Predation on Bufo terrestris Tadpoles: Effects of Cover and Predator Identity

Author(s): Kimberly J. Babbitt and Frank Jordan

Source: Copeia, Vol. 1996, No. 2 (May 16, 1996), pp. 485-488
Published by: American Society of Ichthyologists and Herpetologists (ASIH)
Stable URL: http://www.jstor.org/stable/1446873
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 SHORTER CONTRIBUTIONS 485
reef edge, with minimal water movement at all
stages of the tide.
The fishers relied on a specific and unique
signal each year which indicated the aggrega-
tions were imminent-the smell of the annual
coral spawn, which the fishers believed was the
"smell of the harid" They knew that the long-
nose parrotfish would begin aggregating the
morning after they first smelled them around
sunset. Many fishers gathered for the schooling
in March 1994, near the time of the full moon,
when there was a strong fishlike smell over the
whole island. On this occasion, however, the
fish came in the following month. I investigated
the smell and found that it originated from fresh
slicks of coral spawn. The corals appeared to
have spawned on the first evening after the full
moon. Interestingly, fishers in Samoa and Fiji
rely on the smell of coral spawn to indicate that
the palolo worms will rise within three days;
they believe the smell originates from the worms
(L. Zann, pers. comm.), and Johannes (1981)
reported that Palauan fishers were sometimes
one month early in predicting the time of
spawning of some reef fishes.
Harvesting.-The aggregating parrotfish were
easily harvested. People watched for the schools
from the tops of the coral cliffs from before
sunrise, beginning a few days before the full
moon. On each morning, there were from 300-
500 men and boys waiting on the cliffs; how-
ever, not all participated in the harvesting. As
each school came over the reef crest and into
shallow water, up to 20 men, carrying mono-
filament gill nets, ran over the shallow reef flat
toward the school. Most schools were easily ap-
proached and did not appear to be disturbed
by the approaching people nor by the nets which
enclosed them. This apparent ease with which
the longnose parrotfish were approached and
captured is often observed among schools of
food fishes that are preparing to spawn (Jo-
hannes, 1981). At least one school on each day
escaped and was not harvested. I have no in-
formation on what happened to those schools
that escaped on the day of harvesting because
the fishers were reluctant to allow me to enter
the water. When I dived in the same area one
week later, however, there were still consider-
able numbers of H. harid feeding on the reef
crest and reef slope, just as there were before
the aggregations and harvesting.
Acknowledgments.-Thanks to D. Bellwood, R.
E. Johannes and E. Joubert for helpful com-
ments on earlier versions of this manuscript.
This project was supported at the NCWCD in
Riyadh by A. H. Abuzinada, M. Sulayem, and
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E.Joubert. Assistance at Farasan Island was pro-
vided by G. M. Ali, M. Al-Sanei, and R. Al-
Khuzaiym.
LITERATURE CITED
GLADSTONE, W. 1994. Lek-like spawning, parental
care and mating periodicity of the triggerfish Pseu-
dobalistesflavimarginatus (Balistidae). Env. Biol. Fish.
39:249-257.
JOHANNES, R. E. 1978. Reproductive strategies of
coastal marine fishes in the tropics. Ibid. 3:65-84.
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fornia Press, Berkeley.
PHILLIPS DALES, R. 1976. Annelids. 2d ed. Hutch-
inson Univ. Library, London.
RANDALL, J. E. 1986. Red Sea reef fishes. Immel
Publishing, London.
ROBERTSON, D. R., AND R. R WARNER. 1978. Sexual
patterns in the labroid fishes of the western Carib-
bean, II: the parrotfishes (Scaridae). Smithson.
Contr. Zool. 255:1-26.
THRESHER, R. E. 1984. Reproduction in reef fishes.
TFH Publications, Neptune City, NJ.
National Commission for Wildlife Conservation and
Development, PO Box 61681, Riyadh 11575,
Kingdom of Saudi Arabia. Present address: Great
Barrier ReefMarine Park Authority, PO Box 1379,
Townsville, Queensland 4810, Australia. Sub-
mitted: 13 April 1995. Accepted: 21 July
1995. Section editor: R. Winterbottom.
@ 1996 by the American Society of Ichthyologists and Herpetologists
Copeia, 1996(2), pp. 485-488
Predation on Bufo terrestris Tadpoles:
Effects of Cover and Predator Identity
KIMBERLY J. BABBITT AND FRANK JORDAN
Predation is a major source of mortality in
anuran tadpoles (Calef, 1973; Heyer et al., 1975;
Smith, 1983) and can have a significant effect
on assemblage structure through differential
predation on competitively dominant species
(Morin, 1981, 1983; Wilbur, 1980). Several
mechanisms can function to reduce predation
pressure on tadpoles, including crypsis (Cald-
well, 1982), unpalatability (Formanowicz and
Brodie, 1982), and several behavioral responses
involving alteration of the timing, amount, or
location of activity (Semlitsch, 1987; Stangel and
Semlitsch, 1987; Lawler, 1989). Finally, be-
cause many predators of tadpoles are gape-lim-
 486 COPEIA, 1996, NO. 2
ited, individuals reaching a sufficiently large size
are able to escape predation (Crump, 1984;
Travis et al., 1985; Richards and Bull, 1990;
but see Crump and Vaira, 1991).
Development in structurally complex habi-
tats provides another mechanism through which
predatory effects may be mediated. Increased
habitat complexity (i.e., cover, density, hetero-
geneity) can reduce encounter rates with pred-
ators by providing cover or refuge areas for
prey (Stein and Magnuson, 1976). Predators may
have decreased foraging success in complex
habitats due to difficulties in maneuvering
through dense vegetation or to decreased visual
range (Crowder and Cooper, 1982; Savino and
Stein, 1982; Werner et al., 1983).
Most studies examining the role of habitat
complexity on predator-prey interactions in
aquatic systems have focused on fishes. Consid-
ering the important role of predation on tad-
pole survival and assemblage structure (re-
viewed in Wilbur, 1980), surprisingly little re-
search has focused on the role of cover in me-
diating tadpole-predator interactions. Evidence
that habitat structure is important comes indi-
rectly from studies of amphibian assemblages
where researchers observed tadpoles surviving
in the face of predation through the use of ben-
thic refuge areas (e.g., Morin, 1983, 1986).
However, in one of the only direct tests using
amphibian prey, Figiel and Semlitsch (1991)
found that increased habitat complexity did not
enhance survival of gray treefrog (Hyla chrysos-
celis) tadpoles exposed to crayfish (Procambarus
acutus) predators. In this study, we examine how
varying levels of habitat complexity affect sur-
vival of Bufo terrestris tadpoles exposed to odo-
nate naiads (Anaxjunius) and to aquatic hemip-
terans (Belostoma fluminea).
Materials and methods.-Experiments were car-
ried out at the National Biological Service's
Southeastern Biological Research Center in
Gainesville, Florida. In the laboratory, we filled
180 polyethylene containers (34 x 39 x 14 cm)
with seven liters of well water. Plant stems (Lud-
wigia repens) were added at one of five treatment
levels: none, low (10 g), medium (30 g), high
(60 g), and very high density (90 g). The me-
dium plant density treatment approximated the
plant complexity observed (qualitatively) in the
pond from which we collected both predators
and prey. The two higher plant density treat-
ments are within the range of plant densities
we have observed at other sites where we have
collected both Bufo terrestris tadpoles and pred-
ators. Plant treatments were randomly assigned
to containers, and containers were then ran-
domly assigned to different positions on labo-
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ratory shelves. During the experiment, water
temperatures were 21-23 C, and photoperiod
was maintained at 10L:14D.
We collected prey and predators from a single
pond located at the Southeastern Biological Re-
search Center. We collected three egg masses
of the southern toad (Bufo terrestris) and main-
tained them together in well water in the lab-
oratory until they reached developmental stage
25 (Gosner, 1960). Average snout-vent length
of tadpoles at this time was 4.07 ? 0.08 mm (n
= 30). We then haphazardly assigned 25 tad-
poles to each experimental container. We al-
lowed tadpoles to acclimate to experimental
conditions for 2 h prior to introducing preda-
tors.
We used naiads of the common green darner
(Anax junius) or adult giant water bugs (Belos-
tomafluminea) as predators because both species
are common predators in the vegetated ponds
and marshes of the area. The average length
(? 1 SD) of darner naiads was 44.6 ? 4.6 mm
(n = 15; average wet weight = 1.36 ? 0.22 g),
whereas the average length of giant water bugs
was 20.3 ? 1.1 mm (n = 15; average wet weight
= 0.27 ? 0.04 g). Predators were held individ-
ually without food for 48 h to standardize hun-
ger levels within each species. The experiment
began at 0600 on 27 April 1991 when we added
predators individually to containers. We ter-
minated the experiment after 36 h and deter-
mined survival. In addition to predator treat-
ments, we ran controls without predators to as-
sess whether plant treatment alone affected sur-
vival. Controls were replicated five times per
plant density.
Two species of predator (Anax, Belostoma) were
crossed with five levels of plant density (none,
low, medium, high, very high) in this 2 x 5
randomized factorial design. Treatments were
replicated unevenly, but at least 10 replicates
were used for each treatment combination.
Fixed effect analysis of variance (ANOVA) was
used to test whether predator type, plant den-
sity, or their interaction affected the survival of
Bufo tadpoles. Visual inspection of residuals and
F-max tests indicated that data transformations
were unnecessary (i.e., ANOVA assumptions
were validated). The Bonferonni-Dunn post hoc
method (Day and Quinn, 1989) was used to test
for differences in tadpole survival among the
five plant density treatments. Controls were not
included in the above analyses because there
was 100% survival across all plant density treat-
ments.
Results.-All tadpoles in the no-predator con-
trol containers survived. In experimental con-
tainers, both predator type (F1,,118 = 23.36, P =
 SHORTER CONTRIBUTIONS 487
0.001) and plant density (F4,,,, = 2.96, P = 0.023)
affected tadpole survival. Overall, Belostoma
predators consumed about 54% of the tadpoles,
whereas about 30% were consumed by Anax.
Belostoma predators ate more tadpoles than did
Anax at all plant densities (Fig. 1). Tadpole sur-
vival generally increased with increasing plant
density (Fig. 1); however, only the low and very
high levels differed significantly (P = 0.0044;
Bonferonni-Dunn test). The interaction be-
tween predator type and plant density was not
significant (F,,,,8 = 0.50, P = 0.737), indicating
that predator foraging success was affected sim-
ilarly across plant densities.
Discussion.--Both factors examined in this study,
habitat complexity (i.e., plant density) and pred-
ator species, had significant effects on survival
of Bufo terrestris tadpoles. Similar to results from
studies of fish predators (e.g., Crowder and
Cooper, 1982; Savino and Stein, 1982; Rozas
and Odum, 1988), high cover levels in this study
resulted in increased prey survival. Reduction
in foraging success for odonates probably re-
sulted from interference in stalking behavior
and decreased visual range. Visual detection and,
to a lesser extent, tactile stimulation are im-
portant for prey capture by odonate naiads
(Pritchard, 1965), and high cover levels can in-
terfere with such foraging tactics (Folsom and
Collins, 1984). Belostoma are also visually ori-
ented foragers and were probably affected sim-
ilarly; however, they preyed on more tadpoles
at every cover level. Because all cover levels
resulted in similar interspecific differences in
predation levels, it is likely that interspecific dif-
ferences between Anax and Belostoma are related
more to physiological differences in food intake
requirements or absolute differences in forag-
ing ability rather than to any mechanism related
specifically to habitat complexity. For example,
in experiments where cover levels were fixed,
Gascon (1992) found that aeshnid naiads con-
sumed more tadpoles than libellulid naiads.
In addition to morphological, developmental,
or behavioral mechanisms possessed by tadpoles
of different species, increased habitat complex-
ity may provide a potentially important mech-
anism for decreasing predation rates on tad-
poles. For predators that detect prey through
movement, low activity of tadpoles coupled with
complex habitat structure may further decrease
capture rates. High cover provided increased
protection to Bufo terrestris tadpoles, which are
active foragers (pers. obs.), and we would pre-
dict that high cover levels would provide com-
paratively greater protection for less active spe-
cies.
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25-
O Anax
1 Belostoma
20-
00. 15_
0,
-o
5
0
None Low Medium High Very high
Plant density
Fig. 1. Number of Bufo terrestris tadpoles that sur-
vived predation under different cover levels. Bars are
means ? 1 SE.
The results of this study contrast with those
of Figiel and Semlitsch (1991), who did not find
differences in tadpole survival with increasing
habitat complexity. These investigators used ar-
tificial cover (window screen) which probably
provided low cover at all treatment levels com-
pared to our high cover treatment. Alterna-
tively, the contrasting results of these two ex-
periments may be related to species-specific for-
aging tactics of crayfish as compared to insect
predators.
The relationship between habitat complexity
and predation may be influenced by several fac-
tors such as microhabitat use and foraging ac-
tivities of both prey and predators, leading to
complex results (Stredl and Collins, 1992). In-
teractions among habitat complexity, predator
density, and prey density can lead to complex,
nonlinear relationships that can have an im-
portant influence on species interactions, even
when habitat complexity does not have a large
infuence on tadpole survival (Stredl and Collins,
1992). More research is needed to elucidate the
relationships between tadpoles, predators, and
habitat structural complexity.
Acknowledgments.-We appreciate the support
of the staff of the Southeastern Biological Re-
search Center, Gainesville, Florida. We thank
W. M. Kitchens and G. W. Tanner for their
continuing support. M. L. Crump provided
valuable comments on the manuscript. This pa-
per is Contribution No. R-04842 of the Journal
 488 COPEIA, 1996, NO. 2
Series, Florida Agricultural Experiment Sta-
tion, Gainesville, 32611.
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(KJB) Department of Wildlife Ecology and Conser-
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@ 1996 by the American Society of Ichthyologists and Herpetologists
Copeia, 1996(2), pp. 488-493
Station-Holding by the Mottled Sculpin,
Cottus bairdi (Teleostei: Cottidae),
and Other Fishes
PAUL W. WEBB, CYNTHIA L. GERSTNER, AND
SCOTT T. MINTON
Many small fusiform fish live in strong uni-
directional currents in intertidal, subtidal, and
freshwater lotic habitats. Many of these fishes
are benthic, including members of the Cottidae,
Gobiidae, Blennidae, Loricariidae, and etheos-