Society for the Study of Amphibians and Reptiles

The Use of Agar Models to Study Amphibian Thermal Ecology

Author(s): Carlos A. Navas and Cybele Araujo
Reviewed work(s):
Source: Journal of Herpetology, Vol. 34, No. 2 (Jun., 2000), pp. 330-334
Published by: Society for the Study of Amphibians and Reptiles
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330 SHORTER COMMUNICATIONS

cal lizard, Tropidurusmelanopleurus(Sauria: Tropi- durus itambere(Tropiduridae) in southeastern Bra-

duridae). Copeia 1993:969-976. zil. J. Herpetol. 27:347-351.
POUGH, H. 1973. Lizard energetics and diet. Ecology VITT,L. J. 1991. An introduction to the ecology of Cer-
54:837-844. rado lizards. J. Herpetol. 25:79-90.
ROCHA,C. E D. 1989. Diet of a tropical lizard (Liolae- .1993. Ecology of isolated open formation Tro-
mus lutzae) of Southeastern Brazil. J. Herpetol. 23: pidurus (Reptilia: Tropiduridae) in Amazonian
292-294. lowland rain forest. Can. J. Zool. 71:2370-2390.
.1992. Reproductive and fat body cycles of the . 1995. The ecology of tropical lizards in the
tropical sand lizard (Liolaemuslutzae) of southeast- caatinga of northeast Brazil. Occ. Pap. Oklahoma
ern Brazil. J. Herpetol. 26:17-23. Mus. Nat. Hist. 1:1-29.
.1996. Seasonal shift in lizard diet: the season- ,P. A. ZANI,ANDJ. P. CALDWELL. 1996. Behav-
ality in food resources affecting the diet of Liolae- ioral ecology of Tropidurushispiduson isolated rock
mus lutzae (Tropiduridae). Ciencia e Cultura 48: outcrops in Amazonia. J. Trop. Ecol. 12:81-101.
264-269. VRCIBRADIC, D., AND C. E D. ROCHA.1995. Varia;ao
, ANDH. G. BERGALLO. 1994. Tropidurustorqua- sazonal na dieta de Mabuya macrorhyncha(Sauria,
tus (collared lizard): Diet. Herpetol. Rev. 25:69. Scincdae) na restinga de Barra de Marica, RJ. In:
RODRIGUES,M. T. 1987. Sistematica, ecologia e zo- F A. Esteves (ed.), Oecologia Brasiliensis, vol. 1:
ogeografia dos Tropidurusdo grupo torquatus ao Estrutura, Funcionamento e Manejo de Ecossiste-
sul do Rio Amazonas (Sauria, Iguanidae). Arq. mas Brasileiros, pp. 143-153, Instituto de Biologia
Zool. (Sao Paulo). 31:205-230. de UFRJ,Rio de Janeiro.
1988. Distribution of lizards of the genus Tro-
pidurus in Brazil (Sauria, Iguanidae). In P. E. Van- Accepted: 25 January 2000.
zolini: and W. R. Heyer (eds.), Proceedings of a
Workshop on Neotropical Distribution, pp. 413-
425. Academia Brasileira de Ciencias, Rio de Ja-
neiro.
Journal Vol.34, No. 2, pp. 330-334,2000
of Herpetology,
SAEZ, E., AND A. TRAVESET. 1995. Fruit and nectar Copyright 2000Societyfor the Studyof Amphibiansand Reptiles
feeding by Podarcislilfordi (Lacertidae) on Cabrera
Archipelago (Balearic Islands). Herpetol. Rev. 26:
121-123. The Use of Agar Models to Study
SCHALL,J. J., AND S. RESSEL.1991. Toxic plant com-
pounds and the diet of the predominantly herbiv- Amphibian Thermal Ecology
orous lizard Cnemidophorusarubensis.Copeia 1991: CARLOS A. NAVAS*ANDCYBELE ARAUJODepartamento
111-119. de Fisiologia;Instituto de Biociencias;Unizvrsidadede Sao
SCHOENER,T. W. 1967. The ecological significance of Paulo;Rua do Matao TR 14, No. 321 CEP 05508-900 -
sexual dimorfism in size in the lizard Anolis con- Sao Paulo,SP, Brazil E-mail:navas@usp.br
spersus. Science 155:474-477.
SIEGEL,S. 1956. Nonparametric Statistics for the Be-
havioral Sciences. McGraw-Hill, New York.312 pp. Understanding the thermal ecology of ectothermic
vertebrates is fundamental to studies in areas as di-
SILVA, J. G. DA, AND G. V. SOMNER. 1984. A vegetacao
verse as physiology, ecology, evolution, and conser-
de restinga na Barra de Marica, RJ. In L. D. Lac-
vation (Huey, 1982; Hutchison and Dupre, 1992). Care-
erda, D. S. D. Araujo, R. Cerqueira, and B. Turcq ful
study of thermal relationships of ectotherms al-
(eds.), Restingas: Origem, Estrutura, Processos, pp. lows one to assess:
217-225. Anais do Simp6sio Sobre Restingas Bras- (1) the extent to which they take
advantage of the thermal environment; (2) the range
ileiras, Niter6i, RJ. of body temperatures at which individuals are active;
SOKOL,O. M. 1967. Herbivory in lizards. Evolution 21:
192-194. (3) the implications of microhabitat selection on body
H. 1962. Some remarks on herbivorous liz- temperatures; and (4) the effects of habitat disturbance
SZARSKI, on the thermal ecology of animals. A variety of meth-
ards. Evolution 16:529. ods have been proposed to investigate relationships
TEIXEIRA, R. L., ANDM. GIOVANELLI. 1999. Ecologia de between environmental and body temperatures (Bak-
Tropidurustorquatus(Sauria: Tropiduridae) da res-
ken, 1992). For example, comparisons between opera-
tinga de Guriri, Sao Mateus, ES. Rev. Brasil. Biol. tional temperatures (OT, the temperature that animals
59:11-18. would have if stationary in a given microhabitat, as
TEIXEIRA-FILHO, P., C. E D. ROCHA,ANDS. RIBAS.1996. measured with
termal e uso do habitat tor- copper models) and body tempera-
Ecologia por Tropidurus tures (BT), have made a significant contribution to the
quatus (Sauria: Tropiduridae) em uma area de res- understanding of lizard thermal relationships (Hertz,
tinga do sudeste do Brasil. In J. E. Pefaur (ed.), 1992; Hertz et al., 1993). However, methods developed
Herpetologia Neotropical, pp 255-267, Actas del II to study thermal relationships of lizards are not read-
Congreso Latinoamericano de Herpetologia, II Vol- ily applied to amphibians.
umen, Consejo de Publicaciones, Universidad de Although classical studies of anuran thermoregu-
Los Andes, Merida, Venezuela. lation concentrate on rather large species (Lillywhite,
L.
VALLEJO, R., AND M. S. VALLEJO. 1981. Contribuicao 1970; Valdivieso and Tamsitt, 1974; Pearson and Brad-
ao estudo dos microartr6podos do "litter" na res- ford, 1976; Carey, 1978; Brattstrom, 1979;
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tinga de Barra de Marica, RJ. Rev. Brasil. Biol. 41:
535-543.
VAN SLUYS, M. 1993. Food habits of the lizard Tropi- 'To whom correspondence should be sent.
 SHORTER COMMUNICATIONS 331

1.5

(0 3.6 g
1.0
C

8.6 g
0.5
Q.
E
0)
CO
0.0
5 10 15 20 25 30
Manipulation Time (sec)
FIG.1. Temperatureincreasein agar frog replicas(N = 10) aftermanipulation.The models (two sizes) were
held in one hand during a given time lapse to simulatethe expected change in body temperatureexperienced
by small anuransunder similarcircumstances.Thermocoupleswere insertedin the centerof the models, which
had initial temperaturesranging from 22 to 23 C. Regressioncurves (10-30 sec) are shown. Smallermodels
would be expected to have steeper curves.

Williams, 1991), the small size of many amphibians
makes it difficultand time consumingto obtaindirect
measures of field body temperatures.The probes of
most quick-readingfield thermometersare more than
a millimeterin diameter,thus inappropriateto mea-
sure cloacal body temperaturesin small amphibians.
An alternative,cloacalinsertionof thermocouplewire,
requires manipulationof the animal, and for small
amphibianseven a few seconds delay may have con-
siderable effect on body temperature(Fig. 1). The
measure of surface body temperatureswith either
thermistors(Navas, 1996) or infrared probes (which
do not requiremanipulationof animals)may offerthe
best non-invasive method. However accurate IR
probes are expensive and require placing the sensor
close to the animal, which can be difficult.
One alternativeto the direct measureof amphibian
body temperatureis to estimate OTs using suitable
models (artificialanimal replicas).If OTs match BTs
in amphibians,models may be used to estimate ani-
mal field temperatures.The commonly used copper
models, however, may be inappropriatebecause an-
urans have permeableskin and, from a thermalstand-
point, behave as water saturated bodies. Although
agar replicas do not experiencemovement or blood
circulation,they have been used successfully as null
models (total permeability)in the study of amphibian
water relationships(Spotila and Berman, 1976; Wy-
goda, 1984), and it seems reasonableto use them to
estimate OTs as well. Here we discuss the validity of
using agar models connectedto temperaturedatalog-
gers to estimateBT in small amphibians.We used hy-
lid frogs as an example, and tested the assumptions
that, under similar conditions,agar models and frogs
would: (1) have similar rates of water evaporation(2)
show comparablegradientsof body temperature;and
(3) exhibit similar temperaturesin the field.
This study was carriedout from Januaryto Juneof
1998 using several species of treefrogs in the genus
Scinax (Hylidae). Scinax fuscovarius (BM + SE = 8.37
+ 1.02 g) and S. g. ruber(5.02 + 0.63 g) were used for
most lab experiments.In the field we used two more
species, Scinaxsp. (about2 g) and S. hiemalis(0.74 +
0.09 g). All four species are common in southeastern
Brazilbut occur at differentelevationsand are active
at differing time of year. Scinaxfuscovariusand S. g.
ruberwere captured at ParqueAnhanguera,about 60
km NW of Sao Paulo(SP)in the vegetationsurround-
332 SHORTER COMMUNICATIONS
ing small to medium sized ponds. This type of habitat
is similar to that of Scinax sp., a species found at Cam-
pos do Jordao, 1800 m, about 280 km NE of SP. Scinax
hiemalis, a winter specialist, was found in Atibaia
(1200 m), about 62 km North of SP, in forested areas
with slow running streams.
Agar models were made in two sizes (3.187 + 0.179
g and 8.735 ? 0.216 g) from alginate molds made
from frozen individuals of S. g. ruber and S. fuscovar-
ius. Smaller models can be made but are more fragile.
To make models we anesthetized and killed frogs by
prolonged immersion in urethane 30%, and froze
them (-80 C) in a typical posture. We made the al-
ginate molds by immersing the frozen frogs into fluid
alginate, which had been previously poured into a
small plastic container. After hardening, alginate
molds were filled with liquid agar (lOg/L). To avoid
boring into agar and damaging the models, we placed
temperature sensors in the center of the model before
the agar solidified.
To compare the rates of water loss between models
and frogs we used 10 individuals (five S. r. ruberand
five S. fuscovarius) and 10 agar models, five of each
size. We placed each frog or model inside a perforated
cylindrical plastic container (4 cm tall X 2 cm diam-
eter). Each container was then fitted into an airtight
Plexiglas chamber (9 cm tall X 6 cm diameter) con-
taining 10 g of dry silica gel. Relative humidity equil-
ibrated at about 56%, as monitored with a Stowaway
RH data logger located inside the chamber. The cham-
ber was opened every 15 min and the inner container
weighed to the nearest 0.001 g and returned to the
chamber. The experiment continued until the model
or the frog lost 10% of its initial mass. We did not
detect among experiments differences in relative hu-
midity or temperature (ANOVA: RH, F,,,8 = 0.724, P
= 0.406; Temperature: Fl1,, = 3.977, P = 0.061).
A test of thermal gradients was performed with five
small and five large agar models. Each model was
fitted with 5 thermocouples (0.1 mm diameter) as fol-
lows: 1 mm under the dorsal surface, 1 mm above the
ventral surface, 1 mm inside the model at the place
corresponding to the cloaca in frogs, 1 mm inside the
model "mouth", and in the center of the model. These
models were placed outdoors and their temperatures
recorded every 10 min over two hours, a period that
was large enough to involve significant air tempera-
ture fluctuations (14.8 to 20.6 C). A second set of ex-
periments compared the temperature increase of ani-
mals and small models under brief exposures to a
heating source (150 W incandescent light bulb placed
50 cm above). We placed three thermocouples at the
anterior, posterior, and dorsal positions of the models,
as described before. We anesthesized frogs by immer-
sion in a solution of urethane 30% for 20 min, and
then placed three thermocouples, one in the mouth, a
second in the cloaca, and the third under the dorsal
skin at the central part of the back. Temperatures were
measured at room conditions and after heating epi-
sodes of varying lengths, namely 1, 5, 10, and 15 min,
in that order. Models and frogs were allowed to attain
room temperature between measurements.
The field test compared the temperatures obtained
from agar models connected to HOBO data loggers
connected to high accuracy temperature sensors (On-
set Computer Corporation, accuracy +0.4 C and res-
olution +0.2 C at 20 C), and those obtained from di-
rect measurements of frogs. The more appropriate
model size was used for each species. We placed at
least five agar models in typical microhabitats and
programmed the data-loggers to take one reading ev-
ery 15 min. To characterize microhabitats, animals
from each species were located by visual inspection
between 1800 and 6000 h. Some individuals were fol-
lowed after sunrise to identify daytime retreat sites.
For each frog observed we took note of time of day,
distance to the closest water body, substrate type, and
height above floor level. The typical microhabitats for
each species were: (1) S. fuscovarius N = 38, litter and
soil, perching height (h+) = 9.2 + 3.0 cm; (2) S. g.
ruber,N = 42, twigs and leaves, h+ = 52.3 + 8.2 cm;
(3) S. hiemalis,N = 70, leaves and twigs, h+ = 15.9 +
1.8 cm; and 4) Scinax sp., Campos do Jordao, N = 32;
leaves, h+ = 89.4 + 10.5 cm.
After placing the agar models in the field, we
looked for visible animals to obtain data on field body
temperatures. We performed most measures of surface
body temperature using a Raynger ST2 infrared ther-
mometer applied at less than 1 cm from the animal.
A sample of the two larger species (S. g. ruber and S.
fuscovarius) was measured with a Barnardt 115 ther-
mocouple thermometer quickly applied to the lateral
part of the body of the animal (for details see Navas,
1996). Despite the large number of animals detected
in the field (N > 200), reliable data on body temper-
ature were obtained for only 87 frogs. These numbers
illustrate the difficulties of using a direct measure ap-
proach to study the thermal ecology of small amphib-
ians (many animals escaped or could not be measured
within 10 sec).
The agar models produced a temperature data set
that was much larger than that obtained for real ani-
mals; therefore we paired each datum of frog body
temperature with a temperature reading taken from
an agar model. To pair data we chose the agar model
that was closest to a given measured frog, and select-
ed the temperature reading that best matched the time
of day at which the temperature of that frog had been
measured. This procedure generated a data matrix in
which each frog temperature had a corresponding
temperature obtained from a model. These two sets of
data were compared using correlation and ANOVA.
To test for consistency on temperature data obtained
using different sensors, we compared temperature
readings from thermocouples and HOBO data loggers
with those obtained from high precision mercury
thermometers (used as reference). In a test measuring
water temperature from 0 to 50 C, HOBO loggers re-
corded temperatures up to 5 C different from those of
mercury thermometers. Within 5 and 35 C, however,
temperatures measured with thermocouples, HOBO
data loggers, and thermistors, were statistically indis-
tinguishable (ANOVA, F3,16 = 0.167, P = 0.918; Barlett
test for homogeneity of variances, x2 = 0.084, df = 3,
P = 0.994). The mean temperatures within this range
were (N = 45): mercury thermometer 17.7 - 1.36 C
(Mean + SE); thermocouple: 18.5 + 1.33 C; HOBO1:
18.39 + 1.43 C, and HOBO2:19.0 + 1.42 C.
Both agar models and frogs lost water with time
(F,462
= 9.604, P = 0.02). The rates of water loss, as
indicated from the comparison of slopes, were com-
parable in frogs and models (ANCOVA, F,,41^P <
 SHORTER COMMUNICATIONS 333

25 0*
00
0

a) 20 ?0 / ?
C)
23
0
0)0 0
CD
c, 15 @0 %'.
SO)
0.
e? e
*I*
0W0,-))
-O)
')?)
10 O)~
4J
U. 0 *

I I I I

12 16 20 24

Agar model temperature (?C)

FIG. 2. Bivariate plot and regression line showing the relationship between agar model temperature and
frog temperature in the field. Data of frog temperature were coupled with the temperature readings from the
agar models that were close in time and space to the real frogs measured (see text for explanations).

0.001). Agar models showed small thermal gradients
in field tests. The gradients were caused by dorsal
temperatures only (Scheffe's post-hoc test, P < 0.001),
which were higher than those measured at any other
position of the model (central, ventral, anterior, and
posterior). The differences among dorsal and other
temperatures ranged from 1.2 to 1.5 C, whereas other
positions exhibited a maximum average difference of
only 0.2 C. The effects of artificial radiation on agar
models and immobilized frogs were virtually identi-
cal; no interaction involving type (models vs. frogs),
time of exposure, and thermocouple position was
found to be significant (ANOVA, P > 0.9 for all inter-
action terms). At any given thermocouple position the
maximum difference in temperature increase between
models and frogs was 0.13 C.
The body temperatures of the 87 frogs measured in
the field matched those of agar models. The difference
between the two data sets was undistinguishable from
zero (mean difference = -0.271 + 0.17 C; t = -1.565,
df = 86; P = 0.121); furthermore, data on frog and
model temperatures were highly correlated (Fig. 2; r
= 0.93, F1 85 = 551.9, P < 0.001). For heuristic purposes
we used the agar model method to estimate and com-
pare the activity temperatures of S. fuscovarius (a low
elevation summer specialist) and S. hiemalis (a mid el-
evation winter specialist) during 10 nights of their
peak calling season (February and June respectively).
The estimated temperatures were: S. fuscovarius, N =
698, 22.5 + 0.37 C (Mean + SE), Maximum = 27.1 C,
Minimum = 18.6 C; S. hiemalis N = 243, 12.3 + 0.82
C, Maximum = 22.3 C, Minimum = 7.1 C.
Our data suggest that the overall thermal properties
of frogs and agar models are similar. As a result, frogs
and agar models exhibit comparable temperatures in
the field. We conclude that agar models are appropri-
ate to estimate the ranges of field body temperatures
of small amphibians with "typical" skin permeability.
Departure from the latter assumption (frogs with re-
duced skin permeability because of a waxy secretion)
may cause actual body temperatures to be higher than
those predicted from our models. For such "water
proof" species, copper models may be more appro-
priate. When dealing with species that are active in
humid environments, materials other than agar may
be suitable. For example, in a 48 h test carried out in
the humid and shaded montane-forest environments
used by S. hiemalis we found differences lower than
0.8 C among mean temperatures obtained with plas-
tic, plaster, copper, foam, and agar models positioned
sidewise in the field.
Some cautions with our method are necessary. A
334 SHORTER COMMUNICATIONS
problem may emerge if solar radiation causes thermal
gradients in agar models that are not similar to those
of real animals. This was not the case for the small
hylid frogs used in this study, despite possible differ-
ences in reflectance between models and frogs. It
should be noted, however, that color changes relate to
the thermal biology of some amphibians (Tracy, 1979;
King et al., 1994), so that differences in reflectance be-
tween models and animals may be relevant when
working with some diurnal species. Also, because the
thermal variance is higher on the surface than in the
center of models, for sake of consistency it is advisable
to place all sensors central in the model. Finally, agar
models left in the field should not be allowed to des-
iccate. The rate of water loss will depend on model
size, environmental conditions, and degree of expo-
sure. Our experience suggests that daily replacement
(or rehydration) may be necessary. If two sets of sen-
sors are available for each data logger, a used model
can be easily replaced by a new, fully hydrated agar
model-sensor set.
Acknowledgement.-Financial support to carry out
this research came from a FAPESP research grant to
C.A. Navas (95/9378-6) and a scientific training FA-
PESP fellowship to C. Araujo. We thank H. Lillywhite
and two anonymous reviewers for insightful com-
ments on the manuscript. Permit to collect animals
was granted by IBAMA-Brazil.
LITERATURECITED
BAKKEN, G. S. 1992. Measurement and application of
operative and standard operative temperatures in
ecology. Amer. Zool. 32:194-216.
BRATTSTROM, B. H. 1979. Amphibian temperature reg-
ulation studies in the field and in the laboratory.
Amer. Zool. 19:345-356.
CAREY, C. 1978. Factors affecting body temperature in
toads. Oecologia 35:179-219.
HERTZ, P. E. 1992. Temperature regulation in Puerto
Rican Anolis lizards: a field test using null hypoth-
eses. Ecology 73:1405-1417.
, R. B. HUEY, AND R. D. STEVENSON.1993. Eval-
uating temperature regulation by field-active ec-
totherms: the fallacy of the inappropriate question.
Amer. Natur. 142:796-818.
HUEY, R. B. 1982. Temperature, physiology, and the
ecology of reptiles. In C. Gans, and F H. Pough
(eds.), Biology of the Reptilia. Vol. 12., pp. 25-74.
Academic Press, New York.
HUTCHISON, V. H., AND R. K. DUPRE.1992. Thermo-
regulation. In M. E. Feder, and W. W Burggren
(eds.), Environmental Physiology of the Amphibi-
ans., pp. 206-249. Univ. Chicago Press., Chicago,
Illinois.
KING, R. B., S. HAUFF, AND J. B. PHILLIPS.1994. Phys-
iological color change in the green treefrog: re-
sponses to background brightness and tempera-
ture. Copeia 1994:422-432.
LILLYWHITE,H. B. 1970. Behavioral temperature reg-
ulation in the bullfrog, Rana catesbiena. Copeia
1970:158-168.
NAVAS, C. A. 1996. Implications of microhabitat selec-
tion and patterns of activity on the thermal ecol-
ogy of high elevation neotropical anurans. Oecol-
ogia 108:617-626.
PEARSON, O. P., AND D. F BRADFORD. 1976. Thermo-
regulation of lizards and toads at high altitudes in
Peru. Copeia 1976:155-170.
SPOTILA,J.R., AND E. BERMAN. 1976. Determination of
skin resistance and the role of skin in controlling
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moregulation: discussion. In E. H. Burtt Jr. (eds.),
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Garland Press, New York.
VALDIVIESO,D., AND J. R. TAMSITT.1974. Thermal re-
lationships of the neotropical frog Hyla labialis
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Mus. 26:1-10.
WYGODA, M. L. 1984. Low cutaneous evaporative wa-
ter loss in arboreal frogs. Physiol. Zool. 57:329-337.
,AND A. A. WILLIAMS.1991. Body temperature
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335.
Accepted: 30 January 2000.
Journal Vol.34, No. 2, pp. 334-340,2000
of Herpetology,
Copyright2000Societyfor the Studyof Amphibiansand Reptiles
Prenasal Bones and Snout Morphology
in West Indian Bufonids and the Bufo
granulosus Species Group
JENNIFER B. PRAMUK Division of Herpetology,Natural
History Museum and Biodiversity Research Center,and
Departmentof Ecologyand EvolutionaryBiology,The Uni-
versity of Kansas,Lawrence,Kansas 66045-2454, USA
Anurans of the Bufo peltocephalusspecies group (B.
cataulaciceps,B. empusus, B. fluviaticus, B. fustiger, B.
guentheri,B. gundlachi, B. lemur,B. longinasus, B. pelto-
cephalus, and B. taladai) comprise an assemblage of
toads endemic to the West Indies and range from
western Cuba to the Virgin Islands (Schwartz and
Henderson, 1991; Table 1). The West Indian bufonids
are notable for their unusual snout morphology (Pre-
gill, 1981), their relative scarcity and secretive habits
(Barbour, 1926), and the possible application of their
phylogeny to questions of Caribbean biogeography.
To date, the monophyly of the group remains in ques-
tion and their evolutionary relationships are unre-
solved.
In a description of the cranial morphology of the
West Indian toads, Pregill (1981) cited three putative
cranial synapomorphies as evidence of their mono-
phyly, and resurrected the genus PeltophryneFitzinger,
1843 for the group. Subsequently, immunological dis-
tance data (Hedges, 1996), as well as molecular and
morphological data (Graybeal, 1997) have indicated
that Peltophryneis nested within New World Bufo.
Hedges (1996) synonymized Peltophrynewith Bufo.
Herein, I refer to the West Indian toads as the Bufo
peltocephalusspecies group.
It has been hypothesized that the endemic West In-
dian toads originated from a mainland species occur-