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      Optimal Resource Allocation to Survival and Reproduction in Parasitic Wasps Foraging in Fragmented Habitats

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          Abstract

          Expansion and intensification of human land use represents the major cause of habitat fragmentation. Such fragmentation can have dramatic consequences on species richness and trophic interactions within food webs. Although the associated ecological consequences have been studied by several authors, the evolutionary effects on interacting species have received little research attention. Using a genetic algorithm, we quantified how habitat fragmentation and environmental variability affect the optimal reproductive strategies of parasitic wasps foraging for hosts. As observed in real animal species, the model is based on the existence of a negative trade-off between survival and reproduction resulting from competitive allocation of resources to either somatic maintenance or egg production. We also asked to what degree plasticity along this trade-off would be optimal, when plasticity is costly. We found that habitat fragmentation can indeed have strong effects on the reproductive strategies adopted by parasitoids. With increasing habitat fragmentation animals should invest in greater longevity with lower fecundity; yet, especially in unpredictable environments, some level of phenotypic plasticity should be selected for. Other consequences in terms of learning ability of foraging animals were also observed. The evolutionary consequences of these results are discussed.

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          Most cited references111

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          Ecological and Evolutionary Responses to Recent Climate Change

          Ecological changes in the phenology and distribution of plants and animals are occurring in all well-studied marine, freshwater, and terrestrial groups. These observed changes are heavily biased in the directions predicted from global warming and have been linked to local or regional climate change through correlations between climate and biological variation, field and laboratory experiments, and physiological research. Range-restricted species, particularly polar and mountaintop species, show severe range contractions and have been the first groups in which entire species have gone extinct due to recent climate change. Tropical coral reefs and amphibians have been most negatively affected. Predator-prey and plant-insect interactions have been disrupted when interacting species have responded differently to warming. Evolutionary adaptations to warmer conditions have occurred in the interiors of species' ranges, and resource use and dispersal have evolved rapidly at expanding range margins. Observed genetic shifts modulate local effects of climate change, but there is little evidence that they will mitigate negative effects at the species level.
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            Confounding factors in the detection of species responses to habitat fragmentation.

            Habitat loss has pervasive and disruptive impacts on biodiversity in habitat remnants. The magnitude of the ecological impacts of habitat loss can be exacerbated by the spatial arrangement -- or fragmentation -- of remaining habitat. Fragmentation per se is a landscape-level phenomenon in which species that survive in habitat remnants are confronted with a modified environment of reduced area, increased isolation and novel ecological boundaries. The implications of this for individual organisms are many and varied, because species with differing life history strategies are differentially affected by habitat fragmentation. Here, we review the extensive literature on species responses to habitat fragmentation, and detail the numerous ways in which confounding factors have either masked the detection, or prevented the manifestation, of predicted fragmentation effects. Large numbers of empirical studies continue to document changes in species richness with decreasing habitat area, with positive, negative and no relationships regularly reported. The debate surrounding such widely contrasting results is beginning to be resolved by findings that the expected positive species-area relationship can be masked by matrix-derived spatial subsidies of resources to fragment-dwelling species and by the invasion of matrix-dwelling species into habitat edges. Significant advances have been made recently in our understanding of how species interactions are altered at habitat edges as a result of these changes. Interestingly, changes in biotic and abiotic parameters at edges also make ecological processes more variable than in habitat interiors. Individuals are more likely to encounter habitat edges in fragments with convoluted shapes, leading to increased turnover and variability in population size than in fragments that are compact in shape. Habitat isolation in both space and time disrupts species distribution patterns, with consequent effects on metapopulation dynamics and the genetic structure of fragment-dwelling populations. Again, the matrix habitat is a strong determinant of fragmentation effects within remnants because of its role in regulating dispersal and dispersal-related mortality, the provision of spatial subsidies and the potential mediation of edge-related microclimatic gradients. We show that confounding factors can mask many fragmentation effects. For instance, there are multiple ways in which species traits like trophic level, dispersal ability and degree of habitat specialisation influence species-level responses. The temporal scale of investigation may have a strong influence on the results of a study, with short-term crowding effects eventually giving way to long-term extinction debts. Moreover, many fragmentation effects like changes in genetic, morphological or behavioural traits of species require time to appear. By contrast, synergistic interactions of fragmentation with climate change, human-altered disturbance regimes, species interactions and other drivers of population decline may magnify the impacts of fragmentation. To conclude, we emphasise that anthropogenic fragmentation is a recent phenomenon in evolutionary time and suggest that the final, long-term impacts of habitat fragmentation may not yet have shown themselves.
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              Habitat fragmentation, species loss, and biological control.

              Fragmentation of habitats in the agricultural landscape is a major threat to biological diversity, which is greatly determined by insects. Isolation of habitat fragments resulted in decreased numbers of species as well as reduced effects of natural enemies. Manually established islands of red clover were colonized by most available herbivore species but few parasitoid species. Thus, herbivores were greatly released from parasitism, experiencing only 19 to 60 percent of the parasitism of nonisolated populations. Species failing to successfully colonize isolated islands were characterized by small and highly variable populations. Accordingly, lack of habitat connectivity released insects from predator control.
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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
                2012
                6 June 2012
                : 7
                : 6
                : e38227
                Affiliations
                [1 ]INRA, Sophia Antipolis Cedex, France
                [2 ]Netherlands Institute of Ecology (NIOO-KNAW), Wageningen, The Netherlands
                [3 ]Institute of Ecology, FB 2, Biology, University of Bremen, Bremen, Germany
                California State University Fullerton, United States of America
                Author notes

                Conceived and designed the experiments: EW PC LEMV TH. Analyzed the data: EW PC. Wrote the paper: EW PC. Conceived the original idea: EW TH LEMV. Designed the software used to produce the results: PC EW.

                Article
                PONE-D-12-03441
                10.1371/journal.pone.0038227
                3368906
                22701614
                82ffb86f-6e92-49a7-9b72-a346415f1e97
                Wajnberg et al. 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
                : 1 February 2012
                : 6 May 2012
                Page count
                Pages: 9
                Categories
                Research Article
                Biology
                Ecology
                Community Ecology
                Food Web Structure
                Species Interactions
                Behavioral Ecology
                Population Ecology
                Theoretical Ecology
                Evolutionary Biology
                Behavioral Ecology
                Evolutionary Ecology
                Zoology
                Entomology
                Computer Science
                Computerized Simulations

                Uncategorized
                Uncategorized

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