Ontogenetic Dietary Shifts and Food Resource Partitioning in a Stream-Dwelling Urodela Community: Mechanisms to Allow Coexistence Across Seasons

Javier Sánchez-Hernández, Albert Montori, Gustavo A. Llorente


Trophic interactions are important factors structuring animal communities. Although food resource partitioning among salamanders living in syntopy has often been explored in freshwater systems, the potential influence of ontogenetic shifts and seasonal patterns on resource partitioning is little known. The aim of this study was to explore the seasonal patterns in food resource partitioning among developmental stages of two Urodela species (Calotriton asper and Salamandra salamandra) in a fishless stream. We tested the hypothesis that seasonal patterns in intra- and interspecific food resource partitioning may occur at two different levels: (i) diet composition and (ii) prey size. For the purpose of this study, food resource partitioning was approached at three scales: (i) ontogenetic (larva, immature and adult), (ii) seasonal (winter, spring, summer and autumn) and (iii) species. Inter- and intraspecific competition for food was measured as dietary overlap according to Schoener’s index. The overlap in diet among developmental stages and species was high. However, prey size and the contribution of terrestrial invertebrates to the diets changed among developmental stages of species. This suggests that competition for food may be alleviated by ontogenetic dietary shifts in prey size and terrestrial invertebrates use.


allochthonous resources; aquatic-terrestrial coupling; diet overlap; prey size; trophic ontogeny

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Araujo M. S., Bolnick D. I., and Layman C. I. (2011), «The ecological causes of individual specialisation», Ecol. Lett., 14, 948 – 958.

Avery R. A. (1968), «Food and feeding relations of three species of Triturus (Amphibia, Urodela) during the aquatic phases», Oikos, 19, 408 – 412.

Balogová M., Miková E., Orendáš P., and Uhrin M. (2015), «Trophic spectrum of adult Salamandra salamandra in the Carpathians with the first note on food intake by the species during winter», Herpetol. Notes, 8, 371 – 377.

Bogdan H. V., Covaciu-Marcov S. D., Gaceu O., Cicort-Lucaciu A. S., Ferenţi S., and Sas-Kovács I. (2013), «How do we share food? Feeding of four amphibian species from an aquatic habitat in south-western Romania», Animal Biodivers. Conserv., 36, 89 – 99.

Bolnick D. I., Yang L. H., Fordyce J. A., Davis J. M., and Svanbäck R. (2002), «Measuring individual-level resource specialization», Ecology, 83, 2936 – 2941.

Bowman A. W. and Azzalini A. (1997), Applied Smoothing Techniques for Data Analysis. The Kernel Approach with S-Plus Illustrations, Oxford Univ. Press, Oxford.

Catherine A. T. (1985), «Resource Partitioning in Amphibians and Reptiles», Copeia, 1985(1), 1 – 21.

Chessel D., Dufour A. B., and Thioulouse J. (2004), «The ade4 package. I. One-table methods», R News, 4, 5 – 10.

Costa A., Baroni D., Romano A., and Salvidio S. (2017), «Individual diet variation in Salamandra salamandra larvae in a Mediterranean stream (Amphibia: Caudata)», Salamandra, 53, 148 – 152.

Cudmore W. W. and Bury R. B. (2014), «Resource partitioning in two stream salamanders, Dicamptodon tenebrosus and Rhyacotriton cascadae, in the Cascade Mountains, Oregon», Am. Midland Naturalist, 172, 191 – 199.

Degani G., Goldenberg S., and Warburg M. R. (1980), «Cannibalistic phenomena in Salamandra salamandra larvae in certain water bodies and under experimental conditions», Hydrobiologia, 75, 123 – 128.

Denoël M. (2004a), «Terrestrial versus aquatic foraging in juvenile Alpine newts (Triturus alpestris)», Ecoscience, 11, 404 – 409.

Denoël M. (2004b), «Feeding performance in heterochronic alpine newts is consistent with trophic niche and maintenance of polymorphism», Ethology, 110, 127 – 136.

Denoël M. and Andreone F. (2003), «Trophic habits and aquatic microhabitat use in gilled immature, paedomorphic and metamorphic Alpine newts (Triturus alpestris apuanus) in a pond in central Italy», Belg. J. Zool., 133, 95 – 102.

Denoël M. and Joly P. (2001), «Size-related predation reduces intramorph competition in paedomorphic alpine newts», Can. J. Zool., 79, 943 – 948.

Denoël M. and Schabetsberger R. (2003), «Resource partitioning in two heterochronic populations of Greek Alpine newts, Triturus alpestris veluchiensis», Acta Oecol., 24, 55 – 64.

Denoël M. and Winandy L. (2014), «Fire salamander (Salamandra salamandra) in Larzac plateau: low occurrence, pond-breeding and cohabitation of larvae with paedomorphic palmate newts (Lissotriton helveticus)», Acta Herpetol., 9, 43 – 49.

Farasat H. and Sharifi M. (2014), «Food habit of the endangered yellow-spotted newt Neurergus microspilotus (Caudata, Salamandridae) in Kavat Stream, western Iran», Zool. Stud., 53, 61.

Fasola M. (1993), «Resource partitioning by three species of newts during their aquatic phase», Ecography, 16, 73 – 81.

Fasola M. and Canova L. (1992), «Feeding habits of Triturus vulgaris, T. cristatus and T. alpestris (Amphibia, Urodela) in the northern Apennines (Italy)», Boll. Zool., 59, 273 – 280.

García-París M., Montori A., and Herrero P. (2004), «Amphibia. Lissamphibia», in: Fauna Ibérica. Vol. 24, Museo Nacional de Ciencias Naturales, CSIC.

Gillespie J. H. (2013), «Application of stable isotope analysis to study temporal changes in foraging ecology in a highly endangered amphibian», PLoS ONE, 8(1), e53041.

Gotelli N. J. and Ellison A. M. (2013), EcoSimR. Ver. 1.00, http://www.uvm.edu/~ngotelli/EcoSim/EcoSim.html

Griffiths R. A. (1986), «Feeding niche overlap and food selection in smooth and palmate newts, Triturus vulgaris and T. helveticus, at a pond in mid-wales», J. Animal Ecol., 55, 201 – 214.

Grutter A. S. (2000), «Ontogenetic variation in the diet of the cleaner fish Labroides dimidiatus and its ecological consequences», The Marine Ecol. Progr. Ser., 197, 241 – 246.

Guerrero F., Perez-Mellado V., Gil M. J., and Lizana M. (1990), «Food habits and trophic availability in the high mountain population of the spotted salamander from Spain (Salamandra salamandra almanzoris) (Caudata: Salamandridae)», Folia Zool., 89, 841 – 53.

Guillaume O. (2006), «Role of interspecific communication on spatial segregation in the sympatric salamanders Calotriton asper and Salamandra salamandra (Amphibia: Salamandridae)», Bull. Soc. Herpétol. Fr., 118, 5 – 16.

Jehle R., Bouma P., Sztatecsny M., and Arntzen J. W. (2000), «High aquatic niche overlap in the newts Triturus cristatus and T. marmoratus (Amphibia, Urodela)», Hydrobiologia, 437, 149 – 155.

Keitzer S. C. and Goforth R. R. (2013), «Spatial and seasonal variation in the ecological significance of nutrient recycling by larval salamanders in Appalachian headwater streams», Freshwater Sci., 2013, 1136 – 1147.

Kim S. L., Tinker M. T., Estes J. A., and Koch P. L. (2012). «Ontogenetic and among-individual variation in foraging strategies of northeast Pacific white sharks based on stable isotope analysis», PLoS ONE, 7(9), e45068.

Kopecký O., Vojar J., Šusta F., and Rehák I. (2012), «Composing and scaling of male and female alpine newt (Mesotriton alpestris) prey, with related site and seasonal effect», Ann. Zool. Fenn., 9, 231 – 239.

Krebs J. M. and Turingan R. G. (2003), «Intraspecific variation in gape-prey size relationships and feeding success during early ontogeny in red drum, Sciaenops ocellatus», Environm. Biol. Fishes, 66, 75 – 84.

Kronfeld-Schor N. and Dayan T. (2003), «Partitioning of time as an ecological resource», Ann. Rev. Ecol. Evol. Syst., 34, 153 – 181.

Kuzmin S. L. (1990), «Trophic niche overlap in syntopic postmetamorphic amphibians of the Carpathian Mountains (Ukraine, Soviet Union)», Herpetozoa, 3, 13 – 24.

Maerz J. C., Myers E. M., and Adams D. C. (2006), «Trophic polymorphism in a terrestrial salamander», Evol. Ecol. Res., 8, 23 – 35.

Milanovich J. R. and Hopton M. E. (2014), «Stoichiometry of a semi-aquatic plethodontid salamander: Intraspecific variation due to location, size and diet», Integr. Zool., 9, 613 – 622.

Montori A. (1988), Estudio sobre la biología y ecología del tritón pirenaico Euproctus asper (Dugès, 1852) en La Cerdanya. PhD thesis, University of Barcelona.

Montori A. (1991), «Alimentaciуn de los adultos de Euproctus asper (Dugès, 1852) en la montaña media del Prepirineo catalán (España)», Rev. Esp. Herpetol., 5, 23 – 36.

Montori A. (1992), «Alimentaciуn de las larvas de tritón pirenaico, Euproctus asper, en el prepirineo de la Cerdaña, España», Amphibia–Reptilia, 13, 157 – 167.

Montori A. (1997), «Trophic segregation between the Pyrenean Newt (Euproctus asper) and the Brown Trout (Salmo trutta fario)», in: W. Bohme, W. Bischoff, and T. Ziegler (eds.), Herpetologia Bonnensis, Societas Europaea Herpetologica, Bonn, Germany, pp. 273 – 278.

Montori A. and Llorente G. A. (2014), «Tritón pirenaico — Calotriton asper», in: A. Salvador and I. Martínez-Solano (eds.), Enciclopedia Virtual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid, pp. 1 – 28.

Montori A., Richter-Boix A., Franch M., Santos X., Garriga N., and Llorente G. A. (2012), «Natural fluctuations in a stream dwelling newt as a result of extreme rainfall: a 21-year survey of a Calotriton asper population», Basic Appl. Herpetol., 26, 43 – 56.

Nakazawa T. (2015), «Ontogenetic niche shifts matter in community ecology: a review and future perspectives», Popul. Ecol., 57, 347 – 354.

Nery T. and Schmera D. (2016), «The effects of top-down and bottom-up controls on macroinvertebrate assemblages in headwater streams», Hydrobiologia, 763, 173 – 181.

Petranka J. W. (1984), «Ontogeny of the diet and feeding behavior of Eurycea bislineata larvae», J. Herpetol., 18, 48 – 55.

Pianka E. R. (1981), «Competition and niche theory», in: R. M. May (ed.), Theoretical Ecology, Blackwell Sci. Publ., Oxford, pp. 167 – 196.

R Core Team (2015), R: A Language and Environment for Statistical Computing, R Foundation for Statistical Computing, Vienna.

Ranta E., Tjossem S. F., and Leikola N. (1987), «Female-male activity and zooplankton foraging by the smooth newt (Triturus vulgaris)», Ann. Zool. Fenn., 24, 79 – 88.

Roşca I., Gherghel I., Strugariu A., and Zamfirescu Ş. R. (2013), «Feeding ecology of two newt species (Triturus cristatus and Lissotriton vulgaris) during the reproduction season», Knowl. Manag. Aquatic Ecosyst., 408, 05.

Salvidio S., Oneto F., Ottonello D., Costa A., and Romano A. (2015), «Trophic specialization at the individual level in a terrestrial generalist salamander», Can. J. Zool., 93, 79 – 83.

Salvidio S., Romano A., Oneto F., Ottonello D., and Michelon R. (2012), «Different season, different strategies: Feeding ecology of two syntopic forest-dwelling salamanders», Acta Oecol., 43, 42 – 50.

Sánchez-Hernández J. (2016), «Do age-related changes in feeding habits of brown trout alter structural properties of food webs?» Aquatic Ecol., 50, 665 – 695.

Sánchez-Hernández J., Eloranta A. P., Finstad A. G., and Amundsen P.-A. (2017a), «Community structure affects trophic ontogeny in a predatory fish», Ecol. Evol., 7, 358 – 367.

Sánchez-Hernández J., Glaber H.-M., and Amundsen P.-A. (2016), «Food resource partitioning between stream-dwelling Arctic charr Salvelinus alpinus (L.), Atlantic salmon Salmo salar L. and alpine bullhead Cottus poecilopus Heckel, 1836: an example of water column segregation», Hydrobiologia, 783, 105 – 115.

Sánchez-Hernández J., Glaber H.-M., and Amundsen P.-A. (2017b), «Prey diversity as a driver of resource partitioning between river-dwelling fish species», Ecol. Evol., 7, 2058 – 2068.

Sánchez-Hernández J., Vieira-Lanero R., Servia M. J., and Cobo F. (2011), «Feeding habits of four sympatric fish species in the Iberian Peninsula: keys to understanding coexistence using prey traits», Hydrobiologia, 667, 119 – 132.

Schellekens T., de Roos A. M., and Persson L. (2010), «Ontogenetic diet shifts result in niche partitioning between two consumer species irrespective of competitive abilities», Am. Naturalist, 176, 625 – 637.

Schoener T. W. (1970), «Nonsynchronous spatial overlap of lizards in patchy habitats», Ecology, 51, 408 – 418.

Schoener T. W. (1974), «Resource partitioning in ecological communities», Science, 185, 27 – 39.

Schoener T. W. (1989), «The ecological niche», in: J. M. Cherrett (ed.), Ecological Concepts, Blackwell, Boston, pp. 79 – 107.

Schriever T. A. and William D. D. (2013), «Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient», Freshwater Biol., 58, 223 – 236.

Sih A., Crowley P., Mcpeek M., Petranka J., and Strohmeier K. (1985), «Predation, competition and prey communities: a review of field experiments», Ann. Rev. Ecol. Evol. Syst., 16, 269 – 311.

Thioulouse J., Chessel D., Dolédec S., and Olivier J. M. (1997), «ADE-4: a multivariate analysis and graphical display software», Statist. Comput., 7, 75 – 83.

Toft C. A. (1985), Resource partitioning in amphibians and reptiles», Copeia, 1985, 1 – 21.

Vandermeer J. H. (1972), «Niche theory», Ann. Rev. Ecol. Evol. Syst., 3, 107 – 132.

Vignoli L. and Luiselli L. (2012), «Dietary relationships among coexisting anuran amphibians: a worldwide quantitative review», Oecologia, 169, 499 – 509.

Vignoli L., Bissattini A. M., and Luiselli L. (2017), «Food partitioning and the evolution of non-randomly structured communities in tailed amphibians: a worldwide systematic review», Biol. J. Linn. Soc., 120, 489 – 502.

Vignoli L., Bologna M. A., and Luiselli L. (2007b), «Seasonal patterns of activity and community structure in an amphibian assemblage at a pond network with variable hydrology», Acta Oecol., 31, 185 – 192.

Vignoli L., Bombi P., D’Amen M., and Bologna M. A. (2007a), «Seasonal variation in the trophic niche of a heterochronic population of Triturus alpestris apuanus from the south-western Alps», Herpetol. J., 17, 183 – 191.

Vignoli L., Luiselli L., and Bologna M. A. (2009), «Dietary patterns and overlap in an amphibian assemblage at a pond in Mediterranean Central Italy», Vie Milieu, 59, 47 – 57.

Wallace R. K. (1981), «An assessment of diet-overlap indexes», Trans. Am. Fisher. Soc., 110, 72 – 76.

Wells K. D. (2007), The Ecology and Behaviour of Amphibians, Univ. of Chicago Press, Chicago.

Werner E. E. and Peacor S. D. (2003), «A review of trait-mediated indirect interactions in ecological communities», Ecology, 84, 1083 – 1100.

Zaccarelli N., Bolnick D. I., and Mancinelli G. (2013), «RInSp: an R package for the analysis of individual specialization in resource use», Meth. Ecol. Evol., 4, 1018 – 1023.

DOI: https://doi.org/10.30906/1026-2296-2019-26-3-135-149


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