Competition and response to predation risk: effects on morphological and behavioural defences in anuran larvae
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Abstract
Many freshwater organisms exhibit responses to conspecific density. Although a reduction in antipredator responses under high competition conditions is a well-documented pattern, most studies focus on species from temperate zones, in the case of anurans, primarily in those species that produce eggs in gelatinous masses or gelatinous strings. Research in subtropical regions remains limited. We investigated how competition intensity affects predation risk, shaping the morphological and behavioural defences in Physalaemus cristinae tadpoles, a prolonged breeder species that constructs foam nests. As a prolonged breeder, this species experiences sustained predation pressure over several months, influencing the dynamics of its intra- and interspecific interactions. Therefore, understanding this aspect of its life history is crucial for assessing the universality of this pattern across different ecological contexts. Initially, we assessed alterations in morphology, growth, and development. Variables included larval densities (4 and 8) and predator odour presence (with and without). We measured five morphological traits (body length, tail fin length, body depth, tail fin depth, tail fin muscle depth), growth, and development rates. Furthermore, we evaluated behavioural parameters: tadpole activity recorded before introducing predator odour and at 15, 30 and 45 min post-signal introduction, and tadpole location, calculated as the proportion of larvae in different areas based on the predator odour introduction point. At low densities, tadpoles exposed to predator odour displayed significant increases in all morphological traits compared to those not exposed. Conversely, at high densities, no differences were observed regarding predator odour. The results show that conspecific density can play a significant role in antipredator response. Tadpoles change morphology and behaviour in response to predator odour, but their response is dependent on tadpole density. The understanding of these complex interactions will enable the maintenance of long-term functionality and resilience in aquatic environments, which provide numerous ecosystem services.
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-
Acosta, N.R. (2010). Plasticidad fenotípica en la metamorfosis de larvas de Rhinella arenarum del Valle del Lerma, Salta. — Tesis doctoral, Universidad Nacional de La Plata, La Plata.
-
Addinsoft (2006). XLSTAT Ver. 7.5 for Excel interface. — Addinsoft, Pentraeth.
-
Albecker, M.A., Strobel, S.M.K. & Womack, M.C. (2023). Developmental plasticity in anurans: meta-analysis reveals effects of larval environments on size at metamorphosis and timing of metamorphosis. — Integr. Comp. Biol. 63: 714-729. DOI:10.1093/icb/icad059.
-
Armitage, K.B. (2004). Badger predation on yellow-bellied marmots. — Am. Nat. 151: 378-387.
-
Armúa de Reyes, C. & Estévez, A.L. (2005). Diversidad de heterópteros acuáticos, con especial referencia a las Belostoma (Heteroptera; Belostomatidae). — In: Temas de la biodiversidad del litoral fluvial argentino (Aceñolaza, F.G., ed.). INSUGEO, San Miguel de Tucumán, p. 281-292.
-
Asquith, C.M. & Vonesh, J.R. (2012). Effect of size and size structure on predation and inter-cohort competition in red-eyed treefrog tadpoles. — Oecologia 170: 629-639.
-
Bairos-Novak, K.R., Mitchell, M.D., Crane, A.L., Chivers, D.P. & Ferrary, M.C.O. (2017). Trust thy neighbour in times of trouble: background risk alters how tadpoles release and respond to disturbance cues. — Proc. Roy. Soc. Lond. B: Biol. Sci. 284: 20171465. DOI:10.1098/rspb.2017.1465.
-
Beckerman, A.P., Rodgers, G.M. & Dennis, S.R. (2010). The reaction norm of size and age at maturity under multiple predation risk. — J. Anim. Ecol. 79: 1069-1076.
-
Benard, M.F. (2004). Predator-induced phenotypic plasticity in organisms with complex life histories. — Annu. Rev. Ecol. Evol. Syst. 35: 651-673.
-
Bishop, D.C., Haas, C.A. & Mahoney, L.O. (2012). Response of Lithobates okaloosae, L. clamitans, and L. sphenocephala tadpoles to chemical cues of snake and fish predators. — Fla. Sci. 75: 1-10.
-
Bolnick, D.I. & Preisser, E.L. (2005). Resource competition modifies the strength of trait-mediated predator-prey interactions: a meta-analysis. — Ecology 86: 2771-2779.
-
Bouchard, S.S., O’Leary, C.J., Wargelin, L.J., Rodriguez, W.B., Jennings, K.X. & Warkentin, K.M. (2015). Alternative Competition-induced digestive strategies yield equal growth, but constrain compensatory growth in red-eyed Treefrog Larvae. — J. Exp. Zool. 323: 778-788.
-
Bourdeau, P.E. (2010). An inducible morphological defence is a passive by-product of behaviour in a marine snail. — Proc. Roy. Soc. Lond. B: Biol. Sci. 277: 455-462. DOI:10.1098/rspb.2009.1295.
-
Brodie, E.D. & Formanowicz Jr, D.R. (1987). Antipredator mechanisms of larval anurans: protection of palatable individuals. — Herpetologica 43: 369-373.
-
Bronmark, C. & Miner, J.G. (1992). Predator-induced phenotypic change in body morphology in crucian carp. — Science 258: 1348-1350. DOI:10.1126/science.258.5086.1348.
-
Caro, T. (2005). Antipredator defenses in birds and mammals. — The University of Chicago Press, Chicago, IL.
-
Chalcraft, D.R. & Resetarits, W.J. (2003). Predator identity and ecological impacts: functional redundancy or functional diversity. — Ecology 84: 2407-2418.
-
Crane, A.L. & Ferrari, M.C.O. (2015). Minnows trust conspecifics more than themselves when faced with conflicting information about predation risk. — Anim. Behav. 100: 184-190. DOI:10.1016/j.anbehav.2014.12.002.
-
Darlington, R.B. & Smulders, T.V. (2001). Problems with residuals analysis. — Anim. Behav. 62: 599-602.
-
Davenport, J.M., Seiwert, P.A., Fishback, L.A. & Cash, W.B. (2016). The Interactive effects of fish predation and conspecific density on survival and growth of tadpoles of Rana sylvatica in a subarctic wetland. — Copeia 104: 639-644.
-
Di Rienzo, J.A., Casanoves, F., Balzarini, M.G., Gonzalez, L., Tablada, M. & Robledo, C.W. (2020). Centro de Transferencia InfoStat. — FCA, Universidad Nacional de Córdoba, Cordoba. Available online at http://www.infostat.com.ar
-
Donelan, S.C., Grabowski, J.H. & Trussell, G.C. (2017). Refuge quality impacts the strength of nonconsumptive effects on prey. — Ecology 98: 403-411. DOI:10.1002/ecy.1647.
-
Duggan, I.C., Green, D.J. & Shiel, R.J. (2002). Distribution of rotifer assemblages in North Island, New Zealand, lakes: relationships to environmental and historical factors. — Freshw. Biol. 47: 195-206.
-
Elvidge, C.K., Macnaughton, C.J. & Brown, G.E. (2013). Sensory complementation and antipredator behavioural compensation in acid-impacted juvenile Atlantic salmon. — Oecologia 172: 69-78.
-
Feminella, J.W. & Hawkins, C.P. (1994). Tailed frog tadpoles differentially alter their feeding behavior in response to non-visual cues from four predators. — J. North Am. Benthol. Soc. 13: 310-320.
-
Fernandez, J.M.C. & Oliveira, M.Z.T. (2017). Predation and schooling influence on the primary response on individual of Rhinella ornata (Spix, 1824) (Anura: Bufonidae): an experimental assessment of habitat selection. — S. Am. J. Herpetol. 12: 57-60.
-
Ferrari, M.C.O., Elvidge, C.K., Jackson, C.D., Chivers, D.P. & Brown, G.E. (2010). The responses of prey fish to temporal variation in predation risk: sensory habituation or risk assessment? — Behav. Ecol. 21: 532-536.
-
Frommer, J.G., Hiermes, M. & Bakker, T.C. (2009). Disentangling the effects of group size and density on shoaling decisions of three-spined sticklebacks (Gasterosteus aculeatus). — Behav. Ecol. Sociol. 63: 1141-1148. DOI:10.1007/s00265-009-0767-9.
-
Gómez, V.I. (2019). The influence of tadpole density and predation on the behavioral responses of two Neotropical anurans. — Phyllomedusa 18: 293-298.
-
Gómez, V.I. (2023). Efecto de la temperatura sobre los estadios de vida iniciales de larvas de Trachycephalus typhonius y Scinax nasicus (Anura: Hylidae). — Ecol. Austral. 33: 89-94.
-
Gómez, V.I. & Kehr, A.I. (2011). Morphological and developmental responses of anuran larvae (Physalaemus albonotatus) to chemical cues from the predators Moenkhausia dichoroura (Characiformes: Characidae) and Belostoma elongatum (Hemiptera: Belostomatidae). — Zool. Stud. 50: 203-210.
-
Gómez, V.I. & Kehr, A.I. (2012). The effect of chemical signal of predatory fish and water bug on the morphology and development of Elachistocleis bicolor tadpoles (Anura: Microhylidae). — Biologia 67: 1-6.
-
Gómez, V.I. & Kehr, A.I. (2013). Interaction between competitors and predators and its effects on morphological and behavioural defenses in Scinax nasicus tadpoles. — Behaviour 150: 921-937.
-
Gómez, V.I. & Kehr, A.I. (2019). Effect of soil disturbance by agricultural activities on the life history traits of monkey frog (Pithecopus azureus). — Environ. Monit. Assess. 19: 608. DOI:10.1007/s10661-019-7663-1.
-
Gosner, K.L. (1960). A simplified table for staging anurans embryos and larvae with notes of identification. — Herpetologica 16: 183-190.
-
Guariento, R.D. & Esteves, F.A. (2013). The several faces of fear: ecological consequences of predation risk in a lagoon model system. — Acta Limnol. Bras. 25: 211-223.
-
Guariento, R.D., Carneiro, L.S., Estevesz, S.A., Jorge, J.S. & Caliman, A. (2015). Conspecific density affects predator-induced prey phenotypic plasticity. — Ecosphere 6: 106. DOI:10.1890/ES15-00142.1.
-
Hammer, T.L., Bize, P., Gineste, B., Robin, J.P., Groscolas, R. & Viblanc, V.A. (2023). Disentangling the “many eyes”, “dilution effect”, “selfish herd”, and “distracted prey” hypotheses in shaping alert and flight initiation distance in a colonial sea bird. — Behav. Processes 210: 104919. DOI:10.1016/j.beproc.2023.104919.
-
Hangartner, S., Laurila, A. & Räsänen, K. (2011). Adaptive divergence of the moor frog (Rana arvalis) along an acidification gradient. — BMC Evol. Biol. 11: 366. DOI:10.1186/1471-2148-11-366.
-
Hoverman, J.T., Auld, J.R. & Relyea, R.A. (2005). Putting prey back together again: integrating predator-induced behavior, morphology, and life history. — Oecologia 144: 481-491.
-
Jara, F.G. & Perotti, M.G. (2010). Risk of predation and behavioral response in three anuran species: influence of tadpole size and predator type. — Hydrobiologia 644: 313-324.
-
Johansson, F., Stoks, R., Rowe, L. & De Block, M. (2001). Life history plasticity in a damselfly: Effects of combined time and biotic constraints. — Ecology 82: 1857-1869. DOI:10.1890/0012-9658(2001)082[1857:LHPIAD]2.0.CO;2.
-
Kehr, A.I. (1987). Crecimiento individual en larvas de Hyla pulchella pulchella y Bufo fernandezae en condiciones controladas de coexistencia y densidad (Amphibia Anura). — Cuad. Herpetol. 3: 1-6.
-
Kloskowski, J. (2018). Total non-consumptive effects of fish on Pelobates fuscus and Hyla orientalis tadpoles in pond enclosure experiments. — Zool. Sci. 35: 528-534.
-
Kotler, B.P., Brown, J.S., Slotow, R.H., Goodfriend, W.L. & Strauss, M. (1993). The influence of snakes on the foraging behaviour of gerbils. — Oikos 67: 309-316.
-
Lass, S. & Spaak, P. (2003). Chemically induced anti-predator defences in plankton: A review. — Hydrobiologia 491: 221-239. DOI:10.1023/A:1024487804497.
-
Lima, S.L. (1995). Back to the basics of anti-predatory vigilance: the group-size effect. — Anim. Behav. 49: 11-20. DOI:10.1016/0003-3472(95)80149-9.
-
Lima, S.L. & Bednekoff, P.A. (1999). Temporal variation in danger drives antipredator behavior: The predation risk allocation hypothesis. — Am. Nat. 153: 649-659. DOI:10.1086/303202.
-
Lima, S.L. & Dill, L.M. (1990). Behavioral decisions made under the risk of predation: a review and prospectus. — Can. J. Zool. 68: 619-640. DOI:10.1139/z90-092.
-
Lima, S.L. & Steury, T.D. (2005). Perception of predation risk: the foundation of nonlethal predator-prey interactions. — In: Ecology of predator-prey interactions (Barbosa, P. & Castellanos, I., eds). Oxford University Press, Oxford, p. 166-188.
-
Lucon-Xiccato, T., Chivers, D.P., Mitchell, M.D. & Ferrari, M.C.O. (2016). Making the dead talk: alarm cue mediated antipredator behaviour and learning are enhanced when injured conspecifics experience high predation risk. — Biol. Lett. 12: 20160560. DOI:10.1098/rsbl.2016.0560.
-
Ludwig, D. & Rowe, L. (1990). Life-history strategies for energy gain and predator avoidance under time constraints. — Am. Nat. 135: 686-707.
-
Nagano, M. & Yoshida, T. (2020). Size-selective predation accounts for intra- and inter-specific variation of inducible morphological defense of Daphnia. — Ecosphere 11: 1-13.
-
Neptune, T.C. & Benard, M.F. (2024). Longer days, larger grays: carryover effects of photoperiod and temperature in gray treefrogs, Hyla versicolor. — Proc. Roy. Soc. Lond. B: Biol. Sci. 291: 20241336.
-
Pangle, K.L. & Peacor, S.D. (2006). Nonlethal effect of the invasive predator Bythotrephes longimanus on Daphnia mendotae. — Freshw. Biol. 51: 1070-1078. DOI:10.1111/j.1365-2427.2006.01555.x.
-
Pangle, K.L., Peacor, S.D. & Johannsson, O.E. (2007). Large nonlethal effects of an invasive invertebrate predator on zooplankton population growth rate. — Ecology 88: 402-412. DOI:10.1890/06-0768.
-
Peacor, S.D. (2003). Phenotypic modifications to conspecific density arising from predation risk assessment. — Oikos 100: 409-415.
-
Peacor, S.D. & Werner, E.E. (1997). Trait-mediated indirect interactions in a simple aquatic food web. — Ecology 78: 1146-1156.
-
Peacor, S.D. & Werner, E.E. (2001). The contribution of trait-mediated indirect effects to the net effects of a predator. — Proc. Natl. Acad. Sci. USA 98: 3904-3908. DOI:10.1073/pnas.071061998.
-
Relyea, R.A. (2003). How prey respond to combined predators: A review and an empirical test. — Ecology 84: 1827-1839. DOI:10.1890/0012-9658(2003)084[1827:HPRTCP]2.0.CO;2.
-
Relyea, R.A. (2004). Fine-tuned phenotypes: tadpole plasticity under 16 combinations of predators and competitors. — Ecology 85: 172-179. DOI:10.1890/03-0169.
-
Relyea, R.A. & Auld, J.R. (2004). Having the guts to compete: how intestinal plasticity explains costs of inducible defences. — Ecol. Lett. 7: 869-875.
-
Relyea, R.A. & Rosenberger, D. (2018). Predator effects on metamorphosis: the effects of scaring versus thinning at high prey densities. — Copeia 106: 457-467.
-
Relyea, R.A., Stephens, P.R. & Hammond, J.I. (2021). Phylogenetic patterns of trait and trait plasticity evolution: Insights from tadpoles. — Evolution 75: 2568-2588. DOI:10.1111/evo.14338.
-
Ren, C., Teng, Y., Shen, Y., Yao, Q. & Wang, H. (2021). Altered temperature affect body condition and endochondral ossification in Bufo gargarizans tadpoles. — J. Therm. Biol. 99: 103020. DOI:10.1016/j.jtherbio.2021.103020.
-
Resetarits, W.J., Rieger, J.F. & Binckley, C.A. (2004). Threat of predation negates density effects in larval Gray Treefrogs. — Oecologia 138: 532-538.
-
Reuben, P.L. & Touchon, J.C. (2021). Nothing as it seems: behavioral plasticity appears correlated with morphology and colour, but is not in a Neotropical tadpole. — Proc. Roy. Soc. Lond. B: Biol. Sci. 288: 20210246. DOI:10.1098/rspb.2021.0246.
-
Reznick, D.N., Bryga, H. & Endler, J.A. (1990). Experimentally induced life-history evolution in a natural population. — Nature 346: 357-359. DOI:10.1038/346357a0.
-
Reznick, D.N., Butler, M.J. & Rood, H. (2001). Life-history evolution in guppies. VII. The comparative ecology of high- and low- predation environments. — Am. Nat. 157: 126-140. DOI:10.1086/318627.
-
Ruthsatz, K., Peck, M.A., Dausmann, K.H., Sabatino, N.M. & Glos, J. (2018). Patterns of temperature induced developmental plasticity in anuran larvae. — J. Therm. Biol. 74: 123-132. DOI:10.1016/j.jtherbio.2018.03.005.
-
Schoeppner, N.M. & Relyea, R.A. (2009). Interpreting the smells of predation: How alarm cues and kairomones induce different prey defences. — Funct. Ecol. 23: 1114-1121. DOI:10.1111/j.1365-2435.2009.01578.x.
-
Sheriff, M.J., Peacor, S.D., Hawlena, D. & Traker, M. (2020). Non-consumptive predator effects on prey population size: a dearth of evidence. — J. Anim. Ecol. 89: 1302-1316.
-
Sih, A. & McCarthy, T.M. (2002). Rey responses to pulses or risk and safety: testing the risk allocation hypothesis. — Anim. Behav. 63: 437-443.
-
Skelly, D.K. & Werner, E.E. (1990). Behavioral and life-historical responses of larval American toads to an odonate predator. — Ecology 71: 2313-2322. DOI:10.2307/1938642.
-
Smith, G.R. & Awan, A.R. (2009). The roles of predator identity and group size in the antipredator responses of American toad (Bufo americanus) and bullfrog (Rana catesbeiana) tadpoles. — Behaviour 146: 225-243.
-
Smith, J.A., Donadio, E., Pauli, J.N., Sheriff, M.J. & Middleton, A.D. (2019). Integrating temporal refugia into landscapes of fear: prey exploit predator downtimes to forage in risky places. — Oecologia 189: 883-890. DOI:10.1007/s00442-019-04381-5.
-
Spieler, M. (2005). Can aggregation behavior of Phrynomantis microps tadpoles reduce predation risk? — Herpetol. J. 15: 153-157.
-
SPSS (1997). SYSTAT 7.5 for Windows. — SPSS, Chicago, IL.
-
Stoks, R., De Block, M., Van de Meutter, F. & Johansson, F. (2005). Predation cost of rapid growth: behavioural coupling and physiological decoupling. — J. Anim. Ecol. 74: 708-715. DOI:10.1111/j.1365-2656.2005.00969.x.
-
Stoks, R., De Block, M., Slos, S., Van Doorslaer, W. & Rolff, J. (2006). Time constraints mediate predator-induced plasticity in immune function, condition, and life history. — Ecology 87: 809-815. DOI:10.1890/0012-9658(2006)87[809:TCMPPI]2.0.CO;2.
-
Teplitsky, C. & Laurila, A. (2007). Flexible defense strategies: competition modifies investment in behavioral vs. morphological defenses. — Ecology 88: 1641-1646.
-
Thaker, M., Vanak, A.T., Owen, C., Ogden, M. & Slotow, R. (2010). Group dynamics of zebra and wildebeest in a woodland savanna: effects of predation risk and habitat density. — PLoS ONE 5: e12758. DOI:10.1371/journal.pone.0012758.
-
Thaker, M., Vanak, A.T., Owen, C., Ogden, M., Neimann, S. & Slotow, R. (2011). Minimizing predation risk in a landscape of multiple predators: effects on the spatial distribution of African ungulates. — Ecology 92: 398-407. DOI:10.1890/10-0126.1.
-
Tollrian, R. (1995). Chaoborus crystallinus predation on Daphnia pulex: Can induced morphological changes balance effects of body size on vulnerability? — Oecologia 101: 151-155.
-
Touchon, J.C. & Vonesh, J.R. (2016). Variation in abundance and efficacy of tadpole predators in a neotropical pond community. — J. Herpetol. 50: 113-119.
-
Turner, A.M. (2004). Non-lethal effects of predators on prey growth rates depend on prey density and nutrient additions. — Oikos 104: 561-569.
-
Turner, A.M. & Montgomery, S.L. (2003). Spatial and temporal scales of predator avoidance: experiments with fish and snails. — Ecology 84: 616-622. DOI:10.1890/0012-9658(2003)084[0616:SATSOP]2.0.CO;2.
-
Vavrek, M.A. & Brown, G.E. (2009). Threat-sensitiv responses to disturbance cues in juvenile convict cichlids and rainbow trout. — Ann. Zool. Fenn. 46: 171-180. DOI:10.5735/086.046.0302.
-
Vonesh, J.R. & Warkentin, K.M. (2006). Opposite shifts in size at metamorphosis in response to larval and metamorph predators. — Ecology 87: 556-562. DOI:10.1890/05-0930.
-
Webster, M.M. & Laland, K.N. (2008). Social learning strategies and predation risk: minnows copy only when using private information would be costly. — Proc. Roy. Soc. Lond. B: Biol. Sci. 275: 2869-2876. DOI:10.1098/rspb.2008.0817.
-
Wells, K.D. (2007). The ecology and behavior of amphibians. — The University of Chicago Press, Chicago, IL.
-
Wilbur, H.M. (1997). Experimental ecology of food webs: complex systems in temporary ponds. — Ecology 78: 2279-2302.
-
Wojdak, J.M. & Luttbeg, B. (2005). Relative strengths of trait-mediated and density-mediated indirect effects of a predator vary with resource levels in a freshwater food chain. — Oikos 111: 592-598.
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Competition and response to predation risk: effects on morphological and behavioural defences in anuran larvae
In: BehaviourSearch for other papers by Valeria I. Gómez in
Current site
Google Scholar
PubMed
Search for other papers by Cynthya E. González in
Current site
Google Scholar
PubMed
Search for other papers by Marta I. Duré in
Current site
Google Scholar
PubMed
Abstract
Many freshwater organisms exhibit responses to conspecific density. Although a reduction in antipredator responses under high competition conditions is a well-documented pattern, most studies focus on species from temperate zones, in the case of anurans, primarily in those species that produce eggs in gelatinous masses or gelatinous strings. Research in subtropical regions remains limited. We investigated how competition intensity affects predation risk, shaping the morphological and behavioural defences in Physalaemus cristinae tadpoles, a prolonged breeder species that constructs foam nests. As a prolonged breeder, this species experiences sustained predation pressure over several months, influencing the dynamics of its intra- and interspecific interactions. Therefore, understanding this aspect of its life history is crucial for assessing the universality of this pattern across different ecological contexts. Initially, we assessed alterations in morphology, growth, and development. Variables included larval densities (4 and 8) and predator odour presence (with and without). We measured five morphological traits (body length, tail fin length, body depth, tail fin depth, tail fin muscle depth), growth, and development rates. Furthermore, we evaluated behavioural parameters: tadpole activity recorded before introducing predator odour and at 15, 30 and 45 min post-signal introduction, and tadpole location, calculated as the proportion of larvae in different areas based on the predator odour introduction point. At low densities, tadpoles exposed to predator odour displayed significant increases in all morphological traits compared to those not exposed. Conversely, at high densities, no differences were observed regarding predator odour. The results show that conspecific density can play a significant role in antipredator response. Tadpoles change morphology and behaviour in response to predator odour, but their response is dependent on tadpole density. The understanding of these complex interactions will enable the maintenance of long-term functionality and resilience in aquatic environments, which provide numerous ecosystem services.
| All Time | Past 365 days | Past 30 Days | |
|---|---|---|---|
| Abstract Views | 510 | 188 | 16 |
| Full Text Views | 23 | 11 | 0 |
| PDF Views & Downloads | 48 | 30 | 0 |
