Revisionof the Neotropical

SnakeGenus
Chironiws
Fitzinger
(Serpentes,
Colubridae)
JamesR. Dixon+,John A. IfiestJr.* andJosM. Cei**

'.

iJ:g;;1gg1"t
3i:;91;5;*

Fnheries
sciences.
rexasA&Muli,enbr, coresestation,
-.
*turares.unjveBidad
Naciomr
deRiocuano,
(5800)
Riocuarto.

TABLE OF CONTENTS
Intfoduction
Crtorira Adve isemnrlnfo.mation
Mrreridh rnd Methods
Sinopsisolrhe cenus Ci ianius Fitzjn}e\ 1g26
I{istoricalreriew
Family status
Chafactersummarv
Identfr catian af Chin nius
Key to the Species
Systematicaccounas
(-hitnaid,biarinat , t\t\iedl
Chito ius cai aits (Linnaeus)
Chircnius einatus cainat .' (LinnaeusJ
Chitonius t.tinatusf
aprlrr(Wemerl
Chttontu\, ortnart)s\D,y tHltove]l
Chtrunttt.?'ul?t \ tl ionacusl
Chircniusfl.trolineatus(Jan)
Chi,04td\
fu, " || Innreu(r
Chttun,a,1yuu,,rr*" fLrnnaeu.r.
Chtoniu,tu..ut tcutontt tr. neq .ub.pecres
Chircn iusgt andisquamis (p eterc)
Chirc tus taetico b (Wied)
Chitotiius laurenti, new species
Chitoni s ma ticotaRoze.
I l.t,untu,nuhNettth,Schmidlrnd Wailef
.
Chironius nultil,enttis cochrcnael':rogeand Romano
Ch; t on i d \ n Ltlt tventt t. tnvpaI d, Baite,

Chitu ius multit,ent s multterfir Schmidrand

Walker
Chilouiusmulti,entis seplrrrbnar, new subspecies
Ch;Iont \ qua(tti.annaru\|Boiet

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,

new
spectes
i'" n u' 4,tadi, ari40ttl, quala.r n t,
a, I Boie/
th
u' rLun utu\ (Wasten
Itnt.mnt
ntrnniutrtncenti(Boutenger)

pag.
)
'i
'i

Interspecifi
c relationships
Formationof phJ,logenetic
Tree
aenu/ot(H,.tof) or ( entrutdno
South
Zoogeographic
Considerations
Erol|lIion of Chirc lus
Intergenedcrelationships
Summaryol taxonomicchanges
.
Lite.aturecited
Appendi\ I, specimenskno.$n,not
examined
AppendixIl, specimensexaoined

181
184
188
195

r
,
,
,

200
213
211
221

233

237

231

219

250

INTRODUCTION

zoogeogmphic
It hasrecentlybecomoquite apparentthatcompfehensive
studiesofNeotroplcal feptilescannotbe lerformed efiectivelyuntil many
lhorough,basic,ta-xonomicinvestigations
havebeen completed.This fevision concernsthe genusCrl,"orirqa groupoi snakeswhich havebeenlradi
tionally,ta-{onomically
abused.OfallNeotropicalsnakes!
only Crr'r'orirrhave
10 or 12 scalerows at midbody.Their uniquenessand relativeabundance
makethis genusan interestingand expedientsubjectfor study.
in tryinglo
Ourintereslin Ci,Torrrrdevelopedwh,3nwe becameinvolved
idenrify a number of Peruviansnakes.Confusionin tle interpretationof
folms in the genusCr;/oflirr had beendiscussed
by Bailey(1955)and Wiest
(1978).After usingthe keysofDonoso-Barros(1969)and Petersand OrejasMiranda(1970).
v/esoonrealizedthat existingliteralurewasnotadequatefof
identifyingthesesnakes.Much perplexityalso existedin the allocationof
namesin museumholdingsasseenbythe wideuseof Chiro iuscainatus'dn(l
Chinnius fuscus,bolh Linnean speciesnames,with comparatjvelylittle mention ofthe othorspecies.
Sulprisingly,somemajorlending institutionsof the
world peryeiuatedthe useofsynonymssuchasIIeryetodnras
Cchiro ius)ol
used namesthat do not applyto Crirorirr suchasr efilinatus Oydglet,1824),
which belongsin the genusPrrrrer(Bailey,1955;
Hoge,1960a).
lt wasobviousearlyin thisstudythatwhllotho genusitselfwasuniquedndwell defined,
A fact that fuithei
the specieslimits were quite vagueand misunderstood.
point is that 49.34k
emphasizesthis
oI all Chinniusspecimens
xaminedin the
cou6e ofthe studywereeithermisidentifiodto species
or unidentifiedby the
lending institutions,but only 0.40/0
of the specimensloane.l as Chironius
actuallybelongedto other genera.
on lhe genericrelalionshipsol
Therehavebeenno major presentations
relationships
havebeendiscussed
by
Crilorlrr. Although someinterspecific

10
BaileJ(1955)and Donoso-Barros
(1969),thereare viftually no major works
keatingthe systematics
ofthe entiregenus.Further,the generalrangeofile
genusis known,but the distribulionsofthe specieshave
beenpoorlyundcrstood and noi well deiined.Most ofour loowledgeol stecies,djstriburions
comesli-om regionalstudies,whichareoccasionally
incompleteor unfeliable
asto species
idoniifications.
Theseincludelhe worksofDa CunhaandNascimenro (1978,1983,1984)for eastemAmazonia;Moonen and van DeuNen
(1979)fbf Surinam:Chilpaux (1986)for Frenchcuiana; porez-sanros
and
Moreno(1988)forColombia;Via a era1_
(1988)
for CentGlAmerica;Carrillo
de Estinosa(1989)for Peru.The lackolecologicai,behavioral,biochemical,
and orher specialized
studieshasalsohelpedto impedelaxonomicstability
rvithinthe genus.Speciesof Criforlrr havebeenmentionedin a numberoi
diverseanatomicalrolatedstudies,and for agroup ofsnakesto be so widely
used,ii.is imponantthatiheir systemalics
be trealedto removeexistingcon
fusion in the allocarionof names.
C1l//IONIUS ADVERTISEMENT INFORMATION
The genusCrilorirr canbe definedasamediumto 1aryesized,te[estdal_
arboreal,agl]lhous colubrid.The hemipenishas a singlesulcusand lobe,
nakedpocketproximally,spinouscontrally,calyculatedisralry,andis withoui
capitationor apicaldisks.The lupil is round.Thereis a lull comllenent of
colubridheadscales.
Thereareaboui123ro 196veniralsand96b 2b8subcau_
dals.The midbody domalscalesare in 12 of l0 rows,with lhe presenceor
absenceof keelingand apicalpitson dorsalscales,dependingupon ihe spe_
Crilorirr is diagnosedasthe only Neotropicalsnakegenuswith l0 or 12
scalerowsatmidbody.This characteraloneseparates
C,itori6from ellother
Centraland SouthAmericansnakegenera.
(h,ro,ttu,hd. d \eor-npicaldi.rib. ,ion.e\tendingtiom
lhe -o thcrn
coaslol HondJ-ri In urugui) drd nolhed.t AJgenltna,
Mu.t spe.ie..re
lbund east to the Andes in ot around the Amazon Basin.Chirotli s has
reachedthe Islandofcorgona alongthe PacificcoastofColombia, several
islandsalongihe Atlanticcoastof Brazil,andTrinjdadandSt.Vincentlsland
in the CadbbeanSea.
Habitatprefercnce
vadesextensively
within the genusandwithincertain
species.Lo\rland tropicalrainforestis preferrcdby Crtr,orrrr,bur humid
montaneforests
andsavan
nalopenlormationsarcalsoinhabited.Theelevation nnge ol the genusis from sealevelto 2800m. The rangeand habitatoI
exchtaxon are expldnedin greaterdetailin the speciesaccounts.
A generaldescriptionofthe total sum olvariationin the genusis: Total
lengthvariesfrom 274mm to 2718mm and the tail is 26.9-43.tpercentofthe

I
ll

total lcngth.The ventralsvary from 12l-196in malesand 131-190

in females.
The subcaudalsafe paircd and vary fiom 96-208nl males.and 99-205in
femalcs.The umbilicalscaf is normallythreeventmlslong. The analplate
maybe dividedof entirc.The amountol keelingisquiievadablc,andpreseni
in all species
axseptC,rrrr?/ri, Malesgenerally
havemofe pronounccdkeel,
ing andir somospecies
\Vithnofe scxleforvskeeledthanlemales.Apicalpits
areabsenlin C ,?ortrold,presentor absentin C 4uotiicainatus,arTd,presenl
in all other species.
The nofmal comple cnr ofcolubrid do$al headscalesare:supralabials
nofmally8 or 9;supralabials
entefingorbit usually,l+5+6,4+5.or 5+6;infralabidsnormally10of 1l trvopain of genials;infralabialscontactinggcnielson
one sideusually6 or 7; usualh I preocular;prcoculatusu.lly nol in co tact
wilh iiontal; usually2, somelimes3 postoculars;
uppefpostocularusuallyin
contactwith parietals;
tempofalsvadable,usually1+1or1+2.sometimcsI+l+
I or 1+l+2; usuallyI loreal;usually410 postcephalics.
Thereare24-51aglyphousnaxillarytccth,and26-46dentafyteeth.Thereafel2 or l0 scalerowsat
nidbody, withscdlerow feductionquitevadable,usually12-12-10,l2-12-8,10
10-10,
of l0l0-8.
Hemipenesof ell speciesafe singlelobcd.cylinddcal.and dillefeniiared.
The retfacledlengthofthehemipenisrcachestosubcaudals
5-13;,41.
//r?./r/
/errs ,ragrr.i originatesbetwccnsubcaudals15-32.The sulcusDermaticus
simple(not divided)and usuallyterninatessubapically.
A basalnated pocker
is presentolposite the sulcus.The basalfofiion of lobe may be with ol
without minute spinules.The cntire centralpoflion oflobe containsspines
$hich number4 12ler longltudinalrow.The cntircdistalportionoflobehas
calyceswhich number8-26pcr longitudinalrow-The cdlyceshavefroximal
spinousfringesrhar changeto papiUatefringcs distally.
Inlraspecifi
sally,coloraiionandcoiorpatlernsmayor meynot be variable,
Fofestlorms iend to be dafk, usuallygrcen.olive,or blackdorsally;open
ibrmalion lbrms are uniform yellow brown, olive brown, of havc a stdped
paite .
Speciesof Cir"rtrr are oviparcuq ierrestfial-afbofeal,
mild to vety
aggrcssive
in disposirion,
andgenefallyslcepabovegroundat nighl in treesof
bushes.Thesesnakesareactiveforegersduringdaylightandpfeyalmo$ exc
lusivelyon fiogs.Crn'orlrr, herein,is comprisedof 13species
anda totalof 20
tAxa. Chironiusbia:trhatus (Wied); C. cainatus utiratrr (Linnaeus); C ../1d,
rolrrds (Wemet; C C r"/it(Hal1owe11);C a?1t s (Linnteus); C.Ia\iali neatus (Jan); C. fuscusJuscus(Linnae]ls), C. f. leuu)nelaprr, new subspecies;C
gtafidisqudmis
(PeIefi);C-Laericallis
lNied); C.laurcftarew spccies;Cno,
ticolaRoze;C. nultiwntris rir.irdndr Hogea]1dRomano;C- n.Jovedtustsaileya(- m. mull entis SchmidlandWalker;C m- sQlert"loralir,now subspecies; 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

f
t2
multivnttis, anrl C. vincenti share many uniquely derived chatactef states,
thesespecies
areplacedh the r,r/rn,entnrsleciescomplex,a convenientway
oI showingthe closerelationshipamong thcse speciesbut which has no
nomenclaturalstatus,
In 15of the 20 CriloTrtrrllrxamalesattaina largefsizethan females.The
smallestspecimenexaminedwasajuvenilelnaleC manticola2j4mminLotal
lengrh.The largestspecimenwasa male C grdrdtrquamis2718mm jn total
length.Smallestandlafgestspecimens
examinedfoteach
taxonarcpresented
in the sleciesaccoln|s.

MATERIALS AND METHODS

Ofrhe 2978sDecimens
ol Crilo'irr usedin theseanalyses,
lessthan 400
\yerenotpersonallyex,rmined
by us,Most specimens
wereprocuredon loans
fron museumsand educationalinstituiionsin the United States,Europe,
SouthAmerica,and Cenl|al America,and thesecollcctionsafe listedal the
end ofthis section.A list of lnown bur unavailableCrjlorirr specimens
is
providedin Appcndi\I andslecimensexaminedarclisted(_by
spccies,locality, andmuseumnumbor)inAlpendixll. We examinedor acquiredinforma
tion on 19ofthe28i),pespecimens
olell species
andsubspeciesnamesthatare
associated
with Cir)oaiff. The feDainingninetypescouldeithernot beloca!
ed aftere{ensive searchiigof weropositivelydeierminedto be lost or desscalecounts,aspectsof anatomy,measurements,
ralios,and color pa!ternsofpresefledspecimens
wereutilizedin analyzinginterspecificrelarionshipsandinimspecificvariation.Duelo timefacto|Sassociated
with thisstudy
of to thc poor conditionofcerlain snakes,completedatawerenol obtaincd
fiom all specimens.
Ventralswercountedfollowingthe meihod ofDovling (1951a).
Single
half-ventralswefe ignoredsincethey indicaledvertebralaberaiions (ting,
1959),
but two hat scalesofa dividedvenrfalwefecountedesone.Subcaudals
arepated and\refecountedon ihe left siclesrarlingwith thefirsrscalebehind
rhe analplatc that contacteda scaleon the oppositcside.Fusedsubcaudal
lairs were includedin all counts,as was the unfaired, telrninalscale.
Conditionofthe dorsalheadscalesisconsisteniforallspecies
andwasnot
rccordedtbf mostspecimens.
The llumber ofsupralabials,
supralabials
enterlng the orbit, inffalabials(lastinfralabialdetemined as the lastscaleat least
half coveredby the last supralabial),infralabialscontactinglhe genials,
genials,
pfeoculars,
!rcocularscontactingtheliontal,postocula$,postoculan
contactinglhe parietals,lenporals (scalesbehindlostocularsand berween
the paietalsand supralabials),
loreals,andlostcephalics(scalestouchingthe
postedormargln oftemporalspa elals)were noted.

A slashmark is used to separatecharactercounts taken on oppositesides

ofthe headwith the left sideindilrated
first(e.g.,supralabials
9/8 signifiesnine
supralabialson the lell side and eight on the right). A hyphen hdicates range
in a character(e.9.,venlrals135-155
means135to 155ventrals).Segmenisofa
chalacterananged in a seriesarc separatedby a plus sign (e,9.,temporals1+2
denotesone primary and two secondarytempomis).Teeth on the left maxilla
wereexaminedand countedi, rirr usingthe methodof
ofmany specimens
Myers (1974).The number of dentary teeth (left side) were noted for a lew
specimensof most species.Condition ofthe analplate (singie,divided,or
incompletelydivided),positionofapicalpits,anddegreeofkeelingwerealso
recorded,
Wlen no evertedhemipeniswaspresent,the sexofeach spesimenwas
determined by maling a post-analslit between the anteiior subcaudalpai$
andcheckingfor the presence
ofrr rr, hemipenes.
Hemipenialmorphology,
includinglength(bynumberof subcaudals
involved),shape,conditionof sulcusspennaticus,
and omamentationis describedaccordingto the lermino(1960).1,r,t/ hemipenesofat leastthreespecimlogyolDowlhg and Savage
ensandseveralevertedhemipenes
wereusedin the descriplionfbr eachspecies.Eggspresentin oviductswere countcdby palpationor by incisingthe
midlineoftheposteriorventralsandvisuallyinspectingthem.Witl thislatler
method,the greatestlength and width of the eggswere measured.Snakes
with obviouslyenlargedbodiesin the stomachrcgionhadtheirstomachcontents removedand examined.Noticeableecto-and endoparasites
werecollected and identified.
For consistencyabbreviatedscaleforv formulaswefe stardardizedby
counting the nnmber of dorsal scalerows over the tenth ventral behind the
head.overthe middleventral.and overthetenth ventralanleriorto the anal
p1ate.Completescalerow formulasindicatingthe exactpositionofscalerow
reductions ard additions were constructed for most specimensusing the
melhod (with slight modiflcationt suggestedby Dowling (1951b),and these
(1976).
formulaswercsummarized
followingthemodelpresentedbyThomas
(ir
Ali measuements
millimetres) were taken on the left side where
applicable.Snoutvent and tail lengthswere takenfrom specimens
straightened againsta meter stick. Other measurementstaken with a vemief calipers
weresnoutlength(straightline disiancefrom tip ofsnoutto posteriorendof
interprefrontal suture),frontal scalelength, headlength (shaight line distance
from tip of snoutto posteriormarginofjaw), headwidth betweenthe lateml
marginofthe supraoculars,
headwidth betweenthe e),1emalnares,
rostral
scaleheiglt andwidth,andhorizontaleyediameter.Specimens
in the 300-500
mm snout-vent sizeclassof most speciesof Crilo,'?lrsgenerallyshow ontogenetic changesin color patterns and degree of keeling. Therefore, the term
juvenile is ruther arbitrarily defined herein asspecimensbeiow 400mm snoutvent lensth.

\l"-1..hne i.,: rot,.i.r, (.e...c..r,t.r <,t,.,d(1 .J.,.,,no..I

. e.r..lr'
J<,1.S..r. \.o r\r (.....,heri.r]...:..gcL rro n..t.o.
',1r"'. "] . : m' tD. .t.-c,r'<n.,".o ..i. .i.. il-,i ,n.ce...it.o1.
\iI .rrron.I r. ht\lL
1. . nt tcrrJ ls
The51rorr/rrdi!isionoferchtaxongeneray jnctudes
onlyjuniors!no_
nymsand the dillcren(gcnericcontbinxtionsthal hrve bcen uscdlor ticse
names.The fi^t felifcnce to cilchIel1)ein ihc slnonym).is gi!en,as .epel
I"._r.\ ir. ....r' e.l,rIn.h. \i.-.
-.me,ot ....\;,..nt.Ito1r.rr:1,1rIc/.,..\tr.
..n.(,.tL..
t,e^- jtj(.ntllr(.. . n.r|. r.tr,.n:
the cu.lliDg museum_specjnlennunlbef.condition.sex.rypc localit). an;
collectof.The 1ltuArr.rA.
indiclr esthe e.rsicslwlrs ihat orc laxoncin be dls
lirguishedfionlullothefs.t--achdoronurangc1nat
ir lhe Di!1r?rllldlsecti;I
. te.. l. ,,c.1,\.rr, I
ll-..r on-.t,.c .-,.,.r..i .el..n, ...pe,l
,.,r.' 'lr:je n, :. .rre J b . . , ,rr . l i . nr.r lh .r cderI.o r .r
o !u\\h
rnd Seibeft(1972)lvasl.rrgclyusedio descdbethe hrbiiar or \;gclalional
Tonesocclr|led bl each specics.Ahjludinal lirit$ ofa stecjcsrre based
strlcllyufon elevalionsfccordedrvilh thc specimcndata.
Thr ,cr./l/rid, prcscnlsthc numbefofspecirncns
exaninedfor erchspe
cresor slrDsDectes
andI sumnafizationol thc characicrslof cachtxxon.IncLudedir lhis scctionisinlonnalionon mlnimunrandmaxi]nunrsize,fel|rile
iail lngth.scalccounh and felatoddara.and rhe number ol. matdiaryarld
denral)teeth.Ontogenerjc
changes
in colof.ind/of pltrem and scruajit,
d, _' t'.r . h r...r( . rc.ri,c....t {h. r. n.r ,\1.r-. \ .o.r |n rrh.,1.
r'r. r l,mrt-,., .1e-ro.iJr { . / , . ||J . u u " n I Ie . r .. e g \ . || ..retrlrscnlativctllrlnbcr ol tfcser.vedspecirnens
ofe.rchspeciei.l; pxtlcrned
snrles Ihc numbcrandfolilion ofbandsarcnoted.and lhe lengthind width
ol rhcse ma.kingsare exprcssdir iefns of ihe number of scalerorvs
rn!olved.
Color dcscrittionsol li\,eCrn.rrlr/rarebasedon lteraturc accountsand
rnlirrrnlnionmrde avlritrbleffon thoscwho havehaclfield exfcrjence$,jth
thcsesnr (es.Iliuslr.alions
rnd fhotogfafhs fircijiraicdlhe doclrmenration
of
Ircserredandlivc specimcns.
L la, a .." ri .^n.
m.r.(..,r. ieodr.tLr..r troru.\.....ti..1
_r'rn..,',,| In.r ,t.ci,i..., i.,u
i.. J.et\ J..lemr)c. bj u.\: .i...\.
.- I
crh,,C\..i _th<\t.tr.i.:t \n.h.,\\,cnt15\\r.r,
\mJdh
.170V/6conrurcr.
Asignificance
lcletot 0.0tras Lrscdtbra|
r tes|S.
Thissec_
liof ls info nnt jn bringinglo ljghr possiblc
e!olurionar.ily
distillcrunjts
wrthina specics.
/-/? H/r1r,fnotcsincludea\,rjhble inlirfmftion on beh.rvio
r, iime of aclivily. se$onrl occurferce.relfoduction, cggs,fecdinghabils.incLecto_ancl
endopafasltes.
thc nerrd?lJsectionprovidsa plrre fbr presenting
othcr inte_
resrng.perlrncnt,or rclated$orks nor citeLlelsc$hefein ihis siud),.Com
mentsaboulthc et!.molo$oflhc species
numesa.e usualllinclucled.

Il is udbnunaie, bul nevertheless

a laci thatvefy few majof revisionsarc
ableto drawulon all availablecharaclers,
melhods,and tcchniquesfbr ana
lyzingdalaand lbrmulatingconclusionsHopefully.behavioral,anatomical,
chrcmosomal,biochemical,and othcr studics$'ill somcday bc uscdlo iesi
and amllily ouf resultsand interyretations.
Cited institurionswith thoir co cspondingcuraloN afe abbrcviatcdas
ibllows:
AMNHrAmcrican Museumot NaiuralHistory,New YorklDr. ClnrlesJ.
Cole,Mr. GeoryeW.Foley,Df. ChaflesW. Myem,Dr. RichardG. Zweifel).
ANSP:Acadcmyof NaruralSciences
ofPhiladelphia(Mr. EdmondV. Malnale, Dr. ThomxsUzzell).ASU: Afizona SiateU.iveNity, Temte (Df. M.
JackFouquetle).AU: Auburn University(Dr. Roberi FL MoLrnt).BMNH:
BritishMuseu (NaturalHistory),London (Mr. Andrcw F'.Stimson).CAS:
Calilbrnia Academy of Sciences,San Francisco(Df. Alan E- Levito ).
CENAI: Centro Nacionalde Investigaciones
Iologicas,BuenosAircs (Dr.
Avelino Baffio, Dr. Marta E. Miranda).CM: CarncgicMuscum,Pittsburgh
(Dr. ClarenceJ. Mccol,). COH:C.O.HandleyVenezueldn
Project,at USNM,
Washinglon,D. C..CRE:CosraRicaExpedition,
Univclsityolsouthem Cali
Iornia,Los Anseles(Dr. Cafl S. Lieb, Dr. JayM. Savase).
FML: Fundacion
Miguel Lillo, UniversidadNacionalde Tucuman(Dr. RaymondF. Laureno.
FMNH: Ficld Muscum of Natural History, Chicago(Mr. H,\7men
Marx).
GIUS: Geogmphisches
Institut der Unive|sit;ildesSaaflandes,
Saafbiicken
(Dr. PaulMiiller. Dr. HaraldSchreiber).IRSNB
Institu! RoyaidesScienses
Naturellcsdc tsclgiquc,Bruxelles(Dr. Xavier Misonno, Dr. CasronR. de
Wnte).JAP-GOV:J.A.Pelers- G. OrcesV.EcuadorianColleclioqatUSNM,
Washington,D. C.. KU: Universityof Kansas,Museun of NaturalHistory,
Lawrence(Mf. Joserh T. Collins,Df. Willialn E. Duellnan). LACM: Los
Angeles Counry 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: Museode BiologiaUniversi(Dr.J.Raconis).
dadCnlraldeVenezuola,
Caracas
MCN: MuseodeCiencjas
Naturales.Carecas(Dr. A. R. Lancini V.). MCZ: Mlrseumot Comparadve
Zoology,HarvardUniversity,Cambddge(Ms.PatriciaG.Haneline,Mr. Josd
P. Rosado,Df. ErnestE. Williams).MHNGj Museumd'Histo'rcNatufelle.
Cenave(Dr. Volker Mal}nen).MLINLS:Museode HistodaNaturalLa Salle,
Caracas(Df. C. A. de Lima).MILS: Museodel Instiluio de La Salle.Bogod
(Dr. HermanoNiceloroMaria).MN: MuseuNacional.Universidade
Fedelal
do Rio dsJanedo(Dr. AntenorLeilaode Carvalho).MNHNM:Museo Nacional de Histofia Naiuml, Univenidad de La Republice,Monievideo(Dr.
FodericoAchaval).MNHNPTMuscum Nationald'HistoifeNaturelle,Pads
(Df. R. Roux-Estve.
Df. JeanGuib6). MPM: MihvaukeePublic Museum

aDf.M.\ A. \ic{ersonr._\4
RSN.[4u\eoregiond]e
di 5cientenarurali,I of no
rDr. LtendUa\e :. Dr. franco Andreoner.
VS\U: \4JseoCi\icoo Srofia
ildlurJle UracomoDoria.CenovaIDr.
LitiaCapoccjlr.r.
V\,/: l\,1useurn
of
ve'lebrule/oolog).Uni\ erisr\ofCdlilont.r.
Be"kelev
r\4r.SreDhen
B Rulh,
ur..Krchlrdu.sage.-DrDdvidB. Wrle,. M/l 5p: \4r.eo de,/oologt,r
aa
unt!e^rdadede Sio PaulofDr. pcutoL. VanloLini)\4/Ul : Museodi7oo_
loga 5'slemntica
deUUri\ersilaoi [orno. lorno (col,ec.ion
no\aloanedro
lne vu:eo.Regjonale
di Sf,e[zen,rlurJli.lorinor rDr. |end Gdvelti,Dr.
frdnco {ndreone)M\8. \bturbisrori\ches
\4Lse-m,Ba.etLDf.[ . Raim.
Dr. OlivioRieppel).NMW:NaturhistorischesMuseu-,Wi"nb..
demann).NRM: NaiurhistoriskaRiksmuseet,Stockholm
^".ii"l
Edels_
Gjr. Carl
lam).RM\H Rij".mJseumrrn\aruu-tirkeHisro.ie.f.iae.LOr.Vnuri.
Hoogmoed).
SDNHV: SanD,ego\arururHi,loD Murum lDr. Chdrtes
R.
Crur ) r.S\,1\5:Srtuliches\4u\eumfur \aturkunoe.SluIgan(Dr. Heinz
Wermutl)._TCWC:TexasCooperative
Wildlife Col""tlon, f i'"u, eaU Un;_
\ersirJ..Codege
SlationrV. Jo BoJCt.
Vr. l\4ile \4ccoid/ f\Ha: i;;s
\alJml HisloryCollec.ron.
Uni\ers y of le\a\. AusrinrMJ. CeorgeD
tt.rumgardrer.
Dr. Robe"r[. MJnirr. tU: futJnet n,rer.iil,\en O.jean.
I Dr. tiarordA. Dundee).
UCT(B: t ni\ersiddddeConcepcion,lrstilJtoCen_
.ru1de Biologia
rVr tho"nd( ekdlo\ict. ). U(\4. I niver.irJofC.tlao
Museum.Biuldef(Dr. I. PaulVastinr.UC\4NH:Unr\et5rlJ
olConnecthur
Mu\eum-oiNdlur.rlHi)lory.Sto.rstDr.
Robent. Dubo..r.
UCR:I ni\ers,dad
0e( oslaR ca.VLseodeZoologid.
-RodrigoI dcio..(Dr.
CiudadUni\ersrlafia
Dougla\ C Robin\onr UFiF5V: I nirerisrtof florio4 i.onda Srare
Mlrlgum,
_cainesvi e (Dr. Watter Auffenberg, Mr. f"t,j, e. U"yf*).
UIMNHT Univerjstyof Iltinois Museum of Nttuml History, Urba;;
i;;.
D,ona,df .
Nt<.Doroth) M. Smirhr.UMVT: t ;,i;;.ly;i
_HoFmeiner
vr:brganMuseurnol,/oologJ.
AnnArborI Dr.Arno a C. Kluge,Dr. Ro;ajd
C. \ussbrum).USNN'l:\ational Museumot N,rlLralHisloO.$ashington,
D.C r\4r. RonaldI. Cromoie.Dr. w. RondtdHeler,t\4s.Fr,rnce,
L M;Cut_
iougn.uf.(jeorgeR.Zug)
UIA: I niversil)o,Te\.r(d Ajlinglon(o eclion
ol ve-lelrrdles.
Arlinglonl\4f. JohnL. Drfting.Dr. Willi.jml-. p\bum).I I :
Unne|5rJ,ofuldh.lMu.eum
oJloorog).sdl I akeC t) { Dr.Joh; \1. I egle1.
U/vK: Un \ersiletel\Zoologis"eML,seum.
Kbenhavn(Dr. F. l\ Biaer_
trupl.ZN44:Zodlog(bM-\eum. AmslercdmrDr.
A. Ame,ing,
D-.D Hit.e_
nius). ZMB: Zoologisches
Museum and der Humboldi_Uni;rsitiit, Berlio
fDr. Cunther Peter-).ZII\,lH: Zoologisches
In\lilul und ,zoolog'<che.
!ruseum. Unrref)rtdtHamburg(Dr. Hans_Withelrn
Koeocle,Dr. W.
Ladeesj.,/Mt | :Zoologisda
Vu.ee . I ppsala
Un.rersirer
rDr.I arsWalLint.
/MZ_ ,/oologisches
\4rseum. UniveAit;t lurich rDr. HansjitrgHol,/./
ZS\,IH: Zoologi\che
SlaarssammtJng.
Herpelotogi.che
Abreitrng,Vitn.hen
{DJ.Ulrilh Cruberj.ZVt : ZoologiaVeneorados ( iercid,R .riVNHNM,
Montevideo

l7

SYNOPSISOF THE GENUS Cll1RON1US

FITZ]NGER. i826
Geneic SrnanJ.n!
Crn",r"s F zinser.1826:31.
TrDe{peciesColal.r.drr,ar

Linna.usby odgnal designation.

'lrFe-s!.cicsnot desieraled.
Et?.1.).t)rasBaie.1826:231
]1?rr,.r,drydr
WusGr.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{nccics ard?,rerladrollrr 1\'ied b} 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.

HISTORICAL REVIEW
Snakesof drc colubdd genusClr'rorirr werc odginallyassignedio ftc
genusCotur?/'byLinnaeus(1758),
who namedlbut specie..\
CalubetcatinaLu.t,
C. etuletus, C.fuscus, Lnd C. satumtuus.Nl ol rheseate tecogrlized as ralid
speciesof Clrtirrlrr exceprsatuninus,.rthichhasbeen placedin ihe synonlm-v ot.r6crs herein.Coluberun atus otLinnJeus(1758,U66)wascalled
"ChifonsNatler"by Meffem (1790).
A shorttime laterDonndodT(1798).
\i\,ho
referenced
Merrem's(1790)
worf,,probablyuiilizedMerrem'scommonname
and formulatedColubetchito ius. Donndorfls genus-species
combination
appearedin the litefaturconly onceand Crlrrpr.rdrrdtrr wasagainusedby
Menem (1820)andLichrenstcin(1823).
Fitzinger(1826)stecificallydesignated Linnacus'Colrrel cdtrdlu.tasthe lypc-species
ofhisnew genusClironirr.
Bt' 1827overhallthe presentlyrecognizedspecicsofCrlldrnrs hddbeen
naned bylinnaeus (1758),wicd (1820,1824),
wdgler (1824),
andBoie(1827)
and vere placedin fte generaCrlrrl (Linnaeus,1758;Wied, 1820);Nar?-r
(WdEler,1824),Chhanius (Filzinger, 1826),and t? etakras (.Boic,1826,182'7
).
Latef, spccies,oflhis genuswefe refcfiedto asllcrprrdryds(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).
T he genosH erpetodryasvas widelyusedby manyearlyherpetologisls
and
natumlisls.This namewasprcvalenlintheworksol Schlegel(1837),
Lichtens,
tein (1856),
Gunther(1861),
Jan(1863),
andJanandSordelli(1869).
Cope(1860,
1868,1869,1876,1886a,
I886b,1887e,
1887b,
1893)usedl/e?erodryarin hisNeotropical herpclofaunasludies,greatlyjnfluencingthc use of this generic
name.Boulengef(l894a)causedamisunderstandin
g of Hetpetad
rr as(:C hno

l8

ti,rr, by placing a number ofnames in s).nonymyand delimiting the genusto

five species.Only three of the five are valid species.two are consDecific.and
ore i\ norr Crrorirr. It ,5inlere.ting
thatBor.r.engeas
| -C;lalogue
I l8q4d
of
snakes",considered
a fahlycompletecompendiumofthe thenknom species
ofsnakes, made no mntionot Cc,lubetexotetusLjutlaeus.a vaLialsDeaiesas
recopized herein. A fe$ olher( $ho perperudledlhe nseol H?t;eroJNa\
s ereBoellger| 188).I8c8r. PerdLca
{ t8c5.t8q6.1897a.
l8q7b.tq04),BarUour
(19141915),
Boulenser(1915),
BarbourandNobte(1920),
Seri6(t92ii, ctiesch
(1925),and Schouten(1931).
The transition in the use of Helpetodryasto Chinn rrr was quite apparent
in the Literaturelrom the 1920's.Ruthven (1922)used Crlforlrr for the first
time in nearly 100yearssinceFitzinger (1826),andin a rather obscurefootnote
he called attontion to the fact that Linnaeus' Colrra rdtrdlr., was tvDe-sDe_
ciesof lhe genusCrliontr, b) originatdes.gnarion.
,qmaratr lq2) reiisessed
fhevalidity of Eryetodr|asandHelpetodryasa^d concludedthat Clilorirr haal
priority over the lomer two genera.Chilo iusFltzinger
EryetodtyasBoje
^nd Crtloriur as the
were both named in 1826;Amaral,s reasonfor establishins
\dlid generic
namewa5baseduponthe faclI halfirzingerie.ignareo
a q pespeciesfor Crilorlff, but Boie made no t)?e designation.
A rcview of tle dated Liieraturercvealed further evidencethat Crilorirr
l]as piotjty over Eryetodrr)ar.The date 4 June 1826appearsat the end ofthe
introduction in Fitzinger's 'Neue Classificationder Reptilien". However,this
does not help determinewhether Fitzingels or Boie's work came first
becausethe publishing date ofBoie's paper,Erp6tologie de L'ile de Java.was
only indicatedby the year, 1826.But anothersectionof Fitzinser,sa1826)
_ tbe ne$
inlroducrion
is rran.tated
species
ol my friendHerr B;ie wiu be
published next", in a separatework, which shall appearunder tbe title , Em6_
lologiede Java. and Boiet 1826)
$role 'Findinghose\er wilhin lhe sh;1
heatise, published in Vienne by M. Fitzinger, and entiiled: Neue Classification der Reptilien,.....".Theuseofthe future tenseby Fitzingerandthe past
tenseby Boie indicatesthai Fitzingert work precededBoie,s.and hence
Chironiushas pi'otity o'ter ErpetodrJas.
FAMILY STATUS
A1l recent authors placed Chitunius]n the famtly Colubddae basedprimarily on hemipenial, osteological, and dental characte$. This aenus ;as
includedin lhe highertdxaot lhe folo$ing selected
reterence<.i
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,NatricinaeBoulengcr,1894a- Colubridae,ColubdnaeDunn, 1928- Colubridao,Colubrinae.
Amaral,1929a- Colubridae,Colubrhae.
Roner, 1956- Colubridae,Colubrinae.
Underwood,1967- Colubddae.
( olubrin.e.
Doqling.la6- la-< - Coluondac,
Dtfr\'rrIoN
Crilorirs. medium io large sized,tenestrial,arboreal,
aglt?houscolubdds;hemipeniswith sing:lesulcusand lobe,nakedpocketproximally,spinous centrally,calyculatedistally,andwithout capiiationofapicaldisks;pupil
round;lull complementof colubridheadscales;123-196
ventrals;96-208sub,
caudals;midbody
dorsalscalesin 12ot 10fows;keeljngandapicalpitson dorsal scalspresenlor not.
DLAcNosIS
Crlforlrr is the only Neotropicalsnakegenuswith 10or 12scalerowsal
midbody.This chafacteralone
separutes
Clllo,?/rsftom all oi]ler Centraland
SouthAmericansnakegenera.
DrsrRlBrmoN
Ci,Tdrtrr has a Neotropicaldislribution,extendingfrom the northern
coastofHondurasto Uruguayand northoastAlgentina(Fig.1).Most species
arelound eastofthe Andesin or aroundthe AmazonBasin;Tablel indicatos
thc sleciesklown fiom eachcouniry. Crrorlw has reachedthe Islandof
Gorgonaalongthe Pacificcoastofcolombia,severalislands
alongtheAtlant
ic coastolBrazil, and Trinidad and St.Vincentislandin rho CadbbeanSeaHabitallreferencevarieserlensivelywithin the genusandwithin certain
species-Lowland tropicai rainloresiis preferredby Chiorirr, but humid
montaneforestsandsavanna/open
formalionsarealsoinhabited.The elevation rangeoftho genusis lrom sealevelto 2800m. The rangeand habitatof
eachiaxon are explainedin greaterdeiail in the spociesaccounls.
DtscRltrloN
Total lengthrangesftom 274 mm to 2718mm; tai126.943.3percentof
total length;ventEls 123196in males,131-190
paired.
in females;subcaudals

2Q

XX

!r^Irotl

x><

xxxxxx

x;t
XX

X
X

XXX

XX

xxx

XXXX

XXX

XXX

XX' <
!

;<t

:;

l:

:E::

-1

*,i, *i: i!: s: iii:ir i t

I - Disiribution
ofibe senusCrtoDirrFitzinscr.

22

:l',tiiitii::i,iliiti{
;ili::f,i,J.,ti;.:,,",ilTtJ:
i"Eliii
ilii:!l?fr
rry!ff'l#lifi
#i5:,:TTtH,i",*ff
[i:?:":ln
J,,f*t*la

trffi:ffi::Hf
i;ffiittf;Sitii'#i:#":.fiTH,:i:",,lifi
:ii?T;
;,tii';i]';lti
;Ht:;i::i,xt:ll,i,,i,i,.:.
i;i.,,?i
;;:*Tl*j;:
l{:
.ii;i:,
;',,::il';,
:;;lili',i,i,";ll'illl,,,'i
ilLl'i:i::i;"i;illt;
ji;:il:::Tlx'
,"'.';
i,T.lJ..,lll,1;,:,i"ii'":i,'.#:.::$:1,;
itrifi
; "5i;
;;;i;:;::l;tj.i
:il.Jj
i:::;
::ri::+li:i;i:1.'1,Til::il'.:i::
e,
nur
i, ided,
ancu.u,;.).e',m,.
:ll^.".,.9:l:
ll J', :-tl,.pcrmaricrr,.imI
H';;,i'l:;i
l1t;.ti1iff
l:,".';;i::ii:::;1,"J{:;
:"ffi;:1"xj;l
:",:
il: ::i;::ili::i,liili:;J"1t:iJ",i:"lillL:u,
;T];i;::,J1';T:l
.ron_
I p " !o c u ta rlpteocrrdr
-5ral ,L nol i l Lorl l cl si th

qurre\rfidble..r.uati1t2 t2_,0.t.t_r2.8.
tU t0_t0,ur t0_'|U.8

mcl,,D;nuu(
'finge.chJnpingro papilJare
fihge. ci.tllli

no,
be\a'iab,e
,",.ilT,1;ljl,l,;,1i3:::J
i,:lfflllfl:''-dvo'm:'r
o'',,.n o*. i,", niri;:-,; .;;l";;;.
iH :',,:i::
":]i::r:JJ]::1
Speciesof Crnurirs are ovipafous.telret

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

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

ot 2lt tr\

t Chirantusbkdtitlotut

twrcd\ C cot
...tta,^)t..,v\e".e,r:c.,.,,,'-,,n,',"*",,,,.li,.l"lllliilijl;ilTl;j,i
t ned tu . tJ a E t.a /./,.,/\/r.c r: l L i ).l r cL5r:C I/.,.n,r,/rfr,,
ne$ \-bsD ;ci e,l
I Ftrtnat\quamt\ (petets): a /r.aj, r//l\
tS.ed,: C /r,//,"r,,, ne{ sDe;ies:a.

';:,1::::;l;,:,,1i,:,i:xffi
i;::ifff:::1'.Jii!,,,,i,i];,i";!,!!i,"!ti:
;::";*",:':,7,i';
i;:i;;JJ"r:"T|:
;l,l:;;i?,"
:tii i,1;!!"1,i
;::1,:;
;.:"1i:i:t;
:"'.il.J1i'
ili*1;i;il];:.1",,?fil.T,,..:'"'::;::;l"j:;
reJ,ion'hp smons
.pecie.
rr-ese
bur{ h:.hhxsn;

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

23

E6

':

fl

-l

6
3- e*p

EI
EI

;t

siS:s
Siiirs:isiiiS"

24
CHARACTER SUMMARY
Sizeandp rcpatiol1s.ln tS

of Lhe20 Chirah
size
tranremaes.
rr,esm^rd;
;;;;"#;#ff
n_'t4ti...fu2-tt -r nrJ

#*iX*Xlij;j:?

h rctdllength.Ihe "ge.l,pecim.. r, ;rrr.," i .

r;;r:i
"
./rd,?bI /d mr In tolallengt'.Sma.lcsl
andrJrge\l.lecimen.
e\am,nedtbr
ea.l_rd\on.fe t,-c.errcdn the.pecie.arco.nr.,
I herrl rsroughjJ
d thi-J.orsJigh.l)
more.or lhetolr.,englni0 a.trpecicr
at" ( h;rcntu'(frble 3t.ln( ,l/drlr. lhemerr ofl heLaitasa peicenrage
ot rorr,
length fangesftom 31.3-40.1
percentand variesfittte letween sexi oJ eactr

E
u:

M"ostmtios consistingof numerousmeasuremenrs

raten on the head
weretound to be relativelyconsistentwithin the g"nw una
*"i" of ti.ii"i
taxonomicvalLre.Inall taxa exceptC. carinatr

;:1ififi:,Ji#;if
,"*Ti;
l"n:{*;:i?,,""ma;$ffi
"':::lf

(o i a rn e teor tc J e )/J J/4 \nout

A m o d e d l ee )e
l engl h)uccur.InaLl r,l
,
/e
nut pn rL r. t.td t, -i
0 rttt a n d (. q u a d ntan .!tus.d1dI tdrgeeJe
\ . c r / q r o a u a t r n g
\o,aretcrof
e!
\0 o ,l re n g th )i s o o.,e_5eoby .raul trnt . tr
ol he, \peci es.

i#'J::fiTJJ::lT$Ai;i:1"ffi:'.Tl
",lll;:i:litff
:ljl.'ffi
;J:ilt
lowing prey which move quicHy ;nd erfaticaily.

lefltah Ventral variation is presenredin Table 4. The

lowest ventral
count of 123is_in C snrlet r and the highest numoe$ are
m C: n ?rlDrrlrr

, d . 4 / d r r $ \ ' c h L d -a 5 m a n ra \l q b .C c n erd
).r . tadt,ai , C . n
a t / r a?r7 r.p e L i e .o J I h e n u tri rc n tri , s p eci e,comde(
hdvc the hi ghe(tnum
Def or \ e n .trrt) | h e rci r c o n .i o c rJ b l eo v efl api n the tdnge
ot.\cnt-,1i ro5 ma,el
ir no - c m l te s In ,c d i " r.\o 1 . B ,)e d o r m ea,,. femal e. rcno
ro nare.ti gh.l l

moreventrals
ihanmaLes;however,
in sft ofthe20ta,ramates
averagea
ioi!
thanfemales.Thenumberofvenu;ls oftenvariesal"rg g""pr"pfri. g'."Ai"'"t"

:uocdJ.,ri'\a.e-oJmaJ pa.rcd
e\cepr
for.he,ermrnrt,f
ine
.,,llf.
s nr c n r al]/id^
5 rt.o c o u n l e d
d 1 d i n c rL d e d l h,( chE fa.rer.S ubcauLl al
vcrj ,rl i on

-n
is
shownin_Tabte5. Crilo, iustaetricoltis
hasLhefe\r"rt,ut"ouaJ. *tif, u-tr'rioi_
mxmof96.Thehighest
subcaudalcount
of208isi C..iiliitrt" iri,i"i_
r r - A gd j n . m e m b e rsu t .h e n u t,1 )e n t n,,D e.i e-.o-npte\
"
h" i . ;;" ;;;;;:,

nulr Def u l \u b c d u d a l ..\4 h tc C . ta a t, a tt i \.t..!uadrtari nau.,

andr . t ul nth"
ols ll) c lly ta \e Ih e l o $ e (t c o u n l \. L \te n\rve overl af
occrr i n the rungeoJ

subcaudals
fof malesandfemalesin eachta-{on.sir"a on rn"*", i;;;1",
lc0Jrohd\e fe{er.rb(.rudat.lhir
mrres.
hose\er.In\ir ot lhe20rd\dmates

J v er J ge ol e s \l h x n l e rn J l e .,T h e n u m b e r ot." bcaLdal si eouenl l l

\dne.al ong
geographicg:radients.

7
=2
=

3S1S9133933?933939s3
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gEsogS!3!S ageggE A ig S
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,

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- ;;;i

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; !ii;:ri-:i;i:? ar :? i

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,"i i i.l ! ! i I e,i : ; I ; i : t i

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a aes s :g-3s
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26
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6

9 ? 3 ? e 3 s n si99$33Se9Si
A' o:

Ef?i::if?l::{:?iEiE

? = ?': ?l E E ! e c i E i=q i = : I I
i:Se-!32q;i!q!?e!:=!

333Sss=5:

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+- - .;<j r - j v i 6<i

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? I

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"=Si a ii-nn t
?! i! E! ! : :

; g g g R g-

a3R

5 e=

F = - A h *

ii sr*sE
cs::
ii"

21

;.

^.-.^lq^l

i
E

^^^:

^^^^

i : o : . . t n n o !\q 1 q - - \q 1 q \q n \- c]

Li
-r

ii

E
?

i .i

q..r

-i .i

e? o9 n

\q

!i

-t

n..t

; ic

i;;

E i iEi

i3 G

ti

F 35 -

E
z

s i e : * ri::rili

28
lndlp/ale The conditionofthe anelpiatein all taxais shownin Table6.
Chironiusfusa.$,C.laeriollis,C. mo ticola,ard C.tcrrxrlrr consistentlyhave
an enlireanalscale,and a dividedanalplateis presentmost frequentlyin all
orher specjes.This characteris quite variable in C arolrrr where an entte
anal occurs14.2010,
a divided anal 82.2%,and an incompletelydividedanal
l.60/0.
T,be 6- aoddr on o

1 analllar e n r \ \ u l /

TAXON

t 'o ? L r
Divided

3
I

78

150
155
180
12
453

Anal plate
lncomlletelydivided

20

222
7
33

111
28

89
30

50
44

55

-4
I

130

Keeledscales.Chircnixss.rrfl1lrr is the only specieswhich exhibits no keeling,In all otherspecies

keelingoccursonly on the doftal scales,
alwaysoccurs
on an evennumberofscalerowq alld ismostobvjouson the posteriorpartof
the bodybut mayextendfromiust behindthe headto welldownthe tail. Cljro ius quadicafinatus is consistently the only specieswith more than two
keeledscalerowsand may haveup to 12rowswith keeling.The numberof
scalerowskeeledandthe degreeofkeelingchangeontogenetically
Geespe-

29
ciesaccounis).This chaGctefisalsoscxuallydimoryhic,anddopendingupon
the species.
maleshaveeilher more pfonouncedkeelingand/or more scale
rowskeeledthanfemates(Table7).ln tlxa wherethe paravertebral
scalesare
the only rows keeled,adult mdlesare usuallysircnglykeeledand femalos
weaklykeeled.Inspecies
suchasC Drtdnndrrs,
C rxoletus,C.J16va
lineatus,
C.
gru disquanis, C. lduenti, C. nofiticala, C. quadricarinatus,and C. ti ce ti,
whichmayhavemorethanlwo keeledsca1ercws,
theparavertebml
scalefows
alwayshavethe most prominetukeelsand malesgenerallyhavemore scale
rows keeledthan 1ema1es.
Atical litr. A,picalplIs arclairly conspicuous,conflned only to certain dorsalscalesol thebody andtail,andlocatedatthe rcarofthesescales.
Two pits
pef scaleoccuronsomespecimens,
butthe mosl numerousconditionis only
one pit lef scale.Wherea pit is presenton a keeledscale,the pii is usually
found in the upperportionofthe scaleabovethe keel.Severalslatesofapical
pits occurin Crriorlrt rangingfrom completelyabsentto phs on the dorsal
scalesofthe neckand on the paravertebral
scalesfor the lengil of lhe body
and iai1.Apical pils only on the neck scalesis the most common character
statein mostiaxaof Crilrriur (Tab1e8).Apicalpils areeitherabsent(52.2q0)
oI prcsenlonly on $e neckscales(47.8q0)
in C 4rddti triratus.'fhetendelLcy
for pits to be completelyabsent(95.4q0)
occursin C. monticoLd.
Dotsalheadsa es.The numborand arangementof thesoscalesdo noi
varyin Crtlortrr. A iypicdlcolubridcomplementofscalesarepfeseni,incLuding one rostral,one frontal,and a singlepair ofparietals,supraoculaJs,
pfefrontals,and inlemasalsSupralabials.
Supralabials
rangefrom 7-11in Crtlont r (Table 9)- Most
species
havea9/9condition,bur S/8occu$frequenllyinsometaxa.Crriodrr
quadtininatusis Ihe onlyspecieswhich
hasa higherincidenc
ofeightsupra
labialsratherlhan nine.
Suprclabials
e teri g arbir Thenumberofsupralabials
enleringtheorbit is
prescnl.Thosespe
largelydependentuponrhe total numberof supralabials
cimenswith nine supralabials
usualy havesupralabials4-6
enleringtheorbit,
whefeassnakeswith eight supralabials
haveeithersupralabials
4-5 or 5 6 in
Ihe orbll. Chito iusquadicdrlratwis the only species
in whichsupralabials
46 entedngthe orbit doesnot occur at dl1(Table10).
Infralabials.The trnber ofinftalabials rangeslrom 8-13in Crtloatrr and
variesconsiderably
intmspecifically
wilh the norm usually10of 1l (Tablel1).
Grtdh.Two pairsofgenialsarccharactedstic
of all species
with the only
excepdon being rhree pairs in lwo specimens01 C exoletus('lable 12).

l0
I

tl

E
E

2
3

E
E

F3

ril

,:
I

-l -l

.n
Q

:9:r

"R

:';:i

t9
92

||

R;

-e j

o -s -

!e

E : i,
!r{F

i s:e-!F
* !:i i i

3 -e

:,8
!:i

-:

3l

:o

-r

n;

^o 30

o;

23P

z&i

tttttl

9a

*::iF

.:i
;:!i i
iii! t ! i ! i :
SEEEES

- :i.!

-:

12

&z J

: -sE

z'zz
hE
6

td 3

_d

4z n

a
i

E ; !a

lx

6
p

t-

Trbte 9

Frequcncyofslpralabi.ls nl llxa ol Crrorirr

TAXON

t1

25
11.1
26

2i3
126
261
l :1 5

l
l t5
)2
2

:r69

6
ll

176

2
1

1
1
6
58
ll

52
145
i4

2l

',

2
I
2
5
1
)

90
'14

5
1

2
41
178

1l

Inlidlabialsontadiry gerl.rr. The rungeofthis charactefin Clilontrr is

lburlo eightorcachsideofthechin.
Themostftequent
conditionsin
alltaxa
are sft or seveninffalabialscontaclingthe genialson one side (TableU).
Prco.!/drr.Therc isnormall,\7
a singleprcocularin Crlmrt?A,but two preocularsoccuf in somespeoimensin 1l ofthc 20 ta-\a(Table13).
Pdslo.,r/d. Oncto four posroculars
arepfescntin Cr,rorirs with hYotredominatingin mosttaxa(Tablel3). Membef ofthe mrlt0ertlr spcciescomplex andC.lla"olineatushave
a felativelyhigh incidence(above$qo)ofthrcc
Postoculars.
Tempordls.Thetempot^l
conditionis quile variablewithin the genusdue
io unusualfusionsor splilsofthe basicscales
involved.A totalof 30dill efent

l4

--S

al

- 11

tttl

!t5t+

3\

r^ -!o

':
.!

5
.9

'E

a
g
3

i r u i r 5 i : r i i tlur

l ble ,l - l eq .elo

of Cl f ur

| dor al' r v
'
'

"
INFRALAII

TAXON
2

2),
I

I
l

21
1
25
8
4
3
3

IJ
2

3
l

20
l

\LS

t0

1l

i33
91
81
109
219

75

257
22
l

33
22
81
22
E
52
35
3
54
38
l

l3

12

t62
50
350
32
80
2
50
2A
42
12
19

21
12
123

9
I
I
5
2
I

l0
10
5
23

5l
l

1+2,1+1+1,and1+
cbaracterstatest{'ereobserved:however,four states,1+1,
1+2,occurmostfrequenLlylnChircnius(Tablel4). The condition1+1wirha
diagonalsplit acrcsstho anlerior upper quarter of the primary temporal scale
prolably gavedseto 1+1+I ( f-_ r--l _ t'_ f-- r ), and the samob'pe offission
eaveriseto t+ t+Z from 1+2 { l------1 - v---F=\ Those specieswith 12scale
iows ai midbodygenerallyhave1+2temporals.Exceltionsto ihis areC
'icdtemporals;C eirDlftrr
ot 1+1+2(7 30/0)
I+2 (43.90/0),
/indrrr with 1+1(46.90/0),
andC no'11t
\\ith 1+1(8360/0);
of 1+2(89.8oh);C..flawlineatus
with l+1(7.1q0)
The state1+listhe domin(61.50/0),
or1+1+2(1780/0).
coldwith 1+1(6.5V0),l+2
l0 scalerows(C.tr'at
wilh basicalty
anl conditionin threeolthe four specjes
wilh
l0 scalerowsat mid_
species
The
fourth
C. laevicotlis, C. saltrulrr)
^t\d
(13.60/0),
or 1+1+2
(17.80/0),
t+1+1
(12
1+2
1+1
6q0),
has
body.C.cn disqudnk,
(45.8q0)
tempo|als.

gj
fi

36

trt

6o

ll

I
{

'E

toae

E
E
- llll-

ttl

iir*eci::riii,

l7

;lill

1 6-

t6

3
F

l-

Ll

n
E
6
a
.=

a
5
d

:rei e s t : : i : l * ;

l8
Tablc 14 - Frequencyof temloral scllesin tdu of Crim,?,r.

TAXON

l+i

t+ 2

r21
5
8

119
233
291

'11
r01

898
,]

29
21
52
3
1l
I

38
2

TEMPOTATS
1+1+1
1+1+2
l

20
3
6
7

5
2
29
2

12
1
22
l
23
\2
I

98

22

3
I
5l
I1
90
'11

'j

;
2
5

a
2

5
231

91
1
6

loldk. The number olloreals in Cttlo,?i,lJmngesfrom none to two with

one lorealgreallypredominatingh a[ ta"\a(Table15).
Pastcephalics.
This chamctershowst.emendousvariationwithin each
taxon and rangesftom 4'11in t.hegenus(Tab1e15).There appearsto be atrend
toward tewer postcephalicsin specieswith fewer scalerc\\s (C.Ji.tscus,
C. laert
cc'l|is,
C. scutrulus)N\d/or with 1+l temporals( C.flanlineatus, C.fuscus,
^nd
C. lae|icoLlii,
and C. scwrulrr). This agreeswith an overall trend in Crrlo,rrur
towardgenerallyreducingthe iotal number of scales.
M&xi arJJteeth.All Chitu lrr are aglyphouswith no diastemaor grooved
tegtl on the ma-\illarybone. Mexillary teeth arc recurvedand increased
slightlyin lengthposterioriy.
Variaiionin the numberofma-xillary
teethfor all
ta"\aof Clr,iorirr is presented
in Table16.Thelov/estmzl-\illary
tooth countof
24is in C. exaletusand the highestnumbers occur in C/rrod&rcrr which has
asmany as51.Figure2 illustrateslhe maxillaeolthfee speciesof Crir'otdr.

l9

itr lt

tl

@ ti tdl

- *i

tl

.t

tdr i6_

to

!
;

li

llld

3
a
E

i:

fl

al -l l l l l llllll

i i $ r e eciss::iEt

in lengthposDentaryteeth.Teerhonlhe dentarybonegenerallydecrease
in
each
sna,ke.
Again,
toth
number
ofmaxiliary
leriorlyand aboutequalthe
the
highesl
wiih
26
and
is
in
C
xolelrs
dentary
teeth
lowesl
number
of
the
count of46 occuN in C.trscrsr.!.rr. Vadation in the numbet ofdentary toeth
is shoNn in Tabls 16.
dorAbbrctiatedda6at s.alerot,/o/r?r/dr'The moslcommonabbreviated
Table
17
Thenumare
shown
in
salscalerow fo rmulasin all taxaof Crilotlrs
berofscaleiows andthe typesofreductionlhatlakeplacearevariablewiihin
(are)relalively
ihe genus,but the basicscalerow lbrmula(s)for eachspeciesis
the
consistentand arethereforetdonomically veryuseful.In many species
scalerow formulas(basedonthreestandardcountsexplainedin themethods
thai thisshaseclion)arcsexuallydimoehic,andin thesespeciesitispossible
males
in
severalspecies
As
will
be
sho\dn,
sexlinked.
mcteris chromosomally
tend 10losetwo morescalerowsposteriorlythan $e lomalesThe adaptive
towsposteriorlyis provaluefor femalesto not reducethe numberol scaLe
purnecessary
for reproductive
babh,relatedto maintaininggirth advantages
(e.g.,
eggs).
carrying
loses
in certaintyposofscalerow
Althoughlhereis overlapamongsomespecies
existin this chaftcter 1) Chioniusbicai'
fofmuias,severalintereslingtrends
roduciiontbr
is the only specieswhich hasa fairly consistenl12-12-10
'1dlrr
have
a 12{2-10
species
complex
mrltrverlrir
olthe
sexes.
2).
MembeN
both
in males 3) Clrionius qucttlicainatus gene'
formula in fomales and 12_12-8
raliyreducesto 10postcrioflyinfemalesandto eightin males.This sleciesis
unusualin havinga high incidenceofmore than 12scalefowsfor the firsi or
a very
is considered
antedorsiandardcount;this condition,(i6-13)12-(10-8),
ofdeived andprlmitive
primitiveieaturcin Crirorirr (for lurtler discussion
characterstatesseesectionconcemingfonnationolthe phylogenetictree)
arrdC. mo ticolalendtohavoa l212-10formulaln
4). Chircniusl16t)atineatus
in males,but both reductiontypesmay occur in both
femalesand 12-12-8
a relativelyconsistent12-12sexes.5).
Chiloniusun atusandC. exaletushate
seemsto bfidgeihe gap
laericollis
8 reductionfor both sexes-6). Chnonius
thosewirh 10.This sle
rows
and
12
scale
species
with
basically
betweenthose
cies can have 12 or l0 scalerows at the first standard counl or at midbody.
Femalesreduceto 10 scalerows posteriorlyin snakeswith 12scalefows at
Malesusuallyreducepostemidbodyor showno reductionat all (10-10_10)
dorly to eightscalerows.'7).Chircnius
fusa$ andC r.?mlrrshowno reduc_
tion (10-10{0)in femalesand havea 1010-8fomula in males.8). Clilottrs
grali(lisquaniu is the only specieswhich has a fairly consistonl 10-10-8reduction fof bolh sexes,
Summationol mmplete dotsat scale tot'tfotmula$ Summations fbr completedorsalscalercw fo rmulasareshownfor eachtrxon h Table18 Inallspe-

Fig. 2

Mdillae of CtuD,tun verlrolaleral spect.

cieswith anabbreviated
fofmulastarting
12-12andinC. grundisquamis,Ihe
fusionofscalerow numbers2 and3 areprimarilyinvolvedin the reductions
irom 12to i0 andfrom 10to 8rows.ln C 2r.?,r, C laericollis, C.scutnlus
^n(l
the fusionof scalerow numbem1and2 aremainlyinvolvedin
the reduction
from 10to 8 scalerows.Baseduponihe meanofthe ventralswhere12ssale
rowsreduceto 10and10to 8.thereductionsin femalesofall tLxatendto occur
severalventralsmoreposteriorlythan in males.The first majorreductionin
alsohaveanunusual
all species
occursjust behindmidbody,andmostspecies

42

;;

i. _

-:;l

++++l++n ++ll+
E, 7r
l]:ief
t:

a l: q

-ra :::._ = -,4 2

:1
2=

;^r
"" P

f
?

;i
i:

i-.

f :f

i
1

;i9
s 3:

"

E
r6

t-

ts
f

a
e
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t

i ;i "1?,r:

| ::

" -!

2;,"

;3

E
E

liir*is i r s : i : u e :

'{3

dl'bi'

ffii#h
*'%

ffi|ffi
catyceS

"rr^."ffiffi
ffil'ss
ffiW
'"""',*,r*
b-----,..-WF
s u lc u s
s p e rln a lic u s

1i

Fig. I - Everled hemipenis of ChnahiB fusds iads,

s p in u I e s

U'l/. 1100.shoMlg chdacteristic struc

dd

rrd

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rl

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R*=
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'i.

= F ^, - n= =
+---"

oii.

a ::i

"31-

!.\-r9
r =:
=F3l:aFr
;+fii;+ii

;:l

r ax

R+

!xl
ri
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11:,

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9939

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=ceitE!-i=i] i11- lilir

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50

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=^r

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3::3
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Li :

aa

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9:
33

r
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52

a oe'c
:+.1:
2t

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a
FN
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;AIJ

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r5

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=.

n3a

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gl
EI

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FI
!l

+ +
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-q l

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:
-

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5
l
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EI

lEll

EI

il

zl

EI
5l p
FIfl

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!; ll

l
sl
EI
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EEI
=: l

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Ei seiEEi:t.

il

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d'

+,a

+ 6

;s

9 t

q ?

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?c

gE

TF

1++

t:J x ::i

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.=
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r+i

!E =2!

t
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55

! i ! re H
ZzJ'-d

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56
condition of gainingtwo scalercws somewhelebetweenlhe last standald
count (of the;bbre;iatedscalerow formulx)andihe analplate ll is nol indi
caied;lscwherein this lext, but one specimenin eachofihe followingtaxa
(12'12-6),
C fusus
reducedposteriodyto asfew assjx scalcrcws:C ei.oletus
(10'10-6)
(1010-6),
dhquamis
and
C
sra
lrs.rs
the dorsal
Tditreductionfronsixtoforrscale/olt The subcaudalatwhich
presented
each
ta_xon
for
is
scale
rows
sc, esof th e tail rcducefrom sixto fouf
subcaudals
theseiwo
subcaudalswhele
meanofthe
on Table19.Basedonthe
are 1ost,tbis fealuctionoccuNmosl anteriorlyin C exoletxs,C multi\)et.ls,
and C rircrrtiand mosl posleriorlyin C /der;.o//ir.Membersof the tr'lllt''
in C
btll .hela:l educrron
,r: .pecrcs
conpleragreiin \o mcnJchrraclers.
(
v"1'
d'l
nutttvct'tIt'
pos,ctiorlJ
i'
r.
tler
/arlpnl,occ- . oi..inctllmore
dnQeres.The hemipenesarogenerallydescribedin lhe genericaccounl
accountsSelected
in detaiifor eacht&xonin the species
aboveandpresented
ofthe hemipenesof Cr,Torlrrhavea relativelysimlle moryhology
characters
and arc easilystudiedln theiiretfactedof eve cd condiiions.Figurc3 i1lustraresal eveitedhemipenisof C r"t.urrdr't and showsthe characiedstic
in thissiudy.The only Crirdri[themiponesprcviously
structwesemphasized
of
C cai atus, C fuscus' and C gnnclisquamisbyCope
weie
those
illustmted
(1895,1900),and C lrrdrr bv Vellard(i946)

IDENTIFICATION OF CHIRONI US
arcnotidenli
of Crnonirr' Subspecios
Akeyisprovidedfor the 13species
fof
of
subslecies
number
the
aplropriale,
where
key,
but
fiable u;in;this
rel-erto the
eachsDeci;sale indiceted.For easyrecognitionof subspecies,
aliagoosisand descdplionundof the speciesin queslion Severalspecies
variatlon
morethanonceinrhetrey This rellectsthe $eat intraspocjfic
aDD'ear
most
charactef
nature
of
overlapping
in'Chhoniusand wide mnging and
Other inierent lroblems in conslructinga useful key are due to sexually
dimo$hjc and ;ntogeneticallychangingcharacten.Becauseofthe overlap
to be idenliof cha;acteistagesoiranges,ii is recommendedthat specimens
most keys,
tey
As
in
fi,3dbe laken all the way througheachcoupletin lho
should
properly
Thi!
key
out
canb; identifiedor worked
nol all specimens
juveniles
(specimens
but
snakes,
ad
Lllt
sjzed
be adeq;alefor iclentifyingmost
seneraivunder 400mm snout_ventlength)shouldbe checkedagainstlhe
"aiasnoses.
distributions.and descriptlonsmenlionod in lhe sysiematic
acc;unts.

57
KEY TO TIJE SPECIES
1.

l0 scrlerc\rs at didbody
12 sc.le rows at midbody

2(L ) \n I p d.ee o , "

\iblFLre d inded

.3

3(2), Ar leasllove. Dortior ofslpralabiah liehl color

Al1 supr.Llbilh comlleret blact or dark colored

'l
lodi@ltis

46). No keeli.ei naxilkiy teeih 13.41

turavedebFl scalerows keeiedinaxillrit leetb 39_51(2subspedet

scuttutus

5(1).Anal plale entne

Anal Dlatedivided

/rrrt
6
9

6(5). A1 leastlowei ponior of sq'nlubi.ls light coloredI tempoEh usuellv1+2or olherwls,

1
bul usully not I+i
All sulrllabift conplelely blact or d.rk colorcdr remporils l+1 consistedv
l4erieallis
'
l(6). Noapnalpirsioiienuithlor2pansofbrordblactlateialstdpesonllillofieroorethan
2 dondl scalerowskeeled.
Ariftl pitspresenlon aLleusldorsals@lesoflect resionino biold blackhteralslriDes
8
o; tailaont ldavertebral scalerows kccled
ventalsdd 111_
fcmalesivith131-162
renfals ald 1]&160subcaudals:
8(7).Malessith 123-155
Maleswilh 163196lentralsald 156208subcaudalqfemaleswilh 161190venlialsdd
nuttirennis
157205 subcuudals(4 subspeciet .

bv blact
aDteriorlv
ud boidered
9(5).tsightyelloslertebralstipepresent
lir n: . a- . s
verteb@lstile presenlor not,itpresent,not brighrvellov borderedmteriorlvbl bhck
10
sides abovc the irsr scalerow
10(9).4 or more dosal salc tuws keeled
Ody 2 (paiaveneb6lt dorsaLscalerovs keeled
ll(10). Morc than 3l mdnhry reeh.
Lessthan 38 mdiuary teerh

ll
l5

12
13

DoBun unlfom darkbrcmr 40maxillaryteeth;naleswith lTl venlrolslnd i2 l2_8scale

12(111.
rcw fomulai femalcs wiLh 15?subcaudals;found onlv on St. Vincent lslmd to"nri
DoNun usuauyolivc silh lieht veltelral niili 28_40mdillary leerhi dlles vitl 149_164
vertrah. 125157 subcandalsdd 1212_10scaleiow fofrula; females wilh 154169von-

subcaudalsi
nainlandSouurAmenca
lrals,121-153

i8
I3(t 1). Distinctrcftebol sLripenot presenridoBal sate row fomula pducesto t0 or 8 poste
I o J . bc r r d_ . { ir ! . . r ol1. ^ o. r erI r .
LighLverrebralstripe usuall_\'presert; doBal scalerow fotuuta reducesro t0 posreriorlyi
subcaudalsvirh blact outei tip

14(13).
163-181
renrals:l4l{77 subcaudals
l,l1161verrrals:
104129
subcaudah
(2 subspeciet
v l0' Do o l \.d ' ero { o m u d e d -!e r.o8 .. prorr).
Du q

.c1 e o \

o r r JJ

e .lr r e ,r o r 0 ooc.eFo

16(15).Lesslhd 3lt ventralspius subaudah

More tlun 316ventralsplus subcaudais

20
t7
l9

17(16) Do$rn withDattemofeitherlalgeyelo*ish spotso! t6t scllerow oftail. lichrflecting

o n m o{ do^ar . dle. u r br olo eoor . h - D r o r d . . n p e ,ide o r t J . u b c d u o " t . t e t o r n i
dd ouuinedir blackor darl bro*n (3 subspeciet
cannaus
Patlemnor a ahove,geneiattyuntfom cotor wi$out disiincrspolsor broadlaierat
stripesisubcaudals
nlybeletlowish,buinotconDlctetyour[nedinblacloidarkbrown

18
18{17).163l8l venirals;brosnisholivedorsumwirh mottledvenlolatcEt pattemonDosterior
ofbodya.d on tail.
auren,i
Ul-162 ventralsidorsalcolo! yariablevitt !o uusuul noitled yeatrclaleralpalrem
19(16),Moi ed ventrolateral
latlem on posieriorofbody lnd on lailt subu{jalsl4t-177(aboutlso).
bukntj
No unusualmotlled venirolateral
t56-208(- aboul
lallem; subcaudats
20(15).l-es than 315venhalsplus suboudals
More than 314lenlrals Dlus$bcaudah

2I
23

21(201.Dorsum*ithpattehofeitlerlargeyellowishsporsoniBrscalerovoflait,lishlflec[ing
on mosldosalscdcs,orrwobroadrcddishbroM sl.ipesanreriorlt;sub@udals
yeltowish dd outlinedin btackor darkbrcM(3 subslecie9.
cdnnaus
Dosum not patlcmedasaboveisub.aldalsnay be yellovishibu usualtynotcodpteiery outlinedin llack or dart b
22
22(2I). Motlled lentrolateral patteh on I osrerjor of body and on lait i yenrals 163l8Oin males.
166181in limalesj dislincl vertcbral strile rot prescnr: subcaudalswilhout bhct outei
tips .
t.rtenti
No unuslal noltled ventrolalest pauem; vertrals 149164in males,t5iu69 h femalesj
lrsht verlebralstrile usualltliesenq subcaudals
w h blact ouler tils .
,,carm,,,,
23(20).Lisht venebralstrirc usudly presentiryhire
or letow subcaudals
{lrh blackoutertls
a.d uslally black zig zag Iine ncdially beiweensubcaudalpaisi nales wiih 149t64
vetrtralsand 125157slbcardalsrfenales{ith 154169leDtra]sand t2t-153subcaudats

Ty! e: -1
--.ei
wasdesigna:::
14
number of i: . s]!
eachsideol a: id

and 6 feea Ii::fl

tyle localiii i -'r I
(leagues)o: \ -i- |
ol12 dorsal
]!
-:i:
fowrcduc,ia::if
iormula is ri: .riq
donum \.i:r : ,
stnpes.!e,:;_:ir

59
renrah and jll llli subcatd'lsi
Paltcr. vnriableand nol.s rbovermuleswittr161_196
21
l'll_2oi
subcludals
l9orenlrals
i.d
fenalcs*iLh 16l
111lii ai
24(23).Ilottled vcntrcldrerilrratlcmon postefiorofbodv and on l.ili subcauCals
tauahn
aboutl5o).
(i
180)
25
pattern:
156-208
about
\ubcaudals
ventrolaLenl
.rodled
No unusual
onlton Sl
subcandaLsifound
25(21).Do6un unifom da* btu{ni '10raxillar} leeth:157_158
\,o .. Il;r'
marilho rccthr156208su|caudah;tourd
Doslm vaiuble iacolorand palterniS2_40
o

l.nrdJdd

lm .r o .

ic

.l \n e r ' d ,il

b ierF\'

SYSTEX({TIC ACCOUNTS
Chironius bicarinatns(Wied)
(Fig. 4)
Calubetbi.ninotus.w\ed.l82ll81iWied.1824:colorplate,!a-!eunnumbered.LictE.IsisU(10),
.Vrn zo lini. 19?l: 101.
!l2 0
vdrfiDr./trard (wied): Mercrn, 1820117.
Erretunryt!bnohdtus (N ied):Boie. 1826:231.
lk4)andtlas bt.dtutalut (Wie<l):Wagler.18301180.
bi.atinutalNied): FirzirCer.18411:26
Herpetatlryas
Henelodtrds.ahnatus(ior .l Limeus): Dundnl Bitfon, xnd D!m.ril.1854:201
Herydadryar.annatus\at hi..hndta lwied) Boeftger,1898:55
Chnanirscoinans lnot of Ltnmeut: Anaral. 1925:4
Crto,irs br.drtaM (Wicd):Biiley, 1955:8

Type: Atypespecimenwasnotfound.ltisnotclearwhetheraholotype
briof descriptiondoesindicatea
however,Wied's (1820181)
wasdesignaled;
"one row of keeledscaleson
number oflhis slecies'maiorcharacledsticsl
paifed
and".. itisdark
subcaudalsllT"
ventrals
155.
dorsum,
eachsideofthe
oiivegfeenon thc doNum,yellowon lhe ventef,reachesa lenglhbetween5
and 6 feet".Wied's(1824)colof plateis clearlyindicativeof this speciesThe
Iypelocalityis on a sandbeachofLagoa,nearthe Rio Jucti,within "5lcgoas"
EstadoEslirito Santo,Brazil.
(leagues)of Villa do Espirito-Santo,
with acombination
D iagno si s: Chircniusbicainatusistheonly species
of12dorsalscalerowsat midbody,a dividedanalplate,anda consistentscale
row reductionol 12-I210 in malesandfemales.Unfo unately,lhis reduciion
mlLsibeusedlor sepafofmula is not ahvaysdiagnosticand othef characters
tuling C. hicati atusfrcm somc othef taxa. This specicshas a uniform olive
doBum wilh a light vertcbralstripe(vcrsusreddishbrown anterioflateral
stripes,yello ish spotson first scaletow oftail, or yellowishf'leckhgon all