AMNH expedition focusing on birds of 12)thatcomprisespairedinternasals,paired
NEW TOXICOCALAMUS FROM PAPUA NEW GUINEA N O’Shea et al. 245
Figure2. Satelliteclose-upmapsoftheregionsurroundingtheWangbintypelocality,indicatedbyareddot,ofToxicocalamus
ernstmayrinewspecies,showing(A)itspositiononthesouthernversantoftheStarMountainsandproximitytotheIndonesian
borderinrelationtotheOkTediRiverandFlyRiverdrainages,and(B)close-upviewtoillustratethespecificpositionofthe
localityalongtheOkTediRiver.
246 Bulletin of the Museum of Comparative Zoology, Vol. 161, No. 6
mined(Ne)andknownotallength(TTL),anal(SL),andsupraoculars mintoni 1(1)5500+111611*nodata PF+PR,SO+F single singlepaired6(3–4) 15-15-15194nodatapairedpaired51+nodatanodatanodatanodata
T2.ComparativedataforspeciesinthegenusToxicocalamus,includingT.ernstmayrinewspecies.ExaminedcharacteristicsincludethenumberofspecimensexaABLE(Nk),andmaximumlengthsformalesandfemales.Asterisks(*)denotespecimenswithanincompletetail.Abbreviationsincludesnout–ventlength(SVL),taillength(TL),tplate(AP),anteriortemporals(AT),frontal(F),internasals(IN),postoculars(PO),prefrontals(PF),preoculars(PR),posteriortemporals(PT).subcaudals(SC),supralabials(SO),sensulato(s.l.). TaxonCharacterernstmayrisp.nov.buergersigrandisholopelturuslongissimusloriaes.l. (N)1(1)3(6)1(1)18(18)10(12)211(315)Nek5555=nodata330+35365nodata665+122787590+75665610+836935Length(SVL+TLTTL)555555R1100+1001200567+24591960+801040760+77837715+44759681+47728 HeadcharactersHeadscutefusionnonePF+PR+INnonenonePF+PRnonePOfusedSL,single,orpairedfusedsinglepairedusuallysingleusuallypairedpairedATabsent,singleorpairedsingleabsentsinglesinglesinglesinglePTabsent,singleorpairedpairedsinglepairedpairedpairedsingleorpairedSL(contactorbit)6(3–4)4(2–3)6(3–4)6(3–4)6(3–4)6(3–4) BodycharactersDorsalscount15-15-1515-15-1516-15-1515-15-1517-17-1715-15-15=nodata330nodata226–251244–273150–211VentralscountR203313–319207246–256273–304179–227APentireordividedpairedsinglepairedpairedpairedsingleorpairedSCsingleorpairedpairedpairedpairedsinglepairedpaired=nodata40nodata54–6845–4840–62SubcaudalscountR2924–282737–4124–3223–41=nodata9.6nodata15.0–17.510.4–12.19.9–23.1TailratiotoTTL(%)R8.34.1–6.37.77.9–9.25.3–7.76.5–15.6Tailshapeandterminusstout,conicalflat,roundedstout,conicalelongate,conicalelongate,conicalelongate,conical
NEW TOXICOCALAMUS FROM PAPUA NEW GUINEA N O’Shea et al. 247
spp. 17(21)5332+794115667+38705 noneusuallypairedsingleorpairedusuallypaired6(3–4) 15-15-15168–184178–195singleorpairedpaired30–5720–3411.1–20.87.3–19.7elongate,conical
stanleyanus 31(37)5529+836125667+38705 PF+PRsinglesinglepaired5(2–3) 15-15-15218–245230–281singlepaired35–5122–299.0–21.15.1–9.7elongate,conical
spilolepidotus 2(2)nodata5770+80850 nonepairedsinglesingleorpaired6(3–4) 15-15-15nodata200–205pairedpairednodata33nodata9.4–10.7elongate,conical
n
o
x
Ta preussi 42(55)570+55625544+40784 PF+PR+INsingleabsentsingle5(2–3) 13-13-13263–295283–359singlepaired38–4216–337.9–15.23.5–8.4flat,rounded
57
*
6
achysomus 1(1)5+4654nodata nonesinglesinglepaired6(3–4) 15-15-15171nodatapairedpaired20+nodatanodatanodatanodata
p 0
0
5
68 ed cal
misimae 2(3)5393+754nodata PF+PRsingleorpairsinglepaired6(3–4) 15-15-15227–231nodatapairedpaired46–48nodata13.3–16.0nodataelongate,coni
=R =R =R=R
d
T2.ContinuedABLE Character (N)Nek 5Length(SVL+TLTTL) HeadcharactersHeadscutefusionPOfusedSL,single,orpaireATabsent,singleorpairedPTabsent,singleorpairedSL(contactorbit) BodycharactersDorsalscount Ventralscount APentireordividedSCsingleorpaired Subcaudalscount TailratiotoTTL(%) Tailshapeandterminus
248 Bulletin of the Museum of Comparative Zoology, Vol. 161, No. 6
Figure3. HolotypeofToxicocalamusernstmayrinewspecies(MCZR-145946)in(A)dorsaland(B)ventralview.Scale55cm.
prefrontals, a frontal between the supra- holotype of T. ernstmayri demonstrates
oculars, and paired parietals (Fig. 4A9), six a complete lack of the strongly banded
supralabials, with third–fourth contacting patterningdiagnosticofPapuaNewGuinean
theorbit(Fig. 4C9,D9);dorsalscalesin15– M. i. fasciatus (Fischer, 1884). The pattern-
15–15 rows; 203 ventrals, 29 subcaudals, ing of T. ernstmayri is similar to that of the
mostly paired, a few single; divided anal almost patternless ‘‘yellow phase’’ M. i.
plate, short tail terminating with spinous, ikahekafromWestPapuaProvince,Indonesia
cone-shaped terminal scale (Fig. 4E9). exemplified by the holotype of M. i. ikaheka
Comparisons. The genus Toxicocalamus (MNHN 7669). During the development of
can be distinguished from all other New another project (O’Shea, unpublished data)
Guinea elapids, including M. ikaheka but we examined 132 specimens of M. ikaheka
excepting Pseudonaja textilis (Dume´ril fromacrossthetaxon’sentirerange,confirm-
et al., 1854), by the absence of a temporo- ing the subspecific pattern differences; 15
labial scale between the fifth and sixth vouchers are included in Appendix 1 as
supralabials (see Discussion). From P. examplesforthesepatterns.
textilis it can be differentiated by its We here compare T. ernstmayri with all
relatively small eyes (diameter two-thirds 12 congeners (11 species and one subspe-
the distance from the lower edge of the cies), with relevant characteristics for these
orbittotheloweredgeoftheupperlip)and species given in parentheses. A more expan-
unremarkable supraoculars, whereas the sive listing of comparative characters is
highly active and visually alert P. textilis provided in Table 2. Toxicocalamus ernst-
possesses large eyes (diameter 1.5 times the mayri exhibits the regular colubrid-elapid
distance from the lower edge of the orbit to nine-scute dorsal arrangement (Fig. 4A9)
the lower edge of the upper lip), under that distinguishes it from T. mintoni (fused,
strongly pronounced, shelved supraocular head-wide, frontal-supraocular scute), T.
scales. Toxicocalamus ernstmayri can fur- buergersi, T. preussi preussi, and T. p.
ther be distinguished from M. ikaheka angusticinctus (fused prefrontal-internasal-
(characters in parentheses) by its short tail, preocular scutes), and T. stanleyanus, T.
equivalent to 8.3% TTL (11–14% TTL), its longissimus, T. misimae, and T. mintoni (all
ventralscalecountof202(174–191),andits with fused prefrontal-preocular scutes). The
low subcaudal count of 29 (36–49). The presence of six supralabials distinguishes