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P. Lloyd

Publications and source records attributed to P. Lloyd.

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Postnatal growth rates covary weakly with embryonic development rates and do not explain adult mortality probability among songbirds on four continents

Growth and development rates may result from genetic programming of intrinsic processes that yield correlated rates between life stages. These intrinsic rates are thought to affect adult mortality probability and longevity. However, if proximate extrinsic factors (e.g., temperature, food) influence development rates differently between stages and yield low covariance between stages, then development rates may not explain adult mortality probability. We examined these issues based on study of 90 songbird species on four continents to capture the diverse life-history strategies observed across geographic space. The length of the embryonic period explained little variation (ca. 13%) in nestling periods and growth rates among species. This low covariance suggests that the relative importance of intrinsic and extrinsic influences on growth and development rates differs between stages. Consequently, nestling period durations and nestling growth rates were not related to annual adult mortality probability among diverse songbird species within or among sites. The absence of a clear effect of faster growth on adult mortality when examined in an evolutionary framework across species may indicate that species that evolve faster growth also evolve physiological mechanisms for ameliorating costs on adult mortality. Instead, adult mortality rates of species in the wild may be determined more strongly by extrinsic environmental causes.

American Naturalist

Females increase reproductive investment in response to helper-mediated improvements in allo-feeding, nest survival, nestling provisioning and post-fledging survival in the Karoo scrub-robin Cercotrichas coryphaeus

In many cooperatively-breeding species, the presence of one or more helpers improves the reproductive performance of the breeding pair receiving help. Helper contributions can take many different forms, including allo-feeding, offspring provisioning, and offspring guarding or defence. Yet, most studies have focussed on single forms of helper contribution, particularly offspring provisioning, and few have evaluated the relative importance of a broader range of helper contributions to group reproductive performance. We examined helper contributions to multiple components of breeding performance in the Karoo scrub-robin Cercotrichas coryphaeus, a facultative cooperative breeder. We also tested a prediction of increased female investment in reproduction when helpers improve conditions for rearing young. Helpers assisted the breeding male in allo-feeding the incubating female, increasing allo-feeding rates. Greater allo-feeding correlated with greater female nest attentiveness during incubation. Nest predation was substantially lower among pairs breeding with a helper, resulting in a 74% increase in the probability of nest survival. Helper contributions to offspring provisioning increased nestling feeding rates, resulting in a reduced incidence of nestling starvation and increased nestling mass. Nestling mass had a strong, positive effect on post-fledging survival. Controlling for female age and habitat effects, annual production of fledged young was 130% greater among pairs breeding with a helper, and was influenced most strongly by helper correlates with nest survival, despite important helper effects on offspring provisioning. Females breeding with a helper increased clutch size, supporting the prediction of increased female investment in reproduction in response to helper benefits. ?? 2009 J. Avian Biol.

Journal of Avian Biology

Linking demographic effects of habitat fragmentation across landscapes to continental source-sink dynamics

Forest fragmentation may cause increased brood parasitism and nest predation of breeding birds. In North America, nest parasitism and predation are expected to increase closer to forest edges because the brood-parasitic Brown-headed Cowbird (Molothrus ater) and generalist nest predators often enter the forest from adjoining developed (largely agricultural) habitats. Yet the abundance of brood parasites and nest predators at the patch scale may be strongly constrained by the total area of developed habitat at landscape scales. The scale and extent of landscape effects are unclear, however, because past studies were mostly conducted within local landscapes rather than across independent landscapes. We report replicated studies from 30 independent landscapes across 17 states of the United States that show that nest parasitism is strongly affected by fragmentation at a 20 km radius scale, equivalent to the maximum foraging range of cowbirds. Nest predation is influenced by both edge and landscape effects, and increases with fragmentation at a 10 km radius scale. Predation is additive to parasitism mortality, and the two together yield decreased population growth potential with increasing forest fragmentation at a 10 km radius scale for 20 of 22 bird species. Mapping of population growth potential across continental landscapes displays broad impacts of fragmentation on population viability and allows geographic prioritization for conservation.

Ecological Applications