Society for the Study of Amphibians and Reptiles

A New Subspecies of the Bushmaster, Lachesis muta, from Southeastern Costa Rica
Author(s): Alejandro Solórzano and Luis Cerdas
Source: Journal of Herpetology, Vol. 20, No. 3 (Sep., 1986), pp. 463-466
Published by: Society for the Study of Amphibians and Reptiles
Stable URL: http://www.jstor.org/stable/1564518 .
Accessed: 26/06/2014 10:58

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

Society for the Study of Amphibians and Reptiles is collaborating with JSTOR to digitize, preserve and extend
access to Journal of Herpetology.

http://www.jstor.org

This content downloaded from 163.178.208.25 on Thu, 26 Jun 2014 10:58:16 AM

All use subject to JSTOR Terms and Conditions
 NOTES 463

Acknowledgments.-M. Gillam provided field as-

sistance and the Conservation Commission of the
Northern Territory, National Parks and Wildlife
Service of Australia and the Australian Customs
Service permitted the collection and export of the
specimen from Australia. H. W. Greene provided
the Locam and K. Schwenk assisted in its opera-
tion. M. H. Wake, K. Schwenk and K. de Queiroz
provided useful comments on the manuscript.
LITERATURE CITED
CLIBURN, J. W. 1957. Leaping ability of the glass
lizard. Herpetologica 13:24.
EHMANN,H. 1980. The natural history and con-
servation of the bronzeback (Ophidiocephalus
taeniatusLucas and Frost) (Lacertilia, Pygopod-
idae). In C. B. Banks and A. A. Martin (eds.),
Proc. Melbourne Herpetol. Symp. Pp. 7-13. FIG.1. Holotype of Lachesismuta melanocephala
Dominion Press, Blackburn.
ssp. nov. (MICP 301).
GANS, C. 1974. Biomechanics. J. B. Lippincott,
Philadelphia. 261 pp.
.1984. Slide-pushing-a transitional lo-
comotor method of elongate squamates. In M.
presence in southern Nicaragua, there is very lit-
W. J. Ferguson (ed.), The Structure, Develop- tle information on this species in Nicaragua.
ment and Evolution of Reptiles. Pp. 13-26.
Currently, Lachesismuta includes three subspe-
Symp. Zool. Soc. Lond. 52. cies: (a) L. m. rhombeata,present in the rainforest
, AND H. MENDELSSOHN. 1972. Sidewind- areas of the Atlantic regions of Brazil, from the
ing and jumping progression of vipers. In A. State of Alagoas to Rio de Janeiro; (b) L. m. muta,
de Vries and E. Kochva (eds.), Toxins of Ani- distributed in the equatorial rainforest of Brazil,
mal and Plant Origin. Vol. 1. Pp. 17-38. Gor-
Guyanas, Venezuela, Trinidad, Peru, Ecuador, Co-
don and Breach, N. Y. lombia and Panama (Martinez and Bolafios, 1982);
JENKINS, R., ANDR. BARTELL.1980. A Field Guide and (c) L. m. stenophrys, present in the tropical
to Reptiles of the Australian High Country. rainforest of Costa Rica and Panama. In Costa Rica,
Inkata Press, Melbourne. 278 pp. this subspecies is widely distributed along the At-
KLUGE, A. G. 1974. A taxonomic revision of the lantic slopes, in the northern plains of San Carlos,
lizard family Pygopodidae. Misc. Pub. Mus.
Alajuela Province, and a peculiar population
Zool. Univ. Mich. 147:1-221.
ranges from the region of Puerto Cortes, to Pen-
SWANSON, S. 1976. Lizards of Australia. Angus insula de Osa, Valle de Coto Brus and Valle de
and Robertson, Sydney. 80 pp. Coto Colorado. In Costa Rica, the central moun-
tain range, with elevations of 2000-3000 m, con-
Accepted: 6 November 1985. stitutes a strong geographic barrier that separates
Atlantic and Pacific populations of L. muta as well
as other snakes (Taylor et al., 1974). In the ser-
pentarium at the Instituto Clodomiro Picado we
observed that specimens of L. muta from the Pa-
Journalof Herpetology, Vol. 20, No. 3, pp. 463-466, 1986
Copyright 1986 Society for the Study of Amphibians and Reptiles cific region of Costa Rica display several features
that clearly distinguish them from the typical pat-
tern of L. m. stenophrys. Previously, Bolafios et al.
A New Subspecies of the Bushmaster, (1978) described differences between these two
populations in immunological characteristics of
Lachesis muta, from their venom. Recent morphological studies of the
Southeastern Costa Rica southeastern populations of L. muta from Costa
Rica prompt us to conclude that it represents a
ALEJANDRO SOLORZANO AND LUIS CERDAS, Instituto new subspecies. In this work we describe this
ClodomiroPicado, Facultad de Microbiologia,Univer-
sidad de Costa Rica, San Jose, Costa Rica. subspecies and compared it to other subspecies of
L. muta, especially to L. m. stenophrys.

Lachesis muta (Viperidae: Crotalinae) is distrib- Lachesismuta melanocephalassp. nov.

uted from the southern tropical regions of Brazil
to the Atlantic and Pacific slopes of Costa Rica
(Vial and Jimenez-Porras, 1967; Peters and Orejas-
Miranda, 1970; Hoge and Romano-Hoge, 1981;
Bolafios, 1982). Although Villa (1962) reported its
Holotype.-Museo Instituto Clodomiro Picado
(MICP) 301; an adult male (Fig. 1), 1969 mm snout-
vent length, 2150 mm total length, collected in
tropical rainforest 9 km northern of Ciudad Neily
in southeastern Provincia de Puntarenas, Costa

This content downloaded from 163.178.208.25 on Thu, 26 Jun 2014 10:58:16 AM

All use subject to JSTOR Terms and Conditions
464 NOTES

TABLE 1. Statistics of variation in ventral scales and midbody dorsal scale rows in Lachesis muta
melanocephalaand L. m. stenophrys.

Ventrals Dorsalscale rows

N x + 1 SD Range x + 1 SD Range
L. m. melanocephala
Males 4 213.7 ? 2.63 211-216 37 ? 1.15 36-38
Females 4 213.7 ? 3.3 209-216 39 ? 1.15 36-40
Mean 8 213.7 ? 2.76 209-216 38 + 1.51 36-40
L. m. stenophrys
Males 25 201.9 ? 1.74 198-204 34.6 ? 0.86 33-36
Females 29 204.5 ? 2.47 199-209 35.3 ? 1.03 33-38
Mean 54 203.3 ? 2.49 198-209 35.0 + 1.00 33-38

Rica, on 31 May 1984 by A. Sol6rzano and Luis
Cerdas.
Paratypes. -MICP-302 from Palmar Norte,
MICP-303 from Corcovado National Park in Pen-
insula de Osa; MICP-304 from Golfito, MICP-305
from Palmar Sur, MICP-306 from Sabalito de Coto
Brus, MICP-307 from Alturas del Rio Cot6n, east
from Valle de Coto Brus. All are located in the
province of Puntarenas.
Diagnosis.-The main features that distinguish
this subspecies are the color pattern and the high
number of ventral scales. It can be differentiated
from L. m. stenophrysin that it has a completely
black dorsocephalic region and a deep yellow-col-
ored dorsum. This contrasts with the pale gray-
ish-yellow coloration that characterizes speci-
mens of L. m. stenophrys. Moreover, L. m.
melanocephaladoes not have the postocular band
that typifies L. m. stenophrys,since the black head
cap extends to the postocular region. Neonates of
L. m. melanocephalaare yellowish, having a dark
dorsal pattern and a black dorsocephalic region.
In contrast with adult specimens, neonates have
light, narrow postocular bands in the black head
cap. In contrast, neonates of L. m. stenophrysare
orange, including their head, and have a rhom-
boid pattern on the body and a black postocular
band. We have not found specimens with a mor-
phological pattern intermediate between these two
subspecies.
Description of Holotype.-The specimen is an
adult male (MICP-301), 1969 mm snout-vent
length, 2150 mm total length, and 181 mm tail
length. Dorsally, it is dark yellow with black
rhomdoidal markings. The dorsocephalic region
is totally black down to the postocular area. There
is no distinct temporal band. Ventrally it is light
yellow. Scale counts are: ventrals 216; caudals 44;
rows at midbody 36; supralabials (left, right) 9-8;
infralabials 12-12; scales between supraoculars 10;
scales between infraocular and supralabials 4;
postocular 2 and preoculars 2. The hemipenis is
bifurcate, 47 mm in length, with a basal spinous
region 21 mm in length.
Scale Counts.-The mean number of ventral
scales in L. m. melanocephalais 213.7 (range: 209-

A B

1 cm FIG.3. Distribution of Lachesis muta in Costa

FIG. 2. Left palatine bones in lateral view; Rica. Black dots correspond to L. m. melanocephala,
Lachesismuta stenophrys(B) and Lachesismuta me- and circles to L. m. stenophrys. Each symbol cor-
lanocephala(A). responds to a single specimen.

This content downloaded from 163.178.208.25 on Thu, 26 Jun 2014 10:58:16 AM

All use subject to JSTOR Terms and Conditions
 NOTES
216; N = 8), and the mean number of dorsal rows
at midbody is 38.0 (36-40; N = 8; Table 1). In com-
parison to L. m. stenophrys, the mean number of
ventral scales is conspicuously lower: 203.3 (198-
209; N = 54), and there is a mean of 35.0 dorsal
rows at midbody (33-38; N = 54; Table 1). Ac-
cording to Roze (1966), L. m. muta from Venezuela
has 213-230 ventral scales and 35-37 dorsal rows,
whereas Peters and Orejas-Miranda (1970) report
more than 214 and 226 ventral scales for males
and females, respectively. Although information
concerning L. m. rhombeatais scarce, Peters and
Orejas-Miranda (1970) describe a range of ventral
scales of 214-226. Therefore, it is evident that the
number of ventral scales in L. m. melanocephalais
intermediate between L. m. stenophrys and L. m.
muta.
Dentition.-Another conspicuous variation be-
tween the subspecies lies in the morphology of
palatine bone, with a concave anterior face (Fig.
2) in L. m. melanocephala.
Karyotype.-The karyotype of a single male
specimen of L. m. melanocephalawas studied using
the method of Gutierrez et al. (1979). A diploid
number of 36 chromosomes was observed, with
16 macro- and 20 microchromosomes. Among
macrochromosomes, pairs 1, 3, 4, and 7 are meta-
centrics, pairs 2 and 5 submetacentrics, and pairs
6 and 8 acrocentrics. This karyotype is very sim-
ilar to the ones of L. m. stenophrys (Gutierrez et
al., 1979) and L. m. rhombeata(Becak and Becak,
1969). In general, karyotypes of crotaline snakes
are highly conservative, all having a similar pat-
tern of 36 chromosomes.
Etymology.-The subspecific designation refers
to the black color of the dorsocephalic region.
Distributionaland EcologicalNotes.-Lachesis muta
seems to be restricted in this distribution to re-
gions below 1000 m, which are classified ecolog-
ically as Tropical Moist and Wet Forest (Hold-
ridge, 1964). Such a distribution may be related
to special requirements for incubation of eggs (Vial
and Jimenez-Porras, 1967). The subspecies L. m.
melanocephala is distributed in the southeastern
region of Costa Rica (Fig. 3). Interestingly, several
specimens of this subspecies have been collected
in Alturas de Cot6n (Puntarenas province) at el-
evations from 1000 to 1500 m, in a Premontane
Wet Forest-Rain Forest transition. There are no
reports of collection of other subspecies of L. muta
in regions with similar ecologic characteristics or
with altitudes above 1000 m.
In Costa Rica, the Talamanca range constitutes
a geographic barrier, with elevations up to 3800
m, which extends to Panama. This barrier pre-
vents intergradation between the two subspecies.
Furthermore, there are no reports of L. m. melan-
ocephala in the Panamanian region of Chiriqui
(Martinez, pers. comm.) nor are there specimens
from other regions with characteristics that would
suggest intergradation between L. m. melanoce-
phala and L. m. stenophrys.
Acknowledgments.-We are grateful to Jose Ma-
This content downloaded from 163.178.208.25 on Thu, 26 Jun 2014 10:58:16 AM
All use subject to JSTOR Terms and Conditions
465
ria Gutierrez and Carlos E. Valerio for their con-
tribution in the preparation of this manuscript,
and to Gino Biamonte and Carlos Jimenez for the
photographic work.
LITERATURE CITED
BECAK, W., ANDM. L. BECAK.1969. Cytotaxono-
my and chromosomal evolution in Serpentes.
Cytogenetics 8:247-262.
BOLAIOS, R. 1982. Las serpientes venenosas de
Centro America y el problema del Ofidismo.
Primera parte: aspectos zool6gicos, epidemio-
logicos y biomedicos. Rev. Cost. Cienc. Med.
3:165-184.
G. MUNoz, AND L. CERDAS. 1978. Toxi-
cidad, neutralizaci6n e inmunoelectroforesis
de los venenos de Lachesis muta de Costa Rica
y Colombia. Toxicon 16:295-300.
GUTItRREZ, J. M., R. TAYLOR,AND R. BOLAiNOS.
1979. Cariotipos de diez especies de ser-
pientes costarricenses de la familia Viperidae.
Rev. Biol. Trop. 27:309-319.
HOGE,A. R., AND S. A. R. W. L. ROMANO-HOGE.
1981. Poisonous snakes of the world. Part I:
Check list of the pit vipers: (Viperoidea, Vi-
peridae, Crotalinae). Mem. Inst. Butantan. 42
+ 43:179-310.
HOLDRIDGE, L. R. 1964. Life zone ecology. Trop.
Sci. Center. San Jose, Costa Rica.
MARTfNEZ, V., ANDR. BOLAFNOS.1982. The bush-
master, Lachesismuta muta (Linnaeus) Ophidia:
Viperidae in Panama. Rev. Biol. Trop. 30:100-
101.
PETERS,J. A., AND B. OREJAS-MIRANDA. 1970. Cat-
alogue of the Neotropical Squamata. Part I.
Snakes. Bull. U.S. Natl. Mus. 297:1-397.
ROZE, J. A. 1966. La taxonomia y zoogeografia
de los ofidios de Venezuela. Imprenta Univ-
ersitaria, Caracas. 362 pp.
TAYLOR, R., R. BOLAI,OS, A. FLORES,ANDG. FLORES.
1974. Geographical distribution of Viperidae,
Elapidae and Hydrophiidae in Costa Rica. Rev.
Biol. Trop. 21:383-397.
VIAL, J. L., AND J. M. JIMENEZ-PORRAS. 1967. The
ecogeography of the bushmaster, Lachesismuta,
in Central America. Amer. Midi. Natur. 78:182-
187.
VILLA, J. 1962. Las Serpientes Venenosas de Nic-
aragua. Editorial Novedades, Managua. 90 pp.
Accepted: 15 November 1985.
KEYTOTHESUBSPECIES OFLachesismuta
(modified from Peters and Orejas-Miranda,
1970)
la. Ventrals 209-230 ...................... 2
lb. Ventrals less than 209; dorsum of head
unicolor light yellow, with occasionally
some dark spots; dorsal ground color
466
light yellow, with black rhomboidal pat-
tern; wide postocular stripe; neonates
orange with dark orange or reddish
markings ................... stenophrys.
2a. Dorsum of head covered with dark spots
..................................... 3
2b. Dorsum of head black to postocular re-
gion; dorsal ground color yellow with
black markings; neonates with thin post-
ocular light yellow stripes within black
cap. Ventrals 209-216 ...... melanocephala.
3a. Spots on head large and distinct; wide
postocular stripe, not bordered by light
stripe above; rostral shield triangular;
ground color bright reddish; color of su-
praoculars contrasts strongly with sur-
rounding black spots .......... rhombeata.
3b. Spots on head small; narrow black post-
ocular stripe bordered with white above;
rostral trapezoidal; ground color gray-
ish; color of supraoculars not strongly
contrasting with surrounding black spots
.................................. muta.
Journalof Herpetology,Vol. 20, No. 3, pp. 466-469, 1986
Copyright 1986 Society for the Study of Amphibians and Reptiles
Selection of Home Range Site by
Females of the Lizard,
Sceloporus jarrovi
DOUGLASE. RUBY, Departmentof Bioscienceand Bio-
technology,Drexel University, Philadelphia,Pennsyl-
vania 19104, USA.
The proximate factors that determine the spa-
tial distribution of individuals in a heterogeneous
habitat are not well known in reptiles (Heatwole,
1977). Although habitat selection may have an
evolutionary impact in terms of foraging behav-
ior (Moermond, 1979a), intraspecific and inter-
specific competition (Schoener, 1977), mating sys-
tems (Stamps, 1977, 1983a), or biogeographic
constraints, behavioral aspects of habitat selection
have received little attention (Moermond, 1979b).
During a study on the territorial and mating
behavior of the lizard Sceloporusjarrovi between
1972 and 1974 (Ruby, 1976, 1977, 1978, 1981), dif-
ferences between the sexes in patterns of site te-
nacity became evident. A fall mating cycle in this
species produces a long period of territorial be-
havior (May through August) prior to mating
(Ruby, 1978). Males adjust home range centers
prior to breeding activity to maximize their over-
lap with females. However, female S. jarrovi es-
tablish a relatively stationary home range in the
spring; home range centers of adult females move
very little between or within years (Ruby, 1978).
Females apparently select a site no more than once
a year and perhaps only twice during a lifetime
This content downloaded from 163.178.208.25 on Thu, 26 Jun 2014 10:58:16 AM
All use subject to JSTOR Terms and Conditions
NOTES
(first summer and first spring of life). In this case,
habitat selection and selection of a home range
are indistinguishable (Heatwole, 1977).
Such an important choice raises questions about
the criteria upon which this choice is made. Do
females identify long-term or short-term attri-
butes of areas? Are females responsive to distri-
butions of males in the spring? Is there a rela-
tionship between distribution of ephemeral
resources, such as food, in the spring and at other
times of the year? The lack of female movement
during the year supports the hypothesis that fe-
males respond to some physical or biological pa-
rameters in the spring which correlate with sur-
vival throughout the summer. I investigated five
factors in 1974 that might influence the location
of the home ranges of females: (1) amount of rocky
substrate, (2) the number of crevices suitable for
a lizard to use for predator escape (see Stamps,
1983b), (3) food abundance and consistency, (4)
overlapping home ranges of other lizards of either
sex, and (5) use of the same area in previous years.
The study area was a 0.60 ha site at 1825 m
elevation on the east side of Mt. Graham near
Safford, Arizona (Graham Co.) as described by
Ruby (1976). Thirty nonoverlapping circular plots
(radius ca. 13 m; centers evenly spaced) were
marked on a map of the study area. For each plot,
the percent of rocky substrate was visually esti-
mated and the number of crevices was counted in
the field. The number of adult lizards per plot was
determined 6 times during the year by direct count
in the daily censuses (usually 2 censuses/day)
conducted over 5 day periods.
Food availability was measured by placing a
10 x 10 cm masonite board covered with "Stick-
em Special"? on the ground at the center of each
plot. The traps were set out for a 24 h sampling
period, starting in the early morning, on 6 dates
(2 and 22 June, 10 and 30 July, 19 August, 24 Sep-
tember). Each date was the third day of the five-
day censusing periods mentioned above. The in-
sect sampling period and its corresponding 5-day
census constituted a "trial." All arthropods col-
lected were counted, regardless of size. This pro-
vides a measure of relative arthropod abundance
among traps even though insects vary in size and
caloric value and traps are not as selective as liz-
ards (Simon, 1975; Ballinger and Ballinger, 1979).
Sticky traps also do not capture ants, a major food
item of this species (Ballinger and Ballinger, 1979).
Numbers of arthropods were ranked among plots
for each test date and tested for (1) correlations
between relative arthropod abundance and lizard
density for each trial; (2) for similarity in rank
between trials (i.e., consistency); and (3) for cor-
relations between ranks for specific trials and the
average rank for all trials (i.e., predictability). Since
an ephemeral stream ran through the study area
after sufficiently hard rainfall, the food availabil-
ity in the ten outermost (peripheral) circular areas
were compared with the food availability in the
ten innermost (presumably wetter, central) areas