Society for the Study of Amphibians and Reptiles

Breeding Activity of the Neotropical Treefrog Hyla elegans (Anura, Hylidae)

Author(s): Rogério P. Bastos and Célio F. B. Haddad
Source: Journal of Herpetology, Vol. 30, No. 3 (Sep., 1996), pp. 355-360
Published by: Society for the Study of Amphibians and Reptiles
Stable URL: http://www.jstor.org/stable/1565172
Accessed: 06/11/2009 09:03

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at
http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless
you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you
may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at
http://www.jstor.org/action/showPublisher?publisherCode=ssar.

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.

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
 Vol. 30, No. 3, pp. 355-360, 1996
Journalof Herpetology,
Copyright1996Societyfor the Study of Amphibiansand Reptiles

Breeding Activity of the Neotropical Treefrog

Hyla elegans (Anura, Hylidae)
ROGERIOP. BASTOS'AND CELIO F. B. HADDAD2

de BiologiaGeral,ICB,UFG,CaixaPostal131,74001-970Goidnia,Goids,Brasil,and
'Departamento
de Zoologia,UNESP,CaixaPostal199, 13506-900RioClaro,SaoPaulo,Brasil
2Departamento

ABSTRACT. - Breeding of Hyla elegans was monitored from August 1991to July 1992at a temporarypond
in Ubatuba, State of Sao Paulo, southeastern Brazil. Males began to call as they entered the chorus, and
defended their calling sites from other males, at times with physical interactions. Females, however, were
not aggressive toward either males or other females. We found a positive correlation between the numbers
of females and males in the chorus, but no significant correlation between OSR (number of reproducing
females/number of reproducing males) and the number of males present. OSR was highly male-biased;
on average, there were 10 males for each female; this low OSR may explain low average mating success
of males. Females chose males as mates freely, and males did not attempt to intercept females approaching
other males. Males in amplexus were larger and heavier than unmated, calling males. In addition, snout-
vent lengths of males and females in amplexus were positively correlated, and males were, on average,
0.81 the length of females. Experimentally paired males and females with smaller or larger ratios of SVLs
had a lower percent of fertilization than pairs near the population average.

During the breeding season, many anurans
aggregate in certain areas to reproduce (Wells,
1977). At these aggregations, social interactions
are frequent, and reproductive behavior can be
complex (Arak, 1983). In many species, mating
occurs nonrandomly with respect to variation
in male traits (see Cherry, 1992). For example,
males in amplexus can be larger than solitary
ones (e.g., Lee and Crump, 1981; Ryan, 1983),
or there can be positive assortative mating by
size or mass (e.g., Davies and Halliday, 1977;
Robertson, 1986).
The two mechanisms of sexual selection pro-
posed by Darwin (1871), male-male competition
and female choice, are involved in the genesis
of nonrandom mating patterns (Olson et al.,
to Sao Paulo, Brazil (Lutz, 1973; Frost, 1985),
and from sea level to approximately 800 m (Lutz,
1973). The present study describes the breeding
behavior of H. elegans, in southeastern Brazil,
with emphasis on sexual selection.
Our aims were to: (1) quantify the reproduc-
tive parameters of both sexes (temporal distri-
bution, sex ratio, size, amplexus, and clutches),
(2) determine whether matings are non-random
in relation to male size, (3) verify the occurrence
of assortative mating (i.e., if females select males
that are proportional in size to themselves), and
(4) verify if relative sizes of mates influence
fertilization.
MATERIALS AND METHODS

1986). In anurans, the intensity of each mech-
anism of sexual selection depends, in part, on
the duration of breeding period (Wells, 1977;
Ryan, 1985). In anurans with prolonged breed-
ing seasons, females generally arrive at breed-
ing sites asynchronously, having the opportu-
nity to choose males (Arak, 1983) that defend
territories for vocalization and/or oviposition
(Wells, 1977; Haddad and Cardoso, 1992). In
contrast to prolonged breeders, female anurans
that breed explosively have little opportunity
to choose males (Arak, 1983). In such a pattern,
characteristics that confer greater advantage in
male-male competition will be more important
for reproductive success (Wells, 1977; Halliday,
1983).
We conducted a long-term study of the tree-
frog Hyla elegans Wied-Neuwied, a species that
has a wide geographical distribution along the
Atlantic Forest, from the States of Pernambuco
Hyla eleganswas observed in a temporary pond
at Nuicleo Picinguaba (44?50'W and 23?23'S),
Parque Estadual da Serra do Mar, in the mu-
nicipality of Ubatuba, State of Sao Paulo, Brazil,
between August 1991 and July 1992. The area
of the pond was approximately 2600 m2. Due
to irregular rainfall during 1991 and 1992, the
pond dried more than once. We visited the study
pond either fortnightly or monthly, monitoring
the pond for 2-10 nights during each visit. Field
observations were conducted from 1730 h until
2400 to 0200 h, depending on activity of tree-
frogs. We monitored the pond for a total of 278
h in 73 visits.
We marked 330 males and 75 females by toe
clipping (Martof, 1953), representing approxi-
mately 40% of the population at the pond (Bas-
tos, 1993). We measured the snout-vent length
(SVL) of individuals to the nearest 0.1 mm with
a caliper ruler and weighed individuals to the
356 R. P. BASTOS AND C. F. B. HADDAD
TABLE1. Comparisons (t-values) of mean snout-vent length and mean mass of males and females of Hyla
elegans. * Female mass after oviposition.
Males
N Mean + SD Range
SVL (mm) 258 25.74 + 1.14 22.1-29.6
Mass (g) 223 1.17 ? 0.20 0.7-1.8
nearest 0.1 g with a balance. We counted the
total number of males and females in the pond
and recorded the location of these individuals,
territorial encounters, matings, and number of
clutches.
All pairs found in amplexus were collected
and placed in plastic bags containing water from
the pond until females laid eggs. The clutches
obtained were preserved in 5%formalin. Males
and females were weighed, measured, and re-
leased on the following night.
To test the hypothesis that females choose
males that maximize fertilization success, we
captured pairs in amplexus and unmated, call-
ing males. These individuals were transported
to the laboratory, where the original male of a
pair in amplexus was exchanged with another
of greater or smaller SVL. Clutches produced
by the experimental pairs were maintained in
the laboratory for two days, after which the
number of embryos was counted.
For the statistical analyses of the results we
used the Student's t-test or the Pearson corre-
lation coefficient (Zar, 1984). Data of relative
sizes of mates influencing fertilization were fit-
ted to a second-degree polynomial regression
(Zar, 1984). The significance level considered
was 0.05.
RESULTS
Adult Characteristics.-Females were larger and
heavier after oviposition than males (Table 1).
Males in amplexus were larger and heavier than
unmated males (Table 2). The average (+SD)
mass of clutches was 0.74 ? 0.23 g (N = 14),
which represents 35% of mean females' mass
after oviposition.
Chorus Structure.-Choruses usually lasted
from sunset until approximately six hours. New
males continued to arrive at the chorus until
about three hours after sunset, after which the
TABLE 2. Comparisons (t-values) of mean snout-vent length and mean mass of mated and unmated males
of Hyla elegans.
Matedmales
N Mean + SD
SVL (mm) 53 26.35 ? 1.16
Mass (g) 52 1.23 ? 0.12
Females
N Mean + SD Range t df P
49 32.34 + 1.55 28.8-35.7 28.16 57 <0.01
59 2.01 + 0.26* 1.3-2.6* 21.23 61 <0.01
number of calling males decreased (Fig. 1). The
first females arrived at the chorus one or two
hours after sunset (x + SD = 81 min ? 35, range
= 50-131, N = 6 nights).
The operational sex ratio (or OSR, defined as
the number of reproducing females/number of
reproducing males) was highly male-biased
during the entire study period; estimates of OSR
varied from 0.024 to 0.224 (x + SD = 0.083 +
0.05; N = 23 nights). Number of mating females
was positively correlated with the number of
males (r = 0.818, P < 0.01; N = 23 nights), being
on average 10 males:1 female. Estimates of OSR
were not significantly correlated with the num-
ber of males present (r = 0.307, P > 0.05; N =
23 nights), indicating that females were not pro-
portionally more abundant in larger choruses
(Fig. 2).
Male and FemaleBehaviorin the Chorus.-Male
H. elegans defended calling sites by calling (see
Bastos and Haddad, 1995) and through physical
combat. These territories were not used for ovi-
position or shelter.
Few males (N = 4) adopted a crouched pos-
ture, did not emit calls (their vocal sacs were
not filled), and remained between 10 to 30 cm
from the calling males. There was no correla-
tion between the size of the chorus and the
number of these non-calling males. Aggressive
interactions between females were not ob-
served, and females adopted a crouched posture
similar to that displayed by non-calling males.
Gravid females observed on six occasions
moved slowly among the territories of calling
males, keeping a distance of about 8 to 10 cm
from the males. Other females (N = 3) perched
on branches of emergent vegetation, maintain-
ing a distance of about 20 to 70 cm from groups
of 4 to 5 males. Females sometimes sat near
males for as long as three hours, and in some
cases spent up to one hour near one male before
Unmatedmales
N Mean+ SD t df P
205 25.58 + 1.09 4.5 256 <0.01
171 1.15 + 0.21 3.2 154 <0.01
 REPRODUCTION IN HYLA ELEGANS 357

80' 0.3-

.
60
U) 0.2-
a
z U
0
u. 40 U
O
0.1- U
U * *s
LU
m U
*b
- 20 U
.
Z *

0 1 2 3 4 5 6
HOURSAFTER SUNSET
FIG.1. Mean numbers of calling males (?SD) of
Hyla elegansat different times. Averages calculated
for 10 nights.
moving toward others. However, no female was
observed approaching closely all the males
available at the pond on any particular night.
We also did not observe any male intercepting
a female that was moving toward a calling male.
We observed the initiation of amplexus nine
times. Amplexus is axillary and only occurred
when the female remained very close to a call-
ing male (<5 cm), although no physical contact
between the pair was observed before clasping.
Most males (N = 279) were not observed in
amplexus, 50 were seen clasping once, and only
three males were observed twice in amplexus
(one of these males was captured in amplexus
on. I I
I 50 10 150 200
CHORUSSIZE
FIG.2. Relationshipbetween number of males of
Hyla elegansin the chorus and OSRin 23 nights.
mass of females after oviposition (r = 0.425, P
< 0.01, N = 47). There was a higher percentage
of fertilization when the male SVL/female SVL
rate was closer to the mean value (x = 0.81)
recorded for amplectant pairs found in the field
(Fig. 4). Deviations from this mean rate value
resulted in a reduction in the percentage of fer-
tilization (98% at the mean value found in the
field; Fig. 4).
Six gravid females were marked and recap-
tured once; the average number of days be-
tween successive captures was 55.67 + 40.48
30-
a

on two consecutive nights). 28- * * *

E *
We found a positive correlation (r = 0.415, E U
U
P < 0.01; N = 51) between SVL of males and > 26- *. s
*

*I
females in amplexus, indicating positive as- u)
U . ?? ?
*
sortative mating for body size (Fig. 3A). On mmmm
?
average, SVL of males was 0.81 (SD = 0.04) of 24-
the length of females. There was no assortative
mating for mass (Fig. 3B; r = 0.189, P > 0.05; 22 I I v
N= 51). B 30 32 34 36
FEMALE
SVL (mm)
Oviposition.-After entering amplexus, the pair
remained at the male's calling site until neigh-
boring males ceased their calling or left the site. 1.6'
The durations between the start of amplexus U
and the beginning of oviposition for two pairs * U.. U U
1.4-
were 200 and 215 min, respectively. The am- 0
ua
plectant female walked in circles on the aquatic
U.... U

vegetation and periodically showed abdominal < 1.2- ? * a

??? .
contractions, arching her back and expelling -I
1.
? a ?U ?
her eggs, which were fertilized directly in the
water. The female freed herself from the am- 1.0o ? ?

plectant male by dislodging him with her hind

legs. Ovipositions ranged from 31 to 53 min (x 0.8 .
+ SD = 41 ? 9.38, N = 4). 1.0 1.5 2.0 2.5 3.0
Females deposited an average of 300 + 60 FEMALE
MASS(g)
eggs (N = 47). We found a significant positive FIG.3. Relationship between the snout-vent
correlation between clutch size and SVL (r = lengths (above) and masses (below) of males and fe-
0.382, P < 0.01; N = 46), and clutch size and males of Hyla elegans,capturedin amplexus.
358 R. P. BASTOS AND C. F. B. HADDAD
100-
80-
z
o 60-
< males, as reported for Hyla minuta(Haddad and
40-
t be related to competition for calling sites (Had-
.I
20-
SVL MALE/SVL FEMALE
FIG.4. Curvilinearrelationshipbetween percent-
age of eggs fertilized and the ratioof male to female
body size for pairs of Hyla elegansmated artificially
[Y = -3,673.0 + 9,620.3X - 6,137.3X2, R2= 0.85, P <
0.001].The arrow indicates the mean ratio (0.81) of
male to female body size for pairsfound in the field.
(range = 12-112). The numbers of eggs depos-
ited by two of these females were counted at
first capture and recapture. One female pro-
duced 301 and 241 eggs at first capture and re-
capture, respectively; the other, 538 and 266.
DISCUSSION
Hyla eleganshas a prolonged breeding season,
and males aggregate several nights throughout
the year. These aggregations can be classified
as leks because oviposition occurred away from
territories defended by selected males, and males
provided only sperm to females (Emlen and Or-
ing, 1977; Sullivan and Hinshaw, 1992). The
arrival of males and females at the breeding site
throughout the study was not synchronized.
Nightly estimates of OSRs were strongly male-
biased; on average, ten times as many males as
females were present each night. This estimate
is close to those reported for other species with
prolonged breeding seasons (e.g. Howard and
Kluge, 1985; Dyson et al., 1992). Such low OSRs
may account for why only 17% of the males
succeeded in mating one or more times.
The number of males was positively corre-
lated with the number of females present at the
chorus, as has also been observed for many anu-
rans (e.g., Morris, 1989; Sullivan and Hinshaw,
1992; Tejedo, 1993). This correlation may simply
indicate that the chorus composition is related
to environmental conditions (Green, 1990), be-
ing explained by causes not related to sexual
selection (Tejedo, 1993). Because the temporary
pond studied dried several times and could have
been the only local suitable site for breeding,
the aggregations of H. elegans could also be a
consequence of such transient conditions. There
was no significant correlation between number
of males present and estimates of OSR, indi-
cating that males in larger choruses do not have
a higher probability of mating than males in
smaller choruses.
Male H. elegansbegan calling activity and ter-
ritory establishment before the arrival of fe-
Cardoso, 1992). The earlier arrival of males may
dad and Cardoso, 1992). By obtaining a suitable
call site, a male increases the distance over which
his calls are broadcast.
As observed for other species of anurans
(Kluge, 1981; Perrill, 1984; Telford and Dyson,
1988), male H. elegans can breed on consecutive
nights. Male H. elegans that consistently attend
choruses might increase their chances of repro-
duction (see for other species: Given, 1988; Ritke
and Semlitsch, 1991).
Male H. elegans did not wait for contact by
the female in order to enter in amplexus; similar
observations were made by Morris (1989) on
Hyla chrysoscelis.Arnold (1976) suggested that
males do not wait for physical contact by fe-
males to prevent furtive clasping by satellite
males. The number of supposed satellite males
observed for H. elegans was low, which appar-
ently had little influence on the behavior of the
individuals. We did not observe males of H.
elegans intercepting gravid females moving to-
wards a calling individual. However, males may
attempt to intercept pairs as they move toward
oviposition sites, since females in amplexus only
began to move toward the oviposition site after
the nearest calling males ceased their activities
or left the vicinity. Such a hypothesis is based
on the possibility of interception of the pair and
the displacement of the males already in am-
plexus.
Female H. elegans were not amplexed by the
first male they encountered. However, any fe-
male mate choice was limited and females could
not analyse all the males from the chorus, due
to distance of displacement involved. Never-
theless, it is reasonable to assume that female
choice may occur at the level of local male group
and probably within a single night, in a similar
manner observed for H. chrysoscelis (Morris,
1989) and H. minuta(Haddad and Cardoso, 1992).
Assortative mating in some species of anu-
rans with axillary amplexus results in higher
fertilization success due to the greater precision
of juxtaposition of the cloacae of the partners
(Licht, 1976). Smaller males would be incapable
of adequately reaching the cloaca (and may not
have enough sperm to fertilize the whole
clutch); larger males would not be able to keep
a constant juxtaposition of cloacae throughout
the oviposition (Robertson, 1990; Bourne, 1993).
In H. elegans, the observation that a greater per-
centage of eggs were fertilized when the ratio
of male and female body size, in experimentally
 REPRODUCTION IN HYLA ELEGANS
paired individuals, was closer to the average
observed for natural pairings supports Licht's
hypotesis. We did not observe egg fertilization
in one instance in which the ratio of male to
female body size was high (0.90; Fig. 4). This
result is difficult to explain; however, similar
results were obtained for Uperoleia laevigata
(Robertson, 1990).
Although females of H. elegans may breed
more than once in a season, they seem to require
an interval of approximately two weeks be-
tween laying of two consecutive clutches. Such
an interval is similar to that reported for other
anurans (Wells, 1976; Perrill and Daniel, 1983).
Considering the methodology used in this study,
the interval could be even shorter, since marked
females may have laid eggs in days not included
in our samples in the field. The ability to pro-
duce more than one clutch during the repro-
ductive season seems to be restricted to species
of prolonged breeding (Howard, 1978; Telford
and Dyson, 1990).
Clutch size in H. elegans was positively cor-
related with both the SVL and the mass of fe-
males, as has been found in other species of
anurans (Berven, 1988; Martins, 1988). Within-
individual variation in the size of successive
clutches observed in our study has been re-
ported for other anurans (Perrill and Daniel,
1983; Ritke et al., 1990), and probably is related
to the ability of females to obtain resources
needed to produce another clutch (Kluge, 1981),
in addition to possible temperature influences
(Kaplan, 1987).
Our data on the reproductive behavior of
males and females of H. elegans indicate that
some individuals can breed more than once
during the year. This behavior may be highly
adaptive in coping with unpredictable abiotic
(e.g., rain), and biotic (e.g., competition and pre-
dation; Morin et al., 1990) factors.
Acknowledgments.-We thank A. S. Abe, J. P.
Pombal, Jr., M. J. Ryan, B. K. Sullivan, and two
anonimous reviewers for critical reading of the
manuscript. A. d'Heursel for the help with the
English. J. P. Pombal Jr., E. C. P. Pombal, L. P.
C. Morellato for field assistance. We also thank
the Instituto Florestal of the State of Sao Paulo
for permiting access to the study area and CNPq
for fellowships and financial support (C. F. B.
H. processes no 301281/91-2 and 403221/91-9).
LITERATURE CITED
ARAK,A. 1983. Male-male competition and mate
choice in anuran amphibians. In P. Bateson (ed.),
Mate Choice, pp. 181-210, Cambridge University
Press, Cambridge.
ARNOLD,S. J. 1976. Sexual behaviour, sexual inter-
ference and sexual defence in the salamanders Am-
359
bystomamaculatum,A. tigrinumand Plethodonjor-
dani.Z. Tierpsychol.42:247-300.
BASTOS,R. P. 1993. Reproductive biology of Hyla
elegans (Anura, Hylidae), in Ubatuba, State of Sao
Paulo. Master's Thesis, Universidade Estadual
Paulista, Rio Claro, SP, Brasil (in Portuguese).
,AND C. F. B. HADDAD. 1995. Vocalizations
and acoustic interactions in Hyla elegans (Anura,
Hylidae) during the reproductive activity. Natur-
alia 20:165-176 (in Portuguese).
BERVEN, K. A. 1988. Factors affecting variation in
reproductive traits within a population of Wood
Frogs (Rana sylvatica). Copeia 1988:605-615.
BOURNE, G. R. 1993. Proximatecosts and benefits of
mate acquisition at leks of the frog Ololygon rubra.
Anim. Behav. 45:1051-1059.
CHERRY,M. J. 1992. Sexual selection in the leopard
toad, Bufo pardalis. Behaviour 120:164-175.
DARWIN,C. 1871. The Descent of Man, and Selection
in Relation to Sex. Princeton Univ. Press, Prince-
ton, New Jersey.
DAVIES,N. B., AND T. R. HALLIDAY.1977. Optimal
mate selection in the toad Bufo bufo. Nature 269:
56-58.
DYSON, M. L., N. I. PASSMORE, P. J. BISHOP, AND S. P.
HENZI. 1992. Male behavior and correlates of
mating success in a natural population of African
painted reed frogs (Hyperolius marmoratus). Her-
petologica 48:236-246.
EMLEN,S. T., AND L. W. ORING. 1977. Ecology, sexual
selections and the evolution of mating systems.
Science 197:215-223.
FROST,D. R. (ed.) 1985. Amphibian species of the
world. A taxonomy and geographical reference.
Allen Press and Association Systematics Collec-
tions, Lawrence, Kansas.
GREEN,A. J. 1990. Determinants of chorus partici-
pation and the effects of size, weight and com-
petition on advertisement calling in the tungara
frog, Physalaemus pustulosus (Leptodactylidae).
Anim. Behav. 39:620-638.
GIVEN, M. F. 1988. Territoriality and aggressive in-
teractions of male carpenter frogs, Rana virgatipes.
Copeia 1988:411-421.
HADDAD, C. F. B., AND A. J. CARDOSO.1992. Female
choice in Hyla minuta (Amphibia: Anura). Acta.
Zool. Liolloana 41:81-91 (in Spanish).
HALLIDAY, T. R. 1983. The study of mate choice. In
P. Bateson (ed.), Mate Choice, pp. 3-32, Cambridge
University Press, Cambridge
HOWARD, R. D. 1978. The evolution of mating strat-
egies in bullfrogs, Rana catesbeiana.Evolution 32:
850-871.
, AND A. G. KLUGE. 1985. Proximate mecha-
nisms of sexual selection in wood frogs. Evolution
39:260-277.
KAPLAN, R. H. 1987. Developmental plasticity and
maternal effects of reproductive characteristics in
the frog, Bombinaorientalis. Oecologia (Berlin) 71:
273-279.
KLUGE,A. G. 1981. The life history, social organi-
zation, and parental behavior of Hyla rosenbergi
Boulenger, a nest-building gladiator frog. Misc.
Publ. Mus. Zool., Univ. Mich. 160:1-170.
LEE,J. C., AND M. L. CRUMP. 1981. Morphological
correlates of male mating success in Triprionpe-
360 R. P. BASTOS AND C. F. B. HADDAD
tasatusand Hyla marmorata(Anura: Hylidae). Oec-
ologia (Berlin) 50:153-157.
LICHT,L. E. 1976. Sexual selection in toads (Bufo
americanus).Can. J. Zool. 54:1277-1284.
Lurz, B. 1973. Brazilian Species of Hyla. Univ. Texas
Press, Austin.
MARTINS,M. 1988. Reproductive biology of Lepto-
dactylus fuscus in Boa Vista, Roraima (Amphibia:
Anura). Rev. Brasil. Biol. 48:969-977 (in Portu-
guese).
MARTOF, B. S. 1953. Territoriality in the green frog,
Rana clamitans.Ecology 34:165-173.
MORIN,P. J., S. P. LAWLER, ANDE. A. JOHNSON.1990.
Ecology and breeding phenology of larval Hyla
andersonii:the disadvantages of breeding late. Ecol-
ogy 71:1590-1598.
MORRIS, M. R. 1989. Female choice of large males in
the treefrog Hyla chrysoscelis:the importance of
identifying the scale of choice. Behav. Ecol. So-
ciobiol. 25:275-281.
OLSON,D. H., A. R. BLAUSTEN, AND R. K. O'HARA.
1986. Mating pattern variability among western
toad (Bufo boreas)populations. Oecologia (Berlin)
70:351-356.
PERRILL, S. A. 1984. Male mating behavior in Hyla
regilla. Copeia, 1984:727-732.
, ANDR. E. DANIEL.1983. Multiple egg clutch-
es in Hyla regilla, H. cinereaand H. gratiosa.Copeia
1983:513-516.
RITKE, M. E., J. G. BABB,ANDM. K. RITKE.1990. Life
history of gray treefrog (Hyla chrysoscelis)in west-
ern Tennessee. J. Herpetol. 24:135-141.
,AND R. D. SEMLITSCH. 1991. Mating behavior
and determinants of male mating success in the
gray treefrog, Hyla chrysoscelis.Can. J. Zool. 69:
246-250.
ROBERTSON, J. G. M. 1986. Female choice, male strat-
egies and the role vocalizations in the Australian
frog Uperoleiarugosa.Anim. Behav. 34:773-784.
. 1990. Female choice increases fertilization
success in the Australian frog, Uperoleialaevigata.
Anim. Behav. 39:639-645.
RYAN,M. J. 1983. Sexual selection and communi-
cation in a Neotropical frog, Physalaemuspustulo-
sus. Evolution 37:261-272.
. 1985. The Tungara Frog: a Study in Sexual
Selection and Communication. Univ. Chicago
Press, Chicago, Illinois.
SULLIVAN, B. K., ANDS. H. HINSHAW.1992. Female
choice and selection on male calling behavior in
the grey treefrog Hyla versicolor.Anim. Behav. 44:
733-744.
TEJEDO, M. 1993. Do male natterjack toads join larger
breeding choruses to increase mating success? Co-
peia 1993:75-80.
TELFORD, S. R., ANDM. L. DYSON. 1988. Some deter-
minants of the mating system in a population of
painted reed frogs (Hyperoliusmarmoratus).Behav-
iour 37:29-32.
. 1990. The effect of rainfall on interclutch
interval in painted reed frog (Hyperoliusmarmor-
atus). Copeia 1990:644-648.
WELLS,K. D. 1976. Multiple egg clutches in the green
frog (Rana clamitans). Herpetologica 32:85-87.
. 1977. The social behaviour of anuran am-
phibians. Anim. Behav. 25:660-693.
ZAR,J. H. 1984. Biostatistical Analysis. 2a ed. Pren-
tice-Hall, Englewood Cliffs, New Jersey.
Accepted: 24 April 1996.