Redescription of Nephrurus asper Gunther, and description of N. amyae sp. nov. and N. sheai sp. nov

15 Pages·1994·5.7 MB·English

by  Patrick J Couper

REDESCRIPTIONOFNEPHRURUSASPERGUNTHER. AND DESCRIPTION OF
N.AMYAESP. NOV ANDN SHEAISP. NOV.
PATRICK COUPERANDROBERTA.M GREGSON
J.
Couper, P.J. & Gregson R.A.M. 1994 12 01: Re-description ofNephrurus asperGunlher,
and description ofN, amyae sp, nov, and N sheai sp. nov, Memoirs ofthe Queensland
Museum37(1):67-81 Brisbane. ISSN0079-8835.
Threeallopatricspecies,clearlydefinedbymorphologyandcolourpattern,comprise the»V.
asper species complex. N. asper is confined Jo Queensland (10o5.VS-25*2rS), and hasa
distinctive,boldrybandedcolourrnorphoccurringonCapeYorkPeninsula(extendingsouth
toMt Surprise, 18'21'S);N. amyaesp. nov. tothe south-central Northern TerritoryandN,
sheaisp.nov. totheKimberleyregionofWesternAustraliaandtheArnhcmEscarpmentof
the Northern Territory. All three species occur in open woodland habitats where they feed
onawiderangeofarthropodspecies.Malesareinreproductiveconditionduringthesummer
monthsandgravid femalesarepresentinthepopulationfrommid-summertQoautumn for W.
asperand N. amyae. N. sheaimay be sexually active throughoutthe year. Gekkomdor,
Nephrurus, systematic*, newspecies, distribution, open noodland, diet, reproduction,
Patrick Couper, QueenslandMuseum, PO Box SS00 South Brisbane, Queensland 4WL
%
Australia;RobertA.M.Gregson,DepartmentofPsychofogy.AustralianNationalUniversity,
Canberra,Australian CapitalTerritory02(H), Australia; 10June 1994.
Nephrurus asper was described from a single The current study is augmented by Kol-
specimen (BMNH1946.8.2334) from Peak mogorov SmirnovTwo-sampleTestsand multi-
Downs Stn, mideastem Queensland (22°56'S variate discriminant analyses. These analyses,
i
148305*E). Subsequently, specimens matching performed using body measurements and quan-
the type description have been recorded from titativescalecountsonly,provideanindependent
roost ofnorthern Australia(Cogger, 1992). assessment ofthe morphological identity ofthe
While the distinctive morphology of.V. asper fourgeographic groups identified on colourpat-
has ensured its nomenctatural stability (Cogger tern and/ordegree ofspinosity.
et.aL, 1983),severalauthorshavecommentedon The statisticalanalyseswereconductedby one
geographical variation within the species. Slon of us (RAMG) and the laxonomic analysis was
(1963) noted differences in tail length between conducted bytheother(PJC)
specimens from Western Australia and All measurements were taken using Miluloyo
Queensland. Bauer (1990) observed that electronic callipers. Abbreviations for body
specimens from the south-central Northern Ter- measurements are as follows: snout-vent length
ritory were largerthan those in otherpartsofthe (SVL); axilla to groin (AG); tail, tip toconstric-
range. Wilson and Knowles (1988) recognised tionatbase (T);tail,tiptovent(TV);knobwidth
the broad-banded form of N. asper from Cape (KW);forelimKaxillau>tip nflongestdigit(LI);
YorkPeninsulaasdistinct fromoverpopulations hindlimb, groin totipoflongestdigit (L2); neck
ofthisspecies. length, axilla toposterior edge ofear(NL): head
An examination of all N. asper held in the length, tip ofsnout to dorsal/anterior margin of
collections of major Australian museums ear (HL>; head width, measured between ear
revealedthatthespecies, ascurrentlyrecognised, openings (HW); head depth, from aboveeyes to
couldbedividedintofourgeographicallydistinct ventral margin of maxillae (HD); snout, tip to
groups (Arnhcm Escarpment, NT & Kimberley anterior margin nforbit(S); eye to ear, posterior
region, WA; south-central NT; Queensland & margin "' OfrMl to anterior margin of ear (EB)
Cape York Peninsula) on the basis of colour, Abbreviations forscalecountsandcaudal annuli
pattern,spinosity,andotherscalecharacters.Pat- arcasfollows:numberofgranularscalesindirect
ternandspinosity have been used widely in gck- contact with dorsal edge of rostral scale (R);
konid revisionarv work (Couper ct. uH 1993: numberof interorbital scales counted across the
Storr, 1988, 1989). They have been found to be narrow, bonybridgebetween theorbits, includes
useful in delineating the species in the /V. asper the enlarged series of >caks on either side (J)\
complex, in a way which conforms closely with number ofsupralabial scales counted to exclude
statistical analyses granular scales towards angle of mouth (SL);
1 .
MEMOIRSOFTHF.QUKENSLAND MUSEUM
TABLE 1.DtQcrcntiationbetweengroupsusmgaKS2 Arnhem Escarpment (NT) & Kimberley region
Procedure x~ from the 6 possible comparisons be- WA). eroup 2 = QLD. group 3 = south-central
tweenthe4samples.Nisthenumberofcontrastswith NI T,andgroup4 =CYP.
fK1o3Sr.28t1dh.ex,201stlaetviesltiics>a6b\oTuthe9..2015.lfeovretlhfeor0x01Zll>eaveblouista5b.o9u9tT (K1S2T)he(KKoollmmooggoorroov,vS1m9i4r1;noSvmTiwrnon-vs,amIp9l3e9iTe(sMti
each ofthe 6 pairwise contrasts between groupa.
GROUPS oneachof 17characteristics,givinga total of10
cbhoadryacter 1 vs2 1 vs3 1 vs4 2vs3 2vs4 3vs4 Xpr2ovcaelduuersee,acuhsiwnigthm2idn.fi.mTahlispirsioarnoanspsaurmapmteitonnes
T 6.59 3.69 4.9*1 4.00 0.47 3.88 about the form of frequency distributions of
TV 18.00 5.81 3.24 5.83 0.52 231 measurablecharacteristics (Walsh, 1965).
KW It indicates which characteristics differentiate
6.50 7.53 0.24 22.99 3.88 1.61
TA k.44 430 4.10 0.35 0.59 044 belterlhan al some given probability level, ifthe
characteristicsarcindependent. ThisisnotStrict-
Li 1 135 4.34 1.80 1.05 0.92 44 ly true a priori, and hence conservative prob-
L2 0.52 3.98 ] 81 2.07 0.48 ; abilitylevelsmaybeusedindecidingwhatresults
HW j070 803 i 13 7.16 0.41 to accept. It also will fail to detect second-order
HL 26 7,19 0.59 4.88 1 I ! j 4J measures, suchasratiosordifferencesofcharac-
S 1.31 7.11 2.14 9.20 3.71 : -.7 teristics, which can be non-linear and of use,
NL 6.32 6.17 1.58 1564 1.44 5,07 iunnlmesms thhaevseebaeracnbudiilvtiidnetdobtyheSdVaLta,.tThhoseembeaasseudroens
SL 25.32 24.93 \1 59 1.24 2.55 3.44 scale counts have not. Bilateral measures have
IL 35.80 26.48 15.38 0.73 2.41 1.32 been pooled (Tabic 1) Discriminant analyses
R 3.40 30.75 079 30.13 3.79 9. :. havealsobeenrun becausetheycan utiliselinear
i 19.84 3528 ft iM 7.42 0.20 2.84 (but only linear) combinations ofcharacteristics.
EE : r n-92 7 50 4 19 2.85 732 and hence may separate groups which are not
HD CUJ9 :- i • !98 4.60 2\0 • 78 1 Tdihsetidnigsuicsrhiemdinoanntsianngalleycsheasradctoe,rihsotwicesve(rT,ablmeak1)e.
AG n •: 2.35 I 65 1.37 1 |] 0.26 some distributional assumptions approximating
N 8 9 4 6 - 3 to normality and homosccdascity acrossgroups.
The role ofthe KS2 is also involved with the
number of infralabial scales counted to exclude question of what characteristics to use in Ihe
granularscalestowardsangle ofmouth (IL),and discriminantanalysis,ifnotallthecharacteristics
number ofcaudal annuli counted between basal which have been measured arc used.
constriction and distal knob (TA). 2.litecorrelation matrix on the characteristics
OTHER abbreviations USED: Queensland over the whole 149 complete records used. This
Museum (QM), Australian Museum (AM). showedthat therankofthedata is much lessthan
Museum of Victoria (NMV), South Australian 17;thisindicated,asexpected,thatthereisredun-
Museum (SAM). Western Australian Museum dancy in the morphological data.
<WAM) and the Northern Territory Museum 3. Four discriminant function analyses (Rao,
(NTM Cape YorkPeninsula(CYPl,mideastcrn
). 1952):
Queensland (MEQ)« Queensland (QLD), North 3.1. Using all the characteristics, on raw
cmTerritory (NT), Western Australia (WA).
measures.
Reproductive and dietary data were compiled 3.2. Using a stepwisediscriminantanalysis, on
by examinationofspecimens in which the body raw measures.
cavity had already been opened
) 3 IJsinjj all ihe characteristics, on measures
divided by SVL.
statisticalanalyses
3.4. Usinga stepwisediscnminanl analysis,cm
measures divided by SVL.
The following data analyses have been per- The stepwise procedures are inbuilt to the
formed, Using complete records on 149 animals SPSSX program used;becauseoftheintercorrela-
Someare inlcrnaJ checks on consistency and arc tions noted under part (2) above, the variables
hence not reported in full here. The characters retained in a stepwise analysis will depend on
u.sedintheseanalysesareIhebodymeasurements thoseinlereorrelationsand on therelativesizesof
and scale counts defined above. Group I =. the the four groups, but will be analogous to the
DESCRIPTION OFNEPHRURUS 55
F
u %
n
c o
2- t
O
i o o °o
o
n o
OO a *>
° O
2 °
7 v
o Group 2 %
o
O o jT
° Group4 °
9 A ^»A W °fi O n Group J
o O o * '
o& ©
1-
"I
^
GroupCenlroids
' Group4
-2 © * A O 4 Group 3
* Group2
Function 1 * Group 1
-3
"2
FIG. I.Plotoftheindividualsandgroupccntroidsonthe 1sttwocanonicalvarialcsfromthediscriminantanalysis.
Group 1 =the Arnhem Escarpment(NT) & Kimberley region (WA), group 2 = QLD, group3 =south-central
NT,andgroup4=CYP.
characteristics with larger %~ values in Table 1. they are weights to multiply with corresponding
We present tables derived from (3.4) here; as measurements, and then sum the products and
there are only 11 animals in the smallest group, addtheconstantterm inthecolumn. Asaddition-
the number of variables in the discriminant al specimens become available itis preferable to
analysis has to be reduced to below 10. The incorporate them in a extended discriminant
stepwise procedure is the most appropriate for analysis,thusyieldingnewweights.Theefficien-
this, retaining only those variables with dis- cyofallocation decisionsusingthisdiscriminant
criminativepower; itshowedbyF-teststhatonly analysis issummarised(Table4); itwill be noted
eight variables were legitimately retained in the that the relatively poor separation of group 2
(QLD)andgroup4(CYP)iscompatible withthe
analysis.
Information for the identification of new results from the KS2 analysis.
specimens when allocation is based on body
RESULTS
measurements and scale counts only is not in
itself sufficient (Tables 2, 3). Allocation to
species must also take into consideration colour The plotofindividuals and group centroids on
pattern and degree of spinosity. The relative the First two canonical variates from the dis-
values of the coefficients (Table 3) are of use, criminant analyses (Fig. 1) gives a picture com-
. .
56 MEMOIRSOFTHEQUEENSLAND MUSEUM
TABLE2.Standardiseddiscriminantftinctioncoefficients SYSTEMATIC*
and correlations with discriminant functions from the
=
stepwise analysis with three canonical functions. 1c NephrurusasperGuntber, 1986
coefficientslorthefirstcusenminant,lr=itscorrelations (Fig. 2AJ3; Statistical analysesgroups 2,4)
withdiscriminatingvariables,andsocm.
char Ic Ir lie Ilr nic nir Material Examined
HW .070 .112 511 475 -.106 .415 Queensland: QMJ22I79. QMJ24921, B&Hiaga,
CapeYorkPenirvsula(10^3\S!42o24*EuQMJ57652,
I l 452 3-88 209 ^t;i -1 234 -.201 6.5km E of Heathlands airstrip (11*44.33*5
LI 392 **• -.459 !CC -.472 .107 142'38.28'E);QMJ54644,Heathlandsroad, 1kmfrom
NrL --1]16»4 -20.4.•; .13.91847 473327 »i/iJj0C93 ..119519 mQBaeMaiJgn4lUe1C6aA5pi.ersWtrrcoiiappdafjltu3on*cA0tu5ir*ouSnku!(u14,1r*m54i76d'*-ESw>a;y1,4Q2NM"J4C07*a3Em8)p,;
SL .864 360 -.367 .358 i.j:i i:: Kingsborough (!6"55rS 145'(XTE). QMJ4960. Bul-
TA -1.750 OKI .122 .431 -.956 086 lock Ck, via Cajms (16°55'S 145°46*E); QMJ57993,
Itv sn -.327 -.138 .2"! .124 -.122 QMJ5S850, ElizabethCk, orBulIcringa NP, NofMt
Surprise tl8'00'2rS >45*59'50"E); QMJ2125. '
paliblewithTablesoneandfour.Group2(QLD) NorthernRjvcrs;QMJ44689.NewMoonStn(19*ITS
andgroup4(CYP)arenotsignificantlyseparable 145'44'E); QMJ3443, Charters Towers, Black Jack
(20'08'S, 146*l3'Ei;QMJ420l5, Hughcnden,anprox
on external morphology. The recognition of 2km W of kma-Redeliffe Sin Homestead <2CT59'S
group4 wasbasedoncolour-pattern.Theunifor- l44'33,Ej;QMJ44948,EgeraSinr20'59,SH6*05,EV.
mity in morphology between groups 2 and 4 J143,NatalDowns(21'05'S. J46WE),QMJ45
dovetails well with preliminary genetic data Kuridala, S of Cluneiirry (2T17*S 140'30'E);
whichshowa low levelofdivergence between a QMJJ5565.80kmS\VofMackav(2J'39*S 148*42*E)
broad-banded specimen from Heathlands. CYP QMJ31976, QMJ35040. WinWlon<22*23'S 143'02'Hl;
MEQ QMJ5727. Lwcknow Sin, of Winlon (22*43*5
a(nCdouapsepreAcV.imDeonnnfcrlolmant,heuCnappueblll.adraetag)i.onAolfl QLD I|4407*'5358*,EE));;QQMMJJ281635959,.QCMoJr3k5S5t2n.viCalWerimno:nntn-(:222''5560',SS
material examined in (his study, wilh ihe L-xcep- l42M8*Et,QMJ46720.AramacRanRC,WinhavenStn
tionofthebroad-bandedspecimensfromCYP, is ;22'S7*S 145*41*E): QMJ56088, Capella. 20km Non
consistent with the holotvpc of S. asper Clermont road (22'59'S 147'53'E); QMJ36795.
<BMlNiiHon19(s4e6e.8R.e2m3.a3r4k)sinfobrotM.hcaoslpoeurr)-paantdteirsnhaenrde hMaomopntgogno,oDaalrm\a0,'SWar1o4Sn'l0a4'E(»2.3"Q2M0'JS3761570,'3R2ocT.ki-.
assigned to this laxon. The broad-banded CYP QMMaJr9y9lV2a.lLeonSgtrnc,acYhal(l2e3r'o2i6,vSiaI4B4l*a1c5'kEa)l.lQ(M2J37'93853',S
specimens are regarded as a geographically dis- 146*58'E). QMJ367V4, E of Comet Sin (23'37'S
tinct colour morph of A*, asper. The statistical J48S40'E),QMJ34205-O6.halfmy uplheBlackdown
analysesshowthatgroup I (AmhcmEscarpment, Tahlclandetcarprncnl (23'37'S !49I0*E); QMJ8O80.
g(NsuToiusth&ha-bcleKeniimfrbiaolecrnNleTQy)LDarrecgainomdno,rCpWhYoAPl)ogeriawcnnabdlilrygtreddoiusaptsin3a- sLBUialotyk-gea1s:n2sl3f**u25nE30g)**a4;Sn6Q*Ml(S2J4324W*n33I!395'5W*,6EE-1))74;7,;Q°MM1oQJ8uM'9rJ1>;a33QI24M,5Jk4B7m5l,4Saco1Wk.faiDlnjliluan(omc2nat4ni*fioo2ins6s*noiSalf
singlegroup(Table 1, Fig. I)andalsofromeach Theodore & Gibihi Roads (2(4"38*S ]49*59'E>:
other. Groups I and 3 arc here describedas new QMJ7878, QMJ10526, Jundah (24*5D'S 143*04'E);
allopatiic Uxj. The recognition of those taxa is QMJ4540, Dawson Vallev. Castle Ck (24*50*5
further supported by preliminary genetic data 150'20'E); QMJ6011* Wcathersfield?; QMJ74:
(CTohuepcr"A&tlaDsononfelAluasnt.raulnipaunblR.edsaotuar)c.es' (Anon. lRoc(aIli2t-y4d0aSta;I4A2°M4K0'E1i2;SIA8M3,RBa1i2a8v7i9a6,DoCawinrsn,sW(Ie6n*5l5o*cSk
1952 69) Shows rciKonable conformity between 1J4455*'4s6r*EE)):; AAMMRR3II7173835,2.MlappFrooxx.nr1I4n.g5hkmamW(1o8f61*9M*St
the distribution ofthe N. asper species complex Cooper*, nr Si Paul'svine scrub (20'31'S 146'55'Ei;
and the occurrenceofuplandcountry and heavy, AMk(4l83. 210km S ofGhatfeM lowers i'2r.^*S
NW
often stony soils. However, the zoogcographical 146*55'EH AMR63065. approx. 48km of Clcf-
ismepslsiecdawtiitonhsouotfFitrhsitsesdtiasbtrliibsuhtiinognacpahnytnootgebney afso-r (iHio2itO2eM'tl5,102aS2p^p21r54o'7xS*.31844*70E')2k;3m'AE)NM;RAofM12RD0i01a90m47a,7n0tM3ianNyanoefLCaJlkueensrcmtoSiniotnn
thegenusanddeterminingthedatesofspeciation AMR
(23'33'S I41"22,E), 15107, Yamala (23'35'S
events. Presently, there i.< no fossil record for J48J22*E); AMRI13M6, Diamantina Lakes Stn, SW
Npleepteh.runu and biochemical studies are incom- o(f23W'i4n3t'oSn<12439""44T0*SE)1;41*A1M1*REI);1A0M5R454,57A8M6,RIDua1r0i5n62g,a
DESCRIPTIONOFNEPHRURUS 57
camp, 14km NL of Scott's Tank, Diamantina Lakes, TABLE 3. Fisher linear discriminant function coeffi-
2N,WAoMfRWi1n3d07o2r6a,h (2203k'4m5'WS KoIfMOM'oEu)r;aAoMnR1B3a0uh7i2n1i-a chiiegnhtess.iAfnuinmctailosn,argeivaesnsitghneaibrlmeelaostuhreesmpeenctisescowrirtehcttehde
Downs road <24°37*S 149'47'E); NMVDT-D182 forSVL.
('Donald Thomson Collection), Lower Archer R
O3'30*S 142"00*E); NMVD7527, Ml Cook, Cook- GROUPS
StoAwMnR(1154'02283'S, 12425'k1m5'EN);WNMofVDC2oOo4k0t,owQnue(e1n5s*l2a5n'dS, cHhWar l 2 3 4
145"04*E); SAMR 12594, Cooktown (15'28'S 841.13 841.40 785.09 (881*74
MTtS'B); SAMR15791, Charters Towers (20'05'S 1 -26.45 -134.19 -11654 -95.68
146*I6*E); SAMR1771, Ml Coolon <2I*23'S LI 372.24 330.65 $6] 47 314.64
(12437°'24<3T'ES);1S4A1MWRE4)26;O0S,AM6Rk4m26N0Io-f0D2i,am4akmimnaNStonf NL 3629 67.61 *3 14 ^ ::-
R -37.54 -12.84 -56.61 50.85
Diamantina Stn (23*44'S 141'OrE); SAMR42603,
85km W of Windorah (25t2I ,S Hl'SO'E); SL 3971 52.31 64.69 27.21
WAMR55475-7, 16km W of Charters Towers TA -135.06 -IS.37 -28.78 23.92
(20°08'S 146"07'E); WAMR55552. WAMR55601-2. TV -5.45 47.62 54.14 48.57
F1e4r3m'o0y2*EH)o;mesNtTeaMdR,18I84k4m,SCloefrWmmotnton|2(223'"5100?,SS Owi -1975* -193.83 -184.77 195.18
147"38'E); NTMR266, Rockhampton f23'22'S
150'32'E). mean 16.4), TV - 15.7-27.4 (N 66, mean 22 7)
HL - 27.9-34.8 (N 73, mean 31.3). HW - 23.9-
Diagnosis 30.6 (N 74, mean 27.7). HD - 10.7-15.8 (N 70,
csopNil.noousraes.ppaIettrtiesrindsiosmfteidintigsuuidmis-ghsietidszfe(rddiogmiatn,sVd.usnmhhoedaaesnrdbaeytdetlvhyes 7m120e,.a0mn-e11a53.n.08)1(,5N.S57)-.3,9.m2e-1a2n.712i.N3)7,4.NLme-an10.J41-211),.1EE(N-
digits strongly marked with alternating bandsof Head. Large and deep, covered with small,
brownandwhite);fromN.sheaiandN.amyaeby roundtohexagonal,juxtaposcdscales.Posterior-
its smallersize (max SVL 1 14mm vs 121mmN, ly bearing scattered tubercules intermixed with
sheai, 135mm N. amyae). h is further distin- the smaller scales. Each tubcrcule consists of a
guishedfromjV. amyaeby thespinosityoftheits high central scale, circled by a ring of smaller
lumpandthighs(moderatelyspinosevsextreme- hasal scales. Tubercules mosl prominent on Ihe
ly spinose). The arrangement of the basal scales nape. Dorsal skin co-ossified with skull Nostril
surrounding the tubercules on the rump and small, opening upwards and backwards. Eye
thighs also separates N. asperand N. amyae. In large with vertical pupil. Ear vertically elongate.
N. asperthe basal scales are uniform in size and tympanum deeply recessed. Rostral scale small,
lessthanhalftheheightofthecentral scale. InN, with 8-17 (N 77, mean 12.5) scales in direct
amyaethebasal scalesare irregularin sizeandin contactwithdorsal edge. Interorbitalscales 7-! 2
most specimens examined, some of the basal (N 76. mean 8.9). Supralabial scales 11-17 (N
scales are greater than half the height of the 152,mean 13.5). btftaJiblal scales 10-17 (N 152.
centralscale. mean 13.5).
In the KS2analysisN. asper(excludingbroad- Neck, Broad, less than fuilfwidthofhead.
nbiafnidcaendtlyCsYepParsabpleecfirmoemnsothwehriQchLDarpeopnuoltatisoings- smBaoldly,graSntouulta,rdsocr&slaalttoann.dGvreannturlalarsusrcfaalcesesinwtietrh
of this species) and N. amyae show significant mixed with larger conical tubercules on dorsal
sfoerp;arKa\tVi,onH\oVf,thSe,sNhLa,peR,of&frI.eqAtueanscpyedri(sterxicbluutdiionngs andlateralsurfaces. Mid-dorsaltuberculessmall.
sbhrooawd-sbigannidfeicdanCtYsePparsapteicoinmeonfst)heasnhdap(eftofihferaei- TABLE4.Classificationresultsfor149animals.Actual
quency distributions for; T. TV, KW, TA, NL. groupby predictedgroup membership.
SL,!L&I(TablcI).
Actual No. of PredictedGroupMembership
Description Group Cases 1 2 3 4
SVL(mm): 47-114 (N 74, mean 84.1). Propor- Group 1 59 45 8 4 2
tions,(%SVL): - LI - 36.8-47.8 (N 74. mean Group2 48 7 30 7 9
43.0), L2 - 43.2-55.6 (N 73, mean 49.0), AG - Group3 :! 2 2 25 i
34.4-54.2 (N74,mean44.4),T- 1 1.4-22.2(N66, Group4 ii 2 1 8
58 MEMOIRSOFTHEQUEENSLAND MUSEUM
ramp tubercules moderate. Basal scales sur- Distribution
rounding nimp tubercules uniform and less than From the drier parts of coastal MEQ through
halfashighasthecentralscale. Rumptubercules central and western districts. Also present on
larger than flank tubercules (455ft materia! ex- CYP(Fig. 3).
amined) or subequal to flank tubercules (55% The locality given forQMJ2I25 is ^Northern
material examined). Rump tubercules smaller Rivers', which usually refers toan area in north-
thannapetubercules (51% material examined)or em NSW. The following entry in the register
\ubec|ual to nape tubercules (49% material ex- (QMJ2126) isa specimenofSalman'uscornutus
amined). Lower flanks and ventral surface with from the same locality. Neitherofthese species
Scattered, Slight!} raised rosettes which varybe- occurin NSW,norare Iheysympatrieelsewhere.
tween being pronounced orbarelydtscernihle. N. usper typically occurs in dry open country
Limbs. Long and slender, bearing enlarged while S. cornutus is confined to rainforest
tubercules on dorsal surfaces. Tubercules on habitats. Either the locality "'Northern Rivers'
thighslargest,withuniformbasal scaleslessthan shimId be regarded as an error or it refers to a
region(unknowntotheauthors)ofsouthernCape
half as high as the central scale. Digits short,
York Peninsula where both rainforest and dry
cylindrical, undilated distally and terminating in
a non-retractile claw. Third toe on hindlimb forestsoccur.
longest.
Habitat
Tail. Short, moderately depressed, constricted /V. asper favours rocky substrates in open
at base and terminating in .» globular, kidnev- woodlands (Steve Wilson, Tim Hawkes, pers.
shaped knob (KW 17.8-38.8%T. N 66, mean comm.). ll has been recorded also from stony or
29 I),Usuallyfour,rarelyfiveorsix longitudinal compacting soil plains, whereshelteringsitesare
rows of tubercules present on dorsal and lateral available (Ehmann 1992; Couper, unpvbl, data);
surfaces. Uniform basal scales surrounding from low heath at Heathlands, CYP, and Darwin
caudal tubercules less than half a* high as the Stringvbarkopen forest,atAurukun(Cameron&
central scale. Caudal annuli 9-12 (N68. mean Cogger, 1992..
10.4).
ReproductiveNotes
Clutch size - two eggs. Egg-laying occurs in
Pattern mid-late summer. Female WAMR55476 was
gravid during late February. The two oviducal
Biuiy. In spirit, mid-dark brownr or mid-dark eggs measured 27.1 X 13.8mm and 25.2 X
grey on dorsal surface. Many specimens have a 14.0mm;QMJ54644hadagreatlyenlargedviteJ-
broad black band on the nape. Seven - eight, logenic follicle in each ovary at the time it was
raiely six. narrow irregular crossbands killed in early February, 1992. The largest of
(crossbands broad and continuous in CYP thesemeasured9.67X 7.28mm. Agravid female
specimens)present betweenheadandhindlimbs was brought to Wild World for identification in
Crossbands range from obscure to bold and are mid-April (T. Hawkes, pers. comm). Male
continuous or broken into aseries ofspots, wilh WAMR5S552 was in reproductive condition
each spot covering a tubercule. Fine black lines lepididvmisenlargedand turgid) in late Novem-
form reticulations which become obscure in ber.
larger specimens and can be difficult to sec in
darkerindividuals. Ventral surfacecream,some- Dietary Notes
times faintly marbled with grey or brown. The gut of QMJ56088 contained two cricket
Limbs. Lacking irregular crossbands distally. legs (Orthoptera: Gryllidae). QMJ36795 con-
Digits unhanded oronly obscurely banded. tained a medium-sized centipede (Chilopoda:
Tail. As forbody, with an obscure, broad, pale Scolopendridae) and an ant. QMJ44689 con-
tained fragmentsofa medium-sizedspider,anda
crossband on proximal half.
medium-sized cockroach (Blattodea: Blattidae).
Head. Sometimes slightly lignter than body Two faecal samples were collected at Heath-
Covered dorsally and laterally with fine black lands,CYP,inlateJanuary 1992.Thefirstsample
reiiculations, which form an intricate pattern. containedalargespider,asmallcentipede,alarge
These reticulations are most prominent in coleopteran larva and small to medium-sized
juveniles and may fadein largerspecimens. cockroaches (Blattodea). The second sample
DESCRIPTION OFNEPHRURUS 59
m&-
FIG. 2. Nephrurus asper. A, Moura, QLD; B, Broad-banded form, Heathlands, Cape York Peninsula, QLD.
(Photographs: S. Wilson).
60 MEMOIRS OFTHEQUEENSLAND MUSEUM
FIG.3.DistributionoftheNephrurusaxperspeciescomplex.N.amyae(diamonds),N.asper(circles),N.sheai(squares).
containedlegsfromamedium-sizedgrasshopper ways (18°09\ 144°4D. This narrow-banded
(Acrididae), fragments of a large scorpion specimen isapproximately 75kmNEofthe most
(Urodacussp.)andfragmentsofasmallphasmid. southerly museum record of the broad-banded
form (QMJ57993). Clearly, the range of both
Remarks colour forms overlap latitudinally but, to date,
Aclear photograph oftheholotypeofN. asper they have not been found in sympatry.
(BMNH1946.8.23.34) was provided by Bauer,
(1990). The narrow, irregular dorsal crossbands NephrurusamyaeCoupersp. nov.
and moderate, uniform rump scalation of this (Fig. 4; Statistical analyses group 3)
specimen are consistent with the pattern and
scalationofQMJ56088,aspecimen recentlycol- Nephrurusasper. Cogger, 1992(inpart).
lectedonly 14kmfromthe type locality. Further,
thesespecimensaremorphologically/phenotypt- MaterialExamined
cally consistent with populations of spinose Holotype- AMR 04458,WinneckeGoldfields,Gar-
Nephrurus occurring in Queensland between den Stn, N ofAlice1 Springs, NT (23'17'S 134D25'E).
18°19*S & 25°2)'S. These are regarded here as Foundinminetunnel,February, 1982. Don.M.Robin-
N. asper, The broad-banded formonCYP is also son.
included in N. asper because it cannot be sig- 1P5arkamtySpeEs:ofNGolretnheHrenlenTer(r2i3t°o3rIy':S Q1M3J25nI216E5)0;,
nificantlyseparatedfromthisspeciesbyscalation
QMJ53650, Hugh R, nr Alice Springs (23"49'S
features or body proportions, and preliminary \33°2TE)\ AMR49716, Mt Gillen, Alice Springs
genetic data show a low level ofdivergence be- (23°42'S 133'48'E); AMR50542, Alice Springs
tween a specimen from Heathlands, CYPManEdQa (23'42'S 133"53'E); AMR1I965, Mt Gillen, Alice
specimen from the Capella region of Springs(23'43'S I33°48?E); AMR90198,JayCk, NT
&
(Couper Donnellan, unpnbl. data) (23"50*S 133'29'E); AMR10371, between Hale and
Thebroad-bandedCYPcolourformisnarrow- Plenty Rivers, Central Australia (24°25'S 136"10'E);
YslyoousrtekhpearrHnaetrlepidemitftorlooomfgitithcsearlnaanrSgoreco.iwTe-tibyam)ndHheaadwskfoeprsrmov(aiCtadtpehede NNi3MM3VV"4DD05'5E3)1;891.N9M-V2B1Da,r52r58o3.w797kC,mk,nSro(Af2li1Bc*ae3r2rS*po3rwinC1gk3s3<(°2253r3'4'45E2')'S;S
133°52'E); NMVD12684-5, Central Auslralia;
colour transparencies of a broad-banded SAMR1892, Barrow Ck (2T32'S 133°53,E);
specimen from 6km east of Almaden (17°2rS, SAMR38837, Mt Zcil (23"25*S 132°25'E);
144°43'E) and a narrow-banded specimen from SAMR40561, approx 35km SE of Kings Ck Homes-
approximately 87km south, near the junction of tead (24°26'S 13r49*E); SAMR30523, No Locality
the Kennedy and Gulf Redevelopment^! High- data; NTMR44I, Barrow Ck (21*32*8 133"53'E);
DESCRIPTIONOFNEPHRUBUS M
R14096. Mt Riddock Stn, Dulcie Ra. (22B30'S 10.2-14.8 (N 39, mean 12.9). NL - 10.4-19.9 (N
135'25'E); NTMR33717, Winnecke Goldfields 37, mean 14.7).
123'02'S 134'23'E'i;NTMR33716, Bond Springs Sin
(23'34'S WrS&tOt NTMR12377, 16km N Alice Head. Large and deep, covered with small,
Springs (23*34'S 133'53'E); NTMR33720. Alice roundtohexagonal,juxtaposedscales.Posterior-
Springs Hills, 19 Mile Bore (23'35tS 133*52'E); ly bearing scattered tubercules intermixed with
NTMR33722-3, Alice Springs, Wiglcys Turnoff the smaller scales. Each tubercule consists of a
(23'36'S 133'53'E); NTMR12380. 9km N Alice high central scale, circled by a ring of smaller
Springs (23'37'S 133'53'E); NTMR33721, Alice basal scales Tubercules most prominent on the
Springs,Charles R (23'39'S 133*51*E); NTMR5586, nape. Dorsal skin co-ossified with skull. Nostril
NTMR5969, NTMR1499I, Alice Springs (23°4rS small, opening upwards and backwards. Eye
133'52'E);NTMR33724,NTMR33726.AliceSerines
(23*42'S 133°53'E); NTMR5383. Alice Springs large with vertical pupil. Earvertically elongate,
PowerHouse(23'42'S 133'55'E);NTMR5466.Aiice tympanum deeply recessed. Rostral .scale small,
Springs, Mt Gillen (23"43'S 133'48*E); NTMR2458, with7-18(N39,mean9.7)scalesindirectcontact
Alice Springs, Emily Gap (23'45'S 133S57T); with dorsat edge. Interorbital scales 6-11 (N 40,
NTMR33719, Hermannsburg, Finke Crossing mean 8.0). Supralabial scales 11-17(N78,mean
(23*58*S 132"46'E); NTMR33715, KrichaulT Ra. 13.5).Infralabiaiscales 1 1-17(N78,mean 13.4).
(23'59'S 132'38'E); NTMR33728,Nodata.
Neck. Broad, less than halfwidth of head.
Body. Stout, dorsal and ventral surfaces with
Diagnosis small granular scalation. Granular scales inter-
N. amyaeisthelargest andmost spinose mem- mixed with larger conical tubercules on dorsal
ber of the N. asper species group. Its large size andlateralsurfaces.Mid-dorsaltuberculessmall,
distinguishesiifromothermembersofthisgroup rump tubercules large (extremely pronounced)
(max SVL 135mm vs 1 14mm N, asper 121mm except in juveniles. Basal scales surrounding
/V.sheaf). Itisfurtherdistinguishedfrom.V.asper rump tubercules irregular. In many specimens.
and.V sheai by the spinosity ofthe its rumpand somearegreaterthanhalftheheightofthecentral
thighs (extremely spinose vs moderately scale. Rump tubercules larger than flank tuber-
.spinose), The arrangement of the basal scales cules. Rump tubercules larger than nape tuber-
surrounding the tuhcrcules on the rump and cules (90% material examined) or subequal to
thighsalsoseparatesN. amyaefromAtasperand napetubercules(10% materialexamined).Lower
N.sheai. InN, amyaethebasalscalesarcirregular Hanksandventralsurfacewithscattered, slightly
insizeandinmostspecimensexamined,someof raised rosettes which vary between being
diebasalscalesaregreaterthanhalftheheightof pronouncedorbarelydiscernible.
the central scale. In .V, asper and .V. shear the Umbs. Long and slender, bearing enlarged
basalscalesarcuniform in sizeand lessthan half tubercules on dorsal surfaces. Tubercules on
the height ofthe cenlral scale. thighs largest (extremely pronounced). Basal
In the KS2 analysis .V. amyae and N. asper scales surrounding thigh tubercules irregular. In
(excluding broad-bandedCYPspecimens which manyspecimenssomearegreater halftheheight
arc not significantly separable from other QLD ofthe central scale. Digitsshort, cylindrical, un-
populations of this species) show significant dilaleddistaJlyandterminatinginunon-retractile
separationoftheshapeoffrequencydistributions claw Third toe OH hindlimb longest.
for; KW, HW, S, NL, R & I. M amyae and .V. Tail. Short, moderately depressed, constricted
sheaishow significant separation ofthe shape of
SfLr.eq1uLe.ncRy&dKiTstarbibtuetilo)n*.for;KW,HW,HL,S,\L. •2si7ht,ab6pa)e.sdeFkaonnuodrbtle(orKnmgiWintau2t1di.in.ng1a-li3n5.ra0owgSsl&oTb.ouflNart,u3b6ek,irdcmnueelaven-s
present on dorsal and lateral surfaces. Basal
Description scales surrounding caudal tubercules less than
SVUmm):50-135(N40.mean100.3).Propor- halftheheightofthecentralscale. Caudalannul!
tions, (%SVL): - LI - 38.1-52.4 (N 40. mean 8-13 (N 37, mean 10.4).
42.7). L2 - 42.5-56.1 (N 40. mean 48.4). AG - The measurements and scale counts for the
35.4-53.5(N40,mean45.1),T- 12.3-17.5(N37. holotype (AMR104458) are as follows; SVL
mean 15.0), TV - 15.2-252 (N 37, mean 21.3). 135.0mm. LI 54.1mm. L2 63.1mm. T 19.4mm,
HL - 27.7-34.2 (N 38, mean 30.4), HW - 23-9* TV 23.3mm, HL 40.0raro. HW 34.4mm, HD
30.6 (N 38. mean 26.4), HD - 10.5-15.1 (N 38. 17.4mm, S 12.8mm. EE 18.4mm. NL 14.Vmm.
mean 13.2),S-9.1-12.9(N39,mean 10.5). EE- KW4.5mm.R 8. 1 7.SL 15/15.IL 14/14.TA 10.
62 MEMOIRSOFTHEQUEENSLANDMUSEUM
FIG. 4.Nephrurusamyae. KingsCkStn, southwestern NT.(Photograph: S.Wilson).
This species has been illustrated previously as Distribution
N. asper(e.g. Hoser, 1989: 74). Confinedtothehillcountryofthesouth-central
NT centred around Alice Springs (23°421S,
Pattern I33°53'E), extending north to Barrow Ck
Body. In spirit, fawn lo mid brown on dorsal (21°32'S, 133°53'E)(Fig. 3).
surface. Many specimens with a black band
present on the nape. Some specimens with seven
Habitat
tonineobscure,brokenerossbands.Theseappear
N. amyae favours open ground on rocky sub-
as a series of spots with each spot covering a
tubercule. Others with no indication of banding strates in open woodland habitats. It may be
orwithobscureblotches alongthe vertebral line. foundassociatedwithascatteredTriodiaground-
cover.
Fineblack lines form a reticulated pattern on the
dorsum and upper lateral zone. These lines fade
in largerspecimens. Ventral surface cream. ReproductiveNotes
Limbs. Lacking irregular erossbands distally. Clutch size two eggs. Females (NTMR33722-
Digits unbanded but sometimes obscurely mot- 23, NTMR5466) had small vitellogenic follicles
tled. on their ovaries between mid October - late
Tail. As forbody, with an obscure, broad, pale December(<5.5mmdiameter).NTMR2458,col-
crossband on proximal half. lected in mid March, had a greatly enlarged fol-
Head. Often paler than body. Covered on top licle in each ovary (the largest of these follicles
andsideswithfineblackreticulationswhichform measured 14.58mm X 13.0mm). Bedford and
an intricate pattern. These fade in larger Christian (1993) provide clutch data for a
specimens. specimen from Gardens Station, 75km northeast
of Alice Springs. Two eggs measuring 34.3mm
Etymology X 16.6mmand 37.0mm X 16.1mm were laidon
ForAmy Couper. the 17/18 February 1992. These weighed 5.6g