Society for the Study of Amphibians and Reptiles

Advertisement Call Variation and Observations on Breeding Behavior of Bufo debilis and B.
punctatus
Author(s): Brian K. Sullivan
Source: Journal of Herpetology, Vol. 18, No. 4 (Dec., 1984), pp. 406-411
Published by: Society for the Study of Amphibians and Reptiles
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Journalof Herpetology,Vol. 18, No. 4, pp. 406-411, 1984
Copyright 1984 Society for the Study of Amphibians and Reptiles

Advertisement Call Variation and Observations on Breeding

Behavior of Bufo debilis and B. punctatus

BRIAN K. SULLIVAN

Departmentof Zoology, Arizona State University, Tempe,Arizona 85287, USA

ABsTRAcr.-Advertisementcalls of Bufo debilis and B.punctatus were analyzed to determine if

variation in calls was correlated with body size and temperature.Advertisement call variables
(frequency, pulse rate or call duration) were not significantly correlated with male snout-vent
length in either species. All three call variables, as well as call rate, were significantly correlated
with body temperaturein B.punctatus;analysis of temperature-inducedvariation was not possible
for B. debilis. Observationsrevealed that males of neither species employ satellite or active-search-
ing strategiesto locate females in choruses.To initiate amplexusa female approachesa calling male
and the male continues to vocalize until the female makes physical contact.These results are dis-
cussed in relation to previous work with bufonids.

Breeding behavior of a number of an- undescribed. This report summarizes

urans has recently been subjected to so-
ciobiological analysis (see reviews by
Wells, 1977; Sullivan, 1983a). These and
other studies have generated consider-
able controversy concerning the means
by which non-random mating can arise
in anuran breeding aggregations
(Woodward, 1982; Sullivan, 1983a).
Many workers have hypothesized (1)
that female anurans prefer large males
as mates, and (2) that females assess male
size on the basis of advertisement call
variables which are assumed to vary
predictably with male size (Wilbur et
al., 1978; Ryan, 1980; Fairchild, 1981).
Analysis of advertisement call parame-
ters and observations of male and fe-
male behavior in breeding aggregations
are vital to studies testing these hypoth-
eses. However, detailed investigations
have been conducted for only a handful
of anurans (Kagarise Sherman, 1980;
Kluge, 1981; Ryan, 1983; Sullivan, 1982,
1983a).
The Green toad, Bufo debilis, and the
Red-spotted toad, B. punctatus, are two
bufonids found throughout much of the
arid and semi-arid southwestern United
States. Although systematic relation-
ships of these two species have been in-
vestigated (Blair, 1972), their breeding
behavior and mating systems remain
observations on behavior of males and
females in breeding aggregations and
analyzes variation in advertisement calls
of these two anurans. I explicitly test
the hypotheses: (1) that call variables
are predictably correlated with male
size, and (2) that call parameters vary
significantly with body temperature in
B. punctatus.
STUDYAREAS
Six choruses of B. debilis were ob-
served in rain-formed pools in south-
eastern Arizona and extreme south-
western New Mexico for eight nights
during July of 1981 and 1983. Choruses
were located 20 km north of Douglas on
Route 666, Cochise Co., AZ; at the in-
tersection of Route 80 and Rucker Can-
yon Road, Cochise Co., AZ; and within
5 km of Rodeo, Hidalgo Co., NM. One
population of B. punctatus along Pica-
dillo Creek at its intersection with Route
143, Maricopa Co., AZ, was observed for
ten nights throughout its breeding pe-
riod in 1983 (25 April-30 May).
METHODS
Toads were captured by hand and toe-
clipped for individual identification.
Snout-vent length (SVL) was mea-
sured to the nearest mm with a plastic

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 CALLSAND BREEDINGIN BUFO
rule. Observations were conducted with
a six-volt headlamp, and toads were
usually not disturbed by my presence:
when approached, males in a chorus
either continued or resumed calling
within 1 min. On each night some males
were individually observed for 2.5-5.0
min periods, and whenever females
were located they were observed until
they left the chorus or selected a mate.
Advertisement calls of both species
were recorded in the field with a Uher
recorder (4000 IC; tapespeed = 188 mm/
sec) and Shure Uni-dyne IV (548) mi-
crophone. Body temperatures were tak-
en within ten sec of recording a male's
call by inserting the bulb of a Schul-
theis Quick-Recording thermometer 5
mm into the cloaca. Calls were analyzed
in the lab with a Kay 6061B Sonagraph,
following Zweifel (1968) and Sullivan
(1982). Dominant frequency was esti-
mated from a section taken at call mid-
point. Pulse rate was determined by
counting the number of pulses given in
a 0.5 sec portion of a wide-band (300 Hz
filter) audiospectrogram, also taken at
call mid-point. For B. debilisand B. punc-
tatus dominant frequency and pulse rate
remain constant within an individual
call. Call durations were measured in
seconds from tape recordings using a
stopwatch.
Recordings of 16 males calling with
body temperatures of 21.5 ? 1.0?Cwere
used to assess size-related variation in
calls of B. debilis;for B. punctatus record-
ings of 13 males with body tempera-
tures of 18.5 ? 1.5?C were used. Re-
cordings were selected in this manner
to control for possible temperature-in-
duced variation. Product-moment cor-
relations were calculated for SVL and
each call variable.
For B. debilis there was insufficient
range (20.5-22.5?C) in body tempera-
ture to allow analysis of temperature-
related variation. For B. punctatus, 21
recordings of 15 males recorded on suc-
cessive nights were used to assess vari-
ation associated with small changes in
body temperature (range = 8?C). To de-
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407
termine if observations of large or small
males predominated at high or low
temperatures the distribution of male
calls was evaluated in relation to me-
dian body temperature (18.9?C). Of the
11 calls of males with body tempera-
tures above median temperature, 6 were
of males larger than mean size (52.4
mm) and 5 were of males smaller than
mean SVL. Similarly, of the 10 calls of
males below median temperature, 5
were of large males and 5 were of small
males. Hence, distribution of calls of
large and small males in relation to me-
dian temperature was not significantly
different from random (X2= .01, P >
0.05), and analysis of temperature-in-
duced variation in call attributes was not
biased by possible size-related variance.
RESULTS
Mating Behavior: Bufo debilis.-Males
formed choruses in temporary water
sources for 1-3 nights following sum-
mer rainstorms. Male density was low,
ranging from 0.004 to 0.067 males per
m2 of pond surface. However, calling
males aggregated along one section of
shoreline in larger pools, although
within these aggregations males were
usually separated by 0.5-3.0 m. Occa-
sionally males called within 0.1 m of
one another, but no non-calling satel-
lite or active-searching behaviors were
observed. Males called on shore within
2 m of water, or in extremely shallow
water, or in emergent vegetation with
their hindquarters submerged.
Females were observed (N = 8) only
at one chorus, and of these only six were
observed in amplexus. I saw four of
these females select their mates: each
time the female approached a calling
male and jumped on him following his
vocalization. As the female neared the
male, he remained stationary and con-
tinued giving advertisement calls until
the female made physical contact. Fol-
lowing this the male moved about rap-
idly, apparently attempting to clasp the
female who had jumped on and off his
back. However, if the female moved
408 BRIANK. SULLIVAN

TABLE1. Snout-vent length (SVL) and adver-

by 1.5 to 4 m. A male generally called
tisement call variables of 16 male B. debilis,body
from on shore, or in very shallow water,
temperature 21.5 ? 1.0?C,and 13 male B. puncta- for 1-3 min, then submerged for 0.5-
tus, body temperature 18.5 ? 1.5?C.
1.5 min before emerging for another
Bufodebilis Bufopunctatus
bout of vocalization. When males moved
Fre- Call Fre- Pulse Call
and began calling within approximate-
quen- Pulse dura- quen- rate dura- ly 1 m of one another, they rapidly ap-
SVL cy rate tion SVL cy (p/ tion each other while giving ad-
(mm) (kHz) (p/sec) (sec) (mm) (kHz) sec) (sec)
proached
vertisement calls. They engaged in
37 3.3 126 7.3 47 2.3 28 12.6
38 3.3 132 4.5 47 1.8 38 9.6
amplexus-like wrestling bouts during
which both males gave advertisement
38 3.0 120 2.4 48 2.3 38 7.6
39 3.2 130 5.1 49 2.5 38 8.3 calls. These interactions lasted 0.5 to 6
42 3.3 126 7.5 51 2.3 42 10.9 min and when they separated, both
42 3.1 144 - 52 2.3 38 10.2 males returned to their calling sites.
42 3.1 128 - 52 1.6 38 3.3
42 2.8 138 - 53 2.2 40 10.0 Only eight females were found in
43 3.2 132 4.9 53 1.9 36 6.3 breeding aggregations and never was
43 3.2 138 4.9 54 2.3 38 6.4 more than one female observed at a
44 3.4 132 5.3 54 2.2 36 9.4 chorus. Initiation of amplexus was ob-
44 2.9 128 - 55 2.2 38 9.1 served only once: similar to B. debilisthe
45 3.0 142 4.8 55 2.4 38 6.7
45 2.7 128 - male continued to vocalize until the fe-
-
46 3.3 130 5.3 - - - - male made physical contact. After the
46 3.0 140 8.1 - - male clasped the female the pair moved
out of the chorus area and the female
began oviposition. Females (57.5 mm)
were significantly larger (t = 3.75, P <
more than a few cm the male stopped 0.05, N = 41) than males (52.7 mm).
and gave another advertisement call. Call Analysis.-For B. debilisdominant
During this call the female moved to- frequency was not significantly corre-
ward the male and again made physical lated with male SVL (r = -.24, P = 0.37,
contact: this sequence was repeated un- N = 16), nor was pulse rate (r = .43, P =
til amplexus was achieved. Although 0.09, N = 16) or call duration (r = .27,
sample size was low, mean size of males P = 0.43, N = 11). Thus, there is no evi-
who mated (N = 6; SVL = 41.3 mm) was dence that females can use these vari-
not significantly different (Mann-Whit- ables to determine male size. Similarly,
ney U = 36, P > 0.05, N = 15) from for B. punctatus neither frequency (r=
mean size of unmated males (N = 9; .07, P = 0.81, N = 13), pulse rate (r = .38,
SVL = 42.3 mm). Females (50.3 mm) P = 0.21, N = 13), call duration (r = .42,
were significantly larger (t = 5.1, P < P = 0.15, N = 13) nor call rate (r = .33,
0.05, N = 18) than males (42.2 mm). P = 0.36, N = 10) were significantly
Mating Behavior: Bufo punctatus.- correlated with male SVL. Call vari-
Small choruses of 2-6 males formed ables for both species are given in Table 1.
along Picadillo Creek (1 m wide). I ob- Initial analysis of temperature-relat-
served four choruses in a 200 m transect ed variation in calls of B. punctatus re-
and choruses were separated by 30-75 vealed that dominant frequency was not
m. Choruses formed in the same loca- significantly correlated with tempera-
tion on successive nights, although one ture (r = .39, P = 0.08, N = 21). How-
gradually shifted approximately 4 m ever, one call was recorded from a male
downstream as the study progressed. with his vocal sac partially submerged:
During an evening all males occu- the frequency of his call may have been
pied 1-2 m diameter calling sites en- reduced as a result of a change in the
compassing both shores of the stream; resonance of his submerged vocal sac
within choruses males were separated (see Martin, 1972). When this apparent-

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 CALLSAND BREEDINGIN BUFO 409

2.6-
50-
2A- I-4--
-
^ 2.2- * .~~~~~~ ..
40-
U

u ! Y=2.25X-2.48
30-
,Z 2.0
Y = 0.04X + 1.53 ! Nu 21
. 20-
- 1.8- N =20

1.6- 16 17 18 19 20 21 22 23 24
_ ? BODY TEMPERATURE
(C?)
16 17 18
1 19 20 21 22 23 24
BODY TEMPERATURE
(C?) FIG. 2. Pulse rate (pulses per second) against
FIG.1. Dominant frequency against body tem- body temperature for Bufo punctatus.
perature for advertisement calls of Bufo punctatus.
The call eliminated from analysis is circled (see
text). nents (Martin, 1972) of the vocal tract;
hence, these parameters are expected to
vary with body temperature. However,
ly aberrant recording is eliminated fre- it is presumed that dominant frequency
quency is significantly correlated with is determined by "passive" components
temperature (r = .53, P = 0.01, N = 20; (e.g., vocal cord size, mass, tension).
Fig. 1). Pulse rate (r = .84, P < 0.01, N = Dominant frequency has been found to
21; Fig. 2), call duration (r = -.54, P = correlate (positively) with body tem-
0.01, N = 20; Fig. 3) and call rate (r = perature in some populations of B.
.80, P < 0.01, N = 21) were also corre- americanus, B. speciosus, B. valliceps and
lated with body temperature. B. woodhousei (Porter, 1964; Zweifel,
1968). It may be that in these forms tem-
DISCUSSION
perature significantly influences active
Contrary to hypotheses concerning modulation (via muscle contraction) of
size-related variation in anuran call at- the passive components determining
tributes, call variables of B. debilis and dominant frequency (Martin, 1972). In
B. punctatusdo not vary predictably with B. punctatus pulse rate and call rate in-
male size. Hence, females cannot select crease with temperature, and call du-
large males as mates using these param- ration decreases, suggesting that the
eters. Dominant frequency of advertise- passive and active control of these pa-
ment calls has been found to vary with rameters is coupled (see Green, 1982).
body size in populations of B. america- My observations reveal that females
B. B.
nus, canorus, valliceps and B. wood-
housei (Zweifel, 1968; Kagarise Sher-
man, 1980; Sullivan, 1982), but not in B. 12 Y= -0.52X + 17.68
canaliferus, B. coccifer and B. cognatus N 20
(Porter, 1964; Sullivan, 1983b). Al- 10

though mechanics of vocalization in the 8 .~~~~~~~~~~~~~~~~~~~ *

Bufonidae have been investigated in z

0
some detail (Martin, 1972), no hypoth- 6 * * |~~~~~~~~~~~~~~~
esis has been developed to account for 4
these differences. 4

Temperature has a significant effect

2
on call variables of B. punctatus, as has
been found for a number of bufonids 16 17 18 19 20 2 22 23 24
BODY TEMPERATURE(C?)
(e.g., Zweifel, 1968; Sullivan, 1982).
Pulse rate and call duration are vari- FIG.3. Call duration against body temperature
ables controlled by "active" compo- for Bufo punctatus.

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410 BRIAN K. SULLIVAN
of both B. debilisand B. punctatusare able
to move freely in breeding aggrega-
tions to initiate amplexus with calling
males. Males do not engage in scramble
competition and actively-search for fe-
males. Although the breeding period of
a given population of B. debilisis short,
low male density and asynchronous ar-
rival of females at ponds provides an
opportunity for females to freely select
their mates. For B. punctatus the breed-
ing period is more prolonged, but again,
temporal availability of females results
in highly male-skewed operational sex
ratios. Further study is necessary to de-
termine if in fact females of these two
species do discriminate and actively se-
lect their mates according to some as-
pect of male phenotype. Although sam-
ple size was low, the results of the
present investigation suggest that mat-
ing is random with respect to size in B.
debilis.
Wells (1977) hypothesized that an-
uran breeding behavior is influenced by
breeding period length. Wells suggest-
ed that the high density of males in
breeding aggregations of species with
extremely short breeding periods min-
imizes the opportunity for female
choice. Conversely, female choice could
be a significant factor in breeding ag-
gregations of anurans with prolonged
breeding periods. Wells recognized that
male density is the primary factor lim-
iting female choice: at high densities
males actively search and physically
struggle for possession of females. At
low densities males typically remain
stationary and vocalize to attract fe-
males. Wells' hypothesis concerning in-
fluence of male density has been sup-
ported by studies of a number of
bufonids. In B. debilis and B. punctatus,
as well as B. cognatus and B. woodhousei
(Sullivan, 1983a, b, c), low male density
correlates with the opportunity for fe-
males to initiate amplexus with partic-
ular males, independent of breeding
period duration. Although it has been
useful to characterize anurans as explo-
sive or prolonged breeding forms
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(Wells, 1977), it is variation in male
densities and operational sex ratios that
may significantly influence the action
of sexual selection in breeding aggre-
gations.
Acknowledgments.-I would like to
thank my wife Elizabeth for her valu-
able assistance in the field, and her sup-
port in other aspects of my work. Rob
Bowker, D. Delisle, T. Jones and C. Levy
deserve thanks for help with observa-
tions of B. punctatus. Jack Fouquette is
appreciated for discussions and com-
ments on manuscripts. I thank N. J.
Meffe for typing. This work was sup-
ported in part by the Theodore Roose-
velt Memorial Fund.
LITERATURE CITED
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GREEN,D. M. 1982. Mating call characteristics of
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 CALLS AND BREEDING IN BUFO 411

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Accepted: 31 January 1984.

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