Title: Partitioning the non-consumptive effects of predators on prey with complex life histories

Jon M. Davenport1*, Blake R. Hossack2 and Winsor H. Lowe1

1 Division of Biological Sciences, University of Montana, Missoula, MT, USA 59812

2U.S. Geological Survey, Northern Rocky Mountain Science Center, Aldo Leopold Wilderness Research Institute, 790 E. Beckwith Ave., Missoula MT, USA 59801

*corresponding author:

Appendix A. A table of published references and total number of lines per reference used in the meta-analysis of prey activity. Predators functional groups are based on published references and acronyms are as follows: NGL = non-gape-limited, GL = gape-limited, SAP= sit-and-pursue, ACT = active.

Predator species / Predator Functional group / Prey Species / # of lines / Source
Anax imperator / NGL, SAP / Pelophylax lessonae / 6 / Altwegg 2002
Anax junius / NGL, SAP / Lithobates sphenocephalus / 2 / Babbitt 2001
Aeshna palmata / NGL, SAP / Rana aurora / 1 / Barnett & Richardson 2002
Aeshna palmata / NGL, SAP / Rana pretiosa / 1 / Barnett & Richardson 2002
Aeshna juncea / NGL, SAP / Rana temporaria / 2 / Laurila & Kujasalo 1999
Aeshna spp. / NGL, SAP / Rana arvalis / 2 / Laurila et al. 2006
Triturus vulgaris / GL, ACT / Rana arvalis / 2 / Laurila et al. 2006
Gasterosteus aculeatus / GL, ACT / Rana arvalis / 2 / Laurila et al. 2006
Aeshna spp. / NGL, SAP / Rana temporaria / 1 / Orizaola et al. 2012
Anax spp. / NGL, SAP / Lithobates sylvaticus / 8 / Relyea 2002
Salmo salar / NGL, ACT / Rana temporaria / 2 / Nicieza 2000
Belostoma spp. / NGL, SAP / Agalychnis callidryas / 2 / Vonesh & Warkentin 2006
Anax spp. / NGL, SAP / Anaxyrus americanus / 4 / Skelly & Werner 1990
Gambusia holbrooki / GL, ACT / Litoria aurea / 2 / Hamer et al. 2002
Gambusia holbrooki / GL, ACT / Crinea signifera / 2 / Lane & Mahoney 2002
Gambusia holbrooki / GL, ACT / Limnodynastes tasmaniensis / 2 / Lane & Mahoney 2002
Ambystoma tigrinum / GL, ACT / Hyla versicolor / 2 / Skelly 1992
Perca fluviatilis / NGL, ACT / Lestes sponsa / 2 / Johansson et al. 2001
Salvelinus fontinalis / NGL, ACT / Baetis bicaudatus / 4 / Peckarsky & McIntosh 1998
Aeshna cyanea / NGL, SAP / Coenagrion puella / 8 / Mikolajewski et al. 2005
Gobio gobio / NGL, ACT / Lestes depressa / 4 / Mikolajewski et al. 2007

Literature Cited for Appendix A

Altwegg, R. 2002. Predator-induced life-history plasticity under time constraints in pool frogs.

Ecology 83:2542-2551.

Babbit, K.J. 2001. Behaviour and growth of southern leopard frog (Rana sphenocephala)

tadpoles: Effects of food and predation risk. Canadian Journal of Zoology 79:809-814.

Barnett, H.K., and J.S. Richardson. 2002. Predation risk and competition effects on the life-

history characteristics of larval Oregon spotted frog and larval red-legged frog. Oecologia 132:436-444.

Hamer, A.J., S.J. Lane, and M.J. Mahony. 2002. The role of introduced mosquitofish (Gambusia

holbrooki) in excluding the native green and golden bell frog (Litoria aurea) from original habitats in south-eastern Australia. Oecologia 132:445-452.

Johansson, F., R. Stoks, L. Rowe, and M. De Block. 2001. Life history plasticity in a damselfly:

effects of combined time and biotic constraints. Ecology 82:1857-1869.

Lane, S.J., and M.J. Mahony. 2002. Larval anurans with synchronous and asynchronous

developmental periods: contrasting responses to water reduction and predator presence. Journal of Animal Ecology 71:780-792.

Laurila, A., and J. Kujasalo. 1999. Habitat duration, predation risk and phenotypic plasticity in

common frog (Rana temporaria) tadpoles. Journal of Animal Ecology 68:1123-1132.

Laurila, A., S. Pakkasmaa, and J. Merilä. 2006. Population divergence in growth rate and

antipredator defences in Rana arvalis. Oecologia 147:585-595.

Mikolajewski, D.J., T. Bodin, F. Johansson, and G. Joop. 2005. Phenotypic plasticity in gender

specific life-history: effects of food availability and predation. Oikos 110:91-100.

Mikolajewski, G. Joop, and B. Wohlfahrt. 2007. Coping with predators and food limitation:

testing life history theory for sex-specific larval development. Oikos 116:642-649.

Nicieza, A.G. 2000. Interacting effects of predation risk and food availability on larval anuran

behaviour and development. Oecologia 123:497-505.

Orizaola, G., E. Dahl, and A. Laurila. 2012. Reversibility of predator-induced plasticity and its effects at a life-history switch point. Oikos 121:44-52.

Peckarsky, B.L., and A.R. McIntosh. 1998. Fitness and community consequences of avoiding multiple predators. Oecologia 113:565-576.

Relyea, R.A. 2002. Local population differences in phenotypic plasticity: Predator-induced changes in wood frog tadpoles. Ecological Monographs 72:77-93.

Skelly, D.K. 1992. Field evidence for a cost of behavioral antipredator response in a larval amphibian. Ecology 73:704-708.

Skelly, D.K., and E.E. Werner. 1990. Behavioral and life-historical responses of larval American toads to an odonate predator. Ecology 71:2313-2322.

Vonesh, J.R., and K.M. Warkentin. 2006. Opposite shifts in size at metamorphosis in response to larval and metamorph predators. Ecology 87:556-562.

Appendix B. A table of published references and total number of lines per reference used in the meta-analysis of prey larval period. Predators functional groups are based on published references and acronyms are as follows: NGL = non-gape-limited, GL = gape-limited, SAP= sit-and-pursue, ACT = active.

Predator species / Predator Functional group / Prey Species / # of lines / Source
Anax imperator / NGL, SAP / Pelophylax lessonae / 3 / Altwegg 2002a
Anax imperator / NGL, SAP / Pelophylax lessonae / 5 / Altwegg 2002b
Anax imperator / NGL, SAP / Pelophylax esculentus / 5 / Altwegg 2002b
Anax junius / NGL, SAP / Lithobates sphenocephalus / 2 / Babbitt 2001
Aeshna palmata / NGL, SAP / Rana aurora / 1 / Barnett & Richardson 2002
Aeshna palmata / NGL, SAP / Rana pretiosa / 1 / Barnett & Richardson 2002
Notonecta spp. / NGL, ACT / Anaxyrus boreas / 1 / Chivers et al. 1999
Aeshna brevistyla / NGL, SAP / Limnodynastes peronii / 1 / Kraft et al. 2005
Dytiscus marginalis / NGL, ACT / Rana arvalis / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Bufo bufo / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Rana temporaria / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Hyla arborea / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Pelobates fuscus / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Rana dalmatina / 1 / Lardner 2000
Dytiscus marginalis / NGL, ACT / Bufo calamita / 1 / Lardner 2000
Aeshna juncea / NGL, SAP / Rana temporaria / 2 / Laurila & Kujasalo 1999
Aeshna juncea / NGL, SAP / Rana temporaria / 2 / Laurila et al. 1998
Aeshna juncea / NGL, SAP / Bufo bufo / 2 / Laurila et al. 1998
Aeshna juncea / NGL, SAP / Rana temporaria / 1 / Laurila et al. 2004
Aeshna spp. / NGL, SAP / Rana arvalis / 2 / Laurila et al. 2006
Lissotriton vulgaris / GL, ACT / Rana arvalis / 2 / Laurila et al. 2006
Gasterosteus aculeatus / GL, ACT / Rana arvalis / 2 / Laurila et al. 2006
Aeshna spp. / NGL, SAP / Rana temporaria / 1 / Orizaola et al. 2012
Anax spp. / NGL, SAP / Hyla versicolor / 2 / Relyea & Hoverman 2003
Salmo salar / NGL, ACT / Rana temporaria / 2 / Nicieza 2000
Salmo trutta / NGL, ACT / Lissotriton helveticus / 1 / Orizaola & Bana 2005
Aeshna cyanea / NGL, SAP / Lissotriton spp. / 1 / Van Buskirk & Schmidt 2000
Belostoma spp. / NGL, SAP / Agalychnis callidryas / 2 / Vonesh & Warkentin 2006
Aeshna cyanea / NGL, SAP / Bufo bombina / 2 / Vorndran et al. 2002
Aeshna cyanea / NGL, SAP / Bufo variegata / 2 / Vorndran et al. 2002
Anax spp. / NGL, SAP / Anaxyrus americanus / 4 / Skelly & Werner 1990
Gambusia holbrooki / GL, ACT / Litoria aurea / 2 / Hamer et al. 2002
Taricha granulosa / GL, ACT / Rana aurora / 1 / Kiesecker et al. 2002
Gambusia holbrooki / GL, ACT / Crinea signifera / 2 / Lane & Mahoney 2002
Gambusia holbrooki / GL, ACT / Limnodynastes tasmaniensis / 2 / Lane & Mahoney 2002
Ambystoma macrodactylum / GL, ACT / Ambystoma macrodactylum / 2 / Wildy et al. 1999
Notophthalmus viridescens / GL, ACT / Anaxyrus americanus / 2 / Raffel et al. 2011
Haemopis sanguisuga / NGL, ACT / Rana dalmatina / 1 / Hettney et al. 2011
Nepa spp. / NGL, SAP / Rana dalmatina / 1 / Hettney et al. 2011
Lissotriton vulgaris / GL, ACT / Rana dalmatina / 1 / Hettney et al. 2011
Aeshna cyanea / NGL, SAP / Rana dalmatina / 1 / Hettney et al. 2011
Perca fluviatilis / NGL, ACT / Leucorrhinia dubia / 1 / Johansson 2002
Perca fluviatilis / NGL, ACT / Lestes sponsa / 2 / Johansson et al. 2001
Salvelinus fontinalis / NGL, ACT / Baetis bicaudatus / 4 / Peckarsky & McIntosh 1998
Salvelinus fontinalis / NGL, ACT / Callibaetis ferrugineus hageni / 2 / Caudill & Peckarsky 2003
Toxorhynchites rutilus / NGL, SAP / Aedes triseriatus / 16 / Hechtel & Juliano 1997
Aeshna cyanea / NGL, SAP / Coenagrion puella / 4 / Mikolajewski et al. 2005
Perca fluviatilis / NGL, ACT / Lestes sponsa / 1 / Brodin & Johansson 2002
Lepomis gibbosus / NGL, ACT / Chironomus tentans / 7 / Ball & Baker 1995
Anax junius / NGL, SAP / Lestes dryas / 4 / De Block et al. 2008
Anax junius / NGL, SAP / Lestes forcipatus / 4 / De Block et al. 2008
Anax junius / NGL, SAP / Lestes congener / 3 / De Block et al. 2008
Australopelopia prionoptera / NGL, ACT / Polypedilum australotropicus / 3 / McKie & Pearson 2006
Gobio gobio / NGL, ACT / Lestes depressa / 4 / Mikolajewski et al. 2007
Salvelinus fontinalis / NGL, ACT / Baetis bicaudatus / 2 / Peckarsky et al. 2001

Literature Cited for Appendix B

Altwegg, R. 2002a. Predator-induced life-history plasticity under time constraints in pool frogs.

Ecology 83:2542-2551.

Altwegg, R. 2002b. Trait-mediated indirect effects and complex life-cycles in two European

frogs. Evolutionary Ecology Research 4:519-536.

Babbit, K.J. 2001. Behaviour and growth of southern leopard frog (Rana sphenocephala)

tadpoles: Effects of food and predation risk. Canadian Journal of Zoology 79:809-814.

Ball, S.L., and R.L. Baker. 1995. The non-lethal effects of predators and the influence of food

availability on life history of adult Chironomus tentans (Diptera: Chironomidae). Freshwater Biology 34:1-12.

Barnett, H.K., and J.S. Richardson. 2002. Predation risk and competition effects on the life-

history characteristics of larval Oregon spotted frog and larval red-legged frog. Oecologia 132:436-444.

Brodin, T., and F. Johansson. 2002. Effects of predator-induced thinning and activity changes on life history in a damselfly. Oecologia 132:316-322.

Caudill, C.C., and B.L. Peckarsky. 2003. Lack of appropriate behavioral or developmental responses by mayfly larvae to trout predators. Ecology 84:2133-2144.

Chivers, D.P., J.M. Kiesecker, A. Marco, E.L. Wildy, and A.R. Blaustein. 1999. Shifts in life history as a response to predation in western toads (Bufo boreas). Journal of Chemical Ecology 25:2455-2463.

De Block, M., M.A. McPeek, and R. Stoks. 2008. Life history plasticity to combined time and

biotic constraints in Lestes damselflies from vernal and temporary ponds. Oikos 117:908-916.

Hamer, A.J., S.J. Lane, and M.J. Mahony. 2002. The role of introduced mosquitofish (Gambusia

holbrooki) in excluding the native green and golden bell frog (Litoria aurea) from original habitats in south-eastern Australia. Oecologia 132:445-452.

Hechtel, L.J., and S.A. Juliano. 1997. Effects of a predator on prey metamorphosis: plastic

responses by prey or selective mortality? Ecology 78:838-851.

Hettyey, A., K. Vincze, S. Zsarnóczai, H. Hoi, and A. Laurila. 2011. Costs and benefits of

defences induced by predators differing in dangerousness. Journal of Evolutionary Biology 24:1007-1019.

Johansson, F. 2002. Reaction norms and production costs of predator-induced morphological

defences in a larval dragonfly (Leucorrhinia dubia: Odonata). Canadian Journal of Zoology 80:944-950.

Johansson, F., R. Stoks, L. Rowe, and M. De Block. 2001. Life history plasticity in a damselfly:

effects of combined time and biotic constraints. Ecology 82:1857-1869.

Kiesecker, J.M., D.P. Chivers, M. Anderson, and A.R. Blaustein. 2002. Effect of predator diet on

life history shifts of red-legged frogs, Rana aurora. Journal of Chemical Ecology 28:1007-1015.

Kraft, P.G., R.S. Wilson, and C.E. Franklin. 2005. Predator-mediated phenotypic plasticity in

tadpoles of the striped marsh frog, Limnodynastes peronii. Austral Ecology 30:558-663.

Lane, S.J., and M.J. Mahony. 2002. Larval anurans with synchronous and asynchronous

developmental periods: contrasting responses to water reduction and predator presence. Journal of Animal Ecology 71:780-792.

Lardner, B. 2000. Morphological and life history responses to predators in larvae of seven

anurans. Oikos 88:169-180.

Laurila, A., and J. Kujasalo. 1999. Habitat duration, predation risk and phenotypic plasticity in

common frog (Rana temporaria) tadpoles. Journal of Animal Ecology 68:1123-1132.

Laurila, A., J. Kujasalo, and E. Ranta. 1998. Predator-induced changes in life-history in two

anuran tadpoles: Effects of predator diet. Oikos 83:307-317.

Laurila, A., M. Järvi-Laturi, S. Pakkasmaa, and J. Merila. 2004. Temporal variation in predation

risk: Stage dependency, graded responses and fitness costs in tadpole antipredator defense. Oikos 90-99.