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Society for the Study of Amphibians and Reptiles Society for the Study of Amphibians and Reptiles Predator Mediated Patch Use by Tadpoles (Hyla regilla): Risk Balancing or Consequence of Motionlessness? Author(s): Sarah J. Kupferberg Source: Journal of Herpetology, Vol. 32, No. 1 (Mar., 1998), pp. 84-92 Published by: Society for the Study of Amphibians and Reptiles Stable URL: http://www.jstor.org/stable/1565484 Accessed: 24-04-2015 21:35 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/1565484?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp 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 [email protected]. 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 This content downloaded from 136.152.142.28 on Fri, 24 Apr 2015 21:35:02 UTC All use subject to JSTOR Terms and Conditions Journalof Herpetology,Vol. 32, No. 1, pp. 84-92, 1998 Copyright1998 Society for the Study of Amphibiansand Reptiles PredatorMediated Patch Use by Tadpoles (Hyla regilla): Risk Balancing or Consequence of Motionlessness? SARAH J. KUPFERBERG1 Departmentof IntegrativeBiology, University of California,Berkeley, California 94720, USA ABSTRACT.-TOquantify the impact of garter snakes (Thamnophishydrophilus, formerly T. couchii) on tadpole prey (Hyla regilla),I experimentallyexamined whether: (1) garter snakes have greaterimpact on tadpole numbers or on behavior;(2) tadpole patch choice follows the rule of minimizing the ratio of mor- tality risk, iL, to foraging gain, g; and (3) predator-inducedresource avoidance influences algal production. In replicatedenclosed pools constructedon gravel bars of a northernCalifornia river, gartersnakes did not significantlyreduce the number of tadpoles but did alter patch choice.Tadpoles spent less time in high food quality algae mats (Cladophoraglomerata with epiphytic diatoms)and moretime in low food quality patches (Zygnematalesalgae mats and sediments) when gartersnakes were presentthan when they were absent In mortality risk experiments,however, there were no significant differences in the numbers of tadpoles con- sumed among patch types, and gartersnakes were not size selective. Therefore,patch choice did not follow the rule of minimizing /Ag. The change in patch choice was likely a consequenceof the sublethal effect of garter snakes in which tadpole activity is decreased. For negatively buoyant tadpoles, decreased activity results in sinking away from floating algal resources.This apparentresource avoidance by tadpoles did not affect algal mass over the relatively short durationof the experiments. For anurans, many factors affect the transition be misinterpreted as risk-balancing patch choice from tadpole to frog: temperature, nutrition, if motionlessness causes prey to sink away from competition, predation, and environmental vari- floating or suspended algal food resources. ability. These factors are summarized by two all I studied tradeoffs between predation risk encompassing parameters, A, risk of mortality, and foraging gain in a three trophic level sys- and g, growth rate, which have been used in tem: garter snake predators, tadpole prey, and recent models to predict optimal sizes at which algal resources. In western North America, Pa- ontogenetic changes in niche should occur (Wer- cific treefrogs (Hyla regilla) and garter snakes ner and Gilliam, 1984), and to predict patch (Thamnophisspp.) are extremely common in a choice in the presence of a predator (Gilliam wide range of aquatic habitats. Preliminary ob- and Fraser, 1987; Abrahams and Dill, 1989; Got- servations of western aquatic garter snake ceitas, 1990). The basic premise of risk-balancing (Thamnophishydrophilus, formerly T couchii) dis- models is that growing prey confront tradeoffs tribution and feeding on tadpoles in northern between gaining energy and avoiding preda- California rivers suggested that snakes were tion. Foraging in a rich habitat may provide concentrated around algal patches where tad- great energetic gain but may entail predation poles were foraging. To quantify the impact gar- risk, whereas staying in a refuge may entail risk ter snakes have on tadpole prey, I conducted a of starvation or slow growth, making transition series of replicated field experiments. Specifical- to the next life stage unlikely. When animals oc- ly, I examined whether: (1) garter snakes have cupy habitats which minimize the ratio (,/ g), greater impact on tadpole numbers or on be- population growth is maximized. havior, (2) tadpole patch choice follows the rule Adaptive behaviors that decrease vulnerabil- of minimizing // g; and (3) predator induced ity to predators, such as decreased activity lev- resource avoidance influences algal production. els, can also cause patterns of distribution STUDY SYSTEMNATURAL HISTORY among patches that appear to follow the ,//g rule. In addition to influencing shifts in habitat I conducted this research in the South Fork use, predation risk can decrease prey activity Eel River (Angelo Coast Range Reserve, Men- (Lawler, 1989; Werer, 1991; Semlitsch and Re- docino Co., California), where western aquatic yer, 1992; Skelly, 1995). Immobility can decrease garter snakes and Pacific treefrogs are common. the chance of attack by visual predators (Brodie Garter snakes have long been considered im- et al., 1974; Caldwell et al., 1980; Feder, 1983; portant predators of tadpoles because tadpoles Peterson and Blaustein, 1992) and increase sur- can make up a significant proportion of their vival time (Azvedo-Ramos et al., 1992). For neg- gut contents (Fitch, 1965; Arnold, 1992; Lind atively buoyant prey in aquatic environments, and Welsh, 1994). Garter snakes which forage at however, this predator avoidance behavior could aquatic/terrestrial margins may differentially This content downloaded from 136.152.142.28 on Fri, 24 Apr 2015 21:35:02 UTC All use subject to JSTOR Terms and Conditions TADPOLE PATCH CHOICE 85 prey upon metamorphic anurans (Arnold and effects of predatorchemical cues (Petrankaet al., Wassersug,1978; Jones, 1990) when locomotion 1987; Feminella and Hawkins, 1994), I cleaned is hindered by both tail and legs being present pools between trials by bailing out water and (Wassersug and Sperry, 1977). Some garter allowing them to refill by ground water intru- snakes, however, are specialized for underwater sion. Pools were excavated to compensate for foraging (Drummond, 1983; Schaeffel and de decreasing water table height when necessary. Queiroz, 1990)and consume tadpoles of all free- In all experiments, I stocked pools with ten swimming stages (Drummondand MaciasGar- H. regilla tadpoles that had been collected as cia, 1989). Thamnophishydrophilus in particular, eggs and reared on mixed algal diets in flow has visual system modifications, including a through 12 1 bucket enclosures in the river at a large degree of pupil constriction,allowing it to density of 10 tadpoles/bucket. Garter snakes focus under water at greaterdistances than oth- were caughtby hand, kept without food for 24- er snakes (Schaeffeland De Queiroz, 1990). 48 h, and then randomlyassigned to enclosures. In the South ForkEel River,T. hydrophilus and If mean size varied significantly among treat- tadpoles are found in two adjacenthabitats, the ments, snakes were reassigned. Juvenilesnakes main channel and stranded side pools. During were used in the experimentsbecause snakes of the low flow season the main channel becomes that size naturally spend 50% of their time in lentic and overgrown with the green alga Cla- similar pool habitats (Lind and Welsh, 1994). dophoraglomerata. Turfs of algae break off from Snake size ranged from 20 to 33 cm SVL. Trial rock substratesto form large floatingmats. Gar- durationvaried from experimentto experiment ter snakes often bask on and forage in the warm in part because of weather.Snakes often did not sunlit algae mats which contain invertebrates, forage on cool or cloudy days. fish, and tadpoles. Gartersnakes also forage in Patch Choice,Experiments I, II.-For Experi- shallow, turbid side pools that are cut off from ment I, in 1990,six replicateponds were stocked the river channel and support high densities of with tadpoles and 250 g damp mass Cladophora tadpoles. Tadpoledistribution is very patchy.In and 250 g Mougeotia.Algae were spun in a salad 1 m wide transectsurveys across side pools and spinner 50x before weighing to attain uniform adjacentriver channels,density routinelyvaries water content (Hay 1986).After one day a garter from 0-35/m2 between adjacent m2 plots (Kup- snake was added to each of three randomlycho- ferberg,unpubl. data). sen pools. Tadpolelocation and the numbersur- In the main channel and sidepools, algal viving were spot checked 4-6 times/d on days foods availableto tadpoles vary greatly in qual- 2, 5, and 6. The mean proportionin each algal ity in large part due to the presenceof nutritious type and in the gravel sediment on each day epiphytic diatoms. Tadpoles fed epiphytized was calculated.
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