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      Great Spotted Cuckoo Fledglings Often Receive Feedings from Other Magpie Adults than Their Foster Parents: Which Magpies Accept to Feed Foreign Cuckoo Fledglings?

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          Abstract

          Natural selection penalizes individuals that provide costly parental care to non-relatives. However, feedings to brood-parasitic fledglings by individuals other than their foster parents, although anecdotic, have been commonly observed, also in the great spotted cuckoo ( Clamator glandarius) – magpie ( Pica pica) system, but this behaviour has never been studied in depth. In a first experiment, we here show that great spotted cuckoo fledglings that were translocated to a distant territory managed to survive. This implies that obtaining food from foreign magpies is a frequent and efficient strategy used by great spotted cuckoo fledglings. A second experiment, in which we presented a stuffed-cuckoo fledgling in magpie territories, showed that adult magpies caring for magpie fledglings responded aggressively in most of the trials and never tried to feed the stuffed cuckoo, whereas magpies that were caring for cuckoo fledglings reacted rarely with aggressive behavior and were sometimes disposed to feed the stuffed cuckoo. In a third experiment we observed feedings to post-fledgling cuckoos by marked adult magpies belonging to four different possibilities with respect to breeding status (i.e. composition of the brood: only cuckoos, only magpies, mixed, or failed breeding attempt). All non-parental feeding events to cuckoos were provided by magpies that were caring only for cuckoo fledglings. These results strongly support the conclusion that cuckoo fledglings that abandon their foster parents get fed by other adult magpies that are currently caring for other cuckoo fledglings. These findings are crucial to understand the co-evolutionary arms race between brood parasites and their hosts because they show that the presence of the host's own nestlings for comparison is likely a key clue to favour the evolution of fledgling discrimination and provide new insights on several relevant points such as learning mechanisms and multiparasitism.

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          Most cited references12

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          Individual recognition: it is good to be different.

          Individual recognition (IR) behavior has been widely studied, uncovering spectacular recognition abilities across a range of taxa and modalities. Most studies of IR focus on the recognizer (receiver). These studies typically explore whether a species is capable of IR, the cues that are used for recognition and the specializations that receivers use to facilitate recognition. However, relatively little research has explored the other half of the communication equation: the individual being recognized (signaler). Provided there is a benefit to being accurately identified, signalers are expected to actively broadcast their identity with distinctive cues. Considering the prevalence of IR, there are probably widespread benefits associated with distinctiveness. As a result, selection for traits that reveal individual identity might represent an important and underappreciated selective force contributing to the evolution and maintenance of genetic polymorphisms.
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            Arms races between and within species.

            An adaptation in one lineage (e.g. predators) may change the selection pressure on another lineage (e.g. prey), giving rise to a counter-adaptation. If this occurs reciprocally, an unstable runaway escalation or 'arms race' may result. We discuss various factors which might give one side an advantage in an arms race. For example, a lineage under strong selection may out-evolve a weakly selected one (' the life-dinner principle'). We then classify arms races in two independent ways. They may be symmetric or asymmetric, and they may be interspecific or intraspecific. Our example of an asymmetric interspecific arms race is that between brood parasites and their hosts. The arms race concept may help to reduce the mystery of why cuckoo hosts are so good at detecting cuckoo eggs, but so bad at detecting cuckoo nestlings. The evolutionary contest between queen and worker ants over relative parental investment is a good example of an intraspecific asymmetric arms race. Such cases raise special problems because the participants share the same gene pool. Interspecific symmetric arms races are unlikely to be important, because competitors tend to diverge rather than escalate competitive adaptations. Intraspecific symmetric arms races, exemplified by adaptations for male-male competition, may underlie Cope's Rule and even the extinction of lineages. Finally we consider ways in which arms races can end. One lineage may drive the other to extinction; one may reach an optimum, thereby preventing the other from doing so; a particularly interesting possibility, exemplified by flower-bee coevolution, is that both sides may reach a mutual local optimum; lastly, arms races may have no stable and but may cycle continuously. We do not wish necessarily to suggest that all, or even most, evolutionary change results from arms races, but we do suggest that the arms race concept may help to resolve three long-standing questions in evolutionary theory.
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              Self-referent phenotype matching: theoretical considerations and empirical evidence.

              In most birds and mammals, young are raised in family groups. The phenotypes of nestmates and parents are thus reliable cues for recognition of conspecifics and kin. However, in some species, young develop alone, or in broods of mixed relatedness (e.g. because of multiple paternity or maternity), or among heterospecifics or unrelated conspecifics (brood parasites). Under these circumstances, the best referent (model) for discriminating close from distant kin and heterospecifics from conspecifics might be one's own self. This recognition process is known as self-referent phenotype matching. Here we review recent experimental evidence of self-referencing and suggest that behavioral neuroscience can provide new tools and insights into how it works (its proximate mechanistic and ontogenetic bases) and why it exists (its adaptive significance).
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                1 October 2014
                : 9
                : 10
                : e107412
                Affiliations
                [1 ]Departamento de Zoología, Facultad de Ciencias, Universidad de Granada, Granada, Spain
                [2 ]Grupo Coevolución, Unidad Asociada al Consejo Superior de Investigaciones Científicas (CSIC), Universidad de Granada, Granada, Spain
                [3 ]Department of Biology, Terrestrial Ecology Unit, Ghent University, Gent, Belgium
                University of Lausanne, Switzerland
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MS TPC LDN. Performed the experiments: MS TPC JDIA GR EMS LDN. Analyzed the data: MS TPC JDIA GR LDN. Contributed reagents/materials/analysis tools: MS TPC JDIA GR EMS LDN. Wrote the paper: MS TPC JDIA GR EMS LDN.

                Article
                PONE-D-14-10325
                10.1371/journal.pone.0107412
                4182665
                25272009
                989fa148-cd10-418b-9b53-ae42ac9b25ed
                Copyright @ 2014

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 11 March 2014
                : 14 August 2014
                Page count
                Pages: 10
                Funding
                This work was supported by the Spanish Ministerio de Economía y Competitividad/FEDER (research project CGL2011-25634/BOS). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Ecology
                Evolutionary Ecology
                Evolutionary Biology
                Psychology
                Behavior
                Parenting Behavior
                Zoology
                Animal Behavior
                Custom metadata
                The authors confirm that all data underlying the findings are fully available without restriction. All data are within the paper and its Supporting Information files.

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