Strong contribution of immigration to local population regulation: evidence from a migratory passerine.

A mechanistic understanding of the dynamics of populations requires knowledge about the variation of the underlying demographic rates and about the reasons for their variability. In geographically open populations, immigration is often necessary to prevent declines, but little is known about whether immigration can contribute to its regulation. We studied the dynamics of a Red-backed Shrike population (Lanius collurio) over 36 years in Germany with a Bayesian integrated population model. We estimated mean and temporal variability of population sizes, productivity, apparent survival, and immigration. We assessed how strongly the demographic rates were correlated with population growth to understand the demographic reasons of population change and how strongly the demographic rates were correlated with population size to identify possible density-dependent mechanisms. The shrike population varied between 35 and 74 breeding pairs but did not show a significant trend in population size over time (growth rate 1.002 +/- 0.001 [mean +/- SD]). Apparent survival of females (juveniles 0.06 +/- 0.01; adults 0.37 +/- 0.03) was lower than that of males (juveniles 0.10 +/- 0.01; adults 0.44 +/- 0.02). Immigration rates were substantial and higher in females (0.56 +/- 0.02) than in males (0.43 +/- 0.02), and average productivity was 2.76 +/- 0.14. Without immigration, the Red-backed Shrike population would have declined strongly. Immigration was the strongest driver for the number of females while local recruitment was the most important driver for the number of males. Immigration of both sexes and productivity, but not local recruitment and survival, were subject to density dependence. Density-dependent productivity was not effectively regulating the local population but may have contributed to regulate shrike populations at larger spatial scales. These findings suggest that immigration is not only an important component to prevent a geographically open population from decline, but that it can also contribute to its regulation.

Demographic response to environmental variation in breeding, stopover and non-breeding areas in a migratory passerine.

Demographic rates of migratory species passing through several areas during their annual cycle may be affected by environmental conditions at each of these areas. Recent studies provide evidence that their impact is not necessarily immediate, but can be delayed. We studied survival, reproductive success and arrival date at the breeding grounds of red-backed shrikes Lanius collurio, a trans-Saharan migrant, in relation to weather and vegetation on the breeding grounds, the stopover sites during migration and in the wintering areas. These environmental factors are used as proxy of the shrike's food supply. We analysed detailed demographic data of some 4,600 individuals from 25 years with multistate capture-recapture and mixed models. Survival probabilities of juveniles and breeders of both sexes varied in parallel across time, suggesting that all cohorts were sensitive to similar causes of mortality. Reproductive performance increased with temperature and decreased with rainfall on the breeding area. Moreover, it increased with vegetation cover in the Sahelian stopover area used on autumn migration suggesting a carry-over effect. Arrival date was negatively affected by spring temperatures in the breeding area. Hence, demographic rates were affected by environmental factors on the breeding grounds, but also outside and elsewhere. This suggests that the shrike's population dynamics are driven by environmental factors operating at various scales of space and time. However, only a small amount of the temporal variation in demographic rates is explained by the environmental factors considered, suggesting that additional factors, such as those operating during migration, might be important.

Impact of density and environmental factors on population fluctuations in a migratory passerine.

1. Populations of plants and animals typically fluctuate because of the combined effects of density-dependent and density-independent processes. The study of these processes is complicated by the fact that population sizes are typically not known exactly, because population counts are subject to sampling variance. Although the existence of sampling variance is broadly acknowledged, relatively few studies on time-series data have accounted for it, which can result in wrong inferences about population processes. 2. To increase our understanding of population dynamics, we analysed time series from six Central European populations of the migratory red-backed shrike Lanius collurio by simultaneously assessing the strength of density dependence, process and sampling variance. In addition, we evaluated hypotheses predicting effects of factors presumed to operate on the breeding grounds, at stopover sites in eastern Africa during fall and spring migration and in the wintering grounds in southern Africa. We used both simple and state-space formulations of the Gompertz equation to model population size. 3. Across populations and modelling approaches, we found consistent evidence for negative density-dependent population regulation. Further, process variance contributed substantially to variation in population size, while sampling variance did not. Environmental conditions in eastern and southern Africa appear to influence breeding population size, as rainfall in the Sahel during fall migration and in the south African wintering areas were positively related to population size in the following spring in four of six populations. In contrast, environmental conditions in the breeding grounds were not related to population size. 4. Our findings suggest negative density-dependent regulation of red-backed shrike breeding populations and are consistent with the long-standing hypothesis that conditions in the African staging and wintering areas influence population numbers of species breeding in Europe. 5. This study highlights the importance of jointly investigating density-dependent and density-independent processes to improve our understanding of factors influencing population fluctuations in space and time.