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Beastly Bondage: The Costs of Amplexus in Cane Toads (Bufo marinus)

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 Copeia 2009, No. 1, 29–36

Beastly Bondage: The Costs of Amplexus in Cane Toads (Bufo marinus)

Haley Bowcock1, Gregory P. Brown1, and Richard Shine1

Reproductive activities can impose fitness costs as well as benefits. In most anuran species, males clasp females for
prolonged periods prior to gamete release, and intuition suggests that the male’s presence may impair the female’s
ability to move about and to feed. We tested the prediction that female locomotion and feeding would be reduced
during amplexus in laboratory experiments with Cane Toads, Bufo marinus. Amplexus reduced the female’s locomotor
(sprint and swim) performance, with the degree of locomotor impairment (speed and distance per hop) dependent
upon the body size of the amplectant male for trials of terrestrial locomotion, but not for aquatic locomotion.
Amplexus also reduced feeding rates in females; amplectant males did not feed at all. Overall, our data confirm that
amplexus imposes locomotor and feeding costs to female Cane Toads, and suggest that this distinctive posture may
generate sexual conflict in at least some anuran species.

S
UCCESSFUL reproduction is central to Darwinian
fitness, but for most organisms the process of
reproduction entails substantial potential costs (in
currencies such as energy expenditure and risk) as well as
benefits (the production of descendants carrying parental
genetic material). Energetic costs of reproduction are
important in asexual as well as sexually reproducing species,
but the process of sexual reproduction introduces a broad
suite of additional potential costs. For example, participat-
ing individuals may be injured during intraspecific antago-
nistic interactions or overly forceful attempts at copulation
(Leboeuf and Mesnick, 1991; Lew and Rice, 2005; Linder and
Rice, 2005; Bateman et al., 2006), be infected by sexually
transmitted diseases (Chlamydia psittaci bacterium in koalas:
Brown and Grice, 1984), have a reduced food intake
(Schwarzkopf, 1996), be exposed to high risks of predation
when reproduction takes them to habitats to which they are
poorly suited (sea turtles hauling out to oviposit on the
beach: Erkakan, 1993), or even suffer sexual cannibalism
(females of many spider and mantid species ingesting males
following copulation: Elgar, 1992).
Both the nature and magnitude of costs of reproduction
often differ substantially between males and females
(Trivers, 1972; Clutton-Brock, 1991). Typically, males invest
less in gametes or parental care but face high costs in male–
male competition and courtship, whereas for females, direct
energy investment into gametes may comprise much of the
annual energy budget (Clutton-Brock, 1991; Bonnet et al.,
2002). In an extreme example, male shrew-like marsupials
Antechinus die after a brief breeding season, due to
physiological stress (Lee et al., 1977); similarly, females of
many viviparous snake species may produce only a single
litter during their lifetimes, because of high vulnerability to
predation in the post-partum period (Madsen and Shine,
1992). Strong phylogenetic conservatism in mating systems
and sexual size dimorphism generate differences among
lineages in the nature and magnitude of costs of reproduc-
tion (Shine and Fitzgerald, 1995). For example, injuries
inflicted during male–male combat are significant in many
mammalian and avian groups (Promislow, 1992; Liker and
Szekely, 2005), whereas physical battles between rival males
are rare or absent in many other types of animals (e.g., most
anurans: Shine, 1979), and thus, other types of costs (such as
those associated with prolonged mate-searching) become
more significant in this latter group.
1
School of Biological Sciences A08, University of Sydney, NSW 2006, Australia; E-mail: (HB) haley.bowcock@gmail.com; (GPB)
gp_brown@bigpond.com; and (RS) rics@bio.usyd.edu.au. Send reprint requests to RS.
Submitted: 2 March 2008. Accepted: 11 June 2008. Associate Editor: J. D. Litzgus.
F 2009 by the American Society of Ichthyologists and Herpetologists
One major vertebrate lineage in which costs of reproduc-
tion remain relatively unexplored is the Anura (frogs and
toads). Anurans are speciose (.3000 species: Duellman and
Trueb, 1986; Frost et al., 2006) and frequent subjects of
research on reproductive biology (Cocroft and Ryan, 1995;
Roy, 1997; Kelley, 2004). Some aspects of costs of reproduc-
tion in these animals have been examined, such as the costs
of prolonged calling in males in terms of predation risk
(Ryan et al., 1981; Bernal et al., 2007), missed foraging
opportunities (Woolbright and Stewart, 1987), and energetic
costs (Grafe and Thein, 2001). Energy expenditure to
reproduction in females also has been addressed (Lemckert
and Shine, 1993; Castellano et al., 2004). Overall, reviews on
reproductive costs in amphibians focus on energetic expen-
diture (Pough et al., 1992). Less attention has been paid to
the fact that virtually all anurans show a distinctive posture
associated with reproduction, of a form that would plausibly
confer costs to one or both participants. This is amplexus,
whereby a male seizes a female with his forelegs, and retains
that grip until oviposition and fertilization has been
achieved (Duellman and Trueb, 1986). The duration of
amplexus varies greatly across species; generally, amplexus
lasts from a few hours to a few days (Wells, 1977), but can be
as long as several months (Dole and Durant, 1974).
Although the exact clasp position varies, the ubiquity (but
see Wells, 1977) of the amplexus posture provides a model
behavior through which to investigate the costs of repro-
duction within a major lineage of organisms. The potential
importance of amplexus costs is increased by the high
frequency of misdirected clasps (Sullivan and Wagner, 1988;
Marco and Lizana, 2002), the often prolonged duration of
amplexus, and reports of female anurans drowning during
amplexus with multiple males (Darwin, 1871; Davies and
Halliday, 1979; Sztatecsny et al., 2006).
To date, the only documentations of amplexus costs
involve metabolic costs of amplexus in Hyla versicolor
(McLister, 2003) and feeding behavior in captivity for
Bombina orientalis during amplexus (Mercadal and Ginzo,
2006). Beyond these reports, most authors simply infer that
amplexus is likely to be costly (Aronson, 1944; Blair, 1968).
Intuition suggests that a female anuran will be burdened by
an amplectant male: should we try to robustly test that
(arguably self-evident) prediction? Direct measurement can
be helpful even in such cases; by analogy, researchers
working with reptiles hypothesized that pregnancy would
DOI: 10.1643/CE-08-036
30
impair female locomotor performance (Shine, 1980) but
found that the magnitude of locomotor decrement varied
widely within as well as among species, and locomotor
decrement was linked to relative fecundity in some taxa but
not others (Seigel et al., 1987; Sinervo et al., 1991; Wapstra
and O’Reilly, 2001); indeed, females of some lizard species
ran faster not slower when gravid (Qualls and Shine, 1997).
Also, direct quantification has allowed comparison with
costs of other mobility-reducing phenomena such as a full
stomach, low body temperature, or tail autotomy (Shine,
2003), provided quantitative parameters for modelers (Vitt
and Price, 1982), and facilitated comparisons between
lizards and birds (Schwarzkopf, 1994; Veasey et al., 2001).
Similarly, studies on the costs of mate-guarding in insects
(which often involves amplexus-like behavior) have quan-
tified effects of carrying the partner on a male’s mobility,
predation risk, and energy balance (Fairbairn, 1993; Sparkes
et al., 1996). Direct measures of the costs of amplexus can
facilitate comparisons of the costs and benefits of alternative
reproductive tactics among major lineages.
Using the Cane Toad, Bufo marinus (see Frost et al., 2006
and Pramuk, 2006 for tentative generic reallocations to
Chaunus and Rhinella, respectively), we investigated the
following simple questions. First, does amplexus reduce
locomotor (sprint and swim) performance, and does the
degree of impairment depend upon the body size of the
amplectant male? Second, does amplexus reduce feeding
rates of the male and/or female?
MATERIALS AND METHODS
Study species.—The Cane Toad, Bufo marinus, is a toxic
bufonid native to Central and South America, introduced to
Australia in 1935 to control infestations of sugarcane crop
pests (Lever, 2001); it has now spread widely through
Australia (Sabath et al., 1981; Phillips and Shine, 2004;
Phillips et al., 2007). At maturity, Cane Toads grow to
.100 mm snout–urostyle length (SUL), with adult females
up to twice the mass of adult males (Lever, 2001). Cane
Toads are generalized feeders, but rely primarily on small
invertebrates (Lever, 2001). Based on stomach contents
analysis, female toads continue to feed even when full of
eggs (G. P. Brown, pers. obs.). Amplexus is axillary and (like
spawning) is seen in the field throughout the year at our
study site. Most amplectant pairs are found close to breeding
ponds, but amplexus also occurs in areas distant from water.
We have no data on the duration of amplexus in nature; we
have observed that captive Cane Toads often maintain the
posture for several days. Cane Toads are ideal model
organisms for studies of reproductive costs, as they are
abundant and their reproductive state can be manipulated
hormonally (Tyler, 1999). Also, Cane Toads readily amplex
in laboratory conditions (Bowcock et al., 2008).
Animal collection and induction of breeding behavior.—We
collected adult Cane Toads from the Adelaide River
floodplain, Northern Territory (12u389S, 131u199E) in No-
vember and December 2006. We individually marked all
animals with a unique number on a length of flagging tape
tied around the ‘waist.’ We measured mass and SUL in all
animals. Where amplectant pairs were required, we admin-
istered to each male a subcutaneous injection of 0.5 ml
leuprorelin acetate (LucrinH, Abbott Australasia, Kurnell,
Australia) as 0.25 mg ml21 in amphibian Ringer’s solution,
Copeia 2009, No. 1
at least three hours before the trials began. Leuprorelin
acetate mimics the action of gonadotropin releasing hor-
mone (GnRH), which stimulates gonadal production of sex
steroids. In anuran amphibians, administration of GnRH
stimulates the initiation of amplexus (Propper and Dixon,
1997). Females were not injected, to avoid possible con-
founding effects of hormonal status on the dependent
variables. We housed the females and injected males
communally (six males, six females) in large plastic tubs
(1.09 3 1.09 3 0.63 m) to encourage amplexus behavior
(because courtship behavior increases in groups: Wells,
1977). To stimulate breeding, we filled each tub with 10 L
of water, hay, and rocks, and set the tubs at a slight angle so
that the water pooled at one end. All experiments were
conducted at night, the usual activity period of adult Cane
Toads (Lever, 2001). Below, ‘status’ refers to the animal’s
amplectant condition (either amplectant or non-amplec-
tant).
Does amplexus reduce terrestrial locomotor performance?—We
assessed two aspects of locomotor performance (speed and
hops/meter) of 35 female toads (18 amplectant, 17 non-
amplectant) in a 10 m sprint track. The track was an outdoor
aluminum-sided enclosure (10 3 0.4 3 0.4 m) with paint
markings on the grass at every 2 m along the course. We
encouraged movement by walking behind the female and
gently prodding her hindlegs with a paint roller. To prevent
males from dismounting during a trial, we tethered
amplexed pairs of males and females to each other with
flagging tape around the midsection. To control against any
experimental artifacts of tethering, we also tied tape around
non-amplectant females. If males dismounted, or females
ceased to move (despite stimulation) during the trial, we
retained data recorded up to the last 2-m section traversed
by the pair. Although some females failed to complete the
course (either because the female stopped moving, or the
male dismounted), all managed to complete 4 m, enabling
us to compare performance at 4 m (for all groups) and 10 m
(for some groups). Thus, we assessed locomotor performance
with separate repeated-measures ANOVAs over the first 4 m
and the last 6 m. Because locomotor performance of
ectotherms frequently correlates with ambient temperature
(Huey and Stevenson, 1979), we recorded track temperature
at 30 min intervals using a Raytek infrared thermal-
monitoring gun (Raynger ST2L, Raytek Corp., Santa Cruz,
CA).
Does amplexus reduce aquatic locomotor performance?—We
measured the time required for 15 amplectant and 15 non-
amplectant females to swim around a circular pool (diam-
eter 2.4 m, water depth 0.15 m, fitted with a silicone ‘lane’
0.25 m from the edge) with four equal divisions (‘lengths’)
of 1.9 m marked out along the perimeter. In this way, if the
female changed direction during the trial, we could
calculate the time required to complete four lengths. Thus,
we assessed swim performance as speed (repeated-measures)
over four lengths. We encouraged the females to continue
moving by gentle stimulation of the hindlegs with a rod. We
recorded water temperature (with a thermometer) every
30 min.
Does amplexus reduce feeding rates?—We conducted feeding
trials with either a single, non-amplectant male or female
toad, or an amplectant pair (male–female) of toads (n 5 10
Bowcock et al.—Sexual conflict in anurans
for each treatment). The outdoor feeding arenas were large
plastic tubs (1.09 3 1.09 3 0.63 m) inclined at a slight angle,
with 10 L of water pooled at one end. We introduced ten
adult crickets into each tub and allowed 60 min for foraging.
Rocks in the water provided refugia for crickets, which could
otherwise drown (and would thus be ignored by the foraging
toads: H. Bowcock, pers. obs.). Fine mesh lids excluded other
invertebrate prey from the tubs. To control feeding history,
we deprived toads of food for 24 h prior to their trial. We
filmed the foraging bouts (under red light using a Sony
MiniDV Handycam) so that we could measure feeding rates
(number of crickets eaten through time) and identify which
sex did the foraging (in cases where there were two toads in
the arena). We recorded the time required for toads to eat
some or all of the crickets, then converted this to a feeding
rate (number of crickets consumed per hour). For the trials
involving pairs where the male dismounted before the
allotted one-hour period, we only scored foraging that
occurred while the pair was amplectant.
Data analysis.—Temperature was not significantly correlated
with any of the performance variables that we measured, so
we excluded it from analyses. We used repeated-measures
ANOVA in Statview (Version 5.0, SAS Institute Inc.) to
compare mean sprint and swim speeds, and hopping rates in
the sprint trials. We ran principal component (PC) analyses
in JMP (Version 5.0.1a, SAS Institute Inc.) on female size
measures (SUL and body mass) to extract an overall measure
of female body size. These PC scores differed significantly
between amplectant and non-amplectant females in the
swim (but not sprint) trials, so we included body size as a
covariate in our initial analyses of the former set of trials.
However, because female body size (PC score) did not
significantly influence swimming performance, we deleted
the size measure from our final analyses. To assess the role of
physical burdening on locomotor performance, we regressed
sprint speed and hop rates over the first 4 m against a
measure of relative body size of males (ratio of male to
female mass). We considered the first 4 m only for this
analysis, because all females completed this section of the
track. We performed a similar analysis on swim speeds,
including a similar distance (lengths 1 and 2 combined, 5
1.9 m + 1.9 m 5 3.8 m) to facilitate comparison with the
sprint trials. We compared feeding rates of amplectant and
non-amplectant females (males did not feed when amplec-
tant) using one-way ANOVA in Statview. Because the data
for feeding rates were non-normally distributed, we per-
formed analyses on fourth-root transformed measures of
feeding rates.
RESULTS
Does amplexus reduce terrestrial locomotor performance?—
Amplexus impaired the locomotor performance of female
toads in a sprint track (Fig. 1, Table 1). Non-amplectant
females were faster in the final six meters than the first four
meters, whereas the speeds of amplectant females did not
change significantly over the two distances (significant
status*distance interaction; Fig. 1A, Table 1). The presence
of an amplectant male reduced the distance covered per
hop, though this difference was not significant. Females
took more hops/meter in the first 4 m than the last 6 m
(regardless of status; Fig. 1C, Table 1). Increasing relative
mass of the male (i.e., male mass divided by female mass)
31
was accompanied by a decrease in the female’s speed
(Fig. 1B; n 5 18, R2 5 0.33, P , 0.01) and an increase in
number of her hops/meter (Fig. 1D; n 5 18, R2 5 0.33, P ,
0.01).
Does amplexus reduce aquatic locomotor performance?—Am-
plexus impaired swimming performance (Fig. 2A, Table 1).
Non-amplectant females swam faster than amplectant
females, with both sets of females swimming progressively
more slowly with each length (Fig. 2A). The mass of the
male relative to the female did not affect swim performance
significantly (Fig. 2B; n 5 15, R2 5 0.01, P 5 0.81).
Does amplexus reduce feeding rates?—Males did not feed at
all while in amplexus (Fig. 3). Amplectant females con-
sumed fewer crickets than did non-amplectant females; the
difference was more than sixfold (untransformed means of
6.01 and 39.67 crickets/hour, respectively; one-factor AN-
OVA on fourth-root transformed data: F1,19 5 22.88, P 5
0.0001).
DISCUSSION
Intuition suggests that an amplectant male anuran is likely
to reduce a female’s ability to move about and to feed. Such
effects may be particularly strong in a species where adult
males can equal or exceed the size of their female partners
(such as the Cane Toad: Lever, 2001) and where amplexus is
very prolonged (Dole and Durant, 1974; Wells, 1977).
Frustratingly, we have little field data on the incidence
and duration of amplexus. How species deal with the
possible costs of amplexus likely varies with how much
time they spend in amplexus. Costs may be minimal in
species with short amplexus times, but in species where
males actively search for females (including in areas away
from the breeding site), attempt to dislodge amplectant
males, and/or have long amplexus times, the females may be
expected to incur significant costs. Hence, theory predicts
that adaptations of male and female anurans to cope with
the costs of the amplexus may vary interspecifically and be
correlated with aspects of the mating system that influence
the role of amplexus. Future work could usefully explore the
occurrence and duration of amplexus in the field as a basis
for predicting interspecific variation in adaptive traits
related to cost minimization.
In our study, amplectant male Cane Toads impaired
locomotor speed of females both on land and in water,
reduced the distance covered per hop on land, and reduced
rates of food consumption by females. These results
(together with those of McLister, 2003 and Mercadal and
Ginzo, 2006) suggest that sexual conflict resulting from
amplexus may be widespread in anurans, and not an
occasional phenomenon restricted to females drowning
due to amplexus by multiple males (Darwin, 1871; Davies
and Halliday, 1979; Sztatecsny et al., 2006).
The causal basis of the locomotor decrement due to
amplexus presumably involves a simple physical burden: in
our studies, the largest males can weigh 130% more than the
smallest gravid female. In keeping with this inference, larger
male size relative to female size predicted the magnitude of
locomotor impairment, at least for terrestrial locomotion
(Fig. 1B). No such effect was apparent in aquatic locomo-
tion, perhaps because the water supported much of the
male’s mass (Fig. 2B). Also, the magnitude of the locomotor
decrement differed between these two sets of trials:
32 Copeia 2009, No. 1

Fig. 1. Effects of an amplectant male on terrestrial locomotor speeds of female Cane Toads, Bufo marinus, over a 10 m raceway. The histograms
show (A) mean (6SE) sprint speeds and (C) hops/meter by status (non-amplectant, unfilled bars and amplectant, gray bars). See Table 1 for a
statistical summary. The graphs on the right show relationships between the degree of physical burdening (ratio of male mass to female mass) and
(B) female speed and (D) number of hops per meter. Larger male size impaired female locomotion more significantly than did smaller males.

amplexus decreased sprint speeds by about 25%, but
swimming speeds by almost 50% (Figs. 1A vs. 2A). That is,
amplexus was a greater locomotor impediment to females in
the water than on land, even if the amplectant male was
relatively small. That difference in locomotor impairment
presumably reflects biomechanical differences between
swimming and terrestrial locomotion in anurans (Nauwe-
laerts et al., 2007).
In terms of feeding, the mechanisms for continued
(though reduced) feeding in females, and anorexia in males,
are likely to be physical. For females, the amplectant male
sometimes had a grip in front of her legs, which hampered
movement and thus, foraging ability. The fact that amplec-
tant females do eat while amplectant may offset the large
energy input required for gametogenesis (Mercadal and
Ginzo, 2006). Amplectant males, on the other hand, have
their head firmly pressed against the female’s back and thus
would have difficulty opening their mouths to trap crickets,
and by doing so, may lose their grip on the female.
Presumably for the amplectant male, the risk of losing a
pairing outweighs any costs of short-term anorexia.
Clearly, one critical issue is whether or not these
laboratory measures of the costs of amplexus translate into
actual fitness costs to females in the wild. There are no data
to support this extrapolation, although several authors have
referred to an increased vulnerability to predation of
amplectant pairs relative to solitary males and females
(Blair, 1968; Howard, 1981). If such a vulnerability results
from the reduced mobility due to amplexus, the results of
our sprint trials suggest that relative mass of male relative to
female also may influence vulnerability. This aspect not
only complicates the issue of fitness benefits and costs of
Bowcock et al.—Sexual conflict in anurans 33

Table 1. Summary Statistics for the Effects of an Amplectant Male on Locomotor Performance in Female Cane Toads, Bufo marinus. Sprint speed
and hops/m were measured over a 10 m raceway, in two sections (4 m and 6 m). Swim speed was measured in a circular arena of four 1.9 m
sections. Variables are independent factors in repeated-measures ANOVA: ‘Status’ refers to amplectant or non-amplectant females; ‘Distance’ refers
to the section of the track (sprint speed and hops/m) or circular arena (swim speed); and ‘Interaction’ is the interaction term Status*Distance.

Performance measure Variable df F P

Sprint speed Status 1,26 9.98 0.004
Distance 1,26 2.38 0.14
Interaction 1,26 6.26 0.019
Hops/m on land Status 1,26 1.77 0.20
Distance 1,26 6.20 0.0195
Interaction 1,26 2.33 0.01391
Swim speed Status 1,28 38.12 ,0.0001
Distance 3,28 5.37 0.0019
Interaction 3,28 1.72 0.17

mating with males of different sizes, but fits well with field
observations of female mortality during multiple amplexus
(Davies and Halliday, 1979; Sztatecsny et al., 2006). If male
mass relative to female mass is a critical determinant of the
degree of effective burdening, then a female amplexed
simultaneously by multiple males (Kapfer and Muelfeld,
2006) will be more substantially disadvantaged than a
female amplexed by a single amorous partner. Although
male body size had no significant effect on female
swimming speed in our trials, the mortality risks to
amplexus are likely to be greater in the water than on land.
In our laboratory trials, some females were unable to
complete even the first section of the pool and instead were
forced underwater by the amplectant male. Very large males
(relative to female size) or multiple males (see above) would
increase the risk of females drowning.
High costs of amplexus to females do not necessarily
result in sexual conflict: similarly high costs to the
amplectant male may result in males and females having
similar optimal durations of amplexus. However, costs
plausibly are higher for females than for males, because of
the asymmetry in control: males determine when to initiate

Fig. 2. Effects of an amplectant male on aquatic locomotor speeds of

female Cane Toads, Bufo marinus, over four 1.9 m sections (‘lengths’)
of a circular test arena. Panel (A) shows mean (6SE) swim speed by
status (non-amplectant, unfilled bars and amplectant, gray bars).
Statistics are summarized in Table 1. Panel (B) shows the lack of a
significant relationship between the degree of physical burdening (ratio
of male mass to female mass) and female swimming speed over the
first two lengths of the arena.
Fig. 3. Effects of amplexus on feeding rates (mean 6 SE, calculated
from untransformed data) of amplectant and non-amplectant male and
female Cane Toads, Bufo marinus, offered crickets in the laboratory.
Amplectant males did not feed at all, whereas amplectant females fed
less than did non-amplectant females.
34
and terminate amplexus, whereas females are unable to
escape because they cannot give release calls (Bowcock et al.,
2008). The optimal duration of amplexus often will be brief
for a female (amplexus is costly, and the female only needs
the male to be in position just prior to spawning) but long
for a male (if male mating success is maximized by seizing
any female available and hanging on to her even if
spawning is delayed for a few hours or days). This male–
female conflict over the optimal duration of amplexus (and
the identity of the male partner, in situations where females
gain fitness benefits from mate choice) is resolved in the
male’s favor, because only he can terminate amplexus. If his
costs exceed his benefits, he can simply let go of the female.
If her costs exceed her benefits, however, she cannot rid
herself of the burden. Based on that asymmetry in control,
evolutionary theory predicts that males will engage in
amplexus only when there is a net fitness benefit to that
sex, whereas females will have no option but to engage in
amplexus even at times when the net fitness consequences
to females are negative. That situation will generate sexual
conflict.
In nature, male toads often amplect other males as well as
females (Lever, 2001). Although we were unable to measure
the costs to males being amplexed by other males (because
such claspings did not persist long enough for us to quantify
effects), the same kinds of performance decrements are
likely to apply. Hence we might expect males to be under
intense selection to be able to terminate sex-inappropriate
amplexus attempts; the ‘release call’ presumably evolved
under this situation (Duellman and Trueb, 1986; Bowcock et
al., 2008). The cessation of feeding by amplectant Cane
Toad males in our study also suggests that prolonged
amplexus—for weeks or months—might impose a signifi-
cant energy cost to a reproducing male. We stress that we
have not measured fitness costs, only effects on feeding and
locomotion. It remains possible that in Cane Toads
amplexus costs bear little relationship to fitness because
the posture often is maintained only briefly (hours rather
than days), and these animals may rely on their potent
toxins rather than speed to escape from predators. None-
theless, Cane Toads provide a logistically feasible model
with which to measure costs that for so long have simply
been taken as self-evident. In the process, we have detected
non-intuitive patterns: for example, the difference between
terrestrial and aquatic habitats both in terms of the
magnitude of locomotor impairment due to amplexus, and
in the significance of relative male size for such costs. There
is enormous potential for interspecific comparisons in this
neglected area of amphibian biology. Anurans encompass a
massive diversity of reproductive strategies, involving great
variation in cost-determining parameters such as the
duration of amplexus, the degree of sexual size dimorphism,
and vulnerability to predation. Hence, these animals may
provide ideal model systems with which to explore factors
that influence the intensity of sexual conflict.
ACKNOWLEDGMENTS
The University of Sydney Animal Ethics Committee (AEC)
approved our procedures (AEC number L04/10-2006/1/
4480), and the Australian Research Council funded the
work. Many thanks to M. Elphick, M. Greenlees, T. Child,
and the rest of Team Bufo for assistance in data collection
and manuscript preparation, and the staff of Beatrice Hill
Farm for logistical support.
Copeia 2009, No. 1
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