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Revision of theNeotropical

SnakeGenus

ChironiwsFitzinger

(Serpentes, Colubridae)

James R. Dixon+, John A. IfiestJr.* andJos M. Cei**

-

iJ:g;;1gg1"t

'. 3i:;91;5;*

-.

Fnheriessciences.rexasA&Muli,enbr, coresestation,

*turares.un jveBidad

NaciomrdeRiocuano,(5800)

Riocuarto.

TABLE OF CONTENTS

Intfoduction

Crtorira Adve isem nr lnfo.mation

Mrreridhrnd Methods

Sinopsisolrhe cenus Ci ianius Fitzjn}e\ 1g26

I{istoricalreriew

Familystatus

Chafactersummarv

Identfr catian af Chin nius

Key to the Species

Systematicaccounas

(-hitnaid, biarinat , t\t\iedl Chito ius cai aits (Linnaeus)

Chircnius einatus

Chitoniust.tinatusf

Chttontu\,ortnart)s\D,y tHltove]l

Chtrunttt.?'ul?t \ tl ionacusl

Chircnius fl.trolineatus(Jan)

Chi,04td\ fu,

Chttun,a,1yuu,,rr*"

Chtoniu,tu

cainat .' (LinnaeusJ aprlrr(Wemerl

" || Innreu(r

ut

fLrnnaeu.r

.

neq .ub.pecres

tcutontt tr.

Chircniusgtandisqu amis (p eterc) Chirc tus taetico b (Wied) Chitotiius laurenti, new species Chitonis ma ticotaRoze . I l.t,untu,nuhNettth,Schmidl rnd Wailef

. Chironius nultil,enttis cochrcnael':rogeand Romano

Ch;tonid \ n Ltlttventt t. tnvpaI d,

Baite,

Chitu iusmultit,ent smultterfir SchmidrandWalker

Chilouiusmulti,entis sepl

Ch;Iont\ qua(tti.annaru\ |Boiet

rrrbnar, new subspecies

pag

9

t0

12

t7

I7

l8

24

56

57

59

68

73

17

84

92

105

113

114

t27

135

l,l1

146

155

160

t64

168

t73

178

Chitonius

quadicatinatus nkelatr.nttis,

spectes

new

i'"n u' 4,tadi, ari40ttl, quala.r n a,t, I Boie /

th

tnt.mnt u' rLun utu\ (Wasten

I ntrnniutrtncenti(Boutenger)

Interspecifi c

relationships

Formation of phJ,logenetic Tree

aenu/ot( H,.tof) or ( entrutdnoSouth Zoogeographic Considerations Erol|lIion of Chirc lus

Intergenedc relationships

Summary ol taxonomic changes .

Lite.ature cited

Appendi\ I, specimens kno.$n, not examined

Appendix Il, specimens exaoined

pag.

)

'i

'i

181

184

188

195

r 200

, 213

, 211

, 221

, 233

r 237

, 231

, 219

,

250

INTRODUCTION

It hasrecentlybecomoquiteapparentthatcompfehensivezoogeogmphic studiesofNeotroplcalfeptilescannotbe lerformed efiectivelyuntil many lhorough,basic,ta-xonomicinvestigationshavebeencompleted.This fevi- sionconcernsthegenusCrl,"orirqa groupoi snakeswhichhavebeenlradi tionally,ta-{onomicallyabused.OfallNeotropicalsnakes!onlyCrr'r'orirrhave 10or 12scalerowsat midbody.Their uniquenessand relativeabundance makethis genusan interestingand expedientsubjectfor study. Ourintereslin Ci,Torrrrdevelopedwh,3nwebecameinvolvedin tryinglo idenrifya number of Peruviansnakes.Confusionin tle interpretationof folms in thegenusCr;/oflirrhadbeendiscussedbyBailey(1955)andWiest (1978).After usingthe keysofDonoso-Barros(1969)andPetersandOrejas-

Miranda(1970).v/esoonrealizedthatexistingliteralurewasnotadequatefof

identifyingthesesnakes.Much perplexityalsoexistedin the allocationof

namesin museumholdingsasseenbythewideuseof Chiro iuscainatus'dn(l Chinnius fuscus, bolh Linnean speciesnames,with comparatjvelylittle men- tion ofthe othorspecies.Sulprisingly,somemajorlendinginstitutionsof the worldperyeiuatedtheuseofsynonymssuchasIIeryetodnras Cchiro ius)ol

usednamesthat do not applyto Crirorirr suchasrefilinatus

whichbelongsin thegenusPr rrrer(Bailey,1955;Hoge,1960a).lt wasobvi- ousearlyin thisstudythatwhllothogenusitselfwasuniquedndwelldefined, the specieslimits werequitevagueand misunderstood.A factthat fuithei emphasizesthispointisthat49.34koI allChinniusspecimensxaminedin the cou6eofthe studywereeithermisidentifiodto speciesor unidentifiedbythe lendinginstitutions,but only 0.40/0of the specimensloane.las Chironius actuallybelongedto othergenera. Therehavebeenno majorpresentationson lhe genericrelalionshipsol Crilorlrr. Althoughsomeinterspecificrelationshipshavebeendiscussedby

Oydglet,1824),

10

BaileJ(1955) and Donoso-Barros(1969),thereareviftuallyno majorworks

keatingthesystematicsofthe

entiregenus.Further,thegeneralrangeofile

specieshavebeenpoorlyundcrs-

genusisknown,butthedistribulionsofthe

stecies,djstriburions

unfeliable

astospeciesidoniifications.TheseincludelheworksofDa CunhaandNasci-

menro(1978,1983,1984)for

(1979)fbf Surinam:Chilpaux(1986)for

Moreno(1988)forColombia;Via a era1_(1988)for CentGlAmerica;Carrillo

deEstinosa(1989)for Peru.Thelackolecologicai,behavioral,biochemical,

andorherspecializedstudieshasalsohelpedto

rvithinthegenus. Speciesof Criforlrr havebeenmentionedin a numberoi

diverseanatomicalrolatedstudies,andfor

used,ii.isimponantthatiheirsystemalicsbetrealedto

fusionin the allocarionof names.

tood andnoi well deiined.Most ofour loowledgeol

comesli-om regionalstudies,whichareoccasionallyincompleteor

eastemAmazonia;Moonenandvan DeuNen

Frenchcuiana;

porez-sanros and

impedelaxonomicstability

agroupofsnakesto besowidely

removeexistingcon

C1l//IONIUS ADVERTISEMENT INFORMATION

Thegenus Crilorirr canbedefinedasamediumto1aryesized,te[estdal_

hemipenishasa singlesulcusandlobe,

lupil

2b8subcau_

presenceor

arboreal,agl]lhous colubrid.The

nakedpocketproximally, spinouscontrally,calyculatedisralry,andiswithoui

capitationor apicaldisks.The

colubridheadscales.Thereareaboui123ro 196veniralsand96b

is round.Thereis alull comllenent of

dals.The midbodydomalscalesarein 12of l0 rows,with lhe

absenceof keelingandapicalpitson dorsalscales,dependinguponihe spe_

Crilorirr isdiagnosedastheonlyNeotropicalsnakegenus with l0 or 12

scalerowsatmidbody.Thischaracteraloneseparates C,itori6from ellother Centraland SouthAmericansnakegenera.

\eor-npical di.rib. ,ion.e\tendingtiom lhe -o thcrn

(h,ro,ttu, hd. d

coaslol HondJ-riIn urugui) drd

lbund eastto the Andesin ot aroundthe AmazonBasin.Chirotlis has reachedthe Islandofcorgona alongthe PacificcoastofColombia,several islandsalongihe Atlanticcoastof Brazil,andTrinjdadandSt.Vincentlsland in the CadbbeanSea. Habitatprefercncevadesextensivelywithinthegenus andwithincertain

species.Lo\rland tropicalrainforestis preferrcd

montaneforestsandsavannalopenlormationsarcalsoinhabited.Theeleva-

tion nnge ol thegenus isfrom sealevelto 2800m. The rangeandhabitatoI

exchtaxonareexpldnedin greater detailin the

nolhed.tAJgenltna,Mu.t spe.ie re

by Crtr,orrrr,bur humid

speciesaccounts.

A general descriptionofthe totalsumolvariationin thegenus is:Total

lengthvariesfrom 274mm to 2718mm andthetailis26.9-43.tpercent ofthe

ll

totallcngth.The ventralsvaryfrom 12l-196in malesand131-190in females. The subcaudalsafe paircdand varyfiom 96-208nl males.and 99-205in femalcs.The umbilicalscafis normallythreeventmlslong.The analplate maybedividedof entirc.Theamountol keelingisquiievadablc,andpreseni in allspeciesaxseptC,rrrr?/ri, Malesgenerallyhavemofepronounccdkeel, ingandir somospecies\Vithnofe scxleforvskeeledthanlemales.Apicalpits

areabsenlin C

in all otherspecies. The nofmalcomple cnrofcolubriddo$al headscalesare:supralabials nofmally8 or 9;supralabialsentefingorbitusually,l+5+6,4+5.or 5+6;infra- labidsnormally10of 1ltrvopain of genials;infralabialscontactinggcnielson onesideusually6 or 7; usualhI preocular;prcoculatusu.llynol in co tact wilh iiontal; usually2,somelimes3 postoculars;uppefpostocularusuallyin

contactwithparietals;tempofalsvadable,usually1+1or1+2.sometimcsI+l+

I or 1+l+2;usuallyI loreal;usually410postcephalics.Thereare24-51agly- phousnaxillarytccth,and26-46dentafyteeth.Thereafel2 or l0 scalerowsat

nidbody,withscdlerowfeductionquitevadable,usually12-12-10,l2-12-8,10

,?ortrold,presentor absentin C 4uotiicainatus,arTd,presenl

10-10,of l0l0-8. Hemipenesof ell speciesafesinglelobcd.cylinddcal.anddillefeniiared. Theretfacledlengthofthehemipenisrcachestosubcaudals5-13;,41./ /r?./r/ /errs ,ragrr.i originatesbetwccnsubcaudals15-32.The sulcus Dermaticus simple(notdivided)andusuallyterninatessubapically.A basalnatedpocker is presentolposite the sulcus.The basal fofiion of lobe may be with ol withoutminutespinules.The cntirecentralpoflion oflobe containsspines $hich number4 12 ler longltudinalrow.The cntircdistalportionoflobehas calyceswhichnumber8-26pcrlongitudinalrow-The cdlyceshave froximal spinousfringesrharchangeto papiUatefringcsdistally. Inlraspecifisally,coloraiionandcoiorpatlernsmayormeynotbevariable, Fofestlorms iend to be dafk,usuallygrcen.olive,or blackdorsally;open ibrmalionlbrms areuniform yellowbrown,olivebrown,of havca stdped paite .

Speciesof Cir"rtrr

are oviparcuqierrestfial-afbofeal,mild to vety

aggrcssivein disposirion,andgenefallyslcepabovegroundatnighlin treesof bushes.Thesesnakesareactiveforegersduringdaylightandpfeyalmo$exc

lusivelyonfiogs.Crn'orlrr,herein,iscomprisedof 13speciesandatotalof 20

tAxa.Chironiusbia:trhatus(Wied); C. cainatus utiratrr (Linnaeus); C

rolrrds (Wemet; C Cr"/it(Hal1owe11);C a?1 t s(Linnteus); C. Ia\iali nea-

tus (Jan); C. fuscusJuscus (Linnae]ls), C. f.

gtafidisqudmis(PeIefi);C-LaericallislNied); C.laurcftarew spccies;Cno, ticolaRoze;C.nultiwntrisrir.irdndr Hogea]1dRomano;C-n.Jovedtustsai- leya (- m.mull entisSchmidlandWalker;C m-sQlert"loralir,nowsubspe- cies; C. quddti inalus nda?/r,r,crt]J, new subspecies;C. q. qtradticannaue (tsoio)iC r.?r"/rr (WaSlef);andC rir( rt(Boulengor).SinccC /drfsrli, C

/1d,

leuu)nelaprr, new subspecies;C

t2

multivnttis, anrl C. vincenti sharemany uniquely derived chatactef states, thesespeciesareplacedh ther,r/rn,entnrsleciescomplex,aconvenientway oI showingthe closerelationshipamongthcsespeciesbut which hasno nomenclaturalstatus, In 15of the20CriloTrtrrllrxamalesattaina largefsizethanfemales.The smallestspecimenexaminedwasajuvenilelnaleC manticola2j4mminLotal

lengrh.The largestspecimenwasa maleC grdrdtrquamis2718mm

length.Smallestandlafgestspecimensexaminedfoteachtaxonarcpresented

in the sleciesaccoln|s.

jn

total

MATERIALS AND METHODS

Ofrhe 2978sDecimensol Crilo'irr usedin theseanalyses,lessthan400 \yerenotpersonallyex,rminedbyus,Mostspecimenswereprocured onloans

States,Europe,

fron museumsand educationalinstituiionsin the United

SouthAmerica,andCenl|alAmerica,andthesecollcctionsafelistedal

the

end ofthis section.A list of lnown bur unavailableCrjlorirr specimensis

providedin Appcndi\I

ity,andmuseumnumbor)inAlpendixll.

andslecimensexaminedarclisted(_byspccies,local-

Weexaminedoracquiredinforma

tion on19ofthe28i),pespecimensolell speciesandsubspeciesnamesthatare

associatedwithCir)oaiff.ThefeDainingninetypescouldeithernotbeloca! edaftere{ensivesearchiigof weropositivelydeierminedto belostor des-

scalecounts,aspectsof anatomy,measurements,ralios,and colorpa!- ternsofpresefledspecimenswereutilizedin analyzinginterspecificrelarion- shipsandinimspecificvariation.Duelo timefacto|Sassociatedwiththisstudy of to thc poorconditionofcerlainsnakes,completedatawerenol obtaincd fiom all specimens.

Ventralswer countedfollowingthemeihodofDovling

(1951a).Single

half-ventralswefeignoredsincetheyindicaledvertebralaberaiions(ting, 1959),buttwohat scalesofa dividedvenrfalwefecountedesone.Subcaudals arepatedand\refecountedonihe leftsiclesrarlingwiththefirsrscalebehind rheanalplatcthat contacteda scaleon theoppositcside.Fusedsubcaudal lairs wereincludedin all counts,aswasthe unfaired,telrninalscale. Conditionofthe dorsalheadscalesisconsisteniforallspeciesandwasnot rccordedtbf mostspecimens.Thellumberofsupralabials,supralabialsenter- lngthe orbit,inffalabials(lastinfralabialdeteminedasthelastscaleatleast half coveredby the last supralabial),infralabialscontactinglhe genials, genials,pfeoculars, !rcocularscontactingtheliontal,postocula$,postoculan contactinglhe parietals,lenporals(scalesbehind lostoculars andberween thepaietals andsupralabials),loreals,and lostcephalics (scalestouchingthe postedormarglnoftemporalspa elals)werenoted.

A slashmark is usedto separatecharactercountstaken on oppositesides

oftheheadwiththeleftsideindilratedfirst(e.g.,supralabials9/8signifiesnine

supralabialson the lell sideand eight on the right). A hyphen hdicates range in acharacter(e.9.,venlrals135-155means135to 155ventrals).Segmenisofa chalacteranangedin a seriesarcseparatedby aplus sign(e,9.,temporals1+2 denotesone primary andtwo secondarytempomis).Teeth on the left maxilla ofmany specimenswereexaminedandcountedi, rirr usingthemethodof Myers (1974).The number of dentary teeth (left side) were noted for a lew specimensof mostspecies.Conditionofthe analplate(singie,divided,or incompletelydivided),positionofapicalpits,anddegreeofkeelingwerealso recorded,

Wlen no evertedhemipeniswaspresent,the sexofeachspesimenwas determined by maling a post-analslit between the anteiior subcaudalpai$ andcheckingfor thepresenceofrr rr, hemipenes.Hemipenialmorphology, includinglength(bynumberof subcaudalsinvolved),shape,conditionofsul- cusspennaticus,andomamentationis describedaccordingto thelermino- logyolDowlhg andSavage(1960).1,r,t/ hemipenesofat leastthreespecim- ensandseveralevertedhemipeneswereusedin thedescriplionfbr eachspe- cies.Eggspresentin oviductswerecountcdby palpationor by incisingthe midlineoftheposteriorventralsandvisuallyinspectingthem.Witl thislatler method,the greatestlengthandwidth of the eggsweremeasured.Snakes withobviouslyenlargedbodiesin thestomachrcgionhadtheirstomachcon- tentsremovedand examined.Noticeableecto-andendoparasiteswerecol- lected and identified. For consistencyabbreviatedscaleforv formulaswefe stardardizedby counting the nnmber of dorsal scalerows over the tenth ventral behind the head.overthemiddleventral.andoverthetenthventralanleriorto theanal p1ate. Completescalerowformulasindicatingtheexactpositionofscalerow reductions ard additions were constructed for most specimensusing the

melhod (with slight modiflcationt

formulaswercsummarizedfollowingthemodelpresentedbyThomas(1976).

Ali measuements (ir millimetres) were taken on the left side where applicable.Snoutvent and tail lengthsweretakenfrom specimensstraigh- tenedagainstameter stick.Othermeasurementstakenwith avemief calipers weresnoutlength(straightlinedisiancefrom tipofsnoutto posteriorendof interprefrontal suture),frontal scalelength, headlength (shaightline distance from tip of snoutto posteriormarginofjaw),headwidthbetweenthelateml marginofthe supraoculars,headwidth betweenthe e),1emalnares,rostral scaleheiglt andwidth,andhorizontaleyediameter.Specimensin the300-500

suggestedby Dowling (1951b),and these

mm snout-ventsizeclassof most speciesof Crilo,'?lrs generallyshowontoge-

netic changesin color patterns and degreeof keeling. Therefore, the term juvenile isruther arbitrarily definedherein asspecimensbeiow 400mm snout-

vent lensth.

\l"-1

.

',1r"'. "]

e.r

lr'

h-

ne i.,: rot,.i.r, (.e

r.

tD.

\.o

r\r

(

c

r,t.r

<,t,.,d(1

:

.J.,.,,no

rro n

I

t.o.

it.o1.

J<,1.S

.

:

m'

,heri.r]

i.

.i

gcL

.t.-c,r'<n.,".o

il-,i ,n.ce

\iI .rrron.I r. ht\lL 1

nt tcrrJls

The51rorr/rrdi!isionoferchtaxongeneray

r.\

ir.

\tr.

r'

e.l,rIn.h.

n.(,.tL

\i.-.

l. ,,c.1,\.rr,

'lr:je

I

rreJ

n, :

,

ll-

r

,rr . li .

-.me,ot

jnctudes onlyjuniors!no_

\;,

nt.Ito1r.r-

.

n.r|.

r.tr,.n:

nymsandthe dillcren(gcnericcontbinxtionsthal hrvebcenuscdlor ticse

names.Thefi^t felifcnceto cilchIel1)ein ihc slnonym).isgi!en,as .e pel

I"

r:1,1rIc/.,

the cu.lliDg museum_specjnlennunlbef.condition.sex.rypclocalit).an;

ihat orc laxoncin bedls

lirguishedfionlullothefs.t--achdoronurangc1nat ir

collectof.The 1ltuArr.rA.indiclresthee.rsicslwlrs

t,e^- jtj(.ntllr(

on-.t,.c

nr.r

.-,.,.r

lheDi!1r?rllldlsecti;I

i

.el

n,

pe,l

.r

o

!u\\h

. te

,.,r.'

lh .r cderI.or

rnd Seibeft (1972) lvasl.rrgclyusedio

Tonesocclr|ledbl

strlcllyufon elevalionsfccordedrvilhthc specimcndata.

descdbethe hrbiiar or \;gclalional

lirit$ ofa

stecjcsrre based

erchspe

eachspecics.Ahjludinal

Thr ,cr./l/rid, prcscnlsthcnumbefofspecirncnsexaninedfor

cresor slrDsDectesandI

Ludedir

sumnafizationol thc characicrslof cachtxxon.Inc-

lhisscctionisinlonnalionon mlnimunrandmaxi]nunrsize,fel|rile

andfelatoddara.andrhenumberol.matdiaryarld

.h r

r(.

rc.ri,c

.1e-ro.iJr

t

{h.

{

. /

,

colof.ind/ofpltrem

r. n.r ,\1.r-. \ .o.r |n . ||J .u u" n IIe . r

andscruajit,

rrh.,1.

e g \ .||

-

tfcser.ved specirnensofe.rchspeciei.l; pxtlcrned

ofbandsarcnoted.andlhelengthind width

iefns of ihe number of scalerorvs

iail l ngth.scalccounh

denral)teeth.Ontogenerjcchangesin

d, _' t'.r

r l,mrt-,.,

retrlrscnlativctllrlnbcrol

snrlesIhcnumbcrand folilion

ol rhcsema.kingsare exprcss dir

rn!olved.

r'r.

Colordcscrittionsol li\,eCrn.rrlr/rarebasedon lteraturcaccountsand

avlritrbleffon thoscwho havehaclfieldexfcrjence$,jth

fhotogfafhsfircijiraicdlhe doclrmenrationof

rnlirrrnlnionmrde

thcsesnr(es. Iliuslr.alionsrnd

Ircserredandlivcspecimcns.

L

_r'rn

.-

I

la,

,',,|

a

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ri

.^n.

m.r.(

,

i.,u

i

,r.

ieodr.tLr

r

troru.\

ti

\.

1

In.r ,t.ci,i

i

J.et\ J

lemr)c. bj u.\:

.i

crh,,C\

_th<\t.tr.i.:t

\n.h.,\\,cnt15\\r.r,

\mJdh

.170V/6conrurcr. Asignificancelcletot0.0tras Lrscdtbra|r tes|S.Thissec_

bringinglo ljghrpossiblc e!olurionar.ilydistillcrunjts

wrthina specics. /-/? H/r1r,fnotcsincludea\,rjhble inlirfmftion onbeh.rvior,iimeofacliv-

ily. se$onrl occurferce.relfoduction,cggs,fecdinghabils.incLecto_ancl

endopafasltes.thcnerrd?lJsectionprovid

s aplrre fbr presenting othcrinte_

resrng.perlrncnt, or rclated$orks nor citeLlelsc$hefein ihis siud),.Com

mentsaboulthc et!.molo$oflhc speciesnumesa.eusualllinclucled.

liof ls info

nnt jn

Il isudbnunaie,bul neverthelessalacithatvefyfewmajofrevisionsarc ableto drawulon all availablecharaclers,melhods,andtcchniquesfbr ana lyzingdalaandlbrmulatingconclusions-Hopefully.behavioral,anatomical, chrcmosomal,biochemical,andothcrstudics$'ill somcdaybc uscdlo iesi and amllily ouf resultsand interyretations.

Citedinstiturionswith thoir co cspondingcuraloNafeabbrcviatcdas ibllows:

AMNHrAmcricanMuseumotNaiuralHistory,NewYorklDr. ClnrlesJ. Cole,Mr. GeoryeW.Foley,Df. ChaflesW. Myem,Dr. RichardG.Zweifel).

ANSP:Acadcmyof NaruralSciencesofPhiladelphia(Mr. EdmondV. Mal- nale,Dr. ThomxsUzzell).ASU: AfizonaSiateU.iveNity, Temte (Df. M. JackFouquetle).AU: Auburn University(Dr. RoberiFL MoLrnt).BMNH:

(NaturalHistory),London(Mr. AndrcwF'.Stimson).CAS:

BritishMuseu

CalilbrniaAcademy of Sciences,San Francisco(Df. Alan E- Levito

CENAI: CentroNacionalde InvestigacionesIologicas,BuenosAircs (Dr.

AvelinoBaffio,Dr. MartaE. Miranda).CM: CarncgicMuscum,Pittsburgh (Dr.ClarenceJ.Mccol,).COH:C.O.HandleyVenezueldnProject,atUSNM,

Washinglon,D. C

Iornia,LosAnseles(Dr. Cafl S.Lieb,Dr. JayM. Savase).FML: Fundacion

MiguelLillo, UniversidadNacionaldeTucuman(Dr.RaymondF.Laureno. FMNH: Ficld Muscum of NaturalHistory,Chicago(Mr. H,\7menMarx).

GIUS: GeogmphischesInstitutderUnive|sit;ildesSaaflandes,Saafbiicken (Dr. PaulMiiller. Dr. HaraldSchreiber).IRSNBInstitu!RoyaidesScienses Naturellcsdc tsclgiquc,Bruxelles(Dr. XavierMisonno,Dr. CasronR. de Wnte).JAP-GOV:J.A.Pelers-G.OrcesV.EcuadorianColleclioqatUSNM,

Washington,D. C

Lawrence(Mf. JoserhT. Collins,Df. WillialnE. Duellnan). LACM: Los AngelesCounry Museum (Dr. Robet L. Bezy,Dr. John w' Wright). LSUMZ: LousianaStateUnivemilyMuseumofZoolog,T,tsatonRouge(Dr. DouglasA. ltossman,Df. J.P.l{ichafdThomas).LUZM: LundsUnive|siteis ZoologiskaMuseum(Dr. Bo Stille).MBUCV: MuseodeBiologiaUniversi- dadC nlraldeVenezuola,Caracas(Dr.J.Raconis).MCN: MuseodeCiencjas Naturales.Carecas(Dr. A. R. LanciniV.). MCZ: Mlrseumot Comparadve Zoology,HarvardUniversity,Cambddge(Ms.PatriciaG.Haneline,Mr. Josd P.Rosado,Df. ErnestE. Williams).MHNGj Museumd'Histo'rcNatufelle. Cenave(Dr.VolkerMal}nen).MLINLS:MuseodeHistodaNaturalLa Salle, Caracas(Df. C. A. de Lima).MILS:MuseodelInstiluiodeLa Salle.Bogod (Dr.HermanoNiceloroMaria).MN: MuseuNacional.UniversidadeFedelal doRiodsJanedo(Dr.AntenorLeilaodeCarvalho).MNHNM:Museo Naci- onal de HistofiaNaiuml, Univenidadde La Republice,Monievideo(Dr.

FodericoAchaval).MNHNPTMuscumNationald'HistoifeNaturelle,Pads (Df.

).

CRE:CosraRicaExpedition,UnivclsityolsouthemCali

KU:

Universityof Kansas,Museun of NaturalHistory,

R.

Roux-Est ve.Df. JeanGuib6).MPM: MihvaukeePublicMuseum

aDf.M.\ A.\ic{ersonr \4RSN.[4u\eoregiond]edi5cientenarurali,Iof

rDr.LtendUa\e :.Dr.francoAndreoner.VS\U:

ildlurJle

ve'lebrule /oolog).Uni\erisr\ofCdlilont.r.Be"kelevr\4r.SreDhenB

ur

unt!e^rdadedeSioPaulofDr. pcuto L. VanloLini)\4/Ul

loga5'slemnticadeUUri\ersilaoi

lne

frdnco{ndreone)M\8.

Dr. OlivioRieppel).NMW:NaturhistorischesMuseu-,Wi"nb

demann).NRM: Naiurhistoriska Riksmuseet,Stockholm

lam).RM\H

no

\4Jseo Ci\icoo Srofia

l\,1useurnof

Rulh,

UracomoDoria. CenovaIDr.LitiaCapoccjlr.r.V\,/:

M/l

Krchlrdu.sage.-DrDdvidB.Wrle,.

5p: \4r.eode,/oologt,raa

: Museodi7oo_

(col,ec.ionno\aloanedro

[orno. lorno

vu:eo.Regjonale di Sf,e[zen,rlurJli.lorinorrDr. |end

Gdvelti,Dr.

\bturbisrori\ches \4Lse-m, Ba.etLDf.[ . Raim.

Gjr. Carl ^".ii"l Edels_

Rij".mJseumrrn\aruu-tirkeHisro.ie.f.iae.LOr.V nuri.

lDr.

ChdrtesR.

TCWC: TexasCooperativeWildlifeCol""tlon,

Jo

f i'"u,eaU Un;_

i;;s

BoJCt.Vr. l\4ile\4ccoid/ f\Ha:

of le\a\. AusrinrMJ. CeorgeD

MJnirr. tU: futJnet n,rer.iil,\en O.jean.

Hoogmoed). SDNHV:SanD,ego\arurur Hi,loD Murum

Crur ) r.S\,1\5:Srtuliches\4u\eum fur\aturkunoe. SluIgan(Dr.Heinz

Wermutl)

\ersirJ

\alJml HisloryCollec.ron.Uni\ersy

Codege SlationrV.

tt.rumgardrer.Dr.Robe"r[.

IDr.tiarordA.Dundee).UCT(B:t ni\ersiddddeConcepcion,lrstilJtoCen_

.ru1de BiologiarVr Museum.Biuldef (Dr.

Mu\eum-oiNdlur.rlHi)lory.Sto.rstDr. Robent. Dubo

0e

Dougla\C Robin\onr UFiF5V: I nirerisrtof florio4 i.onda Srare

Mlrlgum, _cainesvi e (Dr. Watter Auffenberg,Mr. f"t,j,

UIMNHTUniverjstyof Iltinois Museumof Nttuml

D,ona,df .

vr:brganMuseurnol,/oologJ. AnnArborIDr.ArnoaC. Kluge,Dr.Ro;ajd

tho"nd( ekdlo\ict. ).U(\4. I niver.irJofC.tlao

I. PaulVastinr.UC\4NH:Unr\et5rlJolConnecthur

r.UCR:I

ni\ers,dad

(Dr.

U"yf*).

History,Urba;; i;;.

t ;,i;;.ly;i

(

oslaRca.VLseodeZoologid.CiudadUni\ersrlafia-Rodrigo I dcio

e.

_HoFmeiner Nt<.Doroth) M. Smirhr.UMVT:

C.\ussbrum). USNN'l:\ational Museumot

D.C

iougn.uf.(jeorgeR.Zug) UIA: I

ol ve-lelrrdles. Arlinglonl\4f.JohnL.

Unne|5rJ,ofuldh.lMu.eumoJloorog).sdlI

U/vK:

N,rlLralHisloO.$ashington,

r\4r.RonaldI.Cromoie.Dr.w. RondtdHeler,t\4s.Fr,rnce,L M;Cut_

niversil)o,Te\.r(dAjlinglon(o

Drfting.Dr.Willi.jm l-.

eclion

p\bum). I I

:

akeC t) {Dr.Joh; \1.I egle1.

F. l\

Un\ersiletel\ Zoologis"eML,seum.Kbenhavn(Dr.

der Humboldi_Uni;rsitiit, Berlio

Hans_Withelrn Koeocle,Dr. W.

Biaer_

trupl.ZN44:Zodlog(b M-\eum.AmslercdmrDr.A.Ame,ing,D-.D Hit.e_

nius).ZMB: ZoologischesMuseumand

fDr. Cunther Peter-).ZII\,lH:ZoologischesIn\lilul und ,zoolog'<che.

!ruseum. Unrref)rtdtHamburg (Dr.

Lad eesj.,/Mt | :ZoologisdaVu.ee. I ppsalaUn.rersirerrDr.I arsWalLint.

/MZ_ ,/oologisches\4rseum. UniveAit;tlurich rDr.

ZS\,IH:Zoologi\cheSlaarssammtJng.Herpelotogi.cheAbreitrng,Vitn.hen

{DJ.Ulrilh Cruberj.ZVt Montevideo

HansjitrgHol,/./

: Zoologia-Veneorados (

iercid,R .riVNHNM,

l7

SYNOPSISOF THE GENUS Cll1RON1USFITZ]NGER. i826

GeneicSrnanJ.n!

Crn",r"s F zinser.1826:31.TrDe{peciesColal.r.drr,ar

Linna.usby odgnal designation.

Et?.1.).t)rasBaie.1826:231 'lrFe-s!.cics notdesieraled.]1?rr,.r,drydrWusGr.1E10180.Endr dlrior ot fQ.rodryas!oic. Tyre srecieslor desigtuled.

,yd.,?rr\!'asrcr,lElOl82'l.rpespec\esCatah.ts4lrtrrtrrLinn.els[(:Cri]?,r^Jr!.rr(tit

naeus)llry monot]'L)r.

trlorrrJ

lJitzinger.1841:26.T)-lc{nccicsard?,rerladrollrr 1\'iedb} oisi.d

designationxnd

Pblbsriz Core.1862:149.T!!ie sleciesPlull,rtal/d!.(.A

lechno"ius quonticahd|1!qru.l

,.d/1r.rA lBoie)lb' oriCinaldesisnalionmd mono!'!t.

HISTORICALREVIEW

Snakesof drc colubddgenus Clr'rorirr wercodginallyassignedio ftc

genusCotur?/'byLinnaeus(1758),whonamedlbut specie

C. etuletus,C.fuscus, Lnd C. satumtuus.Nl ol rheseate tecogrlizedasralid speciesof Clrtirrlrr exceprsatuninus, .rthich hasbeenplacedin ihe syno-

nlm-vot.r6crs herein.Coluberun atusotLinnJeus(1758,U66)wascalled "ChifonsNatler"byMeffem(1790).A shorttimelaterDonndodT(1798).\i\,ho

referencedMerrem's(1790)worf,,probablyuiilizedMerrem'scommonname

and formulatedColubetchito ius.Donndorfls genus-speciescombination

appearedin thelitefaturconlyonceand Crlrrpr.rdrrdtrr wasagainusedby Menem (1820)andLichrenstcin(1823).Fitzinger(1826)stecificallydesignat- edLinnacus'Colrrelcdtrdlu.tasthelypc-speciesofhisnewgenus Clironirr. Bt' 1827overhallthe presentlyrecognizedspecicsofCrlldrnrs hddbeen naned bylinnaeus (1758),wicd (1820,1824),wdgler(1824),andBoie(1827)

andvere placedin fte generaCrlrr l

(WdEler, 1824),Chhanius(Filzinger, 1826),and t? etakras

Latef,spccies,oflhis genusweferefcfiedto asllcrp rrdryds(Wagler.1830),

Maclolr 0Naslef, 1830),Hllophis (Fitzinger,1843),Dendraphis(D\rJet.L

Bibron,a,'1dDum6 l, 1854),Pfu.//or;fd(Cope,1862),.ttlor.r (Petc^,1868),

and Z ,r.tr (Wemer,1696).

ThegenosHerpetod ry asvaswidelyusedbymanyearlyherpetologislsand natumlisls.Thisnamewasprcvalenlintheworksol Schlegel(1837),Lichtens,

tein(1856),Gunther(1861),Jan(1863),andJanandSordelli(1869).Cope(1860,

\CalubetcatinaLu.t,

(Linnaeus,1758;Wied,1820);Nar?-r

(.Boic,

1826,182'7 ).

1868,1869,1876,1886a,I886b,1887e,1887b,1893)usedl/e?erodryarinhisNeo-

tropicalherpclofaunasludies,greatly

name.Boulengef(l894a)causedamisunderstandingof Hetpetadrras(:C hno

jnfluencing

thc use of this generic

l8

ti,rr, by placing anumber ofnames in s).nonymyand delimiting the genus to

five species.Only three of

ore i\ norr Crrorirr. It ,5 inlere.tingthatBor.r.engeasI l8q4d| -C;lalogue of

snakes",consideredafahlycompletecompendiumofthethenknom

ofsnakes, madeno m ntionot Cc,lubetexotetusLjutlaeus.a vaLialsDeaiesas

recopized herein.A fe$ olher( $ho perperudled lhe nseol H?t;eroJNa\ s ereBoellger| 188).I8c8r.PerdLca{t8c5.t8q6.1897a.l8q7b.tq04),BarUour (19141915), Boulenser(1915),BarbourandNobte(1920), Seri6(t92ii,ctiesch (1925),and Schouten(1931).

the five arevalid species.two areconsDecific.and

species

The transition in the use of Helpetodryasto Chinn rrr wasquite

apparent

in the Literaturelrom the 1920's.Ruthven (1922)used Crlforlrr

for the first

time in nearly100yearssinceFitzinger (1826),andin aratherobscurefootnote

he called attontion to the fact that Linnaeus' Colrra rdtrdlr.,

ciesof

wastvDe-sDe_

lhegenus Crliontr, b) originatdes.gnarion.,qmaratrlq2) reiisessed

concludedthat Clilorirr

fhevalidity of Eryetodr|asandHelpetodryasa^d

haal

priority overthe lomer two genera. Chilo iusFltzinger

were both named in 1826;Amaral,sreasonfor establishinsCrtloriur ^nd asthe

\dlidgeneric namewa5baseduponthefaclIhalfirzingerie.ignareoaq pe-

speciesfor Crilorlff,

EryetodtyasBoje

but Boie made no t)?e designation.

A rcview of tle datedLiieraturercvealedfurther evidencethat Crilorirr

l]as piotjty

introduction in Fitzinger's'Neue ClassificationderReptilien".

doesnot help determinewhetherFitzingels or Boie'swork camefirst

becausethe publishing dateofBoie's paper,Erp6tologiedeL'ile deJava.was

only indicatedby the year,1826.But

my friendHerrB;iewiube

overEryetodrr)ar.The date4 June 1826appearsat the end ofthe

However,this

anothersectionof Fitzinser,sa1826)

tbe ne$speciesol

inlroducrionisrran.tated _

published next", in a separatework, which shall appearunder tbe title , Em6_ lologiedeJava. andBoiet 1826)$role'Finding hose\er wilhinlhesh;1 heatise,published in Vienne by M. Fitzinger, and entiiled: Neue Classifica-

futuretenseby Fitzingerandthepast

tion derReptilien, ".Theuseofthe

tenseby Boie indicatesthai Fitzingert work preceded Boie,s.and hence

Chironiushas pi'otity o'ter ErpetodrJas.

FAMILY STATUS

A1l recent authors placed Chitunius]n the famtly Colubddae basedpri-

marily on hemipenial, osteological,and dental characte$. This

includedin lhehighertdxaot lhefolo$ingselectedreterence<.i

aenus ;as

Wagler, 1830- Thecoglossi(family). Fitzirget 1843- Dendrophes (family). Dum6ril,1853- Isodeotiens(famity). Gunther,1858 - Dryadidae.

19

Cope,1886b,1891,1895,1900- Colubddae,Colubrinae. Cope,1887a- Colubddae,Natricinae- Boulengcr,1894a- Colubridae,Colubdnae- Dunn, 1928- Colubridao,Colubrinae. Amaral,1929a- Colubridae,Colubrhae. Roner, 1956- Colubridae,Colubrinae. Underwood,1967- Colubddae.

Doqling.la6- la-<- Coluondac, ( olubrin.e.

Dtfr\'rrIoN

Crilorirs. medium io largesized,tenestrial,arboreal,aglt?houscolu- bdds;hemipeniswith sing:lesulcusandlobe,nakedpocketproximally, spin- ouscentrally,calyculatedistally,andwithoutcapiiationofapicaldisks;pupil round;lull complementof colubridheadscales;123-196ventrals;96-208sub, caudals;midbodydorsalscalesin 12ot 10fows;keeljngandapicalpitsondor- salscal spresenlor not.

DLAcNosIS

Crlforlrr is theonlyNeotropicalsnakegenuswith 10or 12scalerowsal midbody.ThischafacteraloneseparutesClllo,?/rsftom alloi]lerCentraland SouthAmericansnakegenera.

DrsrRlBrmoN

Ci,Tdrtrr hasa Neotropicaldislribution,extendingfrom the northern coastofHondurasto Uruguayand northoastAlgentina(Fig.1).Mostspecies areloundeastoftheAndesin oraroundtheAmazonBasin;Tablel indicatos thc sleciesklown fiom eachcouniry.Crrorlw hasreachedthe Islandof GorgonaalongthePacificcoastofcolombia,severalislandsalongtheAtlant ic coastolBrazil,andTrinidadandSt.Vincentislandin rhoCadbbeanSea-

Habital lreference varieserlensivelywithin thegenusandwithincertain species-Lowlandtropicairainloresiis preferredby Chiorirr, but humid

montaneforestsandsavanna/openformalionsarealsoinhabited.Theeleva-

tion rangeoftho genusislrom sealevelto 2800m. Therangeandhabitatof eachiaxon areexplainedin greaterdeiailin the spociesaccounls.

DtscRltrloN

Total lengthrangesftom 274mm to 2718mm; tai126.943.3percent of totallength;ventEls123196in males,131-190in females;subcaudalspaired.

2Q

!r^Irotl

X

XX

 

xxxxxx

x><

 

X

 

x;t

X

XX

X

 

X

XX

XXX

 

XXXX

 

xxx

3

XXX

X

X

XXX

XX'<

!

 

;<t

- i

:

l:

r

:;

:E::

1 -

*,i,*i:i!:s:iii:

I - Disiributionofibe senus CrtoDirrFitzinscr.

22

i"Eliiiilii:!l?fr :l',tiiitii::i,iliiti{;ili::f,i,J.,ti;.:,,"

rry!ff'l#lifi#i5:,:TTtH,i",*ff[i:?:":lnJ,,f*t*la

trffi:ffi::Hfi;ffiittf;Sitii'#i:#":.fiTH,:i:",,li :ii?T;

;,tii';i]';lti;Ht:;i::i,xt:ll,i,,i,i,.:.

I

i;i.,,?i;;:*Tl*j;:l{:

nol il Lorllcl sith

.ron_

;',,::il';,:;;lili',i,i,";ll'illl,,,'iilLl'i:i::i;"i;illt; .ii;i:,

,"'.';i,T.lJ ,lll,1;,:,i"ii'":i,'.#:.::$:1,;ji;:i l:::Tlx'itrifi

p"!ocutarl pteocrrdr -5ral,L

qurre\rfidble.

.r.uati1t2

t2_,0.t.t_r2.8.tUt0_t0,ur

t0_'|U.8

;"5i; ;;;;:;::l;tj.i:il.Jji:::;

i,ided,ancu.u,;.).e',m,.

::ri::+li:i;i:1.'1,Til::il'.:i::

:ll^.".,.9:l:ll J',:-tl,.pcrmaricrr,.imI e,nuri

H';;,i'l:;il1t;.ti1iffl:,".';;i::ii:::;1,"J{:;:"ffi ;:1"xj;l:",:

il:::i;::ili::i,liili:;J"1t:iJ",i:"lillL:u,

;T];i;::,J1';T:l

no,be\a'iab,e

:',,:i::

mcl,,D;nuu('finge.chJnpingro

papilJare fihge.

ci.tllli

,",.ilT,1;l jl,l,;,1i3:::J

o'',,.no*.i,",niri;:-,;

i,:lfflllfl:''-dvo'm:'r

.;;l";;;.

":]i::r:JJ]::1 iH

"cc,":*" ; ;;;;.ffi;,hHiliii'.i::;il*," ,;l*Ti,##,,f;:3

Speciesof

Crnurirs areovipafous.telret

,.,11H:ii:;,;:il;j;:il,it;tr.'"i,;i*fi :1ij:iJ.::j[)

ot 2lttr\

tta,^)t

t Chirantus bkdtitlotut

twrcd\

C cot

fi:

,v\e".e,r:c.,.,,,'-,,n,',"*",,,,.li,.l"l

tnedtu.tJaEt. a

I

/./,.,/\/r.cr: lLi).lrcL5r:C I/.,.n,r,/rfr,,

a

/r.aj, r//l\ tS.ed,:

ne$ \-bsD;cie,l

sDe;ies: a.

Ftrtnat\quamt\ (petets):

';:,1::::;l;,:,,1i,:,i:xffi

C /r,//,"r,,, ne{

i;::ifff:::1'.Jii!,,,,i,i];,i";!,!!i

i,1;!!"1,i;::1,:;

:'"'::;::;

h:.hhxsn;

;::";*",:':,7,i';i;:i;;JJ"r:"T|:;l,l:;;i?,":t ii

;.:"1i:i:t;:"'.il.J1i'ili*1;i;il];:.1",,?fil.T,,

reJ,ion'hpsmonsrr-ese.pecie. bur{

:l,,iX"l,j,-,;ll,li-."

23

3

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6

b

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E

E

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fl

-l

3-e*p

5 : s iS:s Siiirs:isiiiS"

24

CHARACTERSUMMARY

Sizeandprcpatiol1s.ln

sizetranremaes.rr,esm^rd; ;;;;"#;#ff

n_'t4ti

./rd,?bI

ea.l_rd\on.fe t,-c.errcdn

"

tS of Lhe20 Chirah

#*iX*Xlij;j:?

"

r, ;rrr.," i

.

r;;r:i

fu 2-tt-rn-h rctdllength.Ihe rJ "ge.l,pecim

/dmr Intolallengt'.Sma.lcsland rJrge\l.lecimen.

the.pecie.arco.nr.,

e\am,nedtbr

I herrl

rsroughjJdthi-J.orsJigh.l)more.orlhetolr.,englni0a.trpecicr

3t.ln (

,l/drlr. lhemerroflheLaitasapeicenrage otrorr,

at ( h;rcntu' (frble

lengthfangesftom 31.3-40.1percent andvariesfittteletweensexi oJeactr

M"ostmtios consistingof numerousmeasuremenrsraten

taxaexceptC.carinatr

;:1ififi:,Ji#;if

on the head of ti.ii"i

weretoundto berelativelyconsistentwithin theg"nw una*"i"

taxonomicvalLre.Inall

l"n:{*;:i?,,""ma;$ffi

,

A mode dlee)e -i/e (oiarneter otcJe )/J J/4

"':::lf ,"*Ti;

\nout lenglh)uccur.In aLlr,l

tdrge eJe \o,aretcrof

. trolhe, \pecies.

nutpnrL r. t.tdt, 0rttt and (. quadntan .!tus.d1dI

e!c \. r/q ro auatrng\0o,l rength) is

oo.,e_5eo by .raultrnt

i#'J::fiTJJ::lT$Ai;i:1"ffi:'.Tl

:ljl.'ffi",lll;:i:litff

;J:ilt

lowingpreywhichmovequicHy ;nd erfaticaily.

lefltah

Ventral variation is presenred in Table 4. The lowest ventral

and the highestnumoe$ arem C: n ?rlDrrlrr

count of 123is_in C snrlet

r

,d.4/drr $\'ch Ld-a5manr a\lqb.Ccnerd

a t/ra?r7r.peLie. oJI Defor\en.trrt) | hercir

irno -cmltes In,cdi" r.\o1.

).r

. tadt,ai, C.n

he n utrircnt ri, specie,comde(

hdvcthehighe(tnum

con.iocrJble oveflapin thetdngeot.\cnt-,1i5 ro- ma,el

B,)ed or mea,,. female.rcno

ro nare.tigh.ll

moreventralsihanmaLes;however, insftofthe20ta,ramatesaverageaioi!

thanfemales.Thenumberofvenu;lsoftenvariesal"rg

g""pr"pfri. g'."Ai"'"t"

:uocdJ.,ri'\a.e-oJma J pa.rcde\ceprfor.he,ermrnrt,fine

chEfa.rer. SubcauLlalvcrj,rlion is

*tif, u-tr'rioi_

iri,i"i_

;;"

;;;;;:,

.,,llf.

l]/id^

snrcn ra5

rt.o counled d1dincrLded -n lh,(

shownin_Tabte5.Crilo,

mxmof96.Thehighestsubcaudalcount of208isi " C

iustaetricoltishasLhefe\r"rt,ut"ouaJ.

iiliitrt"

nut,1)entn,,De.ie- .o-npte\ h"i.

!uadrtarinau.,

rr-

nulrDeful \ubcdudal

olsll)clly ta\e Ihe

Agdjn. membersut .he

\4htc C.taa t,atti\.t

andr . t ulnth" in the rungeoJ

lo$e(t counl\. L\ten\rveoverlaf occrr

eachta-{on.sir"a

subcaudalsfof malesandfemalesin

lc0Jrohd\efe{er.rb(.rudat.lhir mrres.hose\er.In\ir otlhe20rd\dmates

on rn"*", i;;;1",

JverJgeoles\lhxn

geographic g:radients.

lernJle., The numberot."bcaLdals ieouenlll \dne.along

3S1S9133933?933939s3

gEsogS!3!SageggEAigS
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sie:* ri ::ri

28

lndlp/ale Theconditionofthe anelpiatein alltaxaisshownin Table6. Chironiusfusa.$,C.laeriollis,C. mo ticola,ard C.tcrrxrlrr consistentlyhave anenlireanalscale,anda dividedanalplateispresentmostfrequentlyin all orher specjes.This characteris quite variable in C arol rrr where an entte analoccurs14.2010,a dividedanal82.2%,and an incompletelydividedanal

l.60/0.

T,be 6- aoddr on o

TAXON

1 anal llare n r\

\ul/

t'o?Lr

Divided

Anal plate

lncomlletelydivided

3

150

I

I

155

180

12

78

453

20

222

7

111

33

 

28

89

 

30

I

-4

 

50

l

I

44

55

;

130

Keeledscales.Chircnixss.rrfl1lrr is the only specieswhich exhibits no keel- ing,In allotherspecieskeelingoccursonlyonthedoftalscales,alwaysoccurs onanevennumberofscalerowqalldismostobvjousontheposteriorpartof thebodybutmayextendfromiustbehindtheheadto welldownthetail.Clj- ro ius quadicafinatus is consistently the only specieswith more than two keeledscalerowsandmayhaveup to 12rowswith keeling.The numberof scalerowskeeledandthedegreeofkeelingchangeontogenetically Gee spe-

29

ciesaccounis).ThischaGctefisalsoscxuallydimoryhic,anddopendingupon the species.maleshaveeilhermorepfonouncedkeelingand/ormorescale rowskeeledthanfemates(Table7).ln tlxa wheretheparavertebralscalesare the only rowskeeled,adultmdlesareusuallysircnglykeeledandfemalos weaklykeeled.InspeciessuchasC Drtdnndrrs,C rxoletus,C.J16valineatus,C. gru disquanis, C. lduenti, C. nofiticala, C. quadricarinatus,and C. ti ce ti, whichmayhavemorethanlwo keeledsca1ercws,theparavertebmlscalefows alwayshavethe mostprominetukeelsandmalesgenerallyhavemorescale

rowskeeledthan1ema1es.

Atical litr. A,picalplIsarclairly conspicuous,conflned only to certaindor- salscalesol thebodyandtail,andlocatedatthe rcarofthesescales.Two pits pefscaleoccuronsomespecimens,butthe moslnumerousconditionisonly onepit lef scale.Wherea pit is presenton a keeledscale,thepii is usually foundin theupperportionofthe scaleabovethekeel.Severalslatesofapical pitsoccurin Crriorlrt rangingfrom completelyabsentto phson thedorsal scalesofthe neckandon theparavertebralscalesfor thelengil of lhe body andiai1.Apicalpils only on the neckscalesis the mostcommoncharacter statein mostiaxaof Crilrriur (Tab1e8).Apicalpilsareeitherabsent(52.2q0) oI prcsenlonlyon $e neckscales(47.8q0)in C 4rd dti triratus.'fhetendelLcy for pitsto be completelyabsent(95.4q0)occursin C. monticoLd.

Dotsalheadsa es.The numborandarangementof thesoscalesdo noi varyin Crtlortrr. A iypicdlcolubridcomplementofscalesarepfeseni,incLud- ing onerostral,onefrontal,anda singlepairofparietals,supraoculaJs,pfef- rontals,and inlemasals-

Supralabials.Supralabialsrangefrom 7-11in Crtlont r (Table9)-Most

specieshavea9/9condition,burS/8occu$frequenllyinsometaxa.Crriodrr

quadtininatusisIheonlyspecieswhichhasahigherincidencofeightsupra labialsratherlhan nine.

Suprclabialse teri garbirThenumberofsupralabialsenleringtheorbitis largelydependentuponrhetotalnumberof supralabialsprescnl.Thosespe

cimenswithninesupralabialsusualyhavesupralabials4-6enleringtheorbit,

whefeassnakeswith eightsupralabialshaveeithersupralabials4-5or 5 6 in Iheorbll.Chito iusquadicdrlratwistheonlyspeciesin whichsupralabials4- 6 entedngthe orbit doesnot occurat dl1(Table10).

Infralabials.The

trnber ofinftalabials rangeslrom

8-13in Crtloatrr and

variesconsiderablyintmspecificallywilh thenormusually10of 1l(Tablel1).

G rtdh.Twopairsofgenialsarccharactedsticof allspecieswiththeonly excepdonbeing rhree pairs in lwo specimens01 C exoletus('lable 12).

l0 I l tl I E E 2 3 I E E ril F3 ,:
l0
I
l
tl
I
E
E 2
3
I
E
E
ril
F3
,:
I
-l-l
.n
Q
:9:r
R;
-:
:';:i
"R
-ej
s
r
t9
92
|
|
|
o
o
o
-s
- €
3
-e
:,8
!e
!:i
E:i,
is:e-!F
3
!r{F
*!:iii

3l

n

f

-r

-

|

:o

n;

|

|

|

|

23P

z&i

tttttl

*::iF

.:i;:!ii

iii!t!i!i:

d SEEEES

|

|

|

|

|

|

^o

|

-:i.!

30

o;

|

9a

-:

12

&zJ

:

-sE

z'zz

a

i

=

p

hE

4zn

E;!a

6

3

6

td3

_d

I

lx

t-

Trbte 9

Frequcncyofslpralabi.lsnl llxa ol Crrorirr

TAXON

 

25

11.1

1

26

l

 

)2

2

6

ll

I

 

2

1

1

1

6

58

ll

2i3

126

261

l:15

lt5

:r69

176

52

145

i4

90

'14

2

41

2l

',

2

I

2

5

1

)

5

1

178 1l

t1

I

Inlidlabialsontadiry gerl.rr. The rungeofthis charactefin Clilontrr is lburloeightorcachsideofthechin.Themostftequentconditionsinalltaxa aresft or seveninffalabialscontaclingthe genialson one side(TableU).

Prco.!/drr.Thercisnormall,\7asingleprcocularin Crlmrt?A,buttwopreo- cularsoccufin somespeoimensin 1l ofthc 20ta-\a(Table13).

Pdslo.,r/d. Oncto fourposrocularsarepfescntin Cr,rorirswithhYo tre- dominatingin mosttaxa(Tablel3).Membefofthe mrlt0ertlr spcciescom- plexandC. lla"olineatushave afelativelyhighincidence(above$qo)ofthrcc Postoculars.

Tempordls.Thetempot^lconditionisquilevariablewithinthegenusdue io unusualfusionsor splilsofthe basicscalesinvolved.A totalof 30dill efent

l4

':

.!

5

.9

;

'E

a

g

3

I

I

3\

I

E 3

--S

al

e

!t5t+

r^-!o

I

-11

tttl

I

r

f

I

i ruir5 i:ri

l

ble ,l

-

l

TAXON

eq.elo

| ' doral' '

2

I

l

3

3

l

rv

of Cl fur

"

INFRALAII

\LS

 

t0

1l

2),

i33

75

I

91

81

t62

109

50

21

219

350

1

32

25

257

80

8

22

2

4

l

50

33

2A

22

IJ

81

42

2

22

12

 

E

19

 

52

l0

3

35

10

 

3

5

20

54

23

l

38

 

l

2

12

21

12

l3

I

123 6

9

I

I

5

2

I

2

5l

l

states,1+1,1+2,1+1+1,and1+

cbaracterstatest{'ereobserved:however,four

1+2,occurmostfrequenLlylnChircnius (Tablel4).The condition1+1wirha

diagonalsplit

prolably gavedseto 1+1+I (f-_ r--l

eaveriseto t+ t+Z from 1+2 {

iowsaimidbody generallyhave1+2temporals.Exceltionsto ihisareC

/indrrrwith 1+1

'icd-

acrcsstho anlerior upper

l------1

quarterof the primary temporal scale

andthe samo b'pe offission Those specieswith 12scale

_ t'_ f--

r ),

- v---F=\

(46.90/0),I+2 (43.90/0),ot 1+1+2(730/0)temporals;C eirDl ftrr

withl+1(7.1q0)of 1+2(89.8oh);C

coldwith1+1(6.5V0),l+2 (61.50/0),or1+1+2(1780/0).Thestate1+listhe

anlconditionin threeolthe fourspecjeswilh basicaltyl0

C. laevicotlis, ^t\d C.saltrulrr)

body.C. cn

flawlineatus\\ith

1+1(8360/0);andC no'11t

domin-

scalerows(C.tr'at

The fourthspecieswilh l0 scalerowsatmid_

disqudnk,has1+1(126q0),1+2(17.80/0),t+1+1 (13.60/0),or 1+1+2

(45.8q0)tempo|als.

36

'E

3

E

E

6o

I

{

j

I

I

6

trt

toae

-llll-

ll

ttl

I

9 ii r*eci ::

l7

;lill

16-

 

|

|

 

6

_

 

|

Ll

-

|

t6

 

3

F

a

 

d

n

E

6

a

.=

 

3

-

a

5

d

I

l-

6

d

|

: €€reies t::i

l8

Tablc14 - Frequencyof temloral scllesin tdu of Crim,?,r.

TAXON

l+i

r21

5

8

'11

r01

29

21

52

3

1l

I

5

5

231

t+2

119

233

291

898

,]

38

2

5l

I1

90

'11

91

6

TEMPOTATS

1+1+1

l

5

2

29

2

I

1

1+1+2

20

3

6

7

98

3

'j

a

12

1

22

l

23

\2

I

22

;

2

5

2

lol dk. The number olloreals in Cttlo,?i,lJmngesfrom none to two with one lorealgreallypredominatingh a[ ta"\a(Table15).

Pastcephalics.This chamctershowst.emendousvariationwithin each

taxonand rangesftom 4'11in t.hegenus(Tab1e15).Thereappearsto be atrend toward tewerpostcephalicsin specieswith fewerscalerc\\s (C.Ji.tscus,C.laert

cc'l|is,