A taxonomic study of the sauritus group of the...

A TAXONOMIC STUDY OF THE SAURITUS

GROUP OF THE GARTERSNAKES, GENUSTHAMNOPHIS FITZINGER

By

DOUGLAS ATHON ROSSMAN

A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF

THE UNIVERSITY OF FLORIDA

IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE

DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

August, 1961

UI^IVERSITY OF FLORIDA

3 1262 08552 2661

f

iio tsuGonoaic study of this nature io possible without

cooporatloa of the curators of musoum and university

coilectioas, and i would llii*j to «xpr«a8 mjf si'îîtude to

the following individuals iiho pomittod mo to oxaalno

saaterlal in their caro (the accompanying abbreviations will

bs usod throuohout this study to rofor to speciaons in

theso collections): Ciiaries M, Bo£;ert and iiichard U»

^weifol, ûacrictui i-iuseUi- oi ^âourai niatory (Ai-H'iH): James

S. iJOiOke and x-daond V, i^îaiiiate, Academy of Natural ^ciencos

of Philadelphia (iHiiip); iviiliao êecher, Chicat;o Academy of

:9Ciences iC-î*); well i». laciuiMsrio , Curi.a^j-o ....... ..... »uo;

Robert k\ Inôr and Hyaon i'iarx, Chica^ô Natural aiotory

iriuseum (CliilK); X. Paul Maslin, University oi" Coioraco

Museum (Coi'i); i'hilip *• ;iaith, Illinois ijtate iiatural

History burvey li.itiii); Rilliaia a. Duellaan, University of

Kansas i>iuseutt of iiaturai liistory l^.U); Cieorê ..• -owrey,

Jr., ix>uisiaaa State University Museum of ^-oolo^^ t;*û*i^);

Willlaa ii« Dickinson, KUwauke* Public Museuta {: lii); J.

îiliso Ciibum, iiiseissippi southern Colio,,a i.-»-j; i^« -•

^ .idy, Mississippi Oaao and Fish Coomission Musoum (. ;;

Smest ii. Vsiiiliacis, tiuseura of Comparative U>oloar (i-iC-i);

iraacie li. Cook, ^National l-iuamim of Canada (iiiiC)j Curator,

ii

N«« Xork Jtato i.useum iLilHii) ; Curator, N^va SOOti* Mumub

of ^Glonco (:;û:i^); Edward 3, Xhonaa, Ohio 3tat« Koawai

(O^hi); u, ixÔiiK iilalTy univoraity ol ie:.a., ..aturai history

Collection (i.wiC); Donald n. Tinkle, Texae Teeiioolo^sical

Colleo«>^ vÂV/j; .>aix>xL >• . tiXidtDe, Tulane Univorolty (i.uj;

F* W* GibeoQf U. .ity ol Aricansas Departsient of ^.oolo^

(UATZ); WlUlani J, ..iooAr, Univeroity of i^lori.',^ Colioctiona

lur>; uerturci J* *>.ai'MOi , ^jajiverait/ oi' uooroi.a i^^î; liobort

ll« Smith, Ualvorsity of lilioois Museum of lia&ural iiiaLory

(UIIU4H); itoêr *., iiarbour, University u. fêntucky v-.^j

itoi'uiau £•• hax*twe£. University of Miciiigsn Museum of 'X>ology

iWii'i^)I harry «»• rroeiuan. University of uouUi taroiiJia

MuseuA (U9CM)} Doris iî.* vuUii-a^., uuûou awac. .uxuâl

MuseuBi (U;»J4M); .a>B8 i\ coll, Utiiversity of V t (uVj,

1 am also plervcxsc^ to ucknot7lad::e tr. ..y of

%tkm foXiAwxoti ,>ersoxis ior Lahh «^xi.t ui ixv^. . . ... , lur

assistance in tne field, ur for .lelpiul iafur.uation: kroig

&• itdler, Walter Auffenber^, ..^^^ .. .•• Axtell, ueevo i'i*

Bailey, uiciiord J» tialdauf , Jonn h« bartloy, J« uaenoan

iileakney, Joiin c\ ij^ranham, C. . C* Corp-onte^ , r'j K.

Clarke, J« .^^^.xÛi^ v/j.ûui^, -orxz *i* ôcû'a;., <~ .. ^,

Collins, Mo(s;er Conant, /loyd i»m Iôwns, Williaia ft, Duuli. . :.i,

HnrolH A, fundee, :•;, J, louquetto, Jr., Frederick U,

Ueitlbaca, Carl ucriiax^ u, itobort a« draat, Ji\, J« Alan

Holaan, iiichaxd >;, Johnson, od iLeiaer, Jr., e, -ayne King,

Cltarles .•>• ..*., ^cnubert L* uee, Jaass ^» ^x..^ , barry

ill

Minton, Jr, , i^bort H, fiount, Charlon , Myera, ..lirrod T,

£iolil, iioyd Ui* i'ottor, Jr* , i.uov'ioii i4« tyke^ Jos jii i'i*

Pylka^ John A* ««uix)by, ^lita J« iocônuitn, Uoxma aiid iinrry

tiotlman, C* ^ôbert l>f)Oo^ , i obart .<• ->.iûu^ r'hiXlp , wealth,

ii» C* Stuart, K.^.i uweot, o{:<i.> a, XQiford, Jr^ , LoaaLd ))»'•

Xirikie, viiiia'a u, Turricr, Tiioraras h, Useeii, Ji\, ;Pd

von cei" i*jL;>|.'j, î^^;... >;.-'i''ju, t..AC .,*"

. ia^ ..^.-. .-.. , ».u "ic. i-':"

i»orthir»i^ton,

leci r« .Xlen r. rared tho drawings of tae ribbonsnako

iiuous wiiXcii uproar au ^^ 1^ •

;»pociaX ti*£Uik8 are duo Arnold £»• Urobaau, uiiiiar «tiO0«

guidance this study was initiated, CAnd Arciae Carr and

..iiii<^ J» iviecaer, ttio ou^ crvised its coiii>>ietioi4| lor

constant aiicour'iêriient aJid ready -'jdvic«# i aiao vdsh to

C« tovoe, >.iijLiaia C, Maasoy, and o, xuiy ^,

ViOixl

tho di&uurtation and olXorod helpful nuêstions*

iv

AClL^i.uiiUi»4u.,i.^^^

UHt U* XAli4-E;>V^

U.;»X 0*' ii-i-^ ^ • • •^^^^

iNTHODUCXlCK •*•

h. u5 &iih . O

v» . -.^wion

3

THE yAUHlTUa GitOUP 6

i ion . . . • • . . 6> tr.o jauritus Group .....•.• 7

'«'.^«v,

tj proportioHG ••• 7

3 • 10'ion ii

. Co -»-^

tt«i f.s ^3pU .Ouat,;;» •••••. l^

1 .. .iti* tu .-*«

Hoiotype \^lieilaitiou ••••• *o

» ^(C

,. . ,..ioa ^iieot.i'«i>iiiiC Vf^rirtt=Aoa 34

TAai£ OF CC Continued

TaxoxuMuy . . , , 58

Key to tiio aubapecios of x:^^,.,.,.

^ro xiam^ 62h>. ^s oi' ii .... ...... o4Kv ui.^wxonax'y i.i- . .> •••«,. o3

i*iaturai Uiscory , , 6^

iiabitat .•,,. ^5iôod cJ6

ii' -01* • '6S

^ - 91

THc. :.iwii.i4i« *aaaoi4SKAi^ <-ji

i\ootype ...,,. yij

xx,xoii • • ^3>3

uion •••••••••••••••,••, 93

3v Limor ••....••.,.. 93Oi. ^ .otxc ..ion yylouzviciaaX Variat^xoa • • • xOOCeoâpiiic Variation . , • • . i04

Xa-xonomy 117

key to tne Subspocles of ThaBiiysi^liia

fffimy^VuB i-L^

3 oi" 013 ii9....—o-Loaary ...w 135

Naturai uiatory 136

Habitat 136iôod 137iic - '.ioa 139t. 140

uvuc«u3l0ttS , li^4i

146

i iixCfiit otsJcTCu ..•«. • LiM

vi

i, individual variation in maxLUary dentition

ill tuo oiauTitua Uroup ,•...«•*•••• "

• • iOd, lieaipeiiis Xeii^ta in the i»tturitu8 uroup . .

3 „ . Oi ..x.>.. J..i veatraia in ya^'tfM?^^*^^

4. Coori"ici©nt3 or »exuai diverêaco in six

species of Tha.ano;;hia ^*>

5. „ ,;4..ior,.ai8ia ia subcaudaLs in TtMJBitfWP^V^g

6« OntOoe:ietic c t . -- <'* 'ai

30

26

7. individual ri..îc cuaractcrs

8. ^'ood or ThAmnoaiiifi fen??Aam^'^

9, body Iea4,tn or newiy bom TiiaffiMBB^^ JBfQJâa • '^^

iO. ^c- . . iiaonmista in vontrals in |^#itf»g^Hij^

j^.,,..'-tua ''

il. s»oxuai dicioiviiii^^ *î aubcaudais in 'îmBIW>ifll4^

auritua • • . . .•...•.. :^^

\j 11 ^ ., . .iu.dta.c ciiaracters

13, *ood oi XâianDDiiifl gttWrttVAfl"•"^

14, i»ody ieui»ta o£ newly bom p^wiij^.îUii a^.^'-'iVÂi»^ • ^^^

vil

i« Head patteniB iii tao two o^<«scxos ol' timJ uroup in doraal vxovv (A) arjd

._ . ^ vicv; (ij},«. ••.•••....• 15

^ 19

Vi»»' V'.tiUii^J. ,.'>,.t«.>., .. > . • . . . . . . . • • • *X

4, C c vai- iii V of

5, C iiic variatio.'i ixi \ of

; . . 40

6, u.' ^. ... -' or

^r-—-'— I' I.- .- ii.tfn.î. -^ . • v5

^^.^crica • •••••••• ^7

Uû.tucl a«*« ;/5

y, v-jOiÛ' '' " -iutiou iii ventral numiter of

ami -^^6

1U» Ooooâ; hie vai'iatlou Xii euOCttucol nui.ioor ofYim;.. ^ :;auritua iu tuo uuxtod utatosaad t^..^c.a ••••• ••• j--^

Vi.Xl

Over helf a century has •lapsed «inc« iiutiwen (190^)

proaucoa iu.& ciansic £aoao4,raph of th« garteranake etinu*

T\>fflMlgP^^ff t A nonuuntai tiork which iiaa Mrvod lui a iaoUol

Tor sub8«ciuent studieo ot goographic variation in ophidians

and for taxoriomic nuOiî^r^îiiB goaerai-ly. in that Gtuui, u«

arrangttd the 12 opacios o£ Yliaiunoahia then recofeiiizcd into

two primary phylofieactic divisions, eacJi of whicî contained

two apQciea tiroups wtiich n« oaaad for tiie ^rii^ciiôi ^.acioa

in eac-i. Trius, his biviaion i cojaprisod tho .*adi-x Group

iuid tii© -/oui-iuus wroup and hia Diviaio.x ^* w.*w -iafcana

Uroup and the ijirtalia Group*

In tha course of hia revision, jcutnven exaaiiiod

approxiiaataiy 3uCA. ^.. .c_. ..^, a .iii...j^^ ...kj.ta, -i i::ipr«»»

•iva at firat iîaixc. , and for tiiat tiae, ia not aaaquata

for detailed variational atudiaa irbon reduced ^„vidlii^

it among 12 separate species* This ia ^^rLi.cuiaxu.y true

when one coriaidera tiiat tiiere actually are aix)ut two dosen

s^-ocAos of IfttttH fff i rrf*- ^^^ ^^ wiiich woare not recoîaed by

iiutnven although hs apparently had % eciiu&jio of all uot

five or td.::. As adt!ili.oiial mater ial accairoljtuu in .'Unerlcaa

museums duriui, tiw first i*alf of tiiQ ûtn cosiûr^', >^^c

refined i^ oi êotyraphic variation became poa:iiulo

for many spocies (^'it;cn, 19/*0j Mittianan, 1949; A. G,

omith, i9V>; ail;.! t.xx. , Xj^'^^ to aoatioa oaiy w .j ^K;t

ooaprehttzusivo studiGs}* lor sociO uiikooim reason, tine

5aurltu8 Group ims been overiookod in this rocoot £lurry o£

activity by students o£ ôrtorsxiako variutioii, exco^t lor

a foM brioT aiid scatterud notaa Uulad in papers oa otuer

subjects* Xiiia ânerai lack oi intuiv,.,v^ in tiio ribbon-

anakea ia eopacioiiy aurprisin^ wiiau ono coxuiidar^ um

extensive r,ooA;x'V-^-i-c ran^.c of Uxq i»roiip ihova -.cotia to

ti.c - - V. xairuj to '.oûa *kAca) and u.

,

..J aaiount oi"

Ui 'VaiXabie for study (loore than «: oclmons),

two facts which would appear to indicate t' "t tr-/5 _ai.ritu3

Viiroup would bo a 4i0oc oubjoct for a dotai^cv ^w.^.y oi

variation, bot)i j,oOi^r -r^hic «ind individual.

î^-V'^;^ <j* u..x^ investigation is t.-oToic : to

prov. aato definition of tlie âuritus uroup with

an eyo to usia it as a aodei on *4".ich to base cc -iva

especially appropriato inaauuch as !• aauritua ia Uia tyije

^^.uv^Qtt) and to (snalyse individual and tâo/'^rapiiic variation

in oach of tiie two Oiôcics boionînc^ to tlê t^roui^.

Owine to liinitations o£ tL.iO aiid to other i>rac'w^c i

considerations, no atter.pt wruj :aa(!e to oxaraino ever^.

ci-

raexi ox ^jifffr iPM-^^ m-^'- ^ -• ..^^QJOfflUff i>re»eiit in i*orth

junorican collactiono. "Man ^00 ap&ciimm Sroiu all

parts of the ran; en of both ar>ecie8 w«r© studied, .8

nuat>er is fiuiiiCiesiu, ^ cQiêve, uo deiiiii^w a<. v^^.i- -woiy tiie

variation occurriiit, in taia 4,roup of snakes*

In addition to tn© .raanrvod material, i have b««a

fortunate in havinji live rxui^). •/ r^'^- '^

the Onitod ;>t?.tQ8 and i*iexioo ax for e,^:araii -,

ijiciuaiiie AxvAiJii reproaentativea of all of t.^*e tsubSi^QCies

recognised in this study, îtijout theoe animals, sovoral

diatii^ctive and diarnostic color-oattem charactiirs wold

have ijone uxinotice \ i. eaaii. tiuauor of articuiatod and

disarticulated tons were prepared for coaparative

purposes, and the left caadlla was removed fron a ntmbcr of

the preserved spocinens for tiie purpose of . -i

counts, bex was detorTJinet'. by dissection, u for tiiose

i,.ai.uii ua*c.i :.. '.-. tuo hoiaii><,i-^ -r:rted.

foil i~ôa«i tt; of coi/aaiic vjCtJ.o6 were

Blade: ;.Arir>.tai. icnbtu, - î ieii^.th ritid tvidth, rauzaie

lenÛi, iiiti; uj^rostrax wuou. - sj.,, a»i.u iicifsal-rostral

sutum iexie.^^^* i'uczxo xctô^î f* ua4»d uvroiii, equals tim

oo«)»iao&aJui tiio ToXlowiii^ ratios: rroiitoi.

langtii/pariGtal iont^tli; frontal iîdth/froz*al Ic^th;

musslo 4.V':., ^ -/*roataA lexn^tii; iuu«^.- .^ o; -..-.i\j.,ui'«j. coatact/

naaal.roctral contact. iÎoaaureaents \mro also laade or head

length, body Icn^^th (iocludin^ the tiead), atid tall loii^th.

Hes^ loiit^th/Xiody Ioiv.,th and tail icixi^.t a/total iei\^tii ratios

were calculatoc-, i-Umy a^jociioens lack tails or are badly

diStOXiôdj a;jucu unia uuabor for liliilcil taxx 4.u.i^tu/total

length values could be obtaiaeci is auâll* .^ few of Ute

ratios (frojital iGi;^-,th/parietal loa-tti a;id froatal width/

i.roatai. isuttÛii aaovwx oo aitîiXi icoat varxatJioii ixuc w^JJL

not be dxacuased furtaar*

citaiidard counts tiere made of such faeristic cxiaracters

as nuMber ol au^ra* and infralabiolai pre- anci ;^.o3tucularS|

teaporala, interôxiials, vontrala, and aubcaudals, aa well

aa of any anomalies occurriix^ in thoae caax^ctors* l\ic

ouiBber of dorsal scale rows was detennlnod at t.^ree |X>lnts

on tno boOyt one head lon^^th boiiind the head, nidbodyi and

above Uie xast Vttiiti*ax scute* iaa ^.o^ ... o* cu^.. .cio

row reductioi^ ou the body was detenained for a a^âli nwabor

of 8peciia«M!Ui f and caudodorsal reduction iii a few

individualo.

The number of teeth on eseh of tlie dentiêixûii bones

was tiotod (counts were oade only on the loft side in any

one indlviduul) » aapty sockets bein^ includad in the total

count*

Xnasiauch as certain olonenta of pattom vary sitîri-

cantly In the <uauritu3 Group, Um co.;citi^n oX' a nt^nber of

wiQ0O •icoian'ta was observod in oach specioea. *.ij^Q

IncludGu relative aiae mud aiiape o£ t/te IXt^xt-colorod

parietal spotn, aîtGnt oi' tha 11 ht crcot on tiio .ostocuiars,

proaexice or aoa4j*ico ol a hw»c^ ôutoiUî^Mi. vatt», wxuta ojl

the vertebral stripe, prciDoace or ^cc o£ a b4.ack

paravertebral stripe, width o£ tne li^ît lateral stripe,

de^r<:;o ol 0pottii^> oetwufû taci Viî't<;$bral aiid lateral stripes,

aiid Width of tiie dark ventrolateral stripe occa^.yiici«j Uie

lateral fiiar(îi oC Uie veii&ral acutea on eacti side.

Color liotes on livii:^ apecicieaa were recordod* In

this ro, ard coasistont attention was given tho labiaio,

vci'ucurai atid lateral atrxpo;^, tau uorsuui, <ixx; Ui<d voiitc^".

In Qost caaee tneae evaluatioaa were subjective, but, i»ith

only two exceptions, at lonst one sijecioon oi' each

£;eOt^raphic race 'mfaa comijared with a color atlas Uâors and

Faul, 1950).

A.

4

u.^.iJ-TUî CauUP

it> has racontiy boen shotm that %• oro-xiauflf

Xozi^;

thoujit to bo a subspecios of aaurltua , is a valid spodes

(iiiosOMnf ^'i^)« 7a<J .âiu'ûua ^roup^ mica xs iiov; coUiAQ^

erod to bo ooiaîosod of two closaXy roXatod v ;, j^,

q|fliiprf.tufi (-Înnaeus) vi;ic ^, , ffl i^flm (ây)» has lori^ boon

recot^nizfiKl as bolOt? noro or ioao dlstlrjct rxx>a tiu) oU;or

i;artersuake spociea* iiolbrook il^'i) placua the 8|.ecieo

aauritua (includini;, but not rocoi;nizi;i«^.« rffiiitoftiff^ ^^ -:^, a ênus q£ sXe.idor, suiiiiaz'boA>al colubrida, bub

he does not include tae otiior t^JiJ^-^^'î^'OS, i"itziiv.er {la43)

rxbixniaiXiLa.c f ol. .; all tÔiôrcM^v^.u^ wûwu.aaiXy .iavo boon

pi.-scari ill thia jôi* „ ic-itar aothoro, .Av:.» ns iiiU.- .g

16;«>^f Luu«v>rxX uiid iiocouTw ^«ciud^ only .sfc-^y/^^;... ^.

i'iiai.u ., tiiu otiior opoclos boin^ ,.l..cod in _ . ,.

4iuthven (X90^) doo3 not pXaco flaurlt.ua in A aopcurato &enua,

out acknotoXoduoa its dietixictivo nature by aakin^^ it Uio

onXy apocies to coustXtute an ontire epecioe ^^rox^p in liis

phyXoi&onotic scietuo*

Tneae are mokes oX* Ui» ^<boub "^j^MMMifi posseeeioi*

ouLpenls tihich ejttooda to the eaveath or oi^^'^^

subcaudai in U» iaverted ijositioti. Ihq mmlMV ol iaa-aiiary

toGth ranges froo d7 to 34 l^'iodo 30j. -.e basaJL iX>rtiou o£

Uid to«t.ue iS rea» tuo i.a*Kuu wips black. Tiie tail ie

long, usually couatitutint, 27 percent or «ore of tiio total

length in adults* The laastlrman number of dorsal scale rowa

rarely ejôeeds 19. aj^© vtwitrais raiijê xroa iu*i i*o i«51 (both

extremes In 1, proxliaua ) , the aubcaudals froa ti2 11.

,iyr?]^,\ff^Hff^ to 136 (1. ftffaritua ). in both apecios the juaiee

iiave a ali^^tly greater number than the resales. Xae

lateral stride ia usually corjfined to dorsal acalo rows 3

and 4. tm aui^raxau. : .vertical black bare.

fonu aiid iroportiona

both apecieo ax'u reiatxvei/ slender, lo*i,ia-taiied

snakes when eotti>cu*ed \iix,ii otaer niiesabera of %ii& ônus.

XfaacanoDhifl rfroximyiS iaay oxcood liiJO uâ. in total ic^.o^x*,

bein;' equaled aoong Uie ^ârtersnmes by 'i'itai>.ao;,-iia.s •-^yîf^-fAfi

^'} t*it; . Ai adult exatiiined measure© only

slightly over one ...uwur, l':â roiatlvoly narrow nead uiakes

up 3.7 to 4.6 (ffloan 4«ii ^tiicoiit oi tiio ijouy io.*,,ui in

atUUts. i. e ffiuaaio ie riarrov., tuo interaasal-roatrol

co.iuî;u/:.-ii.ai-rootral contact ratio exceedirit^ l.CX) in only

one of 29 a'julta QXanino^ Ibr tids cnrxacter (laean 0*6:^ )•

iiiis oarrouacas a- „ .vOXViiwo./ ruxxocwJ. w —aquatic imbito of thoso t. cios (i itci, XjttOi - .

,

i960).

Tub tali la ioitc.sx' Utaxt in any oti^r a^>uclQ8 of

'^t^MinflRtoB ^*^ coastitutcs 25«2 to 33*6 ârcoat of Uio

total length ^> .^Jult !• T?i^li^^\fi and 23.8 to 3«^«^ ^.orcoxft

in adult ^* aauritus . .a in pro,>ortio.âto

tail ier^tn la ovldoat iii botn spoclea,

uatsolo^

vert4ibral coluiai^ &b la CiioractorlsLlc ol natrxcliie si^Ktâ*

(HaXoatd« l;^6i, however , .âu doaorlboci a uow ^ciiua v»iiicJ:i la

apparcuitly cXotKdy reiat^jd to tiie natrxcliio «ônu8

lIlfflftlfiifgWMIr ^^^ lucks uy^;a;^4.4iydau on tiio poat-thoraclc

VGjrtobrae)* The stcull Is similar wu u.iat of ot^êr a^^jclea

or ti\o ôau^« tortalii (!liTerenco8 do OJU-st^ but t.iOLio con

best be analyzed by a coraL;£tX^tlve ntudy ol' Uxo aliuiXs of

ax.. .......... --.I " tiiaU -.: I^.•Li.w.. l>'-.:jOilC Xt.i9

scope of tijiO ;,ro8ont liivoutitâtion* iiOiubora ol tiie

;>aurltu8 Group have UiO ^reatoat nuober of teeth on oaeh

too tii-bear1(^, bone ol* any sôclea. in the i_,ci»iii lor wl^ch

data are available, lao .. . oi rtaxlllary teeUt i

ftoci <id7 to 3<^» ^;.A^ûli«e teetii aa<j>-. ^.y ^w •^^^ ^wu-^^jOld

teeth Trou <^9 to 30, aiid tl(jiitar> tootii Iroc; 33 to 3o«

I'ablo 1 aiiova tue lre<ûeiicy uiotrxbutloa ol iiiaxllXary tooth

.xillory dentition in

Species

10

couats (too lev. count o are aâlioblo for t>» oti^cr boaea

for a froqueney distribution table tu aivj imy si^nlTi.

cance). :o 8«xual diaori^hiaa oay b« present, the ciales

having a sliiJitly ^,rcator niosber of naxiXlary tootii on the

-;v::i-:ioa» '^--^ r.;.i.J.ilar/ uoeta - I: " .'oiy in

leQt,th ôstoriorly, but tbe Xaot tiiroo tneth are Lly

enlarised*

iioiaipenie

lue hamx^^nee oi. i,« (iFiTîlliiim axid i,« aauritua ara

virtually iaoiitical lu all roapocta aiîd are tau snalleat

tijaomx lor tiie ônus* ror taany spocloa ioiorsaatlou » nuX,

available* ~ j lavarbed or^jon uaually oxtouds to tlxe

eiîth subcaudni, rrcquontiy to tho sovonth or to tho

aixta, axMi xa oiU.j oao aj;a&ai;co to Ui^ i;iul;i uaoxo «;^«

Fet^ra (lv60) rK>tod a negative coxrolation bet.voon relative

tail leu^^ aiu) aeciipenia longih in cartain dipaadine

onakosy and it rs tiiat tiô aaae phanosionon occurs in

TAtiui. ^..Miiêaipenia lon^tli iii tno t>auritua uroup

iJLôcics

il

Tho aoat distJUictive -^ '-i^ t.i*c scuiiej-iui-io:* of

the Snivrituo Group is titt) îly •loi't.o nuaiber of .ub-

civ.c. ..- iii both ^,;uv-Qa. Tiieu*J raiiee trom a lainicium of 32

iu X« ii Hi^'^

i^M ^ ^ uiiijJimm oi' 136 in 1. {,;aurivua. i...i»..o3t

always exceeding 100 in T. aâritua suid leîs fi îy

species in too tênus nav« aorc than 100 subcÔal.., ..^lid

uaually tha maabar is ouch loos, Utiter icaijortiuit îC'kio

eaaractara inciudo: a Gu-itâ ^jreocui-ar; '- û^.i-â.uua^^q

anterifig tha orbit; a ;.a--aiaua ol 19 cioraoi acaio rows in

96,o porcont o£ tiio X, pyQ3i.igmfl and 99.3 percent of tho i.,

aauritufl exaalnati; ^dl cJoraal acaios stroisgly Iwciviu, Ui 30

of the outeraost row not t.rQatly ealart;,»c#

Coloration

Botr. 1. Timifii^mil and !• iîj,^:^^^^^ x^t;- vuiniical black

bars on ti» labials and black Qcirkiiit^s oa tiw venter,

altiiontih in all I. aaiyitua and in certain populittiona of

T. n IVfitolMt thora ia an e-ctensioa of brown yi^>u-io o..uO tuo

lateral aarsins of Uio v»îtral acutea to foro a dark ventro-

J..O , .. /.r^. ,. - lateral otripe ia aiwayo present on

....... scale rowa 3 and k (coriiined to row 3 iôaterioA^ly in

oi.c i.xj of oroximua and includes row 2 autoriorly in

northem populations of ûrjûaj. but Um vertebral stripe

ia frequently lacking, in eons areas. > ii lateral atrip© ia

aaually a anado of yello\»> white, or blue, the vertebral

12

stripe & shade of yuJUo.., oraiiô, rtiC, or broxn* Tiie color

betwaeu tae sti'l^x^s i-ixiitân Iro... olivo«4^*^y jUid lif^^t brown

to bl6ick, liioro is a broi^d, vortical yellow bar on each

X* nrir^toHB îsd X« aaurituf fona a closely kxiit

species i:;roup wiiich does a^-^ ^îO>i particularly cloao offinl*

ties to any o£ tiie otlier specioa conijlojuis iii tiio iôuxxB^

althouiJti Uutiwon (190i5) out:^*3st3 a reiaLionaiiip to tiie

possibility CiUi be ^roporly evaiuatt^d oiUy witiiia Oae Xr«i«.io«

M}rk ojC a a>j.i; irohenKivc 5t/Uciy ol ti*o intc:rL.:-ociiic

roi.a(«xoauiUp& oX' tiMi (<«^*wa.A\; o^^'^Liu* t-oi' Uiu ^.'

. ..^ . CoUi

say u«at the relatively lou^, taix^ extreme roOuction in

spotting or Uio cioiû..., .^â transitiottsl syncroiitorian

najcillaj:'y deritition^ iiMiicato tiat tho .^-a o£ tao

îaurituo urou; are lii^*ly apecialissecJ, and that tli-e i^roup

xutioio. occu^x&d an odYsncMda tiVQûuxoiiaxY posiuioiA k..j.uhin

ths genus TtMBBflntt^B- .^^ mimbor of tooth ;}rostmt

A» sauritus and i^roxljatts is n<:.t horo int nr rtcd ac being

a priaitivs couciitiouf but; ratiior a seco^jiia*'/ tuou««xcu'wion«

^Transitional e:'"""" * '"'.'. -â equ^^'Tiiit to the "coutinuous-abrupt" arrange' - - cuorisod . tnate (1^00) and

tiM>u4;ht to ropx*a8ent au evolutioîary traiisition betwoen the;orali8ed syucrantoriau tutd tiê aore specxtLUaod:.nrian conditions.

i3

, t^y to tn« &p««i«s

i»ari©tal spots edmost always present, fused, britsat

sud usually iairiy iar4,6 lii^ûio i.., ieit);

browii pi^snt usually not e-;uo^.c into onto

ventral seutss to fom a dark ventrolateral

stripe on each side or, if present, cwvorlofi

Xsss than 2/5 of tiie area of each scuts

(except in tao oiu^tpas iii,>* J; aean

aussle length/frontal len-th for any given

population usually greater than 0»70 .,,.,.* 9r<?,i^m <i" ^^^

Parietal spots frequer.tly lacking, when prcj.oiuiit

tuii^ iixtj uaoll and rarely fusea or bric^*^

(jri^ûre 1A, riiît); brown pl^nent always

extending; onto ventral scutos and usually

coveriii^ 2/5 or rjr>rQ ox tas arsa of e&th

scute; osan aussls len^th/irwital laui^th

usually lesa than 0.70 ...... jaftttTJitttg d • 9^)

tli^mre X.^^Head . as in %i\9 two Sf^ecies of tl»;>auritu£) uroup in dorsa^. v^ow {a) and lateral view {a).

15

< cr

<CO

< CD

COZ)

ocra.

yar»?&l.agegtf^ffi mi^^mA ^^®$ ^^^'7i ^35

Miflilftail iJftiUsadk îrd & Clrard, 1853: 2^

^tflfflinîiii .ravj^ra,,^ ô^^^, ^.^ji 3^0

fÛt^4flii>nA ^•Uv.^J^rua ôuu-a.., ^;ôx; i^^

Hccordlni^ to uîUi and Xayior (1^45) tiio jniolobyp* is

lost, . Uo not knoA^ Iti wiiica ii\stitutioa it iiad been

de|X>;sjktocl* i.t Mad colioctod Jui aoux-aska at a atoiis cûarry

on tiie >fe0t aide of the iUaaouri aUvox*, 3 iailoj ôovu tlM

aiouta uj. ^'oyer*& tio-vui' (lov«a) by iA>u^*si o^v4>edition to tiio

iU>OKy .uûntaina eitlier in X^lj or 1^20* In tems of

pronontly existinr to\4na, ttio ty ;e iocnlity lios approxl-

ta»wtjxy 3 ailea îi<:^ ioxx ciU..;i>tAa, .<oa usuiity, licur

A large f lon^tailc\: uo^^bor of tlie oonus ThUWTf^rtfVfif

16

17

characterised laiyt JL9*X>-X7 ciordal scale rows; a ainê

praocuJUtr; Vplcaliy S suprai.abi.ai.09 tae TourUi suul iiXtli

eiiiteriuij^ HiiQ orbit; 141 to 16i vaiitralsj 00^ to idI £}ub»

caudais; iateral stripo 011 6or9aX scale rows 3 and 4» at

least anteriorly I labials mid ventrala witiiout black

markin;;s; dark voatrolateral stripe absent or narro^f in

most. j[>o«>uiat/ions; parietal spots fused » briîitx^ culored|

and usually lari^c; hemipenis stiort, usually extending to

the seventh or eighth aubeaudal vibexi inverted} laaxillary

teeth misaorous, usually 30 to a side*

The species is distributed from iM^tham Vîscosîii^

Indii^.:u, and thw *.^!3iim3ippi Vi...._ j west tiirough the Ureat

Plains to soutlioastera Color > o uad eastern lieu i.e .ico^^

south throui^h Gnstam MoicLco to ccatrai Casta jiica (Fiiûres

^ Anu Jim it .. ...b a<>« u«k; Aaciu.4.c l. ~~co

near Ac&pulco, uuerr^ro, and on tiie Istiitaus o£ ieiiuaxitopec

in Oaxaca. A western ribbonsoake {iÛHi' 6179) collected in

tiiC Iduvj'j and label!od oiuciply iîiimesota needs to be

authouticaued by i'lu^taor cx)llcctini^« waeeler (1:^47) records

several speciraens m^ j^» .tqxI^w^ £roai uoiw.i-caatral iiorth

Dakota. Thoaias Ussell has exanined ti:iese speciaotis (uiuxS

74337, 74339-342) at lay request and reports that tiioy are

actually l^mMUQM.^ £Ma^ *---' -,

-•-utAy no valid

records or yrojoau^ in the Croat rlaii^s north of ihux'ston

County, £icbx^skA«

Fl^^uro 2,^-,)

*'i^ur« 3.—J-istribution o£ TJjMexico and Coatral Aj^ierlca, JoiicI ck)

rspres—, -8Ub8p«Ci00«

--> specimensaroas

2Z

Distinct sexual difforGncoa oxist in nuobcr of

ventraie, numbor ol oubcauUaiay and reiativo tail 1 .

7h« anotuit or diuorpliiaia in each oi theso characters varxos

from one local population to tho next, and thorct a^peîro to

b« no .^c corralation.

The diTforerice in ooan ombcr oi' voiitraia ranoos froa

0*3 or a acaô jû jLavar o* i#iMi SLmmaMn^ î b.. . J. Troxa

the ^takad Plaiiâ ot weatorn Toxaa (tiia on^^ lo \iith at

lanst rivo indi^'irû^.l£» of oaCii aex in wliich foiaales Iiave

^ ,.i »^w»iiarv©il W udVo ooro - la tnan tha ..;aiaa), to

laox'a than 5 in Tavor o* tUe -. in cortoiu ouuor arc i,

Tlioso jx>pulationo frwa viliic .» û û...., 15 adulta of oach aox

are available are e. J in iuUle 3. xo is a nctico*

able lack of c©o, . c: vari-itioa in tiie dOoiô of

couori^. ;_.;.- .-- •..:..-.. ....--L, ^y ;^,.j coeiiicient Oi i'-v,--v, -..co.

Furt /be notod by roferrinii to iabio 3 in

Klauber (1943) txiat tlio degree of dlsiorphiam exhibited in

this Ciuuractor by iiMabera of tho oauritua Oroup is ali^;it

in coapariaon to aost of tiie other forris for whicli couâi*.

able data are avaxwu^î. 'Ûii'llfflfiilifclf"*^3c^>yi shows o^^ter

sexual diiQorpaiaa in miabor of voatrols tiion doos Tna. ^

ordinoideaf less dinorphian tlian 1* £îi^ and £• JSJLMftOI*

and about txio cuue <4iiouiit as iiligUUftiiAJUft iiifiiSiAAJUStfltMl ^^^ X*

aauritua (I'ablo 4)*

^^4

TAbUu 4«—Coorxiclontso-

or^enco In six ai^cies

ocieo

25

îoatidiat ii,ra«t«r eoxoal differoncea exist In mean

nuaber of subcaudaXs* TUeew siean diJlTerdaces rai\,8 Uroa

a«d iii Tavor of tlia ftfluilas in ti^ GaiTe&con buy^ 'i'v;.>UiO»

aaai^le (tue otu.y oaa in whicii thd aa^^dS liAVtt l^«iMir •ul>«>

caudaXs tfian Uw jTaaaltta) to a« suon as X%2 in favor of

tho mal0a» Table 5 presents a soaparison of Uioao popxiia*

tions from Miiicii at least 1^ adults oi aacu @€U. are

available, i'he anount of sexual diaorphiaai in subcaudaX

numbers is not great eituer in this ai>ecie8 or in X*

aauritus tên tticir respective coefficients of sexual

divorgenea are eompared with those c£ n!»iy otiier colubrid

snakes for teiicr* o..> .a ore avax-taoxo i&laiôr, 1943;

Auffenbert,, 1955; Duollman, 1958; Dowlint,, 196Q; and

others), Tne coefficients are even leas, in fact than in

otiisr species of lijaâtiot^hifl (fable 4) widch is in keei.ijr*4^

witi2 the hyi>othe8i8 that sexual distor^^hisu in oubcaudals

tends to be t^reator in snoru.taileci tnan in lon4>tailed

species (iUauberi 1943)*

Fetaales aversiê longer tiian raales but usually have a

proi-ortioacilly sxigrter tail* tiiu xixriuGnii female examined

Utiiitiri 761 frora Cafaeron Couut/f texas) nas a body le»4,th of

900 ma* plus a 33ii faa, iacoraplete tail, Tne iarr,cat ciale

{\ik^ IZloimk frota iit. Gijiarl«a iaricwi, -ou-;: - }

Iwoi^th of 553 isv»* and an iiicoaplete tail* Wright and thrift

(1957) report aniaala with greater total leni^ths (126^

fetaale, 1119 am* mal«), but i]o not îve locality c!uca«

AdtUtn of both sexes average well below tiiese oaxljaa.

26

£5

t

135

IQO

'O to• •.> o

«

o3

<nI ^

rtj sO<~4

The isean diiicx-Qiigg xit acoiv^ Caix ji<?u^t«h/total lea^^tih

ratio ranges from 0*CX>^ in favor of tlie feaalts (UaXvostou

liAy, Texaa, bein^, tha only area v;;i©rr! trie ratio for f(&isuL»a

has )>eea obsorved to bo i- u.iaa Uia^ ^.or , .0 U»022

in favor of Uio maX9S« iîsan WiX iî^tii/totai .>

for maiea ran^ê froa 0«iJ75 to 0.3i^» for females from 0»<i70

to U»3i3« -ts axi^rbaued by tuo -uiiixc .lo:v>ii/rroataL iert^^th

ratio 9 fttaaXdd a^^<>ar&utiy iâvd siifîtXy ioocôr nuzzifjo titan

do aaioo (e«s ^« 51)*

Caudodorsai scaio roti reduction rsfieets se^ûai.

dinoi':>hxsia, aalos having; the moro posterior roc actio::, iiiis

j. . .urui.jr a coiisGûeace oi w*q pros«nco 01 t.;o

iid their retractor muacios ia the lal part

of tiio tail. The roduction fonmlas for C-a 3a>^^, .-. ^ uit

fflai(3 irom CoBaa«ê County » Uk Ciw ^Q^^^ ^' ^

foniaio froG! 4>t, Ciiariea County, ^ri, are prosciitod

bsiow as ejw4Ut.j.o3, The fonauiaa iâu axit,uux^ aiOdirio«3 froa

tlie ayotom proposed by Uo)«liii^ (l95i) in t^^t Uie figures

above the line raprasent tt« left sdde of the anidal,

C^ 3Q9i 17;-''^'

i5=^Ûl U*^^ ^^^^§{7}c/*

^^^S4Hv} ^î ^""'} ^r!Hiiîu3)

CAS 5099 1'7«'!444 XSrM'^l I3.r'/U.i x^tsstÛI KpkiMi

?«4Lm 6=#^ 4s#^ ^(99)-•iv3^> -*^lu,) -î^'u)

23

Ont' ic Variation

hov/ly torti ltoxIpu^ have a nroportionaliy aiiort.or

Quot be ra^.-id duriii«^ tue Tirat josu' iuaaaucii «: .its

have eosent!ally tiio sauo tali Iwii^ta/tot'l ian ^h vrj...na

as aduito or tiic aa.iO ôi-uxation,

indivictual Variauioa

-Ch iioitiiyr vary j_,uc3it^rw^4iicaiJLy rtor ohow

soiauiJ. or o.iwUo-i- .wuxauicyii aro diacuasod below. Data

on head scales reior to ono aide oni/» Individual variation

in raeriotic cli.-^'nrr.^rr: is GiiTT :,-\rir^r5f' in Tabic ?•

Oculars

Tiiere is ociiiar a»ci usuaii/ 3 poatocuiars, Viio

raoat fre luont variation is the appoai*anc« of a fourth ^8t-

ocLLLfciT uii uiit. ov u'-'o.. i.j.u'j •, ui i4^v> i, i:V7'rtitffv^ft» o^^y oo<*

iiidividuai has an extra prooc\il<J* and ti*at on on

it ifl iatom-Liii to note tiie ntabiiity oi' ttiia ^

in t»»© waux'xtus wi-'oa^x x;* vxcw oi its c.^w*" ;aA-iviwxxiuy -*i

a nuiiibor of ti^ otitar s^êcios oi , : liiutuvcn, l^Joî

Fitch, 1940).

) anterior tc%oral ant; «: alioiitly aaaixc*'

postorior t' xa is Uie uoual coaritiou, but t»iOro is

j^reater iiicxviuual vai'iui.ioii jji u.âu Ci.amctor than iî L**»y

otiMr, rauat i'r -iy an iiicrcaso in number. Aitiioujti tiiS

29

a.

gO&

%

S-t

3

S"*

cs

^

1

•A

*4-

•

C8

C3

O3B

O

--N vO O»r* O »• • fO %0 -T

?i• • •

•n

H

*>•H

«

3

o

^

3 ^ :iC^ c^ r<\

'jD O w^

r^ r^ r>• • •

^ ON

I3

n

c?

9CI

a

I

^

o

.a

oHW

o

8

I d

30

oo

a

o

>

I

I

31

the oecurrenee o£ Z anterior tostporals is coearaon, onXy oom

o£ tiiom^ the iov.oi'y is over ia contact wxta t.^c poutocuiars*

in adcitioii. to wH iz.ci'caoc in the iiuc:.bGr o£ liCUi-aCf Vim

aj.iwâ'Xji" |/0<^'t<u;;u u* t.;iC: ai«t<isrxor ucu^cr^l a;« jc.CuU«i.o*M4^xy

fra^(î,tcd to lorta 1 or ^ siaall scales Wiici« are raruiy much

Xargex' tiuui tiio ; ootoeuX&r acîleti* Iherc- appears to be a

liiiêx* iucidence oX' this variacXon Xa the CUX&|>{is ai(îU.aud8

than ill any oUier arsa {l^Z .ûrceat of t^^ animaXe examined

huvu W4iia i i'a.jaeutation of tiie anterior tâpor^^ ua. oiie or

both sides oX Uê ueac}.

Eii^^ht is the uauaX nti^>er, the fourth and fii'th enter.

inî Wie orbit* êvea apîaroatXy occura by Xos3 o£ a pre-

orbitaX aupralabiaX, 9 by addition oi* X posterior to the

orbit. XiifrequentXy tiîero are &&suX acaXes wed^êd betufeen Z

oX the tmn^taX scales either at their up^êr or Xowar loareîae*

itareXy aa iiidividuaX eupraXabiaX uiXX be p>artiaXXy dividecl by

XnXraXabiaXa

i'en X& tl» ufiuaX hui<iber» 9 ^^ iX beXii^ eûaXXy iiomuion

as variations* liiiie usuaXXy occurs as ti>e rtêuXt of tiie

eXXittination oX the auture betmwtm tiie tîird and Xoijrth or ths

•ifillth and ninth Inl'raXabiaXa, in that order oX Xro^veacy.

£XeYetl usuaXXy reaulta Xron a u^vision oX Ujo tiiird saura-

XabiaX* kQ in the s%raiabXaX6, soiaXX an<xaaXoud scaXes ar«

occasionaXXy iiiterpoaed betvteen norsMtX ones*

32

'I'hero art, ua ar. . In . ur

rather thar. all btii;i,: in a lis:eai* scries of ciri^CG, - -.-ty

iiavo ffloro ttiaa 2 ii^t -ôlSy .^n uauûaily xiiîi ix:«. o*

te i frr,êiifcatxoiij over 7C> ^.îcc..^ oX Uic acriio

e.. :... . J one or both oT tiiose u^ .ôs*

Anoo-a.ouw c;,xv-îGii» or j-fctru^.x ?. u^vXicatloafi ol ventral

scutes, iait-oiy couaod by jjartiai duplication of vortobrae,

occur la many t^roups :ir nnakea (În* , 1959), and U:e ^.nL.rituB

Gxûp is no e.vCCvtioii* '^11 of tho tyt.oc ...-*.sâ«i-c.

i'etars (iy6o) in hia atuiy of tiiO eubra;riily Ui. .a© occur

in both Rroxiauo aid tûritua . trie ;no8t cocBaoa tyr<j beiiii;

one in wtiicn 1 or -.norQ aiKxttolous partial scutos aro ia6oxn.o<l

between the tsrwixiui voatral and ttia mal piato, Xiu-a tyi-O

occiu*£i Mi j^^ percent of tne uroximua exatainod in ttiio

study, Hii additional i+.3 i.ârceat I including, a lov» of ttiose

just :-jutio:icd) navo uartial scutes st some otnor ix>iat on

the v«..tyr, -aituii \LJ:fo^ i-Oiorta li.iuu xy»j j^.oA'cuui. uf U»

8. ao of i., yr ^c://«^tQi]^ whica he ciwwiaad have

ventral or subcaudoX cuiotualioa.

:53

Of th« ixiw ividuaio q:-. , *:.3 poiXi-ut iic^ve 1 or

oojre acts ol fwced subcuudals I i.e., ->.oro ia uo io:ioituGi-

nal suture separatirig the 2 adjacent laasobei'C o£ n subcaudal

pair)* bartoii's (i9!>6) ca-^.-uts on Uiis ooncitiou ii* j;..

byacfhv&to^ arc zu:.t ;cl above,

OsiXy 4 percofit o* t*iC .-.....:< -j:; o.,_....va-,itit,. uiÂCi- . Ui©

typical ioftiiuia or i9-i'>i7. ^* iax-bO aygority of txi© iiicii-

viciuais iri&vir% aaore tiian 19 rous u- any iXjJuit, on tlva body

arc i'roia tiio daita roiîoa oi txja ki.aaissii>i>i iixvar. i.ioy

raaKe up only « ajnaii part ol taat iXJi^îiatimi, uo^QVor, ia

coaw4-iu\* wu ti« ixî^ -4.ation In the viciriity of Ori«-;ibG,

Veracruz^ in wiiicii 3i i^rcGnt oi tiie spocitîQiis c.^^î .-..ud itave

rodvictions in tho coraal seaXo K>vf jTonauia li.o,, iV-io-i?,

uion iTi the auaber oi* dorsoX scald roâ is, in

all .-wO, x*^^ W.J. u V- -...> ui>aciaoiiS c.iecked for this cix^jractor,

tlie result of loot; oi' row 4 on both sides of tiio b<;>, . v,i

the four exc^f^ticw.s , o:';0 lost row 4 on osm side, 4 or 5

on both ti-»c e.ij oiie loot row 4 on one uix. e>, 5 on tiie oUiarj

and one io»t n-w 5 on both nic'ss,

iuaon^ attxw:i tie i'tiu«cu.w>w t»j-43;i jîiiCc •:._— '>^^. vi- iiWals

a^ tiirouî 1U5, ueuii 93»iJj a*aoa<^ feiuaiea froci vaatrai '/d

2U

turou^..!* iJ7, inuLu 30«iJ, - .^..rc ii,.|^©arj to bo a poaitivo

corroiiitio i.ftt.'Yr.n tilio total wuoibor of v .^.tir-ir? r-^ tr*e

poiitci'lor Uito ivt'uc^loii* Co ^^y^ ^'-^ >>oi.i's/ oX r«,c.w;c«

tioii will oi .i^v.4i.>u*»^j . , _^„*v>^.».a.,—*^ WW i ^cz'taia

citi^rue* XL aliio . sac ^*t; oi' aoA:Mal c-i^cxv

pliia:^ on tiie c .îr jiiicc iiaio3 . . / aavo

îso Oi.^ tiitt ^riotal sôt^s varies iia(iivlUuaiiy« mid

•ome iiidividualo uavo otreakij of iiiît piônt cxtoiKilajl

antariori/ i'xxjixi tae uai aix)ti£j aiaii,, tao i"ro-*U-ij.-

parie\;rti & cuTi aiid occasionally onto ta© Trontai. itaeiX.

>>*ujj or ovua î-'wûiiCU, u*. u*.*; ^.<5t»uoc^ju-ai 4-*^.it Spot »ind

tho black poot^rtirbital vitta v«a*i«3 iMdividually as ;«li as

fiooâivhirnlly* - i'-c.acc or absanco of a row of opots above

tlM lateral uUrlpo ox- ol a blac<{. -i ;xi.oii<, tac vcr^jurai

strips also varioa from individual to iijdividual, — itivo

darkness of the dorsal ^^round color frequently variea in a

oiAilar manher.

Geographic Variation

in an animal tho ran.c of whicli la from soutbem

- it:co..iix»; uo coutrax woîua i»a.ca, a ax^waucu ux auuut 3000

aiiwa, it is not aurpriaiiife to find a coiiai oiit

o£ êotiTapiriic variation, :-.:-ccu.nc t.crc is u lack of

concordance atooii;^ >aoot oX ti.«<a,

xcuxxy vaA'j(j.iit>

«'>

foliowiiH^ discuaiiioa the teiut ioii" is uaeu in a

Ulolo<.,icai aeuiKf and iti JiiotJ syiiOi., ^ .-Uh Uae statistical

te.r«., ^'la..e" or "iocai poiûxatiou"' wuiu oo &wjr^ .......i/

equivalent to "population" aa i am uaiii^ it» ^. tiroui/ of

sui pi*uAOt,icaiiy »iia*^*ii. ^puiatione coiiatltut© a raajor

population, iriiicU is in turn etiuivaient to tiic t.. ^.ic

siibc-ccies* i'.criiJtic c;ic prj ortioaai values (;iac**3Sod

bdXovi ard oa,:o..i ji. ,.---..,-»i.a;<xoii aa:.iirii.ct., iii*.* wiw.^ Xact is to

be inferred '^hoii i-ow actually stated in the taj*. -uch

aataplos w»n of an ad©~^^ta alssa nay reasonably bs ttiq.octed

to repreaent txie popul«*#ju>ii *a îc.* uiay i«9r« co— -^ct,Gc\

Vfntrala

...o Clean ver.trf.l counts are rca:. -/ coJkGtiUit east

of UJn i^..- .......^^^^ .v^wwi, *i.'^.^ wv;ateni Indiana to riorth of

iâKc i onciiartrain iu iouiaiaixa iaeo fi^ûre 4)« -i--'^ .cui.s

for fualGn i'cLii betvxjcn 170,i> and i7X*S> for fojuilos b^t\;oiia

10o»9 aiiU i.u':^»u» .-»o»*ta l- ..dUu oi i-auwc' i wiic.iaitriiiii Xt'i îid

adjacent to tiio i'îowiasii^pi delta retîon tUe nuaber of

Tontrals increaaea alitjiitly, the i oanr. ronrinr irori 171*9 to

174«1 in iaales» and from 16<i.9 to i/i*** -^-i **- • * -^

hi^Tjiest meaaa in both sexes occur near tho asouth of Um

West St, frem the Hiasissippi Valley to %h»

itio Gi- , t::o Goan ntaaber of veiitrals decroaaes

gradually <;iiia Ûi a rttXatiVO-Ly uniAo. .u.i-ou, hnjCQ being

Flgurw k,—"'^''' '• •'''• V-' t-»tJon iii vwatraJL nuoiber of

i^ .4. • -.»*Ok

.. _ -... ^.. i „ , ^ . va lower . i>presoat c^-^ & . x'ar - s, .•.tui a i> inpIao« of ait •' vabons, - ^-^3 taatoo apecino. . J-i t;iC

a ccziiaiii airûiiL ol varicitioii i-ro... oaa population to tue

liowcvoi-, ijiôua uiore tiiuu tiiO e^puCiôu (iA.vurû*cu in t^ât tat

aiitaplii oro&a in ftjMt'«X«)si (ordy 0110 male is iiVi^*.^«e) ia i'roia

7 to V a&iu.oa xc5ij Xfinxu ia j^%iiMij.i»& iix>tu, popuJLu&io^«;i x«)38

than «i> niles distant, it is pei'im^.s ai>jiAi,xca;it Uiat ail

avaii'^biti OTii aw^ôcubutou ta.ta roiict ijrairie aroaa*

jout u, ttio ii.i.0 oraaco Vailuy ta« UiSo,^. vc.tral

coalites u&c. w^ 04;^d,y sxA^^^bxy ibî &ac .^xurra îuuro wiuu^.âX^

but fuxi uxx siiar^'iy ijti tiio caastal plaiu 01' iuiauuiipaa (aot

Figure 5)« Taetx} -.- ... diifarencw of &l;ao8t 10 ventrals in

the ^.uatU9 of £^,.3ilQ5 (about ^ vuntrals in malea) iixxii ttjfi

lower Kio OrruifUi Valley and ti:u>ae froci soutaorn 'ia.aulii>a8,

a diBt(6<..vu jx. i.j,UK.4.M îQr<^ than ZQO mllt;;^, rhoro is

•ssoutially no cîcii^ô in tiie laean for fiaaalos ixid only a

sli.iit decreaoo in that for aiales over the ^00-:nilc ctrctcii

iroa 30ut:tci'a .. ...î-xpua to cc^r^v ^- ci- .: , uuc ;.:i.;,.ijî uae

succeeding, ^ to 7^ ail<js eouta^varu aioiii, Uic couat to Ua

vicinity of Ciuciac Vei^ticruz, tlie >actanu for boUi oexes

decruv^;^^ uy ai;.iOSt 4 scales •

lae uu&bur of vuatrols ia louuat in tne iauaaco-

vu;..^.v.c..v^ lowlands w^xcre the aeau for .aoles io 146.6, for

feiaaleo lVy«:/» l^.^s r. ata a decrease of ^6^

acftles ii- a:id 4jr in fu over tne 30C'-..!:iic di tsmcs

Oaxaca aas courita which inujkCuto tnat ventral nisabur;^ in

a

this rei,ion probably avcra^ê soimûit hii.;her tiian in a< Jft»

cent areas. Fron tiio Tsbanca-Cararêche ii'ea to tr-e r?oit;ii©ra

txp Qi uiQ lacvitaa reuutsiaa Uisro xs a sxji^ît xuc ,

aod aiorit; the iOOO aiiieo of Caribboon coaatland iro:a Coaumel

XsiajQd to Boca ctel -io, Coata Kica, th<» m»an nunbor of

VGntrai'j rc;iiaiiis Qaacatiaiiy conetaat*

The picture is not yet cotspietey howovor^ because tijt«

t»eBt«m ribiwnsnako ia not iimit«d to the coastal plain in

tropical îiaerica, but is also ioiuid in the iiierra i»adro

Oriental at altitudes i^ to 6500 fo©t, in tao Chiapas

ni,c;û.anoa up co /ûu I'eet, a.ô on Ui^ racxxxc axda oi o.iu

Central Aiaerican uplands up to 50U0 foot* Xii9 jnajority of

hifîualtitutlo populations of YftfflimTOtiîtifi fci'n?iiAflHtiffl ^^v<^ "^ ^^^^^^^

number of voiitraXs iîarkoc^ly different from tnat oi tîoxr

t,oo£irajjÂiically adjacoat lowland oounterpartag novôvoi',

ratiior than iiavin^ a decreaseil nuabor as ons might praclict

on tna basis of the to&ipQratureâcutollation stuciies oi iôx

(1946) and ^V)*, uoitioii, a-id Fox {m) on f, jlJLS^kaaa* ^'^^

nunber oi van^raxs increaacs:* i'urt»i»ai\iioVQ^ the aiaount of

incz*oa8« seeeis ro^^hly correlated with tho auôunt of iiicroaoa

in altitude. This seems particularly true of tiirea >o ula.

tious in Houtiwcentral Veracrua, trie a^J^rotLos o£ viîcii ai-a

only 35 airlino ailes apart ^îifi:iich raiiiê in altitude Trom

sea l«jvel to 5500 foot. In the vicinity of Ciudad Veracrus,

i&ales have 153*0 aean ventrals^ females nave 149«3« ^^car

Cordooa (^000*3000 foet) ths ii.e&ns are 5 scales ^^reater in

42

both sexca, in tac ^msxyLo Vtova \Aw Orizaba aroa (4U>X)»

5500 i'iiot) the ;.ioan fox .^<.i...-v, -.o yj^ti,j 1 0CAl4.^J ;,«-»*« w..»u* at

Corcioba; in £o it is ute>ut 2^ scales*

- i'.tion In taa Chia/.-as iii, aiandc has a t,rc5ator

tite dctt'ias ; tiie Central .caii u

pO][)ulation« lloxiKJVor, inaojr.ucii as tho tvio arc vddely

r -lyt Tactors other tlum Oi-tituc'c uay b«

Involved* On the other hanc^, the iueana for soiaploG iroa

the Coiitrai Ataerican Ui)land ^./Oijuiation are 3 to 4 scaioa

,^roator Uian roeane for tiie Caribboan lo^^land ani^nala, and

moans for tho Chiaons 3 series ?rc 13 to 15 cscalos

oox\5 than i'or l.^g i.-u^^CiCo-i. ;c iow*.iJCia poj^^wiatîo.*, 100

alios to the iiiortii a:ic\ saio 7000 leot lovxsr in elevation*

This rolationstiip between hl£;h ventral count and hi£,h

altitude does not iiold true north of Uiq UrisabcuCordoba

arei^* u small series Irou tliS vxcinity ol Jalapa, Veracrus

(3500-4500 ieet), a juvenile foiaals (CliMH 103445) iroLi 6

tailcs o ^'SCualtipauy iildulô^ ajiid a subadult uale {

Idjl'j) fro-ti near Tostiitl .n, i'u^bla (appro>.iciatt;l7 05iJO leet),

latitude*

There are other populations, unfortunately re;^ro8ents4

by only a iiUiuiful of r x.q, wiiich do not fit at all

well izito tiie vai'iationai :>atterz: dOvicribod above* Four

feinalos from eastert^ -<^.. ûia i otosi ** o in their moan

number of veixtrals vdLth U>.o sdoutti-cou&rul Veracruz: ea

rather than with tboao in the interwnlng area, Tm fsraait*

{\jm2. 41555-556) frs© CuatotoXapao in aoutiiSiii xcrîcrut iMve

a. moan alnost 6 scaiss :• than ttuat a£ tlie population

leas tjiiaa >u mi-ias to tiie northwst , and 10 acaies i^riîter

tlun in the TaUasco-CaiiijiacliO iôpuiatiou barely ^00 ailoe to

the east. There i-iivj bn ^'.^^ ai:nilicance in tho fact tuat

tiJL^ -: .^.xit'j' J.G not ûr iiou ijoa Au.^t4.a-^j( ^. xsoiatod

mountain raaâ in th© middle oJf tiie ooaatia plain, notod lor

haYlng faitnal asaociationa strikiji ly dilXeruiit irou tiioaa

of the coastal plain *t;i.c. - t,,

Sixbaaudala

Althou. h th« iêneral pattern of reduction In auJacaudal

i;u;;;i^cr - ..^..ixj^jx uu i.u^ ^.*. ^>iO veiitrîs, many of the

detaiic trikiiij-y dissi , roac iii veiitrals ti»

populations thrûhont tho Kisniosippi Valley have ensoiw

tially the sata© iiiean, aiici reduction in nucauei- occurs south-

westward, in aubcaudala tlio hlêat laeana are centered in

oouthaastem Aûisiona (both to tlie north and south of i-ake

Ponchartrain) and the mnber of suhcaudala decreaaes

fairly uniformly and rapidly northvaard east of the

i.issiaaippi iiiver, northwestward into tho Great llains,

westward throu^jh central Xo-xao, and alon^j tii© tiulf Coast

into l-tOAico (so© Fiîures 6 and 7). Tho mean nuciher of sub-

caudala tu waatorn -i..- .. .. . .:..;. :; Kit 1^ uo j.-, v- x.08 loaa

tiiaa in aouthoasteira U .tral fwanoas are

10 to lî lesej ai»d the oeric, . -,. -.uiiii, t:.o ;msartia

iti(\sxr& 6«-.ueo^rapai.c variation in aubcaudai number o£^MiA ^^ ^^>^ Uoatod 3* ijiacK blotcaoa

/ .^ ^.tTQ&a rrou viSiicli oaLi.>j^>^^> were taxton* iuo up: or.iociated with eaeii blotcii aro th« sample mean auu

xze (i;i p£.. iaea) ior males; tiio lowor numbers*'u KtiiQ saue daâ j.i^r loâlea* iWtan a liîo appears Inol oithsr tiis upper or lowor nuaberoi it aiLîiiies tiiat

jciraens o£ tiiat sex %rere present in the aauple*

Flûra 7«<-->*êoâphic varJLatiou in subcaudal numbor or.wuanQuhi,i^ r^roximua in kexioo arid Central Ai.iorica. ck

io8 r@i raaent ti

. ^^j^yfiv miiabors asu

—

in and sample size (in paiv , i'or

^ sane data j>o<* i«maLos«

îo spacitaons oi'

w.id -:l^ o^cii blotch ai'o ie

or or lower numbers, it siîii^-. o

jax v/ore present in Uio aa:;^;lo»

4d

PIat«au populutloQ lias d to 9 f^wer oean subcaudals la

foaalos (10 to 12 in fitiui.u;;»^, as does th« aaeiple Iroa

aoutiiora .'ojciii,

V.cat or the Focos liiver the nuabor of aubcaudals

tlian on ti.o iwarria ir'lateau _. . ^ . .. . j .Ilea to t»iO

east, iiie Coahuilan doaert iX>pulation ia siiullar to tita

traiis-i'ocos aniaala in subcaudol UMCkiXiZ', aoro ao Ui iuct

than to tiie geoLra,û.cAlly noarar iiuevo Loon poûlatioii*

Aloa^ U..C ûxi ou^wû of Toxaop tioo populationa aiiow a rauier

aaoBialous variaiice irota Uib iâttam doacriuoc atx)Vo« Xae

aamnie .le.iii Tor Uaivooton bay area foaaloa fits rattier wtill

juito tii© aot»^y.:iCu oa rcaucUijî -i.Oi»^ u.ic co .>:xu.* -.jwiS

previously mentioxieu, but ttic :aoan i'or aaieo - 7

acales leas titan £or tiM poptilationa on oitiier side ol tie

lialvoaton Bay araa (it xs avon loss tnan tao iucazi for

fetsalea fron the saoie i>o .ulatioj*). »>carcely aioro tiiai* 150

uî--.. xoi^-waur CO.... . . .-tuation is cojnpletely

rcvorood, - . . -03 iiavG tiio expoctad oocaplafflont of aub-

caud'iÎn v.diilG tlie nean for foraalee ia from î to 3^ scalas

In aoutii-cantral Xoiaaulipaa, at about tlM latitude of

the Tropic of Cancer, tjiere ia a atiarp drop in the number of

Bubcaudaia, Uie ueana decreajiiv, by 6 or ^ Bdixas witliin a

distance of leas than 100 laiût;. rrom the nee aouUiwaxxl and

QabC«^ca'd to w.i^ labaaco-Caqpeche area tlie decline in

aubcaudal nuiaoor attain appears to be fairly uaifoftt and

liraauaj. ibtîo u«i> i>e a fairly ateop drop in the fciean for

aaJUia botwoea aoutiwcentrai. Veracruz aiid iaba;^co-. -.o,

but oiiiy Oiio spocjuaan with a co:r; -iete tail ii^ available

fr<Ma tiio liAtdi" ^_; . .

. -..'îOu> , i..i'~- io'uiiiii* i^>jai» iiv*...iior 01

subcaudaia in i» prv....-,,.^.. . .arefitiy occurs in northom

Xucataxi^ Xh«rc -., ... raarkcd incrfttiKe i'; n;5hca;;r'ni:i iz: tii©

Caribb«aa io^nlûls, but i-x>r*3. co uivfe ;iaU»*i.-il a.;i ^^ ^

froa the Gulf aico of ttue Yucatan reniiisula to de

:>ul»eauiial uujaber ai>pare.itiy is loss well wrrolatod

-.vith altitude than is the number of vontrals. There io

vxiUuCilly io c-ax4^nmce ixi t . r -t oi :uu4.a^ i'roja sea love!

and of tijoae from an altitude of >0U0 feot ixi aouta-central

Varacru2, and only a -i-to 4-scflt'i increase in fcraaiea from

hiî oititudaa* iêtiuales froii^ Uiu uarjûaaaa j.oviJbaii.c;.St

Central /uaerican uplaado, and tne Chiapas iii^M«a*« s^w** «»

aajor differences altbou£i,h tlie hl^li*altltude populations do

h«Te siioiitly ai^tisr cisaiiSt 'a^«« laales freia Uiom po, i>aa-

tioos may demonstrate altitudinal variation more clearly

|

fl\»i.hftu^h once a(»ain it uiiouxo wc iîaatttuOered that the Chiapas

snakss «rs separated froa tiie otiter two i^roupa ^ lically

as well as altitudinaily. T<ia fnct t nat the isthainn Oaxaes

series (all froai less tJian 7^^ - - ^xtitcuîj; nas iiieans

aL-aost identical to those of the iiiji-altitude Chiapas

atuiitoa uiiue*%icures the need for caution in interpretation.

50

total .; ratio) Is^ \^th a Teinr notabXe e<i.ce^(;ionB,

remarkably ujilXoiai over moat of tno rai%0 of X« proxitaua f

aakla^ alXouaacea £or toe usual liitor^xîûlatlon voilatiou*

because maxty |jopulatioaa are re^>rosentod by only u i'oiif

spocioens oi" eitlior so^ itavxi*^ &ii/total leai;th

valuee availabiu, the iollovvin^, diacu&slon caii tîvo at best

only an aoproxlnation of tiîo actual situation*

* ... ,. v*iUi t.»o i>ropoit.i.oii«u.i.y ioiiiôat uaiis ore

round in the lov^x* i^.isoicaippi Valley, tac aeans Tor males

rantiiic i'âi 0.31'i to 0.313 and for fofualeo frau 0,301 to

0.313 (oxcoit tiiosfc £voia aoutiiwtist of l*ew urieaxiu an-C in

the Atchafalaya delta ruiîoxi ûicti iuavc 0.^96). Xlicrc is,

in thi& ^ ..a at least, a stxot:^ corrc^lation between relative

tail leiiijUi aiicl tiie niuuber of eubcaudaxs. . -xs corrciatiou

is wcai. or noiicvastcut in many parte of tiie rr..\,c, ..aat of

nortii to the Oaio uiver. ••uut of Mua iûB&xa&iy^l ivivor,

po^'UlatioiiS uaviut, co:«parablo ueatis extoiKi fro;;. t:.c iov;cr

i-Aissiseippi Valluy aoAâv.ard aîd weutkûxi Uirov.4j^ »iv-

end soutiiorn îl8;iouri to tiic liuite of the Usaxic riau. ^

i<^nv uDvuitij for feoiales, however, Oo docx'oa;^^ ^. — - ;ârk

rcfe,ioii).

bo/oru? this cxrcuMacribcri area, txie snakss have some*

wtiat Si.oi^ :âxl3, tiio ..oauu *i>. -, . ^ .-.Ui but one

except ioi ) -i-in^ bi..u..oou 0..- d 0.308 (usually between

0.296 and 0«30>). Til© OiM notable eACOi^tioa is tae Uiia,.as

hi hlando pOjuiation in viilcu tito iiino faaies ejtaaiiittd nave

vaiud ior lOiaaiea, U.^^yUy aXao occui*» in Wis iA)i;uxatiou»

/emaies iroiu outside oi' tiic "loa,,-tailed" area have âeaua

which ratHê i^rcxa 0#ii7y to g.^;#u (uauiiixy xi-u-^ ^•<^>v9 to

©•-i^^7)i but iii addition to the Chiapas fac. .oady

mantioned, tiiose frora trana-iêcos Texas (also 0.«^70) iâva

Siiorter taile tftan ara presoat in adjacont i^puiatioas.

Relative musszlo lontith as expressed by the muazle

l©r»ijth/iroiitax xqu^&u ratio is apparently subject only to

iatcrpopulation variation throuîout laoat oi' the raiii,e oX

X* i^Cfiiy^ffiUS*"^'^^^ Chiapas Hi^Ma^ds ;^ruiation, hov-jvar, in

this ae in GO aaity othur ioatureij, xa axAxorci-b iiXiiu ai-t

other populations, ^particularly tiiose ad^ceat to it,

Chiapas oales have a mean oi* 0.6^1, Xoiaaies 0,6v>4. iioans

for lour or more aales i'roui oti*er popuiatioiia riMitj,9 xôl.

0,?DB to 0.849 (B«ean 0.744; t ior ieiiiolea xrom 0.71i. Vo 0.638

The fused parietal siX>ta attain ti.eir iartiOst relative

sise in northern #lexico and adjacent Texas; in sone indi-

viduals they occupy .^lalf the l©x*tatii oi tuc iiiterârxeual

suture, as In A;UsH 67336 frora i*uevo hoon, in this area

tii - -v,c> - iiîi incidence of liiSht areas or streaks

extoivdini, forward from tiie parietal spots and forking; out

for varyiziii distaiicoa aJLout tiio rrontai-pariotQi suture,

fused and (iêir anterior edôa cilveri;,o at about a 30<-det,rea

an^la frora t}» interriariotal suturo, îvir^:; the entix*o

dition occurs oporadicaLXy tiirou^îout Uie ranâ in ii^divldual

snakes but it is praaent in 70 ^êrcent of the Chiapas

apocioaus*

in laost ivoîuiations tne r'ostocular linht spot

occupies ali oi i,uv ôwu. ,. û^ waouiiua of tije

aiddie, postocular* in tne soutiiem ^^rt oX* the ranô from

trnr.G-i GcoG To.Las to Coahuiia a:Ad i<uovo l. ou, and in the

coaÔui, p.ûjui -. ^ .. i'ropic of Cancar Uu teatrai .iôt'icu,

tho tX>Gtocui.ar iiiît spot is i jro<iucatly abaout and wxiea

present it is usually not brit,ntiy colored. The spot is

also lackiii£, iii au occasioxial individual £xxhj. uora iiorthem

po2>ulations«

liiQ broad black postorbital vxtta u^^xc^^ ^^ ^,

sauritua is usually lackin«^ in tropical ûaoricaii specimens

O"^' £• iJC22lj^M&* aon prascnt, tnc stripo is Irequcntly

narrow and does not ejctuxic outo tiiO poiûocuxar aculo;>«

Tiiroutîout the rust of tiio rar^ê, tne presence or absence

o£ tho vitta, and its extent, varies individually.

In ttie lovdaAids of ..uxioo and Contral A.jerica and

alot% the coast to the lower kiss issippi Valley « the

vertebral striji'u j.ij utiually broad and incorporate o, in

sdtiition to trie vertebral row of scales, toore than iialf ths

5d

liidtii of eacii paravertebral row. Froquently it covero all

of both paravertebral rows. lliHii 9/^9 from noar Ciudad

..ajitOi TasBauiiptts, t^^^Q the vciv - . - ...ur„^ ;. c...v...-'iii^ c.he

oatire vertebral-paravorttabrai series oruS a sjuail portion

of row V «a wail. Outside of this .- .-. tiie strii^G rarely

iiiciudes more than iialf tiie width of oaca i>araveitQbral row

and it is often 9Q««what luorrowor, particularly in tiie

oxtroîU iiortii«m part of the rauô.

aexlciAii and GoatrcsU. *i-.iorican spot ar© alao

ciiaractei*i"nd by aavirxv:' a narrow lateral strii>Q, t:^ stripe

occupyinti i^ô Ui«ii ,isuj. oi ouc oi w.^q .- .. '-. rov/s on

wiiicli it is oituatad aid varyinti aiuouuts of tu© otiier row,

usually ^-:,-.îtly more than hall*. Fvrthemoru, ttis stripe

bacomes evon narrower boyond the j>oiiit oi dorual seal© row

reduction, tJiw strir>8 then belnti iart,oiy coiifined to txie

tiiird row, oaii usoaxiy covering a asaali portion of row 4.

Tlie Chiapas ilionlande population charactaristically has the

stripa iiivoiving only thn third row ostarioriy. Only about

onewialf of the enaôa lii Uiw *ûcvv •'-' -M-^

reduction of tna latoral stripe, iîorth ol' the ilio Qrawit^Q

the lateral stripe altooat iavariably occupies caore than

half of both rows 3 a.ci k aiul it does not becotue noticeably

reduced posterior to ti» loss of row 4«

iuw dai'K ventrolatei; ax ^w*^s.^w, »ti*w.i -.^ i'ouJftd in all

specimens of X* aûrituS f and is the raost reliablo aUxî^

character for distin uisixiut, between tA© tv» 3;jecies ia tiw

54

oiouG oX' j^« Draxla\m« It is brc i la the tiUj:;, ao

v/jiejT© tt4u v»iduh of ©ach stripe is i/7 to 1/4

ctripe is ciityiitiy uartov^r in tiie Coatrai Air.aricriii u, ..,

ctii.l acirro'S't^' ir. tiê lowlands of Contral A-icric:: zt-.C

iiC-^jLCO, uik. u2iii.«-.y Ciaa^'^..cari> aiUJ^jQw-ur ixjrv-. u.u :.,-wt

or cciitrai taxji'Jt,

Cope (Xdc$5) doscrlbed a ribbonsnake Trot;. Coeuiaol

island 9 ûiiituna aoOf x%.exico, aa iiaving reddiaii oranâ

iabiale* While i ;<ava not aoou a woaturn ribboasnaku wiUi

that âiû4.cu4^r labial color, living, ii^'ociiuens Iroia i^rltisti

iiondura^y Ciiiapas^ Vuraczûz, Oiid oas&ern :>an huls rotoai

do iiaVG yelio;w,oraii/,o iaoiciia as o,-; oscr to tae tXccnisU

Wiixvc or î^Qiiuxau yellow orâ jjI'uscu;; i;^ xiiuxv xcu^j.a iroa

Coaliuilai iûovo i^tton, tftid Ui9 Uiutod ;^tatea«

Due to a doarth of color rloocri^.tions in t^tc litora*

turQ| or acco;apanyin^ rauaeun epticiiucaa, 4uiU to liiîtod

arai lability of series oi' iivo anakea frou tfirout,lx)ut tha

rai^.>j vx u-i'-; :j^ '.Jtictj, uiu xuxlu\i^.i.i, uiscussion caQ oixljT

delixait tiie geOt^rapiiic distribution of ttie different varia*-

tiona in tl\c color of tt-.-' (ôrsura, vortobral at^ri o, aiid

lateral :iwrx..-c xi. . . . „ i- >. ^. . .v. .. ^.. >-o.., '

L.ora . i'ccisfe doliaition of liicai o^toxit of each

color variation will nood t<> '-^ worked out on a local baaia.

u^>«ci^..oii& ivoi.i I'OaauXxaf i<aovo A«uon, uxid aîârsntiy

trana-t <iC08 ioôta (tne lattor %«oro xiot aoeu in lifa) xiave

:>5

an olive-i^rey dor»im, ThroiAt,liout the rest of Mexico and

Cantrai /Ueirlca, tiva doraxua ie luor© o£ aui oiive-brown;

Ciiia^© ditJiXaxi6& axauiplQs are an exception tovin^ a coior

ranging frofe. dark to vsry dark browi, Xuo oiive-br>:>vm oJL'

Câtrai Aoêrican gaakos, as *Bil as sever&i other o£

bro'.^'n, occvir in fsoutlieni arid central Texas janc* aion: t^*e

<K>in;i^v/ard turou«^ thm Xoxas; panhai^cllG to eoutiioastem

Coiorar'o, <^ < ^r-'^^^;n«doroiim oaaitoe carie in contact \«ith

biack orwaa, j/Oiû.Uiti.ons appear to be irite.^ in tlieir

dorsal color, ^ ; black fonn is fount' fron contrpl ^rnsac,

central Oklahocta, and eastern TcvwUii aor&î

tlunoughout the rast of Uie rari^ê.

The orani;;© vcrtobrnl strir.o has ti» moot extensive

tioOt;,rii>awC uistriuûioii, Uui**tj r»jpiacotI by another color

oiiiy in tropical ikierica, on the . 3 Plateau, and in

coastal Texas and liouialana. xiibbonsnakes on and in Uio

vicinity oX" the iidwards Platoau iiave a brii^.t red vertebral

stripe, Jni'iiero broâi-dorsuti iôpuiations occur aiom; the Ouii*

Coast of Louiaiana c^.j^. xw—.,-, oau o«i'-^,..u j.a vûoiiy a (joJxi

or ^.reonisn 4,oic Xxx color, it i4i;"x*etiucntly baa an oran^,©

cast, strii.o coior in rdso i;,olci In tne Chinnas

ajLîlauuv., H rociiiit/Ay colxoctuci s^xjciiiiou iîi l*i»il^J ivom

near i^capuico, Uuorroro, haa a . iich a;>pearo to b«

pals jôUowj the color tnay iiava been ipid in liTc, Snakes

from Cwntf^l A;iiorica amd ti\&t poxtion of i'i«:dco not already

discussed, liO. tiie soutiiem aiid central Gulf Coast and

56

adjoiniavi, upXando, have a .^rcyifuh tan vortcbral :>

Wliich msy rxr^p^-.T tn h.-svr- r. :>-^t.-llic lust-or,

iai -i i»JOij^ oL" tii*-' :. of T, p:p.\ i. , . ,

1 i stri.iO is usually aowo siiado of llîit yoUow,

although it is eeo«iî-. . - in lowland Noxieo arx!

ral .t";<!rica» Two .'spocineas fxvx.i lirazos Comity, Toxaa

(Ui' 12193 J, arn imic'oe in having a L.^tesrii ^ri o t at in

lire was blul34A tî.-\s-^.^ on u.ic .ôtii iniu. „xx:îu.û vi.ii&e

posteriorly. It Xa not known whethor tii ition ia

eharactorirjt.ic of tho ontiro vO ul-ition or acciura --rnly as

cu; i.ici-LViauax vu-"i.(aUJLo.i, j» i«..vj;:.o t'./ .iao

had a tpld-coioroti vertebral sfcrii.oj tii© other had tiio

orantio stripe typical of thio area.

It io obvious t^tat Qucli of the geographic variation

observed in X* i^lV^^Vtff l-a oi: the discorclûit type; almost

axx tiic cuaraetera whicji x aavc (Ucc ...iG'i.: vaiv l;* ^^^ .-Kêiitiy

of oach ot;Uir, atxhe boiri^ coaatant throu«>iOut a.

x.

cal area in which oao or several of the other c.taracters

uuder^ô radical change* This diooordont variation is wiiat

%fOuld bo CA^^cctod, tovwver, unleao all o£ thcae caaractors

are genetically linked, thus requiring; certain coabinations

to bo 6clt)CtQd a& a sin^^lo unit* i'aia does not ooeia likely,

and it ia fiioro i)robabl« that each caaxMCtor is aubjcct to

Xi.. : .at aUXvJC&XOZi Oj tac OaVX..\>... w-iit î-xxuca, lÛXim

Iiierfi aro two areas in iSiicii uiarKuc cii<av^ca do tuê ^..laco

>7

in a nuutber of cJuiracters Anrk-mr.lnllr .-;l--.iuIt:.ui.2aL:niY—.the

as HI ./toi'Ji cit «U>ut Uie

iatitiJde oX* tfte Tronic of r,

Aii vuy va-uv.^ .^ w.i;iv.- .xa.,:*>....... wvor/ character «^Qh

varioa ,

IXj - 'li nu'iboi" aiul .labial

color) t~;--ra on r.:. Ct ."1. - .'- in

adjoin! . -i*o iiito-i-aJl-ii-uae \.:i-..; ^,OiJU4,a-«ic;.4

a;. has beea at least ae^niwiaolated i'rom Uio lowland

populations for a ioaî period o£ tijnf>. Thn r? :ount of fiSJM

flow botv«eii them JHfe ttu_ .. .j .— -.-. . Iliis

iooiation, v/hataor ^^nrtiai or - -e, coUi-l^cJ with ac^-pta-

tioos for tiio riôi:x>uo ;-.istonco imposed b/ iiviii;^, at high

altitudes aad,

5 furtnersd by «êi*3tic drift, ftaa xcd

to tnoao siiakQS b^ccxain^ -^nt ia n&ny nssp^jctii froio

t-.iO sirOCA Xrj.i 'uxii.u.i tuoy iuivtj uwuu uuîUju*

Tuo abrui>t transition of cimractors in aoutiiem

Tatfnaulipas coincides v/it.h tîc tmnrition betwoon tlia

l4eotropical iuui iioarctic l"i*vu»viu. ry.ijii>**s, a;* nr.. -:.-^.iU for

sharp faunal breaks. Voatrai :^ ,

labial color, vertebral strij-^ w^^r, .«iv *at3rai stripe

color and widtii ail ciiaiitj© . ly at this point,

boiitiierri iouiaiaria tai^^ht bo considerf5d a thir*:* arid

ratiior aiiior •.. . oi co.xoruaiico. uoxor if v-ic w.or:jua,

srtobral stripe width, aiid vertebral stri/^-o color c :

over a rolativoly {larrow stori© b«t^^ ' .n tlo coast snri t'"ie

ixiterior.

58

i.u 6UL<jiyixi4^ iêo4:,raphic variatloxi Ui tiio vax'oous

Ciiaracters, otâ ie struck by tiio fact taat tae iiuuÛer ol

ventrals a<*(i aubciuidale is usually much more apt to

undergo Xocal riuetuatioJciB Uiati ore tiie dieskeuts o£ color

pattortt. The reason for tno i^reator oOservaDie variability

o-;. u.;uL.u <.i<u:i*idti.c caaractcrs is not k;K>v<ii« .uu^xux/ Uiu^

are riot eubjoct to aa riôrouis iîTlectivo preasure as is

color jnttern, and coii::QcuGr;tly aro less atabia,

Tae jiopulation samijlwu ditjc.^}3f;t! avjovn ^ôup them-

••Ives Into 0ix fairly well-defiited (oajor i^pulatious which

occupy discretf^ 4, vÂu^.uic areas and app«ar to deserve

taxoaot-ilc rccoîiitioa. v.aile •ocae of tîoê populatioiis

can be ci3tii\,uish0d on the basis of differences in scale

counts aa wq.juL -.. a*, coxor . ../^....r -, ..... .. . -a ii,aij ......i

placod on the latter because it is usually leas observably

variable over wide geOorapiiic areas, i«03aer variability of

^rLuraonts aave boon ^^rosontud Iti recent years, oodta if* tii© pai.es of dJjiPlii:iti£iJ& "ôlo v « boUi for aiid

L..

*^..n« in i 3

jLi^ u.t— or t«iw . .(., _i\,

or oi" ai> ovoi. •

ktit. Juon t) w

t; idualc

,

. ^ . •«*

L . ill Uie uvolutxouary ^.rocc :t istaui ioixed

' ore coa*define the ^ . _ .tone to icrocc on, xu.: L 4 is » CO a arbitraryc;.--. .*

anientto .L .. .-.. , .^^ . ..... .... .^_ describet. . u .re of iCtive ^ : in oimi

v.^'. • - ~ ,'^

, seetuB aiupXu juoUAAj.CuU«î» aoa ^ts&A.*w> wwi** Cvi wiM.' V. / . •u.ftilUfSC} UBS jjD %

59

color pattern* a» opposed to ventral and subcaudai nuabor*

appeajTO to be a cou]mon occurrence in the îêaua ThaâQi.;liia

(fiA^chy l94Xi i J •» vOAuo.i, vui.^ i ov., iw), *wv, fcCittor Morkers

have shOKoi tliat color pattern JU; iîdividualB o£ £• clei-.oîs

subjected tn low ttsia-eraturaa durinâ tiioir sn;brycriic

aiiiiaaXs wbich utiderwout cQVoloêut at iioxiual »

.

.xee*

Ventral and aubcaudal ouiabors of tiio tvô t^roupSp on tlie

otiier iiandy are etrUclAt^ly dllierîit*

Hecent eyste&iatlsts viuo have vK>rked wlta the êrtexw

•oake* tgenamlly Xit iiito one of tvô catejôrlcs: ^•^•yi.iioae

Mho base ta:iuonauic rocoîition o£ intraspoclfic populations

priciarily on color r.sttcm (Fitch and Fox) and those who

defiuci aubiip'isciita i^vôxy oa ta© baaxs oi' ailXoronces in

the nudber of veatrals and eubcaudala (Mllatea^, Mlttltân,

A, G» SffiitM, and H, k, cimith). Xhia dichotoiay of viev/poiiits

applies equally well to laoat othor suuko âaora* A

recent paper on X. s,|,rfrflAlr,g aftrW^4# and its aUios

(Fitcb and Kaslin, 1961 i in uhlch aurober of ventrols and

Bubcaudals is dlsiaissed with tv;o brief aentoiicoa on the

general trortds of ciiiial variation in these characters , and

an •arULer paêr on tmiikmimfk iiiirSAiflllf i^^<u.wi.u..a.;, ^j^ji

uhicii recoi^hisseo tws subapeeles aolely on Ute basii^ of

jaeristic differences, represent tiie extreiaea of tiie tvio

approaches* 'fae fact tiiat variation In color pattern aad

in aeriatic ctiaractera is ireîuently ciisconxioiit u£iôc t<ie£ie

60

diXferencas iii i _ - 'ac4i a Jâl probion,

Gooi^raphic raco3 iwv..-î.».aod by practitioners of tiie tMO

scliooia oT iiiter .ioa r aro not, in aost cases,

CfiuivAleiit. ;t v.-orl'-irr. to study the s.tm ..-.teriali

bo quito dilTor^nt, cvon if tiio .^- oT racos v«r«

recoi:;niaef'—wîia'i io unlikely.

The cUaoa inaerent in mxch a situatioa ^jroviOea

op^)onoats o£ ti^ aubi3pRCie8 coi^ct^pt with ari excelleat ar^:^^

mant Tor di acoiitinuin^ wau trinoiaiaX lyrataci of nomcuiclatura*

Isicitifiar interpz'otuTîvo a^^^^rcach coii bo said to he tiia only

correct one, inaa.T.uch as tiiC Ciaractcri: oi-phazlzcc in each

case ara uoroiy ciiiXuri..:'^ o-^^xoOiJiOiiS oi Oii .-iîc

aySteele however , ii* tite suba^y&ciea is to^

>ui

role in ophidian syatcmatics, a unifona basis for intc^>

protaoion must be aatablisliud • ^-^c oiiiy uiternativv; in to

discard tiie trinomial systea* nitiiouî a co.apra.iJui}^, of

sorts sniît be achieved by applying subspocific names only

to those pc .ono v;hicii display cojjcc- j of color

pattern and norictic charp.ct irs, this occurs no iafroquontly

tiiatr uiiiiy tâtiiictx^ ouiU ô tuirocoiîu.:^;^ »

Conaaqueatly i p. that i>- < les of ^ c

variation, at least in the case of Uis ârters:iakeo,

meristic ciâractori} sucii aa ventrals ai*d subcaudals cîOuld

bs tlu^rouîiiy invor;ti^/<tod, carefuily analysed, and fully

reported, but tuu^ oi... ôfinitlon of geoi^mphic races bs

61

b«s«d («ih«n variation is discordant) on color pattorxi,

which has been shown in sevdral siêcXea of l&MUKL^Mdt ^^^

axperiffiontaXly and in aaturo, to be the ;aoro obccinrabXy

stable ciiaractsr, i'aat [.roceciure haa boon foiiowed

taroiWi.hout t.,«j i^rt;auaw atuuy,

Ino pltfaLJls iiutorexa in an atta»pt to ap^ly an

hyrx)tijesxs bacmd on one srsecicn to otAsr lorma are I'uiiy

recu^J •' ViiO ii»vu..tiâLxoaa titu wnica licwo

^ven liiuch attoatioa to botn types oX Cijardctwra (ir itch,

1941 J tox^ Gonion, and Fox, isii; UiO present study) nava

found coior pattern to have iesa obsorV£ibie variability*

the species thus studied represent tnroe of the four

presently recoînizcd specios i^roapa [soae auuioru ivan

Denburg;h and iiieviii, 1918; i>mith and tluiieoy, IJiO) would

recognize a fifth group to accoaaaodats Yhaninofth,iy

an.-'.ustirQatri,q] , walch leads OG to believe %hat the pre.

ceding coiiClusions on cuarp>cter variability crui reasonably

be extended to include u .v, tâtire i,enus. Xaey uiidoubtodly

appxy to a nuiiioor of otnor siiake êitera as iieli, but each

4,roup aiiould be t^vaiuated ince|;crideni;iy.

The aAA âjov po^ûxatuxiiia vjaxc^* j. î.^. Ot^JiiZiii<_,

herein as fieoîrapiiic racca of thMdiO a. -i- -

characterized below wii^hia tuo rra-;,ev;<jrk of tiieir foriftal

i^Qiicrx^rXfXoa ixu auOi*pcc^vi;:..

the followifit, itcy for dxatintûiaiiin^, Um subspecies

o^ 1« t>roxi<a>|s was espeeially desi^^ned to be used with

62

living:, or freshly killod specimens, i-iîanuch as cartaln

iX oxily prcu^^ v.... ....^^jcXolL is avaxi.able« cons. ie

mapiunBiQ will have to be ijiac<*d on "ideographic probability."

jêy to tlie vûbspedoa ojT'j^'toftlflffififliAff VirailJiiBUB

1* l>orauB black; Vbxâbral stripe onuii^Q and

usually oarrovi; dark, voatrolaterai otripoa

lacking £• ii» iiSoîLiiiti \r» 6^ )

!•« Dorsum olivcs-^roy or aouio siiade ol' brown • • • 2

2(1* )• r ornun dnrk to vory driT-fr bimrn; v "rrtrhral

strxiû oOici, aot broo<i; uark Vuiitix>icitcral

otripo broad, uauaily i/6 to 1/5 of a

ventral scale wide; tail rolativoly siiort

(mean tail len^:,Ui/total leiit^th iii oales 0*275,

fefsal^s 0.«e'/0); isarietal aîota usiualiy

i;-»'jvr34i«.a.

.

••••••• ±,m ^» — ,,.-.tiUf.^ lp«S]L )

2'(l'). UoreuM oiive-.t»rcy or browi* (not dark); c&rk

voatzo lateral stripe relatively nai^row or

absent; tail rolativoly lo^.^^;, Ui<â:^ tail

loij^th/total lc;%,tii usually e.xou(.;ii)t, 0»2>0

in aalo.., ^.-.^w *u feisaltiu); parietal spots

x^rely ctuvrorwsiiai^cd ••••.•••••••• 3

3(2»). Vort<3brai stripe brii:.iit I'cd, not bro .;

V'..iîu4^wurajL Btrx, U.V i<arix>V) or lackinc; • • • •

63

3H2'). jwil st;rl|)o o„,

i taii, or 3oai«

: o£ iioid , 4

4(3' )• Vortiibral stripe îieyia*. â.^, î-uauj ^..ui^j^

^ Uov*-o ; iatarai atrip© liarrow a:iteri-

oriy, furt. ijoatarioriy; laean

or 157 Xa. i'eta^xua i# ii« JQIIuL. ^. » ?!?)

^*(3*). -'-^-^cubriu. stripe :-.r-n, «?. or soine sii3.(3e o£

ii,o jj.il i^-QQuimi v>**^t«« or £>iv . yuiiow

Xabiaiu; lateral strl^ not aarrov;^ at

loa^.^ ^«t$riorly; saean auQb&r of vexitrols

Qxcuooii'icS 162 la tualQB mid 1^ in jTemaloe • • • 5

5(4*). Vartcbrai stripe oraiit;c, rwt broad; dorsum

UiTaailjT oliv<j*t,rey; iatorax iit-rx^)©

frociuontiy reduced posteriorly; uarrow

vantrolateriU. stripe Xrequontiy present , • . •

I. &. to. -^ ip. 72)

5't4M. -— jral stri^^ u8uaj.-.y ôae shade of

geld 9 broad; dorsuxa usually olivo^broiaja;

lateral iitripc raroly reducuU; vuatrolateral

stripe uaually absent . . . j,. ^, ÛCaEitta Cp. 66;

Geo^^;rapiiic variation tux.: ,Lu.rc ,• •; bo,:.i diacussod in

i^:ji,^x.Lf .iOacG oui.y tiiv., ..... -oatwU'*!*** t>erti*iaiit to a i'oxtual

deucrxptioa oi' t£» su,.^ . ,at®<i bolow.

-4

jL ;i SmSi^ var. f„ftAfg,Y<^ lu^êrii, i>ibi'ox., U

Dun^rii, l^^: 5^5 (by ii-iplication]

^hftftWir-ii'ifft âurit^ var, orii i; uoco .rt, io>3:

757

TharaaQrizils £âurl,ta var. fi^jrovl I'uacril it iiocourt, ic393»

75a

Ta^^>c, ,^ . ,. ,. ... .,^ .-.uu.ivuu, 1905: 93

'^'Iut;ai0:-ai3 ... Ji iôeoiaari, *-j

{^QlQty:>o . Tn«r! irX;^.r:-ntini> -rf-JsnntQf? for t;io s.ocies

be Xoat, a nootype itaa not boen dooi^âtod because Uie

oriiiiiial doacriptioa (:>ay, i^<i3) and tiio typo locality aro

adoquato to aasociate Wie natao with a 0pi>cific ^x}t.'ulation«

Lariiiition . r. subo, eclos of ^îtMlttBiirila itrftÂflliB

eiiaractftrized by a blau.. uui iîî, >* narrow oraiiô vertebral

atriiê, aiKi lack of a dark ventrolateral atripo.

Uark>Q » riii3 race ran es Tro..: iac'iaiia oiid oout.*orn

V.iacoixsin aouUi Xii uw i'ixauxssi^^x vaixcy to, out xiou

iacludini^, a;>ati>orn Louiaiana, west tiirouiji aout.iora lov-a

and extrasM eastern iiobraska to central ilaneas, coatral

Oklv , arid oastem lexao (o.'icluaive of the coaut).

Deacriution , 1.. -h aa ti«B holoty^/O ie tiot avails

abxG, the following:, description is basod on UF 12160-1,

65

an ad\U.t sale colloctod 6,5 r.iiie» S.SaffordvllIe, Chastt

County D rîinna&f by >.illiaa ^, liueXlman*

;icut©iiation is as roilo'^m: oculars i+3f toi^porais

l-t-2y su ^: rala iaiaIs w^ ji.£4îv.x.:s.i^jicu.;v ^.v^, xawui'^ônlsls 2^

dorsai scale rovm X^>»X9*i7« voiitrald 167^ mibcaiwiais 106«

body ieii^th is 39*i :xi#, taiJ. ier^tli 166, tail , ./total

The tlorsuiu is black ov«ur laoat oX tiie body, b&t

a v»ry dark browsri posteriorly, dart: brown beiov^ tlsQ lateral

otrip©^"'' iae vertobrai auripo la ^j&l& orau^f^ (tiaerz ^

Paul ii«7)t tiio iauoi'ai atriô li^tt yeiiov*. i'-iU voatcr

1a yallo'wlah <^*vi.;.i v,j.Cii c*. i^-ALQ ora*v.s;. w«i,k*û aiui*t» »'''0

iatoroi laarîu .to, tiie dark vontrolataral tjtrlp©

iackinr: entirely. T. êliow postocuiar n..ot nrru)>ie«

ajLi oi u.ici xc >..-:u>-i-ior uaiX of tiio »lciolo ^;itocular;

tbe fuMd parietal epote are sôdoratoly proalneiit and

bri^^t.

Siacussioi^. i»ioristic and proportional crâractor.

istiica ol tais subiii-Ksciea jaay be auaaaarized as roiiowa: 2

to 4 postocuiars; ^+^ ô ..+> tttaporals; 7 to 9 uui.i\. .-;

d to 11 liikTralabiaiBj 1 to 6 ititeriôaiaioj IJ^ljî;^ to

«il-l^«17 dorsal scale rowa; laales 151-17^^, icnaics i>4-.

176 Vt;i.wruixtjj ... .... :a y^^-i^l, *-.. ...-^... >sj-.x*,u ttauc«uv«a-i,s»

£ialaa raiitâ xroa Id^ to Â«dt5 iîa* Iti body liUitÛi^ 1/4 to

iOl color de;^criptiona la this study are baaed on Inûîlykillao, ireaiily &im6 sêclraeus, uolaas otiiorMlse stated*

66

ratio 0««io4 U> v»i3>» .5 to 6 jo i>ici,

in body ioritith, i77«5 to 956 saa, in total l„,

.t

tail lc:icth/total ^^ .„*vh ratio 0.-^67 to C,336.

int between X* &• nroximua and ©ither 1. î,

i^rilino.. r X. j^, diabolicua have not br^cn 3««n in

lifo, but rz'oja tiie little tiiat can be i-jcoiâiîc

prtsorvad spocisions about the distillbution ol* bxx}\2i or

bl.-.ck dorrâ ij'xarr; t.'ic rnn cs of t'lc aiibn .ecion woulr* be

dXjêctcU to .'..uou, it ajV'-'**^'*' tiiat iiiO uxaci. j^, ^» ^. . j:

intcr^^rados with oao brovct race (. ; over a broad

zone in nouthv/eiJtom Kanaau, wtistern Uklahomai and tho

eastern part o£ tiie Texas pontmndley with still anotiior

brovn fori ( rubrilineatua X ^^^»^\B?3LitC^lfl intorf.rudeo) in

aoi-u.^-cuaural Texa^.^ ^^ Aiiiaîy with j^, ^. i\.uxu.xjîu..L,aa

in oaot«-ccntral Te.::au« rada .iozi between X* U* RFTf ?tiVnMfi

aiK5 ^. ^. araritia la discuconr' tindnr t.!jc inttcr fern,

llQlotVDQ . XU 11764, an acuit fonalo rr<x:i ..,

J. on i^nrisJi, uouisi^.na, coiiGCtcd 2^ >'ebruaiv 1950.

cUaractorizcd by an olive.bro mi dorsua, a brood ôld

vertebral stri^. , .' i.:ck of a rark vei^trolatoral stripe.

^Kefora to the fact that thia men livos alonij the coast.

67

Uftmca ^ Tais s >;ies r?!-'s,:GS i'roin Cat Isiaric mud

iSt« XauBtaany Pariaa, i<oui8iaxia| souUi and we^t uioi^ th»

Guif Confit, |>robr.biv aa far an r^rthom T/iinaulipas.

/pe . .iCutâliati .j.i as; JToilowat

oculars 1+3, tmipor&ilB l+*i, aupralabliiio 3, iniraiabiitla

10i) ititeru^niala 2, dorsal scale r^sivs 19«.19«17, vcntralo

169t subcaudals 106, hody length ia 463 i:ea», tall leiigth

200 iam«, tail ie r,feith/totai ient,th ratio 0»30iJ«

The folloidii^ color doacriptio .jg ou UF X2l6$,

an adult fooale colioctod 3 iniles i^' iôutto, :it, C.uirlos

Parish (approxiasateiy 7^ aiiea W Waîiiaiaan) by HaroL-; ,

LuiicieQ: dorsum oxxvo^&ruvâ C~ lâul l>i'>Vit <uroyi.&a

t»xt>«i& below t£tG lateraL atriê (l^t.4i; vertebral strlpo

îd (13&4)t broad ; latara^ v,w*ip« yallou llOEl); veater

lâla tinmen aiodlaJily (loL3), lat<dral oar^â oraxi^ê (llii6i}

dark veatrolatoral strlpo lackiiî; entirely; Ci*ixi aJLoIds

\^hitiii au^rolabials paid yellovdali ^^reeu, iuitarlor throe

tarmlsh* I'ha palo yollovj po^^tocular spot occupies all o£

the loMjr and middle posto caliLrr> and tiao loading edfcO of the

upper one; tu: i .go*. j^^Luîctax S|.j0t3 are iîotierately iî ,, .icic.t

and bright*

Diacuaaion M Horistie and proportional c^mracterlsties

of thla aubapedea may be suomarlaod aa X'ollov^: 2 to 4

poatoculara; 1+1 to ii+3 tenr -orals: 7 to 3 supra iabirls; 3

to 11 Infralabiala; x uu i* i.ii,uijj<^;;*uw; jL/.i^-i/ wu *^.A-'-.iî.7

..a

dorsal ncalo rows; roaios iOi-li51, feaalco 15A.177 vontrais;

u..*w.y >/.ii:7, r&i;;ai.oe d7»i«di> auucuodaxa. îaioi; X'aiivjO Troa

13y to 5^3 aua. in body iaii^txi, ^43 to 76«i uiu. iji tc .

ienothj aduit tail ieiii.tn/total ioj^*.*. ^c.^x^ w.-dO to 0«328»

f'eciales rano© iro^n i7u to 900 a-i. in body ^ , -H^,i> to

IfO-^-^ imm in total - ., aciuit tali i©%th/totGl ios^.th

ius,M;i j»w73 to u»3<iv«

I have »ooa living i^,jriyaUfl ^ MJSiSJmk iiiter^rnxieo

fron Harris County, Toxas, and St, J«â and nort2»m wt.

Cuarlea larisues, iûisiana* Ail navo vertabral atripoe

ooro or losa typical of ororiup .. The i>t, C.aries .

apeciatens v^* 12161 rrx>x:j 1 aile i» ;it« .^wî *a.o^ -.iu.^ xairly

tyijical browiucioroua orariua to i.i<:iviiJuala Wiiicti aro aiâoat

black, The kcyia ol tije ciorjal scale e sxc brov;ii, but vary-

ing, aaouuLa oi t.yi r-XjOiuiiit, ..i-.,.-. j^ -.^c. oc.ô are c,

iipedaeoa frou iw&iuijÔf ^^ân Patricio, mid Victoria Co ...-^ 3,

Texas, also have varyir^ OtaoiUita ol black pi^raoÛ. on

e£iadutiaU.y broâi acal<ja, but Uiaao ro^>rc&ont variations

rroa a viiolly brovai ciorauâ raUicr t«aii iiiturûcca ixuuMoucn

ae the bladc^doraun nrflilk^m doea not apiêar to raa.^w w..at

Xar weat iiUand. i'ioat o£ ti» '^t* JtXiJbt Parisii iixiiviclualfl

are entirely black*

The axbcûxQ.i x^ .lar.'Ji:; <.ow.i\,j Xw uuo gxou.'*

adulta (UiK 12^03) rzD.u Chatuijlvicw at the iioad o£ Ualvaaton

bay iiHv tiiitiroly black dorsa and Îd vertebral stripoo,

but a aJbadult irou Uvo Bum locality .ra to be an

69

SUBBXiM,' '^-'^ £>ituatioa la furtirior coa^;licatod by five

typical orarl^ Troui 9 miies W Tofuball In northwcatsm

riorris toiuity, tliis ioc^JLity Ueii^t, acveraX miioa farther

iiiiattfi tiian Cuaimeivioi' Hîen^ iiitei^^ratêa occur, T.y:)lcal

tlie east.

Bo live ixitor. radoG with other aiibninninn hsvfs bên

area o£ iatc^. xXon b^tviom this race, g» ^» yubri-,.l..ioatf4i:i,

^^ !• Xi* diaboliqt^. SpoclnienQ fro^; 8outii«o<mtruj.

Taiuaulipaa aji.car to be iiitereraces a orixicipB

nt<fUftr;^S» U4,^Wili49\^g ^^ SiiSiUaS^Jk* or nll U^rco fcrrm.

IhimnftrMffi amftoMg rubrili.iQatuq,3. r^;^ subapocioG

itolotvpa^ UiT 141J6«3« an adult r.iai.«« Iroa tiie ^tat«

fl»h Hatciiery, ii.ii miios HW in^roii;, êrr County, Icxas,

colioctefJ -i6 Juiie i960 by M, J, Fouquotte, ,}r, , arid

iiou^las A, i(oaatQfi»*

P^^ft4Uft<>* *^ aubu^cios of Xhatmio.îi^ rmtMîcharacterised bj uj* olive-brovû wo olive^j^ruy dorsutu, a

briiîii x'od vertobrai stri^j^, tujct a tmrrow dark ?oatrolat<?ral

8trip«« or none at all*

jatUUiS* ûjkS siîiîiQMiii xa cw.aîubut<iu tiUâuiÛout tiio

idwardo *^latoau raôii o£ ceatral Texas aiod nortiiv.m^d an

undetarciined diataiice (probably not mudi farther north

than i>au <tnt,ai.o or '-&aco)«

iieXera îrestiiico of a briijit red vortobrai stripain racc^c^ .^« tiuo »AbspocioQ«

70

Pfa<yji^ff^to 2L agl9lrYy«> 5cutaiiation is as loUows:

iaitori.onials 2, doraaX acalu rows i9.1>.i7, v ,

subcaudalfi 107« i>ociy io^Ûi i& 3^3 iui'icf tail iei^th 14i.

ma«, taii 4.u.i4^L.n/totaI iei%,th ratio 0«^d$,

...J dorsum io olivo-browii {i'laart cs, taul l^J'jj, a

lifolifcer si;ado ol" brov.n tKjiow tluB iauarai atrlro (15-^)

•

lourtiiS oX' t,m body^ orctatô (9îJL) oa t:io xiscK.; tiie JLauoraX

atripe ic t yeiJLow (iO?l), The veistor io ^^vie o*^®*^

(JLSi>4)» tud ^K>5C.0rior t;dott ol eacit veatraX acute cream

yollovj} Uite dark vontrolater;)! atrii^ in laGK,i:i^» ^-^g aiin

•hiolda ar« wiiito» UiO uuîViAvibitiis ^-a^vj ^raoaish viiit«#

The pa-U yoiiow iioatocuiar Si.»ot occupioo aXX oT tiie io^.tsr

aad tiic ior.r'itif: iod g of the raid 'io ~os*ôCui:..r ; tho fusod

paritiW'. .. . oho &'£M iiiUi.M*ix\jiXLf ^jTOuliiUiiL a^^ci tiri^*^L%

^ JS^» iteristic aiid panoportionai CiorciCtoristico

of this subapaciee raay ha 0uxiari»»d as follows: 2 to 4

poBtoculara; 1+1 tx) *^+3 > lia; 7 to 9 su^raiabiais; 9

to ii iiiLra labials; 1 to 4 interganiaia ; 1^^-19-1^ to i>-i9-

io V _.û iv a; i^xvi. iv>-i/4, faoalas 158-175 vantraiflj

malaa vcull^, fooiaioa ';;3-116 oubcauUals, Haxes raxvâ Iroa

<i<13 to 493 ct;, in body ioiiîtn, 319 to 7il r^-i. iri total

^diiialoa raiifO Iron 16^^ to 7^ u.i» iii bo<iy ioi^tix^ ^'Jk to

994 «(i. In total l«n4;tb| adult tail i«ii^tM/r.r3t ni icr^ th

rawio iJ»ôo to 0»3'«^6«

71

6 miles ^ Oak nXLx., aaja Gourity, Xo^cau, aavo tiie usual

tMpii^h- T<^'' vortobral stripe. Tii» stripe on tho otiior speci-

men iui l«^04->i) is oraii^t^ Uirou^liou!; tiio iu:i«^tii oi tite

body* •« 3eri«s oJT Tive anak«8 £xosk ¥9m. baak u^riii^s, uays

County, all have the red stripe as does anot^jsr ocrias o£

five froffi a few miles li\i .luatin, Gravis Count;^« Only red

stripes are present in seven individuals froM acatfcered

locaiitieu jui ^. -. -•:/ vôUiiU^ • ....;..,., . .. s, tiie

type locality in r.err County ... r&l : > (otliors Sdea

at tills locality » but nob colluctod, a^ree in r>tripe color}*

Oxm o£ tio speciaena Trom ti:jo Lovil*s ^vivor la n;>z*taom Val

Verde County a4;êe8 well with the above; in tlie other ^^

X2139->^}» tho vertebral stri^^v l.i oratipi (i4a«r8 ^ raul

3B12} on the anterior half of the body. Tho stripe

thereafter is brit>-it roa (3112). Of a sories of 17

aiJu;iaA5 i.jro*ii j,5 aiioo ^ ôxoraco îw , -.^.>c-,^ wo«jjaiiy|^

14 have oraiit.e vertebral stripQa, 1 iuxi. / oraiitê

stripe, and tiie reiaainin^ 2 have red n^ripea. This series

apparently represents a population of rulirili.«oatu.K. -*

^tâ^ffijA^lAfl i^^tert^raviua but one wiiaa rQdniVQQ a ^^ro^îw

derance of iêsiutic iiifluonce lr<%.i diaboliciusu

4 red«stri;)ed individual from tite edjo of t.ie iialOMis*

iiiacanxaent at U^m braunfoie, Cosaal Couiity, is ro, ortod by

wuuaci il903}a îoyo (l'>3>> ^êsciv c ., - -.zurxj^jiâ

spedLoen ftora near i^loroovillu , wiisun County« Tiiis

iocaiit/ xs •oioe di. :; li' i t..- ..acîr;j£aoat cu^ tlie

ratiior thaa ^ 1

rubrilineatus, :;trecktir (1903) liidieatos th&t ûu i^bbon.

•aakeet in î .ri Coui:j&y uauaiiy tiavo a vq^ \ 1

8tr.i . r . .cjr puEsiini; record of a red-stripod snaka

is t.iuv i-A 4_iV>,.i ^^yOi) wiio re!^,.ui-ta oiic iiu.j i-ccos, i..,". s

County, wiiicii ia ovor iOO ciiics wcat of tiis 'tfâstorjifiioat

verified iocaiitiaa for rcd-ctiri -r' inrfivitiuaia. Ail

ot;.î' x'ccorcs froa t;raiii>-i ucoti 4U.;*is m*'& oi aaiiuais i;>.'w..

aix ortiiiê vcrtcurai otri;;©, Uiuoaa tiiis re. t.io

aborrant iiidividuai Wiiich reacs^biea one subs^^ccioo w}iilo

occuiTii^ witaiii %ia& riji>.jsi of uhoUmv race, x dku reluctant

to accojpt ttie accuracy of tiio locality data uxitii £.c( itiouai

ik>XQty;^3e , UF 1221Q, a largo adult foaale £xoa tha

itio iiariador©«, 3 ailoo w iâriadorws, Coahuila, /isxico,

colj.GCt«d 3 Auiûst i960 by 4;oiyOr Coriant,

HSJkWiSAm* -^ aubsiocios of T;tft;,^tftâ>a ">^^^"*

charactorised by an oiivo-^î .i^ ou ulivcbrown dorsLu:t, aa

orcui^ vertebral 6trii>e, a narrow dark vaatroiatarai

(Stripe, arid frequent reduction in the widtii of tlio lateral

^«in allu2io:: to uiiothor horned innabitant of hot ^>lacu&«The ri... j*a "horns'* hovêver aro strictly pictorial

,

boiii^ iv.. by t - 1

dkt^llwuio x^^ca*

stripe on tiê posterior ])ortiou of the body in tiie eouWioni

itaotie ^ This gi les rajtiês from mo tst-central

Tanauiipas, Hugvo l.oon, .citJtern Coahuiia north-

Siortit.i^w;. i,-j;'cx*t>u t.iC racos vaiiey of Tc-x.:. o>u.. uou ..e.-ti.co

«jid, possibly, the i^takod Flaiiis of v;u3tem Io:uio coid

adjacent Jêv ^lexico to oxtrsme southeastern Colorado (Baca

County)* loolutod populations aay ojcist in tlie itio Grains

Valloy in iôstern llevv iloxlco (aue liscussioa).

Pcscri..tiQfi 2£ iJfiiaSjm- -cutoUatioa is as follows:

oculars 1+3, tesiporais 1+3, suprclabials S, iiiX'ralabials

10, iiitoTtioaiais 2, doraal scale ro>.s i9«l"'i-17, ventrals

309 mta*, toil xefAbta/ôtal leai^tii ratio O.'^^C*

Ths color p^ttom is as followr>: t'omui olive-.^_,reyj

vortubral stripti y«sllov4ai'i orau^ ^^...'^.ux<*;.iixy uoi«ia«a tiis

iiood, a briît orai«^s txioreaX'tor, not bxâd; latoral

stripe reduced ia width posteriorly; venter tyreyisb \iiite

««dially, pale orûit^ ;iiart,inaliy ; dark wantiolateral stripe

occupioc 1/10 t;ic vddth of a ventral scute on each side;

ciiin s.ii.-. .lua ;,.uta; supi.-:-.-:^.'-. ,—.•- ..-'— .'^•-i- -u^:, 'ih6

lii^t postocuiar spot is coitt'ixjcc to the loMsr postocular

and is tioK bristly colored; tho fused parietal spot2& are

jiôminent, briît yellow ia color, and braadly u<.. l

with black; a black postorbital vitto is lacking,*

Because a color atlas was not available at the tia«

the above color notes wero ciade, Uia followiii^ supplementary

.4

notes are ^ resontod fc baduXt fetoaXe (UF I22I7.1)

col. in iiuacteca Ocuiôa, noar ^faata Catorluai .'uuvo

L.^.j.:. , i>j >. J, i-v v«j, Ji',, aiid I*oit.ias A,

u.t^*s/u^-> V J—.v«»i^«v. 1* v.ifciOi'ii v^ I auj. -c^ V V ^ i Vwi'UvLiai. Strips

brii^ht orai.^c (iOiiO)j iatoral stripe U ycilov. (J,„ 1);

vcntar -.ic yclio-..--.;-r':2n (1_ ..

. . xc u;;c ^^roiôrtioaal c Isuics

or this raco Ai&y bo sucvnarizod as ToHows t 3 to 4 post*

>^w--...*^, *+^ »/v^ ,+^ te.iporala; o to , ^w:j.raiabiais; '^ to

X^ irilraiabi&La; i to 4 ixitortôaiais; 19.19<-.i^ to 21-i9-i7

dorsal scaio rov^sj naies 161-171.,, o i!)4-i69 ve^.traia;

:^ y,^«»ix%, *..iUiCL> j/— x-'j"-- ^--iJ^Cv..*.. -âxJ^ **«ii'jM i'oii^^w j. X\>L<

i^ to 41^ cua« iî body i9ii4:th, <^^9 to ^9o ^^« iî total

itii*4^tii; adolt tail io:^tii/totai ieii.^,t:i ratio O.îtSj to 0«307«

i?'oiâio^ r«uit»<9 ixu^ i/:^ to //i. ^.iu* î uo&y i.u..^'b.;, ~4ô to

lubU (a&« iu total len^tii; adult tail leii^ta/total iea^tU

ratio Qm^b6 to 0«304*

i'iiu po^:»uiatio*ia iti i«uovo iiooai Coaauila, traas.rocos

r«xaa, axii ooutiioiii ue .(:o aru typical oT tito L: oiee

aa hero doocrlbod, .^^^^^^^ ^^.oci;aeii (U.-î** û920) froai

Union Coaiity Xii extx-: .. a.):.'t.. .aistex^n l:«ew o alao 8e«aui

to boloii^ v;ith tills i;roup, KU 495>'35 from nartlcy Couiity,

i".j-ui;.-, ;:.id t.ui'i LL^j^-jô'f} iruii iiacu oouiit^ , L'Oioraco, are

a^ tiy reîresentativefl ot populatioaa oT ^^^4^1-.. .^^

altiiou^)! the riomum uiay bo aomeiîA darker tlian ia uaual

Tor this race* Tiie northuaatem iôw iiexico aiKi i'exas

75

individuals were ooiiected on ti'ibutarica of tiic Ca::.irfian

iU.vor« liio9e are tx>th •aatward-i'ioîtt^ sbreaac, yot Ute

snakfis show ciiicii J^sa affinity to ttm jtx>puiation0 ia iaoro

ttttsêru piii-c- .....; ... ^... .iiid Cisiarroa circuutaês tuan

tiicy do u> tUio . ji Uie recoa aiver draiiia^ô* aince

ribbouanakes in arid roîious aru ^êaQraily rcatrictcu vo

Uie iaidadiatâ vicjuii&y oi' perciaiiiiat Hj^riît^ atici v/atercoursos,

tbaae nortoam jxjpuiatioiis oi dxaboiJLCua are robabiy

Xairly etTectiVui^ jc ixuu ti:io^3 ja* wue goos urajoi-

•^ and raproaont r&lXc. ^cU «iare part oX a coutinuoualy

distributed ;x>pulation at a tiiao when oonditiona wore aore

nosic*

A aimilar pattern of diojuact diotributioa in this

saoe general aroa occuro in YbaaaoDtiifl sirtalia (Fitch attj

xiatilia, 1961), Tiiat species ii&a an isolated poiulation in

tiie iiio Uranda Vaiiey of iêw H«:d.co, nnd it is rossibio

that X« fifftîljBIMffl CUiy ij^vo o^vij t. ;. ,..:,.,o, i.,..crc ai'C

•averad old roooxxi& i'or tfiis , but Umm ^z soug doubt

as to thoir validity, irriir.i ^-id Cirard (ltJ53) list two

^.•o;uûg; i'n:j..> .^c .. uo û wuu^«al £urid ^ledinajp and discuss

in tiie taxt soeac S|^citri&n& from Ute iiio virando Vailoy«

Inaasuch as jnoae of tne other apeciaens listed are x'rom

localities auywttfire near Ui& Aio urax^de, x asauias tiiat tne

writers are reierriiii: to the Nevf Me:d.co s: ocirnouu, ^ .;3re

Is a <iiab*i;*u-i. ou Ui. ^ îuu Oi wû rivoi" jux iioi'fc»iern

76

Socorro County , but I aavo boon uiiabio to locate a ;a

anywhere in Nm^ 'oxico; aid i, d, Cnhibacii, who is etudyinii

the iterpetoioiUiia oi ti\e atatU| iiaa iiubnaod rr.c tl^t tm

knov/a ol none, kr« UohXbach also tiiinr^s taciX} i^ a ^mssoI*

bility that the snakes nay actually iiave boon collect ?d in

Texaa at Lâbiual, Uvaldo Couiity, aiid îecitui, . ra Coujityi

ratJier than in fiew iêxico. Iho sêituaus thoaaelves are

&j^âi\;.ôM^ .«j juu.^tji coant. Two iiidividuais (u*.*- v>J751)

JTroui **aaat ol AlbU4UorHua" <aay ba fnxu the êcoo draju^ê

inasciuch as their coiiactor, an outoraolOi'.iat, donatJ to

tiiC ;:;>^;.ju... u^.v -xui SjjCCxcs caJLi^-'CUo^. a;, ii-uu&xaiiS xro.~. icooa,

Lioii i-iiiûel C/ouitt/y wast to .•.ibui.jtuor'ûe iif\ it* Gaiilbacii|

pars, coiiica*). Thus t loro is lio conclusiv-- nvirioiicc that

the wt}stem ribboriSi:*ako occurs in tha ^wio oraado Valley ^ but

toe distribution pattern of X* airtaliâ dfyîa (incor-

rectly ruAOiiJu to îG ^, ^« oynata by Fitch and iâlin,

ly61) with it3 isolfiwod i>opulations ii» ttie Caaadian and

racoa dmina|.08 an \:aXl a:: nloni: tho iiio Urando, leads oe to

tiiirit^. tiiwv xt iây Uo 60«.

^pocitoene froa tite eaatem odjo of the ilaine

appear to bo interi^rados between dlabolicufl and| ifffi^tem

in tho riorta, dlftbollcua m^ rubrU^^*?fiVUg i^^ the south.

Noi^p.-coatral ieâxa populatxozis laay bo iutertiodiato between

ai.x u.ircQ racoa*

later aon between dl<^bolicua. rubriliaeatoi;, and

qriîue . in varyin.. ccx .oi.'iatio.ia, apparently occurs UiiX>Ut,i»-

out laoat ox xiix^nu .Ajuwauxii itsAaiâ ^huxmxuu^ju,^ x^\j..: uâ

77

eastern roothiila of the vîorc'o Madre OrleiitaX in wcat*

central litx; ; (as for south &a Ciudac! Victoria) are

here ooaslcierod to r? ^resent tne southorrtiwst ; o ulation or

djj^beliûe ^ but ^xvx.-^ ....... ... to oc

aA^'\^,44<;>^,g --..Xto iiiteiv. ;, Poasibic ..

bettôns dlabolicuc ^.d rutilorie ere discuosod usurer tlie

latwti^' uubspociee*

ytlMWnîi •*uri3;ua iiJriH^B'^tiff lûon, 1940: 19-^

i...^..,>o..;^^ Bire,jStoVfcff nttUffirAa âmuaa^ «^

Holotvpe . \iJiih. 13900, ar* atUilt female Troixi Cozumel

ielandy vûlntaoa '-oo, .^c.vxco, coxxociuu;a ^j^'i-j uaiiiwa^ i.N,>c>^

by ôrsonael of Uie U«;^« i?laU Coocaiaeioa vosi»«il .Îtja^naa^

(Cocaran, 1961)

•

iiari^.o . i'iUa race exteade irom extrcate southern

laîauiipas oouth aloa^^ tlie coastal plal:i, eaatern ria of

the aierra Madre Oriental, and Central American hliÛlaiids

to Carta^ô, Costa lilca* It occurs, in tho iâcllic lowlaiida

oiily at the istûnus of Tohuantepoc and in central Ouarroro,

There are ao reo9ro« lor Liio ui^îu.-.v .. ji vi«. ..o . la.

PgagffAg^^^ SiL uolotvaQ, ^cutcllatioxi is as Xoliows:

ocular- T+3, temporala 1+1,2 (the left aiiterior toa^xjral ia

fraj^«iiW«ti anteriorly), aupralabials 6, iiaraiabiale 10,

iiitoriôiuals a, donsal ecalo rov/a 19-19-17, vw:xrais 147,

aUbCiUldalS 97 it>^^ uci;iu<_,v;u uuX, cs-pjâvuiiiia.^ CU--^.'Ao-^^ > , â

78

ioa. th is li<iU ..t..., Lali xGa,,tri «ii5 au,, t-.iii iGzij_,th/total

^3^ isei* iUiii Q trai.i t oi' «^X4 îvti* i'tiia diucrc^âjicy in

body taeaeurementfi ia undoubtedly due to shrha^^jâ of tlio

hoiotyi>e durin4, /5 years 3torat,o iii preaorvative,

Xiie holoty;<e is describer" {Co e, lB'6b) as iiaviii^; a

brov.iij.aa»oiAVt; uoi^suuj vui-w-jui-ai swi'ijjC cJULSwiait, ouj.y o. . a

faiiit tracu; iatcraX stripe jôlo olive; voiitor paio olive;

sunralnbiais, iitfra labiais, n--.' fir^-t 3 veatrala rodtUah-

ormîi or auMiyovii a ^;air oi i.it^& ^ârietal «|jota prosc.u,

it ia not clear vnatner taici deacriptioa waa of tiis liviî^

aniual; lience a color description is itsre given of UF 12215,

an adult f€tâle collected 13 oiles WuW Ver&cruz, Vuracruz,

by H. J« i-ouiu^jtte, Jr., ajid iiou,;ias A. Hoaaraan: doraua

stripe; vcrtobrai auripe «^royiab tom^ bxx>ad; i«ateral stxupo

pale yciiov;, iuxrrouf aiicl 3li*;itly roducod in ividth on the

pt,;^>>i;i'-n./r ^yUit Oi t.. >-'-'-,I

V --> .:

. jl.- ^,oxi,o*. v-»i. woci;

bccoiiiint^ pale ^roen poutoiîorxy; dark votitroluterril stripe

narxt>v;, occupyint; 1/10 tlie width of a ventral on each aide;

chili ctiiielde and lorAsr .^ârîn of iiii'ralabi&la v.]nite; au.ra*

IribiaLs and upper iaar^^în of iiurulabials bri^^ft yollou

{tiiu ub.jjt live rutiloria have yellow-orai;û xciuxals)*

The lit.iUt po3tocal\r e^jot occupica t.io lower a:id j.!.

etV o of t.c ...id 'IQ .ostocuiar but is not brirhtiy colored;

79

the fused parietal spot a aro promlnont, brljrit yoIXow^ sad

broadly margined with black* *.iu ux^-^cc. iXjatorbitai vitta

is iârrow, irre^ûlar, and cail'lnGd to tne teu]jorals«

Kiscuaaion, Keristic atxi proportiofial ©iar'\ctt:ri3tics

of X« Sl» ri^tilorjlif aiav oe auaiiari:'/" . > iOi4.o-..a; -i u> :>

postocuiars; 1+1 to <i+3 teCiporaia; 7 to 9 supraiabiaia; 3

to 11 iiifrni-hisis} I to iatartieaiais j 17-lo-17 to

^1-19-17 oj^'..'.'.^ uoalu ix>viG'^ .iiales i4^^*l64| foiiialos i.4x-iuQ

veatrals; aaiea 91«>104« foi;ês iS^-^SB subcaudals, i^îalos

raage from 157*5 to 445 Eti» iî* body ieii^*"** '-*•*- *"^ -*'^5 aaa«

in total l»ii^tki', adult tall lîv^tî/total lax^^tii ratio J«î^

to 0»3O7» ir e;;iaiaK raii^ê Trcua 153 to 556 £3u, in body ien£,th,

*iO»5 to 77o i:*ii« ii.4 vuUcii iui»4,i,i*j UUUU.V* uojux 4*;iû.*/ wjiûj.

lcrit,th ratio xa 0«^75 to 0.306.

Tarcc ::3:voci;.icns i'rrx-:: botu'cna hli i4xinn rtnc* Cha'aal in

ejctretae aout.. ...wa i^-m-îi-^ma iii..4,ai*o.itxy xv)jjrosfe*it li'iter«

grades between Ol.VUffrJ^.a ««* diabolicua, Xh© two fetaaioa

(UI#U 101222, 101906) iiavc u*.^ îuo^w voitebral stripe

diaracteriatîc ol' j^, but in Ufa it alinoct surely

\Rras of so.T.e color otijer tnari tan (rreu.irabiy orange). Ttie

ventraj. cuuuuq (15^* -lô; - vviu.i:i li.je Lion.^J. rmi^

of variation for tiexicai* i^ j^. u/uii 110815, a ..iaio,

has a verttibrnl atrine whicii apparently i - tan aîd 162

vciiTirviAS, tiK( liiiui^r cîaracterlatic of a^^^^î^ît tne

latter of d4afa9,tlffl«,,»

A series of snakes {CKM 114532-5d6, -.-.- *j2a93-894,

Ui***«ii 16)/Aijmjddf u^^^* 1U5305) frua Ui© ^lezTa do i'aiaaulipaa

ao

on tô iropic of Coiicer appoaru to C(x.ibi»e ti^ coloz^ pattern

ol tf^-^r^M (or 2SMJm, - v xabollcua iuler^raCea) >ut.u

vei;trai couxitu which uxu uuch closor to triosu oi' rutiloria «

:- ITora tJw - *uo vicinity oi" ti te n; ; ear

• ratiier hi^^h vtiutrol count iUm TouaLoa huve a vcz^trai

count co...;j'irabiG to tiiosu oi' reaaie rotllori^ *1Q0 :..iici: to

t;iO souuaj* i:U2 ^,o^>ui.auuîUj Xii -ouuteni uan i»uû <vtouX ^lUid

noai' iauîco are coiiaidei'a<: to be rut iiorl

a

>

I'ho ofiXy available roprosontative o£ Uie jossibiy

roiict pO|^>ulation in ooaaLaX Guerarero {iii X«i«Îi) Ua;:> a

vertf^brai stripe whica ie not broad aiif^ dafiniteiy was not

XtiAki J.H, iiX'oy but rati'ier wBrnuin oo ^lavu uoon a âxu û..^;.^,

furtiionaoix!, Uie iaterai striixj is iarj-ôiy ooiifinad to row

3 as in^, ^, ai^ . înuj^ . ^n tno other iiaxjcl, the Siyociiuen

a^.roaa with Xowiaiiid rutilorJ^s ia scutoXiation, color of

dor&iua, width of ventrolateral atripo, and ^ropoxtioi^X

auzzle Xei\^th. Until bu&i time aa additional aaterial

(paxxicularly live syGCXi^^nQ} bccoiaos availabio to co.ii'inn

or dex^ the diatinctu.v^-acaa oi' thia populatiO'i., it vgould be

poiiitlea^ L.(j «îvc it taxonoaic rccocîition.

M a^GCXiiltiU (n;.;ii:' <d0u46} tcou ti« uoiuitaixia oI ooutiw

c.'iitrul oa:i i4uxs irotoa^ near âra^,oza (Ic-i. k-^, ; otosx*

i*xo \<«x'ctii *.uxxi>o.;>. J x:j rnxtikcr iai" i v u ai-o.-. u.*o rî4,o oi"

rutilor^^ aa pr. :ûiitl> unu»rdtooUf and thia ruconi ^Msuda

aubaiântiation.

HolotyLiQ , Ui* i*iî6, an adult raaie frwa tiia tx'out

iiatCiiCi'v "'^ -"^J^ v/x-xiiuooax ~«*i.- ........... s,... .^.... , .-..yAiiXi,

coiioctod i .ût>iet 1950 by i-». «^. I'outiuette, Jr., ûl

Douglas -« itos^'iiian*

uaa, ,„ , I'hia lorui is foufiO iu Wau uiiia^â.. ax^.^- ... ^

JTroia 6art Cristobal i«a3 Casas to Coait«ii) vXt olevatioxis

above ^000 foot.

Dasi ,,iii 2£ |v^Qtvva > iicutoliatiou is aa follov©:

ocul +3, ta;:poral3 1+2 (tjie aiiterior to-; oral is i^roi^-

jaoatfik; . .^.. -..*'.].y o;i oowi JXiiuo qx «*.i« atic.. ,, uj^.-ralubials

3^ i. uiaLs IQ, ixiteri-exiial a ii, dorsal ecalc rows

19-19-17, V9ntr;iiJ3 163, subcaudol aorioa aiui tail iacotiplGto,

i3»ody leri4,tU is k9j '^^^t

Tho dorouci Is dark browii (i'iaer» & i*ttul SLld) ^ olive-

brown bolow tiJO lateral stripe (15«J10)« Xiie vortebral

stripo is tîd (l^jv6) aiid uot broad; taet xat«»rux strijô i.8

iitint yellow tlOiil) overlain with browa pit^tant, narrow,

«uid i^rcauii;, vjuxy oii ruw j> ou u^fc û4terior i>art of tliB body*

Xii© venter is paiB <^roon (17i#4)i iXila yollow at ti*B aeck;

tiKJ dark v»Jitroi"tQml atripa is broc^t:, 1/4 tiio widtii of a

voutral acute o^* vucî ^.^ô ôa t*i«j waxi. «u covui'i^ t.^ o.iiîre

surlace ol tae Subcaud«ils), Xhs cUiii siiiolds axx) witite*

irr«iûlarly apottod witii inxlo orana«; tiie supralabials and

^Ralora to ti.j .xxji altitude at tdiidi 0.4^;^ siJosieciesllvao*

62

iniralablals oro paLo /Bllow.oraiio»» ^'«t; or tie au^^ralabiaJUl

t.KJix: ii? ..iuc.* browni

jîji' -.1* . v'oriyini, *,.i« yellou«ortin4ê

(tao aiiterior 3 pXato iplotely overlain; t.ierc ar«

vrirvinr: jsmounts, irrry \iL a riy distributed, on Uie postorior

^j, i..*»^ xiîC pOiiuoCi-juaj^ apot; occupiv?;: ijli of X.I16 io\'L.r cuiO

the anbtikrior pert of tiw) Liiddio pootocuXar^ aiiti is overlain

with brown pl£,acnt; Uê fuae^^, cii-avrotusliBpttd âriotal

fiôts are ^.roiulii^tt. aud pale yellow, overlaiii with brown

- ...oat,

Piacugaioa* incriatie and proportional ciiaracteristics

or this subapecieii uny be 6ua;*arx2ui as Tollowri: <^ to 4

postocuiars; 1+1 to *:+3 tenôvaiB; 7 to 9 suinixabialaj 9

to U a.iiiraj.auAtiiv*i - i*o > i.iu . - ,_ - -iXaiuj x^-i^-j./ ;*<j <^«.1V—

17 dorsal & rawu; :!U4le8 l>7«lo^, ictaxoa 1^3-lo3 voatrals;

17^ ô 4>> fda* <iikn i>i>u^> lu.^Luy ~**>x«> Uo u<W ^->« in total

Ûh; adult tail leiAÛi/total laii^^tn ratio J«'(^6J to Oî^Ô*

* «;. ales rant^s fxoia 141.^ to 5^3 tsva, in body Xen^h, 194 to

7a am» in total Itâ^th; adult tail lciv»th/total length

ratio J,-i52 to O.^B3.

X iiavc asii sewi ju.wur-^r^. .. .- . - .

•.. - v^... .-..^v^.,. --'

i_ ri& f but UiB/ Dii^ît be g.^^gcuk. to occur at lower

•levacions on Uio northern anr? iîr.tnrn r.io.nn oi' t;-.o Ciiilapss

iiionlaxMici, In supralabial oolor u. .ur»l surxô width

alpinus deooasL racoa its rulatioiiSûp to tiM wii es^'road

yutilorifl . Tite darker color of tin dorsum in aloino;. ^ tiie

d3

r«i5uctlon in width of its vertebral etrijê, aiid the over-

layia^^ oi. it a lii/it araaa (audi aa tae dors&l stri^ês)

viXth Aark pit,raent a^^jjear to be îdaptations to rcCuco

rclTloctioxi vAiA- xncrcii^^ w~)Oiyw*.,** >-.* ^^^...-a radiatioa Xii aa

enviroumaiit in wi^cii tito activity oi actoti*exî8 is

severoiy restricted by tiie iov. tcuiiêratures,

X.iv Aroiiuoat occutrti^cc o* wucu t>Cu.tciiation voria*

tioos as 3 inter^^nials and i'ra4êntatiou o£ tin aiiterior

temporal may Uavc been tli© rcijuit of incroase in

variability of aeal<3 ct&ractora an a result of eitaor

expoauro of tie motijer to low tOirieraturea during tlie

by Fox, 194^) or tti© ©ffa eta of fc,er»etic drift on a iiormally

rare variation in an oasentially isolatct? popui itit5n,

Evolutionary Ireiida

Witn t£ie oxcoptiou of g^jîûa vîicu la even non at

leaat seol-iôlaited at hi^h elevauioiis in Urn Chiapas

Hii^lands, the luanner in \îch the ^êoi^^rapnic raco«s of

li'fttfltfitffii'i'ilAiff trl^^lntyift uuuiei'wont t^x) iaolatxoa prosuiuably

neeeasary for their ciiifor<sJutxatioa iu not kJKJwii, fixtuouiiii

on tiie Uiiuxs ui. huQJui* ^rviSOiit o^;;/wrxuui,uû.> xu wuui.c a^.^: uur

tnat acMue eeolotîcal factors wero, mud for tiiat aâtuer still

aro, involved* w>o^tii% aaido al.î:tua toiich aas almnr'y Leon

discussed I wo f iiid tiiat: nîlorlf is co:J.ia&d U; ^

humid Aîodcan tropica; îabollcua occurs alrooat e;iclusivoly

in hitî, arid roi.lonsj rubrUinaatua ia founc> chiefly on

alf

j.&od to tiiO «>'aii Coaat q£ ^ûloiaua and Xojuls; uîci

nirgîtoiHg ^B difitritmbed widely in tiiat ,j»irt of tlio

iutorior oi' contra! Unibod Jta&od in wliicii ttia avora^ô

procipitat.ion &xc<»ed8 30 inches aiinually.

Xhla distributioa ;>u4û;^t;^ OkAU u.iie featuroa dia^;*

iiostie of eacii of tiie^se imjor ^>opui.atio!i8 ciay iiave bQCOcie

£jtnbiiir?;r:!:' in n r-articular n t\vlvor^\iQiit. ar: a rorjult of

ox* > do not iiamfiiori aurvxvax in q ijaxticular oaviron*

fiwnt UDui. V-..U to bo £>osltivoly soiocted. This aoe^s a

r«aao{iabXe ii' aouidwûxt obvious oxpioiiatioa, but xt reHuXx*08

cautious ap...i.iCiition« X* u» 4-^^boixci^fl occurs aiinost

eiitiroly In arid r^^^^xouBi, viiich iai^* ievû v>..^. wo .j_. ,i>oa«

t:;at its Cii«iructorx3ti.C3 roi'l<act acaptatioa to iiio iri an

arid ex4VirontuQat* Closer o.u&uiuatîoxi rcveaic that tnd

rxouoaii- ...- -V.3 iiot iiCtauxi> aciajji,ci- to w..-. .axxc coi«uiu-.uiia

but occu^ . rxctco xxiciiu, poraisc iii«^ in tiie limited

aaaic situatioiia avallablo in tiict JbAâediato vicinity o£

ponuanoat »vator« i'husti aiiail (ilsjuiict iX>pulatioa8 ciuxrae*

torisie rauch of its ^ roiKint distribution, and tîero la

roaaon wo u«xicve tii«xu j^.; , -plAPJ S. - i^*»co continuously

diatributed wi«n tiiO region in vaiica it now occurs was ouch

njoro neaic*

lii. . ck liornuin oi tiio iia>ii:i".tu raco (au i,\.ix aj tao

narro jf its vertebral atripa) may be an a ion Hor

docroaead refloctlon of aolar ro' i-^tlon in tho cooler.

as

oorthom part of ta© rangp of X* DTPXLmua wiior© the ability

to absorb artt^ r>3t(y.n solar heat iôuld be an asset* Tho

fact tliat tiiQ black raco aLoo occurs in so utli-ceutral

United ^tatQ3 wiiero such an adaptation would ooooi to liavo

little proaent^day value say iixlicote that X» a* nroxjaua

originally evolved in tiie latter area during;; a facial

advjiico, iâen cotiditioiis vouid lave been taucn cooler, and

k bri&l euoLuary uX our KnowxuU^ oi ti« ecolo^^y ol X*

projdiftua la ^ resent&d below* **o claidi is in&<\9 && to xta

comploteaesSf aiai msuiy as^>octa ars trvi&ted in uucli ^roater

detail elaoMioro (Tinkle, 195'/f has produce tlie mos^ coâ*

prehenslve stui^ wu u.ûî*

^y personal experlwice tiaa boon the .ro^diBue is

rather cioooly confined tiaÛtîout its ran^,© to tno vicinity

of pernaiient or se: x r;, ^.-ient bo<iies of viator, ©itiior

running or staiidint^, and uoet literature accounts aubstaiw

tiate this observation, Weill and Allen (1959), hotievor,

collected a.laj::ê adult bezieath a lo^ in a hl^h, dry

savanna iii iiritish iionduras. Open rTasisy aituatioiiS scea

to be preferred, but x .lave axao xooik. k,hq aûcjês jm. uot

woodlands, at a desert waterliole, and in ruaii-llnod porxis|

aiid ituthven (lôti) reports tae oc&arence of ttm ribbonsimke

in oraCi.xo<i , ..,..,_ixjv - :.. .^. at i-rooi'eso, iiucatai** * isw

06

ep«cies« iiii>ecii.i«»n8 oi^ |..n?.'L;...u$ luivo oocn oosenreti in

bushufl i.-i .ritisii :;n;:«ûr.nr (I uaîi £tt*«fcoi, ui**.- ioobraska (Hud&on, i<>4isi), ai*? JL

have coiiected tiiou osvoroi Toot vibovo t«» Mater (o!ic over

6 feot vjkj^^/ Jut ruohos in Chiapas aiifi âxs Fov ;^-,

Food

TlMi diet oi' tia weotora ribboasaako is chielXy

coiiXiiip.^ to aiaphibians nx.d t.<exr iarvao, :i^^^ to flriica.

Jpec^ci . I. arc coiiii*toj.y ioiovû to Mav. ^ *i Ou-Ltji* by

^ , ;iiiaua (otnar tuaji undor laboratory coi^ditionaj

are iiatcr' 1 Table S. Tavior (1392) and Fouquette (1954)

have al^ rucurtied in^ôcts ia Uxt aj«t oX tiie western

ribbonanaicei but it xs iiXely taat tfiese nwre titsi result o£

secondary inîeatiou, inr.Hû uuiu <«Aîw (i,/y7) indicate

tuiA tîOre is an account in tiie ixtorature oi ^« .xo^û^

aati:\, aicc, i boiieve tuijfit ti-ioy i.xty ixavc Dcen i'durrin^;

to i.tx>wbjra.ac(i i-Ljj/if v^.-o ropoxt^ -i..vx.^^ »;..,„a\;.->...

,;4 .. -.. a

iDOuae nest in a iOt^* i'uo oiiuke coataiiied tîyift,

;î..l .-^.... y

however, oot aiico. .<a<:ôtherai.c vort&brutes, lizardn,

eartiiV.oriue I and otfier invertu brutes , all ol vjîicii are

i on by one or more spwciea ol' Li^xtoruiiaKu slxtq

apparoixv^x^ lioi/ vuxuntarily j..io<êtod by X* SXSUliSiHA* ^'tore*

over, ;rf»iio riov^ly trfiriaforrj'3c' toada arc roaciiy accented by

captive r-tihonannkaa, adult bulb aro alaoat alwa/3 rcli;ncd,

it u»> oe i.oul^^l. . .u ue^îOors ox t^^c- ^'xtus uxx>u^ .-c

S7

T/UiA*.J U«——i-OOU Oi i.c... . :.-

;»uecloa

3

not as well a< . xoXoiîcaily as othor ôrtoraîakos

to tolnr~tn to i,.. .;j.c ."rotolc* nncrotions of a<1uit to«is,

coatinuQ fc u.itil it is iiitor. or tiiorc is no

inorw Too . .^mi2fU.in (1959) roijort^ «.»-». a siiiole 2«

pittxLuua ccuitaiiiod 64 motiuuorylaoQixi^ to&Cfi oi an u. d*

fled Srecios; atAd jvioiri (194^^) obo^frvos tiiat rioboiifl-

(as vwix ocvci'al Bfoc-i-ijt. uj. ., v.. . <. ^ foodiiit, on

auions 01 ijLSUf ta(j^x)iea, a.x; iro^s iii pools iol't by

a dro: in water n:V2i oi Lao ^mi iuitoiuo uivcr, „ort,<^ci

L..' .— . Jii ti^ iôXiiit tiiU'v ucu^y ox \*iHii a.ju^.'ow, c, v. cju^xxy

tue Juveiiiiai$| ato tneuisalvoa to dcatn*

Hibbottsnakaa are as active at nl^ht as during the clay,

at least wliaro nocturnal te.<.j>oraturoa |>enaxt, aiid a.^.arently

•] as raefiily at one tifie as ti» otiior, liear tiudad

A'latit^ jji ûuauxxpau, >^* u« i;jusuCttG9 «2r« (^>or3« Co. •>

- <^ !• vxt>>âi4fi etalk a callii;^. jdu ^ kaUtUO^*

When t:^; fro£,'e vocnl nacs rTiOvci!, t :c .1:1:1:0 raovod ciosor,

sto. aach tiuiO taw i^ -vopi.au c. - ^.«

Ueproduction

Xinkle (19^7) nas found tiUit ribbousnakea in soutn.

eastern u>ui9laiis re«cn aeû.^.. ..aturity in approximately 2

yoara and tib a mlniffiUiD body i^.i^tn o£ 4«i^ ^-i* in lomalos

and 410 ctn* in males* Oloyd iljn^) r^ôrta tnat a Kansas

id..iaxe v/xtn a oooy xeî^tii oi. -^î^ 'û* uoi u yuuii^, cu.Ki

.uer (1^:^) noboa tJ:«Q aaue \0uUi1, lor a fettle Izoa

69

Oklfthomi with a body ieu^t.. ^. ;»60 mn. Cianc {I9i^)

reoorcjis a brood froxa a 4J*iaca £mmkl» 1'roi.i itortaern

Louisiana, il' he is roxerririâ to total iait^th aiid if ti«l

aiiJL.K..i coiîj^w*--.; wwv^j., it.i5 ixjdy ie*-o--» i^ prolwU)ly

betiieen '}i>Q and 360 iau* the ao-killest .^^ravid XcsiaalDS mich

i .isvc cx-mincd raeaoire 355 tt^ritish lioîdura;;) , 3^5

(Veracious), aii6 454 (-ân i-uia Potosii) akj. In body ieni^th,

Tlwt fact ti^.t five of tuese aix aLiii&al.s are froia tropical

America aiay iiidicate tnat aoxuai taaturity ia aciiieved at a

aasalier siae ia that region, i^jttriiiKJiatiiic, i'rosi liiikie's

aawa c .."3ticiate w... -©c in floxico aiid Central

^k^rioa probably mature at a body leu^th of 300 iom« or

sligl*iy ioi; or,

I4auiiit» «4-, <•-.-. ^.- ,..,. ^ ctt iioou aitur t^m s^.'ring

•eerêace (iu areas \ . ,. w .^v .> . .;a.»iter iiiactxvity).

Carpenter li95a), iHeUl -^ .-ô.* (1959), -"^^^ Xiiikio (1957)

all iiaviiifc, foaiid ^^ravlo fewaiovi aii uarly aii «^ril, 'iite

youa<s» are uatually bore; in late July or early August « but

dates raotê from 10 July (Xinfcxc^^, xv>j > w-.*o.*4>i 14 Septoidoor

(Carpenter^ l}J^)m i>rood hxzb rmitâQ iro^ 4 (Car^cator,

i95ii) to ii? (ft.loin, 19^*9), tma raeafi lor 34 brooda beiiHi

X^îS UUxXyXiLlAiiXiim *- OX w»4J uroi>y^ tU*it^^ UO -UiCiVUae

with tiiO Qise of tae foiaale, Availîbxc datu ou body leri4,th

of newly born X» rojdqua are aunaiariBed in Table 9, X lere

ia AUCh variation between brooda in aiao of individuals

90

'«*«i>od> iutii^ta oi' aowiy uorn i'j-^j^. ^fitt&ft

91

at birth, but if such sire varies /'©o: Taphicaliy it ia not

birtii, but 0iu&il<)r as

i'araaites

iUrviood (193^) reporta rinding, tae heimiatiio

iunt.a aud stoiaaca, rospectivciy, ol i.î^rr'*fr,raf^ Î'oîauL^,

In oxaainiug MC2 5>39X«^f aa acluit i'eiaale ifrfffi^VM ^li'o^ ''^'^^

ôrldfty Xucatan, 1 d^s covered a aaaiX tick eiubeddod on Uto

left si/o ol' the f»ac* at tisc Juiictioii of tae aiitorior

Gîoriô Auasto^ iuis ideritiilcd It an a ^ . X staô of ti«

geiius XjBQdaq (species undetennliicd}.

tomuar aaurita Llzmaeus, X766: 3i^d

l«afcrix ffûrt^tii ^'Orraa^ lJ<^t i^

frfQB!«9tfl4ll ftflUy^lrHJ iioibrook, Xî*^: 21

litlflWIiyiîl'lilfl aaurltA Fit'/iM<êr, 1343: "^6 jby izti^^lication]

SutAaai^ aackoaii Aemiicotti 1q^ : >^

Jûtaonia aauy^.ta Kcimicofwr lii!Jf^)i 9C |by iBinlicaticn]

PrvanpiatftfiiQn g^Jta^gwa ôte, laou: ^^^

yrOpAt^gflgtma gaVMrJ^littfl Oaruuui, ia^3: ^^3

luutftonla aackflni liavle and îco^ lii83: 39

YtWaOOBlUg flftfiteBîlt ôiuiben:, ld94: 3^^

XtUHUUtfillJLl laurttUB '^t;o;io , i^L/o: I64

ThlMfWritiiB aackani Deckert, 191<>: 31

Sinca the holotype (colieetad in "Carolix^") la pro*

•mad to bo loat, Jcitaidt m^ Coruutt (1>>j-^7) ^îava

deeiâtod ao uootypa« U«;». 73X1>» an adult loiJiulo Irom

15 milea ii»^ Gharleatoa, jôrkalay County, ûth CaroiinO|

colieetad 20 July 19^3 by Joiui ûitiby,

9^2

TtMIWWlThiili eûsiracterisod t>y: I9*l'>»i7 dorsal scale rows;

« allele proocuXar; 7 or 3 mpraXabiauLs, tiio third mid

fourth or fourtii aod TiXth^ reciêctiveiy, eritoririG tli«

orbit; iA3 to JL77 veiitrais; 94 to 136 s^bcaudals; Xatoral

stripe on dorsal scale rows 3 and 4 {vo\i 2 is occasionally

iavolved); labials a»d veutrals v.itiiout black .iiarkiags;

darii vontrolritoraX stripo always proscot mid broad;

parietax spots j. raîuciutiy xac«i.x;;>t» , v. , , ^.- .. .^\. uioy are

aia*llf OQd raL'ely in contact or briiîitly colored; hemi.

penes short, eêndin^^ to the di^^th subcaudax ... .i

iuvorted; umxlilasy t&etiii nuuterousi usually 4(^9 to a oido*

7his species ron^ôs Iroia i-tichiân^ southern Ontario,

and aeutheru i^âjuiu, southward a^û o£ thD iîississippl

iiiver to tl^je northern side oT i^^ Poiicitartrsiiln in

Louisiana 9 and to tl» Florida Keys (Figure S)« Thorc is

an isolated colony in central iôva v>cotia, aiid tl&ro r.ay

be one in northern ivisconsin. Tiio s^^cics apparently la

xîre in much of tiie interior o£ eastern Unitod states*

Variation

Sexual i;itiiorphiatB

Tillinff^ntlln aa^Cî^m "faow wûal diiaorpIu«s3 in the

•ame eharacters as does £• proxiiaua . nn^ to about tl)Q sane

Klgurii ^»»Ûstributlon of Tliaainophia AS^^.^"^ ^^ ^Ûnited otatdo and CauaUa* •ioXid dots ropxêseat . .otio

o/^r. >m circles aro litoraturo records* -ârot -.>wuat the irobabic zonos or Intex'^^raca. êtweon

(iegrae (m« Table 3)* In populations Tor wlixch at least

riv« SQakes of each sex are avallablo, the dirforcncc in

..iuaii nuDber oi voiiur^^. - . u»<ai sc^^c^; ii» lavor oT

âs (CJiarlostoa Coutity, ôuUi Cazt>liria) to %7 in

favor of tim tmlBa^ i'iio ^.roatest dimojrpl^laa occurs iii Uie

wcGuon^ inali o£ peniiisular Florloa; la oti»r areas ttio

dG(^roo of diuiorpiilaca does liot appoor to vary ôc .xc&Lly*

The populations Trofa Mbich at loast 1$ adulta of oacii sox

are available are eoopareci in Xablo 10.

Joxual dioiorpliiam iii subcaiidal naiiUor also ai^roes

well with taat exhibitcu uj^ j^* i'..--. .iS l*aujU; 'jn^ UwU t

aot vary as widely, .v. ar<: lou Milii at least

riVG of r!\ch nc:: In t^-.lrh ryales hav - --..«r Bubcaudals thsn

tiiQ faaalea* i^â iêaii dJL'i'erencoo ratê froci 3«4 (iKûtrii-

ccr.trîl i'iissioai >pi} to lO**^ ( :K>utheastem iênfioylvania ) •

There doos not ..^^Ââr to bo any geograpiiic variation in this

cii^racter* Xiioao f»rAaatio:is for vîcl) at loaat ton adults

of each : j are c<waparcd in Tabio 11,

fei^xc;; ../•;.: i ,^^,q iorôr tiiuii tne uaioa Ujl j ^ro-

portitiitally shorter tail. . .u . '^ feciale e.

(U*" 12i;i0 fro.' DnAo County, i?'lorirfa) naa a body lon^^th of

670 ioT^*, a t«;Ui. Miii.i^J,il of 346 utj* i'ao âriôut <ualo iu.Mi4k

106313) is also froia Uade County atid mm a body >37 i-^*

loa^ and a tail ^3 oia* loxio, ><ri{;ht and uri^nt (r:/5>7)

cite a Uew Jersey osie with a total lenÛi of 6i^ uu*

The iQsau differences in adult tail loi^ta/totul

Iv^.Aûu A'aCtXo ran^^tjî xnj... uûu'/ uu w«ux<Y (in alo. Â^^Mâtions

U4

•^ SI

I

G

^.

o

6

o

S

ra

+1

;3

.-I -I

*3•«

O -4'

ft

^3

l̂A

+1

U

9

Io1

i:

+1

t

"1

-4

+1 4-1

• •

M D

:i 3

Jo

-J

.1

UO<D CS

O-i :••

•«.

3«•••

<s oa n

•I

> o

O

I!

II ^.8

99

aaXes have a proportionally Iozic:er tail than ttid fomalos).

Kean tail xti.';t,t»ii/tot&l ientith vaiueo rauii© froia 0»313 to

G.37iJ ii* ;uaiQ3., O.-s^^; to 0,35C in f©r?*aea.

AltJQOu^ i'emaiee have proportionally louder mu8Sl«s

than thB jsaalos in £ive of oevon populations in wliici^ ausele

lenu,th/lroutal Ica^Ûi values ara availablo for at laast

four aaulta oi' aaen sojc^ u,'>> ,-.o,.;* , i.or .v.-x po^ûiiiw-. -.,..

flieaaurad are alau>8t identical* In . .. . uiiey raxioc ln>a

0.626 to 0,715 (wean 0,666), iu —^ frou 0#607 to 0,707

(sKian Q,6o9),

Caudodorsal acala rov roduction occurs i;u:>re posterl*

orly in aales tiMui in fetaaloa, Tho reduction fomulaa for

UF 12187*4 » an adult male froci Charleoton County, iouth

Carolina^ and UF 12133, an adult foctale frtxa Hyde County,

North Carolina, ;i^ .r.;nai-it,0(i below,

Ui 12187«4 -J-Tfji^ l$2fHP^ I3p2ikl i^^J^

UF 1^183 1/ef^ i^sifb ^^^^Mr ^^^

OntOoîiotic Variation

in certain norûiations tiio i.Qiiii number of vc'i;trals or

auhcauidaia xn ûyauxxâ ah axûai" tiiaii tât ol aouita, in

it io lowori and in still ot; .t ia identical*

iOO

There is apparently no consistency to this variation.

Newly bom T* aauritua hava proportionally shorter tails

than do the adults (Tablo 6), Cooriicient of oatoôuctxc

divereên^ values are comparable with those obtained for one

population of JC* sroxiatua except in tlto case o£ southern

i^lorida Jteoales waich oxiiibit alioost no diaii^ê in pro|X>r->

tioiiai tail ion^j^th Xrotn birth to aduitiiood (i'able 6)«

leiuaxes ox two broods i.it/u souuiioru î xui - ^ ^

Collier Couatiesi are unique iii liavinb proportioxiaxxy loiiêr

tails than tfia males of tlê satê brood; tho situutioa is

In the Groat i«akes re^îon arul iii nortnvx»atem

peninsular i^'lorida, wstiere adults have either a black or vojry

dark broû dorsi^, tite dorsum oi' Juveniles is approximately

the satae shade oI hrovsi as in juveniles froa areas wtiore

tx^rc is no pronouncuu cox'^.uuxi^ ux wiiu dorsuta with age«

xua,^vxaaai variation

Charaoters wiiich neitrter vary &> cally nor show

sexual or ontogenetic variation are discussed below. Data

on head scales rofer to one side only. Individual variation

in Ateristic cj;aracters is suwiarised in I'able 1«2.

Oculars

iione of tho >^7 !• sauritus exauiined has sore than

one preocular. ioetoculars aro usually 3» with 4 tiie ax>st

frouucxit variation.

101

I

as(4

Oo

CI

u

O

>

I

I«

I

H *O>-C^50{3\

U K

o

-. o

iH HIH '—;

—4-1 —\ '~i -^ -< —4 '-4

;i r-4'^ci\^

iNr«»o t^o t>-^ c\c<«-•H —1 »~| 'H H *"! "^ <H "H

155 .'*'> ;:*<• !>• ^'A 'A •'A I's ;>

1"I

—^ . W^ f-oj —^ -~4) *-«^ —1^ .**l| '""..^

r^ ..^ -O "n "M

J A;4

o

«k * • *

Set

•00 OH

4- O f -^t J

o o

,— V -A •)

Co—. a -* ON* «t *

NOsOJ>C^tOtO

<-f H -*•O Q »4'

—•w* ^ p-f '—

•

N f^ r»> *«» « «» •

Hf*NrA'W

* « *

U

<s

x-l4^

ssrrBw s aiBuiaJ

102

Thoro is U3it\ilr i anterior and 2 posterior teaporals.

there is no population wnich nae * iii^ iiicidenco ol' Tra^

mnntation of* tae ajiterior portion of the aziterior teojporai.

The number of supraiabiaLs varies 4ieoi,rup-»icaily;

tlio iaireûeiit iaterpoaitiou of sumll scales betv«eca d of

the Qonaiu, iûâlabiale does t*oi* vtuy -i^* uîj-u ...«;iiiiiur,

Ten is the usual nucaber, 9 the most irec-aoat v&rXa^

tloa* Xnfralabial number is more variable in aaurituq than

^^ mx?.%inu3 . Toa LicUviduals of sauritus have 2 cioro

IxU'ralabiais on one aide of the hoaf^ than on tiie othor; xw

specioens of Droxiquft nave tuau iuj .Uffeiv.

the «^ sides of the hoad*

Tto is tfa usual oo!idition; 3 io ti»e laost frequent

varjuiLxoii, but j^t ooes not ii&vo a hi{>i ixicidoixce of

occurxronce in any population.

Ventgala

Anoroalous scutes between tite tenoinal ventral and tlie

•aal plate occur iit 9,0 percent of tlie speeiaieus exa;:iiued;

4*6 perce«ut^ irtcludiiit. a few of those just oieabiouedy rtavt

partial scutea ux ou Uiv Mû^ti^t

103

Fusion of 1 or inoro pAirs of subeaudals occurs In

^•^ p«rceat oi' th« aat«riaX Qxaatioed*

ûmk ffttiAt ffyn

Ti» trreat majority of individuaia examined isavo the

typiCiu. xy-j.v-i/' i.i>ri.iUxa; o.a^^ 'c,w,o .-^.^wiJia iiavo u ;a<i.<».uau<a

uuMier of rows iii uxcesd oT 19

«

iieduction takQu place by loss o£ row 4 on both aides

oi' tiie body in 36 of the 5^ apecimeos exaciincu for tuis

clttiracter, and tiie fourth row is involved in the reduction

on one side of Urn body in all but \..... T -c. ^ .iÂu^^

iiuiividusls. Those ti^ree loss the f ii'th row on both aides*

One «aakc dm-s rrm 6 on r.no side of the body.

i'lie rttcJuct--.aa occui-g opjX>3it© voatrals 76 tliroubii i06

(asaa d9*3} in aales» 71 through lOX (oean ^,0} in

female 9, Tjnere appears to b© a rouiiit, positive correlation

betweeu total number of veatrals and poitit of dorsal scale

row reduction, but tho correlation does not seera to bo as

strong as that obs<»xêd xa jJSi^jjtiUiiA*

Presence or absonce of iârietal spots, and si.se iii«&

present, var>' iiidividually as does extent of Uie black ix>8t«

orbxtul vxtta oiid widtn oi tm ULack paravfci u.^^ral strides

wnicn border tiê vertobraL stripe*

104

Gdoâplxic Variation

Ai i X« aaurltuff has a aualler ranô than X*

nrrfittgWilli ^^ nevertheless exhibit 53 a eonsic'orablo orount

of geographic variation, aojuo o£ t,aa âUwaras ox CiCir^Cvor

distribution beiii^ evon more coi^ploji tnan those observed

i« ^roxiaua. As in tho dincrannion of L>rpxiqu? , all

iaeriotic ax;d pro]jorwxo*;iu. duwa t'*"^- boLoM are based

on population aa..i^vie3»

^>M^?^ll^l^tllliJUi

Xhrou^tout peninsular i^lorida to tito vicijiity of

iollaiiaaaee iii txie iât, txio Ukefeiu>keo uwa..^ r&\,j.oa in ths

norti*, aiid t.iroat»ii coaotal Uoorîa nortii to JaCi or County,

i.iouth Caroliiia, 91»o t>«i'cerit of tije snakes navt u au^^ô-

labials on boUi sides of tfie head aîd ^^^ ^.ercent ixavo <i on

one sldo. A fatb variation, tiie presoiice of J supra-

labxala, occura oaiy in oautcu'a i-lorlda* iurouohout tiiO

rest of tae raiiô of tnia aôcioa, only 9,1 percent of the

speduens havs 8 on both sides, Anotiior lO*ii iêrcent ;iave

d on one side, 7 on tliA otiier, l.ost all of tiie regaining

30,1 percent have 7 sitDraiabiaia on both sicleo of the head.

'ii.ia x-û i».^;< uu iiou, aave u on one or uo&n axuea auc all

but one of ttiese aniiaala is from îichi^î.

Ventral if

Variation in nuinber of vex^trals is exeeedin^y

coa^>lcx and can best be visualised by relorence to i^'iûrs

9* jîi soutneuatem 4ûiaiaiui sod adjacunt îlsuxacx^î.

Figiirs 9««N.^eo{>rapliic variation in ventral miabsr ofThsrj;^j2&lJl JBViritr^WI ^ ^^^ Unitciid states and Canada* Black

tes roprosont t>ie areas Srock vdiich a^nplos wore taken*jper nwbars awK>elat«d witli aaeh bio^ " ro the

^... -3 laoan and aaaple 8i«e (in parentnesv . r ciales;

tiie* lower nuabars presoat the oacje data for ioiisaies, Whon

a iii.e appears in place of oither the uppor or i^vrer

nu:^bar8|it aignifias tiiat no specimena of tiiat sex i^represent in the •ac^>le*

10?

Bales haw a mean oi 153.7 veutraXa, fe&ial»s 157,5.

^Qj..^:.^.,-,^-i

i..-, ,,,

,„-:_.. ;.!ouUioaafcera tii82>i»«ippi (uîciiu

ivo of the biioxl aroa) a/id tiio Hobiie Hay««?en»acola Bay

area agree cloaoly, but tite aaaites in sout..? -.am

i4isaia3lppi havo a mean oI 6 to 7 atore vontrais in aalos,

54 in fomalea. from thence riortiwaorthoastword to iiortt»m

Indiana (eiicluslvo ox GexitraJi Toiiiieiaee) tn« maan ni»ib«r

or vcatrals in fmalea U aiaost iiivariablo. The maan for

oalfts increaaea aiishtly ovcar this distance aid reacioa

i (io7,6) in ce:-»tral. indiaoa. in nortliara Indiana

t: oaii ia about 2^ scales loss and it decrsaacs

^raduaUy but eteadUy nortawara iu ..xoîXiî to tîO Straita

of Mackinac, the mean in oxtr«ai© aortliora iiichiran being

reduced by aiiotrisr S scutes. T ,e l atral

eount drops by 3 :;c-,.c;; .o;, ....... :-;:r-w.»ura x;.. _., and

soutnom iîichiiiaiA, then reduces aore 4,radualiy north -./ard

being another 5 scales iesa in extreoe xiortiiem wiciiiân.

A aeries ox ieonales (no iaales avaiiabie) i:roia extreîo

•outaern Ontario agrees well with foBialoa of mrtuoraIndiana, and Uiooo from cûoiUA waio oave otuy aiiîitly

lower laeans. iiortiioaatward fi-om tJ-iese population© ti« aeaanwaber of vcntrais in botii sexes decreases abrujjtly, by

about 4 sealee 4.*; popuxatioas south of Lake iirie, and theo

tapers off gradually in weiitorn Hw Xork and eastern

Ontario. A aeries of seven fotaaies (no aaigs available)

frou U.G vicinity of iîttobuxi,h :^x es cloaely with foaalus

lug

Xrcxa tao ooi'tuonsteni Oliio-îiorthv.oatoru k'QimaylyaziLa arMi

in tuoaa ventral .. \ Ono riundre<! miles to the south, in

eastom ^.«c.w Vircinla, Uuu • r or vcii-ruia xz; ijoth

aoxaa decroasoo abruptly; i arc 7^ acalos less in

femalee. There also anpoars to be a ciinal rsduction h\

veatrai iiu:ôcr xt\>u cciitral uhio into .-ost Viri,iiiia and

a lêntucky* The rcc)uctioa in tiiis case is also

„wiu:'t, a decrease of 11| (fosialea) to 13 (fflaies) taking

place ovor a distance o£ less than «^00 ioilea*

Xno siiakes in central Tennessee have oeans at least

9 scales less than those o£ t..u , w^..xationa to t iu utji. o£

the I'ennesaee iiiver« A series irosi tiie vicinity oi' iiiloxi,

.;is3i3sipî, nas a srtttazx of 5 to 6 £wmr vontrala thsn tîe

^^w-ations ouixouiioa-ic* ^^ <iii*vi *a. .,..^\. ^.i,î 50 aileii

V t« ûaf.es i'roa aoatlioasteru iU,.:.^.. i.^ aîd southern

Geortila (nxlusive of the coast) appear to be siailar to

tao bLLij.tU. .. :. i.3 in i.ioaii Viâtrol couiit, bat moro spocl*

aens are neodud to detur<uinc just iiow «oll tiiey a^reo, and

if the tviw uii vi connected by iA/iyaâuxytiu xii auuuawestem

•'labooa north of i^^bile â^

,

One hundrGd-and-fii'ty ..lilor, o.-.at of the Mobiln iiay—

.

i-exisacoxa uay area in ti.^ .. Laca^xcola .t.AVor vaii.«^y Uô

aean nuinoer of ventrols docro. ^/lan 9 scales*

The populations Just to the north in soutriom Goori;ia have

means leas than 4 scales hi4;ner» A sLiolQ adult foioalo

(UF 73o^} frora ;it, Vixiceat*s Islaixi off tiic moutn of the

Apalaehiofdâ Hlver tms J.61 ventrals, alnoat 13 !^re Utan

til* liean for ii.*e iîâ-taciiAooxa KOiie^ eorioti, i av;.j taoace

•asttiarci and aoutiiuanl in the ii^stem coastal ioulaiids to

southern Florida, imd riorthwruw! in the eastern coastal

iowiaiids to £U>out t.**; x«i.*:. . ji i.-a/t;o... •. , - .. -.a

liuaber of veatroLs la reiaarkabiy coiciotant, r

163.3 to i64«d in males, 159*'^ to 160,7 in foiaaios I

of the west«ceatral Florida and males of tiie east«CQiiurul

Florida ;x> sulations haw êans 1 to *J scales Iggs than those

Just ;ao*ii.aoiieaj« xu x,..c cuiiwrai lix^/ijuuius of Florida and

in ti. -ern coastal lowlands norUi of Daytoaa Beach,

ventral coiuits abruptly decrease, the .-^loana rasiijiag froni

156.4 to 157#6 in males,-.!)3,6 to ii>4*9 iii fo :«^.lcj« Vo».wraA

number falls off suaivl/ to the r/>rth in iiatuid aeor^Îa,

uuu <jii whe coast it varios fnxa the tsean rsAiê stated for

northern i^lorida by less than 1 scute in fenales and lees

than 3 in males (lerrêr aeries of aales will uiidoubtedly

reiiuc-. ./. :• y.-u-ioiic. .-. .. . —jj u.*.vju i*ux*ui*«astern i'''lorida to

central i*ew Jorawy, taeacc iulatid up ti. « side of the

Hudson itiver to tiw vicinity of Albany, Iv. k. On or

adjacent to the i'iedaont iraa ti^c wavanoah iu.v@r to soutii*

eastern FomisyIvania, moan vontral counts aro <^ to 4

scales less Uimi on the outer Atlantic Coastal Plain*

In these inland |:>opulations the nurîber of laean veititrals

iacroasos to tho north, but by leas than 1^ scales in

feaaies, ao£.«una& sioro xa sales (too few loales are

10

Avaliablci Iron tho soutiicrn localities), ...joe popuXatioxis

a£re« rattier well in aean nunbcr or vcxitrula with thoso

locatod in ejctx*€nc nortiiem Coor«îa and eaatem '.eat

ViTiîuia,

On irfon^ iax^t.i'. and &x, 8cattez*cd localities î uow

Xork aast of tne iiudson Hivar a a Tar north as JLake George,

the ventral count in fetnalos rises abnx-itiy to a aooi: of

in Uqv i^'itjjisaad famaloa ore aomawtiat Ic ji, i..,.^:: c ._/j og

not api>ear to be aaaoelat'^'^ '«lth any dinal troi^ .., in

cantraX iiew li^nând uaana lor Taiaaiaa rai^â Troii 1^4*4 to

1^6«1| only tho soutUam U&m > ire Svirii.la lias a

auffloient nutabar o£ laaloa to î. ^^ r^lionco m^ uho

reaultirti; niaan (i*e«t 1^3«0], Tliu:;^ tiicac populat

a alit^t inrrnqfi 5.n ^mtral number o'^v the relatively

stable Atl«i««MXc ouattUal Plain sarioai but xî.â tx*oud is

.ly roveraad in aouthaastarn îassachuactts^ ilainei and

Nova Scotia^ utiere the xaaAoa aro botwaan 15^»1 and 154*0

in laalea, 149*3 and XîiÔ in foaaloa*

Variation in nubcaudal nu'r^ber apT>eara to ba alaoat

cnoouxc liiiiiu vxoNuu xxi boi'ius ox xuc ...î-uuM^ tuid a

aeaniit4>'ul evaluation ia furtiior ku <i by aavi;i^ cio Tow

individuals with a complete tail in t ..ority oi U)»

aarias exaininuii* a âiiorui.^ ^Mitwuni côi Lo cix^ d,

but aaa iri4.ure XO lor dotax. • on subo«udal counts in

Figure X0*«»«-Oe<^raphic variatJuoa in subeaudaX nuaiberof yi^apuQopHia Miil^Ssiia ^^ tiio Uaitotl Jtate© and taiuwia.i^Xack bl ly UjiO areas Ir Ich samples tmretaken. .a associated ..— aaai blotch ore th«MBplo i ize {'hi |>areatiie903) lor aaies; tl^a

lower xiu cjOKin data lor laoalaa* aiific .-

,._ ... or tiv^ 2r or lower .*,.».-...oro,

it si. . t iio - .J ol V, . jox were pi'eoeut intha 6

113

iA AAlea almoct always excoed 120 in tho area south of the

Ohio silver on tûu wesu^ suai as Xax* lioixu an so uuiteasiiom

i'eîiisylvaiilay northcxu how Jersey, and possibly Coimecticut

on the east, Witîin tills area Unit immnA exceed 1^1^ iii

ua^teru ¥lre>liaia| coastal ueor^la, %tm aouthem four-

riX'ths or pt}ainaular i'loridai the l'k>bile i>ay.>«.PeasaGola tiaf

area (tiie highest Bean, 131*^ i occtirs nere)^ southweatem

Missinsippiy and nortnern iuisDlosi^ pl« Tiie mean ntsnbor •

docreases to the itorth, males frota tjorrulations in northern

xfmidaa, i-iic. _ , uaârio , :. . .-.:. uot riavAiit, over IX*^^

Feaales from this northern area have loss thaoi 105 £i@an

ventral9« Xlio mean number 1:. . ...ales increases southward

»

rai-ely beiai^ iesj than 112 souUi of liew h jpe, south-

asstem liev. liork, Pennsylvania , Ohio, aad central Indiana,

The jaean exceeds 120 in feiaales fraoi nortlieastera Florida,

southeastern Florida, wost'Ceutral Florida {Una hlGhest

Asan, lii^.3, occjts here), Levy County, Florida, the

Uelativu tail lei^^^th is iairly constant in tno area

lyin^, south of tiw Ohio River, and in southern Pennsylvania,

êw Jersey, &iuj gtoutneastem liev^ Xork, Means of tail

len^tii/total leij^^th values in males usually fall bstwesn

0.345 and 0,355, in feiiuU.e^ . : n 0.335 and 0.345.

Means are In excess of 0*360 in jaales from central

Tennsas<ie and the mid.Savannnli River area, and as hi^ as

U.J.tl

0.376 in a aemii aerios TroK casuorn vest vii\ . .. » xn

UiTOQ Ubv Pixijând oeries th« nean for aaico la 0*343 as

comnnrcd t,3 0,3 52 In Nat; Jnrnr^yj in foroaieo it lias

CGC* iro« J«^4*i to ;;,>>u. > ; oa« male avaiiobie

frora Kova i>cotia U- -i> uunuabcrGt: ) has a value of 0<,310«

Males from wester^ h«' York, northweoteriri Pennsylvania,

Oiriio, cutd soutiieni iiitixa.^ ve ueatis rar^jîiit, fraa 0,3*i7

to 0*336; means in feaaica raaê from u»3l7 to 0»31V» 1"

northern Indiana and Michi^^v^. i..ean8 in loales range fxvn

0«310 to 0*31^, in feeiales froru î^^9^J to 0«303. in oastom

Ontario ti;o mean for females is G,312,

Tije vertebral stripo is frequently poorly devolopod

or lacking in t«ie anakes initabitino poainaular ilorida and

southeastern Cieor£;ia« .^jî^v ûu iiî.i.^nj in this area io tan

in color, often only a shade lighter tiiân tho color of the

dorsiKi, the loss of tlie atrijje is brotiîit about inorcly by

oxoi-uaation oi 6..u . r.-o./, blacK >r«jQra ol" tiiO vertebral

stri. • There is alistoat alwaiys at loaet a faiixt indication

of a stripe for a siiort diatanoe iflamediately in back of the

hoad* fhrou^.(Out the rest of tlie ranê Uie vortebrul

stripe is always present and distinctly taarîncd by Uie

black paravertebral stripes altitoui^ t.>s. matter uiay vary

greatly in width. In the Great Lakes recioa, \;iiero tiie

doraun is usucMly black or nearly blacK, tiic stri^.e is

often uarro'iitiiir umu xhmqx^ ,^ . . . - uxo

115

wldtl) of each p«rair«rtebraX row of ecalea (in additioti, of

coarse, to all of UnQ vertobral row). Tiic voitobral stripe

^ '!• nroiltoill ^0 ^8o fx^qusntly narxxûor In tho aortiiem

part of tho rcuiige*

In p«ain8ular i: lorida and ooutheasteni Georgia ttm

lateral stripe is narrow, always occupyiz^: iosn than half

of oao ol Ui@ t^«o rows ou vtijicii ^t xs 3itu£.tcd9 aacl

usually leas tlian !mlf of botn rows, lu tm reiîoa of tlie

Grtjcit i<ake8, dorsal scale rov^ 2 la frequently izivolvod in

tm lat&rul &trip<2 to a »iJ4)iit (l64;,ree on the auterior por«

tioii of tlie body in adaitioa to tlê xioriual 3 and 4f tiius

parallelint^ Uie condition typical of Xiiaanoptâ j|UiSui^£â «

epecies wliicii ianabits part of tsila »&a» area* tiia con*

dition cannot be aaid to typify eteobers of tho populations

in tliia re^îou, bat i.x, oooa uave a aucn iUi,i^r ^ru.^^-. c^ of

occurronce ki&v& tnan elsewhere in ttie ranê of tiie spociea*

Animals with an aateriorly widerted lateral stripe iiavo also

been observed in samplea i'roia central Xenruôacon ©astern

West Virjjinia, and w^sterxi i^Gims/lvaiiia, wiierc tiiey are

not associated with a black dorsum* CA3 156^0 frou Mut

Vir^îiiifi and iUMi 30^0^ fix>m i'tova Scotia have rows 3t 4»

axid 5 incorporated in tlis lateral striiê*

Tiie dorsum is tan or brown in ponlnsular ilorida eaud

coastal uoor£,ia, except in western i«ovy Cou:ity, I'lorida,

and tîence northwestward aloii^; the coast to a point altaost

..6

UBuaiiy « very dark: oroviâ to biacK in color. It is also

biacK or very dark brov.^* in «ova ;^cotia, .^laino, o.;jf the

urw.û ^.....' 1 <.:îoiia w>oaUiV.îiu tiâxtUoiiouu i;.*b x'v^wU ux tao

raitc^d («i>wcept ;ûuliiiiular iiorioa atic ooastai Cioortîai ti2«

dorsum ia a r.<^' i-a brf»'"'n,

lii riiuuoub.. Irou i^caiuouiar iîorxda axxi iôutu-

uor-.i Lioor<îa ttio veitubrai atri^ iu usually soua uiiade

v.'^ V ..., jccaoionaiJL^ ..^w.4 c. iUtaXIic luster, or in iacklii^*

Out£«X(ju witia are<^ it ia udually aooio di'âdo o£ ycilov.,

occaaioaolly overlain with dark pi<y;iont in tliO uruat itokas

reîoa.

Toe lateral striô Is yollo\j ttu'ouîout aoat of the

ran^jO of the spocies; it is occasional Ly white at Qcattcrod

localities on tne i^'lorida poniiiciula • la w . i«uvy

County 3mA in tho coastal area to trio nort.rnuj^t, tiie

latoral stripe xw <,v>uviî.^ u a.A,^4i\, oxuu .;^ oluish white*

Geo^ra^ûic variation is Tor tijo laoat ^>art o£ tue uia-

corriaiit tjrre dinciâsod ..rcviously in conjuxictior* v/iLii j^,

^..ixj^w^g.ô. ^j^m Wait at>wci.aw, ^. a^j^^rxuaa aas twd oajor

zu.'is,^ ol coucordaiit cliaracter transition, i..iore is an

abru;jt ciMi^^w ^.< supralabiai uuiuber, latoral stri^ widtu,

vartuoral otriô color, ai^:: also a slioi^t cxiaii^â in color

ol the Uoroum batwe^m the peniiisular Moriii^4.-«a0i4tiioautarn

CiSOro''^^ aJk\,(( aJA.1 b.iw *.'<.j^XKjn wJ ui.t; nO(t«h ciiwa WOUt* Xhifl

ii7

traooition may b* rolated to ttie X&ct th«fc oiost o£

p%nLiiuuj.'Ai viorlda is believed to iiave been iaolatod rrom

tlie ttaiiiXmid, ia tne Torm o£ islandOi during; ohiXts in tue

XsvQl or rloistocezMs seas«

ikiQ oUif^ trai^sitiou aoxta appear&i to be associated to

a certain dei^reo with the eouthem iicnita or i Xoi&tocoxie

gXaciatiorty the anicials occupying the previously glaciated

area having a darker dorau&i^ a reiativoly 5huxn;6r tailn

arid iTewer aiibcaudais than are found in the populatiorta

isouU'i of tiic fcât:^ - . ,, , »'" olatiohshlp ustjoks

down in northern rêw Jeraey, xork, and aouthem

êw Sngland wiiore ioriij-taj. brotjn dorniua

&ud aXtJx aui)Cii.\Mial ooux^tG riuhM v^ll iiito «yLacxatoa

territory*

The elcsaents of color pattern have less observable

Ôi^rapiiic variability iin j^« aauritua that* doe* the uuiaber

o£ veatrala or ûbcaudals,

iiio ^^jiâtiou saEsples exaiQUa.iuu u^pciai^ to repareaent

four (oajor populations which replace each otner 4ê0ijraphi*

cally* These are defined on the basis of riifforeaces

ciiiefly iii color pattern^ but ai£io to souie i .. o in

scutellation and proportioas* The salient features of

these populations are discussed below witiîn the framework

of their forctal description as subspecies*

u$

Tha foliov.-in, ::o>' for diatin. uia'-iint^ adultj of UiO

Hour r»cî> <ii. j^, j, was (idui^^^Mci |>x*i;ua:':iJ.y lur aso

. -îi iiv<3 .'uateriad.* i£ ooly pjresorvod a^.aci^touc are

available, attention saould bo paid to êoâphic provenance

of tae saiaplo*

Kay to tho cSubspocioa of Xhai.y aauritua

1, ;5upraiabiaio 7, occanionaiiy 8; vertebral

ai,CXô so.ô Siittcie oi ôxôvv; xai-ux-x aiî-i^-o

tiot narrow, occui^yiiit, i/i^ or iaor::> oi both

rows 3 and 4 '^

L*

,

W.U, rcuaiiiala vi, rar«xy 7; Vtiz*tut>rai sLr^.^^©

taa, or lacking, lateral stripo uox'i-ow,

occupyiat^ lo»6 Uiaii i/<J of oitaer rov. 3 or

4, or boUi •••.••••••.••.•••• 3

«i{i), LorBiaii reddiaa brown; iaueral atripo rarely

;.j.i.Uiiu(, aiitoi'xorx^ ; uu.ju. tvx.-.:uxvui^ i^J'»o

( . .ui >.ail iOii^t.Vô^^ lontitu usually

e.\xe<s<:iiii^ 0*340 in raaloa, 0*330 in

leiualoni J* i2* aaurxl.^ V,;. x.9

2»(lj* -.5u« voxvety black or very dax'k biovvu;

ia*wciax »iv.rip« froquantly widened anteriorly,

involviiue, row 2; tail rolativoly abort

(aMsan tail loutpth/total ien.^th usually

not u .( - ccin^j 0*33^ ii^ iaaxoa, j^^'-u In

fez^.^c..; X. 1. ffi?r<vn"vr4gm>Ug (p. i2i)

311* }• Doroua tan or Dxx>wji, rai^j.^- *v^*j» dark;

lateral atrip© yellow, occiioioiiaily

white X. 1. Aftsioai (p. 12^^

3«(i»)« Iôrsuii very dark browi or bxaac; lateral

stri e bluish i*iito or liîftt blue, occasioiiaUy

vu.;,o I. A. liiias IP. i3Q)

Account* 0.1 iiub9p«cie»

fnfiWTB^^* ftaurltuo ftfturitua (idmiaous)

Hlffl^nAphia ûritua aaurltug rtutaven, 190^1 112 [by

Ixiplication]

!i«otYpO > The data presented for the species (p» 92)

api^ly aere»

Deiinitioa. A subspecies or i^^... ....:.-i &,:.^.£s-^

©haracteriacd by 7 supralabials, a reddia.. brov>u dorsiâ, a

golde.i yvii-ow vertebral stripe, a.-.> ^. * iy loJ**;. taii.

tiiixi^jii . * z:^ fioalaate race rant.os l- .»taerii ^.ow

£afclai-4d, aast«rn lôv. Xork, imw .ier»«y, mw oasterii

i-ewasylvaiua, iv^û uc ;.,,rj....i ôi&iiom uhio aiid aoutaem

indiaxw* to tite iîssi^-x... 1 .iiver, souta to taa uort.ieni

edi>e of wike iônchartrain on t-iE wast aiid to nortri>«3J?tern

iôrida aiid Oeor4,ia (excluti*v*3 ol w . < -i^* w.^-iO w*

the state) on tim east*

. .-'iption ft£ aaatXCa. acutellatlou is «s follows:

oculars 1+3, teiaporals 1+3, aur^ralabialo 7, iiuralabials

%)ata on ocuteUation and proportions are f«>ia :>Ciwaict and

Conant (1956-57).

.20

iOy dor.î-il 3c;Uo roKS X9-19-X7t varitraXs 156| subcaudais

113« isody ionf^tii *, ^^w . , , tail le;.. . . îOii ran,, tail

ioiiota/totai ifiii^th ratio Q»^^^m3'^ i^ carrle. to axiotner

dociuiaJL j^lacoj •

Xoaaaucii ^^ .m..i..î.;i^ .^^w: Coaant*s (X9>i6-57) descrip*

tiou oX ttio oeotyps is based ou a proôrved t^ ju, tlie

XoiiowXx% coioi" notes are ; resentod Tor Ui X2X37-X, axi

au>Aj.t oaXd coxxecbt;u ju*;; avuxôx, uîa^'XwUwo:* uoiUi&yf

ôuta C&roXXiiaf by i«iita J« tioai>faaii aud iôut^«uS A, . nt

dorsum rocidish brovoi (Macrs asv' ' 'uX X4iiX2), a saado

Xii^xter beXow the XataraX etri^je; vertobraX stri^vo t,oXcien

yeXXow (X2Jk); XatcraX stripo Xi^Jit vqXXow (XûX); venter

creaui (9^'l)i >u.aii^ voatrojuuw«i"e<i t»ui.*.^'<; uiuaij wtvu^^Â.i^,

1/4 tne viidth of a veatr;iX scute on eacli sXna; c;iiii jaXoXds

and i:ifr':inbiaic wlîitc, r;ii. ralabiala ycIXovdali white, The

Xxât postocuxar spot 'xu co.îxiôc to tas xovôr pOw;,ocu*.cjr;

tno bXack postorbitaX vxtta ia bxx>ad« Tao tiny parietaX

spots are separato and yoXXow, mar^î.uQ<i iiarrowXy 'with

bXacjr:,

Discuaaion . tioristic aiid proport XoaaX ciiaructer.

istics of ttM fiMBinate race ^nj be ouiauariottd as folXoua:

^ to 4 postocuXara; X+X to 4^-1-3 te;a^raXSi o to 3 supra-

XabXaXa; d to XX irdroiabiaiu; X to 4 intoriôiixaXo; X7«-X<i.

x/ uo x>'-x>-x/ ciorsax acaxo j(X>v<u^ . *.i*^-.x^JO, x<, ... u*

X43-Xo> VtiiitraXs; maXss XoO-Xû, X >o-X30 subcaudaXa,

i'iaXen rr.x\Q I'ro:.-. l^L^f> to /4.5X ;.«;l, ia body Xoii^tii, X85 to

1^1

665 «ni» in total lon^th: I'ûlt tail length /total leii^th

ratio 0.333 to 0«3<33« ire ^ i^^ to 636 am^

iii body leuj^th^ 1^0 to 956 .iiu* in total length; adult tall

lengtn/total l^r^^^th ratio 0.317 to U.3^3*

My coacept oi tlie color pattom in !• 5» JK^ i i»

based on livii\, Bpec'L-ionB from Caw . . >3C'X, iUid ivarrott

Charlestozi Coiuit/i .ôuth Oarolinai tiurbour County, .^vlaba^â;

and Forrest County, Misrinnippl, n& i;nli rn n colvnrtu-^ slid«

ol a 8nak» iroa ilurliii^ôu Courity, «' v, iUMi color

notes on one froiu lolland County, Comiecticut (tho slide

aîd color notes vwro contributed by sioboin, tt, Urant, Jr.,

and iiaia ^W8«t, reopoctively)* Into. ;.ion betwcou j^» jg«

Bg|ur;j^tfHia and 1. j. MjS&SHkt î^ between jmM^%m and

flggtfffttfrAffftftJLAflf iQ di?.cur,sod l.ucr.

yhJUaOTi^hi^fl «ftttr4fcÂa Mii&&m£Mi^ÛkM9^ »«v subspecies

ttolotvpe , UJc' X^l7<)mti^ an adult female Troui iiiciiioo<3b

Hollow, near Ithaca, Toapklns County, Uew York, collected

<^0 ii4ay 1959 ))y ^ôhn A* hartley •

Prf4nJ^tto« A subopocies of 1m, ^^ . ^-JL ftiViTAfeVM

cliaracterised by 7 8Uijralabi<4j.u, a velvety blade or very

dark brown doraua, a yellow vertebral stripe, often over*

lain with brown pigment, axui a relatively siiort tail.

-^Kefers to t,i» nortnern distribution of tiiis race inrelation to tno ran^ê of the species.

U2

and central -aaro luay b« aa iaoXated

coioay in n,- rti'jo~ir;t;nr,i .iscoasln (see Ciscuasioii)*

ocuiara 1+3 » vj.^. .^^-aia 1+^, Oi^^^ralabiala 7t i.'Jrralav>ivJ.o

10, iatertiiaaiale ~, .orsal scale rowi^ *y-^^— ,, ^....w* a

liû, sub.. .• i<oU>' Ôii^Ût is >74 iAU», tail lQUc>^h

4i71 iit;i,, tail leiiLtfi/total ier^^tii ratio U,3*il.

tiio body, vor/ dark &rov.a posteriorly; bolow t.iO lateral

Sori::« t.'io clori:uj3 in dorl: bro:-n (..acrs .inrf Pa»il 7ii:i2}«

vu^'tuwiuuL wwripu *0 yclxun, wtAw oO i^&dVily ovorloiii

vfitli brovm that it is im^.>03..iuie to dotcrîine X,ii& ci^ct

s-îadej ./ u iut«r»* v>«ri|jO i^ .. ij.j^.tw j^^ûtt (I0J1)» The

v«.>iit6r is palo ijmen suTraêd tarou,>u><it witli taa; tac dark

vontrolatcrai atriDe is broad, occusyiri; 1/5 tac width of a

voiiiri--aA ou eucii .>.. , *..'.. cuiii aaxcxca oiiC iii^r . .--._:;

are Wiite, txie supralabic..j .^ pale ^.roenish vtnite* i'he

head is black, hence ra> istorbital vitta is visible; tne

parxtit«l apota are laciiiiô*

biaftLfai^.w.. . i.iOri.ttLic oiid proportional c^âr&cter.

istioe of tills aubapeeiea are aa follow;;. ^ wv. ^ ôst-

oculura; 1+1 to <^+3 tui..^,oraia; 6 to u aui^ralubialc; 7 to

11 infralabiala; 1 to 3 i»itor)êuiaia; lu*l^»17 to l^-«il*i<i

1^

uora;ai seal© row*; aaieo iîi-177t iQijaiee 4..,..-ijy ventral*;

ouiiios 9(5<»X^3« xauudea 94*114 aubcaudals* iîaieii raa4.e farom

1^5,5 to 436 aisi. la body loq^th, 179 to 55^ ru^, in total

iîiÛi; adult tail lorit»Ui/total ioii^tii rati.0 j.^>o to u,333«

Feeialea ranê JCroa 12S to 574 sm« iu body l&ik^th, li}4 to

34^ -»• j-î -otai ierifcth; tail length/total length ratio

Q^dHS to 0.335.

in defiairi^ 1. a. Bffip,tt?y»^r4miMtlr,a «yi<^ i" attempting

to aeiJ^xw xt.i rai:4,e, ^ .âvts aad ac€«&a uu x^vj^îi^ ri^boii-

aiia&ea frou itoiuroe and Xompkiua Coui^tios, £êw Xork;

Itniiabuii CouiJty , Unio; ililiadalc aiid IVaahtonaw Counties,

Mictiiiiaii; ai'id ^'larouaix Couxity, Aii^xiUAi., ...^ —- -i so i. -..-;

slidos ol tMo live iodividualo I'roru Hoacy iiarbour, uatario^

atid color notes oa a aorios o£ apecimena Troâ tioiKloau

Proviiicial ia^*^, Ontario iliio sixdaa and uotaa wera

geoarousiy ^,;rovidacatin^ the s»nc oi' intori^radation bct>ên

^8P\w^vg;i^ffinaA4a ^^d anwy^tytB ^y o-ui..iiUitaoa ox i.ri..;...vud

apeciffifsna is mada dlTi'icult by tao iact tiiat wimt .. a

black dorsum in lUto oay beooâ a dark brovai dorsuQ after

aevoral yaars 8torai,a in proservative, particularly if the

aniraal was a; proacJiinr. ecrSyoia at XAie tiaie of ita preaerva*

i*jLoa# t.Qvaruuexuaa, uy cauu-ijL>uLia.> uAuaoiiit. color âmorod

fron preaarved têiamm and by utilisii^ relative tail

Jl - — _^

1^4

Xengtii values and tao color data available £ro.i live

tavion will be posiîble v&ion ux>r(» live apeciio&as or color

Tno norUiorn IndiaAa ^ôpulatioa dearly boloîû to

aepteftCriotialifl . the ceatr;il Indiana snakes a^poai* to be

close to t:tat suuUj^.ucû;^ iuûîouta,^! i«iiu. .^y possibly bo

iri'w .) ^ and the series Trom soutiiweatora xadXcuiSi^^

ooutiii^ ..."J iiiiiiois citnar r^nrcj^cnt-n sauritus ii

aeuttot/w^- ii.uui't,rv..UQ;i or ûi'*i.v>^ . ivô aUuit .u;-j.0S

. ,>>, lJd09^) iroiii tîe i'luucatatuckc Aiver iii souUi*

eastern Indiaita near the facial boundary apâr to be

qauritv>>3 . - o asoiW oi' iiitort^rauatioxi i^xxjoaJly 3

t-troiAâ c<nitr;u. Uiio, The color , ittern o£ a live ad^^t

JTeutw^'s *i\ti^ Mu.ufwJui^ v/wuikUj ^t: ^4..^ uitXo is typical

oX* saurituS y but the sxiake al:» relatively mart

tail, O'oii 75^3 fro;;! Jackson Coimty in tiîe yoxfi.stim ; r.rt ol

O..J ..A/Ute la ceiiiiAtely ;. .m. ;.-.:rJi--_£« "Oi-^.f ..-,;.v* uaxo

ribboaaiiakes are mecabezâ o£ tne nortiiera raco, a;id so pre*

sumably aru the ^^.w, nations in adjacent Pennsylvania

altiiou«î t.uoir s'âtuu is not clearly urulorstood. .^bbon-

snakes from the vicinity of Pittoburi;;h probably are i;iter»

^rades* ,.xa. ouu ..est Vx^-^x.^â animals examined a^^.>.c<i to

be aam^tuB . witii tiie possible exception ol Oil H^l^J Troai

hrcston County ;«iicii apjjeara to uave a black ,

although ail othsr dnar&fXQm a^ ree with tne nominate forra.

^n4,iiu:ui ribboiisoakss all appear to b« naunltua, but tiiO

southern »ew Hanapfîro aatvie aay bo reorosoata.ivGa o

inter4>rada Ôiûiatlon, i 4*av« Aaciw^tii-ja t..:c ..- ..-^--. ...i'^'-a

jTroit) Molfio in aautentri/inAllg boca.iBO it ^£i tiiat tiiesa

aoifflais hbAj tiavo iiaci a black dormia in USq^ arid at least

oxm o£ thoa l-HC^ 699) shoxm mitorior wideniivi of tt« lateral

stripe* Thin rôuiation ooena to ba disjunct froai tlio main

body of ti.e ra»ib^ or aQ^&orit.r '|,i,^'|[|fti|ffi,bit i^îe literature

records for Essax arMJl Franlclixi Count!og, i»ow York (vcbor,

1926, and ^vanaann, i91cJ» respwctivelyi, probably are b-nod

on aiambera of this race, aiid futurt colloctinjj in c^-.-cl

Maw . iiro and coatrai Veirwnt may woll cloao ti •

BXaakaay (iv5i) notaa tiiat ti.^ ^, aauritus Uc coUoctCN x..

liova »icotia were black, witli yellow atripaa. :- .

stjQCletBn of tiiat series which i-ios a complete tail ii*;it^'i^

uxiGUiioeroci^ uas* Wie xow toix ioiit,4*.v ^^ •-. .. .vu^va voiuo

eharacteristic of X« Jft* aii^JitSa î^-^s populfition

is urKueatloiiably a rolictj ti*'^ r.i _ tiio dia»

JuîCtiou ia diacuOGcd by kiXiajtiimnj il>>oi»

liickiusoii (19^) reports tiiat Am &• i>uzuki observad

a ribbonanaka at baaa lake, i^ârinetta County , WiscoaDin,

but tiiat tna apocinen h&b not j.raaorvad* têvortaalesa,

there ia a apcciawn of X. yayritua in the Kilivau-.o ; uulic

*.uS'/ur;i co^,. c.u. l<i«:9) wnica is xj.stec au .L^yâ^ been

126

colXoctad by i in > X.V.Q County at Baao lAko»

-3 cojifuoion, the opoci'ien ,%;reea fairly v.'ell

with i..ii,Ciiiîui ;jv,^'o j»;.,„ '^ ... ^j. ji^ jî uuvu-k-a oi. JCj.«oxa.oit*xoa,

and xow 2 is cluarXy involved in tho lataral atriô on t.ê

anterior -v^rt of t-is body, Honcc it la .aito ntiior posfiible

u»ai; aii -aoâuoG coloiiy ol ..^-.^..m- ..,..._,--... o-.d.3t in

nortii. coaaiii aiiU tao u^.^.or .."..aiiiaula of iuiciu^ûi,

TftmilWRfli^n aauritup a^ckcnfL (&oni.icotti

TttflmBftifilifi A£UiC4tua var. Mfiimfii^^^ hxa&erll aiid Uoeourt, loV3:

759

[yhfyiffffln!?i^? aauritua aackaoi hutiivon, IÛB: 10?

irnntfWRhJH! Bftijjrltfctffl aafitoflU ^j.^^^c.uivu, xj^-ji iq

tiolotv>>c « Th<d iioloty^Xj, witich was ooli.ectucl in

i'lnrifin by iiaron Osten :iacken, is ap;)arGiitiy lost. Dr,

Loria fit Cocaran nas inCortued ae taat OwîiUi 45C1 (now lost)

is listed in tho U«3« Natioial I-iuaeua catalocûta ao a type

or ii;uta»niA flackBnii, <» This 0|>eciiaen probAbiy oliould be

considered a paratype inaaauch as the original doGcrii>tion

wa:;; based on oiic or nor© spoel'aena in tlio coll«sctioa of

...;-w.. . \ôrii uiixveraxtj , .- -c :^/v*a oui.y relui'c.-cc to tue

o..u,:i'. -..c... oi .. . itioûil aaterial in tiio U«*>« Ittational

i%uOt-:i*. co-.tai..ef* i:i r. focstnoto, aiid it is i»ut certain

w.ioUiur tie cot43^ . .'i-ai. to bu a I'û^t oi t.jc ty^ê

0014.(3 a • > ucoty^ nao not been iiateo bccauao the

oriî4ttl do;*cription (Keniticott, *w^^, and tiie tyie

locality aaenUiA^t^ly oorve to a^ôci«ito tito aa..o to i

127

8p«ciric population* Sciimidt (19^3) restricts Uie Wi'»

locality to Uainoisvilio, Alaciiua County ^ jrlorida, and I ««•

DO raauou to dispute this rf^strictioxi,

elujracteriz<.>d by $ supralabiols, a ta^. to bvovsa doroun^ a

tan vertebrnJ. atirixs, often tdth a . otaliic luster, a

xiavixr.: L..^. .:. -..-, . .-.\.,. • ,t Tdi:'- - Loiii, tail,

tta>v.o . X;us raco is diatributecl tixouôut p©ni«»

auiar Florida (exclusive of wosteru Le^fy Couxity aiid tiie

coastal aroa to tite nortiiwost oi it) west to tam viciîity

or Xalla<iaa8«ft, aiid ix>rth throuîi tha Ukaraxtokea c»waap aiid

coaatai tiaor^yia to «Jas,;Gr Count;-, wôutii Caroliiiiaa Appar*

ontly thore ia ari iaolatad colony oii 'Jt» Viacciit*s lîlanil

juot *#ust oi' ttue uouth of tiae Apalacuicoia Hiver*

JTollowiii^ deacriptioa xb bas&U oa uF 1^:14^4^, ajii adult

iTemale froa ^J ailea i; Moss iJluTf , ;-iarion Cotjitty, i'iorida,

coiloctod by lUiirrod T, êiii a^td cjasa *i» ielTord, Jr«

iicutoiiation is as follows; ocn^lars 143, temiôrals 1+2,

supralabials 6, irifralabials i^, u.awui\ôniai.ij i, uorsal

seals rows 19-l^>«i7, voatrals 151, subcaud.aa il4, iiody

leagth is 456 ani. , tail ionrth xi*i3 a;ri., tail iangth/total

lso£l«a ratio u*3^7*

Xbs dorsum is brov<n, .:) vertebral strips is

ostaJLlic tan, the lateral strips j^>al« yellow and narrow,

title stride occu^^yia^, aliiîtly lesa taaii l/d tiie vddth of a

seal® on rows 3 and 4* •iâe vauter is âlo ^rueuish yellow.

i28

pals i^roen oa taa tail; tae dark voxitrolateral sUri^^ is

bro d, occupying sii^îtiy .aoro tasm i/4 the ;;ic![l,h of a

vo;iC>r<jkX acute ou eaCii axde. i.., cîXii suxui'..^ â. xo^xtr

paz*t oX tîe iiiiraiabiais arc wiiitoi ti«0 su^âiablaia aiui

upper part of tiio i/irrai.abiai» i>&jjs yollou» Tma li&r xot.al

spots aro iackine>»

Diacuaaifta ^ ..jristic aiid proportional cluiractaris-*

tics of this suba^/ecieo caay bo sunnarlsod ao followa: ^ to

5 pootocuiarn; 1+ii to ^+3 teaporais; 7 to 9 ;. . lis;

•^ to 12 irif'raiabinls; G to 3 liitor^'.oniais; 1J-.17-17 to

1V-1>-1V Uoraai scale rov.a; ^.. .^•>-xyii., -i : l^yo-loo

vexitrais; uiaiea 116-134 1 f' 105-131 . *-aie8

raiu.o froa l/»4 to i>37 nii., *" -^ iSiiiith, 215 to S20 mra.

ill total leiH,th; adult tail lo4i^tii/total leii<^th ratio Q«3'^<<

to 0»3.'-, * v;.;al9o ran;,e rrw;i 137 -o 070 uui, iii body

xui^ô.i, liu? to i.ui>^ i...i« j.ii cuoiju. j.ui.4^th; adult tail length/

total leij^th ratio 0»317 to 0.364.

X iiavt; socsji liviuj,, oxcuapics of t.uo cjLn; ccieo froa

Aiacauu, ^v^M^^Cf ttrowaru, coiix«ir| i.uae, Luvî.« i'<uic.i7|

Lake, itevy, naiiatoe, «ârloa, i'ôrtin, axui ruiii uoadi

CountxcSf i^lorida, and a colorsd alide of o/ie frora

Caarltoii Cooiity, Uoorîa, provL-oc! uy uarry uit ...Ji'.ia

tiot.'iinan, X tiavo also exaaiiiec u liviiii, apocx.,cii (iio Xoinêr

A..4.iciux«i **v-i tjuu- iij.«i<^'v.A.i*iiiA, Jasper County ^ ûth

Carolitia, which is oi. k or a aaurxtua x jBftfiiUUiii

intermdo, . .; vert iatorcO, atri.TGs of tixis

aulu<U. ar« uot \jl-x^Jîl,xy c _ . , -iu uao cu*-.. .-"'-n iyore of

129

a tan than a reddish brovai, all o£ taooe itotao i>eix«i, more

characteristic of aa^cpj . On tiie otlier haad, the anaka

lias 7 »Ui,raj.auAUig on boUi aidos of tho hoacJ, a condition

found in iflSJteB&Jl oniy on the poriphoiry of tiie rone,© wUwe

ganatic influence from aaurituo would be exrec.- . i:urtii«r-

aore, tae coior oi v-.e vertcbroju swfjê xs liou u.îuctar-

iatic of Uiat usuaiiy found in oititei' fora, and tiio lateral

atripa ia not ospeciaily narrow. Another Jaapar QouxAy

specimen ioii X^ldo^ frois Ui*s wava*i.A*itA i»iii,io.ic:i îdiifa

^fu^3) ii&Q X,Lie uarro-fj iatarai atrxô typical of aaek^j.*i

.

and tne vorttbrai stripa alao appears closer to ti» tyjâ

found in t.tat raco* Tii» dorsum, attar atoraô in alooliol;

for a yaar, appaars to be a ii./it recklish broiai* iti.:Kiciiaena

fnwB ooaas>ax uoor^xa, <j.qj..,xo. ... .îû xi«cxuo.ico of 7 ou^.*a-

labiala^ appear to ba fairly ty^^lcal of sackeui^ T.. c,

ffêly^ni^

apparently extends aiontr. ti*e coast as far north

aa tiia ijavonnai.* iUvar, intorgradir»tj witi* iiiLî- .-j-.-i.-,.•^'

•xtrame aoutharri iiouth Carolina*

iiixuiiu, a ôriea from iûw <-»k®foiK)koa Jitiaiap ia, i

believo, urûcstiosiabiy aacketai ia cpite of frcjuont occiir-

renee of 7 su-.rn.iabiala. iêill 11954) e;s^:rres.'3e3 tne

opinion tiitit .-, .-. . ...u .._.. ....: xar xiiifiuic iis i,.^<ju.iuui c.ouuU>'

,

Cioor^^j-a , .x.i ijarrimill County , ;iouth Caroiiiia, are cloaor to

MS^SiSSLSk ^t^î to aauriuuBf and that aav^fa><^ti#. do^s not occur

below tiia iâil i<ine ii::^ aaatorn boor^îa; i xiava not a&an

apadiaaaa frou iôanuol County* i>ut x fiKid tne asi:iiA& froffi

130

Just below the k'nil iiiiie la eastorn Georgia moA ac.

oouth Caroiim* to txj fairly typical o£ saûjfituq .

Tv<o uvuiv 4.0 v<"-^-- 4.j6234-.ii35) Trora iiak«

;Aicco3ukett, Uofrerson County, Florida, aro aackaai f aiUiout^h

oao of thesi har> 7 supralabiala or; both sides of tivj nead»

iaey rcjji'uui^<.ii. w..c viosx^criiiiiost iLOcaixty ai. vaixci Lius race

has been taicen, '^ for Ui 73o>, an adult i'coala frx)U

5t, Vincont'js Inl.-iv? i;. i'rn:;;:tin County, i'înr-Lr'n, i';.is

'..u.. OJ^ C :i^XiiX\tlsiJ^jf iA^j^/iiik*. iii t»'. _, , \:k$ U<^t CU.! t»4l

specie A..i;j ..ulch ituvo boua collected oa tiu^ a&Jaceiit ^^lu»

iuric î'-i oôd oxauplos of aaurltus , Only further collecting;

cau ro&oive tnis j^rub^c* .vlx s^>cciiêiiu availaole ix-c;^

30utiiVi.;^tern GecrÂc a,- car to be aaurltua - IntGi ^ . .:.

latter raco,

ThffliHWghlrf gsmfitua Joyk&ftA* »** aubspociaa

Holotvpa ^ y^ i.il50-2, an adult foiaalo froî i''id>onald

aloufîi, i>ta Karka iâtioxiinl ^>i.idlife Kefutiê, tvakulla Couiity,

irloridn, collected on 16 i'lay 1999 by Nita J* Boaaiaan and

Loi^ylas Urn i'<>us . . ...•

p,e V-. A aubapociaa of T^ft,^- . aaurltua

ci'irirnctQriZGd by 3 aural Tibials, a very daric brown or

blac^ iioi'vÛiu, ou ut loât t.^ alitorxur ^ .- ^ v^.^ ^.'^/.y,

ur Ixr^^tin

in

wltti ti» vertebral atripe obscured or ab»o«t, <. wj.uioJa

wiiite or iiii^ht blue iatorcii stride, sujd a rwJL&tiveAy ion^

tallm

jîiBSJim This Swiuî^ccxi^^ jccura xn Uic coaat«ai io^«u

laxida of iK>rthwQQtern ;>oainsuIar Florida frou the

Withlacoochao ulvQr on th© south to th« Wacissa River on

tiiO woat,

E£3cr- 2l! uolotype . ^cuteUation Is as followst

ocular^ ^+^ , «, T-ila ^+-, &i«. î-aXabiaia o, i.uj^.' _.. . u

10, iiiterêiiiais 3, dorsal scale rows l9-i9»17, veutrals

160, aubcaudal aeries aud tail incoiuplet©. i>ody lon&th is

621 mk.

The dorsua is black for a siiort diataiica hi back of

tae head, becoKlit.-. a dark brown .:ostF>riorly (i'iaers arid

l*aul 15i^« at inxauna/i vxui aiucii w*u»cit |.»i«>aoxit ovoriyiii^

the brown, iiio vertobrai stripe is iacKiiiti; ti*e narrow

lateral stritis ie liiîit blue (3401i. r;i« veuter is bluish

Wiiite aatoriorly, boco.iiiit, pale taiiuisii yellow tiiereaiter

(tiie ,^x>sterior i..a2't,in of each scute is ii.iit bluisu t^rcjcn);

the dark veatrolatoral stripe is uroac, occui-yiii^^ l/'j tao

width of a veatral on each side* The ciiia shields are

ifcite, tiiG su, raiabiale li^jiit blue at least posteriorly.

The aoati xg b^ric*^, heace no poatorbitul vitta is viaibie;

the parietal S|X>ta are iackixi^*

Diseuaaion^ îeristic and proportional character,

istics of this suospcciee are as foiiow;3: 3 to <

132

postocuJLarLj -.+~ to 2+2 toMporais; 7 to d supr&Ubiaia;

'> to II ixilrcaabiais; i to 3 intertouiais; i7-.iv-i7 to

ii/-i9.J.7 uorsai scaio rows; males i01^-lo9, feaiaiea ii>o-.

166 voatrals; .... i.t,.^ .i.*.j^xj^f io . j.^î j.u>-i^(j tju4;cc.. ,

iîales ratine i'rotn l*jo to i>34 aw. in body ienisth, 2<iu to 7^0

ffan, ia total icrii^,th; adult tail loiiijth/total lonÛi ratio

^9>j'j to,'j,3><^» i •• '-e . ran<ê iiX)a 14t)«> to 621 cuii. in

body ieu^^t:., .cio to 701 raî. in total leiit,Ui; aciult tail

l«ntth/total length ratio 0,331 to 0.37^.

1 iâvG e.^'.ar.;i;iod liviii^; 8pecl!:tens of tnls race from

c 3on, i.*vy, Taylor, aiif' W?»kulla Cotuiticft, and a

All speciiaens from localities northwiôt of tlia 3uwann^!«

i.ivor ,-^m tyoinnl oi" tîn n-hn n<»iQa as defined above; the

oxto.^wi.uii dU4 i*^vy ôui*t> X8 3o.iiewi*at coiJTusiii^j^. To tlê

tiortimeat of tlua comity , iîta^ occurs oa tiie coast, vrith

ilOifâji, iiUaiid to too txortii and eaet« lioi^'ovcir, ap&cir>.oi;a

colxected iu irU.aj[id iôvy County a& or iêar Vxata, uttor

LruiAKf Uuli ii€v>.uioc-;

i.fit;lis, all aave tne very dark

browa dox*3iu.i <.u.(.. u^^xoa »vi>4.ov^- 4^«/v»ral 8tri^>ci Cia^ ^îûriacic

of z^tae . uir 12l2y iroa ^ cailea i*L i««baaon ;>tation iâb a

black to very dai-k bx\>vi;n dorouo and a bri^ît \«iitc lai^cral

sbrxâ Aaoaa IroA wustera i<cvy uouiity ueu;.T:oa ju.w...' unci

oxia ap^>ear to be a mixBd population. \ik' 12139 and 12143,

adult fliales fr^- ^ mile ui&ner are both fairly typical

jl4lBtAa; ia îf lt^l33 and 1213^, adult fojr.aleo Iroa 3 miles

133

waw SiifiUMr, and in Ui' JU140, aa aduxt facaale fron UikeiiS,

the dorsua la dark browxi for a fcrw inches behind the nead

and brown triaraaJTt&r. I'uo vertebral stripe is clearly

defined in ail taree, - .tj iaterfd stripe in the uikwiia

aâCJuî^-'i* xa uxu^î iv-iiL. , ..it«riorl^ j a*. „..

otXior two it ia a du iite*

Xiiese tioraa aix. î.6 probably be dismissed as

uiorai.y oaji-i^^ vaiua.it^ ui. WiS Lutra wyjî.ca.i. coi^j4;xou (i;Acu^-t

for £4Qlll*a iXj'jdi rci|>ort tiuit cêoar ikoy rxbiuoiis;

fairly typical of sackeiii ( Weill was fauuiiiar wltr. t-u-^ rfark,

blue-stri^ eo fox^ at tiiau uiûs, ia iact re.u>i*ts cui.«.iscUiJ)^

it near the mouth of tîe >>tea.aiiatciiee iiiver^ Llxie County

,

in that saua pai>er)» mu. ^ i^ .:;.ii8tenca of UF 121i4» a

subadult eiale froui k ailes »wV; ^tumner, ¥;;ticii resoubles

aaekani in all features save scutellation. flte dorsim is

dark broun^ t.ê vcrteurai awri^ . i... ;. :* iSid .:.ww.*..ct

throuijiiout the leri4,th of tJ» body, and tiie lateral st-rip©

tBvy definitely a liît yollow* These anisials would

undoubtedly be call(»d iiiter4;rades if t.^re were aackeiii on

the coast with wtiich nitae could iatori:,race, but such a

aackan^ ^lâlatK>)i ap^>ears to be lackii^^ aw u.ju ^reaent

tioM* ileill (I9j>7) has deiiioasi;rated tiiat tiiere has been a

relatively rapid subsidence of t;^ uulf Co.iSt in poat«

i'X,&xs\MQGa.Q tiaes, uao. xx* seeiaa poQ:^xuxe x,iM.Xt ty^îcal

aackani did at oiie tiiae exist to tne wiux, of ££^S(|^, When

the territory viiiich it occupied aaîk beuoata Uijq surface of

134

to iiidicato tnat a coa;>tai aion of ^ ^^ ever ttxieted*

UF i*:130, an nr'uit fpr-iAie Tro- I fidlp' w Ynnkeotovii,

has a l>rov<ii cioraua îii/ ûXte jLot.vgi'«u 0Uri.^.tif Guid

ax30 i . J to b© a saciciax4 a xuta^p iiitor^,. , -a

seems rea»onablo itiaa^^^c. uS âcicq|Q|4 dioen occuj. ..^. w.^

couat to tae uoutA. ' i'roat txio viciiuty oi i<ai(0

fsata a in ^'îti'us County are '- <jr«f* to bo tâciiCiîiA

froiu ic.jl uii . .rotisjoii, i-evy Coiuity (taia 0^.0 ex

wouirt bo ivjfarutiLe t.-. aacisttî eACO;'t lor t-;c c:3Xor ol" tua

An adult rcuaoiOi C!i X^^:;^^ po&cs anotju^r kXxui o£

probie..., *r the colors o£ tae anlual a.* u*-w .ic . « .a*i.

close to taoae j.roaant in the j>reiervori apoci.iOii, it ouid

•ro6?biy qu ilify as a aackcai -^ aitiê ijit-iTiirsde. The

cox-cctiiit, a-:,â, aov.cvur, iaciicat© tuau t..«3 aiia^ce wa;^

takou 40 uiiê.'} > i'allaiiassee in Caiaoua County, x'iorida,

Tlia miioaê îy^ri'i v-ôuirf -,inco thn locnlit-.' at Tciio:. la CroeK,

w«uc.; xa in Liberty v/ou**&/ , raui^r tito** cuuLaoguia î t.;o;>o

data ax-e accurate, tuBii aackoni. ,. or a derivative, i>eno-

tr^tec v.v- i^..,.- -^* v,j.-cal aaurlt u» (wtiich occuru 3 Uiiiee

nosibx'c, a ioCctiity lca& than ^ mileid to tne eûti

witnoat .>viiit^ any iikdication ol" ii*tert,x*<*f atian,

tjj.x; v-a."..04* X14 vû'-'U U^^ ^'1 ijcij u .ju^ ^Vt-kviu^vj^ô , *^^ VOntrviXS, SIIO

A tan verteorai. etri;^; tno ilualord a . .'n {at -jyjd^ a

135

male) nas 7 su.raiabiaLs, IkS ver»tral8, and a^^rees equally

woiwL wxtii typical aauritua Ui all otaisr coopecx,&, ±

believe the locality data to oe iu error*

lîthin tne raiiê of T^ftfimffiimi^S m^SHm mm$•notiier sp«ci«B of 4,arteraiiaiie , i« 3Art«4ia f also liae a

very dark dorsuui and & blue or bluiaii uuite lateral

atripe* Xhie situatioa wj.^^ w ootailed «iu u j.aoux uabu*

Evolutionary Treads

The didtii^ctive fefttures o£ oaca of trie four

geoi^raphic races of XnatiinQDiiia sauritiia £ây reflect

historical factors &e lauch ae odaptatioaa to dxTforont

envix-OJiciGiital aituations* iievertxiolees, the black doraua

of •e&tentrioftf4iap the frequent .iari-owiiit. of its vertebral

stxî-e, and tiio frequent overlay ir^^ of tiiat strip© with

brown r> uj^ô'^-t seesri to iiidicate ada- tat ions for absorption

of aoiax- i'auiai,iou pa*-axioi.j.ii<^ t*K>sit> iouad iii Uk& riortliern

race of X« orojdbaus. lae brown ycuue, are alfiôet identical

with the youn/ of j^, ^, aauritu^ , and it soe^s aiîioat

cQi-\,uxii u '.\§, ropreaeata a aortiicru, aore

cold-tolerarit. , coxuvativc of Uie ncminate race.

Xi.w ..o^^î^wu ûbij^jecioa is v«ide<»rau4,in(^ mid appar-

ently Utrivea under a i^roat ;aauy eaviroiuuontal coiiditions*

It Aay iiave evoxved on tiie iainiand at a tiiiie vmen tiie

aoceatrai stock of ti» a^«t;j.o;^ ôi. v.îxc. ,. .......

j

ia tne nearest i^resenu-day repruaeutative) v&s isolated

on one or ;aorc of tne islands that co.iCtituted Florida in

136

i^leiauoccne tiotos* i. ara unabie to 8U4,4:,Q3t an explanation

as to tiifl afâi)tivo sit^nificance, if aiiy, of Um ver/ dûrk

dorsufit «ju4v v»i«c: iatorai stjfuîts îvscnt in the racu > ij,;>,,

cLy ,

It is entirely [X>ssibie that thorc is none| aiid that tlMi

factor vthlch is boin^; selected is aoâohov; aosocintef' vv-ith

cuzyati.i i.^voxvoc xu ttio ^x'oouct:*ou of pj^^Miitt • i.o iiUaUXi

eyos, at loast, tlis. is auch aoro conspicuous ix; oach

of the 8(i>verai hrtMt <t8 it occupies tiian would be the

c ativoiy av Uj,

A brii^f suBiaary of uur knowied^ê of ttiS ecoio^^y of

jt.« i^'\ari,

tu3 is ^resonted bolovi, <.ui account of

tnis Siêcxes iii one |>art of ito reuitê ^MicnJLâiO s«^s

tarpoater (i>i)-e,a, i>5<;b, ara^ ivi)3)»

Habitat

The eastern ribboasaake utiii:>i :^ a ^«at variety of

^tats; the only baaic requirt^rêut jou..:^ to be associa-

v/Ôii with permaucww vi- SG:uit>crmanont bodies of water,

;itats include uucn diver3e ;3ituitions as sandy

beacti, nait aarsh, rofiiisic o citch, pixic fiatv/oods, i;rassy

and ta:.iaracl. swaui., i navo jêraoaialiy colXoctod i, ^.-.uiî.Uvt^

in most of t^teso habitats; tiie ot/ier records are oiUier

froo the ii,torature or X'rotu fioid nottis aupiAicd by

oolieatûes* i-reûent ruforenco aos been made to txie

137

MKftiarborttai t*nd«fficies o£ tiiis a|;«icJio3, wtixcii, by virtus

o£ its relatively loti.êr tali, is porriaô better adapted

Tor oucti activity tiUA4i is i. ii£5iiife^Mi» Cari>«i-ter (l>ij«ia)

repoi'ta tnat 61 /jorcont of tae iûrituB coiiîCted cJuriix^

ti30 suiouer xiu)i*uuw ^ u.ij jui uufthes*

iôod

Xhe di«t of tiie oasteru riidbonsaake conuists ctiieil/

of adult arid larval a&ipnibiaris, and fiahea. ;»peciea

dellnitely known to iiave been eaten by 'Ji,_ jj^ aaurltua

unfier natural concitioas are liated in Tabif» 13. iieveral

au'w.iOi'3 ^âriaco, 1906; Carft'.juoij xv^-u; ua.xjwvon and

Pollack, 1956) have found invertebrates in the storoach

rîitAntc oi' ribbonajiakos , ar^l Fotflor (100?) tar.na aauritutt

as largely insect ivorous* 'on. tue u^ .t know-

ledtê, i'ovder's atateneiit does not aeoia to be Juetii'ied,

and tne few inataiicea in vîch iiivertebrates actually have

been found in sauyitup are probably caaea of aeooadary or

accidciital liquation* Ihe eastern ribbo.neuaicc does tiot

•eeiB any more i^rone tii«u ita> «r/vf4>we4'i« C(jwâtor|;ax-t to voluo.

tarily eat invertebrates , IxzarxiQ^ or eiuioUieraic vorteir

bratcs,

i»oci>uruai. jf>roya.xa^ and £>iodxnt^ i.s an {s^v(.r«êly coui^.iOu

piiejoomerion in tlie epecles, especially in tiie aoutiiierii |>art

of tiiB ranê aiid duritii; tîe waxia aiontha of the ..oar.

13$

TAiiLiS X3,..Food of YliMlinigfi'llAfl aaurituf

h

-9

lUiproductioa

Carponter (l<>^<ib} naa fouiid ti«eit# fentaJLe ribboASi4i»'v«s

In southern alcaiftait attain sexual .laturity nt an aê of

4^1 Biin, Corrtja]>oiidiat, data for : are aot . rcentcd.

r.cCauley (1945) rc>r,ortn a rraviri i'or^JLo fro.-i Mar,/ land

femal9 aauritua i itave meaaursd Uav« body iea^^ths of 33<<i

(fennsyivania), 3^6 (Oeorisia), 350 l*»8et Virginia), 365

(Mova Jcotia)| 37i («jeori,iai, 374 {m&t Vireîiiia), aad 410

{Qhiol laa., reapectivciy. ruaanopiaia §mî,'^m a^pareiiUy

fltaturea at a miciaiXar body aiso than dooa X« i)pQxiiitua>

«4atX(i^ apparently occurs Ui April ia ti«& iiortaora

part ol the rarit.e veiero a;:.rii;v. cr:i©rj,ence takes pl.-icc in taa

Xtttwoi' i-art Ol iiUiCiij la t.*c '.;..«..;.....; .ouuaera ûr&4.oa ol

the riuitj9 wiiere thare ia tio true wlater uloornation, uatiti^

probably occurs earlier. The younj^ are usually born in

Au^vist in Uio iôxtai^ July in taa ;>outh« alUiOuî tuere la

Mucii overlap. L'ates ranô xrosi 2 July tiurouî 4 October

(parstonal obaervationo). 'iinj ^ciuter data, surprlsii;^^/

enou^), is for a faeiale from northern i^'lorlda, axid taare is

a slii;tit r>osf5iblv- that it mifht r..-ve baoa iicr sccot-.d baroori

Ol U^ yeaJE-, iia«cie (195?) aaa îx.^( . tiiat tuxs

pneno&ienon way have occurred in a raaale ;^:roxiaius froxa

aouthaodteni A«oui8iaiia« TelX'ord (195^) r«;^.ort3 that a

raoiala wnicii ^vo birta to lit uoruâl youn^ on V July bora

4 additioîal youit^ on 16 September of ta^ sau* year. Xh«

lactor '.-er« ;iiaiforr.itJC' and bom dead,

(lolTordi l^XJt UiS iicoi. lor o^.broodo boin^ ii iiidivid*

uala. Thorn a •i.p:r.rn to ha ..rc;:o r.cnrt êoyfapiUc Variation

j.;i urooc ;>j.2u. muaii UvUaiUJi' ol Xiîviduals i3 i5«*J in

tii/eive bix>ods Xroiu i'iorida^ X«i*9 iu nine broods £xx>sa

southern Ontario, iO,7 in aeveu ur?od» £roin «.aryland

( ioy, 194^) » iO«0 ixi live brooda froni soutacrit

iiic-ii. an (Carrâter, i^Sîn), aud 6,0 in five broodn from

.1. . u..^ru ..jLcaxt^^^ iijux'i., x^'sîi), luu oxso oi txe utôd

tends to increase with tae si:^ of the fauaXe^ heiice theat

taciane may not b® strictly coi:.<>arable. Available data on

body leiAt^th of newly born X« gauritus aro suia..arizod in

Table 14* i'iaies »veiat;,e ioaiitedly larger tiian fci&aLus in

c»nc southern part of tho rai^ê^ ailiiiitly, if at all iari,er,

in tiio i«orta, jôun^; X« a«^Hritua taitd to averoâ ai;taller at

bii-Ui tiian i, ^f,ÂmuSi»

*.• wu u i^jtid) haa uu;iJi,'v'ju u.iô b.êro v/oro ti j.v

many aa<bryos in Ute ri«>it oviduct us iii tae leit ox.e m tiie

five foaale X« sauriLuq ane oxaîinod, iuui Xiziklo {X'j^7)

iiaa iouiuj ii aiuilar raiatioiiaiui^ Xii >/ i- ^ ^, ^l.._A..L'i«

Paraaitos

Irfingmann (139<^} roporta aii eaatarn ribbonsuako wiich

waa infccucu v.ô.t uaemo8poridia«

Xi^

tabu; 14,—Body length of newly born ^'UliWgfflTl^ff afturitus

ideality

V'v >« V iwU '^' AUiiO

SAa and uuUoubfceciXy i.s ciori.v©a iryj,! en© xatuer s^vycie*

or itci proLutjfo, Iiia r i iâ* ior tmlieyloe^ i.At tae

j:. diM» wiba its l0ii^j9r tail a.i4i rctknoed nuuber ol auvra-

lablaiSy appears to be the ^lore iii.jeciali»e. *vi.. .

^^peciatioa ciay navo occurreci, aa in kiiHny otiier s^.gcxos

pair?3 (i-îair, Ij^-.î, aa t.»o result o£ divioio.i durir4;

v;:ucn wore coril'iaoci during, tiiaciai periods in Floriclian

cnid :,ixicn:i rniujlf* , respect iveiy. To judge by tlio presont

v;ia'»rii>«Wu>.* of tim tt« spocio^, it v«xiicl . tixat ths

Kissisoippi iiivor (at a tiao v,nen it was wi<êr tmn at

present] piayed an laportaat rt>ic ixi tiie upucj^;o4.ui. procoss,

probably by dtiia/ii%, the ti*ie whan tney ai,ain caê in con-

tact Xbilowini; ttie oxpansioii of tiiCir raa, o;: curi:., tfie

^laCicAi, retrvao, ...lOi. oîu wio ^jopuxcJiCxoiia AXiiaj_L/ oxu c-.)..-0

ia coittact, i.ro&ui:<abiy l'xx*8t in aouthom liiiaois, ôiôtic

isoiati-on either was coiapioto or noariy ac, £lie i^rc^t

ii^jî.Jâjr*L^ ij ^,..1.. C>4.a tk^Q U^jUQ^dii Xki oXt«'w <«ilc ..iA»>*Uit

prt>i.cruacti cauaod aufi.icieitt coU|>atxtioa to prcvont exteiv-

sivtt 3y.ttpat,r^ • x.iu overlap in raii^ês which does exist in

142

i43

Inoiana is pXDbabiy ttta result of &ii eastward moveoient of

proxjuiûs In the rmiric i-'rininnuLa r-.^rini, î^ -».«rotiwrjaic

period isea ««ifc;i, i»/ii t..- .i^tio.m ii* iucianai

and ^.êrhapo buoae in ©astern iiiiiioio suiti soi&ue astern

iîacouein as wc^., , apresont reiictii of tf»at oariier, taore

eastvârd) diatribution*

!• âuiut,u§ saciapni . except i*or its loti^,, tail artd

smali, ^BpikraiQ parteta^ ;>,...,=u.., wv-nrs 2. 3t,riKi.v -^' —-

bi£UiCo to !• proJîatHa, | .^ of uuii anci taribbâw

Kid<.Ue itaioricri. The air-iiarity is so rant tÛAt soa©

ta^^noiiiic co:-iasioii i*Cia ro voiVfcti UijouU u Sijociiaoa of

qêJi^fit4 wtiich beara incorrect locality data (see -,

196l)» Two alternative hypotiieaes present Ui*iaiaeiv«ii;

Qitiêj" a«c>êx4 and rutilorig evolved iiidej^ndently In

respoaae to sir.vilar environno^-itai crs.sditions (v:>;ic-i fails

to take into actvju»iw i^.m uoiv^.-uia iôûiaiiioiis oi ruu,

j.îjrju; J ^

or JL. ^ «as derived fra-u ^ . >-is prior to tne par-

tition of tiiG rrbbona-i-ii-.G iiit.-* f.-^«? sîeciee (

a

^ ip^î i iu this

Ciû-:^ v,o:..jj. ^'j tiiL,uût W maîuUiîii jaost clooaly tiie proto*

type of tiia species Siî^ ., a*Hi rutiloriB t»i0 prototype

of !• aCftfiiJAUfti* ^^^^ coustancy with wrdch ti^ ^reauaably

ajficostrai suprîiibial uuîiber {sii is aiaintained in jk

would B9(mu to indicate toat it is t*te nost priraitive race

of j^, "^.S,J ^-»'-' *t^ *- .-. iniui-i, t.a,u. OA j^» ^.^,

>1?J8it''Vr ry iuodifiCution,

Deriving, aacken> _ ; .rvo to explain how,

iA4

ottier than throu4^h iiêfêndmt evolution, U.^^ i^ôcios

aauritua acquired tho dark veatro lateral strl^« .t iii

iAoyScaii and Central .. orican T« i>xx>,

xlmii|ff but iaclcin, in

t.ioac XMCC;i Va^Ch — : Ui COatiiCt WlUl T, aaui'Xvvw .t u.^

presoat tx-A),

Xhe raiit;<>s of S4*cketdi mid ratllorlB are now

s^paratao by over 1000 aiies .:.. tuo r ol two ot».er

£> ciea, A,» il» iL- il» u» or>i*rlu^ y but nju^jiiy

comparable dlajuact^v...^ c^^* ^.^.-.^-.^v... ^..^ ..cxlcatA >>i.iu

•iajterxi uaAted .iittttea 3ut)5êci.o& ol" LXJi- j4a ^*<*

Jtoreria wff<Âi4t9t4^.^s;HiftVa» altiiou.ii ou^y lu the xatter

iiiatanco has anoti«r racu U4. w.ic . ;>u.u^..uc;xcw u^ôa

iater^;03Gci batwtton Uie pair wiucii bocaue so^ûraUcU (u^t

la, ii y_, ^. o t|)d<:urf;^ of i-iorida lo acra cioscl; rclitod to

nosxlnate race, as ccrtaiii ciot.iuxita of color . attora Q'.iqlx to

Indicate)* When the rlbbonauakes wer« ablo oo. ^^^>^..^ u .oir

raii^&s, ibliov^lit^; tae rot]:*eat of t.ie (^laclerri, it Mas the

deriv-'itivou rather t;ian tlie ori^lnai stock {or, bci ro

accuiaue, fc.io *Gi- w wcarxy juxku tiiu uri^jiiiuj. a;»acivj

wiiich aoved liortnuuxQ a:ic! eaotvâxxl (xn the caao of X*

aroxliaua ) . or north v;arv1 nnr' t;f>.-t.x:ard (T» aa^rltua ).

Tuo wa...rxtutt wruu^ x«i, ucolOt,<*'Caxly s^>oaklxit^, U>tii

ae;.«iai'borc.Ai aiu) i»o. wic, lt:> uci.. bera ixave no ^ oors

iniithlri 1^,.. ,. ^ ..... -.or.ior cateôr^ , «... - itter

uiust burui/ be c<. tau oooloo^col eûivaxeiiCtt In

U5

•aaterri iioxico and eastern United »>i,atofi of tmnui^tji^^

ytje^âirm ol' v>\;ac>eru iiox*Ui utaexica, i;«jv&ral sbuciies

(tarj-oater, l^jt>'^H; rou'aette, 191? J J-.ave co. ttie

diet ,.iiio .iUiwiAutiv 4-.i.*u-*j*"uiiCu OA ^.ii© i*xut>ouuaa<i.c';i w^tî 'U«j aie

of tno otnor speciea oi iâai

.

_ ita wnicn tiiey ai'«

syeipatric, .âic' r.-ivo r;.,^vn tiat the dietti aiici .abitat pre-

rer©acoa oi wi*© i»uvar«* ajîsc* ., re diliortfat oaoUiî tiât

s«rioua ooiapetitlon between t/wwi r-irely aris««, se

studies indicate Ui-w >.* v,»t; -j.j,,..,,avrj.c taûc-u'^ ^, .._—.v -î

is 2tf>«t likQ the ribboi a habitat jQi\c^,

aXthcu^h not in diet*

i-;>55« ** rocoiisidoratiuii ol tao racer, CQXu^r

bairci, ^tûcer ^

.

id^^* ive^tiies oi tirie booîciary. Juxted states aiid

i*ieiJ,caw» ijoufujai'y survey, >iasîii%t;on, pp, i-3^t

1333 • Ca&aloiûa o£ liiorth A.,ericaiJi roptilca in ttm

.iusc-u.i of tho J.iithGoninn Iristitatlou* i art

A,..weri'entw« .î..j.;««tso£«Xeui *>.i.;ic« Coll*, vol*

2, uo* 5, p; . 1-172*

barton, ' , (.'•

I'>36. A statistical study of X iWlSIA

(Copo) with oo.uaeata on tne kinf*ni|j of X*

bu^lsrx \vuÂ.>* i^roc* ij*u-, ^.uc, .ai3.ii..î<w-i,

vol* 6>, piy* Vi-o^«

biair, w* i-rauk

l9i»«i* I. xiitriuwuxoiiui. jjatU' - ... .. . . x.» Oiio

sou.b«t«rn viiij-tuu abates la rulatloii to paat aitd

present envlroiifaBat;.. ôotôoi^caîi^ ^ /^...cr*

Astfoc* Auvauctt* -^ci*, arc* 17, pp* 43^-40o*

146

U7

iy^5» » Key to the anak«8 or the United States

,

Cftiiada, and iowor Caiiibniia, Pap, .ân

Acad, wcx,, Arts and «*Qtt€rrs, vol, 4, iio, 2,

pp. i-05.

Blaakney, J, 3hennan

195i, " uwifir auake recozxl Tor »iova wcotxa, ^;,.,:-i, ..ô^

.^4^

jauritua Bauritûi^ Liiifiaaua. Canadiaa i?'xeid

flat,, vol, 65, no, ., ,jp, U3-U9.

li^po, <• L;oOt,oat,r^ûiCid siûdy oi' tia© aW|/iiii>ians aod

reptilea o£ eastajrri Caiiada, i"»ati« riua, Caxiada

tiuli, lio, if>i>, iî^, 1-119,

uoie, iriedricli

id^y, iiemoricunên ubor iîarrsia'c Vcrouci; eixies

Systoeas d«r Maphiblan, -..via, voi, û, pp.

50a.566.

Bo/«r, I-orotiiy A, .md Alhert .'i, 'UinM

1934, ..n aaiiOt.a;,aa iisi* oi' tiic^ a:;i^;.hibians and

reptiles o£ Jofferison County, .Dioscuri,

Trails, Acad. oci, i»t, Louis, vol. <:<>, iio, 4,

pp, l<>5-^0i,

i>ro»wi, Arthur i^,

1901, A rsviow of tiuti 4,ou«ira az^d sâcios 01 <<.,Qrxcaa

snakes uortn of ixexlco, i roc, acad, A«at, wici.

i^hUadelphla, vol, 53, pp. 10-110,

burt, i^m h9

i9<^d« x'mo reiatioii or bxzq to laaturity iii tiio ^ rirtcr

ii^,). Copula, uo« i66, pp* d-.x<d*

Carpeater, Ciuuries C«

l^>«:a« to-iparat ivo v)coiOijy oX tiie coutuou ârter

aiiake tyjL^-ijiaiftiAf^ SL* SW^ •>»#)» t.^'^ ribbon

faortor snake (yiiaau. Wticri) iii aiixod

popolatioas. .^coi, .iono.r, , vox, <i2, pp,

l^^'i^b* Cirov;tn and »aturity ol tiie tttreo apcciea of

IhauiOODhia in ^Hiciii ...... Copeia, i95>«-, ^v

19^3. A study of hibernecuia and nibomatin^^

Assoclatious o.l îûkes and aephlbiaxis in

Hiciiiiî. Mcoio^;^', vol* }ht tio. i, pp, 74->o0«

iyScJ, iieproiiuctio.i, youn.., c^^.a and food of Oiciaiwuia

CXarK, /i* r«

l:^4y« •Jiiai-:<^<n of trie lAUL purls iies of iOul8i«uc«

Jour. AtiîiUtiatitt AcaU* •icx., voi* -49 i)o« 4»

pp« <C44->«i0i«

149

Cociiraii, Doris M»

iy6i„ ijê specl-iena of reptXlv. ls.' ;;^., aibii-mo in

the Unitwl i^tates fiutioiiai. ....;:.cl., IJ,S.

Nati, Mua. Bull., 210. 220, pp. 1-291.

Coaant, Koger

1951. The r«ptU«s of Osiio. iJiid otUtioii* >v ,.,

Midland itv^v*, W4.« ^0 y no» X, pp. I«»2d4*

Cope, ^ward D«

li>60« CataiOtûe of tiie Colubridao in tiic i^.uacun of

Paiicu'toi^>ida» cârt 3* ^roc, Acad* îat* ;»ci«

Philadalpiiia, vol. 1-i, o^-i^ 553-i»66,

ldd^« « co*.triDutioji to ttim nûc^yQX.Qlo^ of iêxico.

i^roc. A.-aQi'-, ri*ilo, i»oc,, vol, *iiii, pf^, 379-404.

Davis, N« .-•, VA«, aiid Frank ^. itice

ldo3. iiiorth Aiiteriean Batracaia and Ueptilia, round

east of the Missiasiooi Kivor. Bull, iliinoi©

Deckert, iiic .

1918. 3i»t of reptiles from Jacksonvilie, Florida.

Uapeia, no. 54, pp, 30-33.

Dickinson, WiiXiau S,

1950. iiecent additions to tiie records of tiie

distribution of tine re^jtixes in iîscouain.

Xraijs, v*i3cou8in Acad, oci.. Arts and Lattors,

vox. ^w, iio. 1, y^. 71^77,

150

Ditmara, îayaond i**

iJ<>6« Tne sttaices found wittiln fifty alias of Uew

York Cit;, Proc, .jiiinnoan Snp, ;;ov York, no,

DoDîln^^f Herndon 0*

i951» roposed method of expreasirig scale

roc; actions in Gnakos, Copeia, 1551| pp, 131-

134.

I960* i* ta.^.;o„ij.c tiûay oi w.ê ru«i3aa«i03, ^ônua

Klaphe r'itai.it,Gr, ViX, Tho Triaspis wuctioa,

;:ooiri. ica, voi, 45, pt, -J, p],., 53-30,

iiuellaai;, .>xi.^. E.

i95ii, A ooxjo^^rapliic study of tn& oolubrid sna<;e

ênua iêptodeira . bull, Amer^ Mus. ..^w« hiot,,

vol* 114, art* 1, pp* 1.15<^*

Duellaan, Aiiiiau ;.* and AXbert 5cîv«artai

195^* uM^/.iibians «».4v.. i-.vu«^^^ j£ souUiern riorida*

liull* I'lon.da Jtate fiua., vol* 3, r«o* 5, p^.>*

lcJi-324.

IfUBiaril, A* a, u,, u, iîorou, îKx .i« lutuOX'Xx

ld54* urpetoloiîa êneraic ou iiiatoirc naturolla <i«s

ro|)til08* itibrairie i.ncyclopodiquo da icout,

Farla, vol. 7, pt* 1, pp. 1-7^0.

151

Duzaeril, A» ami ^;«»1 inaia bocourt

1693* uecaercuee zooau^x -,u«a ^jyur -.oivja' a l*uj.atoxre

do la iiiHiiUi uQ 1' .j,ue» iltûdos axxr JL-aB

rer>tiiea, iiiiiiiion acioatii'iqua au lioxitiu© et

pp, 1-1012.

Dunii, .xiiaot li*

1940* iôtes on sooto Ajtaerlcaj) llsards and atâkes in

the i>useuist of Gotebor^* Herpetolotiica, vol,

1, pp. 139-194.

EvcMrmann, Barton vs?»

191^. l«ot23 on some AdizDndAck reptiles and «aphi-

olaxis. w..., .1 , iio. iÔ, pp. 4ii-5i.

Fitca, Haary ii.

1940, A bioiioo^raphical atudy of Uio oAliiioides

ai'i,aakrei» of îOrter Siiakea (^.enus Xaaviuopiiia ).

Ui'iiv. Caiiroriiia iûi)i. ii.ool., vol. 44, no. 1,

1>41« UfaOiâpiiic variation in ^ârtar atiakâ of tiio

ap«cies ,4{^^fffft?Hirfi airt&li^ xn tno laciiic

COc.ou ra|,ian o£ iôrth .nu&rxcn, ^^^r, *iiîai<i

i«at., vol. *i6, no. 3, pp. 570-:;»y*^.

îteh« Hanry S, and X, Paul îaalin

1961, . ccurrofico ol tae ârtor atiako, Vna _a

airtallay in the Great Plains and iiocky

Mountains. Univ. Kanaas ?ubl. Mua, rtat. iliat.,

vol. i3, no. 5, M.. '<iSy-30d.

152

XSitJ* vliiuiM* I'ascicuius ^'riiaus,

, Vidima, vol. i, pp. l-.i06«

fouquetts, ... »,, wT.

i'>^4« ^ôod compotitioa a^mn^ four sympatric upcci«0

of i/irtar sriakea, ;,enus I'haaitoohis , Xexae

i^'ovriej. , Henry W«

1907» 'i'ho an;::iuians and rc.tiica ai' Hew Jcr-soy.

-^t PP« ^3, '250, 3:ii-3o5, 367-40O lijart),

Kox, bade

iiJ4o» ^Xrect of toiiirjcrature on dcveioîaQnt of

scutelL;^tion in tim ••.artor anakc, ytif|\^*inî?ftiffi

cj-e . ..:. ....^. uo, c^, i:,^4S, pp. ^52-262,

Fox, loiade, Charles Oordoa, and i'^.arjorie H. i'ox

[_ J T!u> rrAf«r,rtin.'. nf Atypical ecute patterns and

iiareiip ia . 'ay cJCiX)Guro to cool

ta-voraturoa durii^ aubrjoxiic devei >.

^.. ^ rttss, iôoloji4-ca.

(iamon, ii.

l5^<^. A ayiiopsis or tite ruptileo ^nd am^jîibions oT

lXlxnoxt<« L>u.i.i,« Xiiinois ô>.<.^'Vj «.«<^l>« «•..».•

Uiat., vol. 3, i i'. 'ii>-3<Jti.

Garatau, :>.

1^3. Jf"^'** r'*-f>tiio» aiK' bar,rrjCiiiai;s ol' itortn Anarica.

«>t&.u. *^»U8. Coiup. -ooX. , vox. J, ii^m 3, pp. l*ldj^

153

GJLoyd, Ho>»ard K«

X9'>i6« Xno ai^viilblana and rs^.tiitts o£ kranklin

County, KiuisaB. Xraiisi* ii£u;iaau Acad* «^ci.«|

vol. 31, Pi'» Uiîa.

193 5* îono aberrant color pattorus in •mkes. Pap*

Mlciiiân Acarf, 3ci,, Arts and Lettors, -rol,

^0, no. 3^, pp. Uw;>l*66d«

Hamilton, w« j,, Jr., and Joseph A. follack

1956. The rood of ao^Tse colubrici snakes froa Fort

iiGnnix^,Geor4â. iicolo<^ , vol. 37, fio. 3,

pp. 519-5-^6.

Harwood, Paul D.

X9y^m The 2)elzitinti:}S para3itic in t^@ *ua;îiibia and

Hoptilia o£ iioustoa, Taxaa, and vicinity,

rroc. a.u, i.«atl. .iuu., vol. 31, no. 17, pp.

1-71.

Holbrook, Jomi ij.

i<i4^. i«orUi Hiuerican norpetolo^. ^nd eciitiou. Vol.

IV, pp. i-i3d«

Hudson, oeorê ^.

1^4^. ihe u4it;>iiibiaA8 and reptiieo oT uebraska*

iuGoraska Conscrv, Ouii,, no. *-4, pp. 1-146.

Ja;rt, Georges

1663. i:«lonco systîtaatico detail olidi doscritti o

dosegnati per I'lcono^ralia general©, iiiiari,

pp. 1-143.

1%

liîfj, "0UO3 on jr cctxlx. .^;c?tf^y of Lâ^ , and a

deseriptioi^ o£ new species oi sorj.'ents in the

coliectioi* or tiio North Vvoatera Univer..-.k,^ of

iivauacou, Xii« i'x\}c« <^cai.i« ât, ^ci,

i'axiaciOi.puia, vol. li, pp, 93->iGJ«

l^J^^» Vertebra duplicatioa , uu osi.eoi.o<îcai. oîoualy

widespread iti 3uaK0£s» iior .otoio, xcn^ vo^. X^,

Pi'* ci?-»0«i,

i->43* XulLian^^th dUTfereaces ia Siiakes witii uotes

Oik fii^ûal dijuior^tii££{4 aiid tau coeilxclont of

diver^snco* bull, e>ooi* woc. Jasi lieô, uo«

X3| axn.* ^, ^.,.. 1-60*

Kiain, Theodore u,, Jr«

19W. A rocoixi lit tar of XW^^K>,^-n^ JUCkdi^

^r,v ..i:.>u;t» aarûtolOoica, vol, 5» p« 17«

i< in, Uustav

1699* On haemoeporidia in American roptiles and

butracuiai:s« iiui< lork Huo, «iour« for «Iau» 7»

iSy-.s PP. i-i7.

i^!>^» >-"- • coitteata of ôrue aquatic s.iai!Coa fn>a

iâ. ' iicAieoter, tittaourtî County, OKXûjLo;.ta,

i55

îzuuieusy CaroXuB

1766. a naturae. iSditio jiuodacij;ia.

ioermberî Eliiar

1^94, hotcis oil reptilos and batracalaas collected In

Florida iti 139^ and 1693* Froc. ^.^. Natl,

Mus,, vol. 17, no, 1003, pp. 3i7«33;^«

l4cCaul€$y, iiobert ii,, Jr,

194^, Tug rti^^t^xui^ or Ear/land &iuX the District of

Colvuîbia, i rivately publiatitsd, 194 pp*

Maerz, Alo/s J. and .M, Hea x'aul

uilX, iiow Xork, 'C(Xi pp,, ^ pl«

Hainate, ii;d/aond V,

1900* :>y3tc.:i.rolc cilviaion &sid evoluuioti ol tao

colubrld sriako clonus iiatrlx . with coifîiucnts on

the Qubâail/ Natriciriae, Froc, Acad, •<.«&•

Sci» niiladelpoiair vol, 11'^, no, 3» pp« 41«71»

omaticQua ..tiiri.o*- wo* j; Colubridao).

Itotulac: ^âtuxtio, no, 341, pp, 1-7*

Merrea, Bias!us

IJ'iÔ, Vorsiich oines ^ystesiis 69r Aai>.>hibien, Uohann

Onridtiaii h.riMi,©r, «u^burt,, p^, l-lyl.

ii»6

1953. _ i-«^ variation ii\ t/se i^p?rter aitake^

TftflWfr liî? 9y:rXrtf.i.^,tfTt-i^» itj.îu u(ju^, -c^., vol,

5, no. 3, i>p» 34a-37'>>.

kittiur.nn , M, B,

l5***y« -iA xc varXuttioa in *iarcy'a ârtar snake,

y^>a4W„^-LG .:uu-cianua (baird and (iirard).

Bull, Chicaao Acad, ^ci,, vol, J, no, 10, pp*

.249.

Myars, Ch&rlo^ .

19^* ,Aîbiaxui a.;a ruptixe& ox I'lu^xivau^ ^tawc i'ark

oaci vicinity, Dent County, «^^liasouri, tu*^. .\3

Lcn<\, i^ci,, vol, 62, no, -, ,^ » /Q,

£(Oill, HxlIl'CC. 1,

JL9^4« ^«;o axid taxoiiofaic allocations of UMiîxibiana

and roptiles in the aoutîeaatern United states,

.-.^1, iv«8« I'XV, i<<)S3 jU.len*U Xî)^/tilU last,,

vol, 1, no, 7» pp. 75- >6,

1957« iiiatoricai bio4ô(j4,ia4,.iy w*. ^ 4wt»rj*jo-uv.^ Florida,

aull, i'lor^ : itus, , vol, <-, xto, ?»

:;::. 175-^...

x>'>j, *i;o occurrence ol cii.i,..»xb - .- ..- -î*

saltv.atar area:i, and a biblio. . • iûll,

Hariiia Sci« Gulf and Caribbuan, vol, 6, no, i,

pp. 1-97.

i^7

i9i>^< Studies oii the aupiiibiaris and rojptiies ol

Aiieti's Itci/tii© iiist., voi, i, iii>« i, iii>, i-76,

Publ. *'iufi, Zooi,, Univ. iîictiifeaa, no. ii4»

1961 hoacr.ciaturai status of the ueotropical sub-

ûi-xUv*s. . iiotulae uaturaa, ao» 3'*u» p*/* l--^*

[ ] V(ili/!nt.iori of t\uQ specific status of the

i<oas<Ban, Hita J*

l'>^6, toit'tiii 4,rowtUg owi leeuiMi^ recoruii w* a wxxioa

of itj^steru rlUUoi* suaiies, 1-. .^îi

^rQX%>^* « *^w» -^jw—'•-'^-^ ..^^rpotol. woc. Lruii,,

vol* 1» ao« 1} pp« ^$«

itutiiVea, *U,«xtaifiQr G,

t^'ter-aaakoQ* U,J» i«atl« h\x3^ iiull«| uo« 61 ^

pp. 1-.201,

158

1323, In J.%"r>-J

-..nvii, rr. ,nt, of an «jqp«ditioa from

f itvoburji Lo biid '^cky Mountains , jierl'onaed

In th9 yttarii i3i<^, id'^Ô. Vol. I, p|>, l-3^«

ionân. Hurst, Roes, Ovue, axid BroMi, .iA)..<Joa.

;»cnuidt, iârl y«

1^5>3» '« CiitiCk list ol liorth Aadrican aa^/xîibiaaa and

reptlLas* l>ijcth editioa* Auaar* iioc. of

ichtayoioîats aiid ilerpetoiOoists, Cliiciv.^j

licuiiiadt, ivarl .'• aiid ibOôr Coxicînt

1956-57, A. i^ndlx to Opinion 36^. . .ueat tiiat tha

International Coraisiasion on roolo;j.ical

lioaonciatare naould direct that ti« uo:;;inal

species "Colubor oirtaiir." i.innaau3 1753, and

"Colufosr uaurita" i*iiiu.< 'Uô, i76o, a-iouid be

inter, by the nooty:>e8 here d tod

by the preaont authora instead of, ao

nitherto pi\>ôa<iU, by rereroxice to 3iy©cii'iod

previously pubiiaiod fxiiurea and deacri^tiona*

. ^^, —f—jo» Opinion 365* Opiniona and

1 dereu by the iiiternational

laion on <x>oioîcai i*oijtn»ciature, voi»

x-^,t'

'• '^* tj'^xo'j'j u^ {x'u;C-;:> ueiii.twu4„, j^^\j..Ki)tl^

. i-;i-;d3o.

159

w£!lii>t^.t| .iU.uU.iv -'^

1949 • TikQ 3u ôs o£ t,ijQ pialas âtxtor nijikB^

'Ct^-^mm^M ?f:f^^v^» ^^-« Ciiica.ô Acad. Sex.,

voX« <^f no» iâ ^p* '^<^>*300«

Siaith, Hobart 'A»

193s, :*otor^ on rn;.tllc^r: .^jkj aapiiibiaas from Yucatan

Univ. kiciii«.ai:i , iio« 3a<^« pp» X«>22,

îth| llobort, v., .:.;k Ja;,i«8 .^« Hutieey

Acad. OCA,, vox. 03a PP» X^6»X64«

or ilejd-co. U.w, riatl, ;iii3, LuiX,, r.n. lily,

î>. 1.239.

;i>£&XUiy PhXXip W«

X957. n snftXyal* of poat-Wiscoaoin blOtieofii w<^..^ of

tiie âirici ^y&aiaaula r@i^Xoa b&aod on

difiti''ibutionai phena^jteiia tjjaoiv, torresfcriaX

Stout, witA^r

1903. A coXi«cl.jjDu of ru^'biXss and batraciîaiis Troa

Prwc. Mcad. i«at. 3ci. PalXacloXphia, vol. 35>

pp. 53^-54-!:.

^60

Stone I Mtmer

iv . J or

,m\d loiavjji.c, ....or,

iitrackex', Joiii. ., o.,

190^, rnr-tllna c-uiri batric-il-nr^ of ,'icLr. .nan

1909. Kot©« «*. V,..* A»«rpetoXoe;y ^x wurnet Co—.u^

,

uor Uaiv« tiuli.*, vol, 12^ no, 1^

pp. 1-20,

OliJ.'.LtxL\i ^ 1.., /i»

1906, J of -,

?Q.-in.';yiVui:ia .:tia*-::j Da: t, A^;ric,, Tiv, /ôcl,,

iiO, ^, i>^, i>>-.Ûi.>,

I'ayior, t.. -.dâr

l<)9*ii, —ologu© of the aiLiKea '" ">brAaka vit:* rifite*

oi* their .«<..l>Xu3 aruu diaux'îA^Wiozi, ..:dr« iia&.y

vol, *i6, iiopt., pp, 7iî»7y^.

AoiXord| .>', ii« y tir,

l95-i, ..icai survey in tlio viciiiity ol"

X,ako iiai-.p, I'oi:-; to,, riorifla, in:,. Jour,

riox'«u, ^';.>, i.i>>l^>«

rilds.;, J, !&•

1.961, Certiiia oouuavnte ->'' t/:"> subspeclAS problwa,

^yst, ôl,, vol, Iv;, uo. If pp, 17-23,

161

TiuklOf -

1957. Kcolo .7, aiaturation and r -rodvictioxi of

3S, no, 1, pp. o>.77,

1937* wU^û^^col oa3c;:'Vatioas oa am^>;u.;^lan8 Siod

reptilos coiloct;6d in souUieastern Oklalioma

during the suioaor of l^3^« <».w. , .îdiaiul

iiat., vol. l$f no, 2, pp. 2a3«303*

Van Denburghy John aru? Joseph H, Slevln

1913« Tiic 4,ai*tejr-siiat3us ox wcswjTii i-ioniu o .

i*roc» Caiiforaia Acad« oci*, 4tii series;, vol«

8, pp. lC>l-.ii70.

Webor, Jay A,

I9^6m Uorpetolotîcai obs&rva;;ioas in th« Adirondack

Mountains, Haw lork* Copoia, no, 169, pp«

lOvJ-11^.

/uiiar* KidlaiMi .'^.w, , vul, 3<^» no« 1, pp, 16<d»

190.

l>y/* UAîObook of 8ti«iK«s ol tiiie Uai&od ;L>tatea c:- c

Caiwttia. Vol. iX. Co;iStock PublisJjlng

AsuocjuiUaa, x&uaca, new York, pp. ^5*1105«

I.. .- :c:}i

Douglas Athon Kos^taan was born July 4» 1936, at

Maukâba, kîscoasin. in Jiaie, I9^^t ^^^ ^'^^ graduated Irom

i^riuCQtoa iii^.h ^ciooX, i^rixicotou, ^ôw Jeraoy* ia June,

19i>d| he recoivod ti;)a det^<)u of Bacaeior of ^^rta frcsci

doutnera xiijaiois uaiveral&y, Carbotidaie, with a major in

aooXoc^y ^^'xi Uiiriors in antaro]>oiOi^ and botany, xa

êptttat»ar, ivî^t ue eujix>juxcu in Uve Uraciuate ooioox uj. o.iei

Univaruity o£ iriorida. He vomed aa a graduate assistant

in tiie Do artuiont of Biology uiitii Juiu^, 19^^ « £uid as a

âduate i<aiiow uiitxi Uuîu, i'>61, i'ro.^ ^'e^-tOi.ibor, 195^0

until tite praaent tioe he lias puraued his work toward tiM

de^sree Doctor ol Piiilosophy*

Douâo Ataon tlosaiaan ia rârriod to t)ie loxîer liita

J&iiV! Kuster €a\6 will become the fatiieir oi oiio child in

Aiiterxcaa oocxety of Icht^yoloiîsta and Uer^-etolo^îsta, Vcm

aocie.y of Systematic <ôolo,..;i8ts, i'hi Si^ma iiiolo, leal

oocxoty, -ic- ^ ...:,. a «-iii, . . ...u iiuoixsuuc. ^^^.gv'-u

tocmiical pajôra in the lielda o£ iiar^êtoi <i

protozooio^*

162

This disseriiation was prepared under the direction

of the chainnan of the candidate's supervisory coituaittee

and has been approved by all meuibers of that corainittee.

It was subniitted to tiie Dean of the Coiie^e of Arts and

Sciences and to the Graduate Gouncii, arid was approved as

partial fulfillment of the requirements for tne degree of

Doctor of Fnilosophy,

August 12, 1961

Dean, Collegeand iaciences

Dean, Graduate ociiooi

Supervisory Cornmittee:

Ciiairraan

y

31B ' 3

A taxonomic study of the sauritus group of the...

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