Predation pressure during early life stages is often high, but few studies have examined antipredator responses at these stages. We studied the effects of an egg predator (leech, Haemopis sanguisuga) and two tadpole predators (dragonfly larvae, Aeshna spp.; and threespine stickleback, Gasterosteus aculeatus) on the timing of hatching and morphology of hatchlings and young tadpoles in two anuran amphibians [Rana arvalis (RA) and R. temporaria (RT)] in a factorial laboratory experiment. We also compared the responses of two geographically separated RA populations on the Baltic island of Gotland and in Uppland on the Swedish mainland. We found inconsistent evidence for the predictions that the presence of an egg predator induces earlier hatching, and the presence of a larval predator delays hatching. RT hatched later in the presence of stickleback than in the control treatment, but RA hatched earlier, less developed and at smaller size in the leech, dragonfly, and stickleback treatments. There was no indication of predator-induced morphology in hatchlings of either of the species. However, young RA tadpoles had shorter tails and deeper bodies in the stickleback treatment and RT had shorter tails in the leech, dragonfly and stickleback treatments. Irrespective of treatment, RA from Gotland hatched with relatively longer bodies than Uppland individuals and had relatively deeper and short tails as young tadpoles. Our results highlight the diversity of induced responses to predators in anuran amphibians: predator presence affects the timing of hatching and morphology of young tadpoles, but these responses vary depending on the species and predator considered.
Growth and development rates often differ among populations of the same species, yet the factors maintaining this differentiation are not well understood. We investigated the antipredator defences and their efficiency in two moor frog Rana arvalis populations differing in growth and development rates by raising tadpoles in outdoor containers in the nonlethal presence and absence of three different predators (newt, fish, dragonfly larva), and by estimating tadpole survival in the presence of free-ranging predators in a laboratory experiment. Young tadpoles in both populations reduced activity in the presence of predators and increased hiding behaviour in the presence of newt and fish. Older tadpoles from the slow-growing Gotland population (G) had stronger hiding behaviour and lower activity in all treatments than tadpoles from the fast-growing Uppland population (U). However, both populations showed a plastic behavioural response in terms of reduced activity. The populations differed in induced morphological defences especially in response to fish. G tadpoles responded with relatively long and deep body, short tail and shallow tail muscle, whereas the responses in U tadpoles were often the opposite and closer to the responses induced by the other predators. U tadpoles metamorphosed earlier, but at a similar size to G tadpoles. There was no evidence that growth rate was affected by predator treatments, but tadpoles metamorphosed later and at larger size in the predator treatments. G tadpoles survived better in the presence of free-ranging predators than U tadpoles. These results suggest that in these two populations, low growth rate was linked with low activity and increased hiding, whereas high growth rate was linked with high activity and less hiding. The differences in behaviour may explain the difference in survival between the populations, but other mechanisms (i.e. differences in swimming speed) may also be involved. There appears to be considerable differentiation in antipredator responses between these two R. arvalis populations, as well as with respect to different predators.
J. 2004. Temporal variation in predation risk: stage-dependency, graded responses and fitness costs in tadpole antipredator defences. Á/ Oikos 107: 90 Á/99.Temporal variation in predation risk may be an important determinant of prey antipredator behaviours. According to the risk allocation hypothesis, the strongest antipredator behaviours are expected when periods of high risk are short and infrequent. We tested this prediction in a laboratory experiment where common frog Rana temporaria tadpoles were raised form early larval stages until metamorphosis. We manipulated the time a predatory Aeshna dragonfly larva was present and recorded behavioural responses (activity) of the tadpoles at three different time points during the tadpoles' development. We also investigated how tadpole shape, size and age at metamorphosis were affected by temporal variation at predation risk. We found that during the two first time points activity was always lowest in the constant high-risk situation. However, antipredator response in the two treatments with brief high-risk situation increased as tadpoles developed, and by the third time point, when the tadpoles were close to metamorphosis, activity was as low as in the constant high-risk situation. Exposure to chemical cues of a predation event tended to reduce activity during the first time period, but caused no response later on. Induced morphological changes (deeper tail and shorter relative body length) were graded the response being stronger as the time spent in the proximity of predator increased. Tadpoles in the brief risk and chemical cue treatments showed intermediate responses. Modification of life history was only found in the constant high-risk treatment in which tadpoles had longer larval period and larger metamorphic size. Our results indicate that both behavioural and morphological defences were sensitive to temporal variation in predation risk, but behaviour did not respond in the manner predicted by the risk allocation model. We discuss the roles of concentration of predator chemical cues and prey stage-dependency in determining these responses.Prey animals often show highly sensitive responses in behaviour, morphology and life history to variation in predation risk. Although a large number of studies have described and analysed these responses in various systems (Kats and Dill 1998, Lima 1998a, b, reviewed by Tollrian and Harvell 1999, a potentially important element of predator Á/prey interactions remains obscure. In most natural systems predation risk varies temporally: predator activity patterns and densities vary both diurnally and seasonally, and the theory implies that such variation may have a profound impact on prey behaviour (Houston et al. 1993, Clark 1994, Lima and
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