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UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUI"l OF NATURAL HISTORY
Copies of publications may be obtained from th e Publications
Secr et ar y, Museum of Natural Hi story, Un vcrsity of Kansas , Law-
rence, Kansas 66045
Price for this number : $6.00 postpaid
Front cover: The subspeces of thc ridgenose rattlesnake
[Crotalus icillanlt). Clockwise, st arting from the upp er left, G. w.
amalnlis, G. w. meridianalis, G. w. silus, and G. u;. icil lardi. All
photographs by [oseph T. Collins, wit h thc cooperation of the
Dallas Zoo.
UNIVERSITY OF KANSAS
M U S E U ~ [ OF NATURAL H I STORY
S PECIAL PUBLICATIO:'\ No. 5
_________Dcccmber 14, 1979 _
THE NATURAL HI STORY OF
j\/IEXICAN RATTLESNAKES
By BARRY L. ARMSTRONG
Resear ch Associate
and
JAMES B. MURPHY
Curat or
Department of Herpetologij
Dallas Zoo
621 East Claretulon Drioe
Dcllas, Texas 7520.3
UKI\'EI\SITY OF KA:"SAS
L AW RENCE
1979
Ul\"'!VERSITY OF KAN'SAS P UBLICATIONS
MUSEUM OF NATURAL HISTOlW
Editor: E. O, Wil ey
Ca-editor: [ oseph T. Collins
Special Publication No. 5
pp. 1-88; 43 figures
.2 tabl es
Publ ishcd 14 Dcccmb cr 199
COPYIUGHTED 19i9
By
i\ IUSEU), I 0 10' NATUHAL HISTOHY
U :\,I VEHSIT Y 010' K A:\'SAS
LAWHEXCE, K AXSAS 66045
U.S.A.
P II/ :-'-TED By
U:\ I\' ElI SIT Y OF K AKSAS Pruxrrxc SEI\"ICE
L AWRE:-'CE, KA." SAS
ISBN: 0-89338-010-5
To Jonathan A. Campbell
for his encouragement
PREFACE
Beginning in Novcmbcr, 1966, st udies on rattl esnakes (genera
Grata/lis an d Sistrurus and other pit vipe rs were initiated at thc
DalI as Zoo wh ch includcd techniques for mai ntenance an d disease
tr eatments, in conjunction wi th obscrvations on ca ptive anel wi ld
popul at ions. Maintenance tcchniques and elisease treatments have
been pu blish cd in an earlic r co nt ribu tion.
The result s of our studies on th e ecology an d natural h story of
Mcxican ra tt lesnakes are conta ine el in th e present accou nt . Since
nu rnerous heh avioral seque nces wer c diffi cult to record in th e field,
man v rattl csnakes wcre maintaincd in the laboratorv. Over one
huncred aneI twenty-five cap t ive individuals, compris ug over 50
tax a (inc ludi ng fonns ineligenous t o the Un tc d Sta tes) wcrc avail-
able for stuely.
' Ve havo att empted to sho w the value of a multiface tcd ap-
proach to th c stuely of a body of organisms hy beginni ng wi th fielel
obs ervatio ns as a basis for undcrstauding, Ioll owcd b y maintc nance
in th e ca ptive stat e wher cupon specimc ns can be pl aceel upon
eleath in a systernati c mu seum co llectio n. This a rrangement allows
an investi gat or t o exami ne various as pccts of an an imal 's "bcing"
by recording dat a whi ch would be virtuall y impossibl e to record
in th e ficlel. F urthcr, thi s combinccl a pp roach maximizcd our ahili-
ti es as one of us is sorncwhat inco mpc ten t in the ficld aneI the othcr
is an errat ic animal kccper.
The assistance anel coope ra tio n of many perso ns cont rih utc cl to
the eompletion of this study. For va rious courtesics ext ende d to
us , we thank Hay Ashton, James P. Bacon, Hobcrt L. Bezy , Cha rles
~ 1 . Bogert, David Brown, ~ I a r y E. Da\\"SOl I, J.S. Dobhs, Mchael
S. Eelwarels, Thomas H. Fritts. Ja mes c. GilIingham, Honald Goell-
ner, Harr y , V. Creen c, Hcrbert S. l Ia rri s, Charle s l Iocssk-, Terry
Hul se y, J. P. Jo nes, Thomas L. Jarci a n, Joh n E. [ oy. Tommy Lo gan,
Ar thur Lo pez, Danny Lopez, Edwarcl Xlaruska, Hymcn Ma rx,
Robcrt " ' . ~ 1urphy, George R. Pi san i, th e late Loui s Pist ola, Thornas
Portcr, Stcvc J. Prchal, Peter C. Prit ch a rd , W ill iam F. Pyburn,
George B. Rabb, Charl es \V. lt adcl le, Vi ncent D. Rot h, Thomas
Schultz, Hobart ~ 1 . Smith , Bnrney Tornberl in. Tom Va n Devcndcr,
R. Wayne Van Devendcr, James Walk er , Tim W alk c r, John ' V.
Wright, Ri cha rd G. Zweifel , and our many frienels throughout
Mxico.
We are gra t eful t o \Valt er Auffenberg, Charl es M. Bogert ,
Charles C. Carpent er , [ oseph T. Collins, Hoger Co ua nt , James R.
Di xon, Wi lliarn E. Duellma n, th e late Howard K. Cloyel, Mchael
lIerron , Donald \V. Moorc, John A. Sh adduck, and th e late Edward
v
\
H. Taylor for reading and crit iciz ing the manuscript and offeri ng
many helpul sugges to ns. Photographc assis ta nce was offered b y
R. Tcrry Basey, M. Gra nger, Waync Seifert , and Joh n H. Tashjian.
The able sta ff of th e Dalias Zoo Dcpartment of Herpctol ogy, David
G. Barker, Ra ymoncl K. Guese, \ Vill iam E. Lamoreaux, anel Lyndon
A. Mit ch el l have recor clecl numer ou s ob servat ions ancl have con-
tribut ecl to th e overa ll mainte na nce of the rattl csnakcs diseussecl
th rough their enthusiasm and expert ise. The librari an st aff of
Inst it uto Butant an in Sao Paulo assist ed lIS in numcrous ways.
Variou s perso ns associate d wi th th e Univc rs ity of Texas Press,
specifical ly Ann Hidalgo Manl cy, Ph ilip L. \ Vagn er, Rob ert Wau-
chope, and Robert C. \Vest , alIowed us to use figures f rom lland-
hook oi bli dd!e A me rican Indi ans. Ralp h R. \ Voocliwiss an d Ant on
S. Prechtel of th e U.S. Department of Commerce, National Ocean c
an d Atmosphe r ic Admins trnt ion, Washi ng ton , D.C. , p rovid ed cli-
ma t ic infonnation and alIowecl us t o use material from Climai e oi
Mexi co by [ ohn L. Pa ge. Se or Silvino Agu ila r Anguia no , Subdi-
rector Gral. de Geografa y Meteorol oga, sent climat c data and
numerous maps for our use. Memb ers of th e Dall as Zool ogi cal
Soc ietv, through thc efforts of Bernard Brist er, provid ed partial
travel expenses.
The arduous task of t yping ce rt ain parts of th e manuscript was
ch eerfully accompli shed b y Kathryn Campbell, [ anet [ackson ,
Martha, F. Murphy, Verna S. Murphy, ancl Myr a Smit h . Deb
Benn ett skillfulIy execute d th e dra wi ng of th e map of Mexco on
the inside back cover ,
Finally, special recogni ti on must b e extended to Jon athan A.
Campbell, who in so many ways aided us in th e preparation of
thi s manusc ript. His unflagging enthusias m in th e field, th e gener-
O\lS donation of specimens un der his carc, his thou ghtful comments
an d criticisms of th e manu scri pt, his photographi c abilit ies and
overa lI enconragement made the completion of this st udy much
more enjoyable for us o For th ese reasons, we have dcdicated this
study to him.
Barry Lo Arms t rong and James B. Murpln)
Dalia s Zoo
Dalias, Texas
Mnrcli 1979
VI
CONTENTS
PACE
rNTRODUCTION __ _ 1
GENERAL DESCHIPTION OF THE HEGrON 1
MATERIALS AN D :METHODS 3
ACCOUi\'TS OF SPEClES
Crotalus atrox Baird and Cirard _ _ 4
Crot alus hasil iscus (Copc) 6
Crotalus catalinensis Cliff 8
Crotalus cerastes Hallo well 9
Ctotalus durissus Linn 10
Crot alus enya (Cope) 16
Crotalus int ermedius Troschel _ _ 18
Crotalus lenulus ( Kcnnicot t ) 22
Crotalus mitchelli ( Cope ) .____ _ _ 29
Crotalus mol ossus BainI and Girard _ _ 31
Cr otalus potij sti ctus ( Cope) 34
Crotalus pricei Van Dcnbur gh 38
Crotalus pusillus Kl auber __ _____ _____ ._ _ 43
Crotalus ruber Cope _ . 46
Croialus scut ulai us ( Kennicott ) 48
Crotalus steinege ri Dunn _ _ . 50
Crotalus ti gris Kenni cott 50
Crotalus tortugensis Van Denburgh and Slevn 53
Crotalus t.ranscersus Tavlor 53
Croialus triscriatu s ( Wa'gler) .56
Crotalus ciridis ( Hafinesque ) ____ 61
Crotalus iclllarcli Xlcck _ 63
Sisirurus ravll s ( Cope ) 68
DISC USSION i
__ _ ii
APPEi\'DIX: H:\ 1\' GE OH ALTITUDE EXTE1\'SIO:\ S
HEPOHTED Ii': TEXT __ i8
LITEHAT URE CI1'ED i9
INDEX 1' 0 HERPETOLOGI CAL SClE:\TIFl C 86
vii
I NTRODUCTION
Thc view that thc central pl ateau of Mcxico was the center of
di sper sal for ra ttlesnakes is a zoogcograph ical position neccling no
defense (Cloyd 1940, Srni th 1946). Howcver, infor mati on rel at ing
to man y speci es, particularly the pri mi tive monta ne for ms, is limi ted,
Ther e are few observations dcali ng with th e ecology an cl na tural
h story of Mcxican rattl esnakes owing mainly to th e inaccessi bi lit y
of many of thc popul ations. Mexican ra t tles nakes inhabi t many eco-
logical nich os ra ngi ng from xeric to mesic environmeuts. One spe-
ces, Grata/lis triseratus, reaches alti tu des of 4573 m (Klauber 1956).
Man y mai nl and forms are found in th e ce ntra l Mexican plat ean
whcre th ey tencl to be restrictcd to ar d, roc ky situations, Othcr
specics a re found in lowl and desert habitat s, incIuding islaud popu-
lations of the Gulf of Calif ornia.
The pu rpose of thi s study is to: (1) analyze ami dis cus s th e
environme ntal compoue nts (including physiography, vegcta tion, ami
clirnate) of Mexico and their clcct on thc d stribution of rat tl e-
snakes, and (2) recorcl obs erva tions of na tural an d ca ptive po p-
ul ations.
GENERAL DESCRI PTIO OF THE REGl ON
Information recorcled by Goldman (19.51) serves to illustrat e the
ph yscal cha rac teristi cs of Xlcxico (Fig. 1). Barbour (1973) prcsents
a dctailcd dcscri pt ion of the central mesa , an d ~ lorafka (1977) de -
scribes thc Chi huahuan Descrt. I n ordcr to un dcrstand th e currcnt
gc olog ical pattcrns of Mcxico and th c gcohistorical dcvelopment
and pal eogeography, thc rcadc r is ref crr cd to Ma ldonudo-Kocrdcll
(1964) and West (1964). Soil has hccn t rcatccl by St cvens (1964).
The rca der should consult such ll1aps as those includccl in the
works of Cout reras Arias (1942) an d Il crnandez (19:23) for climatic
da ta . I nformat ion on wcathcr an d climate is sununarizc d by Viv
(1964). Sh reve (1944) discussed rai nfalI pa tt erns in Mexi co, no rth
of th e Tropic of Can cel'. Refer to Fig. 2 for th e mean annual
rainfalI in M xico.
Na tural vegeta tion in Middlc America is depictecl in Fi g. 3.
Detail ed accounts of thc vegetati onal relationshi ps for various
Mexican arcas are as foll ows: Bcard (1944), Brand (1936, 1937,
1957), Duell man (1965), Egglcr (1948), Ge nt ry (1942, 1946a, 1946b),
Coldman (1916), Go ldman and Moor e (1946), l lol drdgc (1947,
1964), Leavenworth (1946), Leop old (1950, 1972), Lesueur (1945),
LundelI (1934, 1942), MacDou gal (1808), Ma rt n (1958), Mart ncz
(1945), Mi randa (1942, 1952-19.53), Mi rand a an d Sharp (1950), i\l ul-
ler (1939, 1947), Nelson (1921), Savage (1960), Sharp (1946), Shreve
(1934, 1937a, 1937b, 1939, 1942, 1944), Sta ncllcy (1920-19:26, 1930),
Wagner (1964).
2 SPECIAL OF NATURAL HI STORY
"'
I
I
+-
- -T-
(MI/BBEAN
Fr c . 1. Phvsiograph c region s of Reprinted from Hand book 01
Middlc American l ndians. Copyright 1964 by Uni v. Texas Press. Used by
permission of Robert C. West,
FIC. 2. Mean annnal rai nf all ( rnillimetc rs ) for Mexico. Reproduccd by
pernu ssron [rom the NationaI Oceanic and Atmospheric Administra tion,
Washingt on, n.e.
TIlE NATURAL HISTORY 01' RATTLESNAKES 3
FIG. 3. Na tural vegetat on of Mexico. Format ons class ficd aft er Beard
and ot her s. Reprinted Irom l l andhook 01 Middlc Amcrican l udiaus. Cop y-
rig ht 196-4 by Un v. Texas Pr ess. Use d by permission of Phil ip L. Wagner.
AND
Most of th e localities whic h are list ed in the foJlowing species
accounts are bascd on our own obs ervations un less otherwisc spe -
cified, altho ugh voucher specimen s werc not collecte d in many
instances , Since the primar)' pu'1)OSCof th e st udy was not to collect
rattlesnakes for sys tematic collec tions, man)' of the individuals ob-
scrve d wcre not coJlecte d and in sorne cas es, parturi ent Icmal es
were hel d until th ey gave birth th en rcl ea sed with th e young at th e
sarne locality. Li tters of snakes wcrc 1I 0t measured and wei gh ed
in th e field. In the Iabor at ory, weights wcr c rccordcd with an Ohaus
tripl e-beam bal ance. Measuremeuts werc made on newborn snakes
in most cases , using th e meth od described by QlIinn and [ ones
(1974),
Snakes th at were remo ved fro m wild popuIations have been
deposit ed (or will be up on thei r death) in th e Iollowing vcrtebrat c
collec tions: Unive rsity of Col orado (Ue \ I), Dalias .\ Iuseum of
Natural Hst ory (D.\ INH), I-I erbert S. Harris/ Robert S. Simmons
Prvate Collec tion (HSH- RSS), University of Kansas Mu seum of
Natural Hst or y (KU), and Un iversit y of Texas at Arli ngt on (UTA).
Each museum number ref ers to an individual spe cimen.
THE 0:ATU RAL HI STORY OF ~ I E X I C A N RATTLESi\' AKES 5
sured ca. 1.2 m total length. The courts hi p behavior det ailed below
was typ ical for C. atrox ob ser vations made by uso
0900h The femal e was pl aeed wth th e ma le.
0901 The ma le sensed her pr esenee an d began rapid tongue fliekin g
(1 pe r see ).
0906 The female responded dcfensvely for a Iew minutes, but began
to reaet to maleo
0911 Male raised an terior third of body 5-6 cm above th e surface. Spas-
modic jer king of ma le was apparen t th roughout enti re period (1
every 2 sec) an d ncrease d to 1 per seco
0917 Mal e sta rte d rapid ten gue fHeking on clos est part of body and be-
ga n to inves tigare entir c lat eral and dorsal body surfaee of female.
j erki ng of ma le became more int ens e ( 2 per 3 sec ) . }Olale held head
at 45 anglo wit h menta l scale t ouching dorsum of femal e. Head of
mal e pressed against dorsum of Icmale, and side -to-side mot ions
(approx. 2.5 cm ) were undertaken in clu st ers ( 4-5) at irregular
nterva ls.
0927 By movng in an an terior pl ane, male began to dr ap e lar ge radius
coils over femal e.
0931 Sequen ce Iasted un til most of ma le' s body was in con tac t with
femal e. Male att empt ed to insert hemipe nis but no externa! e versi n
of hernpeni s wa s in evide nce. Both sna kes lift ed the tail s at the
location of the cloacae and occas onallv Ilickcd th e ta ils wit h a
side-to-sde mo tion . The subs t rato was ' not touehed bv th e t ails.
Male cons tant ly moved ta il in an atte mpt to a lign cloacae. When
cloacae werc juxtapos cd, ma le tricd to sliclc t ail under tail of Icmal e.
0951 Cloacae were align ed and ma lc inserl cd. During th e latt er part of
courtship activity , femal e hegan twit ching wi th same Ircqu ency as
ma leo jcrking and twitching stoppe d wh en mule inscrt ed , Cour tship
last ed 20 min and no movemeut ot he r than stcady pulsat on of the
body near male' s tai! wa s in evi dcn ce after penetrat ion. Bot h sexes
would occasonall y drag each ot her slowly by t he attaehed hem-
penis; this behavor perhap s was iniliat ed hy exte rna] st imuli.
Act ua l copula tion last ed from 2-8 hours. Breeding dates wer e
recordcd from 21 .T an uary to 26 .T anuary 1974.
Variation of pre-coital bchavior was observed on 28 Fcbruary
1975. Dur ing th e courtship pcr iod, the mal e was loosel y looped
over the Fema le an d un dulated slowly, In aclcl iti on, th e male tri cd
to positi on his tail und er the cloaca of the female. I n coutras t to
th e beh avior observed previously, th e fcmal e raisecl her tail and
repeat edl y opened her cloaca. \ Vhen th e cloaca e we re juxtaposccl,
th e mal e inserted hi s hei uipcnis but no external ext rusion of the
hcmpcni s was no ticed. The ma le exhib ited pulsating throbs (once
every 3 seconds ) inte rmi t tentlv and th e snakes wcre in copula tio n
nearl y 7 hours. The femal e remainerl pa ssive throngh out the re-
mainder of the scquence. When the snakes wcr e placed in a larger
unit (1.0 X 0.8 X 1.2 m high ), th e femal e assumecl a verti cal an te rior
trunk posture as th e ma le clirected courtship behavior to her. This
may be a femal e rejec tion post ur e. Of nin e capti ve litt er s, t",a
\Ver e born in June, one in .T ul y, three in August , t\Vo in Se ptember
ancl on e in Oct ober. Sexual maturity \Vas reached in 30-36 months.
4 SPECIAL OF NATURAL HISTORY
ACCOUNTS OF SPECIES
Crotalus atrox Baird and Girard
Crotalus atrox is one of th c lar gcst and most aggressvc of the
Mexican rattlesnakcs ( Klauber 1972 : 448 ). Although it has been
confusc d wit h C. scut ulatus ( Klaubcr 1972: 541 ) , C. atrox is gen-
erally wcll known throughout its range. These rattl csnakes are
fou nd in a wide variet y of habitats at varying el evations. Snakes
from southern Oaxaca (Cloyd 1940: 206; Klauber 1952: 102) and
ce nt ral Veracruz (Klauber 1952: 103) present probl cms in C. atrox
di stribution.
Specimens obtained by lIS are from the following local iti es :
BAJA CALI FORi\ IA: Progreso, SOl\' ORA: 4.8 km S Sonoyta,
Caborca, Es queda , CHI HUAHUA: 16-32 km S Chihuahua (4 spe-
cimens), near El Sueco; NUEVO LEN: 16-48 km NE Sabinas
Hidal go (10 specimens).
In Mexco, C. atrox inhab ts deserts, pl ains, grass lands , and Ioot-
hill s fr om ncar sea l evel at La Posa, Sonora (Taylor 1936: 497) to
at lcast 2440 m near Alvarez, Sa n Luis Potos (Klauber 1972: 5:27).
Indivdual s are gcne rally Iound in aricl country and are not neces-
sari ly assoc iated with rocks, Mcsquite, dr'y wash banks, cr eosote
bushes, and burrows provide likely retrcats. Taylor (1936: 497)
found thi s snake common along th c seashore wher e shrubs aneI
sandy beach es met. \Vc fou nd C. at rox especially pl cn t ful in mes-
qu ite grass lands no rth of Sabinas Hidal go in Nu evo Len . Several
snakes have bcen observe d in t his arca along road s at night during
th e hot sumrn er months. Hardy aneI ;"lcDi armid (1969: 213) con-
side red thi s spe cies to be rather un common in thc lowlands of ex-
treme northcrn Sinaloa, and R. T. Busey (pers. comrn.) stated that
C. basiliscus was mu ch more commo n th an C. atrox in thc nearby
lamos arca of southem Sonora. In nort hern Sonora, near Esqueda,
th ese snakes are not un common in sa ndv, creosote habitats.
Throughout its ra nge, thi s ra tt lcs uake is mostly nocturnal , at
lea st dnring th e summer mo nths . xl ost snakes obtaineel by us have
b ccn collccted while crossing roads during th e carly ovening . Some
individ ual s have been see n in th e carly morning or on cloudy days,
but th ey are ge nerally not ac tive and are usuall y coil ed in th e
mo uth of a burrow or under the cover of available brush.
We hav e observed rituali zcd cornbat between mal es in captivity.
Wiley (1929), II ocsslc (1963) and Petzold (1963) have discussed
reproduc tivo bi ology and mating beh avior in this spe cies. We ob-
served cight C. ai r ox courts hip sequc nccs in capt vity and a gen-
eral patt ern wa s apparent. The cnclosure in which obs ervutions
were made measured 80 X 40 X 40 cm hgh, and th e snakcs mea-
6 SPECIAL OF NATU RAL HISTORY
Litter size vared fr om six viable young (4 infertile egg masses) to
l 5 viable young ,
Vegcta t onal characteris t ics associate d with the habitar of G.
ai rox in Mexico inci udc the Season al Fonnation Series, the St eppe,
Thi cket und Scrub Desert as defined by Wagner (1964). Cl oyd
(1937) aneI Lowo (1964) described th c faunal a rcas inhabitcd b y
this species in Ari zona.
Crotalus liasil iscus ( Cope)
Crot alus basili scus basili scus. Crotalu s l). basiliscus is on e of
the Iarges t aneI gent lcs t of t he Mexican Crotalus. Cl oyd (1940: 161),
Bogert a ud Oli ver (1945: 394), and Kl auber (195l : 87) have corn-
mente d on it s sta tus in rela ti onshi p to G. 1Il0/ 0SS11S. As far as is
known, t he clistribut iou of these two species usually does no t ov er-
lap, b ut Hardy an d (1969: l 16) gave reasons for be-
lcving that the two species live sympat rically in northern Sinaloa.
Furthcr nvcst igat ion is necossary in thi s ar ea . Kl au ber (197:2) indi-
ca tes an overla p in the ranges of G. !J . oaxacus an d G. 1/I010sslI s
ni gresccn s in Oaxaca. \ Ve hav e founcl G. !J . bosli scus to be a COIll -
mon snakc (30 specime ns) of Xlexicos west coast wi th in divi duals
being observed in t ropi cal th orn for est s and, to a lesse r degrec, in
tr op ical dcciduou s forest s.
records of G. /J. basiliscus are avai lablc, no doubt owi ng
to th e fact th at thi s ra t tles nake is vcry common. Examples of thi s
snake are kn own from extreme southcrn Sonora, south through
Si naloa, Nayarit , Jali sco, Colima, an d weste m Michoac n where its
ra nge apparent ly t erminar es at t he Ro Balsas. 1t would no t b e
surprising, however. to find thi s ra ttIcs na ke on thc coast al pl ain of
Gue rrero where topographical sirni lari ties to it s known habit at are
apparent. Spccimens ob tained b y li S are fr om the foIlowing 10-
calit ies: SINALOA: 4 km S Santa Luca, 5 km \V Concordia;
l':AYARI T: 4.8 and 8 km E San Bia s; JALI SCO : 8 km 1\ Ciudad
Guzmn, Tamazula , 1'. flCHOACAN: Dos Aguas.
Grata/lis 1J. basiliscus is ge ne ra lly a Iowl and resdcnt inhabiting
the previous ly mentioned t ropical thorn or tropical deciduou s for-
ests. Klaubor (197:2: 5l8) re po rtc d th at a specimc n fro m Apat ziug n,
xli choacn, was coIlecte d at 975 m elev atio n in th e arid tropi cal
scrub for est. 1le fe lt thi s t o be th e IIpper limit s of the alt itudinal
range of this form, but he was no doubt unfami liar w th the speci-
men fr om 19.1 km :t\E Santa Luca in Sinal oa (KV 78966) which
was collected at 1940 III in a humid pine- oak forest o \Ve ca n exte nd
th e altit udinal limits wi th a recentl y co llecte d specimen (VC \ f
51.3 1. '3) [rom near Dos Aguas, 1'. 1ichoac n. This snake was Found in
Jlll y 1974, well wit hin th e co nfines of t he Sierra de Co alcom n at
an e levat ion of :222.5 m. The rat tles na ke was located wit hi n 3 m
of a stream in a canyon bottom in th e humid pine-oak forest. Local
TlIE NATU RAL HISTORY 01" ~ I E X I C A N RATTLESI\' AKES 7
residents rccognized th e individual , but report ed that C. b. liasil iscus
was not ncarly so common as the abundant C. pusillus or the fairly
rarc C. durissus culminatus . Dcspit c the fact that occasional snakes
mal' reach elevations abo ye 2000 m, we consi der such occurrences
rare, for C. h. hasili scus is appa reutly a lowl and fonn. Most of our
specime ns ha ve come f rom bclow 1000 m in various tropical fores ts
(see Hardy and Mcfriarmid 1969: 51-SS). Anothcr snak e (UTA
R-6120) was found on th e south slope of Cerro Bar al osa in the
Sierra de Coalcom n. On 21 july 1976, an individual snake (UTA
R-6071) wa s found coil ed in shade among rocks at 91.5 m at Plomo-
sas, Sinaloa, in a transition pine-oak and tr opi cal decduous fores t
(Fig. 4). This snake was coll ect ed in thc same rock slide as an
example of Crotalus ste jnegeri. C. h, basil iscus appears to be mos t
cornmon, or at least most active , during thc summer rain y season,
and most individuals are located crossing roads. Hardy and Me-
Diarmid (1969: 214) reported tha t these rattlesnakcs hael also been
found in thc dry scason in Sinaloa, al though thi s is thc pcriod of
least acti vit y.
Vegctati ve components associated wi th thi s rattlesnake are the
Arid Tropical Scrub Forest (Duellman 1965) and thc Tropical Thorn
Forest and Tropical Deciduous Forcst as defin ed by Leopold
1"1<; . 4. Habtat of Crot alus ha siliscu s basiliscu s, C. lcpidu s niacu losus and
C. stcincurri near Plomosas, Sinaloa , Mexico, Septe mber 1976. Crota/lis /.
nutcul osu s occurs in t he piue-oak Forest cover inj; the lop of the bl nffs. Crot alu s
st ciuegcri an d C. h, hasiltscu s are Iourul in th e tr op ical dccid uous fores t on
th e lowe r hill sides an d th e transiti on zone at the base of the bluff s, ( Pho to-
graph by Jona than A. Campbcl l. )
8 SPEC IAL PUBLICATI O:,-: - :-. ruSEDr\1 OF NATURAL HISTORY
(1950). According to Goldman (1951), rn esquites, acacias, wild figs,
Spanish cedar, palo mula to (probably Bursera grandifolia), w ld
guava ( Psid iuni guaiac a), Castilla elas t ica, silk-cott on trees aneI
palms grow in t he area inhabite d b y t hi s rattlcsnake.
Crotal us basili scus oaxacus. A Oaxacan subspecies, C. b. oaxa-
Cli S, separatcd ge ographically from C. b. basili scus by the entire
statc of Guerrero, is a ra t he r ncgl ect ed form fr om the interior h igh-
lands of ce nt ra l Oaxaca, Described in 1948 by Clo yd fro m two
specimens, th is subs pecies has rernained rare in syste mat ic collec-
tions. Much of the veget at ion in the area once inhabite d b y C. b,
oaxacus is now eithe r destroyed or near dest ruction. This rattlesnake
was appare ntly quit e cornmon in the Va lley of Oaxaca, but is now
rarely encountered there, mos t individuals now being taken from
the mount a ins surrouncling Oaxaca. These mounta ins exhibir t ypical
pine-oak habitat where roek outc ro ps ar e occasiona lly Iound, aneI
these area s are su pe rficia lly similar to the h bitat of C. uiolossus.
Thcy seem to p ref er the more heavil y for est ed arcas and , accord ing
to th e local pcople, are ofte n scen among rocks where C. intermedius
glo!Jdi is Iou nd. The first specimen ob taine d by us frorn El Tejocote
(also known as Tejocotes ) was located on a ste ep hillside within th e
pi ne-oak forest at 2.438 m. \ Ve had previo usly coll ected in this area
rather extensvcly for t h ree ~ ' e a r s , and this was th e Hrst example
of C. b , oaxacus we liad encountc red. A sccond spec imen was se-
cured along t he Ro Co lorado nea r El Tejocote during the first
wcek in February 1976 at an elc vat ion of 1982 m (J. A. Campbell ,
pers. co mm. ). Anot her was colle cted on 24 March 1976 in a pine-
oak fores t at an elevation of 2285 m (UTA H-6820). An indi vidual
(UTA U-6060) was fo und killed on th e ro ad on 31 [ ul v 1976 at 1030
h in an oak fores t (1932 m) at 3.5.5 km 1\W Telixtlah uaca.
C. b. oaxac us inhabi ts the Montano Fonnation Series aneI Mon-
tane Thcket as defined by Wag ne r (1964) and Pi ne-Oak Forest
(Duellman 1965).
Crotalus catalinensis CJiff
Crotalus cataiine nsi s is fou nd onl y on Santa Catali na Island,
Baja California elel Norte , an d is relat ed (ecologically) mo st closely
to C. ruber lucasensis. The islan d is rocky aneI barren wi th sparse
bru sh anel cacti (Kla uber 1972). The t empera t ures of th e coastal
islanels ar e similar to the adjacent mainland (Nelson 1921),
One of th e ma le snakes mainta ine el by lIS, born duriug September
1975, exh ibited heael-bobbing and tongue-Hcking courtshi p be-
havior toward a nother indivi dual (KU 173096). Another pair copu-
lateel on 15 january 1978. Thi s ra tt lesnake is found in the Seasonal
Forrnation Series anel Cactus Scrub as defin ed b y Wagn er (1964).
TIIE NATUR.AL HI STORY OF MEXICA:-': RATT LESNAKES 9
Crotalu s cerastes Hallowell
Thc two subspecies of thi s ra t tles nake that occur in Xlexico,
C. c. laterorcpeus and C. c. ccrcobo mbus will be rcferred to as C.
cerastes, since their habitat s, habits, an d behavior ar e ess entially th e
samc . Approximatel y 300 individuals have been observed by us on
the eas te rn drainagc of th e Sierra [ arez an d Sierra de San Pedro
M rtir as far south as Pu ert ecitos, a fishing village on the GuIf of
California in Baja California del Xorte. I n Sonora, C. cerastes is
located north an d wes t of th c Nognles-Herrnosi llo-Cuaynuis high-
way (Route 15), with the heaviest concentration of snakes being
Iound in the Desierto de Altar, an arid sandy desert in the extreme
northwest part of the state .
Crotalus cerastes genera lly prefers the sandy arcas of its clesert
environment , alt ho ugh it is not restricted to that particular habitat.
Dammann (1961) reported that C. cerastes populat ions may be af-
fected by the amount of vegeta tion presen t. In Arzona, thc popula-
tion den sit y of thi s rattlesnakc tended to de cr ease as thc vegetntion
increased. Too much vegcta tion may l imit the unique locomotor
abiliti es of thi s spe cies. These rat tlesnakes can be found on hard
stony terrain, such as the Co t tonwood Springs arca of [oshua Tree
Nat ional Monument. These part icular individuals are not as com-
mon as those Iound in sandy si tuations, but th ey appcar to be
slightly larger in body girth. Thi s may be dil e in part to a diff er ent
di et as mammals rather th an li zards ar e more plentiful in these
ar eas. This terrain also has a more luxuriant growt h of desert vege-
tation. Gen erall y, C. cerastes will commonl y be fOUJHI on sandy
alluvial fans, sand dunes, sandy washes, and the fringes of desert
dry lak es. The greates t concen trat ions of these rattl esnakes ar e
usuall y found in areas where wind-blown sand form s srnall mounds
at the bases of creosote bushes (see Brown 1971). \lill er and St eb -
bins (1964) [ound C. cerastes resting undcr bushes. Thc arcas around
San Felipe, Baj a Californ ia, Yumn , Arizona, and ;\!exico Route 2
between Mexicali and Sonoyt a ar e good examples of thi s type of
habitat. In these stuations. C. cerastes is easilv located, usuallv at
the mouth of a burrow, or beneath bushes whe;e shade is availahl e.
One snake from north of Puerto Peiasco, Sonora, was Iound in a
creosote bush about 30 cm aboye th e sandv soil . This individual
appe ared to be basking rathcr than hunting, as th e specimen was
not al ert and did not seem to sense our presence. In th ese ar eas
the sand shifts consta ntlv and C. cerastes is oftcn obs ervcd coilcd
near a bush, parti all y c ncealed by sand. Browu (1971) suggest ed
th at th e unusual crate ri ng beh avior serves primaril y for th cnno-
regul at ion an d sccondari ly for concealment during the da y, whereas
at night the reverso is t rue. Tndviduals have al so been found
wit hin and under th e wreckage of di lapidated buildings wher e as
10 SPECIAL PUBLICATION-i\lUSEm, 1 OF NATURAL HISTORY
ma ny as five specimens have been locatecl in one sea rch. This
rattles nake is mostly nocturnal but may b e seen cluring th e day in
early spring and la t e fa ll o Activt y cycl es seem to b e related pri-
marl y to th ermal cons iderat ions (Brown 1971). Moore (1976) sug-
gestecl that du e t o circula tory adjus tment, ti ghtly coiled, inact ivo
sidewinders (anel C. nutchelli pyrrlws) were ablc to conserve heat
more effect ively th an uncoil ed snakes. Often, ten or more C. cerastes
ma y be encounte re d by clrivi ng ronds at night in reas where they
are common, with as many as 30 occasionall y being seen. Peak ac-
ti vity appe ars to be lat e May and early [une, partcularl y in th e
hours just af te r sun down. Siclewinders will often coil on asphalt
roads whe re au tomobiles ca use hea vy mortality. Brown (1971) felt
that the sna kes us ed roads as a source for irradiated heat.
Sidewinders are ra ther pugnacous and will strike readily. They
will often turn an d bite when restrained, a characte ri st ic usually
associatcd wi th monta ne fonns (C. icillardi, C. poltjstictus ), They
pref er a di et of l zards ( Unta, Dipsosaurus, Cnemulophorus, Uta,
Crotapl ujtus} in capt ivty , and capt ve sidewndcrs will often refuse
mice, Imt rca d ly consume an iguanid lizard. \Ve observed a captve
adult femal e C. C. cercobombus feeding on three of her newborn
young.
The vc ge ta tonal reg imes inhabited b y this species include the
Seasona l Fonnation Series an d Cactus Scrub as defined by \Vagner
(1964). Lowe (1964) dcscrbed th e habtat of this snake in Arizona,
Crotalus duri ssus Lnnaeus
The Neotropica l rattlesuakes, C. durissus, are among th e largest
and some times most aggres sive rattlcsnakes indigenous to Mexico.
\ Vith the exception of an isol at ed subspeces, C. d. iotonacus Found
in Trunaulipas, Qu er taro, Veracruz, and southe rn San Luis Potos ,
snakes of the C. durissus complex a re inhabtants of thc country
south an d east of th e Xlexican Plnteuu, including th e entire Yucat n
peninsula. Despte the fact th at th e common name given C. durissus
suggests a jungle inhabit ant, thesc ra ttles nake s rarel y will be found
in such moi st plac es, much pref erring savannahs or part ally wooded
hi llsi des ,
Crotalus durissus durissus. Crotalus d. tluri ssus is a rcsiclent of
the Xlexi ca n sta tes of Tabasco, Chiapas, central Veracruz, aneI
southeaste rn Oaxac a (Klauber 197:2). Klauber (1952: 65) cite cl difler-
ences in snakes fr om near Jalapa ancl Orizaba, Veracruz, and sug-
gest ec1 tha t with examination of mor c specimens this population
may meri t subspecific recognit ion. Crotalus d. tlurissus has been
recorded by us from th e Followi ng localities: CHIAPAS: 12.8 t o
] 8 km l\E Tapana t epec, OAXACA: 19.3 km W Tehuantepec.
Ths rattlcsnake prefer s d ry savannah habitats with Frequent
rock outcrops. Hartweg and Olive r (1940) characte rzed th e hahitat
12 SPECIAL PUBLI CATI ON- r-. m SEUM OF NATURAL HISTORY
FiG. 5. Habi t at of Crotalus duri ssus culminatus, near [ojutla, Mor elos ,
Mexco. Opcn tropica l deciduous thom forest charact eri zed by limestone
outcrops. (Photograph by M, Gra nger.)
this rattlesnake avoidcd lowl and sit uations, snakes from Copala,
Guerrero, and La Pl acita, Michoac n, have Leen recorded from
near sea level, This lowl and habitat can be descrbed as arid
tropical scrub fores t, characte rize d by Irequent rock outcroppings.
Davis and Smi th (1953: 141) found C. d. culniinat us common in
lowland mountains south of Tepaltzingo, Morclos, on the northern
slopes of t he Ho Bal sas basin. This urea is al so interlaced with
extensive limestone ou tcropp ngs . Duellman (1961: 121) commented
that 18 specimens from El Sab ino, Michoac n, wcr e found near the
upper limi ts of the arid scrub fores t (1050 111) on th e lower slopes
of the Cordi llera Volcnica. A spe cime n from near Morelia,
Mi chonc n, t he northern limit of th e range of C. d. culminatus,
was taken on the edge of a lava flow at 1982 m in a mesquite grass-
land habitat with isolated sta nds of scrubby oaks. Throughout
much of its rango, this subs pecies is gc nc rallv nocturnal , and seems
to reach a peak period of ac tivi ty duriug the summer rainy season,
as do most Mexican Cro talus . A capt ve pair was observed copu-
lating on 4 Februarv 1978.
Cro talus d. cull/l!nalus inha hits the Aricl Tropical Scrub Forest
(DueIlman 1965) and the Temperare Pine-Oak Forest as definecl by
Leopol d (1950).
Crotalus durissus totonacus . The northeastern subspecies, C. d.
totouacus, is probabl y the leasl stu dicd Fonn of C. durissus. Since
its descri ption in 1940 (Cloyd and Kauffelcl), Iew specimens havo
TlIE NATUR.AL HI STORY OF 1IEXICAN RATTLESNAKES 11
of thi s sna ke in the vicnity of Teh uantepec as pl ai ns. Klauber
(1972: 531) suggeste d that th e habit at of G. d . durissus was sorne-
what like th at of th e arid sou t hwcstern Uni ted States. Specimcn s
obt aincd by us from ext re me southeastcrn Oaxaca and southwest-
em Chiapas were fou nd to be very co rnmon in rocky, grassy situa-
ti ons where forest s had been partiall y cIea red. I n late Jun e, three
weeks aft er th c rainy scason had bcgun, ind viduals were found
at night on the road between Tapan at epec and Rizo de Oro. xlost
were obscrved on evenings when rains were in th e fonn of light
drizzl e. Cr otalus d. tlurissus rangos from ncar sca lcvel to at lcast
1585 m ncar Co mit n, Chiapas (Klauber 1972) and is found in th e
Tropical Hain Forcst as defincd by Wagncr (1964). According to
Goldman (1951), th e wild gourd trce, various cacti , Cassia, Acacia
[arnesiana, Prosopis ;uliflora, t wo speces of [atropha, Annona, two
0 1' mor e species of Ficus, l pomoea, two species of Pithe coll ohium,
fan pal ms and the gu sma (Cuozuma ul mij ol ia} ar e found in th e
ar ca inhabited by thi s snakc.
On 8 Decernber 1976, \ V. E. Lamoreaux (pcrs. comrn. ) introduecd
an adult malc G. d. CUII l(l/wnsis into an cnc!osure (100 X 80 X 75 cm
hi gh) co uta ining an aduIt Iern al c G. d. du rissu s. The maIc initiate d
head-bobbing movement s wi th ra pid t ongue-Hicking ove r th e fe-
mal c' s dorsum, The femal e remained passive und 110 intromi ssion
was ob servcd. The nex t day, th c pair was in copulo at 1300 hours.
The coital positi on of thc pai r was unusual for alth ou gh the snak es
were finn ly joined by the hcmi penis, th e tails were pointi ng in
directl y oppos it e pla nes. The ma le' s bod y pu lsat ed near th e ve nt
and th e tai! was curled upward. During intromission, th e male
opened h s mouth intermitteutly and b egan a seri es of vertic al head
and neck jerks with occasional tonguc llicki ng. Parturiti on occurred
on 10 June 1977, and prod uccd on e dcad a nd ninc viable neona t es.
Crotalus durissus culminatus. Crotalus d. culminot us, a Pucific
Coas t ra ee, ra nges From uear sea lcvcl ncar La Pl acita, Mi chonc n,
and Copala, Gucrr ero (Wayne Seifert, pers. comm. ), t o 1982 m near
Xloreli a, Michoac n. Thcse ra tt lesnakes may reach ele va t ions uear
2285 m in th e Sierra de Coalcorn n whcre we have seen tanned
skins, Local rcsid euts have dentifi ed live indi viduals, but this
idcntification should remain tcnt at ive until mo re r eliabl e data ar e
secured. This subspec es occurs fr om near th e Col ima border at
La Pl acita, Michoac n, north ea st to Morclia, east through southe rn
Mor elos and weste rn Pue bla to ext re me southweste rn Oaxaca , in-
cIuding th c cnt ire sta tc of Guerrero. Specimens have b een recorded
by us fr om: ncar La Placi ta , 7 km W Morel ia
(J. A. Campbell, pers . comm.), Morel ia : 5 km W
Jojutl a.
Grata/lis d. cuhni natus is an inhabitant of rou gh, rocky, generall y
arid habit ats (Fig. 5). Although Klaub er (1972: 530) th ought that
TH E NATURAL HI STORY OF MEXICAN RATTLESNAKES 13
reached collections, an d onlv recent lv, with th e con structi on of
new roads , has C. d. t ot onacus hecn ol;served with some frequency
by field invest igators. Records obtained by us are fro m
PAS: 48 km N Soto la Marina, 16 km N Aldama. Four subadu lt
indiv duals were found ki lled on t he road in Tamaulipas: El Carrizo
at Kil omet er Post 26 on 2 Ma rch 1977 (1-:U 17482.5),20.8 km N Soto
la Ma rina on 29 April 1977 (1-: U 174826),38.4 km N Sot o la Marina
on 29 April 1977 (KU 174827), 33.6 km N Sot o la Mari na (1-: U
174828). To pograph cal similariti es suggest that C. d. tot onacus
may be found in the sta te of Hidal go.
Lkc most raees of C. du rissus, C. d. totonacus is typicall y a
lowland inha hi ta nt, although specimens ha ve been report ed fr om
1680 m in th e lower cloud for est of t he Sierra de Guatemal a (Ma rt n
19.58). Throu ghout most of its run gc, th e arca s inhabitcd by thi s
snake are ei the r tr opical thorn or tropical deciduous fores ts markcd
by di stinct wct -dry seasons . Two spccimen s (both juven iles) from
east -cc ntra l Tamaulipas, coll ected by [ ohn E. Jo)' (pers. comrn .) on
7 and 8 August 1975, wcre found active short ly a te r dark during
rain showers (Fig. 6). A juvenil e (UTA R-6707) Iound k llcd on th c
road at Kil omcter Post 17 ncar La Xlari na Viejo on 2 March 1977
containe d rodent hair. Crotalus el . totonacus appcars to pref er arcas
around any type of wat ercou rsc, Loca l rcsid cnts ass ured us th at
this is wherc most of these rattlesnakes wcre seen, hut Dixon el al.
(1972) found this subs peci es in eas te rn Quertaro at 1.585 m in a
,
/'
6. Subadu]t Grata/lis durissus tot onacus. Specimen from 48 km N
Soto la Marina, Tamaulipas, :\ Iexico. ( Photograph by j onathan A. Campbel l.)
14 SPECIAL OF NATURAL HISTORY
pine-oak for est near no st anding wate r. P. Pritchard (pe rs. comm.)
Iound a largo individual on the bank of a small dam. These rattle-
sna kes are common, but as is th e case of most forms of C. durissus,
the peak period of ac tivity seems to be limited to the rainy season.
Growth rutes of captive young are th e mo st rapid of any rattlesnake
taxa mai ntai ne d bv uso
C. d. ( 0( 0 /10 CU; inhab ts the Tropical Thorn and Tropi cal De-
ciduo us Forests, some elements of th e ternperate dcciduous fores t
as dei ned by Leo po ld (1950), and th e Seasonal Formati on Series
and St eppe, Thicket an d Scrub Descrt as defined by Wagn er (1964).
The arca suppo rts mesquite, huisache, ebony a nd other shru bs,
Along th e str ca ms , bald cypresses ( Toxodium} and cottonwoods
are corn rnon (Goldman 1951).
L. A. Mitchell an d J. E. [oy (pers. comm. ) discovercd a mal e
C. d. totonacus measuring 1.7 m total length, in combat with an
nd ult ocelot, Felis pardalis, duri ng February 1977. The animals
wer e bclow a thick canopy of th orn scru b in early aftcrnoon and
the ground beneath th e canopy support ed littl e vegetation. A dis -
tance of 1.3 m se parat ed th e combatants, and th e snake was in a
defensivo posture typi cal of C. durissus, and rattling vigorous ly.
The ocelot remained immobile wit h th e right forcleg rased . When
the snake was exte nded fuIl)' with it s mouth open af ter a strike,
th e ocelot raked wi th it s claws within th e snake's mo uth, th en
wi thdrew th e paw befo re th e suake could close it s mouth. Lat cr
examina t ion of th e oral cavitv al1CI th roat of th e sna ke rcvea led a
number of lacerations in the soft t issue and masse tcr mu scles which
we re intl ict ed b y the ocelot , but the interstitial skin and scales ex-
terior to the lacerat ions wcrc 1I 0t al'ecte d. Lacerat ions werc fou nd
on th e dor sa l as pe ct of th e head and ueck, but not near th e pu nc-
tures wit hi u th e suakc's mout h or throat . The snakc's 10\Ver jaw
was out of alignmen t and the mandibular bone was visible when
the mou th wa s open. A largo female C. d. i otonacus wa s locat ed
15 mi nutes later, w thi n 12 m of the male. The ma le ra t tlesnake
was treated for its injuri es, but succumbed abou t two wee ks later
(1-.: U 174824). The snake regurgi tated a large numbcr of ascarids
during the two weeks prior to its death.
CrotaIus duri ssus tzabean. The Yucatn race of th is spccies,
C. d . tza bca n, unlike C. d . durissus and C. d. culn nat us, is st rictly
a Iowland inhabitant from th e Yuca t n pcninsul a, and ran ges fr om
eastcrn Tabasco to inelude all of Campeche , Qui nt an a Hao, ami
Yucat n. Nei ll and AIIen (1959, HJ60) reported on this ra tt lesnakc
in British Honduras.
Crotalus d. t zabcan is qui te co rn mon throughout th e Yucat n
peninsul a. Ducl lman (196.5: 611) descr bed its habitat as scrub
forest with numerous limeston e out crops. Four specimen s ob taine d
by us \Vere found bet\Veen Kantunil and Chichn llz, Yueatn ,
TIIE NATURAL HI STOHY OF :-' IEXICA:\ RATTLES:\AKES 15
a te r 1600 hours (UTA H-5659-60), fo llowing a two-hou r rainshower,
A mal e, taken on 18 September 1974 betwee n Kantunil and
Chich n It z , Yucat u, and a femal e obtained on 29 August 1969
at Uxmul, Yucat n, were placed togcth er on 11 October 1974. The
Following observations wcre recorded by Terr y Huls ev (pe rs.
comm.). The female exhibi ted nervousn ess an d refused to coil nea r
the mal e. On 24 Oct ob er she was pl aced in isola tion an d again
introduced on 7 No vember, The mal e shed on 29 December and
the femal e was coiled next to the mal e for the first time since they
wer e introduced, Courtship behavior was seen on 30 December.
The mal e began rapid head-bobbi ng aneI tongu e-Hicking. He ap-
pr oached th e female and tapped her do rsum repeatcd ly in the
vicinity of th e 13th and Ith scalc rows . The male' s activity Iast ed
ca. 30 seconds a ud he remained immobile for ca. 30 seconds, At the
end of each period of acti vity, th e ma lc's t ail eucirclcd thc female
in the vici nity of th e vent and he jerked violently. Ceu crall y. the
head-bobbing motions were direct cd antcriorly. Pre-coital activi ty
was continued afte r th e mal e di sconti nued th e cloacal scarching
moti on . The sna kes did no t copulare. 011 2 January lm.5 th e cage
was sprnyed with wat er at OS30 hou rx an d th e male began twit ch ing
shortl y th ereafter. The snakes wcre found in copulo at 1.500 hours
and th e ma le slowly moved hi s tail in a lat erall v c1 irect ed twi tchiug
mot ion, The fe male was wci ghcd periodicall y after co pulat iou anel
the wei ght incr ease was as Iollows : 2,1 Fcbruary 1975 (2. 1 kg); 20
Ap ril (2.6 kg ). 12 (2.7 kg): 2.5 [ ul y (3.0 kg). Twent y-on e youn g
(two dead. l( lj 158547-8) wcrc bOJ"l1 0 11 31 Augu st ami wcre imm edi-
atelv we ighed and mcasurcd, TIJ e ran gc of va riation is as follows:
to ta l len gth 290 -3.50 mm . mean 316: ,, 'eight 18.4-26.S g. mean 23.'1.
Othcr pa rturition dates are 31 Augu st lm.s (LT T.\ H-67:32. GS02). 2
August 1976 (UTA H-67.'3.3) aud 3 Augu st (UTA H-6734-3.5). Figur e
7 re prcscnts on e of th e young at 35 inonths.
Crotalus d. tza bcan is Iound in the Tropi cal Evergree n and
Tropical Hai u Forest as defiucd Lcopold (19.50). Thc arca xup-
ports Entcrolobt nm cijclocarinnn , two or more specics of Fi cus,
Spanis h ccdur, Iogwood. t wo spc cics of silk-cotton, the chico znpo te
(Ac!lras zapoia}, palo mula to (Bu rscra}, t wo or more species of
Cassia, giant nettle (Urera caracasaua} ancI Agave (Coldmuu 19.51).
'rhe subs pecies of C. dllrissus are vcry ll np redi ct able ill telll -
pe ra lllen t. At ti mes th ey are aggressi\"(' aneI stand th ei r grouJl(I,
whereas at other ti mes th ey are placido Our specime ns ex-
hibited both of th ese behaviora l ext re mes. C. dllrisslIs ar e at
Icas t part ially noctuma \, anel al! sce lll to reach peak ac tivi ty during
the Slllllmer rai uy season. Young of thi s species h,1\"e been collected
frolll late Jnne throu gh late August.
16 SPECIAL PUBLICATION-:VWSEUM OF NAT URAL lIISTORY
'j
l
1
..'
r
i r
I .
I r'
FIG. 7. Cratal us durissus tzabca n. Sec texto ( Photograph by [ onathan A.
Campbell. )
Crotalus CIl !JO (Cope)
Crotal us C Il !JO. though wdesprcad throughout Baja Ca lifo rnia,
rernains bi ological ly ncgl cct cd in literat ur c on rattlcsnakcs (Klaubcr
1931). It is characte rized by an unusually small head and rathcr
large ra ttles , also t raits of C. tigris, a sp. -cies not Iound in Baja
Californ ia. Haces of C. C Il !JO are moderately sized rattlesnakes, and
t hroughout much of th e ra nge, are lowlaud or mountain desert
dwell er s. Van De nburgh and Slcvin (1921) found thi s rattl esnake
around human habitations,
Crotalus en yo enyo. The subspeces C. C. e ll !Jo is a rnther com-
mo n rattles ua kc in th e Cape region of Baja Ca lifornia del Sur (Fig.
8). Speci mens are known from El Mrmol , Baja Califo rnia del
No rte, sou th through out th e pen nsul a to th e southcr nmost point.
Specimens obtainc d by liS wer e collectcd at San Antonio (64 km S
La Paz) and 14.4 km N Buen a Vista (B. Tomberli n, pcrs. comm.).
Crotalus c. CIl !JO is an inhabi tunt of arid, ru ggcd deserts and
desert mountains t hrough out its range. Kla ubcr (1972: 531) sug-
gest ed that snakes Iound fro m La Paz to San Jos del Cabo were
from an ari d regi on co ntai ning a profusi n of cacti , xerophytic
shrubs, and rocks. In cent ral Baja Califo rnia, th is ratt lesnakc Iives
in association with la rge boulder s, th e gant ca rdo n cact us (Ccreus
pringjei), the boojum (l dra), and th e Ion g-li ved elophant tr ce. B.
Tornb erlin (pcrs. com m.) comme nted t hat, in th e arca south of La
Paz, thi s subs pecies lives sympatrically with c. rub er lucasensis
18 SI'ECIAL OF NATURAL llISTORY
California, C. e. [ur ous was known fr om Iew spcci me ns, no doubt
du e to the previously poor road condi tions whieh preventcd aeeess
to the opt ma l habi tat of th is snake.
The San Qui nt n Pl ain is characteristic of coastal, ope n, low-
growing scrub habitat s in Baja California del Norte. Klauber (1972:
531) descrbcd the area as muc h like we ste rn San Di ego Co unty,
California, bu t somewhat more rocky and with a Iighter brush
cover. Lowe and Norris (19.'54: .57) char ac te rized the area as con-
taining a markcdly more mesic clima te and biota th an the harsh
euviro nmcnts both southward and in the inter ior of the pen nsul a.
The average annual rainfall is abo ut 1:2.7 cm (Beal 1948), and coast al
fog produces some additional moisture in th e ar ca . On e specimen ,
obtai ncd by us near El Socorro, appcared to come Irom a sandy
hill wh er e vegetat ion was sparse, and ther e was a profusi n of
coastal ground cove r. Xlnny bu rrows we re noted. A specimc n col-
lccted by R. T. Basey in Oct ober 1972 (ambient temper ature 21C)
carne From a light-color ed mud wash wherc vegetation was ex-
tr emely spa rse (pers. connn.). This specime n was a young adult
Iernalc which lat cr gave birth to sevcn youug on :26 Augu st 1974.
To tal length of th e you ng wa s fro m :206-2:2:2 mm, mean :213. Dennis
Bost c (pcrs, comm.) rc la tecl that his scven specime ns of C. e. [ur cu s
from nea r Pun ta Camal were collccted wh cr e t wo alluv ial fa ns
met. Eight additional snakes were taken by ouc of us (BLA) be-
twecn 10-15 October 1977 in the vi c nity of El Rosario.
Crotalus e. [urous appcars to be at least parti all y nocturnal in
that two indi viduals recorded by H. 1', Basey and T. Portcr (pe rs.
eomm. ) wer e collec tcd on a road at night. The type specimen of
this race was taken at noon in the mouth of a small rod ent burrow,
The Scascnal Formation Seri es as defincd b y Wagn er (1964) is
typical of arcas inhabited by C. e. [ur cu s.
Crotalus inter medius Troschel
Thc sma llhead ra ttlesnakes. C. int crmedius, of southe rn M xico
r em an hiologically amoug thc least und er stood specics of rattl e-
snakcs, Li tt le published material on these snakes is availabl e, and
Few specimcns are avai lable in mu seum co llect ons, Most exa mples
of th is rattlesnake are Frorn southcastc rn Mexi co, but Ducllman's
(1961: 121) record of an individua l from Ce rr o Tanctaro , Mi choac n,
is indeed uuusual. \ Vhcn more ficld work is done in ar ca s bctwecn
known local ities, th c taxonom c status and geographic va riation
within C. inter mcdius should hecome more appar ent.
Crotalus interme dius intermedius. Crotalus i. int ermcdius has
been rccorde d from castern Hidal go, wes t cent ral Vera cruz, and
northcastern Puebla (Klaube r 1972: 39-40). Pi anka and Smith
(19;'59) found C. i. inter tnedius 16-24 km W ]apala, Verac ruz. A
single speeimen (KU 155530; Fig. 9) from near Cacaloapan, Pu ebla,
THE NATURAL HISTORY OF MEXICAN RATTLES NAKES 17
"
. , .
.... . . " .' .
# ..... . . ..
F IG. 8. Grata/lis cuyo cuyo. Specimen from14..t km r\ Buena Vista, Baja
Calfornia del Sur, Mexico. ( Photograph by John II . Tashji an. )
and, to a lesser degree, wi tli the spe cklcd rat tlcsnake, C. 111. mitclicll .
He also stated that spccimen s 01' C. C. CIl Y O ar e found in that a rea
from arid llat descr t plains to rocky dcscrt mo untai ns.
Cr otalus e. ell Yo is nocturnal f01' a good portion 01' the year, nnd
seems to rcuch a peak activi ty pcri od in the ea rly fa ll wh en sout hern
Baj a California receiv es a fair amouut 01' its an uual rainfall. At
th is time it is cornmonly Iound crossing roads at night.
Ca pt ive reproduction in C. c. eruto has been rep orterl (Trvon
and Rad cli ffe 1977). On 15 j anuary 1975 court ship activity runoug
snakes mai ntai nc d bv us wa s observcd het wcen 1530-1700 hour s,
The femal e had just ~ h e d and the mal e rubbc d his mental arca on
the Iema lc' s dorsum with convu lsive Iorward jerks in the ma nuer
of C. ioillardi silus (described later in th is pap cr). During this pe-
riod the ma le tr ied vigorously to thrust his tail heneath the cloa ca]
regi on 01' the female, an d in sorne instances, th e malc's tail corn-
pl et el y eucircl ed the Iemal e's cloaca . The mal e would slide Iorward
1-2 cm b y extc nding his looscly draped coils and bracing his body
at the cloacal region of th e female. Lo", int ensity twitch es accorn-
panied th is behavior . Co pnlation wa s not observed,
Crotalus e. ell Yo is an inh abitant of the Sensorial Formaton
Seri es as defined by Wagncr (1964).
Crotalus en yo furvus. Thc Rosario ra tt lesnake, C. e. [ur ous, is
a northern fonn of th is species from the San Quintn Pl ain 01' thc
west coast 01' Baj a California del Norte (Lowe and No rris 1954).
Until recently, with the complet ion of a paved road through Baja
TIIE NATURAL HI STORY OF ~ I E X I C A N RATTLESNAKES 19
ca pturcd in the summer of 1974 by one of us (BLA), extends the
rauge of this fonn some 160 km (airline) to the sou th. This ar ea is
at an elevation of 2195 m and is dominated by high desert vegeta-
ti on where cacti , agave, and yucca ar e prevalent. A rattl csnake of
this mee ob tained by us was tak en at 099.') hours wh en sunlight
was dilTused and humidity wa s high. It was basking in parti al shade
un der a 1arge agave on a southe rn facing hill. Thunder storms oc-
curred the previous evening, but the porous limestone soil surround-
ing the coIl ecting site showed Iittl e evide nce of moi sture. The
vegeta tional h bitat preference of c. i. int ermedius incIudes the
Tropi cal Evergreen For est as defined by Leopold (1950).
Crotalus intermedius gloydi. A Oaxacan subspeci es, C. inter-
mcdius gloydi (Fig, 10), is a montano race from the mounta ins sur-
roun di ng the eity of Oaxaca. 1'0 our knowl edge it has not been
taken from th e desert ar eas of that region . In that respect , it
diller s grcat ly from th e nominate northern subs pecies. Three suakes
obtained by us have come Irom the humid pin e-oak forests aboye
2440 m (Fig. 11) an d another specimc n (KV 155529) from north of
I xtl n de ju rez at 3020 m. This 1att er indi vidual was found on a
wcs t -Facing slope th e day foIl owing a violent hail st orm wh ich left
ab out 5 cm of hai! on the ground. The da y it was eapturecI , how-
ever, was hot (27C) and sunny, The sna kc was fou nd in the shade
of an oak that had begun to sprout new foli age afte r being burned
in a Ei re. J. R. Dixon (pers. comm.) found a speci me n of C. i. gloydi
e
FIG. 9. Crotalu s int ermcdiu s intcrmediu s. Specimen from Cacaloapan,
Puebl a, Mexico. ( Photograph by John 1I. Tashjian. )
20 SPECIAL OF NATURAL HI STORY
cra wling up a vertical ston e wa ll at night in a rainstorm along a
highway cut 57.2 km N Oaxaca at 2286 m on 6 [une 1956. Other
spcci mc ns of C. i. gloydi collcc tcd by us carne fram near El Tcjocot c
(UTA R-4915, 5627, 5637, 7163), which is northwest of Oaxaca, This
ar ca has bc en extens vely logged, an d only sca ttered arcas support
rattl esnake populations, Six additional specirnens were collected
in July 1975 from near Cerro Ma ch n (Sierr a de [u rez), Suchixtepcc
(Sierra ele \ liahuatln) and El Tejocote (Sierr a de Cuatra Vena dos).
Other snakes from th e vicinity of El Tejocote are as follows: (UTA
R-4915) coll ected on 20 August 1975, (UTA R-5627) during August
1975 from a pine-oak, madro o for est at 2440 m, (UTA R-5637) dur-
ing August 1975 in a pine-oak for est at 2285 m, (UTA R-5791-2)
on 17 June 1976 in a pi nc-cak forest at 2285 m, (UTA R-6062-63)
on 31 Jul y 1976 in a pne-oak for est at 2285 m, (UTA R-6122) cluring
[uly 1976 in a pin c-oak for est at 2285 m, (UTA 1\-6229) clur ing
August 1976 in a pin e-oak for est at 2285 m and (UTA 1\-6356) dur-
ing June 1975. \ Ve have obs orved ri tualized combat between male
C. i. gloUdi shortly after capture.
The Montano Formation Series anel Mont ano Thicket as defined
by Wagner (1964) is cha racterist ic of the habitat of C. i. gloUdi,
and Ducl lman (1965) list ed it as a resident of Pine-Oak Forest .
Blue-Ilowered Solanuni , Ceanotlius COCru IC lI s, wild chcrry, and two
01' more spccies of oak s and pines ar e found in the area (Goldman
1951).
Cr otalus intennedius omiltemanus. A Guerre ran subspecies,
C. i. omiltemanus (Figs. 12, 13) is probabl y the most thoraughl y
studi ed raco of the smallhea d ra ttl esnakes. Most cxamples of thi s
suake llave been t aken in the huini d pinc-oak for ests (Pilllls herrerai,
P. pseudostrobus, Quercus sp .) whch surrouu d Omilteme in the
Sierra Madre del Sur (Davis and Dixon 1957, 1959). Nearly all
specimens (35) ob ta ned by us were collected within pine-oak for est
(UTA R-4707-10, 6232-33, 6245, 6821); only one indi vidual , collected
Oll 25 Mav 1975 fram 3.2 km \ V Omilt eme, was taken in an area
of secondary growth in the adjacent hardwood cloud for est at
2653 m by J. A. Campbell (UTA R-5626). This latter snake may havc
reached this area via a power line cut. These rattl esnakes also were
found only during th e major rainy scason which usually began in
\ lay. This ar ea may receive precipita tiou cve ry month, but from
[ anuar y to Ma y, it is rather dry. C. i. omi ltemanus is appa rentIy
a rock-d well er , and none of the specimcns obtained by us wa s
found any distance from a rocky ret reat , with the cxception of th e
cloud Ior est specimen. 111 most inst ances they we re found basking
0 11 racks, espe cially after a rain. One snake (UTA 1\-2813) wa s found
dea d on a grassy hillside on 19 May 1973; cause of dcath was not
de te rmined. \Ve coIl ected snakes fram 2075-2592 m, and doubt
that these rattl esnakes are found near Chlpancin go whi ch has an
THE NATURAL HI STORY OF !,,[EXICAN RATTLESNAKES 21
FIG. 10. Crotalus iut erm edius glor;di. Speci men fra m 31.3 km 0: Ixtl n
de J uar ez, Oaxaca, Mexco. ( Pholograph by John H. Tashjian. )
FIG. 11. Habit at of Croialus int erm edius glor; di, pine-oak forest 31.1 km N
Ixt l n de [uarez, Oaxaca, Mexcn, july 1975. Area had becn partially cut
and burned the pr evious year, characterized by epiphy tic pl ant s. (Photograph
by Jonathan A. Campbell.)
22 SPECIAL PUBLICATION-}, IUSEU},I OF NATURAL HISTORY
elevation of 1372 m (Klaubcr 1972: 533). Although C. i. omi ltema nus
has been definitel y rccorded only from the vici nity of Omiltemc,
it was to be expeete d th at with addi tional field eolleeting othe r
speeimens would be sec ure d throughout the llpper reach es of th e
Sierra Madre del Sur. Supportive evidenee of this sta temen t is in-
di eat ed by th e eolleetion of a specime n (UTA R-5638) on 28 May
1975 fr om 1.6 km SW Filo de Caballo by J. A. Campbell, Another
adult snake, observed approximate ly 1 km N Puerto del Gallo (some
60 km WSW Omil teme), escaped befare being ca ptured. The ele-
va tion at thi s site was 2950 m, and an example of Bothrops barbou ri
was d scovered nearby. Mel ansti c speeimens of C. i, omiltemanus
wcre eolleeted in San Vincentc, Guerrero (UTA R-4538-9). Five
newborn YOllng of this mee were Iound 1.6 km S Orn ilt eme in pine-
oak Forcst at 2286 m on 28 May 1975. Range of varia tion for length
and weight is as Follows: total lengt h 194-212 mm, mean 205;
snout-vc nt 182-195 mm, mean 190; weight 4.9-5.4 g, mean 5.2.
Rituali zcd eombat between adult males was observcd by llS among
four recent ly eaptured snakes .
Cumpbe ll (1977) examinc d the stomuchs of 18 C. i. omiltemanus
Irom Omilteme, und revealed that these snakes fed almost exclu-
sively on lizards (ge nus Scelcporuss wher eas th e stomaehs of ]5
exumplcs of Both rops undulatus from the same ar ca con tained
mostl y rodc nt remains,
Crotal us lepidus (Kenn cot t)
The rock ra tt lesna kes , C. lepidus, are typieal examples of the
various smal l monta no, rack dwel ling ra ttlesnakes inhabiting M xico.
AII Iour subs peces, Icpidus, kluub eri, IIU1CllloSlI S, and //1 orulus, ar e
gencra lly ti m d , ye t rather irritabl e and eurious at tim es. Through-
ou t much of the range of C. lepidus in Mexico , on ly limi ted numbers
of specime ns have been reeorded, probabl y bccau se th e terrai n
inhabited by th ese snakes is gcu erall y steep, rugged , an d inacces-
si ble. More specimens are nee ded from th e southe rn Iimit s of the
ranges to clarify th e relationsh ip bctween C. lepulus und C. tri-
seriat us (Gloyd 1940 : 81).
Crotalus lep idus lepidus, The nominare mee, Crotalus l. lepidus,
is foun d in pinc-oak for est s through mu ch of its range, though dcsert
populations are not uu cornmou. This snake rangos from southeas t-
ern ~ e w Xlcxico and Tran s-Pecos Texas , south th rou gh Coahui la ,
wes t-central Nue vo Len , eas te rn Zaea teeas, and nort hwest ern San
Luis Potos (Klauber 1972: 62).
Cloyd and Smi th (1942: 235) re porte d a spccimen of C. l. lepidus
from thc Sierra del Carmen at 1830 m on an open, southwes tcrn
Iacing slope where junipe rs were present. Tuylor (19.52) reported
a snake from Cerro Pe on Bl an co, San Luis Potos at 2928 m. The
si ngle specimen of th is ra tt les nake, obtaine d by us from La As-
THE NATU RAL HISTORY OF MEXICAN RATTLES NAKES 23
F IG. 12. Crotalu s int ermediu s ami ltem anus. Melanistic spec ime n from
San Vicente, Guerr ero, Mexco, ( Photograph by j oh n H. Tashiian. )
F IG. 13. Crotalus intennedius omiltemanus. Specirnen Irom near Omilt eme,
Gue rrero, Mexico. ( Photogr aph by [ ohn H. Tashj ian. )
2--1 SPECIA L OF HISTORY
ce nsi n, Nue vo Len , was found under similar conditions at 2075
m wh ere junipers, pines, and agaves wer e pr eval ent. The specime n,
a gravid femal e weighing 44 g and me as uriug 435 mm tot al length
(400 mm S-V), appeared to be assigna ble to e. l. morulus according
to Klauber ( 1972: 130) by its havi ng 28 crossbands , However , three
of th e six young born to thi s Iemale on 5 [une 1974 had Icwer th an
25 cross bands , ami three had 25 0 1' more (one mal e had 31 cross-
bands). Range of variatio n for length and wei ght for this brood
is as foIlows : to ta l length 165-190 mm , mean 177; S-V 150-173 mm,
mean 162; wei ght 5.0-5. 7 g, mean 5.2.
Mil st ea d, Mecham, an d (1950) found e. l . Icpidus
in a persimmon, shin-oa k associat ion, and th e Scrub Desert as de-
fiued by Wagner (1964) is its charac terist ic h bit at . Mintou (1959)
deseribecl the hab itat of thi s mee in th e Trans-Pecos area ,
CrotaIus lepidus klauberi. Crotalus l. klaubcri is normally a
resident of moderate elevations in foreste cl mountain s thraughout
most of its ra nge. Sna kes from ext reme west Texas in th e Franklin
Mou nt ai ns are an exception since th is mountain ran ge is ve ry arid
wit h lit tl e vegetat ion. Spcci mcns obta ine d b y us are from: CHI-
HUAHUA: 16 km S Chi hua hua. Arrovo Xlest efi o ami Caon del
Alamo in th e Sierra del Nido, ne ar \ 'il a Matamoro s:
near Villa Ocampo.
Most of th e data on e. l. kl aubcri obta ine d by us have been
from exa mples coIl ected in t he Chirica hua and Huachucn Mountain s
of sout heastcrn Ari zon a, \ Vitlt t he exce pton of th e snakes fr om
th e Sierra del Nido , Chi huahua, spccime ns obta inecl by us from
Mexico were coIlec te d on roads, and aIl carne fr om ge ne ra lIy th e
sa nie t ype of t er rain, primarily desert canyous with rack outcrops
or ledges, and sparse vegetation. e. l. klauher was collecte d at
clevations fro m 915 m sout h of Chihuahua to 1372 m in t he vicin ity
of Villa Ocampo, Durango, where grasses and oa ks fonn a savannah
envi ronmcnt. I n t he Chirica hua, Huachu ca, aud Sierra del Nido
Mo untai ns, e. l. klaubcri is oft en located along rocky st rea m bcds,
aud during the summer rainy scason is obse rved in open pinc-oak
fores ts. These rattlesnakes are most lv diurnal , at least in forest ed
mounta ins, and are most eas ilv on warm, humicl summe r
days wh en th ey reveal t heir prescnce by rattIi ng. Hitu ali zed combar
betwccn males of this subspecies has been obs erved (Carpente r el al.
1976). A fema le specime u, coIlec ted at II erb Xlurtyr Dain in the
Chiri cahun Mountains. Ar izona, gave bi rth to three young on 28
August 1968. Fo ur young were born on 30 Augus t 1975 to anothcr
female spccimen coIlecte cl 4.8 km S Mad er a, Chi huahua , and ran ge
of variation for length an d wei ght is as Iollows: tot al length 181-
196 m111, mean 191; S-V 164-177 mm, mea n 172; weight 6.3-7,0 g,
mean 6.6. A captive pair of e. l. klauheri \Vas observed to br eed
on 21 Febmary 1977. H. K. Guese (pers. eoml11. ) ob served a eaptive
TIIE NATURAL HISTORY OF ~ I E X I C A N RATT LES;" AKES 25
pair of this race from th e Sierra dcl Ni do, Chihuahua, engaged in
courtship acti vity on threc occasions between 14-28 Septemb cr 1977.
The male directed rapid head-bobs (3-5/ 5 sec) on th e dorsum of
the femal e. Tongue-flicking occurred at th e same speed. The
snakcs wer e di sturbed afte r 5 minutes anel di scontinu ed the be-
havior. Another pair of th esc snakes was observed courting on 11
Octobcr 1977, and th e male exhibited the same head-bobbing and
tongue-flicking behavior (D. G. Barker , pers . comm. ).
The Montaue Formation Seri es, Steppe Thickct and Scmb Des-
ert as defined by Wagner (1964) ar e habitats in which C. l. klauberi
typically resides.
Jacob and Alt enbach (1977) described sexual di chromatism in
body coloration in C. l. klauberi from New M xico, Sonora and
Chihuahua. Van Devender and Lowe (1977) reported similar sexual
dichromatism in thi s race from Chihuahua. We have see n twelve
snakes from Arroyo Mest eo (7 ma les, 5 femal es) whi ch exhibi r
sexual dichromatism. A female gave bi rth to five young on 18
August 1977. Two mal e young wer e only sligh tly mottled wi th
black speckling and black bands, whe reas the two femal e young
wer e gr ey, mottl ed wth black speckling aneI had dark grey bands.
The fifth neonate wa s defecti ve, exhibiting a fusio n of thc ven tral
surface in the lower cervical ar ca and the anterior tru nk region
(UTA R-6946).
Crotalus lepidus maculosus. C. l. macul osus (Fi g. 14) is a re-
cently described ra ce of rock ra tt lcsnake (Tanner, e l al., 1972)
whose geographi c distribution rernains vir tually unknown becaus c
much of it s habitat is inacccssibl c. It is a resident of the Pacfc
versant of th e Sierra Madre Occident al , and apparentl y intergrades
with C. l. klauberi on the hi gh Mcxicau Pl ateau in the vic nity of
El Salto, Durango,
Whor e they are most abuncl ant Crotalus l. maculosus aneI an-
other subspecies C. l. morulu s appurcntl y pref er similar habi ta t
conditions. Both races seem to pref er hurnid, pin e-oak for ests with
well-defined rainy seasons (Fi g. 15). We ha ve unsuccessfu lly tri ed
to obtain both subspeci es in the late spring. However , after th e
summer rains bcgin, the two subspeci es apparentl y sur face and
are quit e common. This is unlikc many populat ons of C. l. lepidus
and C. l. klauberi, whose combin cd distributi on divides the ranges
of C. l. maculosu s and C. l. morulus, and have adapted in some
ar eas to des ert or chaparral conditions, These popul at ons ar e active
above grouncl throughout th e year if th e weathe r is suffici entl y
warm (24C). Almost all specime ns of C. l. lIlaCUlOSlIS ob taincd by
us were secured under cloudy or partl y cloudy conclitions, and wer e
partcularly cornrnon just befare and af te r showers during th e rainy
season. On 10 Jul y 1973, a femal e was observed on a southcas tern
facing rock outcrop 4.8 km \V El Alazn, Sinaloa, with el even young
26 SPECLAL OF NATURAL HISTORY
F I G. 14. Crotalus lcp idus maculosus. Spec imen Irom Los Bancos , Durango,
Mevico. ( Photograph by [ ohn H. Tashjian. )
eoiled or draped about her. This rac e exhibits behavor t ypcal of
all subspecies of C. Iepulus in that it is al ert and quick to seek
she ltcr wh cn di scov ered, Crotalus l. ma culosu s posscsses an un-
usuall y small rattl e, and it is barcly audible cven in Iarger spccime ns.
The acce ptcd teehnique hi storicully employed to diseover C. lepidus
in its native habitat (by hearing them rattle) is virtually usel ess
whc n trying to collcct thi s subspccies .
Xlost of the snakes obse rve d by us, which total fort y sightings
(KU 15553:2-534, UTA R-5847), wer e nonnall y loeated by exploring
southe rn facing slopes wh ere an abundance of grass and rack s oceur
in breaks in th e pin e-oak for esto This subs pccies was not found in
rac k slides as were lllan y C. l. lcpitlus and C. l. klouhcri populations.
Howcver, C. l. maculosus is usuallv found near sorne kind of rack
cover. Lizards of the genus Sceloporus appear to be the main food
item of thi s ratt lesnake.
Other ra tt lesnakes found in assoeiation with C. l. maculosus ar e
C. p. pricci , C. 1Il010SS11S nigrcscens, aud possibl y C. icllardi lIle-
ridionalis in the eas tern part of the rangc. Sorne local residents
gave positi vo identification of C. IV. meridonalis from photographs
which we provi ded.
Crotalus l. uuiculosus inhabits thc Pine-Oak Forest and Pine-
Oak \Vood land as defi ned by Leopold (1950).
Croralus lepidus morulus, Crota lus l. morulus (Figs. 16. 17) is
the least known of the subspeeies of C. lepulus. This raee was de-
THE NATURAL HISTORY OF MEXICAN RATTLESNAKES 27
F IG. 15. Hahit at of Crotalus Iepidus maculosus, G. pricei pricei and G.
molossus nigrcscens. Los Ban cos, Du rango, Mexico. St eep hillsi de s covered
with pine-oa k Iore st , Th ese sna kcs werc most Ircqucnt ly cncoun tered in
open arcas of rack outcroppings. (Phot ogr aph by R. Ter ry Basey . )
sc ribcd by Klauber (1952: .52) from specime ns obta ncd fro m P. S.
Mart n, who coIlec tc el in th e Cmez Faras arca of Tarnaulipas.
Mo st of th e known specimcns havo bcen collectcd in th e Cmez
Faras arca , and an add tional specimen is known fr om Chiche (Mar-
tin 19.58: 78), which is northwest of Cuidad Victoria, Tamaulipas.
Rcc entl y, more snakes of this race have becn secured iu th e
Siena ele San Francisco (11. S. lI arri s, per s. comm.), and w have
locatcd an apparen tly isol atcd po pulation (19 spccmc ns) in th e
Sierra Mad re Oriental sorne 24-32 air kilor uctcrs nort hwest of
Galeana near the Nu evo Le n-Coahui la borcl er (KU 159360). No
doubt, mo re C. l. inorulus \ViII be collecte el whi ch \ViII filI th e pres-
ent gaps between th e Icw presentl y known localiti es.
Xlos t of th e ra tt lesnakes collcc tc d bv P. S. Marti n werc locat eel
on th e humiel, eastcrn slopes of the Sierr a de Guatemala at altitudes
ran gin g from 1190-1890 m. These sna kes we re usuall y found along
th o abundant rocky mounta u trai ls in th e Gmcz Faras area. Pines,
oaks, ami firs const tu te th e maj or vcgc tational format ion of thi s
arca, alt ho ugh th crc is an abundance of agave on th e stccp rocky
sides of th e Sierra. Martiu (1958) stated that C. l. morul us was
p resent in the upper part of the Cloud Forcst, but it s presen ce was
not confirme d in the Lo wer CIOlH] Forest. This arca is ext rcmely
ru gged, ami more exte nsi vo coIlec ti ng is needed to delineate the
extent of the ra ngo of C. l. morulus.
28 SPECIAL PUBLICATION-MUSEUM OF NATURAL BISTORY
"
... ~ .
FIC. 16. Crotalus lep idus morulus. Specimen from near San Antonio de
las Alazana s, Coahu ila, Mxico. ( Photograph by John H. Tashjian.)
, -.
"
"
'.'
'.
..*'. ;
,.
", . . ,
"
,
"
.,
,..
,
~ '
FIC. 17, Cro talus lepid us iuorulu s. [uvenile specimen bom to fema lc col-
lecl ed nea r San Antonio de las Alazanas, Coahuila, Mexco. (Photograph by
John H. Tashj an. )
TH E NATURAL HI STOHY 0 1" RATTLESNAKES 29
In lat e August 1973, Dani al Lopez and onc of li S (BLA) collcct cd
16 spccmens of C. l. morulus 24-32 km northwest of Calcana; thc
spe cimcns ran ged from maturc adults to newboru youn g. All were
identified as C. l. morulus in having at least 28 prmary dorsal
blotch es with th e exce ption of au adult femal e which had 19
blotches and no mott ling. This female gave bi rth, severa ] da ys af ter
capture, to Iour young wh ich had th e normal complement of pri-
mary bl ot ches, suggest ing that the Iemale par cnt exhihited an ab er-
rant pattern, not uncommon in the C. lepidu s complex. Dur ing
summe r 1974 an additional indivi dual (UTA R-6123) was tak en
from the same localty in a pine-scrub oak situation on a steep
rock y hill side.
Unlike the specme ns collected by Mart n which wer e associated
with humid pin e-oak for ests at the elevations pr eviously mentioned,
all of the specimc ns obtai ned by liS were locat ed at clcvations be-
tween 2380-2592 m amon gst limestone bedrock whi ch had a south-
ern cxposure , A fcw sca tte rcd pinos and oaks were pr esent, but
most of the vcgctation cons istcd of va rious spcci cs of agavcs and
low growing sh rubs , whic h providcd exce lleut cover for the snakcs ,
These snakes appcared to be extrc mc ly common during thc only
Four hours wc spent collccting (due to inc!ement wc uther) an d
would oft en make their presence known by rattling long bcfor c we
wer e within their vi cw, Thcy would immediatcl y seek shelter
among th e rocks an d agaves, and sevcral escapcd us in this habila t.
Captured snakcs defended tliemselves vigorousl y ami often bil them-
selves 01' the tongs that wcr e uscd to secure thern.
This subspecics has bccn rcport ed Irom the Hu mid Pine-Oak
Forest to Pine-Oak Forest (Mart in 1958).
Crotal us mitch clli (Cope)
The spcckled rat tlesnak c, C. niit chcll , is un in luihi taut of the
Ame rican Southwest , Baja Ca lifamia, islands in the Culf of Ca li-
forui a, ami onc Isla nd, Santa off the Pacific coast of Baja
California del Sur (Klaube r 1972). I t is a hi ghl y variable rattlcs uake,
both in sizc and color , and to a lesser dcgr ee in pat tern. Color an d
pattern sec m to be corr elat cd with the general color of the rock y
terrain whic h any givcn populnti on inha bits.
Crotalus mit chelli mtchc lli. The San Lu can suhs pcc ics, C. I/ /.
nt chclli , rangos from the southcrn hordcr of Baja California del
Norte, south th rou ghout the pen nsula. Island popul ations are 10-
cat cd on Ccrra lvo, Es pirilo Santo, San Jos, Carmen, ami Santa
Margarita lslands (Klaube r 1m2). Thc rango do cs not includc
Santa Cruz Island (Soul ami Sloan 1966). This rattl esnake is COI11 -
mon throughout its range, but is most easil y obscrv ed in thc Ca pe
reg ion. Klauh cr (1936) rccordcd specimcns from Ca pe San Lucas,
San Jos del Cabo, Miralores, Todos Santos, La Paz, Muleg ,
30 SPECIAL OF l\'ATURAL HISTORY
San ta Rosalia , and San Ignaci o. Specimens obtained by us ar e
from 8 km N San Jos del Cabo and 16.1 km N Vista
(KU 173097).
Crotalus m. mitche lli is a dwell er of the rocky portien of Baja
Cali fornia. Rarely \ViII it be found an y di stance from rock y re-
tr eats, and it seems to prefer rocky canyons and rock outcrops
(B. Tomberlin, pers. comm.). This rattl esnake has, however , been
found on desert flat s wher e desert shrubs and burrows provide
shelte r. Crotalus e . c ruj o an d C. ruber lucasen sis are often found
syrnpatr cally wi th C. 111 . mi tchelli, although the first two ar e not
as prevalent in rock y situations. Klauber (1972: 536) found that
C. 111. mitchclli was a rock dwcll er, but also indicated it wa s found
in brushy arca s. Most of th ese ratt lesna kes are nocturnal , since
temperat ures in southern Baj a li mit diu rn al ac tivity. Acti vit y
reaches peak pcriods du ri ng the lat e summer rainy seas on. These
rattlesnakes ar e most oft en observed at ni ght as th ey are crossing
roa ds.
A ca pti ve pair of C. m. mitclielli was observed in copulo on
13 October 1975 at 0800 hours. The right hernipcnis of the maj e
was inserted , and a promincnt bulge was evident which extende d
ten scale rows an terior to the vcnt of th e femal e. The snakes
separated at 1500 hours. On 29 Jun e 1976 th e female gave birth to
one neo na te and one inf ertile egg mass. The newbom snake
measured 275 mm in total length, S-V 263 mm, wei ght 17.5 g
(UTA 1\-6939).
The Seasonal Formation Ser ies and Cactus Scrub and Savannah
as defined by Wagner (1964) are typ cal vegetati ve regimes in-
ha bit cd by C. m. tnit cl ielli . These arca s support mesquite, creosote
bushes (Larrea), cacti and other ar d vegeta tion (Goldman 1951).
Crotalu s mitehclli pyrrhus. A sou thwe stern subspecies, C. 111.
pijrrlius, is a rock-dwelling rattlesnake that ranges in Mxico from
the Californ ia border to the southern portions of Baja California
del Norte. Thi s ra tt lesnake is common on th e desert portions of
the Sierra Ju rez an d Sier ra de San Pedro M rtir. It is also found
on the wcstcrn slopes in chaparral stua t ions , but not commonly.
T\Vo ra ttles na kes obtained by li S from south of Pu ertec tos werc
foun d in th e vcnity of abandone d sul phur mines in extremely
rock y mouu tains , They wer e found just afte r sundown in early
spring, a period of th e year that seems to be a peak activty time
for C. m . pijrrhu s, ' Ve have obser ved ma ny individuals of this
subspeci es north of the U.S. border in southern San Di ego County,
California, during early spri ng. Fewer specimens were found dur-
ing summer an d fall. 1'0 take advantage of favorable ambi ent ,tem-
peratures, this rattl esnake shifts from a diurnal mode of activity
in th e spring and fa ll to noc turnal act vit y in the summer (Moore
1976).
TIIE NATURAL I1ISTORY OF MEX1CAN RATTLESNAKES 31
Crotalus m. Purrllll s inha b ts the Ca lifornia Chaparral, Seasonal
Forrnation Seri es, Cactus Scrub, and Montane Thicket as dcfin ed by
Wagner (1964).
Crot alu s mitch clli mucrtcnsis. A dwarf subspecies, e. m. mller-
tensis, is an ins ular fonn of e. mitchcl li from El Mu erto Island of
th c San Luis group in the Gulf of California. El Muerto is th c
second largest isla nd in thc group, is found ca . 6.4 km southcast
of Hu rfanito Island (thc northcrnmost island in thc San Luis
gro up), and s an extrerncly arid, rock y island supporting sparse
vcgcta tion. Dur ing th e summer , tcrnpcratures on the island reach
high extr emes and limit reptilian act vity.
Sixtccn specimens ob ta incd by us (KU 155535, 174830) were
takcn from th c westcrn and southwestcrn portions of th e island on
30 May 1969 bct wccn 1000-2000 hours , most wer e rel eascd aftcr
captu re. The ma jor ity of these spccimc ns were found shortly after
sunset, sincc dayti me ternperature rea ched approximat el y 38C.
Severa l specimens werc found within 3 m of thc wat er , foraging
in beach deb ris. Othcr spccimens wer e Iocated on rocks, and somc
were fou nd und er sparse bushcs. A single specimcn was recordcd
at 183 m elevation, Two lizards, Uta sianshuriana and Str epiosaurus
mearn si , as wc l! as a spcces of Perouuj scu s, occur on th c El Mu erto
Isl and in lar gc numbers, and probably mak e up a sizable po rtien of
the di et of e. m. IIlll ertell sis.
Th cse dwarf ratt lcsnakcs would rattl e lon g before we were
within 6 m of th ern, indi cati ng a hi gh degrcc of al ert ness. Severa]
ndividuals escape d under massive rockpiles bef ore we wer e able
to observe th cm. Thosc captured resist ed vigor ously, and all
thrashed volen t ly. A1though all wcr e obs erved bcf or e sundown,
it is reasonable to assum e that e. 111. mue rtensi s is mostly nocturnal
because of excess vely high daytime t emperatures.
Two of the sna kes ca ptured by us brcd in captivity on 2.3 March
1977. At the time of collection, thc mal e mcasured 355 mm, and
the fcmal e 280 mm, in tot al lcngth. At th c time of breeding, thc
mal c measurcd 680 mm, and thc femalc 610 mm, in total lcngth.
Two livc an d two dcad young (UTA R-7218-1 9), and thrce infertil e
ma sses, we rc born on 13 Scptcmbcr 1977. Total len gth of thc four
young, measurcd an d weighcd eight da ys after birth, was 143-179
mm (mean 167) an d wcight was 3.3-7.0 g (me an 4.9).
Crotalus m. muertensis inhabits thc Seasonal Formation Series
and Cactus Scrub as dcfined by Wagner (1964) .
Crot alus molossus Ba rd and Girard
The bl ackt ail rattl esnakes, e. molossus, of t he Ameri can south-
west and the cen tral pl at eau of Mexico, are well known th roughout
thci r rangc dcspi tc the fact that they werc at times conf used wi th
e. basi iiscus (Gloyd 1940: 161; Klaubcr 1952: 87). Largc collections
32 SPECLAL PUBLICATlO!\-MUSEUM OF NATURAL HISTORY
of th e subspeces C. m. molossus and C. m. ni grescens have been
ma de in past years, and onl y C. m. cs tc bane nsis, from San Es teban
Island in the Gu lf of California, remains bologica lly unknown .
AH three subspeces of C. niolossus attain mod rate size, and speci-
mens of C. //l. niolossus and C. //l. ni grescens are among th e mo st
handsome rattl esnakes we have had th e opport uni ty to observe.
Ths rattl csnake appe ars to be adaptively succcssful ; th c range and
varety of habitats within its area of distribution are unsurpassed
by most othe r Mexican rattl esnake spcces.
Crotalus molossus molossus, Crotalus m. niolossus lives in di-
verse ha b tats from th e Edwards Plateau in west-central Texas
thro ugh Arizona to the Grand Canyon. I n Mexco, C. //l . molossus
has been reported from th c states of Sonora, Chih ua hua and
Coahuila; it intergrades with C. //l . Ili gre scells in the southern por-
tion of its range. See Klauber (1952: 91) for a discussio n of inte r-
grada t on in th is species. Specimens obtaincd by us have been
recorded from: SO.:\'ORA: 4.8 km S Sonoyta ; CHIHUAHUA: the
Sierra del Nido complex, 16 km S Chihuahua. \Ve have observed
approxrnately 30 indivi duals,
AIthough sornewhat unspccial zcd ecologically (Dammann 1961),
C, //l. molossus is gencraIly considcrcd a montane speci es, pre-
ferring rocky retreats wi thin dominant pinc-oak forests. Typ cal
hab itat nichos for thi s rattl csnake from mountain ranges such as
th e Chricahuas and Huachucas in southe rn Arzona, and th e Sierra
de Ajos and Sierra del Nido of northern Xlcxico, reflects th is prefer-
ence. This snake is oftcn observcd at high elevations (up to 2592 m)
in the Chiricahua Mountains of Arizoua. However, it appcars to
be more common below 2135 m on southe rn or southeastern faci ng
slopes of the Chiricahuas wh er e oak , madroo, agave, and va rious
grasses are thc dominant vcgetaton. Pough (1966) observed many
in th e talus rocks lides wher e C. lepidus klauberi is quite common
and our observa tons corroborate those made by him. Rocky str eam
beds are aIso inha bited by C. 111. inolossus, especiaIly dur ing the
summcr rainy seas on. C. //l . molo ssus is not rcstricted to forcs ted
mountains, however, as Klauber (1972: 537) cited spccime ns from
the valleys an d pl ain s of Cochise County in Ar zona which average
ca. 1220-1525 m clevat ion, \Vc have observcd thi s subspecies in
desert situations north of Scottsclale, Organ Pipe National Monu-
ment, ancl near Ga tcs Pass, south of Tucsou, Arizona. Ta ylor (1936:
497-498) found C. 1/1. //l OIO SSIl S to be common in rocky habitats
nor thwest of Cuaymas at La Posa, whi ch is near sea leve !. Perhaps
thc most unexpected place wh ere thi s snake has bcen found is th c
sand dunes south of Cuida d Jur cz in Ch ihuahua, where a spcc-
men was taken by T. \ Valkcr (pe rs. comm.).
Most C. //l . molossus are gentle in di sposi tion an d generally seek
escape when approache d. They rarcl y rattle, al though those found
THE NATU RAL IlISTO RY OF RATTLES:\'AKES 33
in rockslides will at times reveal their presence by rattling, a char-
acteristic they sha re with C. lepidus and C. pri cei . Allen (1933)
found C. m. molossus in trees ca. 2 III aboye the ground.
C. m. molossus nhab ts t he Seasonal Formation Seri es, Montane
Formation Seri es and Scrub Desert vege tational classifications as
defined by Wagner (1964). See Gloyd (1937) and Lowe (1964) for
a descripti on of the habit at of thi s race in Arizona.
Crotalus molossus nigresccns. The Mexican bl ackt ail rattl e-
snake, C. m. ll igrescells (Fig. 18), is generall y a resident of temperate
pine-oak forests of the central pl ateau of Mexico, anel Gloyd (1940:
164) listed localit ies for every stat e within the pl at ean. \ Ve have
observed twenty snak es from the following localit ies: DURAi'\GO:
17 km N Las Nieves (UTA R-5630), 16 km W Dur ango, Los Bancos,
8.3 km E Coyotes (UTA R-.5700), La Ciudad; JALISCO: W Za-
coa1co; 4.8 km S Ca mpan, Contepec, 7 km W
Morel a, Morelia (UTA R-5112-4), Taccuaro; the
lava beds off th e toll road hetween Mexico Cit y and Cuernavaca:
VERACRUZ: the lava beds near Perot e.
Cr otalus 1/l . nigrescens does not occupy as many varcd habitat s
as C. 111. 111olosSIlS, but is not restri ct ed to a specific environment ,
Indi vidu als were observed by us ncar Las Nieves, Durango, in rock
outcrops at 1220 m wi thi n an oak-grass savannah. They were com-
mon there, and the local residents wer e wcll awa re of their pr csence.
The other pr eviously listed locales in Du rango wer e typicall y pi nc-
oak situa tions whcre broken rock and var ious gr asses pro vided
.
,
t
'..

r
-
-
.

"
'.

"

'.
;...
,
e,
' . .-
...
....
ti
.
-
FIG. 18. Crotalus 1/I 0/OSSI/ S nigrcsccns. Specimen from near Morelia,
Michoac n, Mxico. juvenile. ( Photograph by ] ohn H. Tashjian. )
34 SI'ECIAL OF :-':ATURAL HI STORY
excellent cove r and a var ed food supply (Fig. 15). Most of the
rattl esnakes observed by us from thesc areas in Durango wcr e found
basking in the lat e morni ng duri ng the july-August rany season.
Those from Carapan, Michoac n, Pcrote, Veracruz, and nor th of
Cu ernavaca, Morelos, were a1l locat cd in lava bed areas. Davis
and Smi th (1953: 141) tho ught that the occurrencc of C. m. nigres-
cens in Mor elos was doubtful bascd on geographic grounds, At
Carapa u in Michoac n thcy were extremely comrnon (10 obscrvcd) .
Th e spec imens from Taccuaro and Contepec, Michoac n, wer e in
mesquite grassland, a hab itat in wh ch Klauber (1972: 538) had
previously speculated that thi s subspecies mi ght occur. J. R. Di xon
(pers. comm.) found a specimen in swect-gum, oak Iorcst on thc
east-facing slope of Sierra Madre, 11.2 km ENE Pinal de Amoles,
Quert aro, at 1981 m clevation.
Dunkle and Smith (1937) found a fcmalc C. m. ni grescens with
16 young in a canyon wes t of La Colora do, Zacat ccas. Cop ulation
was observed by us between a captive pair on two occasions (28
May 1973 and 2 Ma rch 1974); no pul sations or othc r movements
were noticed. No young wer e born from these obse rved un ons,
bu t on 9 [ une 1975 five young were born in captivity, and wer e
weighed aud measured. Raugc of variation is as foIlows: tot al
length 291-316 mm (mean 304); S-V 267-290 mm (mean 284); wcight
25.4-27.9 g (mean 26.6).
Two C. 111 . lI ig rescell s were observed copulating on 1 February
1978 at 0920 hours. The diarneter of the fema lc's cloaca was 15 mm
and wa s di stended due to thc male' s hemipen s. A noticcabl e bulge
in the Ieuiale's body extended 35 mm anterior to hcr cloaca. The
shoulder spines of the hemipeni s wcr e visibl e and the organ was
da rk purple in colora tion. Coitus lastcd 105 minutes and det u-
mesccncc of thc hemipenis occ upied 100 seconds. 1ntcnnittent
head-bobbing and tongue-Aicking seque nccs by both snakes oc-
curred during coitus.
An open mouth defensiva postu re was observed in three indi-
viduals, Thc rnouths were hcld open for over five minutes when
the snakcs were provoked,
The subspecies C. 111. ni grcsccII s nhab ts the Temperate Pine-
Oak For cst as defined by Leop old (1950) and Mesqut e Crassland
(Klaubcr 1972).
Crot alus poltj sti ctus (Cope)
Previous accounts of Crotalus polijsti ctus indicated a preference
for marshy situa tions and this snakc was genera lly rcferred to as
thc "aquatic rattl esna kc." This pr csumcd pr efer encc for aquati c
sit uations was rcinforced by specimens, captured in 1919 by Paul
D. R. Ruthling, whi ch wcrc sccurcd in thc tul es of Lake Chapala,
Jalisco (Klauber 1956). However, this was probab ly an unusual
THE NATU RAL IIlSTORY OF .MEXICAN RATTLESNAKES 35
situation since Huthling pointed out that thi s area was, at the time
of coll ection, a flooded swampland. I n thi s ephemeral environment,
it is easy to understand why so many specimens (20-25) we re ob-
served by Ruthl ing, and also wh y the spccies was though t to prefer
an aquati c habitat . It is unlik el y for a snake with a warning mech-
ansm such as a rattle to live in aquati c situa tions where the ratt le
would be render ed useless.
In recent years wc have captured or observed over 100 spec -
mens of C. polusti ctus in the vicnity of the Nevado de Colima in
southern Jali sco (Fig. 21) and northern Michoac n. Smith and
Higareda (1965) commented on specimcns from this area. 1n Jalisco,
this snake ha s be en found on pl at eaus which occur in breaks in
the pine-oak for est at eleva tions of 2075-2317 m. Often these arcas
are interlaced with gentl y Ilowin g streams. This rattlesnake can
be found during most months of the ycar: we have records from
Deccmber , February, and May through September. Snakes ob-
tained by lIS ar e from the following local ities: JALI SCO: near
Chapala (UTA H-4000), Rancho San Francisco (UTA R-4499, 4906-S,
4916,5666-7,6043,6250,6704,6822-23,6927; Figs. 19, 20); :\IICnO-
ACN: Ta ccuaro (KU 155540-541), 8 km \V Morelia, 18 km \V
[ quilpan, .5 km 1'\ Cato de las Esperanzas.
Most C. pohjstl clus are found in rocky situa tions wi th an abun-
dance of tan gra ss called zacaton. Nono of these rattlesnakes have
as yet been di scovered in pine-oak fores t, but thcy muy occur in
tliat hab tat . In Mchoac n, near Ta ccuaro, C. polustictus has been
~ ~ ' ; . ' ~ ' ' ( : . ' : ' : ' : : ' : > ~ ? -: ; ~ : . ~ .
~ -J...
"
. ,'
.. . ~
FIG. 19. Grata/li s pO/!lStict IlS. Specirnen from Rancho San Francisco,
Jalisco, Mexico. (Photograph by [ ohn H. Tas hjian.}
36 SPEClAL PUBLICATION- 1IUSEUM OF NATURAL HISTORY
.-.
"
. ~
I
FIG. 20. Cr otalus pohj sti ctus. juveni le speci me n bom lo female collecte d at
Rancho San Francisco, Ja lisco, Mexco. ( Photograph by [ohn H. Tashj an.)
FIG. 21. Habitat of Grata/lis nalusti ctus. Rancho San Francisco, Jalisco,
Mexco . Sna kes were commonly Iound among boulders or in gophe r burrows
in gra ssy mcadows wt h surrounding pi ne forest on hill sides. (Photograph by
11. Cranger.)
THE NATURAL HI STORY OF ~ I E X I C A I \ RATTLESNAKES 37
collected in mesquite grassland (DuelIman 1965: 655). This area
has rocky outc rops where the species is easily located. We have
also colIected th s speci es in plains-grassland areas, although it does
not appear to be abun dant in this environment ,
Cr otal us pohjstict us is found rather comrnonl y in lat e spring
when summer rains have not yet been sufficiently heavy to con-
tr ibute to growth of grass cover . During th s period, they become
sornewha t noc turnal, active individuals havng been recorded as
late as 2200 hour s. I n late spring, C. polystictlls d spl ays a rather
mild di sposit ion and is usuully quite inoflensive. Sorne spe cimcns
have been observcd attempting to hiele their head s ben eath a coil
of their bod y, behavior similar to t hat exhibited by Lichanura
tri oirgata roseoj usca. Dur ing the summe r, th s specics becomes
muc h mo re aggressve and will gener aIly no t retreat wli en ap-
proached. \ Ve observed an individual which, upon discovery,
ope ne d it s mo uth in a th reat ening pose similar to the hchavior ex-
hibited by Agkis t rodo n piscicoru s. Lampropeltis triau gulum ar-
cijera, a harmless co lub ricl snake, is corn mon in the vicinity of the
Nevado de Co lima, an d ma y be an important pred at or upon these
rattl esnakes.
The kno wn range of C. pol ysti ct lls is decreasing du e to habit at
destructi on. The pl at eados iuhabitcd by these rattl esnakes ar e be-
ing alte red for agricultu ra ! use bccausc they are mo re easily clea red
than adjaeent pi nc-oak for est s.
The preferred habi ta t of C. politstictus is Tempernte Pine-Oak
For est as defined by Leopold (1950), and Mesquite Crassla nd
(Due llman 1965).
Cuesta Terron (1930-31) reported on a brood of 14 newborn
young wh ose average tot al len gth was 12 cm, a figure con sidcrahly
sma lIer than the mean total Icngth of broods measured by uso It is
possibl e that Cues ta Terron ma )' have misi dent ificd his snakes (see
Fi g. 6, p. 54), although we have not seen his specimcns.
Dat a are avai lable to us for six broods born to femal es of th is
spccies, as folIows: Femal e A, ",ei ghing 111 g anel measuring 608 m
tot al length (575 mm S-V), gave b rth to ten young on 20 [une
1974: total length 200-223 mm, mean 211; S-V 184-208 mm, mean
195; weight 6.3-10.9 g, mean 7.2. Fcmal e 13, wei ghing 166 g aneI
measuri ng 660 mm tot al length (620 111m S-V), gave birth to twelv e
young on 13 June 1974: total length 211-287 mm, mean 222; S-V
198-271 mm, mean 208; weight 8.5-11.5 g, mean 9.9. Femal e C
gave birth to sev en young on 26 Ju nc 1975: total length 198-232
mm, mean 216; S-V 185-211 mm, mean 200; wei ght 9.9-10.5 g,
mean 10.1. Fcmale D gave birth to seven neonates on 30 june
1975: tot al length 182-205 mm, mean 197; S-V 167-185 mm, mean
179; weig ht 9.9- 11.1 g, mean 10.6. Fernal e E gave birth to Iive
st illborn young on 30 june 1975: total length 155-203 mm, mean
38 SPECIAL OF "ATURAL HISTORY
177; \Veight 2.4-S.,') g, mean 4.0. F emal e F gave birth to seven
young on 25 June 1975 : total len gth 227-239 mm , mean 232; S-V
201-219 mm, mean 212; weight 9.5-10.9 g, mean 10.2. Th e range
of variation of th e six brood s is as Follows: total length 155-287
mm, mean 2.14; S-V 167-271 mm, mean 201; weight 2.4-11.5 g,
mean 8.9. '1'\\'0 of th c snakes born to Fcmale A were maintained in
captivity an d exhibi tcd eourt ing bc havior on 31 j anuary 1978.
Crotalus pri cei Va n Dcnburgh
Crotalus pricei prieci. Crotalus p . pricei (Fig. 22) is the more
commonly known subs pecies of th is sna kc, Iargc collections having
becn madc in Santa Rit a, Huachuca, Chirica hua , and Graham
Mountains uf sou theastcrn Arizona . Fewcr inclividual s of th is race
have becn rccordcd from Xlexico, yet th ey are ra ther abundant in
i\lexico in pine-oak forcsts at elcvations ra ng ing fro m 2135 to at
least 2745 m. \ Ve have observcd these rat tles nakes fr om thc fol-
lowi ng locali ties in SO?\OHA: Sie rra de Ajos (UTA
1\-6931-34); CHI HUAHUA: th e Sierra del Nido complex: DU-
RA:\GO: Las Adjuntas , near Coyotes, 14 km El\'E El Salto (UTA
1\-2(21), Llano Crande (I\:U 158561), Los Bancos (UTA 1\-6251).
Klauber (1972: 45) suggcstc d that C. J. pricei mal' al so OCCIII" in
eastcrn Sinaloa and no rt hern l\'ayarit, but to 0 11I" knowl cdge, no
spccimens have becn scc urcd in ci t he r of these states.
Crotalus p . pricci is a res idc nt of hi gh rocky pine-covercd slopes
(Fig, 23). I n t hc Unitcd St at es, it secms to pref er talus rock slides
1'1 (;. 22. Cro talus pricci pricei. Spec imen frum near Los Bancos, Durango,
Mexco. (Photograph by [o hn H. Tashjian .)
THE l\'ATURAL HISTORY OF },IEXICA0: RA.TTLESNAKES 39
in th e previously ment oned mountain ranges. This rattl esnake will
ofte n clivul ge its presen ce by ratt ling when approache d. Howevcr,
like many of the montan e ratt les nakes, C. p . pr icei is eurious anel
usuall y ret reat s onl y a short clistnnce. Although genemlly asso-
cia ted with rocks, this ra tt lesnake mal' be founcl also in grassy
mountai n va llcys whe re prcy, such as li zar cls anel small roclents, is
abundant. One aclult sn ake (UTA H-20:21) containecl a juven il e
lizarcl ( Sccloporus poinse tti} . \Ve ha ve frequentl y observecl t his
subs pecies on warm, humid days when clifl usecl sunlight is preva-
lent. 1ndividuals were co llecte cl ne ar rock y retr eats from March to
Novernber , even when a light snow cover was presento No cloub t
th ese retreat s a re th e warmest micro- habi ta ts avai labl e to thi s rat tle-
snake in mountains . On 17 Xlarch 196.5, two C. p. pri ce were founcl
on a sout h-Facing talus slope at Burfoot, some times kn own as Buena
Vis ta Peak, Chi rica hua Mo untains, Arizona at 1:2:20 hours . Snow
surrounclecl th e talus slope, an d t he ambient temperat u re was 11oC.
In Mexico, C. p . pricei is probabl y th e most common rattl esnak e
at hi gher elevations wi thin its range, where it is founcl sympat rically
with C. icillardi, C. lepidus, und C. 1Il 0 10 SS11S. We ha ve collec tecl
C. p. pri cci fro m Ma y through September , sometimes u ncler the
most adverso weather condi ti ons, 1n ] une 1973 we secu red fou r
individuals on a south- faci ng slope nea r Los Ba ncos, Durango (Fig.
FIG. 23. Habitat of Crotalus pricci pricci and G. icillard mcridionalis.
Sparsc pine-oak forest near Lla no Grande, Durango, Mexico. Crotalus p .
pri cci was ge ne rally discovered in rocky areas wh le G. ui, incridtonalis was
col1ectcd in thi ck grass or underbrush, mainly man zanita and scrub oak, 0 11
hill sides. ( Photograph by Lyndon A. Mit chell . }
40 SPECIAL OF l\'ATURAL HI STORY
15), two hou rs afte r a drivin g hail stonn. The hail melted qu ckly
and all Iour snakes were act ive, appare ntly foraging for food.
These Iour snakes exhbitcd an unu sual array of body patt erns .
One indivi dual lia d the typical colora tion of a slate -gray b ack-
ground with brown do rsal spots. The second cxample exhibite d a
blue-gray backgroun d wi th rus ty colored spots. The third spe cimc n
was overall salmon-pin k wi th tan spo tting, and th e Iourth snake
was a completely patternl css light tan. These four spe cime ns were
capturcd within a 45 m ra dius,
A Iemale specimc n of C. p. pricei obtained at Lo s Bancos,
Durango, gave birth to three young 011 24-28 j ul y 1973. Anothcr
Iemale, taken near On ion Saddle, Chiricahua Mouutains, Ari zona,
gave birth to Iour youug on 20 Xlay 1967. Thrce young C. p. pricei
were found on 11 jul y 1973 near Los Bancos, Durango, in a rock
crc vice: th ey had not she d. A ca ptive mating produccd Iour young
on 9 jul y 1971 (J. A. Campbcll. pers . comm.). Three of the four
young were born ali ve al\(l wci ghed 2.7, 3.9, 2.4 g; mean 3.0. Four
female specime ns from Llano Grande, Durango, gave birth in th e
lab or atory on 10, 14, 27, and 29 [ uly 1977, respcct vcl y. Data on
thesc [our broods is as Iollows: Fcmale A produced eight young
(2 sti llborn, less th an 50 mm total length), total Iengt h 157-167 mm,
mean 164; weight 4.0-4.2 g, mean 4.1; Fernale B gave birth to nin e
viable young (four di ed with in two days, UTA R-6935-8), tot al
len gth 152-173 mm, mean 162; weight 3.5-4.0 g, mean 3.6; Fcmale
C gavc bi rth to six via ble young , total length 159-169 mm, mean
162; weight 5. 1-5.6; Femal e D produced six young, tota l Iength
160-1805 mm, mean 168; we igh t 3.6-3.9 g, mea n 3.7. The range of
variat ion an d mean for all litters was as Iollows: total length 152-
185 mm, mean 164; weight 3.5-5. 6 g, mean 4.1. Al! but one of th e
Fernales were collec ted while baskin g in th e early moming on
rocks (L. A. l\litchell and D. G. Barker, pers , corn m.). Kauffeld
(1943a. 1943b) rep orted th at an example of C. p. pricei collected at
Barfoot, Buen a Vis ta Peak, Chiricahua Mountains, Cochise Cou nty,
Arizona, gave b rth to six young on 19 August . He citcd another
record of a brood of th is race born on 3 August 1941. Kcasey (1969)
recorded a bir th of eight young on 23 September 1953. Six young
wer e born on 19 }ul 1' 1971 to a fcmale sec ured at Los Leones,
Chihuahua (Van Devender and Lowe 1977).
C. p. pricei inhabits the Tcmpcrute Pne-Oak association as de-
fined by Leopold (1950). Sce Cloyd (1937) and Lowe (1964 ) for a
description of th c habitar in Arizona.
Crotalus pricei mquihnnnus . Although mu ch litcrature is ava l-
nble ab out C. p. pri cei , Iitt le is known about C. p. miquihuan us
(Figs. 24, 205), no doubt du e to the inabilty of col!ec tors to ge t
into the Iimited areas inh abi ted by it. Specimens of t his rattlesnake
ha\'e been kno wn since 1898 when Ne lson and Goldman succeeded
TlIE NATURAL HISTORY OI' :-'IEXICAi\' RATTLESNAKES 41
in capturing an adult mal e nea r Mi quihuan a, Taumaul ipas. Ho\V-
eve r, it was IIOt placed as a subs pcci es of C. pri cei unt il t he holo-
typ e, a subadult mal e, was sec ured near Calea na , Nu evo Len, in
1938, and described by C loyd (1940: 10:2). Since that t ime, occa-
sona l specimens have been fouud bnt were unfortunatel y lost be-
for e being pl aced in syst ematic collcc tions.
Recent information, still qu ite li mited, presen ts a sornewhat
clearer p ctu rc of th e apparcnt range of C. p. miquihuunus. !JI
1961 an adult snake of th is race was coIlecte d in the Sierra de Los
Amargos, Coahu la, exte ndi ng th e known range some thirty miles
north fr om the type local ity in ?\lIevo Len (Axtel] and Sahath
1963). This \ViII probabl y becomc th e range limit of thi s ra tt le-
snake to th e north, snce th ere is a di st inc: t topographical aud Iaunal
break at thi s sit e wher cin a much mo re arid and desert -t ype hab tat
replaces the pine-oak forest of the Sierra \ fadre Ori ent al . J. H..
Dixon (pers. comm.) secured three specimcns of th is race [rom
14.8-17.2 km E San Anton io de las Alazanas betweeu 1981-280-1 m
elcvatiou during May and August 197:2. AIl wcre takcn I rom a
piou-pin c, aga ve slopc. South from Cerro Potos, the spcci mc n
obtaincd by Nclson aud Go ldman in 1898 is th e onl y knowu record
for thi s subspccies. This is not to sllggest , howcvcr, that the ar ca
surroundiug Tamaul ipas, is th e southcnunost li mit of
the rango of C. p. nquihuanus, Dile to topographi cal , Iaunal , aud
to some degree, geological simi larity, it is possible th at th is sub-
-.
..
i. :'.'
" . " ., .'
-: .,.,
.'-'-
FIG. 2-1 . Crotalus pricci tniq uiltuanus, Spe cime n fro m near San Antonio
de las Alaza nas, Coahuila, Mexi co. ( Photogr aph by [ ohn H. T ash jian. )
42 SPECIAL OF NATURAL HISTORY
Frc , 25. Crotalus pricei miquihuanus, [ uvenle specimen born lo ferna le
collected from Cerro Potos, Xuevo Len, Xlexico. ( Photograph by J01ln II .
Tashj an . )
species will eventua lly be found as far south as the San Lu is
Pot os/ sou thweste rn Tamaulipas border , among th e high for ested
mount ain slopes .
Duri ng our travels thronghont Xlexico from 1967 to 1974, we
sccured four adult C. p. miquihuanus. Two were di scovercd above
3050 m near the t yp e local ty. in mid-july 1974, in a limestone
bedrock, agave, scrub oak h bitat at 1100 hours. They wer e bo th
sunn ing on the limest on e roek and sceme d pl acid ancl ind ifferent
to being captured, unl ike C. p. pricei which reacts defeusively.
The summit of Cerro Potos has been heavily dcforest ed, and a
luxuriant growth of Iow-growing vcgetation, such as agave ancI
scrub oak, ha s rcplaced thc ori ginal pinc forest. Ther e ar e still
man y pines in the ar ea, hut Ce rro Potos is no longer str ctly a
pin e for est. With thi s chango in vegcta tion, the populatiou of
C. p. miquihuonus appears to have increased, possibl y the lo\v-
growing shrubbery offer s more protection to bo th the ratt lcsnakc
and its ma in food prey, lizards of the genus Sceloporus.
A female C. p. miquihuanus, collected on the cast slope of
Cerro Potos, Nuevo Len , at 3203 m, gave birth to five young
(KU 1.5,5,542. 1.57869-871) on 19 Au gust 1974. The range of varia-
tion is as Iollows: total length 130-143 mm, mean 135; S-V 117-130
mm, mean 121; we ight 2.6 g, mean 2.6. Another femal e (UTA
R-6235) captured on 16 Jul y 1976 in a pin e-oak Iorest at 2652 ID,
TH E NATURAL IIISTORY OF ~ l E X I C A N RATTLES:\AKES 43
located 17.2 km E San An tonio de las Alazanas, gave birth to four
young th e Ioll owng day (UTA R-6175-77, 6236).
The habit at type pref erred by G. ]J. mi qui huanus is t he Temper-
ate Pin c-Oak assocato n (Klauber 1956).
Cr otal us pusillus Klauber
Crotal us pusillus is a small rattlesnake Iound in intcrmi tt ent
montane habit at s in southern Jal isco and westcentral Mchoac n
(Figs. 26, 27). Klauber (1972: 163) consdercd thi s tax a to b e on e
of th e most primiti ve Xlcxi can mountain rattl esnakes. Crota lus
pu sillus inhabits the pine-oak forest s at elcvations fr om 1.52.5 m to
at least 2380 m (Sierr a de Coalcom n), and r esides within the
Upper Souoran-Transi tion Zone as defiued by Coldman (1951). I t
occurs sympat rically with G. t. triseriatus throughout its range with
th e exce ption of th e Sierra de Co al com n where possbl y only G.
durissus culminalus and G. !J . basi liscus live in close proxim ty .
Cr otalus pusillus is locall y common, amI 17 spccimens wcrc ob -
served by us (UTA H-4530-1, 5846, (119). We found thcm to be
quite abundant in th e Si er ra de Coalcorn n. in clea red arcas of
humid pine-oak forest (Fig. 28). Some specime ns wcre takcn in th e
early mo rni ng , but this anima] was mo re easily loca ted aroun d
1200 hour s, after th c sun had burned off th c low mo rning clouds .
Duelhna n (1961: 6.58) suggcstcd th at this species was part ially noc-
t urnal. Sevcral snakes were scc ure d in cultiva tcd Iiclds whcre co rn
was plantcd betwcen rock outcrop pi ngs. Others wer e locatcd on
and under fall en logs, an d sorne were found in a rocky, grassy.
agave situat ion. J. R. Di xon (pcrs. comm.) sec urc d spe cim ens from
14.4 and 20.9 km \\' At enquiquc, Jali sco, un der fall en ba rk in an
oa k for est betwccn 1829-2161 m clcv a tion.
Crotalus pu sillus exhibits a typi cal bchavioral pat tern of rut tle-
snakcs from hi gh, humid , monntain arcas. Jt wi ll olten revca l itself
by rattling furiously when approached . but is compara tivc ly Icss
nervous th an G. lepulus in similar enco u nte rs . :\ Ian y 01' ou r speci-
mcns def ended thern sel vcs vigo rously upon discovcry, an d ofte n
would hite thcmsclves 0 1' th e tongs whi ch were uscd to ca pt ure
t hern . Rit ua lized combat betwcen captive males has beeu obse rve d
by us, The stomach of an adult (UTA R-4530 ) containe d an
orthopteran.
A female col lected near Dos Aguas, Xlichoac n, gave birth to
on e l ive and four st ill born young (1\: U 155.545-548). and on e infertile
egg on 23 january 1974. Ran ge of variation is as follows : tot al
len gth 165-179 mm, mean 171; S-V ] .50-162 mm , mean 1;"5.3: weight
3. 0-6.1 g, mean 4.0. The surviving juvenile was maint ained in cap-
44 SPECIAL PUBLICATION-!\1USEUM OF NATURAL HISTORY
FlC. 26. Croial us pusi llus. Specimen from near Dos Aguas, Sierra de
Coa lcomn, Mchoac n, Mexco. ( Photograph by John H. Tashjan.)
, ,
."
Frc. 27. Crotalus pusillus. j uveni le specimen Irom near Dos Aguas, Sierra
de Coal comn, Michoacn, Mexico. ( Photograph by [ohn H. Tashj an.)
THE NATURAL HISTORY 0 1' ~ I E X I C A N RATTLESNAKES 45
FIG. 28. Habit ar of Crotalus pusi llus near Dos Aguas, Sierra de Coal comn ,
M ch oac n, Mexico, j ul y 194. Extensi vo limestone outcropping border cd by
pine-oak forcst. Snakes appcared more abunda nt in arcas prevou sly clea rcd
for cultivation, (Phot ogr aph by [ on athan A. Campbell. )
tivity and bred, and gav e birth to thrce you ng on 15 July 1976.
Two of the young were st llborn wi thin a singl e fetal mcmbrane,
and the mean weight aud mcasnr emeuts for bot h are as Iollows:
total length 111 m, S-V 98 mm, weight 1.1 g. The third neonate
was ali ve at birth, measured 179 mm tot al lcngt h (156 mm S-V) and
wcighcd 6.0 g (UTA H-7164).
On 24 Mnrch 1975 a pair of cnptive C. pusillus was placcd to-
gether and a complete mating seql1 ence was observed.
1130h Male placed in cage wi t h fem ale. Ferna le had prevou sly she d and
dcf ecat ed. Femal e immcdiately begun twi t ching.
1130-1159 Mal c began spas mod c forwar cl jerking ( 2 per sec ) and tengue-
f1 icking (2 per sec) . Mal e loosel y draped his coils over dorsum of
Iema le, bent head lateral ly at 90 an gle to longitu dinal axis of hls
hod y and, by cont racting vi goro usly, pu lled his head approxirnately
2.5 cm post eri orly along hody of fcmale. This ac tion was perfonned
repeat edly. Fe ma le rai scd ta l 11 cm vertically an d oponed cloaca.
Mal e became excited and invest igated Iemal e' s cloaca l reg on. This
entire seque nc e was rc pea tcd 34 times. Fc ma le t witched on ly
occa sionall y.
1200 Male wra pped t ai! a round Icmalc' s head,
1201 Maj e released Iemal c's hcad and hrou ght hi s cl oaca to ve nt of fernalc.
Mat ing was not accomplishe d,
1202- 1220 Both snakes rcmancd qu cscent.
1221 Female moved head, male lost gri p and becarne excted.
1222-1400 Same pa tt em as above,
46 SPECIAL PUBLICATION-\I USEU\I OF NATU RAL IlISTORY
1401 Male and fernal e simulta neous ly raised ta ls vert cally, ti ghtly
wrapped tals. he ld positi on for 1 mi nute, then rel eas ed .
1410 Female raised tai l verti cally ( 11 cm ) for 5 minutes. Mal e con-
tinued to investigatc.
1412 Both ma le and female raised ta i! 11 cm with cloacae juxtap osed
giving appeara nce of a lyre. Distan ce between rattle matrix of
ho th snakes wa s 6 cm. Male released positi on, then slid tail ap-
proxmate ly 2 cm to cloaca of femal e. Mal e' s tai! wa s enci rcled
around ba se of female' s tail. Male slid tai! rapidl y an d repea tedly
along longitud ina l axis of Fcmal e' s ta i!.
1417 Mal e's tai! enci rcle d has e of femal e' s tai! twice.
1419 Mal e lost gri p with his tail.
1124 Fcmale raiscd tai l.
1 3 1 Female continued to ra ise ta i!, male insert ed right hemipeni s. A
noticeable hulge was appa rent cight vent ral sca le rows ante rior to
Iemal e' s cloaca. :-'Ial e con tin ued to pu sh aga nst rcsistance of female
wh ch cause d the fcmale' s ta l to bend backwards over her bod y un-
ti! the ruat rix of her tai! wa s 3.5 cm above her vert ebral column,
1443 Fou r quick pul sati ons at cloacal arca of mal e,
l446 :-'Iale began undul ati ons (2 per sec ) which were withi n a verti cal
pla no and were 5 cm ante rior to the vent . During th s peri od , the
ma le consta ntIy pu shed the Female's tail hackwa rd.
1500 Male tapped men tal arr-a of hi s lowe r jaw in a vert ica l plane on
clorsum of Fema lc .'39 times in a G mi nut e periodo 1\0 tong ue Ilicks
accompanied thi s heh avior unl ess the mal e shf ted POSitiOIl .
1509 Male ta pped his lowe r jaw on dorsum of femal e 17 times, no
to ngue flicks,
153j Pair stiII joi ned, diameter of Female's cl oaca was 12.5 mm wh ch
was dis tended du e to ma lc' s hcmipcn s.
1710 Approx ma tely 6 mm of base of hemipenis visi ble.
1800 Mal e pu shed tai l against Iemal e' s tail , wr ithed for 3 seconds.
183j Same as above.
1841 Bot h tail s moved slowlv for I Dseco
1909 Observat ions
The ncxt day at 0800 hours the snakes had separated.
On 22 April 1975 thc same ma le specime n was placed with a
dil erent femal e. The samc beh avioral pattcrns were observ ed , in-
cluding upraised tails, bending of the Iemal c's tail ba ckwards ove r
hcr body, and pulsat ons. The head tapping behavior of the mal e
wa s also obse rvcd. The rai sed tail of the femal e stimulated the
ma Ie noticeablv,
The habitat prcferrcd by C. pusillus is mxed pine-oak forest
and volca uic rack (Klauber 19.">6) and Pin e-Oak Forest at an alti-
tude of 1.550 to 2300 m (Duellman 1965),
Crotalus rub er Cope
The red diamond ra ttlesnukcs, C. ruher, of extreme southweste m
Ca lifornia and adjacent Baja Ca lifornia, ar e among the largcst and
gc ncrally the most placid ratrl csnakes we have observed. Partl y
becausc of their range amI habitat pr cfer cllce, these snakes have
been collec tcd by one of li S (13 LA) e\'e r)' month of the year. Their
peak pcriod of acti \'i ty is in lat e spring, and the)' are relati\' cl )'
THE NATUHAL HI STOHY OF r-IEXICAN HATTLESl':AKES 47
common throughout their range. I n sou thern California, the stat us
of C. r. ruber popu lati ons is becoming precarious as mor e and more
devel opments claim inland val ley habitats.
Crotalus ruber ruber. The northern subspecies, C. r. rub er,
ra nges from southeastern Los Angel es and Orange, San Di ego ,
wes tern San Bernardino, and River side counties, and extreme south-
west ern Imperi al County in southcrn Ca lifornia, south to Loreto,
Baja Ca lifornia del Sur wher e it intergradcs wi th C. r. lucasensis
(Klauber 1949a: 46). Klauber (1972: 46) also reported indi viduals
fro m Monserrat e, San Marcos, Pond, ngel de la Guarda, and
South San Lo renzo islands in th e Gulf of California. It seems
reasonabl e to assume that this rattl esnake will eve ntua lly be found
on other, smaller islancls that have not yet been th or ou ghl y ex-
pl or ed, It is inte resli ng to note that a series of C. rub er, now known
as C. r. lorenzoensis, from South San Lorenzo I sland in the Univer-
sity of Colorado collection, show tendencies toward rattl e loss
(Radclifle and Mus lin 1975). This conclition was see n on approxi-
rna tely 50 percent of the specimens. C. r. ruber obta ned by us
havo been obs erved from various locations in five counties in south-
ern Ca lifornia, and the Iol lowi ng local it ies in Baj a Calif orni a: 6.4
km S Tij uana, the vic ini ty of Ensenada, th e Hamilt on Ranch (20
specime ns) ncar Colonet, San Quintn, El Socorro (5 speci me ns),
El Rosario, 16 km N Laguna Chapala, San Igna cio, 17.6 km \V
San ta H. osali a, 22.5 km S Mu leg . Crotalus r. ruber appears to be
absen t from the desert east of the Sierra de ]uarez in Baja California
del Norte.
In southern California, C. r. ruh er is a reside nt 01' the hot inlan cl
valleys which include th e Borrego Desert of San Diego County.
It is Iound gene rally in arcas of granite rock ou tcroppings , espe-
c ally cluring win ter months. During these mon ths we have had
success locati ng numbers of these snak es in the hott est part of th e
day. They ar e oft en coiled at the entrance of a gophe r burrow, or
in th e midst of th e rubbl e of a dil ap idated barn. Harcl y are th ey
located any distance from sorne typ c of shelter. Thi s rat tlesnake is
extremely lethargi c during wi nter months, aneI def ensivo behavior
is selclom obs crved eve n wh en it is provokcd . However, during
the lat e spri ng and summer months, C. r. rub er becomes incr easingly
noc turnal, aneI invades th e grasslancls betwcen rock outcrop s where
it probably fecc1s to a large cxtc nt on eotto ntail rabbits, Sy/r; i/aglls
aiuluboni sanctidiegi , and gr ound squi rre ls, Ciiellus !J . be eche ui,
both of which are abundant. In the truc desert regions 01' its range,
this snake is founcl near rocks, and seldom wanders from protective
retreats. In the carl y spring, pairs of red di amond ratt lesnakes ar e
oft en observed basking tog ether in full sun light . Mating was ob-
served in earl y April between a Iarg e pair in Railroad Canyon, just
east of Lake EIsinore, California. The snakes we re not collected.
48 SPECIAL OF i'JATURAL HISTORY
\Vagner (1964) characterized th e vegetatonal eharaete risties of
the arca inhabite d by C. T. ruber as California Chaparral , Seasonal
Fo rma tion Series, and Cactus Scrub,
Crot alus ruber lueasensi s. The southern subspeeies of th e red
diamond ra ttl esnake, C. T. lucasensis, has a geographic range from
the fishi ng vill age of Loreto to th e Cape regi n of Baj a Cali fornia
del Su r. Klauber (1972: 46) reported specirnens of thi s ra ce from
Santa .\ Iargari ta an d San Jos islands. As with C. T. T1I1Jer , C. r .
lucasensis may someday be found on other islands when th ey are
t horoughly explore d, Specime ns ob tain ed by us fr om th e mainland
we re collected at El Triu nf o, 8 km N San Jos del Cabo, and Buena
Vista (B. Tomherlin, pers. comm. ).
Barney Tomberl n (pe rs. comm .) reported th at C. r. lucasen sis
was t he most common snake in th c Cape region of Baja Cal ifornia .
.\ Iost of hi s sna kes wer e taken in Septernbe r when that area re-
ce ved a major portie n of its annual ra infal l. At that time th ey were
nocturnal gcne rally, and specme us wer e eas ily found by night
drivi ng. Although thi s rattl esnake can be found in th e deser t an d
ope n ari d plai ns of southe rn Ba ja California del Sur, it is most corn-
mo n in heavy b rush where roc ks and rocky outcrops are pr evalent .
Th s subspecies inh abits th e Seasona l Formation Seri es an d
Cactus Scrub as defined by Wagn er (1964).
Cr otalus scui ulatus (Kenn cott)
Crotalus scutulat us was on ce confused wi th many of t he othe r
prarie dwell ing rattl esnakes (Klaube r 1972: 541), an d th er efore,
mueh th at has been wrtten about it may, in part, be erroneous .
lt is a rat her large, heavy-boded snake th at has a rango simila r to
that of C. inolossus. Cr otalus scutulat us does no t, however , inhabit
the mult iple habitats within ts rango as does C. molossus. C.
scutulat us is found fr om the Mojave Desert and adjacent Ne vada,
south th rough Arizona, southweste rn Ne\V Xl exico, and Texas, and
thence south int o th e Mexican central pla tean.
Crotalus scutula t us scutulat us, Crotalus s. scutulatus is a corn-
mo n ra tt lesnake wi t hin it s range in the Uni ted States , Klauber
(1972: 47, 541) gave an excellent summary of the localities an d
field data for th s race. In Mexco, we have fou nd C. s. scut ulatus
throughout the Hat , arid cen tra l plnt eau, but nowhere did it seem
part icu larly common. Snakes captured an d obs erved by us (approx.
30) wer e recorde d from: CHIHUAHUA: the va lley between the
Sierra del Nido an d Sierra de Santa Clara at 1982 m (UTA R-4554);
SOI\ORA: 12.8 and 17.6 km S Agua Prieta ; DURANGO: La Zarca,
12.8 km N Rodeo, Sa n Juan del Ro ; NUEVO LEN: Sa nta Fe
(UTA R-4.59.5), 12.8 km S Ga leana, 8 km E San Roberto junction:
COAHUI LA: 3.2 km \V San Anto nio; SAN LUIS POTOSI :
THE NATURAL HISTORY OF MEXICAN RATTLESNAKES 49
Matehuala, 4.8 km E Matehuala, ZACATECAS: 13.3 km E [une-
tion of Mexican Hwy. 45 and 49 (UTA R-2715).
Miller and St ebbins (1964) not ed th at throughout th e northern
portion of its range, e. s. scutulat us appeared to bc ext remely noc -
tumal in behavior. Most of the specimens obtained by us wer e a
result of ni ght driving. In spring, it was not un common to find a
speci men coil ed at th e ba se of a palo ve rde or creosote bush in
early momi ng. During surnrner months, however , these snakes
wer e rarel y encoun tered during dayl ight hour s. Crotalue s. scuiu-
latus seems to prefer the opcn mesqui te-creosote-cacti habitat wi thin
its range. It usually avoids tru e san d desert where Ciotalus cerastes
is th e dominant rattlesnake.
In southem Arizona and adjacent Mexico, e. s. scutulaius is
very common in prairie vallcys (1220-1.52.5 m) between th e numerous
for ested mountain ranges of that region . Southward within the
Mexcan platean thi s snake becomes cre puscular due to th e hi gher
elevations of th e open , arid hab itat. One individual obtaine d by
us from grass land, ju niper plain (1982 m eleva tion) near Santa Clara,
Chihuahua, wa s located crossing a dirt road at 1100 hours in early
]uly. In central Duran go near the town of La Zarca, e. s. scut. ulat us
was oft en seen in mid-afternoon crossing the hi ghway or basking
at the edge of th e pavernent. This ar ca is approxirnatel y 2440 m
elevation, and th e da ytimc tcrnper at urc is somewhat cool. [uniper
and grassland are the dominan t vegetation in th is wind-swept re-
gion. Individuals from th e eastern part of San Luis Potos, cspe -
cially around Xlat chuala, 1372 m, wer e al so crepusc ula r in activi ty
despite a lower elevat ion th an at th e Durango localities. Reese
(1971) found e. s. scut ulatus at a locality 77.2 km S San Roberto
Junction in Nuevo Len, and Banta (1962) found it at Caada
Honda, Aguascali entes. One adult sna ke obtain ed by us (UTA
R-4554) contai ned rodent hair.
The Temperutc Mcsquite-Crusslnnd and Desert as defined by
Leopold (19.50) ar e typ cal cnvironments for e. s. scutulat us . See
Glo yd (1937) and Lowe (1964) for a description of the habitat in
Arizona.
Crotalus scutulatus sal vini, Crotalus s. salc ini (Fig. 29) is also
a resident of op en, hi gh interior plains. It is found in Tlaxcala,
Puebla, and west-ccnt ral Veracruz, and its range is doubtless de-
creasing as these pl ains are cl eared for cultivat ion, The t ype 10-
cality for this rac e, Huamantla, Tlaxcal a (Klauber 1972: 147), at an
elevation of ove r 2440 m, may now possibl y be devoid of enough
suitabl e habitat to sus ta in a population of th ese ratt lesnak es. AH
of our spe cimens wer c coll ect ed with in the la va beds near Perote,
Veracruz, and Tehuac n, Pu ebl a (Fig. 30). Both localities support
luxuriant desert plant gro wth such as yucca, palmettos, and th e
giant, slende r Neobus baumia, which are the mos t common pl ants.
50 SPEC IAL 01" NATURAL HISTORY
\ Vith in th e relati vely undisturbed arcas of it s limited ran ge, C. s.
saloini rema ns a ra ther cornmon sna kc . and, unlike C. s. scutu lat us,
is Found in as soc iation wi th rocks, Thc lava beds of Perote seem
to provide thi s rat tlesnake excellcnt hab it at. 1t ma y, howevcr, be
common there on ly be causc of man's cn croachment upon th e roll -
ing. ope n, rockless plains wherc C. s. saloini was once, no doubt,
th e dominant member of the gen us . Specm ens of C. s. salc ini ob-
tained b y us were '111 locat ed duriug mid-afternoon in July and
Augus t, and were b asking near porous volcanic rocks,
This ra ttlesnake is one of the most aggress ive spccies "ve have
encouute rc d, Sev era] st ru ck so violcntly th at th eir en tire body
appcarcd to be momentaril y air borne. It is common behavior for
them to strike even after bcin g constrained in clo th bags for severa]
weeks. C. s. salcini inhabits the Tcmpcrate Pin e-Oak and Mcsquite-
Grassland vcget ntionul ureas as defiucd by Leop old (1950).
Crotalus steincgcri Dunn
Croialus stcinegeri is a p rimitivo rat tlcsnake Iound in th e moun-
tains of southeaste rn Sinaloa ami sou thwes tern Dura ngo. Klaubcr
(1972: 5-12) reported two spccime ns t ha t ".. . werc found on the
bo rdcr of a pine for est, at th e upper edge of a canyon di ssect ing a
platean." i\lcDiarmid et al. (1976) reporte d an individ ua l [ound
on a road which was 3.3 km west of a pine forest , pr ob abl y in sub-
tro pical dry forcst (Hard y an d 1969). Road tcmperature
was about 27C and air tempcr ature ca. 24"C whcrc th e snake was
collccted , A small mal e (UTA H-.5926) and Iemale we re collecte d at
Plomosas. Sinaloa. at 1067 m on 3 Augu st and 27 August 1976,
res pcct ively, The snakcs were Iound in a trausition arca bct wcen
pine-oak ami tropical deciduous for est (Fig. 4). Vegeta tio n in t his
arca includes slk- cotton tr eos. morning glory t recs, Ente rolobiuni,
Coccololia, nanche (B!lrsollil/la crassiiolta), cass ias , acacia s, and
mimosas (Cold man 19.51). The ma le wus Iou nd in a rock slidc at
the base of a blu/f , an d th e female was secured in a small rodent
burrow in a se mi-opon fielcl. A large mal e (UTA H-6234; Fi g. 31)
was foun d on 1.5 August at Eji do Tcbara, Sinaloa. at 1067 m in a
t ropi cal clcciduous for esto This individual mcasured 638 mm tot al
length, 86 mm tai! length, ami is largor th an the largest snake re -
portcd by et al. (1976).
Crotal us tigris Kenni cott
Crot alus tigris is a m dium to small-si zed desert dwelling ra ttle-
snake with a proport iouutc ly small hcnd and large ra tt le. Thi s
spe cies has a rather limited range, ancl is kuo\Vn only fr um th e
stat es of Ari zo na a nd Sonora. Wright amI \ Vright (19.57: 1001) con-
siderccl this spccies rareo
Cro/alus lgris is a resident of rocky foot hills wit hi n th e Sonaran
THE NATURAL HISTORY OF l\IEXICA N RATTLESNAKES 51
'.
,
, , ,
r .. _ : : ~ . ~ . , ":1..
. .'.
- .. ,,"
FrG. 29. Crotalus seu / lila/lis salci ni . Spccmen Irorn 1.6 km E El Limon
Totalco, Vera cruz, l\l exico. ( Phot ogr aph by John H. Ta shjian.)
FrG. 30. Habitat of Crotalus scui ulat us salci ni and Sis/rurus raous, near
El Liman Tot alco, Verac ruz, M xico. Area is lava low charac terized by
cactus, agave and grasses. (Photograph by R. Terry Basey. )
52 SPECIAL OF NATU RAL HISTORY
FIG. 31. Grata/li s st ei ncgeri. Specimen from Eji do Tebaira, Sinaloa,
Mcxico. ( Pholograph \'r [onath an A. Campbell.)
Desert of Xlexico and the United Sta tes , and rangos from near sea
leve] at La Posa , Sonora, to at least 1465 m elevation (Humphrey
1936). We found an individual snak e at the same elevat ion on th e
west crn slopes of the Santa Rita \Ioull tains, Arizon a. Van Den -
burgh (1922) rccorded the maximurn elevation for thi s spcc ies at
:2440 m. C. tigris is rar ely Iound Far from rocky rct reats, Lowe
( 1964: 173) conside red th s rattlesnake to be strictly a rock dwell er
in rocky canyons, and on the hillsides and bajadas of desert ranges.
Three snakes ob tained by us from 11.2 km l\'E Scottsda le, Arizona
(UTA R-6943, KU 1555:25) werc takcn at ni ght in early [ul y just
af te r a thunderstonn. They were active among gra nite outcrops
where cacti, palo verde, and creosote were abundant. This snake
is nocturnal and rarely venturos Iorth before surnmcr rai ns begin.
Taylor (1936: 49) found two C. tigris at ni ght, south of Hermosillo,
Sonora, coiled in isola ted rack outcro ps in low moun ta ins. \ Ve ob-
served a snake of th is specics from the Tucson Mountains, Ari zona,
at night, as it was investigating a pnc k rat nest. This arca is very
rocky , with much desert vegetation. Dixon el al. (1962: 99) de-
scribed th e hahitat of C. tigris in southem Sonora as scru b deserto
Crotalus ti gr is is beh avioral ly unprcdicta ble. Two specimens
[rom near Scottsda le wer e locat ed when they rattl ed as wc passed
within 3 m of them. This observation occ urre d on an extremely
warm, humid evening, aft er a thunderstorrn. A speci me n from the
Tu cson Mountai ns never ratt led, even while bein g capt ured,
THE NATURAL lIISTORY 01" ~ I E X I C A N RATILESNAKES 53
Klauber (1972: 453) felt th at this rattlesnake was gener all y inofen -
sive, and not pr one to rattl e or strike.
The Seasonal Fonnation Ser ies and Cactus Scrub as deRnecl by
\Vagner (1964) are typical environments inhabit ed by C. tigris.
Crotalus i ortugensis Van Denburgh and Slevin
Klauber (1972) characterized Tortuga Island off Baja California
as rocky nnd barren, more ar id than the mainland, with sparse
brush ancl cac ti. Three specimens of C. iortugensis obtainecl by
liS (KV 174832) were coIlected on 27 Oc tober 1975 in an area of
lava boulders (D. Brown, pers. comm. ). \Vagner (1964) character-
ized th e vegetational charact er of Tortuga Island as Seasonal For-
mation Seri es.
Crotalus iranscersu s Taylor
Crotalus transoersus is, biologicaIly, a poorl y known rattl esnake
both in the Reld and in captivity. Its type locali ty was listed by
Smith and Taylor (1950) as Tres Cumbres , \ Iorclos, but Davis and
Smith (1953) rcassignecl th e type locality to nearby Lagun a Zem-
poala, J\lexico. The lat e E. IT. Taylor (pers. comm.) infonned us
th at th e typ e locality s Tres Cumbres. Further coIlecting may
reveal the presence of C. irauscersu s in northwe stern Morclos and
southwestern Di strito Fe deral du o to the top ogr aphi cal similarity
of these reg ions to that of th e typ e locali ty.
Only three individual s of thi s species have been recorded from
the t ime of it s discovery in 1942 until 1971. Since the n nine speci-
mens of C. transcersus havo been securecl, sx of which are d s-
cusse d here (Figs. 32, 33).
On a tr ip to southern Mexico in the sumrner of 1973, Charles
RadcIiffe and one of us (BLA) succeeded in coIlecting Iive of th ese
snakes (UC\ 1 51421-3, KU 159361-2). An addi tional speci men was
co1Iected in May 1975 (UTA H-3988). AIl were taken at Laguna
Zempoal a, Mexico (Fig. 34) bctween 1 August an d 5 August at
elcvatious ranging from 2896 to 3293 m. The habitat was t emp erare
boreal forest as deRned by Lcopold (1950). Five specimens we re
observcd basking on south-facing slopcs on volcanic rocks in th e
earIy afternoo n. Non e of th e specimens was basking in di rect sun-
ligh t. The snakes were found after mor ni ng rain-showers which
occur almost dail y at thi s locality du ring summer months. Air
t emper at ur e at the time of coIlect ion was bet ween 16-20C.
C. transccrsus is a very inoffensive rattl esnak e, and retreats
rather qu ickl y when di scover ed. l\one of thc capti ve spe cimens
ma intaincd by us rattl cd or struck, unlikc C. t. tris eriatus wi th
which th s species is sympa t ric. T. \ Valkcr (pc rs. comm. ) observed
that only onc of hi s three captive specimens ever rattl ed , and be-
lieves that th s species is not rar e, but is onl y though t to be so
54 SPECIAL PUBLICATION-MUSEUM OF NATURAL HISTORY
" .
F IG. 32. Crotalus transcersus. Speci men from near Laguna Zempoala,
Mexico, Mexico. Reddish color phase wi th distin ct cross-bands. ( Photogr aph by
John 1I. Ta shjian.)
r
......
FIG. 33. Crot alus transcersus. Specimen from nea r Laguna Zempoala,
~ I e x i c o , Mexco. Crayish color phase with indi stinct cross-ha nds, ( Photograph
by j ohn H. Tas lij an.)
THE NATURAL HISTORY OF MEXICAN RATTLESNAKES 55
F IG. 34. Habit at of Crotalus t ransc crsus and G. t riseriatus triscriatus near
Laguna Zcmpoala, M xico, Mexico. Op cn fir-pin c-oak forest wi th unde rstory
of bunchgrass. ( Photograph by ~ 1 . Cranger.)
56 SPECIAL OF 0iATURAL IlISTORY
because of its limited ac tivity periods in its natural ha bitat. An
abundance of lizards und mice in the preferred hab tat of C. trans-
versus probably results in the need Eor only limited forag ing ac-
tivit y. Klauber (l92) men tion ed that one individual contained
l zard scales, and one of our snakes defecated l zard scales, probably
Sce loporus ae ne us.
A numb er of Food items were offcrcd t o captive C. transoersus:
treefrogs (HUla), Izards (Sceloporus, Cerrhonotus, Uta, Plujllo-
dacuj lus, Xantusia, Cnemidophorus, Anolis}, wild rodents (Mus
musculus], snakes (Thamnophis, Nerodia), newborn laborato ry rats,
sparrows, and domesti c gray crickets (Crijllus) : all were refused .
The rodent s we re kill ed by the snakes, but the other potential pr ey
it ems wer e ignor ed.
Captive snakes of thi s species were maintained by us at the
Following thennal Ievels: (1) 26.5C during the day and dr opp ng
to 10C at night; (2) 2.5-27C during the day, 20C at night; (3) low
20C tempcrature at all times. Thcsc snakes may have becn affected
by the change in eleva tion from their nativo habi tar to captive con-
ditions. They fared badly at all three thermal levels.
Crotalus tri seriatus (Wagler)
The dusky ratt lesnakes of south-central Mexco ar e, taxo nomic-
ally, one of the most complcated groups of rattl esnakes yet de -
scribed. \ Ve ha ve mad e over 100 obse rvati ons of both subspecies,
and thi s field work has shed some light on their relationsh ps. It
has becorne apparen t to us th at the two curren tIy recognized sub-
speces, C. t. triseriatus und C. t. aquilus, require a complete review,
and such work is underway by several of our colleagues .
Crotalus triseriatus tri seria tu s. Crota lus t. trtser iatus is a mon-
tane race reaching the highest altitude (4572 m on Mt . Ori zab a,
Veracruz) at which any member of th e genus Crotalus has been
recorded (Klauber 1972: 516, 542). Although Eound at such high
alti tudes , C. t. tri seriatu s is more commonly encountered at eleva-
tions betwecn 2743 m and 3353 m. Specimens have been recordcd
by us from: JALISCO: the mountain valleys N Nevado de Colima
(UTA 1\-4909) (see Campbell , 1979); \ IEX ICO: Laguna Zempoal a;
DISTHITO FEDERAL: La Cima; t\IORELOS: Tres Cumbres,
the lava be ds N Cu ernavaca, Huitzilac (Fig. 35).
Crotalus t. tri seriatus probably occurs in the Mexica n state of
Puebla, bu t we have no records as yet from that ar ca, It also may
be Iouud on Cerro La Malin che, a 4267 m peak in Tlaxcala , Resi -
dent s in this area describ ed to us a snake wh ch appeared to be
C. t. t riser atus, but no specime ns have be en secured. In general,
the range of C. t. triseriatus foll ows a narrow east-we st belt be-
tween the 18t h and 20th parallels. Thi s range includes Cordillera
Volcnica oE souther n Mexco,
THE i\'ATURAL HISTORY OF RATTLESNAKES 57
Crotalus t. tri seriatus is one of th e mo st common fonns of th e
small Mexcan montano rattl esnakcs. Due to th e high clevations
at whi ch it occurs, th s subs pccies is almost entirely diurnal in
activity . Evcn during the wa rmer months of the year, temperatures
fall to level s at nig ht wh ch would no t permit ac tivi ty by this snake.
Consequentl y, th is rattlesnakc is mos t easily obs erved in morn ings
and aftc rnoons when th c sun has sufficicnt ly warmed it s habitat.
Thcse animals are also active during th e rainy season, and it is
not un common to find severa] Iollowing an aft ernoon showe r, bask-
ing on rocks th at are still warm from the carl icr ra ys of the sun.
Dav s and Smith (1953: 141) found C. t. triseriat us common near
Huitzilac, Morclos , along streams th at wcr e bordered by a luxu rant
growth of bunch grass. Ou r observntions agrcc with th crs, this
-snake is rarel y found any di stancc from a rocky, grassy enviren-
mcnt. \Ve have obtaincd snakes on rock slide s near Laguna Zem-
poala, Mexico, but they are not nca r ly so common th er e. Duellman
(1965) mcnt oncd th s spcci cs as an inhabit ant of th e Pine-Oak
Forcst at 1600-3270 m altitudes.
Crotalus t. tri seriatus exh bits beliavior tha t is comparable to
th at of othcr rattl esn akes in montan e habitat s. lt s al ert and ready
to retrcat whcn annoyed , yet cur ious cnough to expose it self to
capture, This subspccies is rclativcly unaggr essve evcn when be-
ing capture d. \ Vc havc obscrved ritual zed combar b etween
captive mal es. C. t. iri seriatus fccds on lizards and small rodents
Fr G. 35. Habitar of Grata/lis t riscriatus triseriatus and Sistrllrlls racuc.
La Cima, Dist rito Fed er al, Mexico. Old lava Ilow covered with bunch grass,
Ior merl y a pine forest. ( Photograph by Gr anger.)
58 SPECIAL 0 1' NATURAL HISTORY
which are pl entiful in it s preferr ed habi tar. Cursory examination of
stools from Freshl y caught specime ns supports this diet pr efer ence.
011 15 jul y 1970, a female from the northern drai nage of Nevado
de Colima, ] alisco, wa s located in a partia lly cIeared pne-oak
forest under a log, along wit h six new- born young. A captive
female pas sed a part ally formed embryo on 1 ]ul y 1976. Four
young wer e born on 30 Oct ober 1975 to a ca ptive Iernale collec ted
at La guna Zempoala. Hange of variation is as follows : total length
1.59-178 mm, mean 168; S-V 141-161 mm, mean 150; weight 4.8-
5.1 g, mean 4.9, A ca ptive pair was obs erved in copulo on 24 April
1975 at 0800 and wer e separated at 1400. The same pai r also bred
on 8 September 1977.
Crotnlus tri seriatus aquilus . Thc northern subspecies of thi s
rattlesnake, C. t. aq uilus (Figs. 36, 37), does not reach the high
altitu des inh abited by C. t. t ri seriat us. lt seems to prefer the open,
grassy habitat s north of the Co rdi llera Volcnica. Speci mens as-
signable to th is subs pecie s havo been recorded by us fro m:
16 km w Jiquilpall ; SAN LUIS POTOS : the
vicinit y of Alvarez; HIDALCO: ]acala , San Vicente, Dur ango, El
Chico (KV 1. 55549-553, 1555.56; UTA 1'-4.540, 6941), La Estanzuela
(UTA R-4675, 6115).
This ratt lesnake is a cornmon montane snake, but. as previously
state d, docs not usuall y occur in extremely high-a lt itude habit at s.
Klauber (197:2: .517) gave 2438 m elevatou, nea r Jacala , Hidalgo,

..
...
.-,
,.
"
" .
... .
"
Ot o .,.
, ,
:.
.; -.. .
-. ,
\ ' .
.
.\
.....
. .. .. ... .. . ......
Frc . 36. Crotalus triseriatus aqu ilus. [u veuile specimen from near El
Chico, Hidalgo, M xico. ( Photograph by [ohn H. Ta shjian.)
THE NATURAL HI STORY OF ~ I E X I C A N RATTLESNAKES 59
.'.
. ;
1'1(; . 37. Cro talus t riseriatu s aquilus. Specimen from near El Chico,
Hidalgo, Mexico. ( Photograph by [ohn 1I. Tas hjian.)
as th e high est point where G. t . af/ II/IIS has been collec ted . II ow-
eve r, we havo locatcd a pop ulat ion near El Chico, Hidal go (Fig.
38) wher c th ese rat tl esnakcs have hcen foumI up to 3110 m in a
fir forest. The snakes from th is popul ation, while fal\ing wcll wi thi n
the known ge og ra phic range 01' C. t. aquilus, are quite t1ifferen t
from 0\11' ot he r specime ns of t his racc, Thc y are largor and exhibit
a gr eenish hue, with mal es far excccdng Jemales in size and
brig htn ess of color. Thcse spec imeus tcnd to be hcavier borlied
t han typical G. t. aquilus and they ar e extrcmely alert an d ag-
gress ive. I n many cha ra ctc rs tics th ey rcs ein bl e G. lcpidus. Furt he r
study 01' thcse sp ecimens is necd cd in arder to uuders tand th cir
status in relation to other popul at ions of G. t . aqui lus.
Cenerally, G. t. aq ui lus is an inhahi tant 01' pinc-oa k forests and
opc n mcsquite-grasslands 01' t he sout hern cen tra l Mexican p latcau
(Fig. 39). Snakes fr om Mi chouc n, t he southe rnmost po rtien of th e
muge of G. t. aquilus, usually inha bi t mes qu te -grass lnnd. No rt h-
ward to Hidal go and San Luis Potos, thi s race tends to occur
more commonly in th e pine-oak fo rcs ts whc re th ere is an abunda ncc
of rack outc roppi ngs .
The behavor of G. t. aquilus in tit e Iicld is much th e sume as
that of G. t. t riser atus, but individual s of th e former teu d to he
some what more aggressi ve and casi ly ngitated. These sunkes often
reveal th eir presence by rattling wh en alarmed, as is typical of
60 SPEClAL PUBLICATION- MUSEUM OF KATURAL HI STORY
FIG, 38. Habi tat of Crotalus t riseriat us aquilus, nea r El Chico, Hidal go,
Xlexico. Fir Iorests cove r hills rea ch ng elevations exceed ng 3110 m. Snakes
most commonly found in open roc ky outcrops, ( Phot ograph by j onathan A.
Carnpbell. )
FIG, 39, l Iabit at of Crotalus triseriatus aquilus, near San Vicente , Hidalgo,
~ [ e x i c o . Stcc p hillsides are cove rcd with pin e-oak foresto Snakes most corn-
mon in ope n limestone bedrock. ( Pho togra ph by H. Terry Basey.)
THE NATURAL BISTORY or RATTLESNAKES 61
e. le pltlus. This race is active in th e momings und afte rn oons, but
unl kc e. t. tri seriatus, it mu y be Iound foraging at ni ght. \ Ve have
found them whil e dri ving at ni ght in th e vicinity of Duran go,
Hidalgo , a somewhat humic1 ar ca wher e night temperatures are
occasionalIy fa vorable for activit y. Ritualized combat b etw een
captive males has been obscrve d by us o
Two Icmales of e. t. aquilus Ir om El Chi co, Hic1algo, gave
bi rth to six and seven young on 7 [une 1974 anel 29 [uly 1974, re -
spectively. Range of va ria tion of th e two broods is as follows:
total lengt h 168-192 mm, mean 178; S-V 1.53-177 mm, mean 163;
wei ght 5.0-8.8 g, mean 6.4. Anothcr female, collecte c1 16 km \V
[ quilpan, Michoac n, ga ve birth to threc young on 29 June 1974.
Ran gc of va riat on is as foIlo ws: tot al length 164-171 mm, mean
167; S-V 1.50-156 mm, mean 1.52; weight .5. 0-6.4 g, mean 5.5. A
captive pair of e. t. aquilus From El Chico was observec1 copulati ng
on 28 April 197.5. One of these snak es later gave bir th to two
via ble and one c1eformec1 young on 27 [ uly 1977, an d the measure-
me nts anel wei ghts ar e as foIlows: total Iength 120-181 mm, mean
1.50; weig ht 4.9-7. 8 g, mea n 6.3. The dcformed snake exhib itcc1 a
fusio n of the ventral surfacc in th e lower cervi ca l ar ca (UTA R-
6940). Ano ther pair of e. t. aqui lus wa s joinec1 in mati ng at 0830
hou rs 011 26 October 1977. The Icmal c's cloaca was disten ded an d
th e male's boc1y pulsatec1 (twice pcr SeCOl}(!) ncar th c ve nt c1 uring
coi tus . The uplift ed tails of th e snakes produccd a ve ntc r-to-vente r
configuration in th e regi on of th c cloaca. Separatiou occurrecl at
0901, and detumescen ce of the hemipeni s occupi ed 7 SCcOllC!S.
Ducllman (HJ6.5) charactc riz ccl the vegctnti onal charac te r of the
habitat of thi s rattl csnake as and Pine-Oak
Forcst at 1600-2000111 altitudes. This racc is found in thc Tcmperate
Mcsquite-Crassland and Pine-Oak Forest as defined by Lcopold
(19.50).
Crotalus viridis (Hnfiucs que)
Crotalus viridis caliginis. Cro tulu s v. caliginis is a ra cc [ound
on Sou th Coronad o I sland off th c northwcst coa st of Baja Cali-
forua . Thi s arca is roc kv with hrush ami cac ti, and two adult
spcc ime ns (KV 1748.35, u'TA 1\-694,5) wcr e collectc d at th c north
end of th e island on 2.5-26june 1D76 (D. Brown, pers. COIl1I11.). The
female passcd one partiall y fonned cmbryo on 14 Augnst 1976.
This subspccies is found in the California Chaparral as defined
by Wagn cr (1964).
Crotalus viridis hell eri . Crotalus v. hclleri is a subspccics found
in southe rn Californ ia and nort hcrn Baj a California, und is a resi-
dcnt of almost every hab it at wi thin its range. As with e ruber,
with which it shures a great dcal of its range, a numher of e. v.
lielleri ha ve [icen observed by lIS every month of the ye ar. These
62 SPECIAL 01' NATU HAL HI STOHY
snakes vary greatly in color and pattern within a give n population.
As individuals 01' th is rattlcs nake age, th ey apparently lose th eir
pattern 01' it fades great ly, a common t ra it 01' all races 01' C. oirulis.
Mos t largor specimens 01' C. v. hcllcri are hlack 01' very durk with
li tt le pnt tcrn, Many young 01' thi s race have dstinctl y patt erned
heads, but thi s fades at an age 01' two ycars ou most individuals.
C. v. liclleri ranges from San Lu is Obispo and Kern count es
south th rou gh southern California (avoi cling only th e true deserts
in th e eas te rn part 01' th e state) aud con t inues into Baj a California
as Iar south as Playa .\I ara Ba1' on th e west coast, and ncar Baha
de Los ngeles on th e eas t coast (Klauber 1949b : 8.3). Xlurray
(19.'5.'5: 47) reported a specimcn 3:2 km SE Mcsquital , sorne 88 km
south 01' Klauber' s (1949h: 83) southcrumost record. Smith et al .
(1971) Found two spee imens 69 km \\'i\'W Bah a de Los nge les
anel 16 km S El Arco. Specimens [rom these southc rnmos t locations
ar e rare, altho ugh it is possihl e for C. v. liclleri to inlia h t th e ar ca
as Ia r south as San Ign acio in ce ntral Baja Californ ia. Hattl esnakes
ob tained by us come fro m many localiti es in southern California,
but we have Few records from Baj a Califo rnia. Individual speci -
mens from lexico have hccn observc d Irom Pun ta Camal, north
01' Ensenada, an d th e westem slopcs and int erior peaks withi n th e
Sierra Jurez.
Crotalus v. hcll cr i is couunon within its range, although its
num bers seem to dccr easc in the southe rn portions. On the 10\Ver
slope s 01' the San Gabriel Mount ains in Los Ang eles Co un ty, Cali -
fornia, the populations 01' thi s race are as co nccutratcd as any 01'
the genlls known to us. Thi s is a rugged, rocky arca with chaparral,
and the ra ny seasou occurs in th c wiuter an d earl y sp ring. Hatt le-
snakes havc hcen Iound fro m th e edges of the Sa n Ga briel Hiver
to th e hot , sparscly vcgetatcd, south-Iucing slopcs and pine-oak
slope s facll g th e north. Altho ugh individuals of C. v. liell cri in thi s
arca do not de n 01' hibernate as such, th ey do congreg atc in rock
slides 01' rock outc rops which [ace th e south 01' eas t during rain)',
wint er months. '1'0 the south in Or.mge, Hiverside, and San Di ego
counties. thi s subspecics is Iou ud from th e Pacific coast in rolli ng,
grassy savannahs to at least 3200 JI} in a pine-fir forest 011 '\[ t. Sa n
Jac into. Throughout mos t of thc se southern California count ies,
C. initclielli JUrrlllls anel C. r. T111Jcr live sympa tr icnl lv wi th C. v.
llelleri . \Ve have observed al1 three ra tt Icsnakes wi thin 45 m of
caeh other near \ Varner Bot Springs, jusl 1I0rlh 01' th e Borrego
Desert. 111 lhat particular ar ca, all three species sepm eq ua ll1'
abunda nt. Elevalion in th is arca is approximalel1' 1068-1:220 m.
111 1967 oue 01' us (BLA) found an apparent h1'hrid , C. v. helleri X C.
r. mber, on lh e road in th is s:lIne ge neral ar ca. Hep orts of hybrids
fronl thi s arca ar e nol 1II1COIllnHlIl.
In Baja California, C. v. ' IClleri is Illllch less comll1on amI is
THE NATU RAL HI STOHY 01" },IEXICA0: RATTLES0:AKES 63
apparent ly rcp laccd by e. ciuj o [ur cu s, e. r . ruber, and e. mit clielli
pijrrhus. A snake Irom Camul was observed in sc rubby, coasta l
vegetati on approximatcl y 9 m Irom th e sho reli ne . Another example
from th e west ern slopes of th e Si erra ju rez was Iound baskin g in
diffused sunlight under a manzanita bush, Klauber (1949b : 83) re-
ported a specimen 16 km 1\'W Ba h a de Los Angeles in desert
mountaius in an elephant trec association.
Crotalu s v. hellcri t ends to be a ra ther aggressi\' e Io nn and is
cas ily agitated, b ch avi or reportcd by Kl a uber (197:2: 454 ). Most
adults wi ll dcfend th emsel ves vigorously,
\ Vagn er (1964) cha racte rizcd Ihe vegcta tional charactcr of the
habit ut of e. v. liel lcri as Cu lifornin Chaparral and Xlont an e For-
maton Seri es, und Leopold (19.')0) cha racte rizeel it as Pinc-Ouk
Woodlund nnd Pion-junip er \ Vooellanel.
Crotalus icillarili Xleek
T he ridgenose ra t tl esn uke. e. icillardi, is an exaru p le of a smull,
montano spec ies whose hiol ogy has rcmaincd somewhat obscure,
))0 doubt due to diffi culties cuc ountc rc d in reach ing possihl e Xlexi-
ca n collcct ion localiti es. Littl e is known of it s huhi ts, hut it eloes
not appear to b e rare wth in its rauge. The four raccs 01' thi s spe-
ces collective ly inhabit th e Sie rra .\I aelre Occi denta l Irom south -
easte rn Ari zona aud so nthwcst crn 1\ 0 \\' ~ Iexico, southeas tc rlv to
southcm Durango and wes t c ru Zacatccas (Kla uber 1919c: U .5).
Crot alus willur d wil lardi , The northern subspccies. e. l e.
icillardi , is a resident of th e pine-oa k Io n -st s of t he l Iuuchuca and
Santa Hit a Xlountains in Ari zouu, a nd th e Sierra de Ajos and Cerro
Azul in So nora, Xl exco. Thi s ra tt lesnakc scems to pn-fc r th e more
humd cauyon bottoms of pine-o ak habit ats as opposed to exposed
arid slopes wher e e. lepulus ami e. pricci are conuuon. B. Toruber-
lin and H. T. Basey (pcrs . COUlIl!. ) rr-portcd that iud ivid ua ls Irom
t hc Si erra ele Ajos (UTA H-66S9, H-GB:2S-30) werc conu no n in thc
wide, rock y st rc am beds whcrc pool s of wat er r emained a fter su m-
mer rains. Thi s was between HJS2 ni uud 2287 m in elcva tion, '1'''' 0
lizurds (ScclapaTl/s jarrad and S. uiululatus were cornmon in t hi s
st rcum hed hahitat. and no do uht co nt ribute a ma jor portien to
th e di ct of thi s snake. Kl au bc r (1972: 61:2) mentioncd a spccinu-n
th a t conta inc d a brush mousc ( Pcronujscus houli}. Alt ho ugh prc-
fcrring th e coo ler, hu mid cnnyons, e. w. willardi is in no \ \ ' a ~ ' re -
st rided to this habitat. A sna kc from upper .\I adera Call yon at
2593 m in t he Sa nta Rit a .\ \ ollntains was obs erveo on an east -faci ng
slope amidst brok en rock a nd sc m b oak . It ",ould ll ave been O\' cr-
looked hao it not ra tt led, fo r lil e a rca ",here it was foumI is steep
amI the sc m b oak nearly impenetrable. A specimen of e. pricei
\Vas locatcd nearb\' . e. IV. Icillan li is nlos t acti\'c on \\',lI'm, humid
days whcn the sl;nlight is in tcnnitl en t. It is es pecially COllllllon
64 SPECIAL OF NATURAL HISTORY
befare aneI af ter an a te m oon shower when the temperature is
hetween 24-29 C.
Crotalus lV. icillardi is typica l of th e small, montane rattlesnakes
in tha t it is nervous an d irri t abl e, although not to th e degree ex-
hihited bv C. lepidus or C. pri cei . At times it reve als its presence
by ra ttling, and w ll cru wl to sa fe ty upon th e 1cast provocation.
Captured spec mcns a re more apt to turn and hite, as opposcd to
typical ratt les nake stri ki ug behavior. Kaufleld (1943b : 3.57) aneI
Stehhins (19.54: 484) aIso found this to he true. Aft er a shor t period
of time, captivc spccimens of C. lV . icillardi gen er all y calm down
aud maintain a peacef ul d isposi to n.
Ma rt n (197.5'1, 197,5h) di scussed reproduction in C. te. ioillardi.
011 18 Apri l 1971 a captve puir was observe d b y us in capillo at
OSOO hours, and they wcrc stil\ join ed at 1700 hours when obse rva-
tions were discontinucd. There wcrc no res ults fro m thi s mating.
Crotal us le. icillardi is fonn d in th e Xlonta ne Formati on Seri es
ami Xlontanc Thieket as defined by Wagner (1964) and Pine-Oak
Forest and Pine-Oak Woodland as dcincd b y Leopold (19.50). : See
Cloyd (1837 ) ancl Lowc (1964) for a clcscri ption of th e habitat in
Ari zona.
Crotalus willurdi amabilis. Fin adult (KV 178794 ; Fig. 40)
a ncI Iou r su hadult C. te. amahihs (UTA H.-7162) were taken in
Arroyo Meste o. Sierra del Nido , Chi huahua, at 2440 m 011 18, 19,
21 JI'tly 1877 (L. A. Xl tche ll aud D. G. Barker, pers. comm.). Most
of the snakes were loeat ed at th e bases of small tr ees in pil es of
twigs and leaf litt er ncar strearns , one was Iound in a shin-oak
association, The arca li ad bcen logged three times in twelve ye ars,
ami the las t Iogging hacl occurrcd four ycars ea rlier. Al! snakcs
wcre vis ible, sorne coi Ied ami others crawling, and onl y two rattl ed
whcn captured. I nt ermitt ent thund er storm ac tivity had oecurred
for two weeks pri or to 18 Jul y, and oeeasional showers oeeurred
during the collceting period. The ambi ent temperature was ca.
21C, und the snakes were foune! throughout th e day. T\Vo C. p.
pri cei , 011 e wirhin .5 m of an C. IV. amobilis, and two C. Iepidus
klouber, one in seruh-oak '1]](1 th e other in a talus slope near shin-
oak , werc also foun d . Two C. te . atnahi lis def eeat ed rodent hair in
th e Iaboratorv. H. K. Cuese (pers. eomm.) obs erve d a ca ptive mal e
eourting a f emale on 8-9 September 1977 . The Iemale rernaincd
in a resting coi l and th e malc direete e! head-bohbing and tongue-
f1i cking across the female's do rsum (3-.5/.5 seconds ). The male
moved his uplifted tai l in hoth a hori zontal amI vertical pl an e with
a n lIndlllati1l g mo tion, Physieal conta d with th e side of the en-
c10sure or th e sllhst ra te stillllllatc d t his hehavior whi eh \Ve interpret
as TaiJ Seareh Copu latory At te lllp ts. The seq ue nee last ed .5 minut es.
Crotalus willar di Illcridionalis . The sOllthermnost sllbs peeies,
C. le. Ill cridi ollalis, inhabi ts a limit ed area in southern Dumngo amI
TIl E l'iATU RAL HISTORY OF },IEXI CAN R.ATTLESNAKES 65
ext re me southwe stern Zncatec as, ano few specimens are ava ila ble
for study (Cloyd 1940; Klauber 19(2). Like it s siste r mees, C. tO.
merulionulis is a montan c inhub itnnt ra rcly Iouncl bel ow 2440 m.
This subspecics is not genera lly a canyon-bo t tom dwell er. The
arca inhabitcd by it is hi gh (2440 m) pine-cover ed pl ateau s where
op en mcadows previd o breaks in th e for esto Five indi vidual s ob-
tained by us from near Llano Grande, Duran go (Fig. 23; UTA
R-5639-41, 6124-2.5), wc re collec tc d on 4 Aug ust 1975 on a partiall y
opcn hillside which slopcd into a gra ssy meadow. The slope had
a northern to northeast ern exposllre. AII specimens were locat ed
at ca. 2.560 III a t a tcmp cratur e of 21-24C betwcen 0830 an d 1000
hours. 1t liad raned the previou s evcuing a nd cont inuc d until
0600 whcn th e sun appcared. The cnvironme nt was hurn id when
th e rattl esnakes wcre ca ptu rcd, an d none was found afte r th e high
clouds liad di sappcared . The hab itar, alt ho ugh not a canyon, was
simila r to th at occupi ed by othcr ra ces of C. u.illurtl i. It was com-
posed of sparse short-needled pines mi xed w th equally sparse oa k
and madro o. Rock s wer e scat ter ccl throughout the scrub-oak, and
grassy grollmI cover provid cd cxc ellcn t ret reats. AII five specime ns
were ncwborn YOl lllg and had sh ee! t heir skin at Icast once. They
exh ibitc d gClltle di sp ositions and re fusc d to bit e upon capt ure. A1I
adult mal e C. ti; . mcridionulis was [ound 011 18 [ uly 1976 a t th e
eres t of a partiall y dcnudcd hill with a Iow den se covc ring of
scruhby brush . It was hasking in dircct sunlight at 1100 hours. An
adult femal e, opaque-col orcd pri or to shcdd ing, was discovercd at
Fi c . 40. Cro/U/IIS usi ltardi amahilis. Specimen from Arr oyo Mest e o, Sierra
del Nido, Chi huahua, Mexico. ( Photograph by Jonathan A. Campbell.)
66 SPECIAL OF NATU RAL HISTORY
1000 hours on 12 Jul y 1977 during an intermittent rainstonn. Hea vy
fog shroud ed the ar ea in the early morning. The ambient tempera-
ture wa s ca. 21C, and the snake was resting in shade. Seven addi-
ti onaI sna kcs (one preser vcd , KU 178975) werc taken in th e upper
reg ons of th e mountains at a ranch near th e city of Durango (S.
Pr chal , pers . cornm.).
Ths snake inhabit s th e Montano Fon nat ion Series as defined by
\Vagner (1964).
CrotaIus will ardi silus. Crotalus w. silus (Fig. 41) is aIso a resi-
dent of the temper at e, pine-ouk for ests in the Anima s Mountains
of Ne\V :\l exico (Bogert and Degenhar dt 1961), the adjacent Sierra
de San Luis, the Sierra de la Purica (Sierra de Nacozari) in Sonora,
and the mounta ins to the immediate east and south of Chihuahua
(Fig. 42). Crotalus w. silus is ver)' similar to C. w. ioillardt in its
habi tat prcf cr ence. Snak es from the Animas and San Lu is ra nges ,
howcver, cxist in a much drier climate since these mount ains re-
ceive less ra infaIl than mountain s which support other C. w illardi
popul ations. Three indivdu als observed by us in the Animas
Mountai ns we re founcl in lat e Oct ober and earIy May whe n it was
very dry. AH th ree specimcns werc located at an elcvation of 2120
m on east-faci ng sIopes wher e pi nes, scrub oak, and manzanita
comp rised th e major vegetat ion . None of th e snakes rattIed or
tried to escape , nnlike other C. w. silus from humid habitats.
\ Ve observ ed approxima tely 40 C. w. si/us bet:ween 1970 and
1974 in the Sierra de la Pur ica in northem Sonora (KU 155554-555,
,.
FrG. 41. Crotalus willardi si/uso Specimen from Sierra de Nacozari, Sonora,
( Photcgraph by [ohn 11. Tashjian.)
THE NATUR.AL HI STORY 0 1" RATTLESNAKES 67
158562, UTA R-6942). No other member of the genus has been
observcd in thi s rangc, which contains perfect C. icillan]! habitat
consisti ng of lon g, large canyons wit h a south to north strcam-Ilow
wher e wate r is present th rou ghout th e yeaL Pines and oaks dorni -
nate the vegetation wi th syca morcs in stands along th e wide can-
yons. Fems and poison oak provi de ground cover in th e rocky
stream beds, where C. IV . silus is mo st preval ent . Snakcs have been
see n from April t hrough October , wit h the greatest uumbers being
obse rvc d in th e [ ul y-August rui uy scason, At th is time. gcneral ly
cloudy skies and hi gh hu midi ty provi de excelle nt cou di tio us for
these snakes. On on e morni ng in carly Jul y wc obs ervcd ele ven
specimc ns in a two hour pc riocl while hiking in a single cauyon at
elcvations of 1630 111 to 2300 11 1. Xlost of the suakcs appcarcd to
be foragillg, since only on e of the eleve n wa s observed in a coiled,
basking position. Of th e four xpecimc ns coll ected that morning,
on e gravid femal e gavc hirth to Iive viab le young on 7 August 1970.
Another cont ained four fully clcvelop cd embryos.
Ten femal es and one mal e were observcd in the Sierra de la
Puri ca betwecn 15-2.3 [ ul y 1978. One pair was copuluting on 15
[ul y, and two adult femal es appcarcd as though th ey had just gi ven
birth. Auothcr femal e gave birth to [our young 179025) on
10 August 1978. Total length of the Iour ranged from 146-218 mm,
mean 190; S-V 129-193 mm, mean 160; weight 4.9-6.9 g, mean 6.0.
FIG. 42 . Sierra de la Pur ica ( Sierra de Nacozari }, Son ora , M xico. Habit at
of Crota lus icillardi silus. Can yon bol toms ru n sou th to nor th, pine-oa k forest
wi th Ierns , poison oak, sycamore tr ees. Aspcns in up pe r regi ous. ( Photograph
by H. Terry Basey. )
68 SPECIAL OF NATURAL HISTORY
On 17 April 1971, captive breeding was observe d, The femal e
was placed with th e male at OSOO hours. The mal e appearecl to
recogni ze th e presence of the female within 10 seconds , began
twitchi ng (1 twitch pcr second), and tested the en ti re dor sal surface
of Fcmalc's body with hi s tonguc (2 tongue-flicks per seconcl). The
mal e rubbed alon g th e dorsal surface of the female for approx-
matel y 5 mi nutes by ho lding his head at 30 angle and sl ding th e
arca of th e mental scale Iorward for 1 cm. Other mating dates
recordcd were 19 Apri l 1971 and 29 [ anuary 197.3. The ma ti ngs
were unsuccessful.
An interesting observat iou was made by us concerni ng the
brown coloration of indivdu als of C. le. silus from the Sie rra ele
la Puri ca , Those specimc ns colle ctcel or observed aboye th e wa te r-
falls (2070 m) tended to bc light brown, rcsembling th e dominant
pine nccdl e littcr. Othcr specimens frorn below the waterfalls
tended to he a much darkcr brown, rescmbli ng th e more dominant
oak lcaf littcr. Specimcn sizc was apparently no t a fac tor in this
diff erence in color.
Sistrurus rallS (Cope)
Sistrurus racus is a resideut of the tablelancl of south-central
Mexico. lt occurs in a vari ety of habitats wi thin that tablcland,
ranging from ncarly 3050 JII in pine-oak forcs t near Huitzilac,
Xlorcl os, to about 1525 111 in agavc-cacti habitat of the valleys
which surround th e cty of Oaxaca in central Oaxaca , Specimens
have been recorded by us from: PUEl3LA: Zacapoaxtl a, Cacaloa-
pan, Pu ebla (J. A. Campbel l, pers. comm.), TLAXCALA: near San
Dion isio; DISTRITO FEDERAL: La Cima; :\lORELOS: I-Iuitzilac,
th e lava beds north of Cuc rnavaca; OAXACA: near Mirl a, Ixtl n
de Ju r ez .
\ Vith n it s known range, S. racus is a very common ra tt lcs na ke.
It see ms to be rather gr egarious, at least in mountain va lley habitats.
\Ve have observcd over 100 S. racus under mauy vnried conditio ns,
a nd it appears to be a ver y adaptable sp eci es . For cxample, near
Hui tzilac, Mor clos, a largo for ested area had bcen clcared, perhaps
30 years ago, and no", consists of taIl bunch grass (Fig. 43). Sistrurus
Ta ll S ha s occupied thi s ar ea and has become so common th at the
local peopl c ar e extremc ly awnrc of its prcsence. The population
from th is parti cular ar ca (at an clevatiou of 2866 m) conta ins the
lar gest examples of thi s snake of aIl the localiti es whcre we havo
collccted th cm. Several males measured 76 cm and engaged in
ritualizc d combat in ca pt ivity. Many of these mal es tend to be
mc lanist ic, a characte rist ic that Klauber (1952: 114) not cd in thcse
snakes from Tres Cumbres, Mor elos.
Al tho ugh S. Ta llS scc ms to prcfcr grassy mountnin val leys , it
TIIE NATURAL llISTORY 0 1' RATTLESi\'AKES 69
is not restrictccl to that habitnt . It is common in the lava bed s north
of Cu cmavaca, Morclos, where it is sympatric with C. t. t riseriatus
and C. ui olossus nigrescens. S. ra Gll Sfrom this arca are muc h sma ller
than the Huitzilac specimen s. I n parts of Pueb la, Veracru z, and
adjaccnt Oaxaca, S. racu s is found at lower elcvations in arcas of
grass, agaves, cacti, and small , stuntcd oaks (Fig. 30). I ndi viduals
i n thi s h bitat appcar to be mor e disper sed and wer e encountered
less frequcntly.
In the spring of 1974, Jonathan A. Campbell and one of us (BLA)
located an un rcported di sjun ct pop ulati ou of Sist r urus in the central
highlands of Guerrero. Thi s populati on is removed sorne 330 ai r
kilomet ers from the ran go of S. rm)llS as pl ottcd by Klauber (1956:
58; 1972: 64), and is isolntcd from othe r popula tions by the Rio
Bal sas basin to the north and the Chilpancingo gap to the east.
A complete descr iption of th is Cuerreran pop ulat on is for thcom-
ing (Camphe ll and Annst rong, HJ79). However, we think it proper
to describ e her e the habit ar and habit s of these snak es. They
ar e residents of the piue-oak forests of th e Sierra Madre del Sur,
and all of our specimc ns were scc ured in largo, brushy Ilood plains
at vur ious eleva tions bet wccn 295-3105 m. The first sna kcs wcr e
securcd in mi tl-May duri ng the warmer parts of the day. This re-
gion recei ves ca. 1500 mm of rai nfull from through October
(Page 1930). Accordin g to the local inhabitan ts, frost does not occur
at these clevations in the Sierra Madre del Sur, a large rnountain
FIG. 43. Hutzilac, Morelos, :-' Iexico. Habit at of Si st rurus racu s. Snakes
extromcly abundant in bu nchgrass mea dows bor dered by fir-pinc-ouk forest o
( Photograph by Cran ger . )
70 SPECIAL OF l\;ATURAL HISTORY
range th at is m ggecl and stcep, anel would seem to offer Iittl e
sui ta ble h bit at for grassland elwellers such as ra ttles nakes of the
genus Sistrurus.
The habits of Guerreran 5. raVll S are comparable to those fr om
the southern central platean. They are ale rt an cl qu ck to retreat
when encounterecl, but will stand their groll nel whcn escape is im-
possible. Like most Sistrurus, they are more apt t o turn and bit e
rather than eoil and st rike . Stoo ls of some specimcns eo nta ine el
roelent hair.
A female of th is species from thc vicinit y of Hui tzi lac, :\ Iorel os,
gave b irth to seven young on 1 Junc 1974. Two other females
from the same localit y gave birth on May 1975 to nine an d th ree
you ng respectivel y. The range of va riat ion of the three broocls is
as foll ows: total len gth 160-208 mm, mean 183; S-V 138 -19.2 mm,
mean 166 ; weight 3.9-7.4 g, mean 5.4. A captivc pair from l l uitzil ac
was observcd in copulo on 1 May 1976.
This rattl csnakc inhabits th e Tcmperate Pine-Oak Forest as de-
fined b y Leopold (1950).
DISCUSSION
The foll owin g observations and cliscussion pr esen t OUT vi ews on
various asp ccts of th e biology of Mexican rat t lesnakes , and indicate
arc as where research is neeelecl concerning th e ecology an d be-
havior of th ese animals.
Habitat. Mo st mo ntane specics from Xlexico were clcscly asso-
ciateel with rock y arcas ancl such mi crohabitats allowcd m ximum
protecti on. Cr otalus l. lepidus, e. l. klauberi, e. p. pricei , and e. m.
mol ossus were often locatcd in talus slopcs . Othcr mori ta ne rntt lc-
snakes were founcl in pine-oak Iorcst s, mcsquite grassla nds an d
hi ghlanel deserts but usually rocky outcroppings were the dominant
Ieatu re . Crotalus irunscersus oc currecl in arcas of volcanic rock,
wher eas e. pricei niiquihuunus was founel in situ ati ons wi th lime-
stone bcdrock, Sorn e of the lowland taxa such as e. tluri ssus culm i-
uatus inhabited arid tropical scrub for est with roc ky outcroppi ngs.
Rattlesnakes wcre rarely [ound in cloud for ests .
\ Vhen first eliscovereel, smalle r montano taxa such as e. inlcr-
metlius, e. lepidus, e. pricei , e. pusi ll us, e. trausocrsus, ancl e.
ioi llard! wo uld often rattl c and qui ckl y cruwl to safety wit hin rocks.
Many of th ese ratt les nakc taxa sccmed to uti liz o specific retreats,
for in somc cases the snakes crawled toward the coll ector in thei r
endeavors to escape , when sui ta blc retreats were available in th e
opposite direction.
OUT fielel experience indicat ed that Mexica n ra tt lesnakes fro m
higher clcvati ons were most oftcn encou nt ercd in microhabi ta ts
with a southern exposu re (C. i. int crmcdius, e. le pidus kl au beri,
e. l. morulus, e. m . molossu s, e. p. pricei, e. t ranscersus} . A few
THE NATURAL HISTORY OF RATTLESNAKES 71
taxa that appeared to vary from this mode were mo re frequently
found in rnicrohabitats with southeastcrn (e . lepidus maculosus,
e. m. molossus, e. oiridi s he lleri ), easte rn (e . pricei mi quiliuonus,
e. w. u:illarili, e. te. silus], northern an d nort heastern (e. iciltardi
me ridonalis), and west ern exposures (e. int ermedius gloy di ).
Ncarl v all rno ntane ra tt lcsuakes were found aboye a 1800 m
contour.. Lowland t axa which in vade the Mcxican highlalJ(]s in-
clude d e. du rissus, e. molossus, aud e. li asili scu s. Crotalus molos-
sus and e. scut ulatus i nhabi tcd 10\Ver cleva tions in th c nort hcrn
part of th e range. A narrow ver tical dist ri bution at rc lut ively low
elevati on characterized the habt nt 01 e. steincgcri (and perhaps
e. lannoinii. Sist rurus racu s avoided steep hillsi des, but was found
on gentle slopes or in valle ys. It was no t nec essari l y associated
with rocky outcroppings, and was Iound also in lava beds. bunch
grass bordercd b y fir Iorcst s, pine-oak forests, and areas of grass,
agave or cact us.
Degree 01 slope incl inat on dclineat ed the habitat of sorne
can rattl esnakes. Crotalus pri cei , C. pusillus, C. inte nurrlius, C.
t ranscersus and e. lepulus oftcn iu habit cd cxt re mely stcep slo pes
whereas C. ici lardi was Iound in inoun tain vallcyx. uxuallv wi th
permuncnt water. Crotalus poltjst ict us and Sistrurus rllCtlS prccrrcd
relutively flat arcas.
Cround cover is ev idcnt ly a significa nt characte ris t ic in the
habitat of sorne Mcxi can ra t tlcsnakc ta xa, aIHI vnri ed Irom light
(e . poly stictu s) to intermedi ate (e . pusi llus, e. triseriatus. S. racus}.
Crotalu s truusccrsus and C. t riset atus (in some urcux) in ha h ite d
re la tivcl y open situ at ions with a heuvv canopy.
D ai ly activity, Daily erne rgence al' \I exiean ra tt les nakes from
re trcats, particularl v montane ta xa , was ass oc iate d wi t l: dillused
sunlight and h igh atmospheric lnu u id it y. hnmediatel v Following
thunderstorm act ivit y, ruttl csu akcs hegan to elne rgc an rl wcre
often encountcrr-d basking 011 rock x which werc st ill moi st. In
gene ral, hi ghland forms t ended to he din rna I and th e lowland
taxa, es pecially dnring summcr 1I1011ths, werc nocturnal . The fol-
10\Vin g Mexican ratt les nak e ta xa were Iound uhroad day: e. b.
basiiiscus, C. h. Oa,HJCIiS, e. cerastes. C. tlurissus tzalrcau, C. i. ini cr-
mcdius, C. i. gloUdi, C. t. omi lten ia nu, C. l. lc pidus, C. l. klauberi ,
e. l. niaculosus, C. /. moru lus, C. n t chc lli niuercusis, C. 111. ni otos-
Sils, e. IJI. nigrcscc us, C. polust iclus, C. J, pricc i, C. p. niiquhuauus,
e. pusillus, e. r. ru[;er, C. s. se/Jt u/atus. C. s. sa/vill, C. stcillcgeri,
C. /ortli gensi s, e. traIl S; CrSUS, e. l. /riserialus, e. t. (Ului/!JS. e.
viridis !ld/cr, e. u;. lVil/ardi , e. le. a11w[;i1is, e. l e. J/l cridiollalis,
e. le. Si/liS, am] Sistrurus ra;ll s. The foIlo\Ving \ Ie xiean rattl esnake
taxa were lOlllld as \\ 'e re pro",lill g by ni ght : C. atro.\' . e.
cerasles, e. d. dllrisslls, C. d. cu/milla/us, C. ti. tololla cus , C. c. eIl UO,
C. e. jurr.;us, e. l. lep idus, C. 1Il . lIl ilCfl clli, C. lll . PUrrlllls, C. 111.
2 SPECIAL OF NATURAL HISTORY
mol ossus, e. polij stictus, e. price i miquihuanus, e. r. ruber, e. r.
lucasensis, e. s. scut ulat us, e. tigris, and C. tri scriatus aquilus.
Adult snakes of three t axa (C. iute rtne dius omiltc ma nus, e. l.
lepidus, e. t. tri scriatus} remained in t he sanie location for a period
of ovc r two weeks and utilized the same rock for basking dail y.
Ju venil es of th ese tax a were cncount ered prowling by night, per-
haps to avoid predation while dispersing.
During the rainy season, pairs of e, interniedius omlltc manus
wer e euco untc rcd in the field.
Seasonal Oeeurrenee. The onset of the rain y season in Mexco
stimulates rat tlesnake scasonal activity, For example, 55 speci-
men s of Crotalus pohjst ictus from sou t hweste rn Jali sco wer e ob-
se rve d during a three day period in th e rain)' season, whereas only
three snakes were secn during a comparab le period in May b efore
the beginll ing of th e rai ns. Seve ntce n speci me ns of e. lepidus
niorulus were Iouncl during ou e da)' after th e first major rain near
Caleana in the Sierra Madre Orienta l; only three iudividuals were
seen during June. The seasoual activi ty of most other Mexi can rattle-
sna ke taxa corr esponded to the patt ern no te d aboye. Except ons
included e, l. lepidus and e. l. kl au beri which we re found th rou gh-
out th e year if th e t em perature was warm. These two ta xa have
aclapted to a more arid environment,
Faunal and Climat ic Assembl age s. Kendei gh (1!).'54) an d Duell-
man (196.5) revi ewed metho ds used in int erpret ing Iaunal assem-
blages and di scus sed the limitations inherent in th ese concepts. The
Life -Zone Concept of Ierr iam (1890, 1894) was envsione d as a
la ti tudinal transcontinen tal li elt or vertical zonation belt in mo un-
tain ous arcas whch incorporated fa unal an d flora l informat ion
with additional climatic data, particularly temperature. The Lfe-
Zon e Co ncept was used by Go ldman (1951) and appl ied to th e
zoogeog raphic charac t eris tics of Mcxico in order to evaluare the
dis tri b ution of birds, mammals, and the fl ora communi ty. Stuart
(196--1 ) fclt that Coldman's summary tended to reflect th e Bioti c
Pr ovince Concepto The Bioti c Province (Ves ta l 1914; Dice 1943,
19.52) was defined by Duell man (196.5) as ".. . a consi derable and
cont inuous geographic arca th at is characterized by th e occurre nce
of one 01' more important eco log ica l associations th at di fler, at least
in prop ortional ar ca covcred, fro m th e associations of adjacent
provinces." Smith (1939), ut ilizing th e lizard genus Scc lopotus, at-
temptcd to map biotic provinces in Mexico. Later (1940, 1949) he
modified h s position con cerning the ap plica ton of the biot ic prov-
inc e concept (see also Pet ers 19.5.5), Smith (1960) no te d, ".. . ex-
tensive ag ree me nt of range limits of numerous species is a ncces-
sary assllmption in acceptance of th e \'al idity of th e biotic province
concept." It is llnlikel y that an)' t\Vo pl ant 0 1' an imal species are
affec ted idenlica lly by al! en\'ronmental compo nents; conseque ntly,
THE NATURAL HISTORY OF RATTLES01AKES 73
no two spcci es or subspecies can be expec ted to have identieal
ranges.
13earing th es e dificulties in mind, we attempted to delineate
th e pattcr ns of di stribution of 42 t axa of Me xican rattl esnak es by
co nsult ing two fa unal an d two clirnati c assemblages: Life-Zone
Conccpt of Goldman (1951). Biot ic Province Concept (Stuart 1964),
Temperature and Ra infall Pr ov inces (Page 1930). an d th e Koeppen
climat ic classi ficat on sys tc m. The last syste m, out line d b y Koeppen
(1900, 1948), Koeppen and Geiger (1930-39) and summarized by
Viv (1964), rou ghl y delineates the climatic types and ut iliz cs data
from annual and monthl y avc rages of ternpe rature and precipita-
tion, inc1uding th e variable cl ec t upon plant grow th through
evaporat ion.
Frequency of occurrence for di strib ut ional overl ap is presented
in Tabl c 1. Two ta xa (C. atrox, C. 1/I 010 SSll S nigresccns} occurred
in six Koeppen provinccs. Crotalus atrox and C. d. tlurissus in-
habited six and five biotic provinces, respectively . Four Life-Zone
Provinces were applicahl e to C. t . triscr a tus. Crotalus 1/l 01 0 SS11S
ni gresccn s and C. s. scut ulai us occurrcd in five ra infall provinccs,
and C. airox inhahited ivc tcmperaturc provinces.
The validity of physiographic, climati c or bioti c pr ovinces, at
least when uscd to explain th c d istrihut ion of rattl esnak cs in lerrain
as compl ex ph ysiographi call y as Mexico , is highl y t cnuou s, The
dis crcpancies Failed t o produce an cxpla nat ion of dist ribu tion that
ca n be considered to be either va lid or useful. Hence, an analysis
of the cco log ica l requiremc nts of rattl esnakes mus t be concerne d
primaril y with th e microhabi tat or hab ita r nic ho (Dammann 1961).
Food. Our field expericncc suggcs te d th at th e occurrence of
montan e ratt les na kcs in a gi vcn habitat was re lated to the avai l-
ability of mcmhers of th e lizarc1 genus Sceloporus. The den sity of
rattl esnak e populations was co nsiderahl y greate r when an area
supportc d a large number of li zards. especially species of Sccloporus
which a re largc1y saxicolous . \ Ve sus pect that the Iccdi ng patterns
of th e smalle r montan o rattl csnakcs are corrc la ted wi th th e dium al
TA B!.E 1. Freq ueucy of occ urrcnc e for di st ri buti onal overlap in 42, taxa of
Mexcan rat t lesnakes accord ng t o bioli c or climatic c1assificati ons. Per cc nta ge
foll o\\' ed by mll llbe r of taxa ill pare ntheses.
Numbcr of Bi otie or Climac Prot:irl ces 111
,, ' hiell Ta xa Occur
Classiiicat on Systelll 1 :2 3 4 5 6
Temperatu re ( Page 1930 ) 67(2,8 ) 24( 10)
5( 2, ) 0(0) 5( 2 ) 0(0)
Rainfall ( Page 1930) 50(21) Hl ( 8)
2, 1(9 ) 5(2,) 5(2,) 0(0)
Lif c-Zonc ( ( ;oldman 1951) 74( 31 ) 19( 8)
5( 2,) 2, (1 ) 0(0 ) 0(0 )
Bioti c Provi uce ( St uart 196,' ) 5:2 ( 2, 2, ) 19( 8 )
2,1 (9) 2, (1 ) 2(1) 2(1 )
e li mat e ( Koeppe n ] 900, ] 948 ) 33(14 ) 33(14 )
]4( 6) ] 2,(5) 0(0 ) 7(3)
4 SPECIAL 01' l'\ATURAL HI STORY
activi ty patterns of va rio us species of Scelopo rus. On one occasion
in th e ea rly afte rnoon a t Llano Crande. Dura ngo, a Sceloporus
poinsctti cl nr te d fr om a clurn p of rocks an d diecl a Iew minutes later.
After five mi nu tes , un adult e. p. pri cei emerged , trai led th e lizard
anc1 cous uined it.
Captive e. pr icei , e. int crmctli us, e. lcpidus, e. iriseriatus, and
e. icllardi showe d a mark ed prcf eren ce for lizards ra ther than
rodents. The snakes us ua llv struck the lizards in th e th oracic re-
gion ancl held outo them until movement ceased. \ Vhen roclents
werc st ruck. thcy we re, in most ca ses. relcasccl, Captive juvcni le
examplcs of the larger lowl and rattl csuakc taxa p ref erred appro-
priat e size d lab oratory mice .
Rit ual ized Mule Combato Crotalus atrox , e, intermedtus gloydi,
e. i. omiltc num us, e. lcpulus klaubcri , e. pusil lus, e. t. t risc riatus,
e. t . aquilus, anc] Sist rurus racu s engagccl in rituuliz ecl maje COIll -
bat in cap tiv ity. A dctailecl dcscription has bccn pub lishecl for
e. lc pulu (Carpente r ct al. 1976) an d a comparat ive an al ysis of
th e other taxa \ViII be Iorthcomi ng (rn anus cript in prep. ).
Courtship aud Mat ing Behavior. The Iollowing :\ Iexiean ra tt lc-
su akcs wcre observccl engaging in courtship a nd / or mating be -
havior i n captiv ity : Crot alus at rox , e, cat alinen si s, e. d urissus cul-
niiuat us, e, d . tzabca n, e, durissus ssp.. e. e . CIl Y O, e. intc rme dius
omiltcmanus, e. l. lepi du s. C. l. klaubcri, e. l. morulus, e. 1/1.
muchclt ', e, 11l. mu crtcnsis, e. molossus nigrcscens, e. ]J. pricei,
e. pu sillus, C. t . tr iscr ia us, e. t. aq ui lus , e. IC. ici ilard t, e. w.
arnabilis, e. ru, silus. nnc] Sisi rurus m Cll S.
The general pattcm of courts h ip and copula t ion is summari zc d
in th e foIl o\Ving accou nt , Shcdding b y th e Icm alc often stimula te c1
malo co urts h ip ac t ivity and the Iemal e Irequcntl y dcf ecatcd. pre-
sumably to evacuatc th e cl oaca. Thc male cli rcctecl head-bobbing
ami tongue-Ilicking se quc nccs upon the head and dorsum of th e
Iemale. [f th e f em a le was in a resti ng coi l, th e male might vi gor-
ous ly p ress hi s 10\Ver jaw agai ns t her bod y in order to stimulate
her to cra wl. This action allowcd for alignme nt of th e bodi cs. The
heacl-neck a ng le of th e maj e vari cd bet ween 30-4.5 and the male's
menta l rcgiou was pressecl on the Iemnl e's do rsum, The dorsa l acl-
va nee movements by th e male, either sagi t ta lly, para-sagi t ta ll y or
laterall y, iuclurlecl a head-bobhing Illoti on \Vhich va ri ec1 iute rspc-
eifiea lly fro m 110 lat eral mo\'emellt to a pronouneec1 sicle-to-sic1e
mo tioll . Female e. al rox as sullle d a rai sed head and ante rior tru nk
stanee 011 oceasion: possibl y thi s \\"as a rej ect ion posh lJ"e (manus eript
in prep.). As t he mal e di reete cl courtshi p act ivities to\\"ard th e fe-
maIe. sIl(' raised her tail ancl ga wd tb e cl oaca (as in e. alrox, e, rl .
tlurissll S, e. JllsilhI S) \\"hich of teu stim ulate d t he ma le. The male en-
eireIecI tb e female's tail \Vith bis tail a1HI rapid ly slid hi s caudal con-
figuration iJl anteriorly ami post eri or!y c1irected movem cn ts (as in e.
THE NATURAL HISTORY OF 75
ilurissus tzubcan , C. C!. duri ssus, C. c. CIl YO, C. pusillus]. This be-
hnvior has been t ermecl th e "st rokc-cycle" (Chiszar ct al . 1976).
The mal e draped lar gc-radius coi ls ove r th e Iemal c's bocly an d at-
temptecl to place the cloucae in apposit ion by sldiug his t ail under
the tai! of the female. The malc mi ght pu sh backwards agai ns t
the resistance of th e Icmale's tai l as he inscrted (as in C. d . durissus,
C. pusillus). Onl y on e hem ipenis was inserte cl, anel eversin of
th e hernipen is pri or to ins ertion was ne vcr obs erved. The tact ile-
chase anel tactile-ali gmncnt pa t terns va ried fro m 4 minutes in
C. ioillarili to over 3 hours in C. pusi llus. The range of var iat ion
mi ght be clue to fe male recept ivit y. Co itus va ried from :2 hours
in C. airox to 9 hours in C. w. icill onl . j erking aud twitching gen-
erallv cea secl when intromission occurred, but often t he male
mai;tained a steady pul sat ion of the bocly near t he tail (as in C.
atrox, C. pusillus). Thc Fcmal c mi ght drag th e ma le backwards
by the attached hemip enis (observe d in C. atrox, C. d . d urissus
until separution occu rrcd.
Rcproduction. The rainy se ason in Xl cxico usual lv occurs be-
tween May and Oct ober, aud th e dry season betwecn Dccemb cr
and April. In parts of southeru Xl cxico, a short dry season occ urs
during the summer raiuy periodo During lat e [ une ami carlv [ul y,
th e raiuy se ason normall y bcgius in northern Xl cxico. 111 lat e Jul y
and Septcmbcr, th e raiu y scason st arts in cent ral and southern
Mcxico. The northwest arca of Baja Californ ia receives wiuter
rains whereas th e southern scction rcccives sumrncr rains.
In compiling literaturc references and our observations on
parturition in wi lrl Xlexican ratt lcsuakcs. wc acc umulated th e fol -
lowing elata on broods: 11 lit ters were bo rn in [un, 1:2 iu Jul y.
16 in Au gu st , :2 in September , 1 in Oct ob cr , aud 1 in [ anuary.
Compil ed obscrvatious on our capt ive indi vidual s rcvcalcd 5 litters
boru in [une, 4 in [ul y, S in All gn st , .! in Sept ember, and 1 in
October. Data on these ra tt lcsnake taxa are listecl in Table 2. .
Parturition in .\ rexican rattl csnakcs st ud ied to dat e coincides wi th
th e rainy seaso n, possihl y due to a greate r abundancc of potcntial
Iood it eins ava lable at th is time.
'Varnillg Mcchanisms ami Defeu sive Act ions. I n addition to
th e use of tit e ra tt le as a warniug mechanism anel th e striking coil,
a number of other cle fcnsivc hchaviors were observc d by us in
Xlcxican rattl csnak cs. \ Vhcn scized by colle ctillg tongs, munv of
th e smaller montano taxa turned and bit in lieu of assull1ing a
st riking coil. Some , es pec iall>' C. ill/ Cfl Il Cdi ll S, hegan spinning on
a longitudinal axis. Adult C. dllris,\'ll S /;:..a!Jcall rais cd th c antc rior
portion of the body in an aIlllo st \'cr tical COlUlllll with a lateral
benel in th e ncck, al1(l faccd the intruder. Captive jU\'ellil e C. d.
t ;:a{ ca/l flattcncd and ro tated th eir head anel cerdcal region as
much as 90 so that th e do rsal aspect, with the co nt ras ting pattern
76 SPECIAL OF NATURA L HISTORY
on the hcad and the dark paravcrtebral neck st ripes, was vi sible.
Young cap t vc G. tlurissus t ot onacu s in fl ated th eir trunk , exh b ite d
convulsive movements and, on rare occasions, Hippcd over upon
the dorsum when eonfrontcd by a human obs crver. Crotalus poly-
st ict us usu all y thrashed wildly wi th pronounccd lateral undul ati on s
whc n handl cd , this snake some t imos retumcd to a striking coil
when cornered, An adult G. s. sc ut ulat us ancl sorne exa mples of
Sistruru s ravlls rcpcatcd ly Ilattenccl the ir heads and t runks on th e
substrate, then lift cd the tail s ve rt ically and waved th em slowly
in response to the presence of human s. The snakes did not attempt
to rattle. Sorn e of th e smallc r ra t tlcs na kes responded to th e pres-
euce of live mice and rats by launching into a st riking co il, th en
st ruc k th e rod ents with closc d mo uth. Hattl esnakes react ed t o
kin gsnakcs ancl othcr ophiophagous snakcs by forming pronounced
body-u ridges which wcre us cd to strike the predators . Dermal
substances of ophiophagus snakes eli ci ted bocly-brid ging responses
From a number of Mexican rat tl esnake taxa (Weldon an d Burgharclt,
1979 ). Crotalus toriugcnsis, when touched, used vertical body-
bridges 01' lateral Ilexures to stri ke an annoying object, Frequen tly,
th e [orce \Vas grea t enough to knock Ireshly killed rocle nts fro m
the Ieeding Iorceps.
Crotalus ste incgeri verti call y elcvutcd its tail and a portien of
th e pos terior trunk ancl held th cin rigi d ly as it ass umcd a st riki ng
coil. Althou gh Kl auber (19.56) \Vas un aware of an open mo uth
2. Dal a on rat tlesnake ltters wi th viable you ng showing number of
young, length of young, an d weight of young. Numbers in paren theses are
rnea ns.
X lI lII te r T ot al lcngtli Weight
Tax on oi uoung (//l //l ) (g)
C. atrox 6--25( 14.3)
C. durissus ssp. 10
C. d. tzahcan 21 290-350( 316) 18.4-26.8(23.4)
C. e. [u r cu s 7 206--222(213)
C. int crmcd ius omi ltcmauus 5 19-1-212(205) 4.9- 5.4 ( 5.2)
C. l. Lcnidus 6 165-190(177) 5.0- 5.7( 5.2)
C. l. klau hcr! 3--- 5(4) 181-196(191) 6.3--- 7.0( 6.6)
C. l. niorulu s 4
C. //l. mitclt r[l i 1 275 17.5
C. //l . mucrt cnsis 4 143---179( 167) 3.3--- 7.0( 4.9)
C. 711010 .1.\ 11.1 ni grcscens 5 291-316( 304) 25.-1-27.4(26.6)
C. ict l/ S 5-12(S) 155-287(214) 2.4- 1I. 5( 8.9)
C.
1'.
ni eei 3--- 9(5 .3) 152-1S5( 16-1) 2.7- 5.6( 4.1)
C.
1'.
nq u h utuvus
4- 5( 1.5) 130-143( 135) 2.6( 2.6 )
C. nusillus 3- 5( 4) 165-179( 171) 8.0- 6.1( 4.0)
C. l. triscriat l/ S 4- 6(5) 159-178( 168 )
,1. 8- 5. 1( 4.9)
C.
t. aq ulus
2- 7(4.7) 120-192( 16S) 4.9- 8.S( 6.1)
C. ici llardi .\ i/ I/ S 4- 5(4.,'5 ) 146--21S( 190) 4.9- 6.9( 6.0)
Si.llrurll s rac us 3--- 9( 6.3) 160-20S(183) 3.9- 7.4( 5.4)
TIlE NATURAL IlISTOH. Y 01" MEXICAN RATTLESNAKES 77
threat ening posture in ra t tlesnakcs, three C. molossus nigrescens
obse rved by us rep eat cdly cxh bited this beh avior for pcrods as
long as Iivc minutes. On e C. polusti cius also assurned thi s posture.
Crotalus atrox, C. lepidus, C. molossus, C. scutulatus, and Sis-
trurus racus emitte d mu sk upon capture . An adult C. duri ssus di s-
chargcd a stream al' mu sk wh ilc bcing restraincd.
RESUMEN
Se presenta informacin sobre la cco loga e hist oria natural de
4.'5 taxa de cascabel es mejicanos, ge ne ros Crotalus y Sistrurus.
Observacio nes de la fisiografa, geologa, clima, y la vegetacin
de .',Ixico son regi stradas en lo qu e se refier e a los cascabeles de
esa regin. Datos ele obs er vaciones tomad as direct as de poblacion es
na turales y ejemplares capt vos son list ad os, incluyendo los pr edil ec-
ciones de habit acin, extc ncion cs de area y elevacin, rc produc in,
periodos de actividad, y comportamiento.
La mayora de los cascabeles monteses habitan bo sques de
pino y encino, llanos cubierto de hi erba, y des iertos de al tur a.
Generalmente los aflorami entos rocosos son un factor deter minante.
Algunas taxa de ti erra baja se enc uent ran en bosqu es tropi cal es
ri dos con afloramientos rocosos. Refugios especficos son a menudo
util izad os por cascabeles mont eses. \ Iicroh ab it aci ones con or ien-
ta cin hacia el sur son habitadas con mas Ircqucncia por cascabel es
de los altos , scqui do por el sudeste , este. no rte, nor est e, y oes te
en el orden de pref er enci a. Casi todas las taxa mo nteses se
encuentran a 1800 metros o m s de a ltura, Una dis tr ibucin
li mi tad a en elevaciones rel at ivmuente bajas caracterizan la habi -
tacin dc Crotalus st cjnc ger (y talvez C. lannomi; Sistrurus Ta CU S
es un a excepcin ya qu e este se encue ntre en asociaciones variadas.
El grado de inclinacin de la fal da es importante. La den sidad de
la vegetacin va de leve a moderado.
Hayos del sol difusos y alta humidad atmosfr ico con tribuyen
a la ac tivi dad diaria, especialmente en las cu lebras qu e habitan
regiones monta osas. Especi es de tierras alt as tiend en a ser diurnas
y las de ti erras bajas nocturnas, especialme nte en el ve rano. Treinta
dos taxa fueron encontrados activas durante el da y diez y ocho
fueron cazadas de noche. Serpientes adultos (C. intcrin edius, C.
lcpi tlus, C. triseriatus} pueden permanecer en el mis mo lu gar por
lo me nos dos semanas y ut ilizar la misma pe a par a tomar el sol.
Ejemplares juveniles de estas ta xa fueron encontradas de noche.
La activi dad de cascabe les en M xico dep ende de la epoca
llu viosa. Pocas cul ebras se enc ue ntran durant e la epoca seca.
Los pat rones de di st ribucin de cas cabeles mejicanos no se
explican por uso de asambl eas faunal es ni climti cas. La frecuencia
de traslape di stribucional involuc rabe n tanto como seis provi nci as
78 SPECIAL PUBLI CATION-MUSEUM OF NATURAL HISTORY
faunales O climticos para una forma de cascabel. Estas discrepen-
cias sug ieren que los requisitos ecol gicos de los cascabeles ti enen
quc ser primariamente evalua dos co n un anlisi s dc microhabitacin
o ni cho habitacional.
La ocurren cia de las culeb ras montanescs se relaciona, en
parte, a la di sponabilidad de especies dc Sceloporus. Muchas de
las culebras peque as de altura mu estran una decidida preferencia
por la gartijas como comida y la s cascabeles jvenes de tierra baja
prefieren roedores pequeos.
Combate ritua li zado cutre machos fuc obs ervado cn G. atrox ,
G. intermedius gloljdi, G. i, omilte manus, G. lepidus klauberi, G.
pu sillus, G. t. tri seriatus, G. t. aquilus y Sisirurus ravlls.
Los aspectos de cortejo y/ o brama fueron registrados en
di eci nu eve di st int as cascabeles ca ptivos, Un patrn ge neralizado
de compo rta miento fuc apare nte y esta descrito.
Parturaci n en los cascabeles meji canos estudiados hasta la
fecha coincide con la epoca llu viosa. Informaci n reproductiva de
19 ta xa es present ada.
Compo rta miento de adverten cia, adicional al curollamiento,
ataque, y resonami euto, inclu ye lo siguiente : torneando y mor-
di endo, rotando en su axis lon gitudinal , rotacin dc la cabeza y
cuello, elevacin vertical dcl cue rpo anterior, inflacin del cue rpo,
movimi entos convulsos, aplasta miento en el suc Io y undulacin
lente dc la cola, lanzando co n boca ce rrada, amenanza con boca
abier ta, ele vac i n ve rtical r gida de la cola y emision de almizcl e.
APPENDIX
Range or altitu de ext cnsious for Mexican ratt lcsnakc taxa reported
in texto
Cr otalus b , basili scus
Cr ot alus bnsili scu s oaxac us
Crotalus durissus culminat us
Crota lus durissus tot onacus
Crotalus i. int crni edius
Cro talus intermedi us gloljdi
l\lICHOACN: near Dos
Aguas in th e Sierra de
Coalcom n (2225 m)
OAXACA: 35.5 km N\V
Telixtlahuaca
7 km \V
Morelia
20.8-48 km N
Soto la Marina
PUEBLA: near Cacaloapan
OAXACA: near Cerro Machn
(Sierra de Jurcz), Suchixtepec
(Sierra de Mahuatln)
THE NATURAL HISTOHY OF r-.IEXICAN HATTLESNAKES 79
Crotalus int ermedius omiltemauus GUERRERO: 1.6 km S\V Filo
de Caballo, San Vicente, 1 km
N Puerto del Gallo
Crotalus lepidus morulus ;\,UEVO LEN: 24-32 km N\V
Galcana
Crotalus steinegeri SINALOA: Ejido Tebaira
Crotalus triscriatus aquilus HIDALGO: El Chi co (3110 m)
Sistrurus ravus GUERRERO: mountains of
Central Guerr ero
LI TEHATURE CITED
AU. E:"", ~ L J.
1933. Hep ort on a cc llcct on of a mphibians and rep til es From Sonora,
Mexico, wth a descrpti on of a new Iizard . Occas, Papo Mus. Zoo!.
Univ, Mi ch., 259 : 1-15.
AXT ELL, H. \ V. and SAIlATII , M. D.
1963. Crot alus pricci miquiliuanus From the Sierra Mad re of Coahuila,
Mexico, Copei a, 1963 : IGI -16'1.
B A:\ TA, B . H .
1962. The amphbans and reptiles frorn l he State of Agu uscali cntes,
Mexico, in the collcc tions of the Ca liforn ia Aca demy of Scienccs.
Wasmann J. Bio!. , 20 :99- 105.
B AIIIlOUII, C. D.
1973. A bi ogeographi cal hst or y of Chirosto ina ( Piscos : Ath crndae ) :
a spccies lock Irom lit e Mexican Plat ean. Copeia , 197 3:533-556 .
BEAL, C. B.
1918. Hec onnaissance of lhc geology aud oil possbilitcs of Ba ja Ca li-
fornia, Mex co . Ceol. Soco Amcr, Me m., :'\ 0. 31: 1-138 .
BEARD, J. S.
1914. Clmax vcgetation in t ropical Amerca. Ecology, 25: 125- 158.
BOGEIlT, C. xt. amI D EGE:\'II All ll T, w. C.
1961. An addition lo the fauna of th e Unitc d Sta tcs, the Chihuahua
ridgc-nos ed rattlesnake in New Mcxico, Am. Mux. Xovi t .,
2061:1- 15.
BOGEIlT, C. M, and O LlVEII , J. A.
1915. A pre liminary analys s of th e hcrpeto fa u na of Sonora. Bull , Am.
~ I u s . i':at. l li st ., 83 :297-126.
BIIA:\'D, D . D.
1936. Not es lo accomp an y a vcgct ati o map of nort hwest Mexico. Univ.
x.x r. Bul!. Biol, Ser. , 280: 16-22.
1937. Tho natural lund scupe of nort hwest crn Chi huahua. Uni\' . N.!\!.
Bull, Bol, Ser., 3 ](j .
1957. Coastul sludy of sout hwest Mcxico, 2 vols. Dep t. of Ceog., Univ,
Texas.
H IIOW:\' , 'f. \ V.
1971. Autecology of lhe side windc r (Grota/lls cerastes) at Kelso Dunes,
Mojave Desert, Ca lifornia. Di ss. Abst r, 3 I B:6336-6337 .
CAXIl'Il ELL, J. A .
1977. The distrbut ion, vuri ation, ami na tural hs tory of th e Mi ddle
American hi ghl aml pitvipers, Botlirops harh ou ri and Bot lirops
godmaui. Ma ster's t hes is, The Univ, Texas at Ar lington. pp . 152.
D.\\'IS, \V.
1957 .
1959.
80 SPECIAL PUBLICA OF NATURAL HISTORY
1979. A new rattlesnakc ( Rep tilia, Serpentes, Vip eridae ) from Jalisco,
Mexico. Trans. Kansas Acad, Sci., 81 ( 4) :365-369.
C.UlI'BELL, J. A. and Anxrsrnoxc, B. L.
1979. Geographic varation in the Mexican pi gmy rattlesnake, Sistrurus
racus (Reptilia, Serpentes, Viperi dae ) , with a description of a
new subs pe cies. lIerpeto logica, in press.
CARPE:\'TEII, C. C. , GILLIl':GIlAl\f, J . C. and J. B.
1976. The combat ritu al of th e rock ratt lesnake (Crotalus lcpidus).
Copel a, 1976:764-780.
Cr nszxn, D. , SCUIlDER, K., 11. and RAOCLIFFE, C. \ V.
1976. Ob servati ons of court ship behavior in the wes tcrn massasauga
(Stst rurus cate uatus tergeminus). Hcrpe tologica, 32:337-338.
CO:'.' TIIEIIAS Anrxs, A.
1942 . Map a de las provinci as climatologicas de la Republica Mexicana.
Dir. Ceorg., Met er o., Hdro., I nsto. Ceog., Mexi co, D. F., p . 1-27.
CUESTA TEIIIIO:'.' , C.
1930- 31. Los Crotalianos Mexicanos. Su Clasificac in, Ecologa y Dis-
tribucin Geografa. An . Inst. Biol. Mex., 1 (3) : 187-199, 2 (1 ) :
,17-72.
A. E .
1961. Sorne factors affecting th e distribution of sympat ric specos - of
ratt lesna kes (genus Crotalus ) in Ati zona. Ph .D. thesis, Uni v.
Mi ch ., Ann. Arh or , Mch . Lbrary of Congress Card Mlc 61-2743.
B. and DIXO:-; , J . R.
Notes on Icxican snakes (Ophidia). Southwest. ""at., 2 : 19-27.
Sn akes of th e Chilpancingo Region, Mexco, Proc. Biol. Soc o
Wash. , 72: 79-92.
DA\'IS, \V. B. an d H.
1953. Snakes of th e Mexican state of Morelos. Herpetologi ca,8: 133-143.
DICE, L. R.
1943. The bot c provinces of North America. Univ. Michigan Pr css,
Ann Arhor.
1952. Natural communities. Univ. Mchigan Pr css.
DI XO:-;, J. R. , KETCIlERSID, C. A. and LIEB, C. S.
1972. The herpetofauna of Qu crtaro, Mexico, wth remarks 0 11 taxo-
nomic problems. Southwest . Na t., 16 :225- 237.
DI XO:-;, J . R. , SABBATII, xr, and \VORTIII.:'o'GTO:-;, R.
1962. Comments on suakes from cen tra l an d wcst cm Mexco, Herpeto-
logi ca, 18:91-100.
DUELDIA:-;, \V. E.
1961. The amphb ans and reptiles of Michoac n, Mexico. Univ, Kansas
Publ. 1111s. Nat. Hi st. , 15 :1-148.
1965. A bi ogeogr aph c ac count of the herpetofauna of Mehoac n, Mex-
ico. Un iv. Kan sas Publ. Nat. Hist., 15 :627- 709.
1965. Amphlbians and reptiles from the Yucatan Pennsula, M xico.
Univ. Kan sas Publ. Mu s. Nat. Hist., 15: 577-614.
DU:'.'KLE, D. H. and Svrrru, H. M.
1937. Not es on sorne Mexican ophidans. Occas. Pap . Mu s. Zoo!. Un v.
Mi ch ., 363:1-15.
EGGLER, \V. A.
J948. Plant communtics in the vcinity of the volca no El Parcutin,
M xico, aft er two and a half years of erupt on . Ecology, 29 :
415-436.
CE?\TRY, H. S.
1912. A stu dy of th e flor a and vegetation of the valley of th e Ro Ma yo,
Sonora. Ca megie In st. Washington Publ., 527.
GLOYD, JI.
1940.
Cr.ovo, JI.
1912.
GOLIlXIA:\',
19lG.
II AII IJY, 1..
1969.
THE l\'ATURAL HISTORY OF RATTLESNAKES 81
1946a. Si erra Tacuich amona-c- a S na loan pl ant local e. Bul l. Torre)' Bol.
Club, 73: 356-362.
1946b. Notes on th c vcget ati on of Sierra Suro ta to in nortb ern Sinaloa .
Bull. Torre)' Bol. Club , 73 :451-462.
CLan>, H. K.
1936. Tbe subspec ies of Crotalus lepidus. Occas, Pap , Mus, Zool. Uni v.
337: 1-5.
1936. A Mexican subspe cies of GTota /I/S 1IIo/ OSSI/S Bai rd and Girard.
Occas, Pap o Zool. Un v. Mi ch ., 325 :1 -5.
1937. A berpet ological considera tion of [aunal reas of Southern Ari zona.
Bull . Chic. Acad. Sci., 5 ( 5) : 79-136.
1910. The rattlesnakes, gen era SistruTUS und Crotalus. Chic. Aca d. Sci.,
Spec, Publ. , 4:266 pp .
1948. Description of a neg lect ed subs pc ces of rattlesna ke fr om M xico.
xat. I1isl. Msc., 17 : 1-4.
K. and KAUFFELD, C. F.
A ncw rattl esnakc from Bul l. Chic. Acad, Sci ., 6 ( 2): 11-14.
K. an d SXlITII, 11 . :-'1.
Amphbi ans and repti les f rom th e Carmen Ionnt ains, Coahuila .
Bull. Chic . Acad, Sei., 6(13 ) :23 1-235.
GOLDXI A:\', E, A.
1916. Plant rccords of an expe dition to Lower Calif ornia. Contrib. U.S.
l\' at!. 16 :309-371.
1951. Biological invest gations in Mcxico. Smithson . Mise, Collect. \ ' 0 1.
115.
E. A. and R. T.
The bi oti c provin ces of Mexico. J . Mammal. , 26:347 360.
11 1. and t\lcDIAlIXIID, R. w.
The amphbl ans an d reptiles of Sin aloa, M xico. Uni v. Kan sas Publ.
Nat , lIi st. , 18 :39-252.
II AlIT\\'EG, ;'IJ . and OU\' EII, J . A.
1940. A con tr bution to t he herpetology of th e Isthmus of Tehuantepec.
IV. Mise, Pub!. Mu s. Zoo!. Univ, Mi ch ., -l7: 1-31.
11ElI:\'A:\'/lEZ, J .
1923. The tcmper ature of Mcxico. Monthl y Weathcr Hev. , Suppl., 23.
l Io usst.u. C.
1963. A hr eeding pair of wcst ern di amondback rattl esnakes Crotalus
atrox. Bull. Phil . llerpetol. Soc., 11 :65-66.
lIoulI\JIl GE, 1.. H.
19,17. Det enninati on of world plant formati ons fr om simple clirnatic
d ata. Sc ience, 105 ( 227 ) :367-368.
1964. Life zone ecology. Tropi cal Science Center, Sa n [ ose.
II UXI PIlll EY, H. H.
1936. Notes on altitudinal di stribution of rattlesnakcs. Ecology, 17:
328- 329.
JACOIl, J. S. an d ALTE:\'IlACH, J. S.
1977. Sexua l color dimorphism in Crotalus lcnul us klaub cri Cloyd
( Heplilia, Serpcntcs, Vip cridae ). J . Herpet ol. , 11 :81-84.
KAUFFEL/l, C. F.
1913a. Crowth and feeding of uew-born Price's and green rock ra t tle-
snukes. Am. xn.n. xat., 29 : 607-614.
1943b. Fi eld not es on sorne Arizona repti les and amphbaus. Am. Mdl ,
Na t., 29 :342-358.
KEASEY, S.
1969. Sume record s of repti les at the Ari zona-Sonoran Dcsert Muscum.
Int. Zoo. Yearb., 9: 16-17.
82 SPECIAL OF NATURAL HISTORY
KE:'\DEIGH, S. C.
1954. Il ist ory a nd evalua tion of varou s concepts of pl ant and animal
communi ti es in Nor th Amorica . E col ogy , 35: 152-171.
KLAUIlEII, L. ,\1.
1930. Diff er ential eharac teristics of southweste rn ra ttlesna kes allied to
Crot alus at rox. Bul !. Zoo!. Soco San Di ego 6: 1-70.
1931. Grota lus t igris and Grotalus cuyo, two lit tle known ra tt lesn akes of
the Sou thwcst . Trans . San Diego Soco :\at. Ilist. , 6:353-370.
1936. Crotal us mi t clicllii, th e speckled ratt lesnak e. Trans. Sa n Di ego
Soc o i\' at. Il ist. , 8 : 149-184.
19,19a. The re la tiouship of Crotalus rub cr and Crotalus lucascn sis. Tran s,
Sa n Di ego Soco Nat. lIi st. , 11 :57 -60 .
191 9h. Sorne new a nd revved suhspecics of rat tlesnakes. Tran s, San
Di ego Soco Nat. Jl ist ., 11 : 6 1- 116.
19H)c. Thc subs pecies of the ridge-nosed ratt lesna kc, Crotalus iciilardi.
Trans, San Di ego Soco Nat. Jl ist., 11:121-140.
1952. Tuxonomi c st udies of th e ratt lcsnakcs of mainland Xlexi co. Bull .
Zoo!. Soco San Di ego, 2Ci: 1-143.
195Ci. Rattlesnakcs, thcir ha hts, life hi stories, a mi inl1uence on man -
kiud . Un\'. California Pres s, Be rkel ey and Los An gel es.
1972. Rattlcsn akes, their hal iit s, life histories, an d inl1uence on inan-
kind. Second edn. Univ, California Press, Berkel cy and Los
Angeles.
KOEPPEX, W.
1900. Ve rsu cli eine r Klassifika ti on der Klimat , vorzugsweisc nach ihrcn
Beze hungeu ZIIT Pla nzenwelt . Ceog. Zeit. , 6:5 93 -6 11, 657-679.
Leipzig .
1948. Climato loga. M xico.
KOEPPE:'\, \ \'. an d C EIGEn, R. eds.
1930-1939. Il undbuch der Klimatol ogl e. 4 vo ls, Bcr lin.
LEA\' E:,\WOIlTII, \ V. C.
] 9 16. A p reluninary study of th c vcgctuti ou of t he regi n betwecn Cerro
Tan c taro and th e Ro Tepalcatcpec, Michoac n, Mexico. Am,
'\ Iidl. :-\at. , 36: 137-206.
LEOPOLD, A. S.
1950. Vegcration zones of '\ Iexico . Ecology, 31 :507-518.
1972. Wi ld lfe of Mcxico. The gar ue l urds and ma mmals. Unv. Cali-
fornia Press . Berkel ey, Los Angeles , London,
LESUEUII, Il .
1945. Thc ccology of the vegetation of Chi huahua , Mexico, north of
paral lel twenty-e ght. Univ. Texas Publ ., 452 1: 1-92.
LO"' E, C. 11. ed .
1964. The ve rteb rares of Arizona. Univ. Arizona Press, Tucson .
LO"' E, C. 11., [n. ami NOHlUS, K. S.
1954 . Analysi s of the herpetofn una of Baj a California, Mexico. Tra ns.
San Di ego Soco Nat. Ili st. , 12:47-64.
LU:'\DELL, C. L.
1934. Prcl n nary ske tch of lh e phytogecgraphy of the Yucat an penin-
sula. Ca rnegi e In st. Washington Puhl., 436: 257- .321.
1942. Flora of easte rn Tabasco ami adjacen t Mexican areas. St udi es of
Amer. Sp c rmutophyt es-Ll I. Coutrib, Univ, Mch gan Herb. , 8:1-95 .
H, \ V., COPi', J. F. ami BItEEDLO\' E, D. E.
1976. Not e on the her petofau na of westc rn Mexco : new ree ords from
Sinaloa and the T res Mar ias Islands. Los. Ang, Co . Co nt rib.
Sci ., 275: 1-17 .
THE NATURAL HISTORY 0 1" HATTLES:'\AKES 83
D . T.
1908. Bol anical fealmes of North Ameri can dcsert s. Carncgi e I nst ,
Washington Pub l., 99 : 1-1 11.

196-1. Ccohistory and pa leogeography of Middle Ameri ca. p. 3-32. 10::
Il andbook of Mddle American l ndians. Univ. Texas Press, Austin .
B. E.
1975a. :\'otes on a hrood of the Arizona ridge-nosed ra tt lesna ke (Crotalus
icillardi uiiiiardi). Bull. Herp. Soc., 11: G-1-65.
19i5b. A broo d of Arizonu rid ge-noscd ratt lesnakes (Crot alus icillardi
icillard ) bred an d born in cuptivity. Bull. Md. Herp. Soc., ll:
187- 189 .
Mxnrt x, P. S.
1958. A biogeography of reptiles and amphibians in the Cmez Faras
reg ion, Tamaulpas, M xico. Mise. Publ. Mus, Zool. Univ. Mich.,
101 : 1-102 .

1945. Las pinaceas Mexican as. Anales ln st. Biol. ( i\lexico, D. F. ) 16:
1-315.
C. H.
1890. Resul ts of a l .iologlcal survev of the San Fran cisco mounta n
region and desert of the Littlc' Colora do in Arizona. :\orth Amer .
Fau na , 0:0. 3: 1- 136.
189-1. Laws of ternperntu re control of the gcographic di st ribuli on of
terrestrial animals and plan ts, Xu t. Ceog. Mug., 6 :229-238.
A. H. and STEIl Ill :\' S, H. C.
1964. The lives of deserl anlmals in [ oshua Trec Xat onal Monurnent .
Univ. Californi a Pr ess, Bc rke lcy.
\ V. \ V., J. S. ami i\IcCLI :\' TO CK, 11.
1950 . The amphibi ans and reptiles of t he Stockton Platean in nor the rn
Terrell Count y, Texas. Tex. J. Sci., 2 ( -1 ) :5-13-562.
S. A., JII.
1959. Observat ons on amphi hiaus ami rept iles of the Big Bend reg ion
of Texas. Sou th west . Na t., 3 : 28-54.
F.
1912. Es tudios sobre la ve getuci n de M xico-Ll l . Xot as generales sobre
la vegetaci n del S. O. Es tado de Puebla, especialme nte de la zona
de I tzoc n de Matru nora s. Anales ln st. Biol. , ] 3:-117--150.
1952-] 953. La vegetaci n de Chiapas. 2 vols. Tuxtl a Culierrez, Dep t.
Prensa y Turismo.
F. and S IlAIIP, A. J.
1950 . Chnracterist cs of Ihe vcgctut ion in certain temperare reg icns of
easte ru Mexico. Ecology, 31 :3 13-333.
Mooms R. C.
] 976 . Seasona l and dail y act ivity patt ems and thcrmoregul ntion in thc
southwes te rn spr-ckh-d rnt t lesnake (erota/ lis mit cl cll i ),yrrllUs) and
tho Colorado dcsert sidowi nde r (Croi al us cerastes lat cror epcns).
Diss, Abstr . 37B: I 099-1I00.
D. J.
1977. A biogcogr aphlcal an al ysis of the Chihuahua n Desert th rou gh its
herpetofauna. Dr. \\' . junk B. V., Publishers, The ll ague.
C. H.
1939. Hel ations of th e vegetati on and c!imali c types in Nuevo Len,
Mcxico. Am. j\ ut. , 21 : GS7-7:29.
1947. Veget ation and cl na tc of Coahuila, l\lexico. Mad roo, 9:33-57 .
84 SPECIA1. OF NATURAL HISTORY
K. F.
1955. Herpetological collections from Baja California. Hcrpet ologica,
11 : 33-48.
1\EILL, 'V. T . an d ALI.E:-: , E. R.
1959. Additions to the Bri tish Hond ur as herpetofauna list. Herpet ologica,
1.'5: 23,5-240.
1960 , Xot eworthy snakes from British Hondur as. Herpet ologica, 16:
14.'5- 162.
:\ELSO:-: , E. 'V.
1921. Lower Calif orn ia and its na tural resources. 1\at . Aca d. Sci.
Memoirs, 16: 1-194.
PAGE, J. L.
1930. Climat e of Mexi co. Monthly Weath er Hcv, Suppl., 33.
P ET Ens , J. A.
19.55. Use and mi suse of th e bi ot ic provi nce concept. Amer, Na t.,
89:21-28 .
PETZOLD, II. C.
1963. Noti zen zur Fortpflanzungs-hiologi e un d jugcndcntwicklung
zwe ier Crubenott ern ( Se rpentes : Cro tali da e : Crotalus atrox und
Agki st rodou ]l . pisciwrlls) . Bijdr. Derk., 33: 61-69.
PIA:'o: KA, E. R. and H.
1959 . Di strihutional reco rds for ce rta in Mexican and Cua temala n reptiles.
I1erp etologica, 15 :119- 120 .
1937a.
1937b .
1939.
1912 .
POL' GII, H.
1966.
Qui xx. H.
1974 .
R-\DCLl FFE,
197.'5.
H EESE, H.
1971 .
S.-\\'AGI:, J.
1960.
SIIAI1I' , A.
1916.
S11I11-:\'1: , F.
1931.
191-1.
S.\llTII. 11.
1939 ,
1910.
Ecologi cal rclationships of ratt lesnakes in southcast ern Arizo na
wi th not es on other species. Cop ei a, 1966:676-683.
and [oxs s, J. P.
Squeeze box t echnique Ior measuri ng snakes. Herp. Bevew,
.'5 (2 ) :35.
C. ' V. and T. P.
A new subspcc ies of th e red rnt tlesnake, Crotalus rub cr, from
San Lorenzo Sur Island, Baja Calif ornia Nort e, Mexico. Copela,
197.'5:490-493 .
W.
Notes on a small herpet ologcal collection Irom northcastern
Mexico. J. Herpet ol., .'5 :67-69.

Evolut on of a peninsular herpetofauna. Syst , ZooJ., 9 : 184-212.
J.
Informe prelimi n r sobre algunos estudios fitogeogr flcos efectuados
en M xico y Cuatcni ala . Rev. SOl'. Mex. Hist. Na t ., 7 :35-40.
Vegetat ion of the northwe st ern coas t of Mexico. Bul!. Torrey Bot.
Club, 61 :373-380.
The vegetut ion of th e cape regi n of Baja Ca liforn ia . Madroo,
4 :10,5-113.
Lowland vegctu tion of Sinaloa. Bul!. Torrey Bol. Club, 64:605-613.
Ob ser vati ons on the vc ge tation of Chihuahua. Mad ro o, 5: 1-13.
Crassland and rcla ted vcgeta ton in northern Mexico. Madro o,
6 : 180-198.
Rainfal! of nort hern Mexico. Ecology, 25 : 105- 111.

The Mexican and Ce nt ral American lzards of the genus Sccloporus.
Zool. Ser . Fi eld 1\at. Hist. , 26: 1-397.
AII analysis of th e biot ic provinces of Mexico. as indicated by the
di stributi on of th e Iizards of th e genus Sccloporus. Ana l. Esc. Nae .
Cien. Bio\. , 2 :95-110.
1960.
SXIITII, H.
1965 .
THE NATURAL HlSTORY 01" RATTLESNAKES 85
1916. Pr elminary not es a nd spcculat ons on the Tr iseriaius grou p of
rattl esnakes in Xlexco. Univ. Kan sas Sci. Bull. , 31 :75-10l.
1919. Herpet ogen y in and Cuat em al a. Ann. Assoc, Amcr. Ceog.,
39 :2 19- 238.
An evalua tion of th e b ot c province concept. Syst . Zool. , 9:41-44 .
and 1IIGAREDA, C. P.
A range ext en sion of th e lance-head ed rattlesnnke, Grata/lis pO/Y-
st ict us. J. Ohio Herpctol . Soc., 5: 56 .
SXIlTIl . JI. a nd TAYLOII, E . JI .
1950. Type localities of Mexican repti les an d am ph b an s, Un iv. Kansas
Sci. Bull., 33:313-380.
SXIITIl, H. :-- 1., IIOLLA:"D, R. L. and BIIO\\':-:, R. L.
1971. Thc prairie ra tt lcsna kc in Baj a California del Sur. J. Hcrpet ol .,
5 :200 .
Sour. , an d SLOA:", A. J.
196G. Biogeography and d st rb ut on of rep ti les and am ph bans on slands
in the Culf of Califo rnia, Mexico. Trans. San Diego Soc. Nat.
JIi st., 14 :137-156.
STA:"I>LEY, P. C.
1920- 1926. Trces and shruhs of M xico. Contr ib . U.S. Na tl. Her b ., \'01.
23 (5 part s ) :1-172l.
1930 . Flora of Yuca tan , Fi eld Mu s. Nat . Hi st . Publ. Bol. Ser ., 279:
157-492.
STEllIlI:"S, R. C.
1954. Amphibians and rep til es of weste rn No rth Arnerica.
New Yor k.
S'rxvs xs, H. L.
1961. Th e sols of Middle Amcri ca and th cir relation lo ludian peoples
an d cultures. p. 265-3 15. IN: Handbook of Middle American
I ndians. Univ, Texas Prcss, Austin.
STUAIIT, L. C.
1964. Faun a of Middl e Am e rica. p. 3 16-362. 1:\ : JI andbook of Mddle
Ameri can lndans. Univ. Texas Pr ess, Au st in.
TA:":"EI\, W. W., DIXO:" , J. R. and II.\III\1S, H. S., [n,
1972. A new su bs pec es of Crotalus leuidus from we stc rn Mexico . Creat
Basin Xa t. , 32: 16-24.
TAYLOII, E . JI.
193G. :\otes on th e hcrpct ological fa una of th e Mcxica n statc of Sonora.
Ulli\. Kansas Sci . Bull., 24 :475-503.
1914. Two new species of c rota lid snakes Irom Mexi co. Univ. Kansas
Sci . BIIII., 30:47-56.
1952. Third contr ibution to thc herpcto logy of San Lui s Potos. Univ.
Kausas Sci . BIIII., 34:793-8 15 .
T II\' O:-'- , B. \ V. and RADCLlFFE, C. \ V.
1977 . Hcproduct ion in ca p tive Lower Ca lifornia ratt lesnakes, Grata/lis
CI/YO CI/ YO (Cope ) . Il erp. Hcv. , 8 (2 ) :34- 36 .
VA:" D E:"Il UIl r.II , J.
1922 . The repti les of we stern Nort h Amer ica . Oc cas , Pap o Calf, Acad.
Sci. , 1\0. 10, 2 vols: 1-1028.
VA:" D E:"D UII GlI , J. and SLE\'I:", J. R.
1921. A Iist of the aruphibi uns a ud rept iles of the Pen nsu la of Lo we r
California, w th no tes Oll thc spccies in the collcc tion of thc
Academy. Proc. Calif. Acad. Sc i., 4 ser. 11:49-72.
VA:" DE\' E:>:DEII , T . R. an d LO\\'I-:, C. JI. , Jn.
19i7. Amphbians and reptiles of Yep mera, Chihuahua, Mexico. J.
Herpet ol. , 1l :4 1-50 .
86 SPECIAL OF I\'ATURAL HISTORY
VESTAL, A. G.
1914. Int ern al relati ons of tcrrestri al associations. Amer. Nat. , 48:
413-445.
Vrv, J. A.
1964. Weath er and climat e of Mexco and Central Ameri ca. p . 187-215.
IN: Handbook of Mddl e American Ind ians. Univ, Texas Pr ess,
Austin.
WAC:-:ER, P. L.
196-1. Natural vcgctat on of Mlddle Ame rica. p. 216-264. II\': Hand-
book of Middle Ame rican Indians. Univ, Texas Press, Austin.
\VELOO:-:, P. J. and BURCHARDT, G. 111.
1979. The ophiophage defensivo response in crotalinc snakes: extensi n
to new taxa, j . Chcm. Eco!. , 5(1):141-151.
WEST, R. C.
1964. Surface configurat on and associated geology of Mddle Arncri ca.
p. 33-83. Il\': Ilandbook of Middl o American Indi ans. Univ.
Texas Pr ess, Austin.
WI LEY, G. O.
1929. Not es on th e Texas rattl esnakc in captivi ty, wi th special reference
to the bir th of a litter of young. Bul!. Ant iven in Inst. Am. , 3:8-14.
\VmcIIT, A. Il . and \VmeIIT, A. A.
1957. Handbook of snakes of the Un ted Sta tes and Cunada . Comstock
Pub!. Co., Ithaca, N.Y.
INDEX TO HERPETOLOGICAL SCIE NTI FIC NAMES
PACE
Agki strodo n pisci corus _ _ __ 37
Ano lis . __.. . .__ __ 56
Botli rops barbouri .__ __ __ .__ .____________ __ . . . 22
Both rops un dulatu s . .__ .__..__ __ __ ._ .. __ .. . 22
Cucmidop horu s .__ . . __ .. .__ . .__ . .... __ ..... 10, 56
Crotalus __ . .. .. __.. ........ __ __ .__ __ . o 6, 12, 56, 77
at rox o 4, 71, 73, 71 , 75, 76, 77, 78
basiliscus __ . _. __ . __ .. ..__ . . 4, 6, 3 1, 71
luts liisc u hasil iscu s o .____ 6, 7, 8, 43, 71, 78
hasiliscus oaxacus __.. . __ . .. ________ __ __ __ __ __ .. __ 6, 8, 71, 78
catalinr nsis .__ . __ . .__ ........ __ .. 8, 74
cerastes ... __... ._ .... ......__ ...... _.. .__ . . . _.... ... 9, 10, 49, 71
ce rastes ccrcobombus . __ o __ .. .. 9, 10
ce rastes lat crorcpcns .. __ .. . 9
du rissns . 10, 12, 13, 14, 15, 71, 7:>, 7j, 75, 7(j, 77
durissu s cul nnat us _ __ o 7, 11, 12, I -l, 13, 70, 71, 74, 78
d uri ssus cU1lla ncns is .. __ .. 11
tl urissus d uriss us __ __ 10, 11, 14, 71, 73, 71 , 75
durissus t ot ollacu s 10, 12, 13, 11, 71, 7B, 78
d uris sus tzab can .. __ __ .__ 14, 15, 16, 71, 74, 75, 76
cnyo . ._ . __ __ _.. __.. __ . 16
c rujo Cll Y O __ lG, 17, 30, 71, 71 , 75
cnyo [urcus __ ..__ ..__ . 17, 18, 63, 71, 76
int crmcdiu s .. __ __ .. __ 18, 70, 71, 74, , J , 77
int er mrdius gloyd i __. . . __ .... __ . 8, 19, 20, 21, 71, 74, 78
int ermctlus int crmedius __ 18, 19, 70, 71, 78
intcrmcdiu s omi lt cmanus ... . 20, 22, 23, 71, 72, 74, 75, 76, 78, 79
lauuomi .. .. __ __. .... _. . . __ . ... __ .. 71, 77
lcpidus . 22, 26, 29, 33, 39, 43, 59, 61, 63, 64, 70, 71, 74, II
THE NATURAL HISTORY OF 11EXICAN RATTLES NAKES 87
69, 70, 77
77, 78, 79
31
70, 72, 74, 76,
61, 6:2, 63,
10, 39, 63, 64, 65, 66, 67, 70, 71, 74,
______________ ___________ ______ 64, 65, 71,
26, 39, 64, 6.' 5,
17, Ci 6, 67, 68, 71, 74,
63, 64, 65, 66, 71, 74,
PACE
lepidu klauberi 22, 24, 25, 26, 32, 61, 70, 71, 72, 71, 75, 76, 78
lcpidus lcpidu 22, 24, 25, 26, 70, 7 ] , 72, 74, 75, 76
lcpidus maculosus 7, 22, 25, 26, 27, 71
Icpidus motulus 22, 2-1, 25, 26, 27, 28, 29, 70, 71, 72, 74, 76, 79
mclielli 29, 31
mit chclli lIlitchclli _____ 17, 29, 30, 71, 74, 76
mit cliclli tnu crtcn sis 31, 71, 74, 76
mu clicll )lyrrllllS 10, 30, 31, 62, 63, 71
molossus 6, 8, 31, 32, 39, 48, 7 ], 77
molossu s cstc bunensis 32
molossu s lII ol osSIIS 32, 33, 70, 71, 72
niolossu s nigresccus 6, 26, 27, .'3 2, 3.'3, .'31, 69, 71, 73, 74, 76, 77
nolustictus 10, 34, 35, 36, 37, 71, 72, 76, 77
pricei 33, 38, 41, 63, 64, 70, 71, 74
pricei )lricei 26, 27, 38, 39, 40, 42, 64, 70, 71, 74, 76
pricei uiiquihuanus _ 40, 41, 42, 43, 70, 71, 72, 76
pu sillus 7, 43, 44, 45, 46, 70, 71, 74, 75, 76, 78
rubcr 16, 47, 61
rubcr lorenz oen sis 47
ruber lu cascn sis __ _ 8, 16, 30, 47, -18, 71, 72
ruber ruber ___ 47, 48, 62, 63, 71, 72
scut ulat us _ _ 4, ,18, 71, 77
scut ulatus salcini __ 49, 50, 51, 71
sc ut ulat us scut ulatus __ 48. 49, 50, 7L 72, 73, 76
st cj ncgeri 7, 50, 52, 71, 76, 77, 79
ti gris ]6, 50, 52, 53, 71, 72
tortu gen sis 53, 71, 76
transc ersu s 53, 5 1, 55, 56, 70, 71
triscriatus _ 1, 22, 5Ci , 71, 74, 77
tr iscriutus aquil us 56, 58, 59, 60, 61, 71, 72 , 74, 7Ci , 78, 79
tri scriatus triscriat us 43, 53, 55, 56, 57, 58, 59, 61, 69
71, 72, 73, 71, 76, 78
__ ___ _______________________ 61, 62
61
71
75
74
71
76
75
10
10
__ 56
56
37
37
56
56
___ _ 22, 26, 42, 56, 72, 73, 71 , 78
56
_ 63
39, 74
63
ciridts
oiridis caliuiuus
c irul is hcllcri
tcillardi _
icillard anuthilis
icillurdi mcri di onalis _ _ _
willardi s il us
uiillardi icillardi _
Crotapluttu s
Di)l soSallTll s _
Ccrthon ot us _
llyla _
Lanniron clt is t t iaugulu ui arciicra
Liclianu ra t rici rgai a roscojusca _
Ncrodia
Plnjilodacujlu
Sc cloporu s
l1Cn ClI S _
jarrod _
poi nsc tt i _ _
undulat us _ _
SistTIl rus _ _
t c _ _ 51, 57, 68, 69,
Strcplosaurns III Cl1T11 si _ _ _
88 SPECIAL OF NATURAL HI STORY
PACE
Tliamnophis _ . .__.. .__..._.. ._. ... 56
Uma _ 10
Uta __ ---- 10, 56
stan sburiaua .__ 31
Xantusia .... ..._. ...__. . ._. .._._..__. . ._... . . 56
St ates of Mexico
1. Aguascalien tes ].
Morelos
2. Baja California del No rte ]8 .
l\' ayar it
3. Baja Ca lifornia del Sur i o. Xu cvo Leon
4. Campeche 20. Oaxaca
5. Chiap as 21. Pueb la
6. Ch ihu ahua 22. Quer etaro
. Coahuila 23. Quintana Roo
8. Co lima 2-1. San Lui s Pot osi
9. Di st rit o Federal
r -
Sinaloa _ ;J .
10. Durango 26. Sonora
]1.
Cua najuato
r-
Tabasco z t ,
12. Guc rrcro 28. Tarnaul pas
] 3. Hidal go 29. Tlaxcal a
14. Jali sco 30. Ver acruz
]5. l\l exico 31. Yucata n
]6. Mchoacan 32. Zacatecas