It's about Her: Male Within-Season Movements Are Related to Mate Searching in a Songbird.

AbstractIn species with resource-defense mating systems (such as most temperate-breeding songbirds), male dispersal is often considered to be limited in both frequency and spatial extent. When dispersal occurs within a breeding season, the favored explanation is ecological resource tracking. In contrast, movements of male birds associated with temporary emigration, such as polyterritoriality (i.e., defense of an additional location after attracting a female in the initial territory), are usually attributed to mate searching. We suggest that male dispersal and polyterritoriality are functionally related and that mate searching may be a unifying hypothesis for predicting the within-season movements of male songbirds. Here, we test three key predictions derived from this hypothesis in Wood Warblers (Phylloscopus sibilatrix). We collected data on the spatial behavior of 107 males between 2017 and 2019 and related male movements to a new territory (in both a dispersal and a polyterritorial context) to mating potential in the current territory. Most males dispersed from their territories within days or weeks after failing to attract a female, despite occupying territories in apparently suitable habitat. Probability of polyterritoriality by paired males increased after the peak fertile period of their mate. Males never dispersed following nest predation if the female remained to renest. Thus, our data are consistent with the hypothesis that both movement types are functionally related to mate searching.

Wood warbler population dynamics in response to mast seeding regimes in Europe.

Mast seeding is the episodic, massive production of plant seeds synchronized over large areas. The resulting superabundance of seeds represents a resource pulse that can profoundly affect animal populations across trophic levels. Following years of high seed production, the abundance of both seed consumers and their predators increase. Higher predator abundance leads to increased predation pressure across the trophic web, impacting nonseed consumers such as the wood warbler Phylloscopus sibilatrix through increased nest predation after tree mast years. Over the past 30 years, the frequency of tree seed masts has increased, while wood warbler populations have declined in several regions of Europe. We hypothesized that increasing mast frequencies may have contributed to the observed population declines by creating suboptimal breeding conditions in years after masting. We measured reproductive output in four study areas in central Europe, which was between 0.61 and 1.24 fledglings lower in the years following masting than nonmasting. For each study area, we used matrix population models to predict population trends based on the estimated reproductive output and the local mast frequencies. We then compared the predicted with the observed population trends to assess if the frequency of mast years had contributed to the population dynamics. In Wielkopolska National Park (PL) and Hessen (DE), masting occurred on average only every 4 years and populations were stable or nearly so, whereas in Jura (CH) and Bialowieza National Park (PL), masting occurred every 2 and 2.5 years, respectively, and populations were declining. The simple matrix population models predicted the relative difference among local population trends over the past 10-20 years well, suggesting that the masting frequency may partly explain regional variation in population trends. Simulations suggest that further increases in mast frequency will lead to further declines in wood warbler populations. We show that changes in a natural process, such as mast seeding, may contribute to the decline in animal populations through cascading effects.

Habitat detection, habitat choice copying or mating benefits: What drives conspecific attraction in a nomadic songbird?

Conspecific attraction during habitat selection is common among animals, but the ultimate (i.e. fitness-related) reasons for this behaviour often remain enigmatic. We aimed to evaluate the following three hypotheses for conspecific attraction during the breeding season in male Wood Warblers (Phylloscopus sibilatrix): the habitat detection hypothesis, the habitat choice copying hypothesis and the female preference hypothesis. These hypotheses make different predictions with respect to the relative importance of social and nonsocial information during habitat assessment, and whether benefits accrue as a consequence of aggregation. We tested the above hypotheses using a combination of a 2-year playback experiment, spatial statistics and mate choice models. The habitat detection hypothesis was the most likely explanation for conspecific attraction and aggregation in male Wood Warblers, based on the following results: (1) males were attracted to conspecific song playbacks, but fine-scale habitat heterogeneity was the better predictor of spatial patterns in the density of settling males; (2) male pairing success did not increase, but instead slightly decreased, as connectivity with other males (i.e. the number and proximity of neighbouring males) increased. Our study highlights how consideration of the process by which animals detect and assess habitat, together with the potential fitness consequences of resulting aggregations, are important for understanding conspecific attraction and spatially clustered distributions.

Accounting for predator species identity reveals variable relationships between nest predation rate and habitat in a temperate forest songbird.

Nest predation is the primary cause of nest failure in most ground-nesting bird species. Investigations of relationships between nest predation rate and habitat usually pool different predator species. However, such relationships likely depend on the specific predator involved, partly because habitat requirements vary among predator species. Pooling may therefore impair our ability to identify conservation-relevant relationships between nest predation rate and habitat. We investigated predator-specific nest predation rates in the forest-dependent, ground-nesting wood warbler Phylloscopus sibilatrix in relation to forest area and forest edge complexity at two spatial scales and to the composition of the adjacent habitat matrix. We used camera traps at 559 nests to identify nest predators in five study regions across Europe. When analyzing predation data pooled across predator species, nest predation rate was positively related to forest area at the local scale (1000 m around nest), and higher where proportion of grassland in the adjacent habitat matrix was high but arable land low. Analyses by each predator species revealed variable relationships between nest predation rates and habitat. At the local scale, nest predation by most predators was higher where forest area was large. At the landscape scale (10,000 m around nest), nest predation by buzzards Buteo buteo was high where forest area was small. Predation by pine martens Martes martes was high where edge complexity at the landscape scale was high. Predation by badgers Meles meles was high where the matrix had much grassland but little arable land. Our results suggest that relationships between nest predation rates and habitat can depend on the predator species involved and may differ from analyses disregarding predator identity. Predator-specific nest predation rates, and their relationships to habitat at different spatial scales, should be considered when assessing the impact of habitat change on avian nesting success.

Numerical response of mammalian carnivores to rodents affects bird reproduction in temperate forests: A case of apparent competition?

Resource pulses such as mast seeding in temperate forests may affect interspecific interactions over multiple trophic levels and link different seed and nonseed consumers directly via predation or indirectly via shared predators. However, the nature and strength of interactions among species remain unknown for most resource pulse-driven ecosystems. We considered five hypotheses concerning the influence of resource pulses on the interactions between rodents, predators, and bird reproduction with data from northern Switzerland collected between 2010 and 2015. In high-rodent-abundance-years (HRAYs), wood warbler (Phylloscopus sibilatrix) nest survival was lower than in low-rodent-abundance-years, but rodents were not important nest predators, in contrast to rodent-hunting predators. The higher proportion of nests predated by rodent-hunting predators and their increased occurrence in HRAYs suggests a rodent-mediated aggregative numerical response of rodent-hunting predators, which incidentally prey on the wood warbler's ground nests. There was no evidence that rodent-hunting predators responded behaviorally by switching prey. Lastly, nest losses caused by nonrodent-hunting predators were not related to rodent abundance. We show that wood warblers and rodents are linked via shared predators in a manner consistent with apparent competition, where an increase of one species coincides with the decrease of another species mediated by shared predators. Mast seeding frequency and annual seed production appear to have increased over the past century, which may result in more frequent HRAYs and generally higher peaking rodent populations. The associated increase in the magnitude of apparent competition may thus at least to some extent explain the wood warbler's decline in much of Western Europe.

Rodent-avoidance, topography and forest structure shape territory selection of a forest bird.

BACKGROUND: Understanding the factors underlying habitat selection is important in ecological and evolutionary contexts, and crucial for developing targeted conservation action in threatened species. However, the key factors associated to habitat selection often remain poorly known. We evaluated hypotheses related to abiotic and biotic factors thought to affect territory selection of the wood warbler Phylloscopus sibilatrix, a passerine living in an unpredictable environment owing to irregular rodent outbreaks and showing long-term declines particularly in Western Europe. RESULTS: Comparing breeding territories to unoccupied areas located close-by revealed that territory occupancy in north-western Switzerland was positively related to slope steepness (topographic hypothesis supported) as well as to numbers of tussocks and trees, respectively, while it showed a unimodal relationship to cover of herb layer (forest structure hypothesis supported). Furthermore, a strong negative correlation between breeding territory occupancy and rodent numbers was found, suggesting that wood warblers avoid areas with high rodent densities (rodent-avoidance hypothesis supported). Comparing breeding territories to abandoned territories showed that breeding territories were located on steeper slopes (topography hypothesis supported), at larger distance from the forest edge (anthropogenic disturbance hypothesis supported) and harboured more trees (forest structure hypothesis supported) than abandoned territories. CONCLUSIONS: Aside from structural and topographic features of the habitat, wood warblers are affected by rodent numbers when settling, making habitat selection unpredictable from year to year. Forestry practices promoting relatively high tree densities, few bushes and an intermediate low-growing ground vegetation cover would enhance habitat quality for this declining passerine. In contrast, forestry practices aiming at increasing light in forests (selective thinning, group-felling) or keeping forest stands permanently covered with shrubs, bushes and trees of various sizes (continuous cover forestry) do not benefit the wood warbler.

Empirical evidence for source-sink populations: a review on occurrence, assessments and implications.

Assessing the role of local populations in a landscape context has become increasingly important in the fields of conservation biology and ecology. A growing number of studies attempt to determine the source-sink status of local populations. As the source-sink concept is commonly used for management decisions in nature conservation, accurate assessment approaches are crucial. Based on a systematic literature review of studies published between 2002 and 2013, we evaluated a priori predictions on methodological and biological factors that may influence the occurrence of source or sink populations. The review yielded 90 assessments from 73 publications that included qualitative and quantitative evidence for either source or sink population(s) for one or multiple species. Overall, sink populations tended to occur more often than source populations. Moreover, the occurrence of source or sink populations differed among taxonomic classes. Sinks were more often found than sources in mammals, while there was a non-significant trend for the opposite to be true for amphibians. Univariate and multivariate analyses showed that the occurrence of sources was positively related to connectivity of local populations. Our review furthermore highlights that more than 25 years after Pulliam's widely cited publication on 'sources, sinks, and population regulation', in-depth assessments of the source-sink status of populations based on combined consideration of demographic parameters such as fecundity, survival, emigration and immigration are still scarce. To increase our understanding of source-sink systems from ecological, evolutionary and conservation-related perspectives, we recommend that forthcoming studies on source-sink dynamics should pay more attention to the study design (i.e. connectivity of study populations) and that the assessment of the source-sink status of local populations is based on λ values calculated from demographic rates.

Reproductive Performance of a Declining Forest Passerine in Relation to Environmental and Social Factors: Implications for Species Conservation.

Identifying factors influencing a species' ecological niche and demography is a prerequisite for species conservation. However, our understanding of the interplay between demographic rates and biotic/abiotic factors is still poor for most species of conservation concern. We evaluated relevance of eight hypotheses relating to timing of breeding, temporal nest exposure, nest concealment, topography, tree structure, predation risk and disturbance, density dependence and weather for explaining variation in reproductive performance of the declining wood warbler Phylloscopus sibilatrix in northern Switzerland. Reproductive performance was monitored with cameras at 136 nests from 2010 to 2012 and was associated to temporal exposure, timing of breeding and concealment of nests. Daily nest survival was positively related to the number of grass and sedge tussocks, nest concealment and nest age. Clutch size and number of fledglings decreased, the later in the season a nest was initiated. Nest survival over an average nesting period of 31 days was 46.9 ± 0.07% (mean ± SE), daily nest survival rate was 0.976 ± 0.002. As for many ground-breeding birds, nest predation was the principal cause of nest failure, accounting for 79% of all nest losses. Conservation measures should aim at increasing the area of relatively homogenous forest stands featuring suitable habitats characterized by abundant and accessible grass and sedge tussocks. In managed forests, such conditions can be found in stands of middle age (i.e. pole wood) with little to no shrub layer.

Estimating Population Size for Capercaillie (Tetrao urogallus L.) with Spatial Capture-Recapture Models Based on Genotypes from One Field Sample.

We conducted a survey of an endangered and cryptic forest grouse, the capercaillie Tetrao urogallus, based on droppings collected on two sampling occasions in eight forest fragments in central Switzerland in early spring 2009. We used genetic analyses to sex and individually identify birds. We estimated sex-dependent detection probabilities and population size using a modern spatial capture-recapture (SCR) model for the data from pooled surveys. A total of 127 capercaillie genotypes were identified (77 males, 46 females, and 4 of unknown sex). The SCR model yielded a total population size estimate (posterior mean) of 137.3 capercaillies (posterior sd 4.2, 95% CRI 130-147). The observed sex ratio was skewed towards males (0.63). The posterior mean of the sex ratio under the SCR model was 0.58 (posterior sd 0.02, 95% CRI 0.54-0.61), suggesting a male-biased sex ratio in our study area. A subsampling simulation study indicated that a reduced sampling effort representing 75% of the actual detections would still yield practically acceptable estimates of total size and sex ratio in our population. Hence, field work and financial effort could be reduced without compromising accuracy when the SCR model is used to estimate key population parameters of cryptic species.

Habitat quality and geometry affect patch occupancy of two Orthopteran species.

Impacts of habitat loss and fragmentation on distribution and population size of many taxa are well established. In contrast, less is known about the role of within-patch habitat quality for the spatial dynamics of species, even though within-patch habitat quality may substantially influence the dynamics of population networks. We studied occurrence patterns of two Orthopteran species in relation to size, isolation and quality of habitat patches in an intensively managed agricultural landscape (16.65 km(2)) in the Swiss lowland. Occurrence of field crickets (Gryllus campestris) was positively related to patch size and negatively to the distance to the nearest occupied patch, two measures of patch geometry. Moreover, field crickets were more likely to occur in extensively managed meadows, meadows used at low intensity and meadows dominated by Poa pratensis, three measures of patch quality. Occurrence of the large gold grasshopper (Chrysochraon dispar) was negatively related to two measures of patch geometry, distance to the nearest occupied patch and perimeter index (ratio of perimeter length to patch area). Further, large gold grasshoppers were more likely to occupy patches close to water and patches with vegetation left uncut over winter, two measures of patch quality. Finally, examination of patch occupancy dynamics of field crickets revealed that patches colonized in 2009 and patches occupied in both 2005 and 2009 were larger, better connected and of other quality than patches remaining unoccupied and patches from which the species disappeared. The strong relationships between Orthopteran occurrence and aspects of patch geometry found in this study support the "area-and-isolation paradigm". Additionally, our study reveals the importance of patch quality for occurrence patterns of both species, and for patch occupancy dynamics in the field cricket. An increased understanding of patch occupancy patterns may be gained if inference is based on variables related to both habitat geometry and quality.

New support for an old hypothesis: density affects extra-pair paternity.

Density has been suggested to affect variation in extra-pair paternity (EPP) in avian mating systems, because increasing density promotes encounter rates and thus mating opportunities. However, the significance of density affecting EPP variation in intra- and interspecific comparisons has remained controversial, with more support from intraspecific comparisons. Neither experimental nor empirical studies have consistently provided support for the density hypothesis. Testing the density hypothesis is challenging because density measures may not necessarily reflect extra-pair mating opportunities, mate guarding efforts may covary with density, populations studied may differ in migratory behavior and/or climatic conditions, and variation in density may be insufficient. Accounting for these potentially confounding factors, we tested whether EPP rates within and among subpopulations of the reed bunting (Emberiza schoeniclus) were related to density. Our analyses were based on data from 13 subpopulations studied over 4 years. Overall, 56.4% of totally 181 broods contained at least one extra-pair young (EPY) and 37.1% of totally 669 young were of extra-pair origin. Roughly 90% of the extra-pair fathers were from the adjacent territory or from the territory after the next one. Within subpopulations, the proportion of EPY in broods was positively related to local breeding density. Similarly, among subpopulations, proportion of EPY was positively associated with population density. EPP was absent in subpopulations consisting of single breeding pairs, that is, without extra-pair mating opportunities. Our study confirms that density is an important biological factor, which significantly influences the amount of EPP within and among subpopulations, but also suggests that other mechanisms influence EPP beyond the variation explained by density.

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.

Behavioral plasticity allows short-term adjustment to a novel environment.

Many species are currently experiencing anthropogenically driven environmental changes. Among these changes, increasing noise levels are specifically a problem for species relying on acoustic communication. Recent evidence suggests that some species adjust their acoustic signals to man-made noise. However, it is unknown whether these changes occur through short-term and reversible adjustments by behavioral plasticity or through long-term adaptations by evolutionary change. Using behavioral observations and playback experiments, we show that male reed buntings (Emberiza schoeniclus) adjusted their songs immediately, singing at a higher minimum frequency and at a lower rate when noise levels were high. Our data showed that these changes in singing behavior were short-term adjustments of signal characteristics resulting from behavioral plasticity, rather than a long-term adaptation. However, more males remained unpaired at a noisy location than at a quiet location throughout the breeding season. Thus, phenotypic plasticity enables individuals to respond to environmental changes, but whether these short-term adjustments are beneficial remains to be seen.

Small and large wetland fragments are equally suited breeding sites for a ground-nesting passerine.

Large habitat fragments are generally thought to host more species and to offer more diverse and/or better quality habitats than small fragments. However, the importance of small fragments for population dynamics in general and for reproductive performance in particular is highly controversial. Using an information-theoretic approach, we examined reproductive performance and probability of local recruitment of color-banded reed buntings Emberiza schoeniclus in relation to the size of 18 wetland fragments in northeastern Switzerland over 4 years. We also investigated if reproductive performance and recruitment probability were density-dependent. None of the four measures of reproductive performance (laying date, nest failure probability, fledgling production per territory, fledgling condition) nor recruitment probability were found to be related to wetland fragment size. In terms of fledgling production, however, fragment size interacted with year, indicating that small fragments were better reproductive grounds in some years than large fragments. Reproductive performance and recruitment probability were not density-dependent. Our results suggest that small fragments are equally suited as breeding grounds for the reed bunting as large fragments and should therefore be managed to provide a habitat for this and other specialists occurring in the same habitat. Moreover, large fragments may represent sinks in specific years because a substantial percentage of all breeding pairs in our study area breed in large fragments, and reproductive failure in these fragments due to the regularly occurring floods may have a much stronger impact on regional population dynamics than comparable events in small fragments.

Ecological and social effects on reproduction and local recruitment in the red-backed shrike.

Numerous hypotheses have been proposed to explain variation in reproductive performance and local recruitment of animals. While most studies have examined the influence of one or a few social and ecological factors on fitness traits, comprehensive analyses jointly testing the relative importance of each of many factors are rare. We investigated how a multitude of environmental and social conditions simultaneously affected reproductive performance and local recruitment of the red-backed shrike Lanius collurio (L.). Specifically, we tested hypotheses relating to timing of breeding, parental quality, nest predation, nest site selection, territory quality, intraspecific density and weather. Using model selection procedures, predictions of each hypothesis were first analysed separately, before a full model was constructed including variables selected in the single-hypothesis tests. From 1988 to 1992, 50% of 332 first clutches produced at least one fledgling, while 38.7% of 111 replacement clutches were successful. Timing of breeding, nest site selection, predation pressure, territory quality and intraspecific density influenced nest success in the single-hypothesis tests. The full model revealed that nest success was negatively associated with laying date, intraspecific density, and year, while nest success increased with nest concealment. Number of fledglings per successful nest was only influenced by nest concealment: better-camouflaged nests produced more fledglings. Probability of local recruitment was related to timing of breeding, parental quality and territory quality in the single-hypothesis tests. The full models confirmed the important role of territory quality for recruitment probability. Our results suggest that reproductive performance, and particularly nest success, of the red-backed shrike is primarily affected by timing of breeding, nest site selection, and intraspecific density. This study highlights the importance of considering many factors at the same time, when trying to evaluate their relative contributions to fitness and life history evolution.

Genetic and environmental influences on natal dispersal distance in a resident bird species.

We analyzed more than 1,600 dispersal events from two populations of a North American cooperatively breeding woodpecker species to determine what factors influence natal dispersal distance and whether distance traveled affects reproduction later in life. We found significant heritability of natal dispersal distance, in both males and females, indicating substantial additive genetic variance for this behavioral trait. Natal dispersal distance additionally was affected by social and ecological factors: individuals dispersing in their first year of life moved longer distances than those staying on their natal site as helpers for a prolonged time prior to dispersal, and increasing territory isolation led to longer dispersal distances. Successful dispersers incurred fitness costs, with lifetime fledgling production (in both sexes) and lifetime production of recruits to the breeding population (in females only) decreasing with increasing natal dispersal distance. We conclude that natal dispersal distance has a genetic basis but is modulated by environmental and social factors and that natal dispersal distance in this species is (currently) under selection.

Inbreeding and experience affect response to climate change by endangered woodpeckers.

In recent decades, female red-cockaded woodpeckers (Picoides borealis) have laid eggs increasingly earlier in response to a changing climate, as has been observed in several other bird species breeding at north temperate latitudes. Within each year, females that lay earlier are more productive than females that lay later. However, inexperienced females, experienced females who change mates and inbred birds have not adjusted to the changing climate by laying earlier, and have suffered reproductive costs as a result. Failure to respond to global climate change may be a further example of the reduced ability of inbred animals to respond to environmental challenges.